Title: The Evolution of Man — Volume 1
Author: Haeckel, Ernst
Language: English
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THE EVOLUTION OF MAN

A POPULAR SCIENTIFIC STUDY


BY

ERNST HAECKEL

VOLUME 1.

HUMAN EMBRYOLOGY OR ONTOGENY.


TRANSLATED FROM THE FIFTH (ENLARGED) EDITION BY JOSEPH MCCABE.


[ISSUED FOR THE RATIONALIST PRESS ASSOCIATION, LIMITED.]



WATTS & CO.,
17, JOHNSONS COURT, FLEET STREET, LONDON, E.C.
1912.



CONTENTS OF VOLUME 1.


LIST OF ILLUSTRATIONS.

GLOSSARY.

TRANSLATOR'S PREFACE.

TABLE: CLASSIFICATION OF THE ANIMAL WORLD.


CHAPTER 1.1. THE FUNDAMENTAL LAW OF ORGANIC EVOLUTION.


CHAPTER 1.2. THE OLDER EMBRYOLOGY.


CHAPTER 1.3. MODERN EMBRYOLOGY.


CHAPTER 1.4. THE OLDER PHYLOGENY.


CHAPTER 1.5. THE MODERN SCIENCE OF EVOLUTION.


CHAPTER 1.6. THE OVUM AND THE AMOEBA.


CHAPTER 1.7. CONCEPTION.


CHAPTER 1.8. THE GASTRAEA THEORY.


CHAPTER 1.9. THE GASTRULATION OF THE VERTEBRATE.


CHAPTER 1.10. THE COELOM THEORY.


CHAPTER 1.11. THE VERTEBRATE CHARACTER OF MAN.


CHAPTER 1.12. THE EMBRYONIC SHIELD AND GERMINATIVE AREA.


CHAPTER 1.13. DORSAL BODY AND VENTRAL BODY.


CHAPTER 1.14. THE ARTICULATION OF THE BODY.


CHAPTER 1.15. FOETAL MEMBRANES AND CIRCULATION.



LIST OF ILLUSTRATIONS.

PORTRAIT OF ERNST HAECKEL FROM THE PAINTING BY FRANZ VON LEUBACH, 1899

FIGURE 1.1. THE HUMAN OVUM.

FIGURE 1.2. STEM-CELL OF AN ECHINODERM.

FIGURE 1.3. THREE EPITHELIAL CELLS.

FIGURE 1.4. FIVE SPINY OR GROOVED CELLS.

FIGURE 1.5. TEN LIVER-CELLS.

FIGURE 1.6. NINE STAR-SHAPED BONE-CELLS.

FIGURE 1.7. ELEVEN STAR-SHAPED CELLS.

FIGURE 1.8. UNFERTILISED OVUM OF AN ECHINODERM.

FIGURE 1.9. A LARGE BRANCHING NERVE-CELL.

FIGURE 1.10. BLOOD-CELLS.

FIGURE 1.11. INDIRECT OR MITOTIC CELL-DIVISION.

FIGURE 1.12. MOBILE CELLS.

FIGURE 1.13. OVA OF VARIOUS ANIMALS.

FIGURE 1.14. THE HUMAN OVUM.

FIGURE 1.15. FERTILISED OVUM OF HEN.

FIGURE 1.16. A CREEPING AMOEBA.

FIGURE 1.17. DIVISION OF AN AMOEBA.

FIGURE 1.18. OVUM OF A SPONGE.

FIGURE 1.19. BLOOD-CELLS, OR PHAGOCYTES.

FIGURE 1.20. SPERMIA OR SPERMATOZOA.

FIGURE 1.21. SPERMATOZOA OF VARIOUS ANIMALS.

FIGURE 1.22. A SINGLE HUMAN SPERMATOZOON.

FIGURE 1.23. FERTILISATION OF THE OVUM.

FIGURE 1.24. IMPREGNATED ECHINODERM OVUM.

FIGURE 1.25. IMPREGNATION OF THE STAR-FISH OVUM.

FIGURES 1.26 AND 1.27. IMPREGNATION OF SEA-URCHIN OVUM.

FIGURE 1.28. STEM-CELL OF A RABBIT.

FIGURE 1.29. GASTRULATION OF A CORAL.

FIGURE 1.30. GASTRULA OF A GASTRAEAD.

FIGURE 1.31. GASTRULA OF A WORM.

FIGURE 1.32. GASTRULA OF AN ECHINODERM.

FIGURE 1.33. GASTRULA OF AN ARTHROPOD.

FIGURE 1.34. GASTRULA OF A MOLLUSC.

FIGURE 1.35. GASTRULA OF A VERTEBRATE.

FIGURE 1.36. GASTRULA OF A LOWER SPONGE.

FIGURE 1.37. CELLS FROM THE PRIMARY GERMINAL LAYERS.

FIGURE 1.38. GASTRULATION OF THE AMPHIOXUS.

FIGURE 1.39. GASTRULA OF THE AMPHIOXUS.

FIGURE 1.40. CLEAVAGE OF THE FROG'S OVUM.

FIGURES 1.41 TO 1.44. SECTIONS OF FERTILISED TOAD OVUM.

FIGURES 1.45 TO 1.48. GASTRULATION OF THE SALAMANDER.

FIGURE 1.49. SEGMENTATION OF THE LAMPREY.

FIGURE 1.50. GASTRULATION OF THE LAMPREY.

FIGURE 1.51. GASTRULATION OF CERATODUS.

FIGURE 1.52. OVUM OF A DEEP-SEA BONY FISH.

FIGURE 1.53. SEGMENTATION OF A BONY FISH.

FIGURE 1.54. DISCOID GASTRULA OF A BONY FISH.

FIGURES 1.55 AND 1.56. SECTIONS OF BLASTULA OF SHARK.

FIGURE 1.57. DISCOID SEGMENTATION OF BIRD'S OVUM.

FIGURES 1.58 TO 1.61. GASTRULATION OF THE BIRD.

FIGURE 1.62. GERMINAL DISK OF THE LIZARD.

FIGURES 1.63 AND 1.64. GASTRULATION OF THE OPOSSUM.

FIGURES 1.65 TO 1.67. GASTRULATION OF THE OPOSSUM.

FIGURES 1.68 TO 1.71. GASTRULATION OF THE RABBIT.

FIGURE 1.72. GASTRULA OF THE PLACENTAL MAMMAL.

FIGURE 1.73. GASTRULA OF THE RABBIT.

FIGURES 1.74 AND 1.75. DIAGRAM OF THE FOUR SECONDARY GERMINAL LAYERS.

FIGURES 1.76 AND 1.77. COELOMULA OF SAGITTA.

FIGURE 1.78. SECTION OF YOUNG SAGITTA.

FIGURES 1.79 AND 1.80. SECTION OF AMPHIOXUS-LARVAE.

FIGURES 1.81 AND 1.82. SECTION OF AMPHIOXUS-LARVAE.

FIGURES 1.83 AND 1.84. CHORDULA OF THE AMPHIOXUS.

FIGURES 1.85 AND 1.86. CHORDULA OF THE AMPHIBIA.

FIGURES 1.87 AND 1.88. SECTION OF COELOMULA-EMBRYOS OF VERTEBRATES.

FIGURES 1.89 AND 1.90. SECTION OF COELOMULA-EMBRYO OF TRITON.

FIGURE 1.91. DORSAL PART OF THREE TRITON-EMBRYOS.

FIGURE 1.92. CHORDULA-EMBRYO OF A BIRD.

FIGURE 1.93. VERTEBRATE-EMBRYO OF A BIRD.

FIGURES 1.94 AND 1.95. SECTION OF THE PRIMITIVE STREAK OF A CHICK.

FIGURE 1.96. SECTION OF THE PRIMITIVE GROOVE OF A RABBIT.

FIGURE 1.97. SECTION OF PRIMITIVE MOUTH OF A HUMAN EMBRYO.

FIGURES 1.98 TO 1.102. THE IDEAL PRIMITIVE VERTEBRATE.

FIGURE 1.103. REDUNDANT MAMMARY GLANDS.

FIGURE 1.104. A GREEK GYNECOMAST.

FIGURE 1.105. SEVERANCE OF THE DISCOID MAMMAL EMBRYO.

FIGURES 1.106 AND 1.107. THE VISCERAL EMBRYONIC VESICLE.

FIGURE 1.108. FOUR ENTODERMIC CELLS.

FIGURE 1.109. TWO ENTODERMIC CELLS.

FIGURES 1.110 TO 1.114. OVUM OF A RABBIT.

FIGURES 1.115 TO 1.118. EMBRYONIC VESICLE OF A RABBIT.

FIGURE 1.119. SECTION OF THE GASTRULA OF FOUR VERTEBRATES.

FIGURES 1.120 TO 1.123. EMBRYONIC SHIELD OF A RABBIT.

FIGURE 1.124. COELOMULA OF THE AMPHIOXUS.

FIGURE 1.125. CHORDULA OF A FROG.

FIGURE 1.126. SECTION OF FROG-EMBRYO.

FIGURES 1.127 AND 1.128. DORSAL SHIELD OF A CHICK.

FIGURE 1.129. SECTION OF HIND END OF A CHICK.

FIGURE 1.130. GERMINAL AREA OF THE RABBIT.

FIGURE 1.131. EMBRYO OF THE OPOSSUM.

FIGURE 1.132. EMBRYONIC SHIELD OF THE RABBIT.

FIGURE 1.133. HUMAN EMBRYO AT THE SANDAL-STAGE.

FIGURE 1.134. EMBRYONIC SHIELD OF RABBIT.

FIGURE 1.135. EMBRYONIC SHIELD OF OPOSSUM.

FIGURE 1.136. EMBRYONIC DISK OF A CHICK.

FIGURE 1.137. EMBRYONIC DISK OF A HIGHER VERTEBRATE.

FIGURES 1.138 TO 1.142. SECTIONS OF MATURING MAMMAL EMBRYO.

FIGURES 1.143 TO 1.146. SECTIONS OF EMBRYONIC CHICKS.

FIGURE 1.147. SECTION OF EMBRYONIC CHICK.

FIGURE 1.148. SECTION OF FORE-HALF OF CHICK-EMBRYO.

FIGURES 1.149 AND 1.150. SECTIONS OF HUMAN EMBRYOS.

FIGURE 1.151. SECTION OF A SHARK-EMBRYO.

FIGURE 1.152. SECTION OF A DUCK-EMBRYO.

FIGURES 1.153 TO 1.155. SOLE-SHAPED EMBRYONIC DISK OF CHICK.

FIGURES 1.156 AND 1.157. EMBRYO OF THE AMPHIOXUS.

FIGURES 1.158 TO 1.160. EMBRYO OF THE AMPHIOXUS.

FIGURES 1.161 AND 1.162. SECTIONS OF SHARK-EMBRYOS.

FIGURE 1.163. SECTION OF A TRITON-EMBRYO.

FIGURES 1.164 TO 1.166. VERTEBRAE.

FIGURE 1.167. HEAD OF A SHARK-EMBRYO.

FIGURES 1.168 AND 1.169. HEAD OF A CHICK-EMBRYO.

FIGURE 1.170. HEAD OF A DOG-EMBRYO.

FIGURE 1.171. HUMAN EMBRYO OF THE FOURTH WEEK.

FIGURE 1.172. SECTION OF SHOULDER OF CHICK-EMBRYO.

FIGURE 1.173. SECTION OF PELVIC REGION OF CHICK-EMBRYO.

FIGURE 1.174. DEVELOPMENT OF THE LIZARD'S LEGS.

FIGURE 1.175. HUMAN-EMBRYO FIVE WEEKS OLD.

FIGURES 1.176 TO 1.178. EMBRYOS OF THE BAT.

FIGURE 1.179. HUMAN EMBRYOS.

FIGURE 1.180. HUMAN EMBRYO OF THE FOURTH WEEK.

FIGURE 1.181. HUMAN EMBRYO OF THE FIFTH WEEK.

FIGURE 1.182. SECTION OF TAIL OF HUMAN EMBRYO.

FIGURES 1.183 AND 1.184. HUMAN EMBRYO DISSECTED.

FIGURE 1.185. MISS JULIA PASTRANA.

FIGURES 1.186 TO 1.190. HUMAN EMBRYOS.

FIGURE 1.191. HUMAN EMBRYOS OF SIXTEEN TO EIGHTEEN DAYS.

FIGURES 1.192 AND 1.193. HUMAN EMBRYO OF FOURTH WEEK.

FIGURE 1.194. HUMAN EMBRYO WITH ITS MEMBRANES.

FIGURE 1.195. DIAGRAM OF THE EMBRYONIC ORGANS.

FIGURE 1.196. SECTION OF THE PREGNANT WOMB.

FIGURE 1.197. EMBRYO OF SIAMANG-GIBBON.

FIGURE 1.198. SECTION OF PREGNANT WOMB.

FIGURES 1.199 AND 1.200. HUMAN FOETUS AND PLACENTA.

FIGURE 1.201. VITELLINE VESSELS IN GERMINATIVE AREA.

FIGURE 1.202. BOAT-SHAPED EMBRYO OF THE DOG.

FIGURE 1.203. LAR OR WHITE-HANDED GIBBON.

FIGURE 1.204. YOUNG ORANG.

FIGURE 1.205. WILD ORANG.

FIGURE 1.206. BALD-HEADED CHIMPANZEE.

FIGURE 1.207. FOETAL MEMBRANES AND CIRCULATION.

FIGURE 1.208. FEMALE GORILLA.

FIGURE 1.209. MALE GIANT-GORILLA.


GLOSSARY.

ACRANIA: animals without skull (cranium).

ANTHROPOGENY: the evolution (genesis) of man (anthropos).

ANTHROPOLOGY: the science of man.

ARCHI-: (in compounds) the first or typical--as, archi-cytula,
archi-gastrula, etc.

BIOGENY: the science of the genesis of life (bios).

BLAST-: (in compounds) pertaining to the early embryo (blastos = a
bud); hence:--
Blastoderm: skin (derma) or enclosing layer of the embryo.
Blastosphere: the embryo in the hollow sphere stage.
Blastula: same as preceding.
Epiblast: the outer layer of the embryo (ectoderm).
Hypoblast: the inner layer of the embryo (endoderm).

BRANCHIAL: pertaining to the gills (branchia).

CARYO-: (in compounds) pertaining to the nucleus (caryon); hence:--
Caryokineses: the movement of the nucleus.
Caryolysis: dissolution of the nucleus.
Caryoplasm: the matter of the nucleus.

CENTROLECITHAL: see under LECITH-.

CHORDARIA and CHORDONIA: animals with a dorsal chord or back-bone.

COELOM or COELOMA: the body-cavity in the embryo; hence:--
Coelenterata: animals without a body-cavity.
Coelomaria: animals with a body-cavity.
Coelomation: formation of the body-cavity.

CYTO-: (in compounds) pertaining to the cell (cytos); hence:--
Cytoblast: the nucleus of the cell.
Cytodes: cell-like bodies, imperfect cells.
Cytoplasm: the matter of the body of the cell.
Cytosoma: the body (soma) of the cell.

CRYPTORCHISM: abnormal retention of the testicles in the body.

DEUTOPLASM: see PLASM.

DUALISM: the belief in the existence of two entirely distinct
principles (such as matter and spirit).

DYSTELEOLOGY: the science of those features in organisms which refute
the "design-argument."

ECTODERM: the outer (ekto) layer of the embryo.

ENTODERM: the inner (ento) layer of the embryo.

EPIDERM: the outer layer of the skin.

EPIGENESIS: the theory of gradual development of organs in the embryo.

EPIPHYSIS: the third or central eye in the early vertebrates.

EPISOMA: see SOMA.

EPITHELIA: tissues covering the surface of parts of the body (such as
the mouth, etc.)

GONADS: the sexual glands.

GONOCHORISM: separation of the male and female sexes.

GONOTOMES: sections of the sexual glands.

GYNECOMAST: a male with the breasts (masta) of a woman (gyne).

HEPATIC: pertaining to the liver (hepar).

HOLOBLASTIC: embryos in which the animal and vegetal cells divide
equally (holon = whole).

HYPERMASTISM: the possession of more than the normal breasts (masta).

HYPOBRANCHIAL: underneath (hypo) the gills.

HYPOPHYSIS: sensitive-offshoot from the brain in the vertebrate.

HYPOSOMA: see SOMA.

LECITH-: pertaining to the yelk (lecithus); hence:--
Centrolecithal: eggs with the yelk in the centre.
Lecithoma: the yelk-sac.
Telolecithal: eggs with the yelk at one end.

MEROBLASTIC: cleaving in part (meron) only.

META-: (in compounds) the "after" or secondary stage; hence:--
Metagaster: the secondary or permanent gut (gaster).
Metaplasm: secondary or differentiated plasm.
Metastoma: the secondary or permanent mouth (stoma).
Metazoa: the higher or later animals, made up of many cells.
Metovum: the mature or advanced ovum.

METAMERA: the segments into which the embryo breaks up.

METAMERISM: the segmentation of the embryo.

MONERA: the most primitive of the unicellular organisms.

MONISM: belief in the fundamental unity of all things.

MORPHOLOGY: the science of organic forms (generally equivalent to
anatomy).

MYOTOMES: segments into which the muscles break up.

NEPHRA: the kidneys; hence:--
Nephridia: the rudimentary kidney-organs.
Nephrotomes: the segments of the developing kidneys.

ONTOGENY: the science of the development of the individual (generally
equivalent to embryology).

PERIGENESIS: the genesis of the movements in the vital particles.

PHAGOCYTES: cells that absorb food (phagein = to eat).

PHYLOGENY: the science of the evolution of species (phyla).

PLANOCYTES: cells that move about (planein).

PLASM: the colloid or jelly-like matter of which organisms are
composed; hence:--
Caryoplasm: the matter of the nucleus (caryon).
Cytoplasm: the matter of the body of the cell.
Deutoplasm: secondary or differentiated plasm.
Metaplasm: secondary or differentiated plasm.
Protoplasm: primitive or undifferentiated plasm.

PLASSON: the simplest form of plasm.

PLASTIDULES: small particles of plasm.

POLYSPERMISM: the penetration of more than one sperm-cell into the ovum.

PRO- or PROT: (in compounds) the earlier form (opposed to META); hence:--
Prochorion: the first form of the chorion.
Progaster: the first or primitive stomach.
Pronephridia: the earlier form of the kidneys.
Prorenal: the earlier form of the kidneys.
Prostoma: the first or primitive mouth.
Protists: the earliest or unicellular organisms.
Provertebrae: the earliest phase of the vertebrae.
Protophyta: the primitive or unicellular plants.
Protoplasm: undifferentiated plasm.
Protozoa: the primitive or unicellular animals.

RENAL: pertaining to the kidneys (renes).

SCATULATION: packing or boxing-up (scatula = a box).

SCLEROTOMES: segments into which the primitive skeleton falls.

SOMA: the body; hence:--
Cytosoma: the body of the cell (cytos).
Episoma: the upper or back-half of the embryonic body.
Somites: segments of the embryonic body.
Hyposoma: the under or belly-half of the embryonic body.

TELEOLOGY: the belief in design and purpose (telos) in nature.

TELOLECITHAL: see LECITH-.

UMBILICAL: pertaining to the navel (umbilicus).

VITELLINE: pertaining to the yelk (vitellus).

***



PREFACE.

[BY JOSEPH MCCABE.]

The work which we now place within the reach of every reader of the
English tongue is one of the finest productions of its distinguished
author. The first edition appeared in 1874. At that time the
conviction of man's natural evolution was even less advanced in
Germany than in England, and the work raised a storm of controversy.
Theologians--forgetting the commonest facts of our individual
development--spoke with the most profound disdain of the theory that a
Luther or a Goethe could be the outcome of development from a tiny
speck of protoplasm. The work, one of the most distinguished of them
said, was "a fleck of shame on the escutcheon of Germany." To-day its
conclusion is accepted by influential clerics, such as the Dean of
Westminster, and by almost every biologist and anthropologist of
distinction in Europe. Evolution is not a laboriously reached
conclusion, but a guiding truth, in biological literature to-day.

There was ample evidence to substantiate the conclusion even in the
first edition of the book. But fresh facts have come to light in each
decade, always enforcing the general truth of man's evolution, and at
times making clearer the line of development. Professor Haeckel
embodied these in successive editions of his work. In the fifth
edition, of which this is a translation, reference will be found to
the very latest facts bearing on the evolution of man, such as the
discovery of the remarkable effect of mixing human blood with that of
the anthropoid ape. Moreover, the ample series of illustrations has
been considerably improved and enlarged; there is no scientific work
published, at a price remotely approaching that of the present
edition, with so abundant and excellent a supply of illustrations.
When it was issued in Germany, a few years ago, a distinguished
biologist wrote in the Frankfurter Zeitung that it would secure
immortality for its author, the most notable critic of the idea of
immortality. And the Daily Telegraph reviewer described the English
version as a "handsome edition of Haeckel's monumental work," and "an
issue worthy of the subject and the author."

The influence of such a work, one of the most constructive that
Haeckel has ever written, should extend to more than the few hundred
readers who are able to purchase the expensive volumes of the original
issue. Few pages in the story of science are more arresting and
generally instructive than this great picture of "mankind in the
making." The horizon of the mind is healthily expanded as we follow
the search-light of science down the vast avenues of past time, and
gaze on the uncouth forms that enter into, or illustrate, the line of
our ancestry. And if the imagination recoils from the strange and
remote figures that are lit up by our search-light, and hesitates to
accept them as ancestral forms, science draws aside another veil and
reveals another picture to us. It shows us that each of us passes, in
our embryonic development, through a series of forms hardly less
uncouth and unfamiliar. Nay, it traces a parallel between the two
series of forms. It shows us man beginning his existence, in the ovary
of the female infant, as a minute and simple speck of jelly-like
plasm. It shows us (from analogy) the fertilised ovum breaking into a
cluster of cohering cells, and folding and curving, until the
limb-less, head-less, long-tailed foetus looks like a worm-shaped
body. It then points out how gill-slits and corresponding
blood-vessels appear, as in a lowly fish, and the fin-like extremities
bud out and grow into limbs, and so on; until, after a very clear
ape-stage, the definite human form emerges from the series of
transformations.

It is with this embryological evidence for our evolution that the
present volume is concerned. There are illustrations in the work that
will make the point clear at a glance. Possibly TOO clear; for the
simplicity of the idea and the eagerness to apply it at every point
have carried many, who borrow hastily from Haeckel, out of their
scientific depth. Haeckel has never shared their errors, nor
encouraged their superficiality. He insists from the outset that a
complete parallel could not possibly be expected. Embryonic life
itself is subject to evolution. Though there is a general and
substantial law--as most of our English and American authorities
admit--that the embryonic series of forms recalls the ancestral series
of forms, the parallel is blurred throughout and often distorted. It
is not the obvious resemblance of the embryos of different animals,
and their general similarity to our extinct ancestors in this or that
organ, on which we must rest our case. A careful study must be made of
the various stages through which all embryos pass, and an effort made
to prove their real identity and therefore genealogical relation.

This is a task of great subtlety and delicacy. Many scientists have
worked at it together with Professor Haeckel--I need only name our own
Professor Balfour and Professor Ray Lankester--and the scheme is
fairly complete. But the general reader must not expect that even so
clear a writer as Haeckel can describe these intricate processes
without demanding his very careful attention. Most of the chapters in
the present volume (and the second volume will be less difficult) are
easily intelligible to all; but there are points at which the line of
argument is necessarily subtle and complex. In the hope that most
readers will be induced to master even these more difficult chapters,
I will give an outline of the characteristic argument of the work.
Haeckel's distinctive services in regard to man's evolution have been:

1. The construction of a complete ancestral tree, though, of course,
some of the stages in it are purely conjectural, and not final.

2. The tracing of the remarkable reproduction of ancestral forms in
the embryonic development of the individual. Naturally, he has not
worked alone in either department.

The second volume of this work will embody the first of these two
achievements; the present one is mainly concerned with the latter. It
will be useful for the reader to have a synopsis of the argument and
an explanation of some of the chief terms invented or employed by the
author.

The main theme of the work is that, in the course of their embryonic
development, all animals, including man, pass roughly and rapidly
through a series of forms which represents the succession of their
ancestors in the past. After a severe and extensive study of embryonic
phenomena, Haeckel has drawn up a "law" (in the ordinary scientific
sense) to this effect, and has called it "the biogenetic law," or the
chief law relating to the evolution (genesis) of life (bios). This law
is widely and increasingly accepted by embryologists and zoologists.
It is enough to quote a recent declaration of the great American
zoologist, President D. Starr Jordan: "It is, of course, true that the
life-history of the individual is an epitome of the life-history of
the race"; while a distinguished German zoologist (Sarasin) has
described it as being of the same use to the biologist as spectrum
analysis is to the astronomer.

But the reproduction of ancestral forms in the course of the embryonic
development is by no means always clear, or even always present. Many
of the embryonic phases do not recall ancestral stages at all. They
may have done so originally, but we must remember that the embryonic
life itself has been subject to adaptive changes for millions of
years. All this is clearly explained by Professor Haeckel. For the
moment, I would impress on the reader the vital importance of fixing
the distinction from the start. He must thoroughly familiarise himself
with the meaning of five terms.

BIOGENY is the development of life in general (both in the individual
and the species), or the sciences describing it.

ONTOGENY is the development (embryonic and post-embryonic) of the
individual (on), or the science describing it.

PHYLOGENY is the development of the race or stem (phulon), or the
science describing it.

Roughly, ontogeny may be taken to mean embryology, and phylogeny what
we generally call evolution.

Further, the embryonic phenomena sometimes reproduce ancestral forms,
and they are then called PALINGENETIC (from palin = again): sometimes
they do not recall ancestral forms, but are later modifications due to
adaptation, and they are then called CENOGENETIC (from kenos = new or
foreign).

These terms are now widely used, but the reader of Haeckel must
understand them thoroughly.

The first five chapters are an easy account of the history of
embryology and evolution. The sixth and seventh give an equally clear
account of the sexual elements and the process of conception. But some
of the succeeding chapters must deal with embryonic processes so
unfamiliar, and pursue them through so wide a range of animals in a
brief space, that, in spite of the 200 illustrations, they will offer
difficulty to many a reader. As our aim is to secure, not a
superficial acquiescence in conclusions, but a fair comprehension of
the truths of science, we have retained these chapters. However, I
will give a brief and clear outline of the argument, so that the
reader with little leisure may realise their value.

When the animal ovum (egg-cell) has been fertilised, it divides and
subdivides until we have a cluster of cohering cells, externally not
unlike a raspberry or mulberry. This is the morula (= mulberry) stage.
The cluster becomes hollow, or filled with fluid in the centre, all
the cells rising to the surface. This is the blastula (hollow ball)
stage. One half of the cluster then bends or folds in upon the other,
as one might do with a thin indiarubber ball, and we get a vase-shaped
body with hollow interior (the first stomach, or "primitive gut"), an
open mouth (the first or "primitive mouth"), and a wall composed of
two layers of cells (two "germinal layers"). This is the gastrula
(stomach) stage, and the process of its formation is called
gastrulation. A glance at the illustration (Figure 1.29) will make
this perfectly clear.

So much for the embryonic process in itself. The application to
evolution has been a long and laborious task. Briefly, it was
necessary to show that ALL the multicellular animals passed through
these three stages, so that our biogenetic law would enable us to
recognise them as reminiscences of ancestral forms. This is the work
of Chapters 1.8 and 1.9. The difficulty can be realised in this way:
As we reach the higher animals the ovum has to take up a large
quantity of yelk, on which it may feed in developing. Think of the
bird's "egg." The effect of this was to flatten the germ (the morula
and blastula) from the first, and so give, at first sight, a totally
different complexion to what it has in the lowest animals. When we
pass the reptile and bird stage, the large yelk almost disappears (the
germ now being supplied with blood by the mother), but the germ has
been permanently altered in shape, and there are now a number of new
embryonic processes (membranes, blood-vessel connections, etc.). Thus
it was no light task to trace the identity of this process of
gastrulation in all the animals. It has been done, however; and with
this introduction the reader will be able to follow the proof. The
conclusion is important. If all animals pass through the curious
gastrula stage, it must be because they all had a common ancestor of
that nature. To this conjectural ancestor (it lived before the period
of fossilisation begins) Haeckel gives the name of the Gastraea, and
in the second volume we shall see a number of living animals of this
type ("gastraeads").

The line of argument is the same in the next chapter. After laborious
and careful research (though this stage is not generally admitted in
the same sense as the previous one), a fourth common stage was
discovered, and given the name of the Coelomula. The blastula had one
layer of cells, the blastoderm (derma = skin): the gastrula two
layers, the ectoderm ("outer skin") and entoderm ("inner skin"). Now a
third layer (mesoderm = middle skin) is formed, by the growth inwards
of two pouches or folds of the skin. The pouches blend together, and
form a single cavity (the body cavity, or coelom), and its two walls
are two fresh "germinal layers." Again, the identity of the process
has to be proved in all the higher classes of animals, and when this
is done we have another ancestral stage, the Coelomaea.

The remaining task is to build up the complex frame of the higher
animals--always showing the identity of the process (on which the
evolutionary argument depends) in enormously different conditions of
embryonic life--out of the four "germinal layers." Chapter 1.9
prepares us for the work by giving us a very clear account of the
essential structure of the back-boned (vertebrate) animal, and the
probable common ancestor of all the vertebrates (a small fish of the
lancelet type). Chapters 1.11 to 1.14 then carry out the construction
step by step. The work is now simpler, in the sense that we leave all
the invertebrate animals out of account; but there are so many organs
to be fashioned out of the four simple layers that the reader must
proceed carefully. In the second volume each of these organs will be
dealt with separately, and the parallel will be worked out between its
embryonic and its phylogenetic (evolutionary) development. The general
reader may wait for this for a full understanding. But in the meantime
the wonderful story of the construction of all our organs in the
course of a few weeks (the human frame is perfectly formed, though
less than two inches in length, by the twelfth week) from so simple a
material is full of interest. It would be useless to attempt to
summarise the process. The four chapters are themselves but a summary
of it, and the eighty fine illustrations of the process will make it
sufficiently clear. The last chapter carries the story on to the point
where man at last parts company with the anthropoid ape, and gives a
full account of the membranes or wrappers that enfold him in the womb,
and the connection with the mother.

In conclusion, I would urge the reader to consult, at his free library
perhaps, the complete edition of this work, when he has read the
present abbreviated edition. Much of the text has had to be condensed
in order to bring out the work at our popular price, and the beautiful
plates of the complete edition have had to be omitted. The reader will
find it an immense assistance if he can consult the library edition.

JOSEPH MCCABE.

Cricklewood, March, 1906.

***


HAECKEL'S CLASSIFICATION OF THE ANIMAL WORLD.

UNICELLULAR ANIMALS (PROTOZOA).

1. Unnucleated.

Bacteria.
Protamoebae.

Monera.

2. Nucleated.

2A. Rhizopoda.

Amoebina.
Radiolaria.

2B. Infusoria.

Flagellata.
Ciliata.

3. Cell-Colonies.

Catallacta.
Blastaeada.

MULTICELLULAR ANIMALS (METAZOA).

1. COELENTERIA, COELENTERATA, OR ZOOPHYTES.
Animals without body-cavity, blood or anus.

1A. Gastraeads.

Gastremaria.
Cyemaria.

1B. Sponges.

Protospongiae.
Metaspongiae.

1C. Cnidaria (Stinging Animals).

Hydrozoa.
Polyps.
Medusae.

1D. Platodes (Flat-Worms).

Platodaria.
Turbellaria.
Trematoda.
Cestoda.

2. COELOMARIA OR BILATERALS.
Animals with body-cavity and anus, and generally blood.

2A. Vermalia (Worm-Like).

Rotatoria.
Strongylaria.
Prosopygia.
Frontonia.

2B. Molluscs.

Cochlides.
Conchades.
Teuthodes.

2C. Articulates.

Annelida.
Crustacea.
Tracheata.

2D. Echinoderms.

Monorchonia.
Pentorchonia.

2E. Tunicates.

Copelata.
Ascidiae.
Thalidiae.

2F. Vertebrates.

2F.1. Acrania-Lancelet (Without Skull).

2F.2. Craniota (With Skull).

2F.2A. Cyclostomes. ("Round-Mouthed").

2F.2B. Fishes.

Selachii.
Ganoids.
Teleosts.
Dipneusts.

2F.2C. Amphibia.

2F.2D. Reptiles.

2F.2E. Birds.

2F.2F. Mammal.

Monotremes.

Marsupials.

Placentals:--
Rodents.
Edentates.
Ungulates.
Cetacea.
Sirenia.
Insectivora.
Cheiroptera.
Carnassia.
Primates.

(This classification is given for the purpose of explaining Haeckel's
use of terms in this volume. The general reader should bear in mind
that it differs very considerably from more recent schemes of
classification. He should compare the scheme framed by Professor E.
Ray Lankester.)

***


THE EVOLUTION OF MAN.


CHAPTER 1.1. THE FUNDAMENTAL LAW OF ORGANIC EVOLUTION.

The field of natural phenomena into which I would introduce my readers
in the following chapters has a quite peculiar place in the broad
realm of scientific inquiry. There is no object of investigation that
touches man more closely, and the knowledge of which should be more
acceptable to him, than his own frame. But among all the various
branches of the natural history of mankind, or anthropology, the story
of his development by natural means must excite the most lively
interest. It gives us the key of the great world-riddles at which the
human mind has been working for thousands of years. The problem of the
nature of man, or the question of man's place in nature, and the
cognate inquiries as to the past, the earliest history, the present
situation, and the future of humanity--all these most important
questions are directly and intimately connected with that branch of
study which we call the science of the evolution of man, or, in one
word, "Anthropogeny" (the genesis of man). Yet it is an astonishing
fact that the science of the evolution of man does not even yet form
part of the scheme of general education. In fact, educated people even
in our day are for the most part quite ignorant of the important
truths and remarkable phenomena which anthropogeny teaches us.

As an illustration of this curious state of things, it may be pointed
out that most of what are considered to be "educated" people do not
know that every human being is developed from an egg, or ovum, and
that this egg is one simple cell, like any other plant or animal egg.
They are equally ignorant that in the course of the development of
this tiny, round egg-cell there is first formed a body that is totally
different from the human frame, and has not the remotest resemblance
to it. Most of them have never seen such a human embryo in the earlier
period of its development, and do not know that it is quite
indistinguishable from other animal embryos. At first the embryo is no
more than a round cluster of cells, then it becomes a simple hollow
sphere, the wall of which is composed of a layer of cells. Later it
approaches very closely, at one period, to the anatomic structure of
the lancelet, afterwards to that of a fish, and again to the typical
build of the amphibia and mammals. As it continues to develop, a form
appears which is like those we find at the lowest stage of mammal-life
(such as the duck-bills), then a form that resembles the marsupials,
and only at a late stage a form that has a resemblance to the ape;
until at last the definite human form emerges and closes the series of
transformations. These suggestive facts are, as I said, still almost
unknown to the general public--so completely unknown that, if one
casually mentions them, they are called in question or denied outright
as fairy-tales. Everybody knows that the butterfly emerges from the
pupa, and the pupa from a quite different thing called a larva, and
the larva from the butterfly's egg. But few besides medical men are
aware that MAN, in the course of his individual formation, passes
through a series of transformations which are not less surprising and
wonderful than the familiar metamorphoses of the butterfly.

The mere description of these remarkable changes through which man
passes during his embryonic life should arouse considerable interest.
But the mind will experience a far keener satisfaction when we trace
these curious facts to their causes, and when we learn to behold in
them natural phenomena which are of the highest importance throughout
the whole field of human knowledge. They throw light first of all on
the "natural history of creation," then on psychology, or "the science
of the soul," and through this on the whole of philosophy. And as the
general results of every branch of inquiry are summed up in
philosophy, all the sciences come in turn to be touched and influenced
more or less by the study of the evolution of man.

But when I say that I propose to present here the most important
features of these phenomena and trace them to their causes, I take the
term, and I interpret my task, in a very much wider sense than is
usual. The lectures which have been delivered on this subject in the
universities during the last half-century are almost exclusively
adapted to medical men. Certainly, the medical man has the greatest
interest in studying the origin of the human body, with which he is
daily occupied. But I must not give here this special description of
the embryonic processes such as it has hitherto been given, as most of
my readers have not studied anatomy, and are not likely to be
entrusted with the care of the adult organism. I must content myself
with giving some parts of the subject only in general outline, and
must not enter upon all the marvellous, but very intricate and not
easily described, details that are found in the story of the
development of the human frame. To understand these fully a knowledge
of anatomy is needed. I will endeavour to be as plain as possible in
dealing with this branch of science. Indeed, a sufficient general idea
of the course of the embryonic development of man can be obtained
without going too closely into the anatomic details. I trust we may be
able to arouse the same interest in this delicate field of inquiry as
has been excited already in other branches of science; though we shall
meet more obstacles here than elsewhere.

The story of the evolution of man, as it has hitherto been expounded
to medical students, has usually been confined to embryology--more
correctly, ontogeny--or the science of the development of the
individual human organism. But this is really only the first part of
our task, the first half of the story of the evolution of man in that
wider sense in which we understand it here. We must add as the second
half--as another and not less important and interesting branch of the
science of the evolution of the human stem--phylogeny: this may be
described as the science of the evolution of the various animal forms
from which the human organism has been developed in the course of
countless ages. Everybody now knows of the great scientific activity
that was occasioned by the publication of Darwin's Origin of Species
in 1859. The chief direct consequence of this publication was to
provoke a fresh inquiry into the origin of the human race, and this
has proved beyond question our gradual evolution from the lower
species. We give the name of "Phylogeny" to the science which
describes this ascent of man from the lower ranks of the animal world.
The chief source that it draws upon for facts is "Ontogeny," or
embryology, the science of the development of the individual organism.
Moreover, it derives a good deal of support from paleontology, or the
science of fossil remains, and even more from comparative anatomy, or
morphology.

These two branches of our science--on the one side ontogeny or
embryology, and on the other phylogeny, or the science of
race-evolution--are most vitally connected. The one cannot be
understood without the other. It is only when the two branches fully
co-operate and supplement each other that "Biogeny" (or the science of
the genesis of life in the widest sense) attains to the rank of a
philosophic science. The connection between them is not external and
superficial, but profound, intrinsic, and causal. This is a discovery
made by recent research, and it is most clearly and correctly
expressed in the comprehensive law which I have called "the
fundamental law of organic evolution," or "the fundamental law of
biogeny." This general law, to which we shall find ourselves
constantly recurring, and on the recognition of which depends one's
whole insight into the story of evolution, may be briefly expressed in
the phrase: "The history of the foetus is a recapitulation of the
history of the race"; or, in other words, "Ontogeny is a
recapitulation of phylogeny." It may be more fully stated as follows:
The series of forms through which the individual organism passes
during its development from the ovum to the complete bodily structure
is a brief, condensed repetition of the long series of forms which the
animal ancestors of the said organism, or the ancestral forms of the
species, have passed through from the earliest period of organic life
down to the present day.

The causal character of the relation which connects embryology with
stem-history is due to the action of heredity and adaptation. When we
have rightly understood these, and recognised their great importance
in the formation of organisms, we can go a step further and say:
Phylogenesis is the mechanical cause of ontogenesis.* (* The term
"genesis," which occurs throughout, means, of course, "birth" or
origin. From this we get: Biogeny = the origin of life (bios);
Anthropogeny = the origin of man (anthropos); Ontogeny = the origin of
the individual (on); Phylogeny = the origin of the species (phulon);
and so on. In each case the term may refer to the process itself, or
to the science describing the process.--Translator.) In other words,
the development of the stem, or race, is, in accordance with the laws
of heredity and adaptation, the cause of all the changes which appear
in a condensed form in the evolution of the foetus.

The chain of manifold animal forms which represent the ancestry of
each higher organism, or even of man, according to the theory of
descent, always form a connected whole. We may designate this
uninterrupted series of forms with the letters of the alphabet: A, B,
C, D, E, etc., to Z. In apparent contradiction to what I have said,
the story of the development of the individual, or the ontogeny of
most organisms, only offers to the observer a part of these forms; so
that the defective series of embryonic forms would run: A, B, D, F, H,
K, M, etc.; or, in other cases, B, D, H, L, M, N, etc. Here, then, as
a rule, several of the evolutionary forms of the original series have
fallen out. Moreover, we often find--to continue with our illustration
from the alphabet--one or other of the original letters of the
ancestral series represented by corresponding letters from a different
alphabet. Thus, instead of the Roman B and D, we often have the Greek
Beta and Delta. In this case the text of the biogenetic law has been
corrupted, just as it had been abbreviated in the preceding case. But,
in spite of all this, the series of ancestral forms remains the same,
and we are in a position to discover its original complexion.

In reality, there is always a certain parallel between the two
evolutionary series. But it is obscured from the fact that in the
embryonic succession much is wanting that certainly existed in the
earlier ancestral succession. If the parallel of the two series were
complete, and if this great fundamental law affirming the causal
connection between ontogeny and phylogeny in the proper sense of the
word were directly demonstrable, we should only have to determine, by
means of the microscope and the dissecting knife, the series of forms
through which the fertilised ovum passes in its development; we should
then have before us a complete picture of the remarkable series of
forms which our animal ancestors have successively assumed from the
dawn of organic life down to the appearance of man. But such a
repetition of the ancestral history by the individual in its embryonic
life is very rarely complete. We do not often find our full alphabet.
In most cases the correspondence is very imperfect, being greatly
distorted and falsified by causes which we will consider later. We are
thus, for the most part, unable to determine in detail, from the study
of its embryology, all the different shapes which an organism's
ancestors have assumed; we usually--and especially in the case of the
human foetus--encounter many gaps. It is true that we can fill up most
of these gaps satisfactorily with the help of comparative anatomy, but
we cannot do so from direct embryological observation. Hence it is
important that we find a large number of lower animal forms to be
still represented in the course of man's embryonic development. In
these cases we may draw our conclusions with the utmost security as to
the nature of the ancestral form from the features of the form which
the embryo momentarily assumes.

To give a few examples, we can infer from the fact that the human ovum
is a simple cell that the first ancestor of our species was a tiny
unicellular being, something like the amoeba. In the same way, we
know, from the fact that the human foetus consists, at the first, of
two simple cell-layers (the gastrula), that the gastraea, a form with
two such layers, was certainly in the line of our ancestry. A later
human embryonic form (the chordula) points just as clearly to a
worm-like ancestor (the prochordonia), the nearest living relation of
which is found among the actual ascidiae. To this succeeds a most
important embryonic stage (acrania), in which our headless foetus
presents, in the main, the structure of the lancelet. But we can only
indirectly and approximately, with the aid of comparative anatomy and
ontogeny, conjecture what lower forms enter into the chain of our
ancestry between the gastraea and the chordula, and between this and
the lancelet. In the course of the historical development many
intermediate structures have gradually fallen out, which must
certainly have been represented in our ancestry. But, in spite of
these many, and sometimes very appreciable, gaps, there is no
contradiction between the two successions. In fact, it is the chief
purpose of this work to prove the real harmony and the original
parallelism of the two. I hope to show, on a substantial basis of
facts, that we can draw most important conclusions as to our
genealogical tree from the actual and easily-demonstrable series of
embryonic changes. We shall then be in a position to form a general
idea of the wealth of animal forms which have figured in the direct
line of our ancestry in the lengthy history of organic life.

In this evolutionary appreciation of the facts of embryology we must,
of course, take particular care to distinguish sharply and clearly
between the primitive, palingenetic (or ancestral) evolutionary
processes and those due to cenogenesis.* (* Palingenesis = new birth,
or re-incarnation (palin = again, genesis or genea = development);
hence its application to the phenomena which are recapitulated by
heredity from earlier ancestral forms. Cenogenesis = foreign or
negligible development (kenos and genea); hence, those phenomena which
come later in the story of life to disturb the inherited structure, by
a fresh adaptation to environment.--Translator.) By palingenetic
processes, or embryonic recapitulations, we understand all those
phenomena in the development of the individual which are transmitted
from one generation to another by heredity, and which, on that
account, allow us to draw direct inferences as to corresponding
structures in the development of the species. On the other hand, we
give the name of cenogenetic processes, or embryonic variations, to
all those phenomena in the foetal development that cannot be traced to
inheritance from earlier species, but are due to the adaptation of the
foetus, or the infant-form, to certain conditions of its embryonic
development. These cenogenetic phenomena are foreign or later
additions; they allow us to draw no direct inference whatever as to
corresponding processes in our ancestral history, but rather hinder us
from doing so.

This careful discrimination between the primary or palingenetic
processes and the secondary or cenogenetic is of great importance for
the purposes of the scientific history of a species, which has to draw
conclusions from the available facts of embryology, comparative
anatomy, and paleontology, as to the processes in the formation of the
species in the remote past. It is of the same importance to the
student of evolution as the careful distinction between genuine and
spurious texts in the works of an ancient writer, or the purging of
the real text from interpolations and alterations, is for the student
of philology. It is true that this distinction has not yet been fully
appreciated by many scientists. For my part, I regard it as the first
condition for forming any just idea of the evolutionary process, and I
believe that we must, in accordance with it, divide embryology into
two sections--palingenesis, or the science of recapitulated forms; and
cenogenesis, or the science of supervening structures.

To give at once a few examples from the science of man's origin in
illustration of this important distinction, I may instance the
following processes in the embryology of man, and of all the higher
vertebrates, as palingenetic: the formation of the two primary
germinal layers and of the primitive gut, the undivided structure of
the dorsal nerve-tube, the appearance of a simple axial rod between
the medullary tube and the gut, the temporary formation of the
gill-clefts and arches, the primitive kidneys, and so on.* (* All
these, and the following structures, will be fully described in later
chapters.--Translator.) All these, and many other important
structures, have clearly been transmitted by a steady heredity from
the early ancestors of the mammal, and are, therefore, direct
indications of the presence of similar structures in the history of
the stem. On the other hand, this is certainly not the case with the
following embryonic forms, which we must describe as cenogenetic
processes: the formation of the yelk-sac, the allantois, the placenta,
the amnion, the serolemma, and the chorion--or, generally speaking,
the various foetal membranes and the corresponding changes in the
blood vessels. Further instances are: the dual structure of the heart
cavity, the temporary division of the plates of the primitive
vertebrae and lateral plates, the secondary closing of the ventral and
intestinal walls, the formation of the navel, and so on. All these and
many other phenomena are certainly not traceable to similar structures
in any earlier and completely-developed ancestral form, but have
arisen simply by adaptation to the peculiar conditions of embryonic
life (within the foetal membranes). In view of these facts, we may now
give the following more precise expression to our chief law of
biogeny: The evolution of the foetus (or ontogenesis) is a condensed
and abbreviated recapitulation of the evolution of the stem (or
phylogenesis); and this recapitulation is the more complete in
proportion as the original development (or palingenesis) is preserved
by a constant heredity; on the other hand, it becomes less complete in
proportion as a varying adaptation to new conditions increases the
disturbing factors in the development (or cenogenesis).

The cenogenetic alterations or distortions of the original
palingenetic course of development take the form, as a rule, of a
gradual displacement of the phenomena, which is slowly effected by
adaptation to the changed conditions of embryonic existence during the
course of thousands of years. This displacement may take place as
regards either the position or the time of a phenomenon.

The great importance and strict regularity of the time-variations in
embryology have been carefully studied recently by Ernest Mehnert, in
his Biomechanik (Jena, 1898). He contends that our biogenetic law has
not been impaired by the attacks of its opponents, and goes on to say:
"Scarcely any piece of knowledge has contributed so much to the
advance of embryology as this; its formulation is one of the most
signal services to general biology. It was not until this law passed
into the flesh and blood of investigators, and they had accustomed
themselves to see a reminiscence of ancestral history in embryonic
structures, that we witnessed the great progress which embryological
research has made in the last two decades." The best proof of the
correctness of this opinion is that now the most fruitful work is done
in all branches of embryology with the aid of this biogenetic law, and
that it enables students to attain every year thousands of brilliant
results that they would never have reached without it.

It is only when one appreciates the cenogenetic processes in relation
to the palingenetic, and when one takes careful account of the changes
which the latter may suffer from the former, that the radical
importance of the biogenetic law is recognised, and it is felt to be
the most illuminating principle in the science of evolution. In this
task of discrimination it is the silver thread in relation to which we
can arrange all the phenomena of this realm of marvels--the "Ariadne
thread," which alone enables us to find our way through this labyrinth
of forms. Hence the brothers Sarasin, the zoologists, could say with
perfect justice, in their study of the evolution of the Ichthyophis,
that "the great biogenetic law is just as important for the zoologist
in tracing long-extinct processes as spectrum analyses is for the
astronomer."

Even at an earlier period, when a correct acquaintance with the
evolution of the human and animal frame was only just being
obtained--and that is scarcely eighty years ago!--the greatest
astonishment was felt at the remarkable similarity observed between
the embryonic forms, or stages of foetal development, in very
different animals; attention was called even then to their close
resemblance to certain fully-developed animal forms belonging to some
of the lower groups. The older scientists (Oken, Treviranus, and
others) knew perfectly well that these lower forms in a sense
illustrated and fixed, in the hierarchy of the animal world, a
temporary stage in the evolution of higher forms. The famous anatomist
Meckel spoke in 1821 of a "similarity between the development of the
embryo and the series of animals." Baer raised the question in 1828
how far, within the vertebrate type, the embryonic forms of the higher
animals assume the permanent shapes of members of lower groups. But it
was impossible fully to understand and appreciate this remarkable
resemblance at that time. We owe our capacity to do this to the theory
of descent; it is this that puts in their true light the action of
heredity on the one hand and adaptation on the other. It explains to
us the vital importance of their constant reciprocal action in the
production of organic forms. Darwin was the first to teach us the
great part that was played in this by the ceaseless struggle for
existence between living things, and to show how, under the influence
of this (by natural selection), new species were produced and
maintained solely by the interaction of heredity and adaptation. It
was thus Darwinism that first opened our eyes to a true comprehension
of the supremely important relations between the two parts of the
science of organic evolution--Ontogeny and Phylogeny.

Heredity and adaptation are, in fact, the two constructive
physiological functions of living things; unless we understand these
properly we can make no headway in the study of evolution. Hence,
until the time of Darwin no one had a clear idea of the real nature
and causes of embryonic development. It was impossible to explain the
curious series of forms through which the human embryo passed; it was
quite unintelligible why this strange succession of animal-like forms
appeared in the series at all. It had previously been generally
assumed that the man was found complete in all his parts in the ovum,
and that the development consisted only in an unfolding of the various
parts, a simple process of growth. This is by no means the case. On
the contrary, the whole process of the development of the individual
presents to the observer a connected succession of different
animal-forms; and these forms display a great variety of external and
internal structure. But WHY each individual human being should pass
through this series of forms in the course of his embryonic
development it was quite impossible to say until Lamarck and Darwin
established the theory of descent. Through this theory we have at last
detected the real causes, the efficient causes, of the individual
development; we have learned that these mechanical causes suffice of
themselves to effect the formation of the organism, and that there is
no need of the final causes which were formerly assumed. It is true
that in the academic philosophies of our time these final causes still
figure very prominently; in the new philosophy of nature we can
entirely replace them by efficient causes. We shall see, in the course
of our inquiry, how the most wonderful and hitherto insoluble enigmas
in the human and animal frame have proved amenable to a mechanical
explanation, by causes acting without prevision, through Darwin's
reform of the science of evolution. We have everywhere been able to
substitute unconscious causes, acting from necessity, for conscious,
purposive causes.* (* The monistic or mechanical philosophy of nature
holds that only unconscious, necessary, efficient causes are at work
in the whole field of nature, in organic life as well as in inorganic
changes. On the other hand, the dualist or vitalist philosophy of
nature affirms that unconscious forces are only at work in the
inorganic world, and that we find conscious, purposive, or final
causes in organic nature.)

If the new science of evolution had done no more than this, every
thoughtful man would have to admit that it had accomplished an immense
advance in knowledge. It means that in the whole of philosophy that
tendency which we call monistic, in opposition to the dualistic, which
has hitherto prevailed, must be accepted.* (* Monism is neither purely
materialistic nor purely spiritualistic, but a reconciliation of these
two principles, since it regards the whole of nature as one, and sees
only efficient causes at work in it. Dualism, on the contrary, holds
that nature and spirit, matter and force, the world and God, inorganic
and organic nature, are separate and independent existences. Cf. The
Riddle of the Universe chapter 12.) At this point the science of human
evolution has a direct and profound bearing on the foundations of
philosophy. Modern anthropology has, by its astounding discoveries
during the second half of the nineteenth century, compelled us to take
a completely monistic view of life. Our bodily structure and its life,
our embryonic development and our evolution as a species, teach us
that the same laws of nature rule in the life of man as in the rest of
the universe. For this reason, if for no others, it is desirable, nay,
indispensable, that every man who wishes to form a serious and
philosophic view of life, and, above all, the expert philosopher,
should acquaint himself with the chief facts of this branch of
science.

The facts of embryology have so great and obvious a significance in
this connection that even in recent years dualist and teleological
philosophers have tried to rid themselves of them by simply denying
them. This was done, for instance, as regards the fact that man is
developed from an egg, and that this egg or ovum is a simple cell, as
in the case of other animals. When I had explained this pregnant fact
and its significance in my History of Creation, it was described in
many of the theological journals as a dishonest invention of my own.
The fact that the embryos of man and the dog are, at a certain stage
of their development, almost indistinguishable was also denied. When
we examine the human embryo in the third or fourth week of its
development, we find it to be quite different in shape and structure
from the full-grown human being, but almost identical with that of the
ape, the dog, the rabbit, and other mammals, at the same stage of
ontogeny. We find a bean-shaped body of very simple construction, with
a tail below and a pair of fins at the sides, something like those of
a fish, but very different from the limbs of man and the mammals.
Nearly the whole front half of the body is taken up by a shapeless
head without face, at the sides of which we find gill-clefts and
arches as in the fish. At this stage of its development the human
embryo does not differ in any essential detail from that of the ape,
dog, horse, ox, etc., at a corresponding period. This important fact
can easily be verified at any moment by a comparison of the embryos of
man, the dog, rabbit, etc. Nevertheless, the theologians and dualist
philosophers pronounced it to be a materialistic invention; even
scientists, to whom the facts should be known, have sought to deny
them.

There could not be a clearer proof of the profound importance of these
embryological facts in favour of the monistic philosophy than is
afforded by these efforts of its opponents to get rid of them by
silence or denial. The truth is that these facts are most inconvenient
for them, and are quite irreconcilable with their views. We must be
all the more pressing on our side to put them in their proper light. I
fully agree with Huxley when he says, in his "Man's Place in Nature":
"Though these facts are ignored by several well-known popular leaders,
they are easy to prove, and are accepted by all scientific men; on the
other hand, their importance is so great that those who have once
mastered them will, in my opinion, find few other biological
discoveries to astonish them."

We shall make it our chief task to study the evolution of man's bodily
frame and its various organs in their external form and internal
structures. But I may observe at once that this is accompanied step by
step with a study of the evolution of their functions. These two
branches of inquiry are inseparably united in the whole of
anthropology, just as in zoology (of which the former is only a
section) or general biology. Everywhere the peculiar form of the
organism and its structures, internal and external, is directly
related to the special physiological functions which the organism or
organ has to execute. This intimate connection of structure and
function, or of the instrument and the work done by it, is seen in the
science of evolution and all its parts. Hence the story of the
evolution of structures, which is our immediate concern, is also the
history of the development of functions; and this holds good of the
human organism as of any other.

At the same time, I must admit that our knowledge of the evolution of
functions is very far from being as complete as our acquaintance with
the evolution of structures. One might say, in fact, that the whole
science of evolution has almost confined itself to the study of
structures; the evolution of FUNCTIONS hardly exists even in name.
That is the fault of the physiologists, who have as yet concerned
themselves very little about evolution. It is only in recent times
that physiologists like W. Engelmann, W. Preyer, M. Verworn, and a few
others, have attacked the evolution of functions.

It will be the task of some future physiologist to engage in the study
of the evolution of functions with the same zeal and success as has
been done for the evolution of structures in morphogeny (the science
of the genesis of forms). Let me illustrate the close connection of
the two by a couple of examples. The heart in the human embryo has at
first a very simple construction, such as we find in permanent form
among the ascidiae and other low organisms; with this is associated a
very simple system of circulation of the blood. Now, when we find that
with the full-grown heart there comes a totally different and much
more intricate circulation, our inquiry into the development of the
heart becomes at once, not only an anatomical, but also a
physiological, study. Thus it is clear that the ontogeny of the heart
can only be understood in the light of its phylogeny (or development
in the past), both as regards function and structure. The same holds
true of all the other organs and their functions. For instance, the
science of the evolution of the alimentary canal, the lungs, or the
sexual organs, gives us at the same time, through the exact
comparative investigation of structure-development, most important
information with regard to the evolution of the functions of these
organs.

This significant connection is very clearly seen in the evolution of
the nervous system. This system is in the economy of the human body
the medium of sensation, will, and even thought, the highest of the
psychic functions; in a word, of all the various functions which
constitute the proper object of psychology. Modern anatomy and
physiology have proved that these psychic functions are immediately
dependent on the fine structure and the composition of the central
nervous system, or the internal texture of the brain and spinal cord.
In these we find the elaborate cell-machinery, of which the psychic or
soul-life is the physiological function. It is so intricate that most
men still look upon the mind as something supernatural that cannot be
explained on mechanical principles.

But embryological research into the gradual appearance and the
formation of this important system of organs yields the most
astounding and significant results. The first sketch of a central
nervous system in the human embryo presents the same very simple type
as in the other vertebrates. A spinal tube is formed in the external
skin of the back, and from this first comes a simple spinal cord
without brain, such as we find to be the permanent psychic organ in
the lowest type of vertebrate, the amphioxus. Not until a later stage
is a brain formed at the anterior end of this cord, and then it is a
brain of the most rudimentary kind, such as we find permanently among
the lower fishes. This simple brain develops step by step,
successively assuming forms which correspond to those of the amphibia,
the reptiles, the duck-bills, and the lemurs. Only in the last stage
does it reach the highly organised form which distinguishes the apes
from the other vertebrates, and which attains its full development in
man.

Comparative physiology discovers a precisely similar growth. The
function of the brain, the psychic activity, rises step by step with
the advancing development of its structure.

Thus we are enabled, by this story of the evolution of the nervous
system, to understand at length THE NATURAL DEVELOPMENT OF THE HUMAN
MIND and its gradual unfolding. It is only with the aid of embryology
that we can grasp how these highest and most striking faculties of the
animal organism have been historically evolved. In other words, a
knowledge of the evolution of the spinal cord and brain in the human
embryo leads us directly to a comprehension of the historic
development (or phylogeny) of the human mind, that highest of all
faculties, which we regard as something so marvellous and supernatural
in the adult man. This is certainly one of the greatest and most
pregnant results of evolutionary science. Happily our embryological
knowledge of man's central nervous system is now so adequate, and
agrees so thoroughly with the complementary results of comparative
anatomy and physiology, that we are thus enabled to obtain a clear
insight into one of the highest problems of philosophy, the phylogeny
of the soul, or the ancestral history of the mind of man. Our chief
support in this comes from the embryological study of it, or the
ontogeny of the soul. This important section of psychology owes its
origin especially to W. Preyer, in his interesting works, such as The
Mind of the Child. The Biography of a Baby (1900), of Milicent
Washburn Shinn, also deserves mention. [See also Preyer's Mental
Development in the Child (translation), and Sully's Studies of
Childhood and Children's Ways.]

In this way we follow the only path along which we may hope to reach
the solution of this difficult problem.

Thirty-six years have now elapsed since, in my General Morphology, I
established phylogeny as an independent science and showed its
intimate causal connection with ontogeny; thirty years have passed
since I gave in my gastraea-theory the proof of the justice of this,
and completed it with the theory of germinal layers. When we look back
on this period we may ask, What has been accomplished during it by the
fundamental law of biogeny? If we are impartial, we must reply that it
has proved its fertility in hundreds of sound results, and that by its
aid we have acquired a vast fund of knowledge which we should never
have obtained without it.

There has been no dearth of attacks--often violent attacks--on my
conception of an intimate causal connection between ontogenesis and
phylogenesis; but no other satisfactory explanation of these important
phenomena has yet been offered to us. I say this especially with
regard to Wilhelm His's theory of a "mechanical evolution," which
questions the truth of phylogeny generally, and would explain the
complicated embryonic processes without going beyond by simple
physical changes--such as the bending and folding of leaves by
electricity, the origin of cavities through unequal strain of the
tissues, the formation of processes by uneven growth, and so on. But
the fact is that these embryological phenomena themselves demand
explanation in turn, and this can only be found, as a rule, in the
corresponding changes in the long ancestral series, or in the
physiological functions of heredity and adaptation.


CHAPTER 1.2. THE OLDER EMBRYOLOGY.

It is in many ways useful, on entering upon the study of any science,
to cast a glance at its historical development. The saying that
"everything is best understood in its growth" has a distinct
application to science. While we follow its gradual development we get
a clearer insight into its aims and objects. Moreover, we shall see
that the present condition of the science of human evolution, with all
its characteristics, can only be rightly understood when we examine
its historical growth. This task will, however, not detain us long.
The study of man's evolution is one of the latest branches of natural
science, whether you consider the embryological or the phylogenetic
section of it.

Apart from the few germs of our science which we find in classical
antiquity, and which we shall notice presently, we may say that it
takes its definite rise, as a science, in the year 1759, when one of
the greatest German scientists, Caspar Friedrich Wolff, published his
Theoria generationis. That was the foundation-stone of the science of
animal embryology. It was not until fifty years later, in 1809, that
Jean Lamarck published his Philosophie Zoologique--the first effort to
provide a base for the theory of evolution; and it was another
half-century before Darwin's work appeared (in 1859), which we may
regard as the first scientific attainment of this aim. But before we
go further into this solid establishment of evolution, we must cast a
brief glance at that famous philosopher and scientist of antiquity,
who stood alone in this, as in many other branches of science, for
more than 2000 years: the "father of Natural History," Aristotle.

The extant scientific works of Aristotle deal with many different
sides of biological research; the most comprehensive of them is his
famous History of Animals. But not less interesting is the smaller
work, On the Generation of Animals (Peri zoon geneseos). This work
treats especially of embryonic development, and it is of great
interest as being the earliest of its kind and the only one that has
come down to us in any completeness from classical antiquity.

Aristotle studied embryological questions in various classes of
animals, and among the lower groups he learned many most remarkable
facts which we only rediscovered between 1830 and 1860. It is certain,
for instance, that he was acquainted with the very peculiar mode of
propagation of the cuttlefishes, or cephalopods, in which a yelk-sac
hangs out of the mouth of the foetus. He knew, also, that embryos come
from the eggs of the bee even when they have not been fertilised. This
"parthenogenesis" (or virgin-birth) of the bees has only been
established in our time by the distinguished zoologist of Munich,
Siebold. He discovered that male bees come from the unfertilised, and
female bees only from the fertilised, eggs. Aristotle further states
that some kinds of fishes (of the genus serranus) are hermaphrodites,
each individual having both male and female organs and being able to
fertilise itself; this, also, has been recently confirmed. He knew
that the embryo of many fishes of the shark family is attached to the
mother's body by a sort of placenta, or nutritive organ very rich in
blood; apart from these, such an arrangement is only found among the
higher mammals and man. This placenta of the shark was looked upon as
legendary for a long time, until Johannes Muller proved it to be a
fact in 1839. Thus a number of remarkable discoveries were found in
Aristotle's embryological work, proving a very good acquaintance of
the great scientist--possibly helped by his predecessors--with the
facts of ontogeny, and a great advance upon succeeding generations in
this respect.

In the case of most of these discoveries he did not merely describe
the fact, but added a number of observations on its significance. Some
of these theoretical remarks are of particular interest, because they
show a correct appreciation of the nature of the embryonic processes.
He conceives the development of the individual as a new formation, in
the course of which the various parts of the body take shape
successively. When the human or animal frame is developed in the
mother's body, or separately in an egg, the heart--which he regards as
the starting-point and centre of the organism--must appear first. Once
the heart is formed the other organs arise, the internal ones before
the external, the upper (those above the diaphragm) before the lower
(or those beneath the diaphragm). The brain is formed at an early
stage, and the eyes grow out of it. These observations are quite
correct. And, if we try to form some idea from these data of
Aristotle's general conception of the embryonic process, we find a dim
prevision of the theory which Wolff showed 2000 years afterwards to be
the correct view. It is significant, for instance, that Aristotle
denied the eternity of the individual in any respect. He said that the
species or genus, the group of similar individuals, might be eternal,
but the individual itself is temporary. It comes into being in the act
of procreation, and passes away at death.

During the 2000 years after Aristotle no progress whatever was made in
general zoology, or in embryology in particular. People were content
to read, copy, translate, and comment on Aristotle. Scarcely a single
independent effort at research was made in the whole of the period.
During the Middle Ages the spread of strong religious beliefs put
formidable obstacles in the way of independent scientific
investigation. There was no question of resuming the advance of
biology. Even when human anatomy began to stir itself once more in the
sixteenth century, and independent research was resumed into the
structure of the developed body, anatomists did not dare to extend
their inquiries to the unformed body, the embryo, and its development.
There were many reasons for the prevailing horror of such studies. It
is natural enough, when we remember that a Bull of Boniface VIII
excommunicated every man who ventured to dissect a human corpse. If
the dissection of a developed body were a crime to be thus punished,
how much more dreadful must it have seemed to deal with the embryonic
body still enclosed in the womb, which the Creator himself had
decently veiled from the curiosity of the scientist! The Christian
Church, then putting many thousands to death for unbelief, had a
shrewd presentiment of the menace that science contained against its
authority. It was powerful enough to see that its rival did not grow
too quickly.

It was not until the Reformation broke the power of the Church, and a
refreshing breath of the spirit dissolved the icy chains that bound
science, that anatomy and embryology, and all the other branches of
research, could begin to advance once more. However, embryology lagged
far behind anatomy. The first works on embryology appear at the
beginning of the sixteenth century. The Italian anatomist, Fabricius
ab Aquapendente, a professor at Padua, opened the advance. In his two
books (De formato foetu, 1600, and De formatione foetus, 1604) he
published the older illustrations and descriptions of the embryos of
man and other mammals, and of the hen. Similar imperfect illustrations
were given by Spigelius (De formato foetu, 1631), and by Needham
(1667) and his more famous compatriot, Harvey (1652), who discovered
the circulation of the blood in the animal body and formulated the
important principle, Omne vivum ex vivo (all life comes from
pre-existing life). The Dutch scientist, Swammerdam, published in his
Bible of Nature the earliest observations on the embryology of the
frog and the division of its egg-yelk. But the most important
embryological studies in the sixteenth century were those of the
famous Italian, Marcello Malpighi, of Bologna, who led the way both in
zoology and botany. His treatises, De formatione pulli and De ovo
incubato (1687), contain the first consistent description of the
development of the chick in the fertilised egg.

Here I ought to say a word about the important part played by the
chick in the growth of our science. The development of the chick, like
that of the young of all other birds, agrees in all its main features
with that of the other chief vertebrates, and even of man. The three
highest classes of vertebrates--mammals, birds, and reptiles (lizards,
serpents, tortoises, etc.)--have from the beginning of their embryonic
development so striking a resemblance in all the chief points of
structure, and especially in their first forms, that for a long time
it is impossible to distinguish between them. We have known now for
some time that we need only examine the embryo of a bird, which is the
easiest to get at, in order to learn the typical mode of development
of a mammal (and therefore of man). As soon as scientists began to
study the human embryo, or the mammal-embryo generally, in its earlier
stages about the middle and end of the seventeenth century, this
important fact was very quickly discovered. It is both theoretically
and practically of great value. As regards the THEORY of evolution, we
can draw the most weighty inferences from this similarity between the
embryos of widely different classes of animals. But for the practical
purposes of embryological research the discovery is invaluable,
because we can fill up the gaps in our imperfect knowledge of the
embryology of the mammals from the more thoroughly studied embryology
of the bird. Hens' eggs are easily to be had in any quantity, and the
development of the chick may be followed step by step in artificial
incubation. The development of the mammal is much more difficult to
follow, because here the embryo is not detached and enclosed in a
large egg, but the tiny ovum remains in the womb until the growth is
completed. Hence, it is very difficult to keep up sustained
observation of the various stages in any great extent, quite apart
from such extrinsic considerations as the cost, the technical
difficulties, and many other obstacles which we encounter when we
would make an extensive study of the fertilised mammal. The chicken
has, therefore, always been the chief object of study in this
connection. The excellent incubators we now have enable us to observe
it in any quantity and at any stage of development, and so follow the
whole course of its formation step by step.

By the end of the seventeenth century Malpighi had advanced as far as
it was possible to do with the imperfect microscope of his time in the
embryological study of the chick. Further progress was arrested until
the instrument and the technical methods should be improved. The
vertebrate embryos are so small and delicate in their earlier stages
that you cannot go very far into the study of them without a good
microscope and other technical aid. But this substantial improvement
of the microscope and the other apparatus did not take place until the
beginning of the nineteenth century.

Embryology made scarcely any advance in the first half of the
eighteenth century, when the systematic natural history of plants and
animals received so great an impulse through the publication of
Linne's famous Systema Naturae. Not until 1759 did the genius arise
who was to give it an entirely new character, Caspar Friedrich Wolff.
Until then embryology had been occupied almost exclusively in
unfortunate and misleading efforts to build up theories on the
imperfect empirical material then available.

The theory which then prevailed, and remained in favour throughout
nearly the whole of the eighteenth century, was commonly called at
that time "the evolution theory"; it is better to describe it as "the
preformation theory."* (* This theory is usually known as the
"evolution theory" in Germany, in contradistinction to the "epigenesis
theory." But as it is the latter that is called the "evolution theory"
in England, France, and Italy, and "evolution" and "epigenesis" are
taken to be synonymous, it seems better to call the first the
"pre-formation theory.") Its chief point is this: There is no new
formation of structures in the embryonic development of any organism,
animal or plant, or even of man; there is only a growth, or unfolding,
of parts which have been constructed or pre-formed from all eternity,
though on a very small scale and closely packed together. Hence, every
living germ contains all the organs and parts of the body, in the form
and arrangement they will present later, already within it, and thus
the whole embryological process is merely an evolution in the literal
sense of the word, or an unfolding, of parts that were pre-formed and
folded up in it. So, for instance, we find in the hen's egg not merely
a simple cell, that divides and subdivides and forms germinal layers,
and at last, after all kinds of variation and cleavage and
reconstruction, brings forth the body of the chick; but there is in
every egg from the first a complete chicken, with all its parts made
and neatly packed. These parts are so small or so transparent that the
microscope cannot detect them. In the hatching, these parts merely
grow larger, and spread out in the normal way.

When this theory is consistently developed it becomes a "scatulation
theory."* (* "Packing theory" would be the literal translation.
Scatula is the Latin for a case or box.--Translator.) According to its
teaching, there was made in the beginning one couple or one individual
of each species of animal or plant; but this one individual contained
the germs of all the other individuals of the same species who should
ever come to life. As the age of the earth was generally believed at
that time to be fixed by the Bible at 5000 or 6000 years, it seemed
possible to calculate how many individuals of each species had lived
in the period, and so had been packed inside the first being that was
created. The theory was consistently extended to man, and it was
affirmed that our common parent Eve had had stored in her ovary the
germs of all the children of men.

The theory at first took the form of a belief that it was the FEMALES
who were thus encased in the first being. One couple of each species
was created, but the female contained in her ovary all the future
individuals of the species, of either sex. However, this had to be
altered when the Dutch microscopist, Leeuwenhoek, discovered the male
spermatozoa in 1690, and showed that an immense number of these
extremely fine and mobile thread-like beings exist in the male sperm
(this will be explained in Chapter 2.7). This astonishing discovery
was further advanced when it was proved that these living bodies,
swimming about in the seminal fluid, were real animalcules, and, in
fact, were the pre-formed germs of the future generation. When the
male and female procreative elements came together at conception,
these thread-like spermatozoa ("seed-animals") were supposed to
penetrate into the fertile body of the ovum and begin to develop
there, as the plant seed does in the fruitful earth. Hence, every
spermatozoon was regarded as a homunculus, a tiny complete man; all
the parts were believed to be pre-formed in it, and merely grew larger
when it reached its proper medium in the female ovum. This theory,
also, was consistently developed in the sense that in each of these
thread-like bodies the whole of its posterity was supposed to be
present in the minutest form. Adam's sexual glands were thought to
have contained the germs of the whole of humanity.

This "theory of male scatulation" found itself at once in keen
opposition to the prevailing "female" theory. The two rival theories
at once opened a very lively campaign, and the physiologists of the
eighteenth century were divided into two great camps--the
Animalculists and the Ovulists--which fought vigorously. The
animalculists held that the spermatozoa were the true germs, and
appealed to the lively movements and the structure of these bodies.
The opposing party of the Ovulists, who clung to the older "evolution
theory," affirmed that the ovum is the real germ, and that the
spermatozoa merely stimulate it at conception to begin its growth; all
the future generations are stored in the ovum. This view was held by
the great majority of the biologists of the eighteenth century, in
spite of the fact that Wolff proved it in 1759 to be without
foundation. It owed its prestige chiefly to the circumstance that the
most weighty authorities in the biology and philosophy of the day
decided in favour of it, especially Haller, Bonnet, and Leibnitz.

Albrecht Haller, professor at Gottingen, who is often called the
father of physiology, was a man of wide and varied learning, but he
does not occupy a very high position in regard to insight into natural
phenomena. He made a vigorous defence of the "evolutionary theory" in
his famous work, Elementa physiologiae, affirming: "There is no such
thing as formation (nulla est epigenesis). No part of the animal frame
is made before another; all were made together." He thus denied that
there was any evolution in the proper sense of the word, and even went
so far as to say that the beard existed in the new-born child and the
antlers in the hornless fawn; all the parts were there in advance, and
were merely hidden from the eye of man for the time being. Haller even
calculated the number of human beings that God must have created on
the sixth day and stored away in Eve's ovary. He put the number at
200,000 millions, assuming the age of the world to be 6000 years, the
average age of a human being to be thirty years, and the population of
the world at that time to be 1000 millions. And the famous Haller
maintained all this nonsense, in spite of its ridiculous consequences,
even after Wolff had discovered the real course of embryonic
development and established it by direct observation!

Among the philosophers of the time the distinguished Leibnitz was the
chief defender of the "preformation theory," and by his authority and
literary prestige won many adherents to it. Supported by his system of
monads, according to which body and soul are united in inseparable
association and by their union form the individual, or the "monad,"
Leibnitz consistently extended the "scatulation theory" to the soul,
and held that this was no more evolved than the body. He says, for
instance, in his Theodicee: "I mean that these souls, which one day
are to be the souls of men, are present in the seed, like those of
other species; in such wise that they existed in our ancestors as far
back as Adam, or from the beginning of the world, in the forms of
organised bodies."

The theory seemed to receive considerable support from the
observations of one of its most zealous supporters, Bonnet. In 1745 he
discovered, in the plant-louse, a case of parthenogenesis, or
virgin-birth, an interesting form of reproduction that has lately been
found by Siebold and others among various classes of the articulata,
especially crustacea and insects. Among these and other animals of
certain lower species the female may reproduce for several generations
without having been fertilised by the male. These ova that do not need
fertilisation are called "false ova," pseudova or spores. Bonnet saw
that a female plant-louse, which he had kept in cloistral isolation,
and rigidly removed from contact with males, had on the eleventh day
(after forming a new skin for the fourth time) a living daughter, and
during the next twenty days ninety-four other daughters; and that all
of them went on to reproduce in the same way without any contact with
males. It seemed as if this furnished an irrefutable proof of the
truth of the scatulation theory, as it was held by the Ovulists; it is
not surprising to find that the theory then secured general
acceptance.

This was the condition of things when suddenly, in 1759, Caspar
Friedrich Wolff appeared, and dealt a fatal blow at the whole
preformation theory with his new theory of epigenesis. Wolff, the son
of a Berlin tailor, was born in 1733, and went through his scientific
and medical studies, first at Berlin under the famous anatomist
Meckel, and afterwards at Halle. Here he secured his doctorate in his
twenty-sixth year, and in his academic dissertation (November 28th,
1759), the Theoria generationis, expounded the new theory of a real
development on a basis of epigenesis. This treatise is, in spite of
its smallness and its obscure phraseology, one of the most valuable in
the whole range of biological literature. It is equally distinguished
for the mass of new and careful observations it contains, and the
far-reaching and pregnant ideas which the author everywhere extracts
from his observations and builds into a luminous and accurate theory
of generation. Nevertheless, it met with no success at the time.
Although scientific studies were then assiduously cultivated owing to
the impulse given by Linne--although botanists and zoologists were no
longer counted by dozens, but by hundreds, hardly any notice was taken
of Wolff's theory. Even when he established the truth of epigenesis by
the most rigorous observations, and demolished the airy structure of
the preformation theory, the "exact" scientist Haller proved one of
the most strenuous supporters of the old theory, and rejected Wolff's
correct view with a dictatorial "There is no such thing as evolution."
He even went on to say that religion was menaced by the new theory! It
is not surprising that the whole of the physiologists of the second
half of the eighteenth century submitted to the ruling of this
physiological pontiff, and attacked the theory of epigenesis as a
dangerous innovation. It was not until more than fifty years
afterwards that Wolff's work was appreciated. Only when Meckel
translated into German in 1812 another valuable work of Wolff's on The
Formation of the Alimentary Canal (written in 1768), and called
attention to its great importance, did people begin to think of him
once more; yet this obscure writer had evinced a profounder insight
into the nature of the living organism than any other scientist of the
eighteenth century.

Wolff's idea led to an appreciable advance over the whole field of
biology. There is such a vast number of new and important observations
and pregnant thoughts in his writings that we have only gradually
learned to appreciate them rightly in the course of the nineteenth
century. He opened up the true path for research in many directions.
In the first place, his theory of epigenesis gave us our first real
insight into the nature of embryonic development. He showed
convincingly that the development of every organism consists of a
series of NEW FORMATIONS, and that there is no trace whatever of the
complete form either in the ovum or the spermatozoon. On the contrary,
these are quite simple bodies, with a very different purport. The
embryo which is developed from them is also quite different, in its
internal arrangement and outer configuration, from the complete
organism. There is no trace whatever of preformation or in-folding of
organs. To-day we can scarcely call epigenesis a THEORY, because we
are convinced it is a fact, and can demonstrate it at any moment with
the aid of the microscope.

Wolff furnished the conclusive empirical proof of his theory in his
classic dissertation on The Formation of the Alimentary Canal (1768).
In its complete state the alimentary canal of the hen is a long and
complex tube, with which the lungs, liver, salivary glands, and many
other small glands, are connected. Wolff showed that in the early
stages of the embryonic chick there is no trace whatever of this
complicated tube with all its dependencies, but instead of it only a
flat, leaf-shaped body; that, in fact, the whole embryo has at first
the appearance of a flat, oval-shaped leaf. When we remember how
difficult the exact observation of so fine and delicate a structure as
the early leaf-shaped body of the chick must have been with the poor
microscopes then in use, we must admire the rare faculty for
observation which enabled Wolff to make the most important discoveries
in this most difficult part of embryology. By this laborious research
he reached the correct opinion that the embryonic body of all the
higher animals, such as the birds, is for some time merely a flat,
thin, leaf-shaped disk--consisting at first of one layer, but
afterwards of several. The lowest of these layers is the alimentary
canal, and Wolff followed its development from its commencement to its
completion. He showed how this leaf-shaped structure first turns into
a groove, then the margins of this groove fold together and form a
closed canal, and at length the two external openings of the tube (the
mouth and anus) appear.

Moreover, the important fact that the other systems of organs are
developed in the same way, from tubes formed out of simple layers, did
not escape Wolff. The nerveless system, muscular system, and vascular
(blood-vessel) system, with all the organs appertaining thereto, are,
like the alimentary system, developed out of simple leaf-shaped
structures. Hence, Wolff came to the view by 1768 which Pander
developed in the Theory of Germinal Layers fifty years afterwards. His
principles are not literally correct; but he comes as near to the
truth in them as was possible at that time, and could be expected of
him.

Our admiration of this gifted genius increases when we find that he
was also the precursor of Goethe in regard to the metamorphosis of
plants and of the famous cellular theory. Wolff had, as Huxley showed,
a clear presentiment of this cardinal theory, since he recognised
small microscopic globules as the elementary parts out of which the
germinal layers arose.

Finally, I must invite special attention to the MECHANICAL character
of the profound philosophic reflections which Wolff always added to
his remarkable observations. He was a great monistic philosopher, in
the best meaning of the word. It is unfortunate that his philosophic
discoveries were ignored as completely as his observations for more
than half a century. We must be all the more careful to emphasise the
fact of their clear monistic tendency.


CHAPTER 1.3. MODERN EMBRYOLOGY.

We may distinguish three chief periods in the growth of our science of
human embryology. The first has been considered in the preceding
chapter; it embraces the whole of the preparatory period of research,
and extends from Aristotle to Caspar Friedrich Wolff, or to the year
1759, in which the epoch-making Theoria generationis was published.
The second period, with which we have now to deal, lasts about a
century--that is to say, until the appearance of Darwin's Origin of
Species, which brought about a change in the very foundations of
biology, and, in particular, of embryology. The third period begins
with Darwin. When we say that the second period lasted a full century,
we must remember that Wolff's work had remained almost unnoticed
during half the time--namely, until the year 1812. During the whole of
these fifty-three years not a single book that appeared followed up
the path that Wolff had opened, or extended his theory of embryonic
development. We merely find his views--perfectly correct views, based
on extensive observations of fact--mentioned here and there as
erroneous; their opponents, who adhered to the dominant theory of
preformation, did not even deign to reply to them. This unjust
treatment was chiefly due to the extraordinary authority of Albrecht
von Haller; it is one of the most astonishing instances of a great
authority, as such, preventing for a long time the recognition of
established facts.

The general ignorance of Wolff's work was so great that at the
beginning of the nineteenth century two scientists of Jena, Oken
(1806) and Kieser (1810), began independent research into the
development of the alimentary canal of the chick, and hit upon the
right clue to the embryonic puzzle, without knowing a word about
Wolff's important treatise on the same subject. They were treading in
his very footsteps without suspecting it. This can be easily proved
from the fact that they did not travel as far as Wolff. It was not
until Meckel translated into German Wolff's book on the alimentary
system, and pointed out its great importance, that the eyes of
anatomists and physiologists were suddenly opened. At once a number of
biologists instituted fresh embryological inquiries, and began to
confirm Wolff's theory of epigenesis.

This resuscitation of embryology and development of the
epigenesis-theory was chiefly connected with the university of
Wurtzburg. One of the professors there at that time was Dollinger, an
eminent biologist, and father of the famous Catholic historian who
later distinguished himself by his opposition to the new dogma of
papal infallibility. Dollinger was both a profound thinker and an
accurate observer. He took the keenest interest in embryology, and
worked at it a good deal. However, he is not himself responsible for
any important result in this field. In 1816 a young medical doctor,
whom we may at once designate as Wolff's chief successor, Karl Ernst
von Baer, came to Wurtzburg. Baer's conversations with Dollinger on
embryology led to a fresh series of most extensive investigations.
Dollinger had expressed a wish that some young scientist should begin
again under his guidance an independent inquiry into the development
of the chick during the hatching of the egg. As neither he nor Baer
had money enough to pay for an incubator and the proper control of the
experiments, and for a competent artist to illustrate the various
stages observed, the lead of the enterprise was given to Christian
Pander, a wealthy friend of Baer's who had been induced by Baer to
come to Wurtzburg. An able engraver, Dalton, was engaged to do the
copper-plates. In a short time the embryology of the chick, in which
Baer was taking the greatest indirect interest, was so far advanced
that Pander was able to sketch the main features of it on the ground
of Wolff's theory in the dissertation he published in 1817. He clearly
enunciated the theory of germinal layers which Wolff had anticipated,
and established the truth of Wolff's idea of a development of the
complicated systems of organs out of simple leaf-shaped primitive
structures. According to Pander, the leaf-shaped object in the hen's
egg divides, before the incubation has proceeded twelve hours, into
two different layers, an external serous layer and an internal mucous
layer; between the two there develops later a third layer, the
vascular (blood-vessel) layer.* (* The technical terms which are bound
to creep into this chapter will be fully understood later
on.--Translator.)

Karl Ernst von Baer, who had set afoot Pander's investigation, and had
shown the liveliest interest in it after Pander's departure from
Wurtzburg, began his own much more comprehensive research in 1819. He
published the mature result nine years afterwards in his famous work,
Animal Embryology: Observation and Reflection (not translated). This
classic work still remains a model of careful observation united to
profound philosophic speculation. The first part appeared in 1828, the
second in 1837. The book proved to be the foundation on which the
whole science of embryology has built down to our own day. It so far
surpassed its predecessors, and Pander in particular, that it has
become, after Wolff's work, the chief base of modern embryology.

Baer was one of the greatest scientists of the nineteenth century, and
exercised considerable influence on other branches of biology as well.
He built up the theory of germinal layers, as a whole and in detail,
so clearly and solidly that it has been the starting-point of
embryological research ever since. He taught that in all the
vertebrates first two and then four of these germinal layers are
formed; and that the earliest rudimentary organs of the body arise by
the conversion of these layers into tubes. He described the first
appearance of the vertebrate embryo, as it may be seen in the globular
yelk of the fertilised egg, as an oval disk which first divides into
two layers. From the upper or animal layer are developed all the
organs which accomplish the phenomena of animal life--the functions of
sensation and motion, and the covering of the body. From the lower or
vegetative layer come the organs which effect the vegetative life of
the organism--nutrition, digestion, blood-formation, respiration,
secretion, reproduction, etc.

Each of these original layers divides, according to Baer, into two
thinner and superimposed layers or plates. He calls the two plates of
the animal layer, the skin-stratum and muscle-stratum. From the upper
of these plates, the skin-stratum, the external skin, or outer
covering of the body, the central nervous system, and the
sense-organs, are formed. From the lower, or muscle-stratum, the
muscles, or fleshy parts and the bony skeleton--in a word, the motor
organs--are evolved. In the same way, Baer said, the lower or
vegetative layer splits into two plates, which he calls the
vascular-stratum and the mucous-stratum. From the outer of the two
(the vascular) the heart, blood-vessels, spleen, and the other
vascular glands, the kidneys, and sexual glands, are formed. From the
fourth or mucous layer, in fine, we get the internal and digestive
lining of the alimentary canal and all its dependencies, the liver,
lungs, salivary glands, etc. Baer had, in the main, correctly judged
the significance of these four secondary embryonic layers, and he
followed the conversion of them into the tube-shaped primitive organs
with great perspicacity. He first solved the difficult problem of the
transformation of this four-fold, flat, leaf-shaped, embryonic disk
into the complete vertebrate body, through the conversion of the
layers or plates into tubes. The flat leaves bend themselves in
obedience to certain laws of growth; the borders of the curling plates
approach nearer and nearer; until at last they come into actual
contact. Thus out of the flat gut-plate is formed a hollow gut-tube,
out of the flat spinal plate a hollow nerve-tube, from the skin-plate
a skin-tube, and so on.

Among the many great services which Baer rendered to embryology,
especially vertebrate embryology, we must not forget his discovery of
the human ovum. Earlier scientists had, as a rule, of course, assumed
that man developed out of an egg, like the other animals. In fact, the
preformation theory held that the germs of the whole of humanity were
stored already in Eve's ova. But the real ovum escaped detection until
the year 1827. This ovum is extremely small, being a tiny round
vesicle about the 1/120 of an inch in diameter; it can be seen under
very favourable circumstances with the naked eye as a tiny particle,
but is otherwise quite invisible. This particle is formed in the ovary
inside a much larger globule, which takes the name of the Graafian
follicle, from its discoverer, Graaf, and had previously been regarded
as the true ovum. However, in 1827 Baer proved that it was not the
real ovum, which is much smaller, and is contained within the
follicle. (Compare the end of Chapter 2.29.)

Baer was also the first to observe what is known as the segmentation
sphere of the vertebrate; that is to say, the round vesicle which
first develops out of the impregnated ovum, and the thin wall of which
is made up of a single layer of regular, polygonal (many-cornered)
cells (see the illustration in Chapter 1.12). Another discovery of his
that was of great importance in constructing the vertebrate stem and
the characteristic organisation of this extensive group (to which man
belongs) was the detection of the axial rod, or the chorda dorsalis.
There is a long, round, cylindrical rod of cartilage which runs down
the longer axis of the vertebrate embryo; it appears at an early
stage, and is the first sketch of the spinal column, the solid
skeletal axis of the vertebrate. In the lowest of the vertebrates, the
amphioxus, the internal skeleton consists only of this cord throughout
life. But even in the case of man and all the higher vertebrates it is
round this cord that the spinal column and the brain are afterwards
formed.

However, important as these and many other discoveries of Baer's were
in vertebrate embryology, his researches were even more influential,
from the circumstance that he was the first to employ the comparative
method in studying the development of the animal frame. Baer occupied
himself chiefly with the embryology of vertebrates (especially the
birds and fishes). But he by no means confined his attention to these,
gradually taking the various groups of the invertebrates into his
sphere of study. As the general result of his comparative
embryological research, Baer distinguished four different modes of
development and four corresponding groups in the animal world. These
chief groups or types are: 1, the vertebrata; 2, the articulata; 3,
the mollusca; and 4, all the lower groups which were then wrongly
comprehended under the general name of the radiata. Georges Cuvier had
been the first to formulate this distinction, in 1812. He showed that
these groups present specific differences in their whole internal
structure, and the connection and disposal of their systems of organs;
and that, on the other hand, all the animals of the same type--say,
the vertebrates--essentially agreed in their inner structure, in spite
of the greatest superficial differences. But Baer proved that these
four groups are also quite differently developed from the ovum; and
that the series of embryonic forms is the same throughout for animals
of the same type, but different in the case of other animals. Up to
that time the chief aim in the classification of the animal kingdom
was to arrange all the animals from lowest to highest, from the
infusorium to man, in one long and continuous series. The erroneous
idea prevailed nearly everywhere that there was one uninterrupted
chain of evolution from the lowest animal to the highest. Cuvier and
Baer proved that this view was false, and that we must distinguish
four totally different types of animals, on the ground of anatomic
structure and embryonic development.

Baer's epoch-making works aroused an extraordinary and widespread
interest in embryological research. Immediately afterwards we find a
great number of observers at work in the newly opened field, enlarging
it in a very short time with great energy by their various discoveries
in detail. Next to Baer's comes the admirable work of Heinrich Rathke,
of Konigsberg (died 1860); he made an extensive study of the
embryology, not only of the invertebrates (crustaceans, insects,
molluscs), but also, and particularly, of the vertebrates (fishes,
tortoises, serpents, crocodiles, etc.). We owe the first comprehensive
studies of mammal embryology to the careful research of Wilhelm
Bischoff, of Munich; his embryology of the rabbit (1840), the dog
(1842), the guinea-pig (1852), and the doe (1854), still form
classical studies. About the same time a great impetus was given to
the embryology of the invertebrates. The way was opened through this
obscure province by the studies of the famous Berlin zoologist,
Johannes Muller, on the echinoderms. He was followed by Albert
Kolliker, of Wurtzburg, writing on the cuttlefish (or the
cephalopods), Siebold and Huxley on worms and zoophytes, Fritz Muller
(Desterro) on the crustacea, Weismann on insects, and so on. The
number of workers in this field has greatly increased of late, and a
quantity of new and astonishing discoveries have been made. One
notices, in several of these recent works on embryology, that their
authors are too little acquainted with comparative anatomy and
classification. Palaeontology is, unfortunately, altogether neglected
by many of these new workers, although this interesting science
furnishes most important facts for phylogeny, and thus often proves of
very great service in ontogeny.

A very important advance was made in our science in 1839, when the
cellular theory was established, and a new field of inquiry bearing on
embryology was suddenly opened. When the famous botanist, M.
Schleiden, of Jena, showed in 1838, with the aid of the microscope,
that every plant was made up of innumerable elementary parts, which we
call cells, a pupil of Johannes Muller at Berlin, Theodor Schwann,
applied the discovery at once to the animal organism. He showed that
in the animal body as well, when we examine its tissues in the
microscope, we find these cells everywhere to be the elementary units.
All the different tissues of the organism, especially the very
dissimilar tissues of the nerves, muscles, bones, external skin,
mucous lining, etc., are originally formed out of cells; and this is
also true of all the tissues of the plant. These cells are separate
living beings; they are the citizens of the State which the entire
multicellular organism seems to be. This important discovery was bound
to be of service to embryology, as it raised a number of new
questions. What is the relation of the cells to the germinal layers?
Are the germinal layers composed of cells, and what is their relation
to the cells of the tissues that form later? How does the ovum stand
in the cellular theory? Is the ovum itself a cell, or is it composed
of cells? These important questions were now imposed on the
embryologist by the cellular theory.

The most notable effort to answer these questions--which were attacked
on all sides by different students--is contained in the famous work,
Inquiries into the Development of the Vertebrates (not translated) of
Robert Remak, of Berlin (1851). This gifted scientist succeeded in
mastering, by a complete reform of the science, the great difficulties
which the cellular theory had at first put in the way of embryology. A
Berlin anatomist, Carl Boguslaus Reichert, had already attempted to
explain the origin of the tissues. But this attempt was bound to
miscarry, since its not very clear-headed author lacked a sound
acquaintance with embryology and the cell theory, and even with the
structure and development of the tissue in particular. Remak at length
brought order into the dreadful confusion that Reichert had caused; he
gave a perfectly simple explanation of the origin of the tissues. In
his opinion the animal ovum is always a simple cell: the germinal
layers which develop out of it are always composed of cells; and these
cells that constitute the germinal layers arise simply from the
continuous and repeated cleaving (segmentation) of the original
solitary cell. It first divides into two and then into four cells; out
of these four cells are born eight, then sixteen, thirty-two, and so
on. Thus, in the embryonic development of every animal and plant there
is formed first of all out of the simple egg cell, by a repeated
subdivision, a cluster of cells, as Kolliker had already stated in
connection with the cephalopods in 1844. The cells of this group
spread themselves out flat and form leaves or plates; each of these
leaves is formed exclusively out of cells. The cells of different
layers assume different shapes, increase, and differentiate; and in
the end there is a further cleavage (differentiation) and division of
work of the cells within the layers, and from these all the different
tissues of the body proceed.

These are the simple foundations of histogeny, or the science that
treats of the development of the tissues (hista), as it was
established by Remak and Kolliker. Remak, in determining more closely
the part which the different germinal layers play in the formation of
the various tissues and organs, and in applying the theory of
evolution to the cells and the tissues they compose, raised the theory
of germinal layers, at least as far as it regards the vertebrates, to
a high degree of perfection.

Remak showed that three layers are formed out of the two germinal
layers which compose the first simple leaf-shaped structure of the
vertebrate body (or the "germinal disk"), as the lower layer splits
into two plates. These three layers have a very definite relation to
the various tissues. First of all, the cells which form the outer skin
of the body (the epidermis), with its various dependencies (hairs,
nails, etc.)--that is to say, the entire outer envelope of the
body--are developed out of the outer or upper layer; but there are
also developed in a curious way out of the same layer the cells which
form the central nervous system, the brain and the spinal cord. In the
second place, the inner or lower germinal layer gives rise only to the
cells which form the epithelium (the whole inner lining) of the
alimentary canal and all that depends on it (the lungs, liver,
pancreas, etc.), or the tissues that receive and prepare the
nourishment of the body. Finally, the middle layer gives rise to all
the other tissues of the body, the muscles, blood, bones, cartilage,
etc. Remak further proved that this middle layer, which he calls "the
motor-germinative layer," proceeds to subdivide into two secondary
layers. Thus we find once more the four layers which Baer had
indicated. Remak calls the outer secondary leaf of the middle layer
(Baer's "muscular layer") the "skin layer" (it would be better to say,
skin-fibre layer); it forms the outer wall of the body (the true skin,
the muscles, etc.). To the inner secondary leaf (Baer's "vascular
layer") he gave the name of the "alimentary-fibre layer"; this forms
the outer envelope of the alimentary canal, with the mesentery, the
heart, the blood-vessels, etc.

On this firm foundation provided by Remak for histogeny, or the
science of the formation of the tissues, our knowledge has been
gradually built up and enlarged in detail. There have been several
attempts to restrict and even destroy Remak's principles. The two
anatomists, Reichert (of Berlin) and Wilhelm His (of Leipzic),
especially, have endeavoured in their works to introduce a new
conception of the embryonic development of the vertebrate, according
to which the two primary germinal layers would not be the sole sources
of formation. But these efforts were so seriously marred by ignorance
of comparative anatomy, an imperfect acquaintance with ontogenesis,
and a complete neglect of phylogenesis, that they could not have more
than a passing success. We can only explain how these curious attacks
of Reichert and His came to be regarded for a time as advances by the
general lack of discrimination and of grasp of the true object of
embryology.

Wilhelm His published, in 1868, his extensive Researches into the
Earliest Form of the Vertebrate Body,* (* None of His's works have
been translated into English.) one of the curiosities of embryological
literature. The author imagines that he can build a "mechanical theory
of embryonic development" by merely giving an exact description of the
embryology of the chick, without any regard to comparative anatomy and
phylogeny, and thus falls into an error that is almost without
parallel in the history of biological literature. As the final result
of his laborious investigations, His tells us "that a comparatively
simple law of growth is the one essential thing in the first
development. Every formation, whether it consist in cleavage of
layers, or folding, or complete division, is a consequence of this
fundamental law." Unfortunately, he does not explain what this "law of
growth" is; just as other opponents of the theory of selection, who
would put in its place a great "law of evolution," omit to tell us
anything about the nature of this. Nevertheless, it is quite clear
from His's works that he imagines constructive Nature to be a sort of
skilful tailor. The ingenious operator succeeds in bringing into
existence, by "evolution," all the various forms of living things by
cutting up in different ways the germinal layers, bending and folding,
tugging and splitting, and so on.

His's embryological theories excited a good deal of interest at the
time of publication, and have evoked a fair amount of literature in
the last few decades. He professed to explain the most complicated
parts of organic construction (such as the development of the brain)
in the simplest way on mechanical principles, and to derive them
immediately from simple physical processes (such as unequal
distribution of strain in an elastic plate). It is quite true that a
mechanical or monistic explanation (or a reduction of natural
processes) is the ideal of modern science, and this ideal would be
realised if we could succeed in expressing these formative processes
in mathematical formulae. His has, therefore, inserted plenty of
numbers and measurements in his embryological works, and given them an
air of "exact" scholarship by putting in a quantity of mathematical
tables. Unfortunately, they are of no value, and do not help us in the
least in forming an "exact" acquaintance with the embryonic phenomena.
Indeed, they wander from the true path altogether by neglecting the
phylogenetic method; this, he thinks, is "a mere by-path," and is "not
necessary at all for the explanation of the facts of embryology,"
which are the direct consequence of physiological principles. What His
takes to be a simple physical process--for instance, the folding of
the germinal layers (in the formation of the medullary tube,
alimentary tube, etc.)--is, as a matter of fact, the direct result of
the growth of the various cells which form those organic structures;
but these growth-motions have themselves been transmitted by heredity
from parents and ancestors, and are only the hereditary repetition of
countless phylogenetic changes which have taken place for thousands of
years in the race-history of the said ancestors. Each of these
historical changes was, of course, originally due to adaptation; it
was, in other words, physiological, and reducible to mechanical
causes. But we have, naturally, no means of observing them now. It is
only by the hypotheses of the science of evolution that we can form an
approximate idea of the organic links in this historic chain.

All the best recent research in animal embryology has led to the
confirmation and development of Baer and Remak's theory of the
germinal layers. One of the most important advances in this direction
of late was the discovery that the two primary layers out of which is
built the body of all vertebrates (including man) are also present in
all the invertebrates, with the sole exception of the lowest group,
the unicellular protozoa. Huxley had detected them in the medusa in
1849. He showed that the two layers of cells from which the body of
this zoophyte is developed correspond, both morphologically and
physiologically, to the two original germinal layers of the
vertebrate. The outer layer, from which come the external skin and the
muscles, was then called by Allman (1853) the "ectoderm" (outer layer,
or skin); the inner layer, which forms the alimentary and reproductory
organs, was called the "entoderm" (= inner layer). In 1867 and the
following years the discovery of the germinal layers was extended to
other groups of the invertebrates. In particular, the indefatigable
Russian zoologist, Kowalevsky, found them in all the most diverse
sections of the invertebrates--the worms, tunicates, echinoderms,
molluscs, articulates, etc.

In my monograph on the sponges (1872) I proved that these two primary
germinal layers are also found in that group, and that they may be
traced from it right up to man, through all the various classes, in
identical form. This "homology of the two primary germinal layers"
extends through the whole of the metazoa, or tissue-forming animals;
that is to say, through the whole animal kingdom, with the one
exception of its lowest section, the unicellular beings, or protozoa.
These lowly organised animals do not form germinal layers, and
therefore do not succeed in forming true tissue. Their whole body
consists of a single cell (as is the case with the amoebae and
infusoria), or of a loose aggregation of only slightly differentiated
cells, though it may not even reach the full structure of a single
cell (as with the monera). But in all other animals the ovum first
grows into two primary layers, the outer or animal layer (the
ectoderm, epiblast, or ectoblast), and the inner or vegetal layer (the
entoderm, hypoblast, or endoblast); and from these the tissues and
organs are formed. The first and oldest organ of all these metazoa is
the primitive gut (or progaster) and its opening, the primitive mouth
(prostoma). The typical embryonic form of the metazoa, as it is
presented for a time by this simple structure of the two-layered body,
is called the gastrula; it is to be conceived as the hereditary
reproduction of some primitive common ancestor of the metazoa, which
we call the gastraea. This applies to the sponges and other zoophyta,
and to the worms, the mollusca, echinoderma, articulata, and
vertebrata. All these animals may be comprised under the general
heading of "gut animals," or metazoa, in contradistinction to the
gutless protozoa.

I have pointed out in my Study of the Gastraea Theory [not translated]
(1873) the important consequences of this conception in the morphology
and classification of the animal world. I also divided the realm of
metazoa into two great groups, the lower and higher metazoa. In the
first are comprised the coelenterata (also called zoophytes, or
plant-animals). In the lower forms of this group the body consists
throughout life merely of the primary germinal layers, with the cells
sometimes more and sometimes less differentiated. But with the higher
forms of the coelentarata (the corals, higher medusae, ctenophorae,
and platodes) a middle layer, or mesoderm, often of considerable size,
is developed between the other two layers; but blood and an internal
cavity are still lacking.

To the second great group of the metazoa I gave the name of the
coelomaria, or bilaterata (or the bilateral higher forms). They all
have a cavity within the body (coeloma), and most of them have blood
and blood-vessels. In this are comprised the six higher stems of the
animal kingdom, the annulata and their descendants, the mollusca,
echinoderma, articulata, tunicata, and vertebrata. In all these
bilateral organisms the two-sided body is formed out of four secondary
germinal layers, of which the inner two construct the wall of the
alimentary canal, and the outer two the wall of the body. Between the
two pairs of layers lies the cavity (coeloma).

Although I laid special stress on the great morphological importance
of this cavity in my Study of the Gastraea Theory, and endeavoured to
prove the significance of the four secondary germinal layers in the
organisation of the coelomaria, I was unable to deal satisfactorily
with the difficult question of the mode of their origin. This was done
eight years afterwards by the brothers Oscar and Richard Hertwig in
their careful and extensive comparative studies. In their masterly
Coelum Theory: An Attempt to Explain the Middle Germinal Layer [not
translated] (1881) they showed that in most of the metazoa, especially
in all the vertebrates, the body-cavity arises in the same way, by the
outgrowth of two sacs from the inner layer. These two coelom-pouches
proceed from the rudimentary mouth of the gastrula, between the two
primary layers. The inner plate of the two-layered coelom-pouch (the
visceral layer) joins itself to the entoderm; the outer plate
(parietal layer) unites with the ectoderm. Thus are formed the
double-layered gut-wall within and the double-layered body-wall
without; and between the two is formed the cavity of the coelom, by
the blending of the right and left coelom-sacs. We shall see this more
fully in Chapter 1.10.

The many new points of view and fresh ideas suggested by my gastraea
theory and Hertwig's coelom theory led to the publication of a number
of writings on the theory of germinal layers. Most of them set out to
oppose it at first, but in the end the majority supported it. Of late
years both theories are accepted in their essential features by nearly
every competent man of science, and light and order have been
introduced into this once dark and contradictory field of research. A
further cause of congratulation for this solution of the great
embryological controversy is that it brought with it a recognition of
the need for phylogenetic study and explanation.

Interest and practice in embryological research have been remarkably
stimulated during the past thirty years by this appreciation of
phylogenetic methods. Hundreds of assiduous and able observers are now
engaged in the development of comparative embryology and its
establishment on a basis of evolution, whereas they numbered only a
few dozen not many decades ago. It would take too long to enumerate
even the most important of the countless valuable works which have
enriched embryological literature since that time. References to them
will be found in the latest manuals of embryology of Kolliker,
Balfour, Hertwig, Kollman, Korschelt, and Heider.

Kolliker's Entwickelungsgeschichte des Menschen und der hoherer
Thiere, the first edition of which appeared forty-two years ago, had
the rare merit at that time of gathering into presentable form the
scattered attainments of the science, and expounding them in some sort
of unity on the basis of the cellular theory and the theory of
germinal layers. Unfortunately, the distinguished Wurtzburg anatomist,
to whom comparative anatomy, histology, and ontogeny owe so much, is
opposed to the theory of descent generally and to Darwinism in
particular. All the other manuals I have mentioned take a decided
stand on evolution. Francis Balfour has carefully collected and
presented with discrimination, in his Manual of Comparative Embryology
(1880), the very scattered and extensive literature of the subject; he
has also widened the basis of the gastraea theory by a comparative
description of the rise of the organs from the germinal layers in all
the chief groups of the animal kingdom, and has given a most thorough
empirical support to the principles I have formulated. A comparison of
his work with the excellent Text-book of the Embryology of the
Vertebrates (1890) [translation 1895] of Korschelt and Heider shows
what astonishing progress has been made in the science in the course
of ten years. I would especially recommend the manuals of Julius
Kollmann and Oscar Hertwig to those readers who are stimulated to
further study by these chapters on human embryology. Kollmann's work
is commendable for its clear treatment of the subject and very fine
original illustrations; its author adheres firmly to the biogenetic
law, and uses it throughout with considerable profit. That is not the
case in Oscar Hertwig's recent Text-book of the Embryology of Man and
the Mammals [translations 1892 and 1899] (seventh edition 1902). This
able anatomist has of late often been quoted as an opponent of the
biogenetic law, although he himself had demonstrated its great value
thirty years ago. His recent vacillation is partly due to the timidity
which our "exact" scientists have with regard to hypotheses; though it
is impossible to make any headway in the explanation of facts without
them. However, the purely descriptive part of embryology in Hertwig's
Text-book is very thorough and reliable.

A new branch of embryological research has been studied very
assiduously in the last decade of the nineteenth century--namely,
"experimental embryology." The great importance which has been
attached to the application of physical experiments to the living
organism for the last hundred years, and the valuable results that it
has given to physiology in the study of the vital phenomena, have led
to its extension to embryology. I was the first to make experiments of
this kind during a stay of four months on the Canary Island,
Lanzerote, in 1866. I there made a thorough investigation of the
almost unknown embryology of the siphonophorae. I cut a number of the
embryos of these animals (which develop freely in the water, and pass
through a very curious transformation), at an early stage, into
several pieces, and found that a fresh organism (more or less
complete, according to the size of the piece) was developed from each
particle. More recently some of my pupils have made similar
experiments with the embryos of vertebrates (especially the frog) and
some of the invertebrates. Wilhelm Roux, in particular, has made
extensive experiments, and based on them a special "mechanical
embryology," which has given rise to a good deal of discussion and
controversy. Roux has published a special journal for these subjects
since 1895, the Archiv fur Entwickelungsmechanik. The contributions to
it are very varied in value. Many of them are valuable papers on the
physiology and pathology of the embryo. Pathological experiments--the
placing of the embryo in abnormal conditions--have yielded many
interesting results; just as the physiology of the normal body has for
a long time derived assistance from the pathology of the diseased
organism. Other of these mechanical-embryological articles return to
the erroneous methods of His, and are only misleading. This must be
said of the many contributions of mechanical embryology which take up
a position of hostility to the theory of descent and its chief
embryological foundation--the biogenetic law. This law, however, when
rightly understood, is not opposed to, but is the best and most solid
support of, a sound mechanical embryology. Impartial reflection and a
due attention to paleontology and comparative anatomy should convince
these one-sided mechanicists that the facts they have discovered--and,
indeed, the whole embryological process--cannot be fully understood
without the theory of descent and the biogenetic law.


CHAPTER 1.4. THE OLDER PHYLOGENY.

The embryology of man and the animals, the history of which we have
reviewed in the last two chapters, was mainly a descriptive science
forty years ago. The earlier investigations in this province were
chiefly directed to the discovery, by careful observation, of the
wonderful facts of the embryonic development of the animal body from
the ovum. Forty years ago no one dared attack the question of the
CAUSES of these phenomena. For fully a century, from the year 1759,
when Wolff's solid Theoria generationis appeared, until 1859, when
Darwin published his famous Origin of Species, the real causes of the
embryonic processes were quite unknown. No one thought of seeking the
agencies that effected this marvellous succession of structures. The
task was thought to be so difficult as almost to pass beyond the
limits of human thought. It was reserved for Charles Darwin to
initiate us into the knowledge of these causes. This compels us to
recognise in this great genius, who wrought a complete revolution in
the whole field of biology, a founder at the same time of a new period
in embryology. It is true that Darwin occupied himself very little
with direct embryological research, and even in his chief work he only
touches incidentally on the embryonic phenomena; but by his reform of
the theory of descent and the founding of the theory of selection he
has given us the means of attaining to a real knowledge of the causes
of embryonic formation. That is, in my opinion, the chief feature in
Darwin's incalculable influence on the whole science of evolution.

When we turn our attention to this latest period of embryological
research, we pass into the second division of organic
evolution--stem-evolution, or phylogeny. I have already indicated in
Chapter 1.1 the important and intimate causal connection between these
two sections of the science of evolution--between the evolution of the
individual and that of his ancestors. We have formulated this
connection in the biogenetic law; the shorter evolution, that of the
individual, or ontogenesis, is a rapid and summary repetition, a
condensed recapitulation, of the larger evolution, or that of the
species. In this principle we express all the essential points
relating to the causes of evolution; and we shall seek throughout this
work to confirm this principle and lend it the support of facts. When
we look to its CAUSAL significance, perhaps it would be better to
formulate the biogenetic law thus: "The evolution of the species and
the stem (phylon) shows us, in the physiological functions of heredity
and adaptation, the conditioning causes on which the evolution of the
individual depends"; or, more briefly: "Phylogenesis is the mechanical
cause of ontogenesis."

But before we examine the great achievement by which Darwin revealed
the causes of evolution to us, we must glance at the efforts of
earlier scientists to attain this object. Our historical inquiry into
these will be even shorter than that into the work done in the field
of ontogeny. We have very few names to consider here. At the head of
them we find the great French naturalist, Jean Lamarck, who first
established evolution as a scientific theory in 1809. Even before his
time, however, the chief philosopher, Kant, and the chief poet,
Goethe, of Germany had occupied themselves with the subject. But their
efforts passed almost without recognition in the eighteenth century. A
"philosophy of nature" did not arise until the beginning of the
nineteenth century. In the whole of the time before this no one had
ventured to raise seriously the question of the origin of species,
which is the culminating point of phylogeny. On all sides it was
regarded as an insoluble enigma.

The whole science of the evolution of man and the other animals is
intimately connected with the question of the nature of species, or
with the problem of the origin of the various animals which we group
together under the name of species. Thus the definition of the species
becomes important. It is well known that this definition was given by
Linne, who, in his famous Systema Naturae (1735), was the first to
classify and name the various groups of animals and plants, and drew
up an orderly scheme of the species then known. Since that time
"species" has been the most important and indispensable idea in
descriptive natural history, in zoological and botanical
classification; although there have been endless controversies as to
its real meaning.

What, then, is this "organic species"? Linne himself appealed directly
to the Mosaic narrative; he believed that, as it is stated in Genesis,
one pair of each species of animals and plants was created in the
beginning, and that all the individuals of each species are the
descendants of these created couples. As for the hermaphrodites
(organisms that have male and female organs in one being), he thought
it sufficed to assume the creation of one sole individual, since this
would be fully competent to propagate its species. Further developing
these mystic ideas, Linne went on to borrow from Genesis the account
of the deluge and of Noah's ark as a ground for a science of the
geographical and topographical distribution of organisms. He accepted
the story that all the plants, animals, and men on the earth were
swept away in a universal deluge, except the couples preserved with
Noah in the ark, and ultimately landed on Mount Ararat. This mountain
seemed to Linne particularly suitable for the landing, as it reaches a
height of more than 16,000 feet, and thus provides in its higher zones
the several climates demanded by the various species of animals and
plants: the animals that were accustomed to a cold climate could
remain at the summit; those used to a warm climate could descend to
the foot; and those requiring a temperate climate could remain
half-way down. From this point the re-population of the earth with
animals and plants could proceed.

It was impossible to have any scientific notion of the method of
evolution in Linne's time, as one of the chief sources of information,
paleontology, was still wholly unknown. This science of the fossil
remains of extinct animals and plants is very closely bound up with
the whole question of evolution. It is impossible to explain the
origin of living organisms without appealing to it. But this science
did not rise until a much later date. The real founder of scientific
paleontology was Georges Cuvier, the most distinguished zoologist who,
after Linne, worked at the classification of the animal world, and
effected a complete revolution in systematic zoology at the beginning
of the nineteenth century. In regard to the nature of the species he
associated himself with Linne and the Mosaic story of creation, though
this was more difficult for him with his acquaintance with fossil
remains. He clearly showed that a number of quite different animal
populations have lived on the earth; and he claimed that we must
distinguish a number of stages in the history of our planet, each of
which was characterised by a special population of animals and plants.
These successive populations were, he said, quite independent of each
other, and therefore the supernatural creative act, which was demanded
as the origin of the animals and plants by the dominant creed, must
have been repeated several times. In this way a whole series of
different creative periods must have succeeded each other; and in
connection with these he had to assume that stupendous revolutions or
cataclysms--something like the legendary deluge--must have taken place
repeatedly. Cuvier was all the more interested in these catastrophes
or cataclysms as geology was just beginning to assert itself, and
great progress was being made in our knowledge of the structure and
formation of the earth's crust. The various strata of the crust were
being carefully examined, especially by the famous geologist Werner
and his school, and the fossils found in them were being classified;
and these researches also seemed to point to a variety of creative
periods. In each period the earth's crust, composed of the various
strata, seemed to be differently constituted, just like the population
of animals and plants that then lived on it. Cuvier combined this
notion with the results of his own paleontological and zoological
research; and in his effort to get a consistent view of the whole
process of the earth's history he came to form the theory which is
known as "the catastrophic theory," or the theory of terrestrial
revolutions. According to this theory, there have been a series of
mighty cataclysms on the earth, and these have suddenly destroyed the
whole animal and plant population then living on it; after each
cataclysm there was a fresh creation of living things throughout the
earth. As this creation could not be explained by natural laws, it was
necessary to appeal to an intervention on the part of the Creator.
This catastrophic theory, which Cuvier described in a special work,
was soon generally accepted, and retained its position in biology for
half a century.

However, Cuvier's theory was completely overthrown sixty years ago by
the geologists, led by Charles Lyell, the most distinguished worker in
this field of science. Lyell proved in his famous Principles of
Geology (1830) that the theory was false, in so far as it concerned
the crust of the earth; that it was totally unnecessary to bring in
supernatural agencies or general catastrophes in order to explain the
structure and formation of the mountains; and that we can explain them
by the familiar agencies which are at work to-day in altering and
reconstructing the surface of the earth. These causes are--the action
of the atmosphere and water in its various forms (snow, ice, fog,
rain, the wear of the river, and the stormy ocean), and the volcanic
action which is exerted by the molten central mass. Lyell convincingly
proved that these natural causes are quite adequate to explain every
feature in the build and formation of the crust. Hence Cuvier's theory
of cataclysms was very soon driven out of the province of geology,
though it remained for another thirty years in undisputed authority in
biology. All the zoologists and botanists who gave any thought to the
question of the origin of organisms adhered to Cuvier's erroneous idea
of revolutions and new creations.

In order to illustrate the complete stagnancy of biology from 1830 to
1859 on the question of the origin of the various species of animals
and plants, I may say, from my own experience, that during the whole
of my university studies I never heard a single word said about this
most important problem of the science. I was fortunate enough at that
time (1852 to 1857) to have the most distinguished masters for every
branch of biological science. Not one of them ever mentioned this
question of the origin of species. Not a word was ever said about the
earlier efforts to understand the formation of living things, nor
about Lamarck's Philosophie Zoologique which had made a fresh attack
on the problem in 1809. Hence it is easy to understand the enormous
opposition that Darwin encountered when he took up the question for
the first time. His views seemed to float in the air, without a single
previous effort to support them. The whole question of the formation
of living things was considered by biologists, until 1859, as
pertaining to the province of religion and transcendentalism; even in
speculative philosophy, in which the question had been approached from
various sides, no one had ventured to give it serious treatment. This
was due to the dualistic system of Immanuel Kant, who taught a natural
system of evolution as far as the inorganic world was concerned; but,
on the whole, adopted a supernaturalist system as regards the origin
of living things. He even went so far as to say: "It is quite certain
that we cannot even satisfactorily understand, much less explain, the
nature of an organism and its internal forces on purely mechanical
principles; it is so certain, indeed, that we may confidently say: 'It
is absurd for a man to imagine even that some day a Newton will arise
who will explain the origin of a single blade of grass by natural laws
not controlled by design'--such a hope is entirely forbidden us." In
these words Kant definitely adopts the dualistic and teleological
point of view for biological science.

Nevertheless, Kant deserted this point of view at times, particularly
in several remarkable passages which I have dealt with at length in my
Natural History of Creation (chapter 5), where he expresses himself in
the opposite, or monistic, sense. In fact, these passages would
justify one, as I showed, in claiming his support for the theory of
evolution. However, these monistic passages are only stray gleams of
light; as a rule, Kant adheres in biology to the obscure dualistic
ideas, according to which the forces at work in inorganic nature are
quite different from those of the organic world. This dualistic system
prevails in academic philosophy to-day--most of our philosophers still
regarding these two provinces as totally distinct. They put, on the
one side, the inorganic or "lifeless" world, in which there are at
work only mechanical laws, acting necessarily and without design; and,
on the other, the province of organic nature, in which none of the
phenomena can be properly understood, either as regards their inner
nature or their origin, except in the light of preconceived design,
carried out by final or purposive causes.

The prevalence of this unfortunate dualistic prejudice prevented the
problem of the origin of species, and the connected question of the
origin of man, from being regarded by the bulk of people as a
scientific question at all until 1859. Nevertheless, a few
distinguished students, free from the current prejudice, began, at the
commencement of the nineteenth century, to make a serious attack on
the problem. The merit of this attaches particularly to what is known
as "the older school of natural philosophy," which has been so much
misrepresented, and which included Jean Lamarck, Buffon, Geoffroy St.
Hilaire, and Blainville in France; Wolfgang Goethe, Reinhold
Treviranus, Schelling, and Lorentz Oken in Germany [and Erasmus Darwin
in England].

The gifted natural philosopher who treated this difficult question
with the greatest sagacity and comprehensiveness was Jean Lamarck. He
was born at Bazentin, in Picardy, on August 1st, 1744; he was the son
of a clergyman, and was destined for the Church. But he turned to seek
glory in the army, and eventually devoted himself to science.

His Philosophie Zoologique was the first scientific attempt to sketch
the real course of the origin of species, the first "natural history
of creation" of plants, animals, and men. But, as in the case of
Wolff's book, this remarkably able work had no influence whatever;
neither one nor the other could obtain any recognition from their
prejudiced contemporaries. No man of science was stimulated to take an
interest in the work, and to develop the germs it contained of the
most important biological truths. The most distinguished botanists and
zoologists entirely rejected it, and did not even deign to reply to
it. Cuvier, who lived and worked in the same city, has not thought fit
to devote a single syllable to this great achievement in his memoir on
progress in the sciences, in which the pettiest observations found a
place. In short, Lamarck's Philosophie Zoologique shared the fate of
Wolff's theory of development, and was for half a century ignored and
neglected. The German scientists, especially Oken and Goethe, who were
occupied with similar speculations at the same time, seem to have
known nothing about Lamarck's work. If they had known it, they would
have been greatly helped by it, and might have carried the theory of
evolution much farther than they found it possible to do.

To give an idea of the great importance of the Philosophie Zoologique,
I will briefly explain Lamarck's leading thought. He held that there
was no essential difference between living and lifeless beings. Nature
is one united and connected system of phenomena; and the forces which
fashion the lifeless bodies are the only ones at work in the kingdom
of living things. We have, therefore, to use the same method of
investigation and explanation in both provinces. Life is only a
physical phenomenon. All the plants and animals, with man at their
head, are to be explained, in structure and life, by mechanical or
efficient causes, without any appeal to final causes, just as in the
case of minerals and other inorganic bodies. This applies equally to
the origin of the various species. We must not assume any original
creation, or repeated creations (as in Cuvier's theory), to explain
this, but a natural, continuous, and necessary evolution. The whole
evolutionary process has been uninterrupted. All the different kinds
of animals and plants which we see to-day, or that have ever lived,
have descended in a natural way from earlier and different species;
all come from one common stock, or from a few common ancestors. These
remote ancestors must have been quite simple organisms of the lowest
type, arising by spontaneous generation from inorganic matter. The
succeeding species have been constantly modified by adaptation to
their varying environment (especially by use and habit), and have
transmitted their modifications to their successors by heredity.

Lamarck was the first to formulate as a scientific theory the natural
origin of living things, including man, and to push the theory to its
extreme conclusions--the rise of the earliest organisms by spontaneous
generation (or abiogenesis) and the descent of man from the nearest
related mammal, the ape. He sought to explain this last point, which
is of especial interest to us here, by the same agencies which he
found at work in the natural origin of the plant and animal species.
He considered use and habit (adaptation) on the one hand, and heredity
on the other, to be the chief of these agencies. The most important
modifications of the organs of plants and animals are due, in his
opinion, to the function of these very organs, or to the use or disuse
of them. To give a few examples, the woodpecker and the humming-bird
have got their peculiarly long tongues from the habit of extracting
their food with their tongues from deep and narrow folds or canals;
the frog has developed the web between his toes by his own swimming;
the giraffe has lengthened his neck by stretching up to the higher
branches of trees, and so on. It is quite certain that this use or
disuse of organs is a most important factor in organic development,
but it is not sufficient to explain the origin of species.

To adaptation we must add heredity as the second and not less
important agency, as Lamarck perfectly recognised. He said that the
modification of the organs in any one individual by use or disuse was
slight, but that it was increased by accumulation in passing by
heredity from generation to generation. But he missed altogether the
principle which Darwin afterwards found to be the chief factor in the
theory of transformation--namely, the principle of natural selection
in the struggle for existence. It was partly owing to his failure to
detect this supremely important element, and partly to the poor
condition of all biological science at the time, that Lamarck did not
succeed in establishing more firmly his theory of the common descent
of man and the other animals.

Independently of Lamarck, the older German school of natural
philosophy, especially Reinhold Treviranus, in his Biologie (1802),
and Lorentz Oken, in his Naturphilosophie (1809), turned its attention
to the problem of evolution about the end of the eighteenth and
beginning of the nineteenth century. I have described its work in my
History of Creation (chapter 4). Here I can only deal with the
brilliant genius whose evolutionary ideas are of special interest--the
greatest of German poets, Wolfgang Goethe. With his keen eye for the
beauties of nature, and his profound insight into its life, Goethe was
early attracted to the study of various natural sciences. It was the
favourite occupation of his leisure hours throughout life. He gave
particular and protracted attention to the theory of colours. But the
most valuable of his scientific studies are those which relate to that
"living, glorious, precious thing," the organism. He made profound
research into the science of structures or morphology (morphae =
forms). Here, with the aid of comparative anatomy, he obtained the
most brilliant results, and went far in advance of his time. I may
mention, in particular, his vertebral theory of the skull, his
discovery of the pineal gland in man, his system of the metamorphosis
of plants, etc. These morphological studies led Goethe on to research
into the formation and modification of organic structures which we
must count as the first germ of the science of evolution. He
approaches so near to the theory of descent that we must regard him,
after Lamarck, as one of its earliest founders. It is true that he
never formulated a complete scientific theory of evolution, but we
find a number of remarkable suggestions of it in his splendid
miscellaneous essays on morphology. Some of them are really among the
very basic ideas of the science of evolution. He says, for instance
(1807): "When we compare plants and animals in their most rudimentary
forms, it is almost impossible to distinguish between them. But we may
say that the plants and animals, beginning with an almost inseparable
closeness, gradually advance along two divergent lines, until the
plant at last grows in the solid, enduring tree and the animal attains
in man to the highest degree of mobility and freedom." That Goethe was
not merely speaking in a poetical, but in a literal genealogical,
sense of this close affinity of organic forms is clear from other
remarkable passages in which he treats of their variety in outward
form and unity in internal structure. He believes that every living
thing has arisen by the interaction of two opposing formative forces
or impulses. The internal or "centripetal" force, the type or "impulse
to specification," seeks to maintain the constancy of the specific
forms in the succession of generations: this is heredity. The external
or "centrifugal" force, the element of variation or "impulse to
metamorphosis," is continually modifying the species by changing their
environment: this is adaptation. In these significant conceptions
Goethe approaches very close to a recognition of the two great
mechanical factors which we now assign as the chief causes of the
formation of species.

However, in order to appreciate Goethe's views on morphology, one must
associate his decidedly monistic conception of nature with his
pantheistic philosophy. The warm and keen interest with which he
followed, in his last years, the controversies of contemporary French
scientists, and especially the struggle between Cuvier and Geoffroy
St. Hilaire (see chapter 4 of The History of Creation), is very
characteristic. It is also necessary to be familiar with his style and
general tenour of thought in order to appreciate rightly the many
allusions to evolution found in his writings. Otherwise, one is apt to
make serious errors.

He approached so close, at the end of the eighteenth century, to the
principles of the science of evolution that he may well be described
as the first forerunner of Darwin, although he did not go so far as to
formulate evolution as a scientific system, as Lamarck did.


CHAPTER 1.5. THE MODERN SCIENCE OF EVOLUTION.

We owe so much of the progress of scientific knowledge to Darwin's
Origin of Species that its influence is almost without parallel in the
history of science. The literature of Darwinism grows from day to day,
not only on the side of academic zoology and botany, the sciences
which were chiefly affected by Darwin's theory, but in a far wider
circle, so that we find Darwinism discussed in popular literature with
a vigour and zest that are given to no other scientific conception.
This remarkable success is due chiefly to two circumstances. In the
first place, all the sciences, and especially biology, have made
astounding progress in the last half-century, and have furnished a
very vast quantity of proofs of the theory of evolution. In striking
contrast to the failure of Lamarck and the older scientists to attract
attention to their effort to explain the origin of living things and
of man, we have this second and successful effort of Darwin, which was
able to gather to its support a large number of established facts.
Availing himself of the progress already made, he had very different
scientific proofs to allege than Lamarck, or St. Hilaire, or Goethe,
or Treviranus had had. But, in the second place, we must acknowledge
that Darwin had the special distinction of approaching the subject
from an entirely new side, and of basing the theory of descent on a
consistent system, which now goes by the name of Darwinism.

Lamarck had unsuccessfully attempted to explain the modification of
organisms that descend from a common form chiefly by the action of
habit and the use of organs, though with the aid of heredity. But
Darwin's success was complete when he independently sought to give a
mechanical explanation, on a quite new ground, of this modification of
plant and animal structures by adaptation and heredity. He was
impelled to his theory of selection on the following grounds. He
compared the origin of the various kinds of animals and plants which
we modify artificially--by the action of artificial selection in
horticulture and among domestic animals--with the origin of the
species of animals and plants in their natural state. He then found
that the agencies which we employ in the modification of forms by
artificial selection are also at work in Nature. The chief of these
agencies he held to be "the struggle for life." The gist of this
peculiarly Darwinian idea is given in this formula: The struggle for
existence produces new species without premeditated design in the life
of Nature, in the same way that the will of man consciously selects
new races in artificial conditions. The gardener or the farmer selects
new forms as he wills for his own profit, by ingeniously using the
agency of heredity and adaptation for the modification of structures;
so, in the natural state, the struggle for life is always
unconsciously modifying the various species of living things. This
struggle for life, or competition of organisms in securing the means
of subsistence, acts without any conscious design, but it is none the
less effective in modifying structures. As heredity and adaptation
enter into the closest reciprocal action under its influence, new
structures, or alterations of structure, are produced; and these are
purposive in the sense that they serve the organism when formed, but
they were produced without any pre-conceived aim.

This simple idea is the central thought of Darwinism, or the theory of
selection. Darwin conceived this idea at an early date, and then, for
more than twenty years, worked at the collection of empirical evidence
in support of it before he published his theory. His grandfather,
Erasmus Darwin, was an able scientist of the older school of natural
philosophy, who published a number of natural-philosophic works about
the end of the eighteenth century. The most important of them is his
Zoonomia, published in 1794, in which he expounds views similar to
those of Goethe and Lamarck, without really knowing anything of the
work of these contemporaries. However, in the writings of the
grandfather the plastic imagination rather outran the judgment, while
in Charles Darwin the two were better balanced.

Darwin did not publish any account of his theory until 1858, when
Alfred Russel Wallace, who had independently reached the same theory
of selection, published his own work. In the following year appeared
the Origin of Species, in which he develops it at length and supports
it with a mass of proof. Wallace had reached the same conclusion, but
he had not so clear a perception as Darwin of the effectiveness of
natural selection in forming species, and did not develop the theory
so fully. Nevertheless, Wallace's writings, especially those on
mimicry, etc., and an admirable work on The Geographical Distribution
of Animals, contain many fine original contributions to the theory of
selection. Unfortunately, this gifted scientist has since devoted
himself to spiritism.* (* Darwin and Wallace arrived at the theory
quite independently. Vide Wallace's Contributions to the Theory of
Natural Selection (1870) and Darwinism (1891).)

Darwin's Origin of Species had an extraordinary influence, though not
at first on the experts of the science. It took zoologists and
botanists several years to recover from the astonishment into which
they had been thrown through the revolutionary idea of the work. But
its influence on the special sciences with which we zoologists and
botanists are concerned has increased from year to year; it has
introduced a most healthy fermentation in every branch of biology,
especially in comparative anatomy and ontogeny, and in zoological and
botanical classification. In this way it has brought about almost a
revolution in the prevailing views.

However, the point which chiefly concerns us here--the extension of
the theory to man--was not touched at all in Darwin's first work in
1859. It was believed for several years that he had no thought of
applying his principles to man, but that he shared the current idea of
man holding a special position in the universe. Not only ignorant
laymen (especially several theologians), but also a number of men of
science, said very naively that Darwinism in itself was not to be
opposed; that it was quite right to use it to explain the origin of
the various species of plants and animals, but that it was totally
inapplicable to man.

In the meantime, however, it seemed to a good many thoughtful people,
laymen as well as scientists, that this was wrong; that the descent of
man from some other animal species, and immediately from some ape-like
mammal, followed logically and necessarily from Darwin's reformed
theory of evolution. Many of the acuter opponents of the theory saw at
once the justice of this position, and, as this consequence was
intolerable, they wanted to get rid of the whole theory.

The first scientific application of the Darwinian theory to man was
made by Huxley, the greatest zoologist in England. This able and
learned scientist, to whom zoology owes much of its progress,
published in 1863 a small work entitled Evidence as to Man's Place in
Nature. In the extremely important and interesting lectures which made
up this work he proved clearly that the descent of man from the ape
followed necessarily from the theory of descent. If that theory is
true, we are bound to conceive the animals which most closely resemble
man as those from which humanity has been gradually evolved. About the
same time Carl Vogt published a larger work on the same subject. We
must also mention Gustav Jaeger and Friedrich Rolle among the
zoologists who accepted and taught the theory of evolution immediately
after the publication of Darwin's book, and maintained that the
descent of man from the lower animals logically followed from it. The
latter published, in 1866, a work on the origin and position of man.

About the same time I attempted, in the second volume of my General
Morphology (1866), to apply the theory of evolution to the whole
organic kingdom, including man.* (* Huxley spoke of this "as one of
the greatest scientific works ever published."--Translator.) I
endeavoured to sketch the probable ancestral trees of the various
classes of the animal world, the protists, and the plants, as it
seemed necessary to do on Darwinian principles, and as we can actually
do now with a high degree of confidence. If the theory of descent,
which Lamarck first clearly formulated and Darwin thoroughly
established, is true, we should be able to draw up a natural
classification of plants and animals in the light of their genealogy,
and to conceive the large and small divisions of the system as the
branches and twigs of an ancestral tree. The eight genealogical tables
which I inserted in the second volume of the General Morphology are
the first sketches of their kind. In Chapter 2.27, particularly, I
trace the chief stages in man's ancestry, as far as it is possible to
follow it through the vertebrate stem. I tried especially to
determine, as well as one could at that time, the position of man in
the classification of the mammals and its genealogical significance. I
have greatly improved this attempt, and treated it in a more popular
form, in chapters 26 to 28 of my History of Creation (1868).* (* Of
which Darwin said that the Descent of Man would probably never have
been written if he had seen it earlier.--Translator.)

It was not until 1871, twelve years after the appearance of The Origin
of Species, that Darwin published the famous work which made the
much-contested application of his theory to man, and crowned the
splendid structure of his system. This important work was The Descent
of Man, and Selection in Relation to Sex. In this Darwin expressly
drew the conclusion, with rigorous logic, that man also must have been
developed out of lower species, and described the important part
played by sexual selection in the elevation of man and the other
higher animals. He showed that the careful selection which the sexes
exercise on each other in regard to sexual relations and procreation,
and the aesthetic feeling which the higher animals develop through
this, are of the utmost importance in the progressive development of
forms and the differentiation of the sexes. The males choosing the
handsomest females in one class of animals, and the females choosing
only the finest-looking males in another, the special features and the
sexual characteristics are increasingly accentuated. In fact, some of
the higher animals develop in this connection a finer taste and
judgment than man himself. But, even as regards man, it is to this
sexual selection that we owe the family-life, which is the chief
foundation of civilisation. The rise of the human race is due for the
most part to the advanced sexual selection which our ancestors
exercised in choosing their mates.

Darwin accepted in the main the general outlines of man's ancestral
tree, as I gave it in the General Morphology and the History of
Creation, and admitted that his studies led him to the same
conclusion. That he did not at once apply the theory to man in his
first work was a commendable piece of discretion; such a sequel was
bound to excite the strongest opposition to the whole theory. The
first thing to do was to establish it as regards the animal and plant
worlds. The subsequent extension to man was bound to be made sooner or
later.

It is important to understand this very clearly. If all living things
come from a common root, man must be included in the general scheme of
evolution. On the other hand, if the various species were separately
created, man, too, must have been created, and not evolved. We have to
choose between these two alternatives. This cannot be too frequently
or too strongly emphasised. EITHER all the species of animals and
plants are of supernatural origin--created, not evolved--and in that
case man also is the outcome of a creative act, as religion teaches,
OR the different species have been evolved from a few common, simple
ancestral forms, and in that case man is the highest fruit of the tree
of evolution.

We may state this briefly in the following principle--The descent of
man from the lower animals is a special deduction which inevitably
follows from the general inductive law of the whole theory of
evolution. In this principle we have a clear and plain statement of
the matter. Evolution is in reality nothing but a great induction,
which we are compelled to make by the comparative study of the most
important facts of morphology and physiology. But we must draw our
conclusion according to the laws of induction, and not attempt to
determine scientific truths by direct measurement and mathematical
calculation. In the study of living things we can scarcely ever
directly and fully, and with mathematical accuracy, determine the
nature of phenomena, as is done in the simpler study of the inorganic
world--in chemistry, physics, mineralogy, and astronomy. In the
latter, especially, we can always use the simplest and absolutely
safest method--that of mathematical determination. But in biology this
is quite impossible for various reasons; one very obvious reason being
that most of the facts of the science are very complicated and much
too intricate to allow a direct mathematical analysis. The greater
part of the phenomena that biology deals with are complicated
HISTORICAL PROCESSES, which are related to a far-reaching past, and as
a rule can only be approximately estimated. Hence we have to proceed
by INDUCTION--that is to say, to draw general conclusions, stage by
stage, and with proportionate confidence, from the accumulation of
detailed observations. These inductive conclusions cannot command
absolute confidence, like mathematical axioms; but they approach the
truth, and gain increasing probability, in proportion as we extend the
basis of observed facts on which we build. The importance of these
inductive laws is not diminished from the circumstance that they are
looked upon merely as temporary acquisitions of science, and may be
improved to any extent in the progress of scientific knowledge. The
same may be said of the attainments of many other sciences, such as
geology or archeology. However much they may be altered and improved
in detail in the course of time, these inductive truths may retain
their substance unchanged.

Now, when we say that the theory of evolution in the sense of Lamarck
and Darwin is an inductive law--in fact, the greatest of all
biological inductions--we rely, in the first place, on the facts of
paleontology. This science gives us some direct acquaintance with the
historical phenomena of the changes of species. From the situations in
which we find the fossils in the various strata of the earth we gather
confidently, in the first place, that the living population of the
earth has been gradually developed, as clearly as the earth's crust
itself; and that, in the second place, several different populations
have succeeded each other in the various geological periods. Modern
geology teaches that the formation of the earth has been gradual, and
unbroken by any violent revolutions. And when we compare together the
various kinds of animals and plants which succeed each other in the
history of our planet, we find, in the first place, a constant and
gradual increase in the number of species from the earliest times
until the present day; and, in the second place, we notice that the
forms in each great group of animals and plants also constantly
improve as the ages advance. Thus, of the vertebrates there are at
first only the lower fishes; then come the higher fishes, and later
the amphibia. Still later appear the three higher classes of
vertebrates--the reptiles, birds, and mammals, for the first time;
only the lowest and least perfect forms of the mammals are found at
first; and it is only at a very late period that placental mammals
appear, and man belongs to the latest and youngest branch of these.
Thus perfection of form increases as well as variety from the earliest
to the latest stage. That is a fact of the greatest importance. It can
only be explained by the theory of evolution, with which it is in
perfect harmony. If the different groups of plants and animals do
really descend from each other, we must expect to find this increase
in their number and perfection under the influence of natural
selection, just as the succession of fossils actually discloses it to
us.

Comparative anatomy furnishes a second series of facts which are of
great importance for the forming of our inductive law. This branch of
morphology compares the adult structures of living things, and seeks
in the great variety of organic forms the stable and simple law of
organisation, or the common type or structure. Since Cuvier founded
this science at the beginning of the nineteenth century it has been a
favourite study of the most distinguished scientists. Even before
Cuvier's time Goethe had been greatly stimulated by it, and induced to
take up the study of morphology. Comparative osteology, or the
philosophic study and comparison of the bony skeleton of the
vertebrates--one of its most interesting sections--especially
fascinated him, and led him to form the theory of the skull which I
mentioned before. Comparative anatomy shows that the internal
structure of the animals of each stem and the plants of each class is
the same in its essential features, however much they differ in
external appearance. Thus man has so great a resemblance in the chief
features of his internal organisation to the other mammals that no
comparative anatomist has ever doubted that he belongs to this class.
The whole internal structure of the human body, the arrangement of its
various systems of organs, the distribution of the bones, muscles,
blood-vessels, etc., and the whole structure of these organs in the
larger and the finer scale, agree so closely with those of the other
mammals (such as the apes, rodents, ungulates, cetacea, marsupials,
etc.) that their external differences are of no account whatever. We
learn further from comparative anatomy that the chief features of
animal structure are so similar in the various classes (fifty to sixty
in number altogether) that they may all be comprised in from eight to
twelve great groups. But even in these groups, the stem-forms or
animal types, certain organs (especially the alimentary canal) can be
proved to have been originally the same for all. We can only explain
by the theory of evolution this essential unity in internal structure
of all these animal forms that differ so much in outward appearance.
This wonderful fact can only be really understood and explained when
we regard the internal resemblance as an inheritance from common-stem
forms, and the external differences as the effect of adaptation to
different environments.

In recognising this, comparative anatomy has itself advanced to a
higher stage. Gegenbaur, the most distinguished of recent students of
this science, says that with the theory of evolution a new period
began in comparative anatomy, and that the theory in turn found a
touch stone in the science. "Up to now there is no fact in comparative
anatomy that is inconsistent with the theory of evolution; indeed,
they all lead to it. In this way the theory receives back from the
science all the service it rendered to its method." Until then
students had marvelled at the wonderful resemblance of living things
in their inner structure without being able to explain it. We are now
in a position to explain the causes of this, by showing that this
remarkable agreement is the necessary consequence of the inheriting of
common stem-forms; while the striking difference in outward appearance
is a result of adaptation to changes of environment. Heredity and
adaptation alone furnish the true explanation.

But one special part of comparative anatomy is of supreme interest and
of the utmost philosophic importance in this connection. This is the
science of rudimentary or useless organs; I have given it the name of
"dysteleology" in view of its philosophic consequences. Nearly every
organism (apart from the very lowest), and especially every
highly-developed animal or plant, including man, has one or more
organs which are of no use to the body itself, and have no share in
its functions or vital aims. Thus we all have, in various parts of our
frame, muscles which we never use, as, for instance, in the shell of
the ear and adjoining parts. In most of the mammals, especially those
with pointed ears, these internal and external ear-muscles are of
great service in altering the shell of the ear, so as to catch the
waves of sound as much as possible. But in the case of man and other
short-eared mammals these muscles are useless, though they are still
present. Our ancestors having long abandoned the use of them, we
cannot work them at all to-day. In the inner corner of the eye we have
a small crescent-shaped fold of skin; this is the last relic of a
third inner eye-lid, called the nictitating (winking) membrane. This
membrane is highly developed and of great service in some of our
distant relations, such as fishes of the shark type and several other
vertebrates; in us it is shrunken and useless. In the intestines we
have a process that is not only quite useless, but may be very
harmful--the vermiform appendage. This small intestinal appendage is
often the cause of a fatal illness. If a cherry-stone or other hard
body is unfortunately squeezed through its narrow aperture during
digestion, a violent inflammation is set up, and often proves fatal.
This appendix has no use whatever now in our frame; it is a dangerous
relic of an organ that was much larger and was of great service in our
vegetarian ancestors. It is still large and important in many
vegetarian animals, such as apes and rodents.

There are similar rudimentary organs in all parts of our body, and in
all the higher animals. They are among the most interesting phenomena
to which comparative anatomy introduces us; partly because they
furnish one of the clearest proofs of evolution, and partly because
they most strikingly refute the teleology of certain philosophers. The
theory of evolution enables us to give a very simple explanation of
these phenomena.

We have to look on them as organs which have fallen into disuse in the
course of many generations. With the decrease in the use of its
function, the organ itself shrivels up gradually, and finally
disappears. There is no other way of explaining rudimentary organs.
Hence they are also of great interest in philosophy; they show clearly
that the monistic or mechanical view of the organism is the only
correct one, and that the dualistic or teleological conception is
wrong. The ancient legend of the direct creation of man according to a
pre-conceived plan and the empty phrases about "design" in the
organism are completely shattered by them. It would be difficult to
conceive a more thorough refutation of teleology than is furnished by
the fact that all the higher animals have these rudimentary organs.

The theory of evolution finds its broadest inductive foundation in the
natural classification of living things, which arranges all the
various forms in larger and smaller groups, according to their degree
of affinity. These groupings or categories of classification--the
varieties, species, genera, families, orders, classes, etc.--show such
constant features of coordination and subordination that we are bound
to look on them as genealogical, and represent the whole system in the
form of a branching tree. This is the genealogical tree of the
variously related groups; their likeness in form is the expression of
a real affinity. As it is impossible to explain in any other way the
natural tree-like form of the system of organisms, we must regard it
at once as a weighty proof of the truth of evolution. The careful
construction of these genealogical trees is, therefore, not an
amusement, but the chief task of modern classification.

Among the chief phenomena that bear witness to the inductive law of
evolution we have the geographical distribution of the various species
of animals and plants over the surface of the earth, and their
topographical distribution on the summits of mountains and in the
depths of the ocean. The scientific study of these features--the
"science of distribution," or chorology (chora = a place)--has been
pursued with lively interest since the discoveries made by Alexander
von Humboldt. Until Darwin's time the work was confined to the
determination of the facts of the science, and chiefly aimed at
settling the spheres of distribution of the existing large and small
groups of living things. It was impossible at that time to explain the
causes of this remarkable distribution, or the reasons why one group
is found only in one locality and another in a different place, and
why there is this manifold distribution at all. Here, again, the
theory of evolution has given us the solution of the problem. It
furnishes the only possible explanation when it teaches that the
various species and groups of species descend from common stem-forms,
whose ever-branching offspring have gradually spread themselves by
migration over the earth. For each group of species we must admit a
"centre of production," or common home; this is the original habitat
in which the ancestral form was developed, and from which its
descendants spread out in every direction. Several of these
descendants became in their turn the stem-forms for new groups of
species, and these also scattered themselves by active and passive
migration, and so on. As each migrating organism found a different
environment in its new home, and adapted itself to it, it was
modified, and gave rise to new forms.

This very important branch of science that deals with active and
passive migration was founded by Darwin, with the aid of the theory of
evolution; and at the same time he advanced the true explanation of
the remarkable relation or similarity of the living population in any
locality to the fossil forms found in it. Moritz Wagner very ably
developed his idea under the title of "the theory of migration." In my
opinion, this famous traveller has rather over-estimated the value of
his theory of migration when he takes it to be an indispensable
condition of the formation of new species and opposes the theory of
selection. The two theories are not opposed in their main features.
Migration (by which the stem-form of a new species is isolated) is
really only a special case of selection. The striking and interesting
facts of chorology can be explained only by the theory of evolution,
and therefore we must count them among the most important of its
inductive bases.

The same must be said of all the remarkable phenomena which we
perceive in the economy of the living organism. The many and various
relations of plants and animals to each other and to their
environment, which are treated in bionomy (from nomos, law or norm,
and bios, life), the interesting facts of parasitism, domesticity,
care of the young, social habits, etc., can only be explained by the
action of heredity and adaptation. Formerly people saw only the
guidance of a beneficent Providence in these phenomena; to-day we
discover in them admirable proofs of the theory of evolution. It is
impossible to understand them except in the light of this theory and
the struggle for life.

Finally, we must, in my opinion, count among the chief inductive bases
of the theory of evolution the foetal development of the individual
organism, the whole science of embryology or ontogeny. But as the
later chapters will deal with this in detail, I need say nothing
further here. I shall endeavour in the following pages to show, step
by step, how the whole of the embryonic phenomena form a massive chain
of proof for the theory of evolution; for they can be explained in no
other way. In thus appealing to the close causal connection between
ontogenesis and phylogenesis, and taking our stand throughout on the
biogenetic law, we shall be able to prove, stage by stage, from the
facts of embryology, the evolution of man from the lower animals.

The general adoption of the theory of evolution has definitely closed
the controversy as to the nature or definition of the species. The
word has no ABSOLUTE meaning whatever, but is only a group-name, or
category of classification, with a purely relative value. In 1857, it
is true, a famous and gifted, but inaccurate and dogmatic, scientist,
Louis Agassiz, attempted to give an absolute value to these
"categories of classification." He did this in his Essay on
Classification, in which he turns upside down the phenomena of organic
nature, and, instead of tracing them to their natural causes, examines
them through a theological prism. The true species (bona species) was,
he said, an "incarnate idea of the Creator." Unfortunately, this
pretty phrase has no more scientific value than all the other attempts
to save the absolute or intrinsic value of the species.

The dogma of the fixity and creation of species lost its last great
champion when Agassiz died in 1873. The opposite theory, that all the
different species descend from common stem-forms, encounters no
serious difficulty to-day. All the endless research into the nature of
the species, and the possibility of several species descending from a
common ancestor, has been closed to-day by the removal of the sharp
limits that had been set up between species and varieties on the one
hand, and species and genera on the other. I gave an analytic proof of
this in my monograph on the sponges (1872), having made a very close
study of variability in this small but highly instructive group, and
shown the impossibility of making any dogmatic distinction of species.
According as the classifier takes his ideas of genus, species, and
variety in a broader or in a narrower sense, he will find in the small
group of the sponges either one genus with three species, or three
genera with 238 species, or 113 genera with 591 species. Moreover, all
these forms are so connected by intermediate forms that we can
convincingly prove the descent of all the sponges from a common
stem-form, the olynthus.

Here, I think, I have given an analytic solution of the problem of the
origin of species, and so met the demand of certain opponents of
evolution for an actual instance of descent from a stem-form. Those
who are not satisfied with the synthetic proofs of the theory of
evolution which are provided by comparative anatomy, embryology,
paleontology, dysteleology, chorology, and classification, may try to
refute the analytic proof given in my treatise on the sponge, the
outcome of five years of assiduous study. I repeat: It is now
impossible to oppose evolution on the ground that we have no
convincing example of the descent of all the species of a group from a
common ancestor. The monograph on the sponges furnishes such a proof,
and, in my opinion, an indisputable proof. Any man of science who will
follow the protracted steps of my inquiry and test my assertions will
find that in the case of the sponges we can follow the actual
evolution of species in a concrete case. And if this is so, if we can
show the origin of all the species from a common form in one single
class, we have the solution of the problem of man's origin, because we
are in a position to prove clearly his descent from the lower animals.

At the same time, we can now reply to the often-repeated assertion,
even heard from scientists of our own day, that the descent of man
from the lower animals, and proximately from the apes, still needs to
be "proved with certainty." These "certain proofs" have been available
for a long time; one has only to open one's eyes to see them. It is a
mistake to seek them in the discovery of intermediate forms between
man and the ape, or the conversion of an ape into a human being by
skilful education. The proofs lie in the great mass of empirical
material we have already collected. They are furnished in the
strongest form by the data of comparative anatomy and embryology,
completed by paleontology. It is not a question now of detecting new
proofs of the evolution of man, but of examining and understanding the
proofs we already have.

I was almost alone thirty-six years ago when I made the first attempt,
in my General Morphology, to put organic science on a mechanical
foundation through Darwin's theory of descent. The association of
ontogeny and phylogeny and the proof of the intimate causal connection
between these two sections of the science of evolution, which I
expounded in my work, met with the most spirited opposition on nearly
all sides. The next ten years were a terrible "struggle for life" for
the new theory. But for the last twenty-five years the tables have
been turned. The phylogenetic method has met with so general a
reception, and found so prolific a use in every branch of biology,
that it seems superfluous to treat any further here of its validity
and results. The proof of it lies in the whole morphological
literature of the last three decades. But no other science has been so
profoundly modified in its leading thoughts by this adoption, and been
forced to yield such far-reaching consequences, as that science which
I am now seeking to establish--monistic anthropogeny.

This statement may seem to be rather audacious, since the very next
branch of biology, anthropology in the stricter sense, makes very
little use of these results of anthropogeny, and sometimes expressly
opposes them.* (*This does not apply to English anthropologists, who
are almost all evolutionists.) This applies especially to the attitude
which has characterised the German Anthropological Society (the
Deutsche Gesellschaft fur Anthropologie) for some thirty years. Its
powerful president, the famous pathologist, Rudolph Virchow, is
chiefly responsible for this. Until his death (September 5th, 1902) he
never ceased to reject the theory of descent as unproven, and to
ridicule its chief consequence--the descent of man from a series of
mammal ancestors--as a fantastic dream. I need only recall his
well-known expression at the Anthropological Congress at Vienna in
1894, that "it would be just as well to say man came from the sheep or
the elephant as from the ape."

Virchow's assistant, the secretary of the German Anthropological
Society, Professor Johannes Ranke of Munich, has also indefatigably
opposed transformism: he has succeeded in writing a work in two
volumes (Der Mensch), in which all the facts relating to his
organisation are explained in a sense hostile to evolution. This work
has had a wide circulation, owing to its admirable illustrations and
its able treatment of the most interesting facts of anatomy and
physiology--exclusive of the sexual organs! But, as it has done a
great deal to spread erroneous views among the general public, I have
included a criticism of it in my History of Creation, as well as met
Virchow's attacks on anthropogeny.

Neither Virchow, nor Ranke, nor any other "exact" anthropologist, has
attempted to give any other natural explanation of the origin of man.
They have either set completely aside this "question of questions" as
a transcendental problem, or they have appealed to religion for its
solution. We have to show that this rejection of the rational
explanation is totally without justification. The fund of knowledge
which has accumulated in the progress of biology in the nineteenth
century is quite adequate to furnish a rational explanation, and to
establish the theory of the evolution of man on the solid facts of his
embryology.


CHAPTER 1.6. THE OVUM AND THE AMOEBA.

In order to understand clearly the course of human embryology, we must
select the more important of its wonderful and manifold processes for
fuller explanation, and then proceed from these to the innumerable
features of less importance. The most important feature in this sense,
and the best starting-point for ontogenetic study, is the fact that
man is developed from an ovum, and that this ovum is a simple cell.
The human ovum does not materially differ in form and composition from
that of the other mammals, whereas there is a distinct difference
between the fertilised ovum of the mammal and that of any other
animal.

(FIGURE 1.1. The human ovum, magnified 100 times. The globular mass of
yelk (b) is enclosed by a transparent membrane (the ovolemma or zona
pellucida [a]), and contains a noncentral nucleus (the germinal
vesicle, c). Cf. Figure 1.14.)

This fact is so important that few should be unaware of its extreme
significance; yet it was quite unknown in the first quarter of the
nineteenth century. As we have seen, the human and mammal ovum was not
discovered until 1827, when Carl Ernst von Baer detected it. Up to
that time the larger vesicles, in which the real and much smaller ovum
is contained, had been wrongly regarded as ova. The important
circumstance that this mammal ovum is a simple cell, like the ovum of
other animals, could not, of course, be recognised until the cell
theory was established. This was not done, by Schleiden for the plant
and Schwann for the animal, until 1838. As we have seen, this cell
theory is of the greatest service in explaining the human frame and
its embryonic development. Hence we must say a few words about the
actual condition of the theory and the significance of the views it
has suggested.

In order properly to appreciate the cellular theory, the most
important element in our science, it is necessary to understand in the
first place that the cell is a UNIFIED ORGANISM, a self-contained
living being. When we anatomically dissect the fully-formed animal or
plant into its various organs, and then examine the finer structure of
these organs with the microscope, we are surprised to find that all
these different parts are ultimately made up of the same structural
element or unit. This common unit of structure is the cell. It does
not matter whether we thus dissect a leaf, flower, or fruit, or a
bone, muscle, gland, or bit of skin, etc.; we find in every case the
same ultimate constituent, which has been called the cell since
Schleiden's discovery. There are many opinions as to its real nature,
but the essential point in our view of the cell is to look upon it as
a self-contained or independent living unit. It is, in the words of
Brucke, "an elementary organism." We may define it most precisely as
the ultimate organic unit, and, as the cells are the sole active
principles in every vital function, we may call them the "plastids,"
or "formative elements." This unity is found in both the anatomic
structure and the physiological function. In the case of the protists,
the entire organism usually consists of a single independent cell
throughout life. But in the tissue-forming animals and plants, which
are the great majority, the organism begins its career as a simple
cell, and then grows into a cell-community, or, more correctly, an
organised cell-state. Our own body is not really the simple unity that
it is generally supposed to be. On the contrary, it is a very
elaborate social system of countless microscopic organisms, a colony
or commonwealth, made up of innumerable independent units, or very
different tissue-cells.

In reality, the term "cell," which existed long before the cell theory
was formulated, is not happily chosen. Schleiden, who first brought it
into scientific use in the sense of the cell theory, gave this name to
the elementary organisms because, when you find them in the dissected
plant, they generally have the appearance of chambers, like the cells
in a bee-hive, with firm walls and a fluid or pulpy content. But some
cells, especially young ones, are entirely without the enveloping
membrane, or stiff wall. Hence we now generally describe the cell as a
living, viscous particle of protoplasm, enclosing a firmer nucleus in
its albuminoid body. There may be an enclosing membrane, as there
actually is in the case of most of the plants; but it may be wholly
lacking, as is the case with most of the animals. There is no membrane
at all in the first stage. The young cells are usually round, but they
vary much in shape later on. Illustrations of this will be found in
the cells of the various parts of the body shown in Figures 1.3 to
1.7.

Hence the essential point in the modern idea of the cell is that it is
made up of two different active constituents--an inner and an outer
part. The smaller and inner part is the nucleus (or caryon or
cytoblastus, Figure 1.1 c and Figure 1.2 k). The outer and larger
part, which encloses the other, is the body of the cell (celleus,
cytos, or cytosoma). The soft living substance of which the two are
composed has a peculiar chemical composition, and belongs to the group
of the albuminoid plasma-substances ("formative matter"), or
protoplasm. The essential and indispensable element of the nucleus is
called nuclein (or caryoplasm); that of the cell body is called
plastin (or cytoplasm). In the most rudimentary cases both substances
seem to be quite simple and homogeneous, without any visible
structure. But, as a rule, when we examine them under a high power of
the microscope, we find a certain structure in the protoplasm. The
chief and most common form of this is the fibrous or net-like "thready
structure" (Frommann) and the frothy "honeycomb structure" (Butschli).

(FIGURE 1.2. Stem-cell of one of the echinoderms (cytula, or "first
segmentation-cell" = fertilised ovum), after Hertwig. k is the nucleus
or caryon.)

The shape or outer form of the cell is infinitely varied, in
accordance with its endless power of adapting itself to the most
diverse activities or environments. In its simplest form the cell is
globular (Figure 1.2). This normal round form is especially found in
cells of the simplest construction, and those that are developed in a
free fluid without any external pressure. In such cases the nucleus
also is not infrequently round, and located in the centre of the
cell-body (Figure 1.2 k). In other cases, the cells have no definite
shape; they are constantly changing their form owing to their
automatic movements. This is the case with the amoebae (Figures 1.15
and 1.16) and the amoeboid travelling cells (Figure 1.11), and also
with very young ova (Figure 1.13). However, as a rule, the cell
assumes a definite form in the course of its career. In the tissues of
the multicellular organism, in which a number of similar cells are
bound together in virtue of certain laws of heredity, the shape is
determined partly by the form of their connection and partly by their
special functions. Thus, for instance, we find in the mucous lining of
our tongue very thin and delicate flat cells of roundish shape (Figure
1.3). In the outer skin we find similar, but harder, covering cells,
joined together by saw-like edges (Figure 1.4). In the liver and other
glands there are thicker and softer cells, linked together in rows
(Figure 1.5).

The last-named tissues (Figures 1.3 to 1.5) belong to the simplest and
most primitive type, the group of the "covering-tissues," or
epithelia. In these "primary tissues" (to which the germinal layers
belong) simple cells of the same kind are arranged in layers. The
arrangement and shape are more complicated in the "secondary tissues,"
which are gradually developed out of the primary, as in the tissues of
the muscles, nerves, bones, etc. In the bones, for instance, which
belong to the group of supporting or connecting organs, the cells
(Figure 1.6) are star-shaped, and are joined together by numbers of
net-like interlacing processes; so, also, in the tissues of the teeth
(Figure 1.7), and in other forms of supporting-tissue, in which a soft
or hard substance (intercellular matter, or base) is inserted between
the cells.

(FIGURE 1.3. Three epithelial cells from the mucous lining of the
tongue.

FIGURE 1.4. Five spiny or grooved cells, with edges joined, from the
outer skin (epidermis): one of them (b) is isolated.

FIGURE 1.5. Ten liver-cells: one of them (b) has two nuclei.)

The cells also differ very much in size. The great majority of them
are invisible to the naked eye, and can be seen only through the
microscope (being as a rule between 1/2500 and 1/250 inch in
diameter). There are many of the smaller plastids--such as the famous
bacteria--which only come into view with a very high magnifying power.
On the other hand, many cells attain a considerable size, and run
occasionally to several inches in diameter, as do certain kinds of
rhizopods among the unicellular protists (such as the radiolaria and
thalamophora). Among the tissue-cells of the animal body many of the
muscular fibres and nerve fibres are more than four inches, and
sometimes more than a yard, in length. Among the largest cells are the
yelk-filled ova; as, for instance, the yellow "yolk" in the hen's egg,
which we shall describe later (Figure 1.15).

Cells also vary considerably in structure. In this connection we must
first distinguish between the active and passive components of the
cell. It is only the former, or active parts of the cell, that really
live, and effect that marvellous world of phenomena to which we give
the name of "organic life." The first of these is the inner nucleus
(caryoplasm), and the second the body of the cell (cytoplasm). The
passive portions come third; these are subsequently formed from the
others, and I have given them the name of "plasma-products." They are
partly external (cell-membranes and intercellular matter) and partly
internal (cell-sap and cell-contents).

The nucleus (or caryon), which is usually of a simple roundish form,
is quite structureless at first (especially in very young cells), and
composed of homogeneous nuclear matter or caryoplasm (Figure 1.2 k).
But, as a rule, it forms a sort of vesicle later on, in which we can
distinguish a more solid nuclear base (caryobasis) and a softer or
fluid nuclear sap (caryolymph). In a mesh of the nuclear network (or
it may be on the inner side of the nuclear envelope) there is, as a
rule, a dark, very opaque, solid body, called the nucleolus. Many of
the nuclei contain several of these nucleoli (as, for instance, the
germinal vesicle of the ova of fishes and amphibia). Recently a very
small, but particularly important, part of the nucleus has been
distinguished as the central body (centrosoma)--a tiny particle that
is originally found in the nucleus itself, but is usually outside it,
in the cytoplasm; as a rule, fine threads stream out from it in the
cytoplasm. From the position of the central body with regard to the
other parts it seems probable that it has a high physiological
importance as a centre of movement; but it is lacking in many cells.

The cell-body also consists originally, and in its simplest form, of a
homogeneous viscid plasmic matter. But, as a rule, only the smaller
part of it is formed of the living active cell-substance (protoplasm);
the greater part consists of dead, passive plasma-products
(metaplasm). It is useful to distinguish between the inner and outer
of these. External plasma-products (which are thrust out from the
protoplasm as solid "structural matter") are the cell-membranes and
the intercellular matter. The internal plasma-products are either the
fluid cell-sap or hard structures. As a rule, in mature and
differentiated cells these various parts are so arranged that the
protoplasm (like the caryoplasm in the round nucleus) forms a sort of
skeleton or framework. The spaces of this network are filled partly
with the fluid cell-sap and partly by hard structural products.

(FIGURE 1.6. Nine star-shaped bone-cells, with interlaced branches.

FIGURE 1.7. Eleven star-shaped cells from the enamel of a tooth,
joined together by their branchlets.)

The simple round ovum, which we take as the starting-point of our
study (Figures 1.1 and 1.2), has in many cases the vague, indifferent
features of the typical primitive cell. As a contrast to it, and as an
instance of a very highly differentiated plastid, we may consider for
a moment a large nerve-cell, or ganglionic cell, from the brain. The
ovum stands potentially for the entire organism--in other words, it
has the faculty of building up out of itself the whole multicellular
body. It is the common parent of all the countless generations of
cells which form the different tissues of the body; it unites all
their powers in itself, though only potentially or in germ. In
complete contrast to this, the neural cell in the brain (Figure 1.9)
develops along one rigid line. It cannot, like the ovum, beget endless
generations of cells, of which some will become skin-cells, others
muscle-cells, and others again bone-cells. But, on the other hand, the
nerve-cell has become fitted to discharge the highest functions of
life; it has the powers of sensation, will, and thought. It is a real
soul-cell, or an elementary organ of the psychic activity. It has,
therefore, a most elaborate and delicate structure. Numbers of
extremely fine threads, like the electric wires at a large telegraphic
centre, cross and recross in the delicate protoplasm of the nerve
cell, and pass out in the branching processes which proceed from it
and put it in communication with other nerve-cells or nerve-fibres (a,
b). We can only partly follow their intricate paths in the fine matter
of the body of the cell.

Here we have a most elaborate apparatus, the delicate structure of
which we are just beginning to appreciate through our most powerful
microscopes, but whose significance is rather a matter of conjecture
than knowledge. Its intricate structure corresponds to the very
complicated functions of the mind. Nevertheless, this elementary organ
of psychic activity--of which there are thousands in our brain--is
nothing but a single cell. Our whole mental life is only the joint
result of the combined activity of all these nerve-cells, or
soul-cells. In the centre of each cell there is a large transparent
nucleus, containing a small and dark nuclear body. Here, as elsewhere,
it is the nucleus that determines the individuality of the cell; it
proves that the whole structure, in spite of its intricate
composition, amounts to only a single cell.

(FIGURE 1.8. Unfertilised ovum of an echinoderm (from Hertwig). The
vesicular nucleus (or "germinal vesicle") is globular, half the size
of the round ovum, and encloses a nuclear framework, in the central
knot of which there is a dark nucleolus (the "germinal spot").

FIGURE 1.9. A large branching nerve-cell, or "soul-cell," from the
brain of an electric fish (Torpedo), magnified 600 times. In the
middle of the cell is the large transparent round nucleus, one
nucleolus, and, within the latter again, a nucleolinus. The protoplasm
of the cell is split into innumerable fine threads (or fibrils), which
are embedded in intercellular matter, and are prolonged into the
branching processes of the cell (b). One branch (a) passes into a
nerve-fibre. (From Max Schultze.))

In contrast with this very elaborate and very strictly differentiated
psychic cell (Figure 1.9), we have our ovum (Figures 1.1 and 1.2),
which has hardly any structure at all. But even in the case of the
ovum we must infer from its properties that its protoplasmic body has
a very complicated chemical composition and a fine molecular structure
which escapes our observation. This presumed molecular structure of
the plasm is now generally admitted; but it has never been seen, and,
indeed, lies far beyond the range of microscopic vision. It must not
be confused--as is often done--with the structure of the plasm (the
fibrous network, groups of granules, honey-comb, etc.) which does come
within the range of the microscope.

But when we speak of the cells as the elementary organisms, or
structural units, or "ultimate individualities," we must bear in mind
a certain restriction of the phrases. I mean, that the cells are not,
as is often supposed, the very lowest stage of organic individuality.
There are yet more elementary organisms to which I must refer
occasionally. These are what we call the "cytodes" (cytos = cell),
certain living, independent beings, consisting only of a particle of
plasson--an albuminoid substance, which is not yet differentiated into
caryoplasm and cytoplasm, but combines the properties of both. Those
remarkable beings called the monera--especially the chromacea and
bacteria--are specimens of these simple cytodes. (Compare Chapter
2.19.) To be quite accurate, then, we must say: the elementary
organism, or the ultimate individual, is found in two different
stages. The first and lower stage is the cytode, which consists merely
of a particle of plasson, or quite simple plasm. The second and higher
stage is the cell, which is already divided or differentiated into
nuclear matter and cellular matter. We comprise both kinds--the
cytodes and the cells--under the name of plastids ("formative
particles"), because they are the real builders of the organism.
However, these cytodes are not found, as a rule, in the higher animals
and plants; here we have only real cells with a nucleus. Hence, in
these tissue-forming organisms (both plant and animal) the organic
unit always consists of two chemically and anatomically different
parts--the outer cell-body and the inner nucleus.

In order to convince oneself that this cell is really an independent
organism, we have only to observe the development and vital phenomena
of one of them. We see then that it performs all the essential
functions of life--both vegetal and animal--which we find in the
entire organism. Each of these tiny beings grows and nourishes itself
independently. It takes its food from the surrounding fluid;
sometimes, even, the naked cells take in solid particles at certain
points of their surface--in other words, "eat" them--without needing
any special mouth and stomach for the purpose (cf. Figure 1.19).

Further, each cell is able to reproduce itself. This multiplication,
in most cases, takes the form of a simple cleavage, sometimes direct,
sometimes indirect; the simple direct (or "amitotic") division is less
common, and is found, for instance, in the blood cells (Figure 1.10).
In these the nucleus first divides into two equal parts by
constriction. The indirect (or "mitotic") cleavage is much more
frequent; in this the caryoplasm of the nucleus and the cytoplasm of
the cell-body act upon each other in a peculiar way, with a partial
dissolution (caryolysis), the formation of knots and loops (mitosis),
and a movement of the halved plasma-particles towards two mutually
repulsive poles of attraction (caryokinesis, Figure 1.11.)

(FIGURE 1.10. Blood-cells, multiplying by direct division, from the
blood of the embryo of a stag. Originally, each blood-cell has a
nucleus and is round (a). When it is going to multiply, the nucleus
divides into two (b, c, d). Then the protoplasmic body is constricted
between the two nuclei, and these move away from each other (e).
Finally, the constriction is complete, and the cell splits into two
daughter-cells (f). (From Frey.))

FIGURE 1.11. Indirect or mitotic cell-division (with caryolysis and
caryokinesis) from the skin of the larva of a salamander. (From
Rabl.).
   A. Mother-cell (Knot, spirema), with Nuclear threads (chromosomata)
      (coloured nuclear matter, chromatin), Cytosoma, Nuclear membrane,
      Protoplasm of the cell-body and Nuclear sap.
B. Mother-star, the loops beginning to split lengthways (nuclear
membrane gone), with Star-like appearance in cytoplasm, Centrosoma
(sphere of attraction), Nuclear spindle (achromin, colourless matter)
and Nuclear loops (chromatin, coloured matter).
C. The two daughter-stars, produced by the breaking of the loops of
the mother-star (moving away), with Upper daughter-crown, Connecting
threads of the two crowns (achromin), Lower daughter-crown and
Double-star (amphiaster).
D. The two daughter-cells, produced by the complete division of the
two nuclear halves (cytosomata still connected at the equator)
(Double-knot, Dispirema), with Upper daughter-nucleus, Equatorial
constriction of the cell-body and Lower daughter-nucleus.)

The intricate physiological processes which accompany this "mitosis"
have been very closely studied of late years. The inquiry has led to
the detection of certain laws of evolution which are of extreme
importance in connection with heredity. As a rule, two very different
parts of the nucleus play an important part in these changes. They
are: the chromatin, or coloured nuclear substance, which has a
peculiar property of tingeing itself deeply with certain colouring
matters (carmine, haematoxylin, etc.), and the achromin (or linin, or
achromatin), a colourless nuclear substance that lacks this property.
The latter generally forms in the dividing cell a sort of spindle, at
the poles of which there is a very small particle, also colourless,
called the "central body" (centrosoma). This acts as the centre or
focus in a "sphere of attraction" for the granules of protoplasm in
the surrounding cell-body, and assumes a star-like appearance (the
cell-star, or monaster). The two central bodies, standing opposed to
each other at the poles of the nuclear spindle, form "the double-star"
(or amphiaster, Figure 1.11, BC). The chromatin often forms a long,
irregularly-wound thread--"the coil" (spirema, Figure A). At the
commencement of the cleavage it gathers at the equator of the cell,
between the stellar poles, and forms a crown of U-shaped loops
(generally four or eight, or some other definite number). The loops
split lengthwise into two halves (B), and these back away from each
other towards the poles of the spindle (C). Here each group forms a
crown once more, and this, with the corresponding half of the divided
spindle, forms a fresh nucleus (D). Then the protoplasm of the
cell-body begins to contract in the middle, and gather about the new
daughter-nuclei, and at last the two daughter-cells become independent
beings.

Between this common mitosis, or indirect cell-division--which is the
normal cleavage-process in most cells of the higher animals and
plants--and the simple direct division (Figure 1.10) we find every
grade of segmentation; in some circumstances even one kind of division
may be converted into another.

The plastid is also endowed with the functions of movement and
sensation. The single cell can move and creep about, when it has space
for free movement and is not prevented by a hard envelope; it then
thrusts out at its surface processes like fingers, and quickly
withdraws them again, and thus changes its shape (Figure 1.12).
Finally, the young cell is sensitive, or more or less responsive to
stimuli; it makes certain movements on the application of chemical and
mechanical irritation. Hence we can ascribe to the individual cell all
the chief functions which we comprehend under the general heading of
"life"--sensation, movement, nutrition, and reproduction. All these
properties of the multicellular and highly developed animal are also
found in the single animal-cell, at least in its younger stages. There
is no longer any doubt about this, and so we may regard it as a solid
and important base of our physiological conception of the elementary
organism.

Without going any further here into these very interesting phenomena
of the life of the cell, we will pass on to consider the application
of the cell theory to the ovum. Here comparative research yields the
important result that EVERY OVUM IS AT FIRST A SIMPLE CELL. I say this
is very important, because our whole science of embryology now
resolves itself into the problem: "How does the multicellular organism
arise from the unicellular?" Every organic individual is at first a
simple cell, and as such an elementary organism, or a unit of
individuality. This cell produces a cluster of cells by segmentation,
and from these develops the multicellular organism, or individual of
higher rank.

When we examine a little closer the original features of the ovum, we
notice the extremely significant fact that in its first stage the ovum
is just the same simple and indefinite structure in the case of man
and all the animals (Figure 1.13). We are unable to detect any
material difference between them, either in outer shape or internal
constitution. Later, though the ova remain unicellular, they differ in
size and shape, enclose various kinds of yelk-particles, have
different envelopes, and so on. But when we examine them at their
birth, in the ovary of the female animal, we find them to be always of
the same form in the first stages of their life. In the beginning each
ovum is a very simple, roundish, naked, mobile cell, without a
membrane; it consists merely of a particle of cytoplasm enclosing a
nucleus (Figure 1.13). Special names have been given to these parts of
the ovum; the cell-body is called the yelk (vitellus), and the
cell-nucleus the germinal vesicle. As a rule, the nucleus of the ovum
is soft, and looks like a small pimple or vesicle. Inside it, as in
many other cells, there is a nuclear skeleton or frame and a third,
hard nuclear body (the nucleolus). In the ovum this is called the
germinal spot. Finally, we find in many ova (but not in all) a still
further point within the germinal spot, a "nucleolin," which goes by
the name of the germinal point. The latter parts (germinal spot and
germinal point) have, apparently, a minor importance, in comparison
with the other two (the yelk and germinal vesicle). In the yelk we
must distinguish the active formative yelk (or protoplasm = first
plasm) from the passive nutritive yelk (or deutoplasm = second plasm).

(FIGURE 1.12. Mobile cells from the inflamed eye of a frog (from the
watery fluid of the eye, the humor aqueus). The naked cells creep
freely about, by (like the amoeba or rhizopods) protruding fine
processes from the uncovered protoplasmic body. These bodies vary
continually in number, shape, and size. The nucleus of these amoeboid
lymph-cells ("travelling cells," or planocytes) is invisible, because
concealed by the numbers of fine granules which are scattered in the
protoplasm. (From Frey.))

In many of the lower animals (such as sponges, polyps, and medusae)
the naked ova retain their original simple appearance until
impregnation. But in most animals they at once begin to change; the
change consists partly in the formation of connections with the yelk,
which serve to nourish the ovum, and partly of external membranes for
their protection (the ovolemma, or prochorion). A membrane of this
sort is formed in all the mammals in the course of the embryonic
process. The little globule is surrounded by a thick capsule of
glass-like transparency, the zona pellucida, or ovolemma pellucidum
(Figure 1.14). When we examine it closely under the microscope, we see
very fine radial streaks in it, piercing the zona, which are really
very narrow canals. The human ovum, whether fertilised or not, cannot
be distinguished from that of most of the other mammals. It is nearly
the same everywhere in form, size, and composition. When it is fully
formed, it has a diameter of (on an average) about 1/120 of an inch.
When the mammal ovum has been carefully isolated, and held against the
light on a glass-plate, it may be seen as a fine point even with the
naked eye. The ova of most of the higher mammals are about the same
size. The diameter of the ovum is almost always between 1/250 to 1/125
inch. It has always the same globular shape; the same characteristic
membrane; the same transparent germinal vesicle with its dark germinal
spot. Even when we use the most powerful microscope with its highest
power, we can detect no material difference between the ova of man,
the ape, the dog, and so on. I do not mean to say that there are no
differences between the ova of these different mammals. On the
contrary, we are bound to assume that there are such, at least as
regards chemical composition. Even the ova of different men must
differ from each other; otherwise we should not have a different
individual from each ovum. It is true that our crude and imperfect
apparatus cannot detect these subtle individual differences, which are
probably in the molecular structure. However, such a striking
resemblance of their ova in form, so great as to seem to be a complete
similarity, is a strong proof of the common parentage of man and the
other mammals. From the common germ-form we infer a common stem-form.
On the other hand, there are striking peculiarities by which we can
easily distinguish the fertilised ovum of the mammal from the
fertilised ovum of the birds, amphibia, fishes, and other vertebrates
(see the close of Chapter 2.29).

(FIGURE 1.13. Ova of various animals, executing amoeboid movements,
highly magnified. All the ova are naked cells of varying shape. In the
dark fine-grained protoplasm (yelk) is a large vesicular nucleus (the
germinal vesicle), and in this is seen a nuclear body (the germinal
spot), in which again we often see a germinal point. Figures A1 to A4
represent the ovum of a sponge (Leuculmis echinus) in four successive
movements. B1 to B8 are the ovum of a parasitic crab (Chondracanthus
cornutus), in eight successive movements. (From Edward von Beneden.)
C1 to C5 show the ovum of the cat in various stages of movement (from
Pfluger); Figure P the ovum of a trout; E the ovum of a chicken; F a
human ovum.)

The fertilised bird-ovum (Figure 1.15) is notably different. It is
true that in its earliest stage (Figure 1.13 E) this ovum also is very
like that of the mammal (Figure 1.13 F). But afterwards, while still
within the oviduct, it takes up a quantity of nourishment and works
this into the familiar large yellow yelk. When we examine a very young
ovum in the hen's oviduct, we find it to be a simple, small, naked,
amoeboid cell, just like the young ova of other animals (Figure 1.13).
But it then grows to the size we are familiar with in the round yelk
of the egg. The nucleus of the ovum, or the germinal vesicle, is thus
pressed right to the surface of the globular ovum, and is embedded
there in a small quantity of transparent matter, the so-called white
yelk. This forms a round white spot, which is known as the "tread"
(cicatricula) (Figure 1.15 b). From the tread a thin column of the
white yelk penetrates through the yellow yelk to the centre of the
globular cell, where it swells into a small, central globule (wrongly
called the yelk-cavity, or latebra, Figure 1.15 d apostrophe). The
yellow yelk-matter which surrounds this white yelk has the appearance
in the egg (when boiled hard) of concentric layers (c). The yellow
yelk is also enclosed in a delicate structureless membrane (the
membrana vitellina, a).

As the large yellow ovum of the bird attains a diameter of several
inches in the bigger birds, and encloses round yelk-particles, there
was formerly a reluctance to consider it as a simple cell. This was a
mistake. Every animal that has only one cell-nucleus, every amoeba,
every gregarina, every infusorium, is unicellular, and remains
unicellular whatever variety of matter it feeds on. So the ovum
remains a simple cell, however much yellow yelk it afterwards
accumulates within its protoplasm. It is, of course, different, with
the bird's egg when it has been fertilised. The ovum then consists of
as many cells as there are nuclei in the tread. Hence, in the
fertilised egg which we eat daily, the yellow yelk is already a
multicellular body. Its tread is composed of several cells, and is now
commonly called the germinal disc. We shall return to this
discogastrula in Chapter 1.9.

(FIGURE 1.14. The human ovum, taken from the female ovary, magnified
500 times. The whole ovum is a simple round cell. The chief part of
the globular mass is formed by the nuclear yelk (deutoplasm), which is
evenly distributed in the active protoplasm, and consists of numbers
of fine yelk-granules. In the upper part of the yelk is the
transparent round germinal vesicle, which corresponds to the nucleus.
This encloses a darker granule, the germinal spot, which shows a
nucleolus. The globular yelk is surrounded by the thick transparent
germinal membrane (ovolemma, or zona pellucida). This is traversed by
numbers of lines as fine as hairs, which are directed radially towards
the centre of the ovum. These are called the pore-canals; it is
through these that the moving spermatozoa penetrate into the yelk at
impregnation.

FIGURE 1.15. A fertilised ovum from the oviduct of a hen. the yellow
yelk (c) consists of several concentric layers (d), and is enclosed in
a thin yelk-membrane (a). The nucleus or germinal vesicle is seen
above in the cicatrix or "tread" (b). From that point the white yelk
penetrates to the central yelk-cavity (d apostrophe). The two kinds of
yelk do not differ very much.

FIGURE 1.16. A creeping amoeba (highly magnified). The whole organism
is a simple naked cell, and moves about by means of the changing arms
which it thrusts out of and withdraws into its protoplasmic body.
Inside it is the roundish nucleus with its nucleolus.)

When the mature bird-ovum has left the ovary and been fertilised in
the oviduct, it covers itself with various membranes which are
secreted from the wall of the oviduct. First, the large clear
albuminous layer is deposited around the yellow yelk; afterwards, the
hard external shell, with a fine inner skin. All these gradually
forming envelopes and processes are of no importance in the formation
of the embryo; they serve merely for the protection of the original
simple ovum. We sometimes find extraordinarily large eggs with strong
envelopes in the case of other animals, such as fishes of the shark
type. Here, also, the ovum is originally of the same character as it
is in the mammal; it is a perfectly simple and naked cell. But, as in
the case of the bird, a considerable quantity of nutritive yelk is
accumulated inside the original yelk as food for the developing
embryo; and various coverings are formed round the egg. The ovum of
many other animals has the same internal and external features. They
have, however, only a physiological, not a morphological, importance;
they have no direct influence on the formation of the foetus. They are
partly consumed as food by the embryo, and partly serve as protective
envelopes. Hence we may leave them out of consideration altogether
here, and restrict ourselves to material points--TO THE SUBSTANTIAL
IDENTITY OF THE ORIGINAL OVUM IN MAN AND THE REST OF THE ANIMALS
(Figure 1.13).

Now, let us for the first time make use of our biogenetic law; and
directly apply this fundamental law of evolution to the human ovum. We
reach a very simple, but very important, conclusion. FROM THE FACT
THAT THE HUMAN OVUM AND THAT OF ALL OTHER ANIMALS CONSISTS OF A SINGLE
CELL, IT FOLLOWS IMMEDIATELY, ACCORDING TO THE BIOGENETIC LAW, THAT
ALL THE ANIMALS, INCLUDING MAN, DESCEND FROM A UNICELLULAR ORGANISM.
If our biogenetic law is true, if the embryonic development is a
summary or condensed recapitulation of the stem-history--and there can
be no doubt about it--we are bound to conclude, from the fact that all
the ova are at first simple cells, that all the multicellular
organisms originally sprang from a unicellular being. And as the
original ovum in man and all the other animals has the same simple and
indefinite appearance, we may assume with some probability that this
unicellular stem-form was the common ancestor of the whole animal
world, including man. However, this last hypothesis does not seem to
me as inevitable and as absolutely certain as our first conclusion.

This inference from the unicellular embryonic form to the unicellular
ancestor is so simple, but so important, that we cannot sufficiently
emphasise it. We must, therefore, turn next to the question whether
there are to-day any unicellular organisms, from the features of which
we may draw some approximate conclusion as to the unicellular
ancestors of the multicellular organisms. The answer is: Most
certainly there are. There are assuredly still unicellular organisms
which are, in their whole nature, really nothing more than permanent
ova. There are independent unicellular organisms of the simplest
character which develop no further, but reproduce themselves as such,
without any further growth. We know to-day of a great number of these
little beings, such as the gregarinae, flagellata, acineta, infusoria,
etc. However, there is one of them that has an especial interest for
us, because it at once suggests itself when we raise our question, and
it must be regarded as the unicellular being that approaches nearest
to the real ancestral form. This organism is the amoeba.

For a long time now we have comprised under the general name of
amoebae a number of microscopic unicellular organisms, which are very
widely distributed, especially in fresh-water, but also in the ocean;
in fact, they have lately been discovered in damp soil. There are also
parasitic amoebae which live inside other animals. When we place one
of these amoebae in a drop of water under the microscope and examine
it with a high power, it generally appears as a roundish particle of a
very irregular and varying shape (Figures 1.16 and 1.17). In its soft,
slimy, semi-fluid substance, which consists of protoplasm, we see only
the solid globular particle it contains, the nucleus. This unicellular
body moves about continually, creeping in every direction on the glass
on which we are examining it. The movement is effected by the
shapeless body thrusting out finger-like processes at various parts of
its surface; and these are slowly but continually changing, and
drawing the rest of the body after them. After a time, perhaps, the
action changes. The amoeba suddenly stands still, withdraws its
projections, and assumes a globular shape. In a little while, however,
the round body begins to expand again, thrusts out arms in another
direction, and moves on once more. These changeable processes are
called "false feet," or pseudopodia, because they act physiologically
as feet, yet are not special organs in the anatomic sense. They
disappear as quickly as they come, and are nothing more than temporary
projections of the semi-fluid and structureless body.

(FIGURE 1.17. Division of a unicellular amoeba (Amoeba polypodia) in
six stages. (From F.E. Schultze.) the dark spot is the nucleus, the
lighter spot a contractile vacuole in the protoplasm. The latter
reforms in one of the daughter-cells.)

FIGURE 1.18. Ovum of a sponge (Olynthus). The ovum creeps about in a
body of the sponge by thrusting out ever-changing processes. It is
indistinguishable from the common amoeba.)

If you touch one of these creeping amoebae with a needle, or put a
drop of acid in the water, the whole body at once contracts in
consequence of this mechanical or physical stimulus. As a rule, the
body then resumes its globular shape. In certain circumstances--for
instance, if the impurity of the water lasts some time--the amoeba
begins to develop a covering. It exudes a membrane or capsule, which
immediately hardens, and assumes the appearance of a round cell with a
protective membrane. The amoeba either takes its food directly by
imbibition of matter floating in the water, or by pressing into its
protoplasmic body solid particles with which it comes in contact. The
latter process may be observed at any moment by forcing it to eat. If
finely ground colouring matter, such as carmine or indigo, is put into
the water, you can see the body of the amoeba pressing these coloured
particles into itself, the substance of the cell closing round them.
The amoeba can take in food in this way at any point on its surface,
without having any special organs for intussusception and digestion,
or a real mouth or gut.

The amoeba grows by thus taking in food and dissolving the particles
eaten in its protoplasm. When it reaches a certain size by this
continual feeding, it begins to reproduce. This is done by the simple
process of cleavage (Figure 1.17). First, the nucleus divides into two
parts. Then the protoplasm is separated between the two new nuclei,
and the whole cell splits into two daughter-cells, the protoplasm
gathering about each of the nuclei. The thin bridge of protoplasm
which at first connects the daughter-cells soon breaks. Here we have
the simple form of direct cleavage of the nuclei. Without mitosis, or
formation of threads, the homogeneous nucleus divides into two halves.
These move away from each other, and become centres of attraction for
the enveloping matter, the protoplasm. The same direct cleavage of the
nuclei is also witnessed in the reproduction of many other protists,
while other unicellular organisms show the indirect division of the
cell.

Hence, although the amoeba is nothing but a simple cell, it is
evidently able to accomplish all the functions of the multicellular
organism. It moves, feels, nourishes itself, and reproduces. Some
kinds of these amoebae can be seen with the naked eye, but most of
them are microscopically small. It is for the following reasons that
we regard the amoebae as the unicellular organisms which have special
phylogenetic (or evolutionary) relations to the ovum. In many of the
lower animals the ovum retains its original naked form until
fertilisation, develops no membranes, and is then often
indistinguishable from the ordinary amoeba. Like the amoebae, these
naked ova may thrust out processes, and move about as travelling
cells. In the sponges these mobile ova move about freely in the
maternal body like independent amoebae (Figure 1.17). They had been
observed by earlier scientists, but described as foreign
bodies--namely, parasitic amoebae, living parasitically on the body of
the sponge. Later, however, it was discovered that they were not
parasites, but the ova of the sponge. We also find this remarkable
phenomenon among other animals, such as the graceful, bell-shaped
zoophytes, which we call polyps and medusae. Their ova remain naked
cells, which thrust out amoeboid projections, nourish themselves, and
move about. When they have been fertilised, the multicellular organism
is formed from them by repeated segmentation.

It is, therefore, no audacious hypothesis, but a perfectly sound
conclusion, to regard the amoeba as the particular unicellular
organism which offers us an approximate illustration of the ancient
common unicellular ancestor of all the metazoa, or multicellular
animals. The simple naked amoeba has a less definite and more original
character than any other cell. Moreover, there is the fact that recent
research has discovered such amoeba-like cells everywhere in the
mature body of the multicellular animals. They are found, for
instance, in the human blood, side by side with the red corpuscles, as
colourless blood-cells; and it is the same with all the vertebrates.
They are also found in many of the invertebrates--for instance, in the
blood of the snail. I showed, in 1859, that these colourless
blood-cells can, like the independent amoebae, take up solid
particles, or "eat" (whence they are called phagocytes =
"eating-cells," Figure 1.19). Lately, it has been discovered that many
different cells may, if they have room enough, execute the same
movements, creeping about and eating. They behave just like amoebae
(Figure 1.12). It has also been shown that these "travelling-cells,"
or planocytes, play an important part in man's physiology and
pathology (as means of transport for food, infectious matter,
bacteria, etc.).

The power of the naked cell to execute these characteristic
amoeba-like movements comes from the contractility (or automatic
mobility) of its protoplasm. This seems to be a universal property of
young cells. When they are not enclosed by a firm membrane, or
confined in a "cellular prison," they can always accomplish these
amoeboid movements. This is true of the naked ova as well as of any
other naked cells, of the "travelling-cells," of various kinds in
connective tissue, lymph-cells, mucus-cells, etc.

We have now, by our study of the ovum and the comparison of it with
the amoeba, provided a perfectly sound and most valuable foundation
for both the embryology and the evolution of man. We have learned that
the human ovum is a simple cell, that this ovum is not materially
different from that of other mammals, and that we may infer from it
the existence of a primitive unicellular ancestral form, with a
substantial resemblance to the amoeba.

The statement that the earliest progenitors of the human race were
simple cells of this kind, and led an independent unicellular life
like the amoeba, has not only been ridiculed as the dream of a natural
philosopher, but also been violently censured in theological journals
as "shameful and immoral." But, as I observed in my essay On the
Origin and Ancestral Tree of the Human Race in 1870, this offended
piety must equally protest against the "shameful and immoral" fact
that each human individual is developed from a simple ovum, and that
this human ovum is indistinguishable from those of the other mammals,
and in its earliest stage is like a naked amoeba. We can show this to
be a fact any day with the microscope, and it is little use to close
one's eyes to "immoral" facts of this kind. It is as indisputable as
the momentous conclusions we draw from it and as the vertebrate
character of man (see Chapter 1.11).

(FIGURE 1.19. Blood-cells that eat, or phagocytes, from a naked
sea-snail (Thetis), greatly magnified. I was the first to observe in
the blood-cells of this snail the important fact that "the blood-cells
of the invertebrates are unprotected pieces of plasm, and take in
food, by means of their peculiar movements, like the amoebae." I had
(in Naples, on May 10th, 1859) injected into the blood-vessels of one
of these snails an infusion of water and ground indigo, and was
greatly astonished to find the blood-cells themselves more or less
filled with the particles of indigo after a few hours. After repeated
injections I succeeded in "observing the very entrance of the coloured
particles in the blood-cells, which took place just in the same way as
with the amoeba." I have given further particulars about this in my
Monograph on the Radiolaria.)

We now see very clearly how extremely important the cell theory has
been for our whole conception of organic nature. "Man's place in
nature" is settled beyond question by it. Apart from the cell theory,
man is an insoluble enigma to us. Hence philosophers, and especially
physiologists, should be thoroughly conversant with it. The soul of
man can only be really understood in the light of the cell-soul, and
we have the simplest form of this in the amoeba. Only those who are
acquainted with the simple psychic functions of the unicellular
organisms and their gradual evolution in the series of lower animals
can understand how the elaborate mind of the higher vertebrates, and
especially of man, was gradually evolved from them. The academic
psychologists who lack this zoological equipment are unable to do so.

This naturalistic and realistic conception is a stumbling-block to our
modern idealistic metaphysicians and their theological colleagues.
Fenced about with their transcendental and dualistic prejudices, they
attack not only the monistic system we establish on our scientific
knowledge, but even the plainest facts which go to form its
foundation. An instructive instance of this was seen a few years ago,
in the academic discourse delivered by a distinguished theologian,
Willibald Beyschlag, at Halle, January 12th, 1900, on the occasion of
the centenary festival. The theologian protested violently against the
"materialistic dustmen of the scientific world who offer our people
the diploma of a descent from the ape, and would prove to them that
the genius of a Shakespeare or a Goethe is merely a distillation from
a drop of primitive mucus." Another well-known theologian protested
against "the horrible idea that the greatest of men, Luther and
Christ, were descended from a mere globule of protoplasm."
Nevertheless, not a single informed and impartial scientist doubts the
fact that these greatest men were, like all other men--and all other
vertebrates--developed from an impregnated ovum, and that this simple
nucleated globule of protoplasm has the same chemical constitution in
all the mammals.


CHAPTER 1.7. CONCEPTION.

The recognition of the fact that every man begins his individual
existence as a simple cell is the solid foundation of all research
into the genesis of man. From this fact we are forced, in virtue of
our biogenetic law, to draw the weighty phylogenetic conclusion that
the earliest ancestors of the human race were also unicellular
organisms; and among these protozoa we may single out the vague form
of the amoeba as particularly important (cf. Chapter 1.6). That these
unicellular ancestral forms did once exist follows directly from the
phenomena which we perceive every day in the fertilised ovum. The
development of the multicellular organism from the ovum, and the
formation of the germinal layers and the tissues, follow the same laws
in man and all the higher animals. It will, therefore, be our next
task to consider more closely the impregnated ovum and the process of
conception which produces it.

The process of impregnation or sexual conception is one of those
phenomena that people love to conceal behind the mystic veil of
supernatural power. We shall soon see, however, that it is a purely
mechanical process, and can be reduced to familiar physiological
functions. Moreover, this process of conception is of the same type,
and is effected by the same organs, in man as in all the other
mammals. The pairing of the male and female has in both cases for its
main purpose the introduction of the ripe matter of the male seed or
sperm into the female body, in the sexual canals of which it
encounters the ovum. Conception then ensues by the blending of the
two.

We must observe, first, that this important process is by no means so
widely distributed in the animal and plant world as is commonly
supposed. There is a very large number of lower organisms which
propagate unsexually, or by monogamy; these are especially the sexless
monera (chromacea, bacteria, etc.) but also many other protists, such
as the amoebae, foraminifera, radiolaria, myxomycetae, etc. In these
the multiplication of individuals takes place by unsexual
reproduction, which takes the form of cleavage, budding, or
spore-formation. The copulation of two coalescing cells, which in
these cases often precedes the reproduction, cannot be regarded as a
sexual act unless the two copulating plastids differ in size or
structure. On the other hand, sexual reproduction is the general rule
with all the higher organisms, both animal and plant; very rarely do
we find asexual reproduction among them. There are, in particular, no
cases of parthenogenesis (virginal conception) among the vertebrates.

Sexual reproduction offers an infinite variety of interesting forms in
the different classes of animals and plants, especially as regards the
mode of conception, and the conveyance of the spermatozoon to the
ovum. These features are of great importance not only as regards
conception itself, but for the development of the organic form, and
especially for the differentiation of the sexes. There is a
particularly curious correlation of plants and animals in this
respect. The splendid studies of Charles Darwin and Hermann Muller on
the fertilisation of flowers by insects have given us very interesting
particulars of this.* (* See Darwin's work, On the Various
Contrivances by which Orchids are Fertilised (1862).) This reciprocal
service has given rise to a most intricate sexual apparatus. Equally
elaborate structures have been developed in man and the higher
animals, serving partly for the isolation of the sexual products on
each side, partly for bringing them together in conception. But,
however interesting these phenomena are in themselves, we cannot go
into them here, as they have only a minor importance--if any at
all--in the real process of conception. We must, however, try to get a
very clear idea of this process and the meaning of sexual
reproduction.

In every act of conception we have, as I said, to consider two
different kinds of cells--a female and a male cell. The female cell of
the animal organism is always called the ovum (or ovulum, egg, or
egg-cell); the male cells are known as the sperm or seed-cells, or the
spermatozoa (also spermium and zoospermium). The ripe ovum is, on the
whole, one of the largest cells we know. It attains colossal
dimensions when it absorbs great quantities of nutritive yelk, as is
the case with birds and reptiles and many of the fishes. In the great
majority of the animals the ripe ovum is rich in yelk and much larger
than the other cells. On the other hand, the next cell which we have
to consider in the process of conception, the male sperm-cell or
spermatozoon, is one of the smallest cells in the animal body.
Conception usually consists in the bringing into contact with the ovum
of a slimy fluid secreted by the male, and this may take place either
inside or out of the female body. This fluid is called sperm, or the
male seed. Sperm, like saliva or blood, is not a simple fluid, but a
thick agglomeration of innumerable cells, swimming about in a
comparatively small quantity of fluid. It is not the fluid, but the
independent male cells that swim in it, that cause conception.

(FIGURE 1.20. Spermia or spermatozoa of various mammals. The
pear-shaped flattened nucleus is seen from the front in I and sideways
in II. k is the nucleus, m its middle part (protoplasm), s the mobile,
serpent-like tail (or whip); M four human spermatozoa, A spermatozoa
from the ape; K from the rabbit; H from the mouse; C from the dog; S
from the pig.

FIGURE 1.21. Spermatozoa or spermidia of various animals. (From Lang).
a of a fish, b of a turbellaria worm (with two side-lashes), c to e of
a nematode worm (amoeboid spermatozoa), f from a craw fish
(star-shaped), g from the salamander (with undulating membrane), h of
an annelid (a and h are the usual shape).

FIGURE 1.22. A single human spermatozoon magnified 2000 times; a shows
it from the broader and b from the narrower side. k head (with
nucleus), m middle-stem, h long-stem, and e tail. (From Retzius.))

The spermatozoa of the great majority of animals have two
characteristic features. Firstly, they are extraordinarily small,
being usually the smallest cells in the body; and, secondly, they
have, as a rule, a peculiarly lively motion, which is known as
spermatozoic motion. The shape of the cell has a good deal to do with
this motion. In most of the animals, and also in many of the lower
plants (but not the higher) each of these spermatozoa has a very
small, naked cell-body, enclosing an elongated nucleus, and a long
thread hanging from it (Figure 1.20). It was long before we could
recognise that these structures are simple cells. They were formerly
held to be special organisms, and were called "seed animals"
(spermato-zoa, or spermato-zoidia); they are now scientifically known
as spermia or spermidia, or as spermatosomata (seed-bodies) or
spermatofila (seed threads). It took a good deal of comparative
research to convince us that each of these spermatozoa is really a
simple cell. They have the same shape as in many other vertebrates and
most of the invertebrates. However, in many of the lower animals they
have quite a different shape. Thus, for instance, in the craw fish
they are large round cells, without any movement, equipped with stiff
outgrowths like bristles (Figure 1.21 f). They have also a peculiar
form in some of the worms, such as the thread-worms (filaria); in this
case they are sometimes amoeboid and like very small ova (Figure 1.21
c to e). But in most of the lower animals (such as the sponges and
polyps) they have the same pine-cone shape as in man and the other
animals (Figure 1.21 a, h).

When the Dutch naturalist Leeuwenhoek discovered these thread-like
lively particles in 1677 in the male sperm, it was generally believed
that they were special, independent, tiny animalcules, like the
infusoria, and that the whole mature organism existed already, with
all its parts, but very small and packed together, in each
spermatozoon (see Chapter 1.2). We now know that the mobile
spermatozoa are nothing but simple and real cells, of the kind that we
call "ciliated" (equipped with lashes, or cilia). In the previous
illustrations we have distinguished in the spermatozoon a head, trunk,
and tail. The "head" (Figure 1.20 k) is merely the oval nucleus of the
cell; the body or middle-part (m) is an accumulation of cell-matter;
and the tail (s) is a thread-like prolongation of the same.

Moreover, we now know that these spermatozoa are not at all a peculiar
form of cell; precisely similar cells are found in various other parts
of the body. If they have many short threads projecting, they are
called ciliated; if only one long, whip-shaped process (or, more
rarely, two or four), caudate (tailed) cells.

Very careful recent examination of the spermia, under a very high
microscopic power (Figure 1.22 a, b), has detected some further
details in the finer structure of the ciliated cell, and these are
common to man and the anthropoid ape. The head (k) encloses the
elliptic nucleus in a thin envelope of cytoplasm; it is a little
flattened on one side, and thus looks rather pear-shaped from the
front (b). In the central piece (m) we can distinguish a short neck
and a longer connective piece (with central body). The tail consists
of a long main section (h) and a short, very fine tail (e).

The process of fertilisation by sexual conception consists, therefore,
essentially in the coalescence and fusing together of two different
cells. The lively spermatozoon travels towards the ovum by its
serpentine movements, and bores its way into the female cell (Figure
1.23). The nuclei of both sexual cells, attracted by a certain
"affinity," approach each other and melt into one.

The fertilised cell is quite another thing from the unfertilised cell.
For if we must regard the spermia as real cells no less than the ova,
and the process of conception as a coalescence of the two, we must
consider the resultant cell as a quite new and independent organism.
It bears in the cell and nuclear matter of the penetrating
spermatozoon a part of the father's body, and in the protoplasm and
caryoplasm of the ovum a part of the mother's body. This is clear from
the fact that the child inherits many features from both parents. It
inherits from the father by means of the spermatozoon, and from the
mother by means of the ovum. The actual blending of the two cells
produces a third cell, which is the germ of the child, or the new
organism conceived. One may also say of this sexual coalescence that
the STEM-CELL IS A SIMPLE HERMAPHRODITE; it unites both sexual
substances in itself.

(FIGURE 1.23. The fertilisation of the ovum by the spermatozoon (of a
mammal). One of the many thread-like, lively spermidia pierces through
a fine pore-canal into the nuclear yelk. The nucleus of the ovum is
invisible.

FIGURE 1.24. An impregnated echinoderm ovum, with small homogeneous
nucleus (e k). (From Hertwig.))

I think it necessary to emphasise the fundamental importance of this
simple, but often unappreciated, feature in order to have a correct
and clear idea of conception. With that end, I have given a special
name to the new cell from which the child develops, and which is
generally loosely called "the fertilised ovum," or "the first
segmentation sphere." I call it "the stem-cell" (cytula). The name
"stem-cell" seems to me the simplest and most suitable, because all
the other cells of the body are derived from it, and because it is, in
the strictest sense, the stem-father and stem-mother of all the
countless generations of cells of which the multicellular organism is
to be composed. That complicated molecular movement of the protoplasm
which we call "life" is, naturally, something quite different in this
stem-cell from what we find in the two parent-cells, from the
coalescence of which it has issued. THE LIFE OF THE STEM-CELL OR
CYTULA IS THE PRODUCT OR RESULTANT OF THE PATERNAL LIFE-MOVEMENT THAT
IS CONVEYED IN THE SPERMATOZOON AND THE MATERNAL LIFE-MOVEMENT THAT IS
CONTRIBUTED BY THE OVUM.

The admirable work done by recent observers has shown that the
individual development, in man and the other animals, commences with
the formation of a simple "stem-cell" of this character, and that this
then passes, by repeated segmentation (or cleavage), into a cluster of
cells, known as "the segmentation sphere" or "segmentation cells." The
process is most clearly observed in the ova of the echinoderms
(star-fishes, sea-urchins, etc.). The investigations of Oscar and
Richard Hertwig were chiefly directed to these. The main results may
be summed up as follows:--

Conception is preceded by certain preliminary changes, which are very
necessary--in fact, usually indispensable--for its occurrence. They
are comprised under the general heading of "Changes prior to
impregnation." In these the original nucleus of the ovum, the germinal
vesicle, is lost. Part of it is extruded, and part dissolved in the
cell contents; only a very small part of it is left to form the basis
of a fresh nucleus, the pronucleus femininus. It is the latter alone
that combines in conception with the invading nucleus of the
fertilising spermatozoon (the pronucleus masculinus).

The impregnation of the ovum commences with a decay of the germinal
vesicle, or the original nucleus of the ovum (Figure 1.8). We have
seen that this is in most unripe ova a large, transparent, round
vesicle. This germinal vesicle contains a viscous fluid (the
caryolymph). The firm nuclear frame (caryobasis) is formed of the
enveloping membrane and a mesh-work of nuclear threads running across
the interior, which is filled with the nuclear sap. In a knot of the
network is contained the dark, stiff, opaque nuclear corpuscle or
nucleolus. When the impregnation of the ovum sets in, the greater part
of the germinal vesicle is dissolved in the cell; the nuclear membrane
and mesh-work disappear; the nuclear sap is distributed in the
protoplasm; a small portion of the nuclear base is extruded; another
small portion is left, and is converted into the secondary nucleus, or
the female pro-nucleus (Figure 1.24 e k).

The small portion of the nuclear base which is extruded from the
impregnated ovum is known as the "directive bodies" or "polar cells";
there are many disputes as to their origin and significance, but we
are as yet imperfectly acquainted with them. As a rule, they are two
small round granules, of the same size and appearance as the remaining
pro-nucleus. They are detached cell-buds; their separation from the
large mother-cell takes place in the same way as in ordinary "indirect
cell-division." Hence, the polar cells are probably to be conceived as
"abortive ova," or "rudimentary ova," which proceed from a simple
original ovum by cleavage in the same way that several sperm-cells
arise from one "sperm-mother-cell," in reproduction from sperm. The
male sperm-cells in the testicles must undergo similar changes in view
of the coming impregnation as the ova in the female ovary. In this
maturing of the sperm each of the original seed-cells divides by
double segmentation into four daughter-cells, each furnished with a
fourth of the original nuclear matter (the hereditary chromatin); and
each of these four descendant cells becomes a spermatozoon, ready for
impregnation. Thus is prevented the doubling of the chromatin in the
coalescence of the two nuclei at conception. As the two polar cells
are extruded and lost, and have no further part in the fertilisation
of the ovum, we need not discuss them any further. But we must give
more attention to the female pro-nucleus which alone remains after the
extrusion of the polar cells and the dissolving of the germinal
vesicle (Figure 1.23 e k). This tiny round corpuscle of chromatin now
acts as a centre of attraction for the invading spermatozoon in the
large ripe ovum, and coalesces with its "head," the male pro-nucleus.
The product of this blending, which is the most important part of the
act of impregnation, is the stem-nucleus, or the first segmentation
nucleus (archicaryon)--that is to say, the nucleus of the new-born
embryonic stem-cell or "first segmentation cell." This stem-cell is
the starting point of the subsequent embryonic processes.

Hertwig has shown that the tiny transparent ova of the echinoderms are
the most convenient for following the details of this important
process of impregnation. We can, in this case, easily and successfully
accomplish artificial impregnation, and follow the formation of the
stem-cell step by step within the space of ten minutes. If we put ripe
ova of the star-fish or sea-urchin in a watch glass with sea-water and
add a drop of ripe sperm-fluid, we find each ovum impregnated within
five minutes. Thousands of the fine, mobile ciliated cells, which we
have described as "sperm-threads" (Figure 1.20), make their way to the
ova, owing to a sort of chemical sensitive action which may be called
"smell." But only one of these innumerable spermatozoa is
chosen--namely, the one that first reaches the ovum by the serpentine
motions of its tail, and touches the ovum with its head. At the spot
where the point of its head touches the surface of the ovum the
protoplasm of the latter is raised in the form of a small wart, the
"impregnation rise" (Figure 1.25 A). The spermatozoon then bores its
way into this with its head, the tail outside wriggling about all the
time (Figure 1.25 B, C). Presently the tail also disappears within the
ovum. At the same time the ovum secretes a thin external yelk-membrane
(Figure 1.25 C), starting from the point of impregnation; and this
prevents any more spermatozoa from entering.

Inside the impregnated ovum we now see a rapid series of most
important changes. The pear-shaped head of the sperm-cell, or the
"head of the spermatozoon," grows larger and rounder, and is converted
into the male pro-nucleus (Figure 1.26 s k). This has an attractive
influence on the fine granules or particles which are distributed in
the protoplasm of the ovum; they arrange themselves in lines in the
figure of a star. But the attraction or the "affinity" between the two
nuclei is even stronger. They move towards each other inside the yelk
with increasing speed, the male (Figure 1.27 s k) going more quickly
than the female nucleus (e k). The tiny male nucleus takes with it the
radiating mantle which spreads like a star about it. At last the two
sexual nuclei touch (usually in the centre of the globular ovum), lie
close together, are flattened at the points of contact, and coalesce
into a common mass. The small central particle of nuclein which is
formed from this combination of the nuclei is the stem-nucleus, or the
first segmentation nucleus; the new-formed cell, the product of the
impregnation, is our stem-cell, or "first segmentation sphere" (Figure
1.2).

(FIGURE 1.25. Impregnation of the ovum of a star-fish. (From Hertwig.)
Only a small part of the surface of the ovum is shown. One of the
numerous spermatozoa approaches the "impregnation rise" (A), touches
it (B), and then penetrates into the protoplasm of the ovum (C).

FIGURES 1.26 AND 1.27. Impregnation of the ovum of the sea-urchin.
(From Hertwig.) In Figure 1.26 the little sperm-nucleus (sk) moves
towards the larger nucleus of the ovum (ek). In Figure 1.27 they
nearly touch, and are surrounded by the radiating mantle of
protoplasm.)

Hence the one essential point in the process of sexual reproduction or
impregnation is the formation of a new cell, the stem-cell, by the
combination of two originally different cells, the female ovum and the
male spermatozoon. This process is of the highest importance, and
merits our closest attention; all that happens in the later
development of this first cell and in the life of the organism that
comes of it is determined from the first by the chemical and
morphological composition of the stem-cell, its nucleus and its body.
We must, therefore, make a very careful study of the rise and
structure of the stem-cell.

The first question that arises is as to the two different active
elements, the nucleus and the protoplasm, in the actual coalescence.
It is obvious that the nucleus plays the more important part in this.
Hence Hertwig puts his theory of conception in the principle:
"Conception consists in the copulation of two cell-nuclei, which come
from a male and a female cell." And as the phenomenon of heredity is
inseparably connected with the reproductive process, we may further
conclude that these two copulating nuclei "convey the characteristics
which are transmitted from parents to offspring." In this sense I had
in 1866 (in the ninth chapter of the General Morphology) ascribed to
the reproductive nucleus the function of generation and heredity, and
to the nutritive protoplasm the duties of nutrition and adaptation.
As, moreover, there is a complete coalescence of the mutually
attracted nuclear substances in conception, and the new nucleus formed
(the stem-nucleus) is the real starting-point for the development of
the fresh organism, the further conclusion may be drawn that the male
nucleus conveys to the child the qualities of the father, and the
female nucleus the features of the mother. We must not forget,
however, that the protoplasmic bodies of the copulating cells also
fuse together in the act of impregnation; the cell-body of the
invading spermatozoon (the trunk and tail of the male ciliated cell)
is dissolved in the yelk of the female ovum. This coalescence is not
so important as that of the nuclei, but it must not be overlooked;
and, though this process is not so well known to us, we see clearly at
least the formation of the star-like figure (the radial arrangement of
the particles in the plasma) in it (Figures 1.26 to 1.27).

The older theories of impregnation generally went astray in regarding
the large ovum as the sole base of the new organism, and only ascribed
to the spermatozoon the work of stimulating and originating its
development. The stimulus which it gave to the ovum was sometimes
thought to be purely chemical, at other times rather physical (on the
principle of transferred movement), or again a mystic and
transcendental process. This error was partly due to the imperfect
knowledge at that time of the facts of impregnation, and partly to the
striking difference in the sizes of the two sexual cells. Most of the
earlier observers thought that the spermatozoon did not penetrate into
the ovum. And even when this had been demonstrated, the spermatozoon
was believed to disappear in the ovum without leaving a trace.
However, the splendid research made in the last three decades with the
finer technical methods of our time has completely exposed the error
of this. It has been shown that the tiny sperm-cell is NOT
SUBORDINATED TO, BUT COORDINATED WITH, the large ovum. The nuclei of
the two cells, as the vehicles of the hereditary features of the
parents, are of equal physiological importance. In some cases we have
succeeded in proving that the mass of the active nuclear substance
which combines in the copulation of the two sexual nuclei is
originally the same for both.

These morphological facts are in perfect harmony with the familiar
physiological truth that the child inherits from both parents, and
that on the average they are equally distributed. I say "on the
average," because it is well known that a child may have a greater
likeness to the father or to the mother; that goes without saying, as
far as the primary sexual characters (the sexual glands) are
concerned. But it is also possible that the determination of the
latter--the weighty determination whether the child is to be a boy or
a girl--depends on a slight qualitative or quantitative difference in
the nuclein or the coloured nuclear matter which comes from both
parents in the act of conception.

The striking differences of the respective sexual cells in size and
shape, which occasioned the erroneous views of earlier scientists, are
easily explained on the principle of division of labour. The inert,
motionless ovum grows in size according to the quantity of provision
it stores up in the form of nutritive yelk for the development of the
germ. The active swimming sperm-cell is reduced in size in proportion
to its need to seek the ovum and bore its way into its yelk. These
differences are very conspicuous in the higher animals, but they are
much less in the lower animals. In those protists (unicellular plants
and animals) which have the first rudiments of sexual reproduction the
two copulating cells are at first quite equal. In these cases the act
of impregnation is nothing more than a sudden GROWTH, in which the
originally simple cell doubles its volume, and is thus prepared for
reproduction (cell-division). Afterwards slight differences are seen
in the size of the copulating cells; though the smaller ones still
have the same shape as the larger ones. It is only when the difference
in size is very pronounced that a notable difference in shape is
found: the sprightly sperm-cell changes more in shape and the ovum in
size.

Quite in harmony with this new conception of the EQUIVALENCE OF THE
TWO GONADS, or the equal physiological importance of the male and
female sex-cells and their equal share in the process of heredity, is
the important fact established by Hertwig (1875), that in normal
impregnation only one single spermatozoon copulates with one ovum; the
membrane which is raised on the surface of the yelk immediately after
one sperm-cell has penetrated (Figure 1.25 C) prevents any others from
entering. All the rivals of the fortunate penetrator are excluded, and
die without. But if the ovum passes into a morbid state, if it is made
stiff by a lowering of its temperature or stupefied with narcotics
(chloroform, morphia, nicotine, etc.), two or more spermatozoa may
penetrate into its yelk-body. We then witness polyspermism. The more
Hertwig chloroformed the ovum, the more spermatozoa were able to bore
their way into its unconscious body.

(FIGURE 1.28. Stem-cell of a rabbit, magnified 200 times. In the
centre of the granular protoplasm of the fertilised ovum (d) is seen
the little, bright stem-nucleus, z is the ovolemma, with a mucous
membrane (h). s are dead spermatozoa.)

These remarkable facts of impregnation are also of the greatest
interest in psychology, especially as regards the theory of the
cell-soul, which I consider to be its chief foundation. The phenomena
we have described can only be understood and explained by ascribing a
certain lower degree of psychic activity to the sexual principles.
They FEEL each other's proximity, and are drawn together by a
SENSITIVE impulse (probably related to smell); they MOVE towards each
other, and do not rest until they fuse together. Physiologists may say
that it is only a question of a peculiar physico-chemical phenomenon,
and not a psychic action; but the two cannot be separated. Even the
psychic functions, in the strict sense of the word, are only complex
physical processes, or "psycho-physical" phenomena, which are
determined in all cases exclusively by the chemical composition of
their material substratum.

The monistic view of the matter becomes clear enough when we remember
the radical importance of impregnation as regards heredity. It is well
known that not only the most delicate bodily structures, but also the
subtlest traits of mind, are transmitted from the parents to the
children. In this the chromatic matter of the male nucleus is just as
important a vehicle as the large caryoplasmic substance of the female
nucleus; the one transmits the mental features of the father, and the
other those of the mother. The blending of the two parental nuclei
determines the individual psychic character of the child.

But there is another important psychological question--the most
important of all--that has been definitely answered by the recent
discoveries in connection with conception. This is the question of the
immortality of the soul. No fact throws more light on it and refutes
it more convincingly than the elementary process of conception that we
have described. For this copulation of the two sexual nuclei (Figures
1.26 and 1.27) indicates the precise moment at which the individual
begins to exist. All the bodily and mental features of the new-born
child are the sum-total of the hereditary qualities which it has
received in reproduction from parents and ancestors. All that man
acquires afterwards in life by the exercise of his organs, the
influence of his environment, and education--in a word, by
adaptation--cannot obliterate that general outline of his being which
he inherited from his parents. But this hereditary disposition, the
essence of every human soul, is not "eternal," but "temporal"; it
comes into being only at the moment when the sperm-nucleus of the
father and the nucleus of the maternal ovum meet and fuse together. It
is clearly irrational to assume an "eternal life without end" for an
individual phenomenon, the commencement of which we can indicate to a
moment by direct visual observation.

The great importance of the process of impregnation in answering such
questions is quite clear. It is true that conception has never been
studied microscopically in all its details in the human
case--notwithstanding its occurrence at every moment--for reasons that
are obvious enough. However, the two cells which need consideration,
the female ovum and the male spermatozoon, proceed in the case of man
in just the same way as in all the other mammals; the human foetus or
embryo which results from copulation has the same form as with the
other animals. Hence, no scientist who is acquainted with the facts
doubts that the processes of impregnation are just the same in man as
in the other animals.

The stem-cell which is produced, and with which every man begins his
career, cannot be distinguished in appearance from those of other
mammals, such as the rabbit (Figure 1.28). In the case of man, also,
this stem-cell differs materially from the original ovum, both in
regard to form (morphologically), in regard to material composition
(chemically), and in regard to vital properties (physiologically). It
comes partly from the father and partly from the mother. Hence it is
not surprising that the child who is developed from it inherits from
both parents. The vital movements of each of these cells form a sum of
mechanical processes which in the last analysis are due to movements
of the smallest vital parts, or the molecules, of the living
substance. If we agree to call this active substance plasson, and its
molecules plastidules, we may say that the individual physiological
character of each of these cells is due to its molecular
plastidule-movement. HENCE, THE PLASTIDULE-MOVEMENT OF THE CYTULA IS
THE RESULTANT OF THE COMBINED PLASTIDULE-MOVEMENTS OF THE FEMALE OVUM
AND THE MALE SPERM-CELL.* (* The plasson of the stem-cell or cytula
may, from the anatomical point of view, be regarded as homogeneous and
structureless, like that of the monera. This is not inconsistent with
our hypothetical ascription to the plastidules (or molecules of the
plasson) of a complex molecular structure. The complexity of this is
the greater in proportion to the complexity of the organism that is
developed from it and the length of the chain of its ancestry, or to
the multitude of antecedent processes of heredity and adaptation.)


CHAPTER 1.8. THE GASTRAEA THEORY.

There is a substantial agreement throughout the animal world in the
first changes which follow the impregnation of the ovum and the
formation of the stem-cell; they begin in all cases with the
segmentation of the ovum and the formation of the germinal layers. The
only exception is found in the protozoa, the very lowest and simplest
forms of animal life; these remain unicellular throughout life. To
this group belong the amoebae, gregarinae, rhizopods, infusoria, etc.
As their whole organism consists of a single cell, they can never form
germinal layers, or definite strata of cells. But all the other
animals--all the tissue-forming animals, or metazoa, as we call them,
in contradistinction to the protozoa--construct real germinal layers
by the repeated cleavage of the impregnated ovum. This we find in the
lower cnidaria and worms, as well as in the more highly-developed
molluscs, echinoderms, articulates, and vertebrates.

In all these metazoa, or multicellular animals, the chief embryonic
processes are substantially alike, although they often seem to a
superficial observer to differ considerably. The stem-cell that
proceeds from the impregnated ovum always passes by repeated cleavage
into a number of simple cells. These cells are all direct descendants
of the stem-cell, and are, for reasons we shall see presently, called
segmentation-cells. The repeated cleavage of the stem-cell, which
gives rise to these segmentation-spheres, has long been known as
"segmentation." Sooner or later the segmentation-cells join together
to form a round (at first, globular) embryonic sphere (blastula); they
then form into two very different groups, and arrange themselves in
two separate strata--the two primary germinal layers. These enclose a
digestive cavity, the primitive gut, with an opening, the primitive
mouth. We give the name of the gastrula to the important embryonic
form that has these primitive organs, and the name of gastrulation to
the formation of it. This ontogenetic process has a very great
significance, and is the real starting-point of the construction of
the multicellular animal body.

The fundamental embryonic processes of the cleavage of the ovum and
the formation of the germinal layers have been very thoroughly studied
in the last thirty years, and their real significance has been
appreciated. They present a striking variety in the different groups,
and it was no light task to prove their essential identity in the
whole animal world. But since I formulated the gastraea theory in
1872, and afterwards (1875) reduced all the various forms of
segmentation and gastrulation to one fundamental type, their identity
may be said to have been established. We have thus mastered the law of
unity which governs the first embryonic processes in all the animals.

Man is like all the other higher animals, especially the apes, in
regard to these earliest and most important processes. As the human
embryo does not essentially differ, even at a much later stage of
development--when we already perceive the cerebral vesicles, the eyes,
ears, gill-arches, etc.--from the similar forms of the other higher
mammals, we may confidently assume that they agree in the earliest
embryonic processes, segmentation and the formation of germinal
layers. This has not yet, it is true, been established by observation.
We have never yet had occasion to dissect a woman immediately after
impregnation and examine the stem-cell or the segmentation-cells in
her oviduct. However, as the earliest human embryos we have examined,
and the later and more developed forms, agree with those of the
rabbit, dog, and other higher mammals, no reasonable man will doubt
but that the segmentation and formation of layers are the same in both
cases.

But the special form of segmentation and layer formation which we find
in the mammal is by no means the original, simple, palingenetic form.
It has been much modified and cenogenetically altered by a very
complex adaptation to embryonic conditions. We cannot, therefore,
understand it altogether in itself. In order to do this, we have to
make a COMPARATIVE study of segmentation and layer-formation in the
animal world; and we have especially to seek the original,
PALINGENETIC form from which the modified CENOGENETIC (see Chapter
1.1) form has gradually been developed.

This original unaltered form of segmentation and layer-formation is
found to-day in only one case in the vertebrate-stem to which man
belongs--the lowest and oldest member of the stem, the wonderful
lancelet or amphioxus (cf. Chapters 2.16 and 2.17). But we find a
precisely similar palingenetic form of embryonic development in the
case of many of the invertebrate animals, as, for instance, the
remarkable ascidia, the pond-snail (Limnaeus), and arrow-worm
(Sagitta), and many of the echinoderms and cnidaria, such as the
common star-fish and sea-urchin, many of the medusae and corals, and
the simpler sponges (Olynthus). We may take as an illustration the
palingenetic segmentation and germinal layer-formation in an
eight-fold insular coral, which I discovered in the Red Sea, and
described as Monoxenia Darwinii.

(FIGURE 1.29. Gastrulation of a coral (Monoxenia Darwinii). A, B,
stem-cell (cytula) or impregnated ovum. In Figure A (immediately after
impregnation) the nucleus is invisible. In Figure B (a little later)
it is quite clear. C two segmentation-cells. D four
segmentation-cells. E mulberry-formation (morula). F blastosphere
(blastula). G blastula (transverse section). H depula, or hollowed
blastula (transverse section). I gastrula (longitudinal section). K
gastrula, or cup-sphere, external appearance.)

The impregnated ovum of this coral (Figure 1.29 A, B) first splits
into two equal cells (C). First, the nucleus of the stem-cell and its
central body divide into two halves. These recede from and repel each
other, and act as centres of attraction on the surrounding protoplasm;
in consequence of this, the protoplasm is constricted by a circular
furrow, and, in turn, divides into two halves. Each of the two
segmentation-cells thus produced splits in the same way into two equal
cells. The four segmentation-cells (grand-daughters of the stem-cell)
lie in one plane. Now, however, each of them subdivides into two equal
halves, the cleavage of the nucleus again preceding that of the
surrounding protoplasm. The eight cells which thus arise break into
sixteen, these into thirty-two, and then (each being constantly
halved) into sixty-four, 128, and so on.* (* The number of
segmentation-cells thus produced increases geometrically in the
original gastrulation, or the purest palingenetic form of cleavage.
However, in different animals the number reaches a different height,
so that the morula, and also the blastula, may consist sometimes of
thirty-two, sometimes of sixty-four, and sometimes of 128, or more,
cells.) The final result of this repeated cleavage is the formation of
a globular cluster of similar segmentation-cells, which we call the
mulberry-formation or morula. The cells are thickly pressed together
like the parts of a mulberry or blackberry, and this gives a lumpy
appearance to the surface of the sphere (Figure E).* (* The
segmentation-cells which make up the morula after the close of the
palingenetic cleavage seem usually to be quite similar, and to present
no differences as to size, form, and composition. That, however, does
not prevent them from differentiating into animal and vegetative
cells, even during the cleavage.)

When the cleavage is thus ended, the mulberry-like mass changes into a
hollow globular sphere. Watery fluid or jelly gathers inside the
globule; the segmentation-cells are loosened, and all rise to the
surface. There they are flattened by mutual pressure, and assume the
shape of truncated pyramids, and arrange themselves side by side in
one regular layer (Figures F, G). This layer of cells is called the
germinal membrane (or blastoderm); the homogeneous cells which compose
its simple structure are called blastodermic cells; and the whole
hollow sphere, the walls of which are made of the preceding, is called
the blastula or blastosphere.* (* The blastula of the lower animals
must not be confused with the very different blastula of the mammal,
which is properly called the gastrocystis or blastocystis. This
cenogenetic gastrocystis and the palingenetic blastula are sometimes
very wrongly comprised under the common name of blastula or vesicula
blastodermica.)

In the case of our coral, and of many other lower forms of animal
life, the young embryo begins at once to move independently and swim
about in the water. A fine, long, thread-like process, a sort of whip
or lash, grows out of each blastodermic cell, and this independently
executes vibratory movements, slow at first, but quicker after a time
(Figure F). In this way each blastodermic cell becomes a ciliated
cell. The combined force of all these vibrating lashes causes the
whole blastula to move about in a rotatory fashion. In many other
animals, especially those in which the embryo develops within enclosed
membranes, the ciliated cells are only formed at a later stage, or
even not formed at all. The blastosphere may grow and expand by the
blastodermic cells (at the surface of the sphere) dividing and
increasing, and more fluid is secreted in the internal cavity. There
are still to-day some organisms that remain throughout life at the
structural stage of the blastula--hollow vesicles that swim about by a
ciliary movement in the water, the wall of which is composed of a
single layer of cells, such as the volvox, the magosphaera, synura,
etc. We shall speak further of the great phylogenetic significance of
this fact in Chapter 2.19.

A very important and remarkable process now follows--namely, the
curving or invagination of the blastula (Figure H). The vesicle with a
single layer of cells for wall is converted into a cup with a wall of
two layers of cells (cf. Figures G, H, I). A certain spot at the
surface of the sphere is flattened, and then bent inward. This
depression sinks deeper and deeper, growing at the cost of the
internal cavity. The latter decreases as the hollow deepens. At last
the internal cavity disappears altogether, the inner side of the
blastoderm (that which lines the depression) coming to lie close on
the outer side. At the same time, the cells of the two sections assume
different sizes and shapes; the inner cells are more round and the
outer more oval (Figure I). In this way the embryo takes the form of a
cup or jar-shaped body, with a wall made up of two layers of cells,
the inner cavity of which opens to the outside at one end (the spot
where the depression was originally formed). We call this very
important and interesting embryonic form the "cup-embryo" or
"cup-larva" (gastrula, Figure 1.29, I longitudinal section, K external
view). I have in my Natural History of Creation given the name of
depula to the remarkable intermediate form which appears at the
passage of the blastula into the gastrula. In this intermediate stage
there are two cavities in the embryo--the original cavity (blastocoel)
which is disappearing, and the primitive gut-cavity (progaster) which
is forming.

I regard the gastrula as the most important and significant embryonic
form in the animal world. In all real animals (that is, excluding the
unicellular protists) the segmentation of the ovum produces either a
pure, primitive, palingenetic gastrula (Figure 1.29 I, K) or an
equally instructive cenogenetic form, which has been developed in time
from the first, and can be directly reduced to it. It is certainly a
fact of the greatest interest and instructiveness that animals of the
most different stems--vertebrates and tunicates, molluscs and
articulates, echinoderms and annelids, cnidaria and sponges--proceed
from one and the same embryonic form. In illustration I give a few
pure gastrula forms from various groups of animals (Figures 1.30 to
1.35, explanation given below each).

(FIGURES 1.30 TO 1.35. In each figure d is the primitive-gut cavity, o
primitive mouth, s segmentation-cavity, i entoderm (gut-layer), e
ectoderm (skin layer).

FIGURE 1.30. (A) Gastrula of a very simple primitive-gut animal or
gastraead (gastrophysema). (Haeckel.)

FIGURE 1.31. (B) Gastrula of a worm (Sagitta). (From Kowalevsky.)

FIGURE 1.32. (C) Gastrula of an echinoderm (star-fish, Uraster), not
completely folded in (depula). (From Alexander Agassiz.)

FIGURE 1.33. (D) Gastrula of an arthropod (primitive crab, Nauplius)
(as 32).

FIGURE 1.34. (E) Gastrula of a mollusc (pond-snail, Linnaeus). (From
Karl Rabl.)

FIGURE 1.35. (F) Gastrula of a vertebrate (lancelet, Amphioxus). (From
Kowalevsky.) (Front view.))

In view of this extraordinary significance of the gastrula, we must
make a very careful study of its original structure. As a rule, the
typical gastrula is very small, being invisible to the naked eye, or
at the most only visible as a fine point under very favourable
conditions, and measuring generally 1/500 to 1/250 of an inch (less
frequently 1/50 inch, or even more) in diameter. In shape it is
usually like a roundish drinking-cup. Sometimes it is rather oval, at
other times more ellipsoid or spindle-shaped; in some cases it is half
round, or even almost round, and in others lengthened out, or almost
cylindrical.

I give the name of primitive gut (progaster) and primitive mouth
(prostoma) to the internal cavity of the gastrula-body and its
opening; because this cavity is the first rudiment of the digestive
cavity of the organism, and the opening originally served to take food
into it. Naturally, the primitive gut and mouth change very
considerably afterwards in the various classes of animals. In most of
the cnidaria and many of the annelids (worm-like animals) they remain
unchanged throughout life. But in most of the higher animals, and so
in the vertebrates, only the larger central part of the later
alimentary canal develops from the primitive gut; the later mouth is a
fresh development, the primitive mouth disappearing or changing into
the anus. We must therefore distinguish carefully between the
primitive gut and mouth of the gastrula and the later alimentary canal
and mouth of the fully developed vertebrate.* (* My distinction (1872)
between the primitive gut and mouth and the later permanent stomach
(metagaster) and mouth (metastoma) has been much criticised; but it is
as much justified as the distinction between the primitive kidneys and
the permanent kidneys. Professor E. Ray-Lankester suggested three
years afterwards (1875) the name archenteron for the primitive gut,
and blastoporus for the primitive mouth.)

(FIGURE 1.36. Gastrula of a lower sponge (olynthus). A external view,
B longitudinal section through the axis, g primitive-gut cavity, a
primitive mouth-aperture, i inner cell-layer (entoderm, endoblast,
gut-layer), e external cell-layer (outer germinal layer, ectoderm,
ectoblast, or skin-layer).

The two layers of cells which line the gut-cavity and compose its wall
are of extreme importance. These two layers, which are the sole
builders of the whole organism, are no other than the two primary
germinal layers, or the primitive germ-layers. I have spoken in the
introductory section (Chapter 1.3.) of their radical importance. The
outer stratum is the skin-layer, or ectoderm (Figures 1.30 to 1.35 e);
the inner stratum is the gut-layer, or entoderm (i). The former is
often also called the ectoblast, or epiblast, and the latter the
endoblast, or hypoblast. FROM THESE TWO PRIMARY GERMINAL LAYERS ALONE
IS DEVELOPED THE ENTIRE ORGANISM OF ALL THE METAZOA OR MULTICELLULAR
ANIMALS. The skin-layer forms the external skin, the gut-layer forms
the internal skin or lining of the body. Between these two germinal
layers are afterwards developed the middle germinal layer (mesoderma)
and the body-cavity (coeloma) filled with blood or lymph.

The two primary germinal layers were first distinguished by Pander in
1817 in the incubated chick. Twenty years later (1849) Huxley pointed
out that in many of the lower zoophytes, especially the medusae, the
whole body consists throughout life of these two primary germinal
layers. Soon afterwards (1853) Allman introduced the names which have
come into general use; he called the outer layer the ectoderm
("outer-skin"), and the inner the entoderm ("inner-skin"). But in 1867
it was shown, particularly by Kowalevsky, from comparative
observation, that even in invertebrates, also, of the most different
classes--annelids, molluscs, echinoderms, and articulates--the body is
developed out of the same two primary layers. Finally, I discovered
them (1872) in the lowest tissue-forming animals, the sponges, and
proved in my gastraea theory that these two layers must be regarded as
identical throughout the animal world, from the sponges and corals to
the insects and vertebrates, including man. This fundamental "homology
[identity] of the primary germinal layers and the primitive gut" has
been confirmed during the last thirty years by the careful research of
many able observers, and is now pretty generally admitted for the
whole of the metazoa.

As a rule, the cells which compose the two primary germinal layers
show appreciable differences even in the gastrula stage. Generally (if
not always) the cells of the skin-layer or ectoderm (Figures 1.36 c
and 1.37 e) are the smaller, more numerous, and clearer; while the
cells of the gut-layer, or entoderm (i), are larger, less numerous,
and darker. The protoplasm of the ectodermic (outer) cells is clearer
and firmer than the thicker and softer cell-matter of the entodermic
(inner) cells; the latter are, as a rule, much richer in yelk-granules
(albumen and fatty particles) than the former. Also the cells of the
gut-layer have, as a rule, a stronger affinity for colouring matter,
and take on a tinge in a solution of carmine, aniline, etc., more
quickly and appreciably than the cells of the skin-layer. The nuclei
of the entoderm-cells are usually roundish, while those of the
ectoderm-cells are oval.

When the doubling-process is complete, very striking histological
differences between the cells of the two layers are found (Figure
1.37). The tiny, light ectoderm-cells (e) are sharply distinguished
from the larger and darker entoderm-cells (i). Frequently this
differentiation of the cell-forms sets in at a very early stage,
during the segmentation-process, and is already very appreciable in
the blastula.

We have, up to the present, only considered that form of segmentation
and gastrulation which, for many and weighty reasons, we may regard as
the original, primordial, or palingenetic form. We might call it
"equal" or homogeneous segmentation, because the divided cells retain
a resemblance to each other at first (and often until the formation of
the blastoderm). We give the name of the "bell-gastrula," or
archigastrula, to the gastrula that succeeds it. In just the same form
as in the coral we considered (Monoxenia, Figure 1.29), we find it in
the lowest zoophyta (the gastrophysema, Figure 1.30), and the simplest
sponges (olynthus, Figure 1.36); also in many of the medusae and
hydrapolyps, lower types of worms of various classes (brachiopod,
arrow-worm, Figure 1.31), tunicates (ascidia), many of the echinoderms
(Figure 1.32), lower articulates (Figure 1.33), and molluscs (Figure
1.34), and, finally, in a slightly modified form, in the lowest
vertebrate (the amphioxus, Figure 1.35).

(FIGURE 1.37. Cells from the two primary germinal layers of the mammal
(from both layers of the blastoderm). i larger and darker cells of the
inner stratum, the vegetal layer or entoderm. e smaller and clearer
cells from the outer stratum, the animal layer or ectoderm.

FIGURE 1.38. Gastrulation of the amphioxus, from Hatschek (vertical
section through the axis of the ovum). A, B, C three stages in the
formation of the blastula; D, E curving of the blastula; F complete
gastrula. h segmentation-cavity. g primitive gut-cavity.))

The gastrulation of the amphioxus is especially interesting because
this lowest and oldest of all the vertebrates is of the highest
significance in connection with the evolution of the vertebrate stem,
and therefore with that of man (compare Chapters 2.16 and 2.17). Just
as the comparative anatomist traces the most elaborate features in the
structures of the various classes of vertebrates to divergent
development from this simple primitive vertebrate, so comparative
embryology traces the various secondary forms of vertebrate
gastrulation to the simple, primary formation of the germinal layers
in the amphioxus. Although this formation, as distinguished from the
cenogenetic modifications of the vertebrate, may on the whole be
regarded as palingenetic, it is nevertheless different in some
features from the quite primitive gastrulation such as we have, for
instance, in the Monoxenia (Figure 1.29) and the Sagitta. Hatschek
rightly observes that the segmentation of the ovum in the amphioxus is
not strictly equal, but almost equal, and approaches the unequal. The
difference in size between the two groups of cells continues to be
very noticeable in the further course of the segmentation; the smaller
animal cells of the upper hemisphere divide more quickly than the
larger vegetal cells of the lower (Figure 1.38 A, B). Hence the
blastoderm, which forms the single-layer wall of the globular blastula
at the end of the cleavage-process, does not consist of homogeneous
cells of equal size, as in the Sagitta and the Monoxenia; the cells of
the upper half of the blastoderm (the mother-cells of the ectoderm)
are more numerous and smaller, and the cells of the lower half (the
mother-cells of the entoderm) less numerous and larger. Moreover, the
segmentation-cavity of the blastula (Figure 1.38 C, h) is not quite
globular, but forms a flattened spheroid with unequal poles of its
vertical axis. While the blastula is being folded into a cup at the
vegetal pole of its axis, the difference in the size of the
blastodermic cells increases (Figure 1.38 D, E); it is most
conspicuous when the invagination is complete and the
segmentation-cavity has disappeared (Figure 1.38 F). The larger
vegetal cells of the entoderm are richer in granules, and so darker
than the smaller and lighter animal cells of the ectoderm.

But the unequal gastrulation of the amphioxus diverges from the
typical equal cleavage of the Sagitta, the Monoxenia (Figure 1.29),
and the Olynthus (Figure 1.36), in another important particular. The
pure archigastrula of the latter forms is uni-axial, and it is round
in its whole length in transverse section. The vegetal pole of the
vertical axis is just in the centre of the primitive mouth. This is
not the case in the gastrula of the amphioxus. During the folding of
the blastula the ideal axis is already bent on one side, the growth of
the blastoderm (or the increase of its cells) being brisker on one
side than on the other; the side that grows more quickly, and so is
more curved (Figure 1.39 v), will be the anterior or belly-side, the
opposite, flatter side will form the back (d). The primitive mouth,
which at first, in the typical archigastrula, lay at the vegetal pole
of the main axis, is forced away to the dorsal side; and whereas its
two lips lay at first in a plane at right angles to the chief axis,
they are now so far thrust aside that their plane cuts the axis at a
sharp angle. The dorsal lip is therefore the upper and more forward,
the ventral lip the lower and hinder. In the latter, at the ventral
passage of the entoderm into the ectoderm, there lie side by side a
pair of very large cells, one to the right and one to the left (Figure
1.39 p): these are the important polar cells of the primitive mouth,
or "the primitive cells of the mesoderm." In consequence of these
considerable variations arising in the course of the gastrulation, the
primitive uni-axial form of the archigastrula in the amphioxus has
already become tri-axial, and thus the two-sidedness, or bilateral
symmetry, of the vertebrate body has already been determined. This has
been transmitted from the amphioxus to all the other modified
gastrula-forms of the vertebrate stem.

Apart from this bilateral structure, the gastrula of the amphioxus
resembles the typical archigastrula of the lower animals (Figures 1.30
to 1.36) in developing the two primary germinal layers from a single
layer of cells. This is clearly the oldest and original form of the
metazoic embryo. Although the animals I have mentioned belong to the
most diverse classes, they nevertheless agree with each other, and
many more animal forms, in having retained to the present day, by a
conservative heredity, this palingenetic form of gastrulation which
they have from their earliest common ancestors. But this is not the
case with the great majority of the animals. With these the original
embryonic process has been gradually more or less altered in the
course of millions of years by adaptation to new conditions of
development. Both the segmentation of the ovum and the subsequent
gastrulation have in this way been considerably changed. In fact,
these variations have become so great in the course of time that the
segmentation was not rightly understood in most animals, and the
gastrula was unrecognised. It was not until I had made an extensive
comparative study, lasting a considerable time (in the years 1866 to
1875), in animals of the most diverse classes, that I succeeded in
showing the same common typical process in these apparently very
different forms of gastrulation, and tracing them all to one original
form. I regard all those that diverge from the primary palingenetic
gastrulation as secondary, modified, and cenogenetic. The more or less
divergent form of gastrula that is produced may be called a secondary,
modified gastrula, or a metagastrula. The reader will find a scheme of
these different kinds of segmentation and gastrulation at the close of
this chapter.

By far the most important process that determines the various
cenogenetic forms of gastrulation is the change in the nutrition of
the ovum and the accumulation in it of nutritive yelk. By this we
understand various chemical substances (chiefly granules of albumin
and fat-particles) which serve exclusively as reserve-matter or food
for the embryo. As the metazoic embryo in its earlier stages of
development is not yet able to obtain its food and so build up the
frame, the necessary material has to be stored up in the ovum. Hence
we distinguish in the ova two chief elements--the active formative
yelk (protoplasm) and the passive food-yelk (deutoplasm, wrongly
spoken of as "the yelk"). In the little palingenetic ova, the
segmentation of which we have already considered, the yelk-granules
are so small and so regularly distributed in the protoplasm of the
ovum that the even and repeated cleavage is not affected by them. But
in the great majority of the animal ova the food-yelk is more or less
considerable, and is stored in a certain part of the ovum, so that
even in the unfertilised ovum the "granary" can clearly be
distinguished from the formative plasm. As a rule, the formative-yelk
(with the germinal vesicle) then usually gathers at one pole and the
food-yelk at the other. The first is the ANIMAL, and the second the
VEGETAL, pole of the vertical axis of the ovum.

(FIGURE 1.39. Gastrula of the amphioxus, seen from left side
(diagrammatic median section). (From Hatschek.) g primitive gut, u
primitive mouth, p peristomal pole-cells, i entoderm, e ectoderm, d
dorsal side, v ventral side.)

In these "telolecithal" ova, or ova with the yelk at one end (for
instance, in the cyclostoma and amphibia), the gastrulation then
usually takes place in such a way that in the cleavage of the
impregnated ovum the animal (usually the upper) half splits up more
quickly than the vegetal (lower). The contractions of the active
protoplasm, which effect this continual cleavage of the cells, meet a
greater resistance in the lower vegetal half from the passive
deutoplasm than in the upper animal half. Hence we find in the latter
more but smaller, and in the former fewer but larger, cells. The
animal cells produce the external, and the vegetal cells the internal,
germinal layer.

Although this unequal segmentation of the cyclostoma, ganoids, and
amphibia seems at first sight to differ from the original equal
segmentation (for instance, in the monoxenia, Figure 1.29), they both
have this in common, that the cleavage process throughout affects the
WHOLE cell; hence Remak called it TOTAL segmentation, and the ova in
question holoblastic, or "whole-cleaving." It is otherwise with the
second chief group of ova, which he distinguished from these as
meroblastic, or "partially-cleaving ": to this class belong the
familiar large eggs of birds and reptiles, and of most fishes. The
inert mass of the passive food-yelk is so large in these cases that
the protoplasmic contractions of the active yelk cannot effect any
further cleavage. In consequence, there is only a partial
segmentation. While the protoplasm in the animal section of the ovum
continues briskly to divide, multiplying the nuclei, the deutoplasm in
the vegetal section remains more or less undivided; it is merely
consumed as food by the forming cells. The larger the accumulation of
food, the more restricted is the process of segmentation. It may,
however, continue for some time (even after the gastrulation is more
or less complete) in the sense that the vegetal cell-nuclei
distributed in the deutoplasm slowly increase by cleavage; as each of
them is surrounded by a small quantity of protoplasm, it may
afterwards appropriate a portion of the food-yelk, and thus form a
real "yelk-cell" (merocyte). When this vegetal cell-formation
continues for a long time, after the two primary germinal layers have
been formed, it takes the name of the "after-segmentation."

The meroblastic ova are only found in the larger and more highly
developed animals, and only in those whose embryo needs a longer time
and richer nourishment within the foetal membranes. According as the
yelk-food accumulates at the centre or at the side of the ovum, we
distinguish two groups of dividing ova, periblastic and discoblastic.
In the periblastic the food-yelk is in the centre, enclosed inside the
ovum (hence they are also called "centrolecithal" ova): the formative
yelk surrounds the food-yelk, and so suffers itself a superficial
cleavage. This is found among the articulates (crabs, spiders,
insects, etc.). In the discoblastic ova the food-yelk gathers at one
side, at the vegetal or lower pole of the vertical axis, while the
nucleus of the ovum and the great bulk of the formative yelk lie at
the upper or animal pole (hence these ova are also called
"telolecithal"). In these cases the cleavage of the ovum begins at the
upper pole, and leads to the formation of a dorsal discoid embryo.
This is the case with all meroblastic vertebrates, most fishes, the
reptiles and birds, and the oviparous mammals (the monotremes).

The gastrulation of the discoblastic ova, which chiefly concerns us,
offers serious difficulties to microscopic investigation and
philosophic consideration. These, however, have been mastered by the
comparative embryological research which has been conducted by a
number of distinguished observers during the last few
decades--especially the brothers Hertwig, Rabl, Kupffer, Selenka,
Ruckert, Goette, Rauber, etc. These thorough and careful studies,
aided by the most perfect modern improvements in technical method (in
tinting and dissection), have given a very welcome support to the
views which I put forward in my work, On the Gastrula and the
Segmentation of the Animal Ovum [not translated], in 1875. As it is
very important to understand these views and their phylogenetic
foundation clearly, not only as regards evolution in general, but
particularly in connection with the genesis of man, I will give here a
brief statement of them as far as they concern the vertebrate-stem:--

1. All the vertebrates, including man, are phylogenetically (or
genealogically) related--that is, are members of one single natural
stem.

2. Consequently, the embryonic features in their individual
development must also have a genetic connection.

3. As the gastrulation of the amphioxus shows the original
palingenetic form in its simplest features, that of the other
vertebrates must have been derived from it.

4. The cenogenetic modifications of the latter are more appreciable
the more food-yelk is stored up in the ovum.

5. Although the mass of the food-yelk may be very large in the ova of
the discoblastic vertebrates, nevertheless in every case a blastula is
developed from the morula, as in the holoblastic ova.

6. Also, in every case, the gastrula develops from the blastula by
curving or invagination.

7. The cavity which is produced in the foetus by this curving is, in
each case, the primitive gut (progaster), and its opening the
primitive mouth (prostoma).

8. The food-yelk, whether large or small, is always stored in the
ventral wall of the primitive gut; the cells (called "merocytes")
which may be formed in it subsequently (by "after-segmentation") also
belong to the inner germinal layer, like the cells which immediately
enclose the primitive gut-cavity.

9. The primitive mouth, which at first lies below at the lower pole of
the vertical axis, is forced, by the growth of the yelk, backwards and
then upwards, towards the dorsal side of the embryo; the vertical axis
of the primitive gut is thus gradually converted into horizontal.

10. The primitive mouth is closed sooner or later in all the
vertebrates, and does not evolve into the permanent mouth-aperture; it
rather corresponds to the "properistoma," or region of the anus. From
this important point the formation of the middle germinal layer
proceeds, between the two primary layers.

The wide comparative studies of the scientists I have named have
further shown that in the case of the discoblastic higher vertebrates
(the three classes of amniotes) the primitive mouth of the embryonic
disc, which was long looked for in vain, is found always, and is
nothing else than the familiar "primitive groove." Of this we shall
see more as we proceed. Meantime we realise that gastrulation may be
reduced to one and the same process in all the vertebrates. Moreover,
the various forms it takes in the invertebrates can always be reduced
to one of the four types of segmentation described above. In relation
to the distinction between total and partial segmentation, the
grouping of the various forms is as follows:--

1. Palingenetic (primitive segmentation)

1.1. Equal segmentation (bell-gastrula).

1.1.A. Total segmentation (without independent food-yelk).

2. Cenogenetic segmentation (modified by adaptation).

2.2. Unequal segmentation (hooded gastrula).

2.2.A. Total segmentation (without independent food-yelk).

2.3. Discoid segmentation (discoid gastrula).

2.3.B. Partial segmentation (with independent food-yelk).

2.4. Superficial segmentation (spherical gastrula).

2.4.B. Partial segmentation (with independent food-yelk).

The lowest metazoa we know--namely, the lower zoophyta (sponges,
simple polyps, etc.)--remain throughout life at a stage of development
which differs little from the gastrula; their whole body consists of
two layers of cells. This is a fact of extreme importance. We see that
man, and also other vertebrates, pass quickly through a stage of
development in which they consist of two layers, just as these lower
zoophyta do throughout life. If we apply our biogenetic law to the
matter, we at once reach this important conclusion. "Man and all the
other animals which pass through the two-layer stage, or
gastrula-form, in the course of their embryonic development, must
descend from a primitive simple stem-form, the whole body of which
consisted throughout life (as is the case with the lower zoophyta
to-day) merely of two cell-strata or germinal layers." We will call
this primitive stem-form, with which we shall deal more fully later
on, the gastraea--that is to say, "primitive-gut animal."

According to this gastraea-theory there was originally in all the
multicellular animals ONE ORGAN with the same structure and function.
This was the primitive gut; and the two primary germinal layers which
form its wall must also be regarded as identical in all. This
important homology or identity of the primary germinal layers is
proved, on the one hand, from the fact that the gastrula was
originally formed in the same way in all cases--namely, by the curving
of the blastula; and, on the other hand, by the fact that in every
case the same fundamental organs arise from the germinal layers. The
outer or animal layer, or ectoderm, always forms the chief organs of
animal life--the skin, nervous system, sense-organs, etc.; the inner
or vegetal layer, or entoderm, gives rise to the chief organs of
vegetative life--the organs of nourishment, digestion,
blood-formation, etc.

In the lower zoophyta, whose body remains at the two-layer stage
throughout life, the gastraeads, the simplest sponges (Olynthus), and
polyps (Hydra), these two groups of functions, animal and vegetative,
are strictly divided between the two simple primary layers. Throughout
life the outer or animal layer acts simply as a covering for the body,
and accomplishes its movement and sensation. The inner or vegetative
layer of cells acts throughout life as a gut-lining, or nutritive
layer of enteric cells, and often also yields the reproductive cells.

The best known of these "gastraeads," or "gastrula-like animals," is
the common fresh-water polyp (Hydra). This simplest of all the
cnidaria has, it is true, a crown of tentacles round its mouth. Also
its outer germinal layer has certain special modifications. But these
are secondary additions, and the inner germinal layer is a simple
stratum of cells. On the whole, the hydra has preserved to our day by
heredity the simple structure of our primitive ancestor, the gastraea
(cf. Chapter 2.19.)

In all other animals, particularly the vertebrates, the gastrula is
merely a brief transitional stage. Here the two-layer stage of the
embryonic development is quickly succeeded by a three-layer, and then
a four-layer, stage. With the appearance of the four superimposed
germinal layers we reach again a firm and steady standing-ground, from
which we may follow the further, and much more difficult and
complicated, course of embryonic development.

SUMMARY OF THE CHIEF DIFFERENCES IN THE OVUM-SEGMENTATION AND
GASTRULATION OF ANIMALS.

The animal stems are indicated by the letters a-g: a Zoophyta. b
Annelida. c Mollusca.
d Echinoderma. e Articulata. f Tunicata. g Vertebrata.

1. Total Segmentation. Holoblastic ova. Gastrula without separate
food-yelk. Hologastrula.

1.1. Primitive Segmentation. Archiblastic ova. Bell-gastrula
(archigastrula.)
a. Many lower zoophyta (sponges, hydrapolyps, medusae, simpler
corals).
b. Many lower annelids (sagitta, phoronis, many nematoda, etc.,
terebratula, argiope, pisidium).
c. Some lower molluscs.
d. Many echinoderms.
e. A few lower articulata (some brachiopods, copepods: Tardigrades,
pteromalina).
f. Many tunicata.
g. The acrania (amphioxus).

1.2. Unequal Segmentation. Amphiblastic ova. Hooded-gastrula
(amphigastrula).
a. Many zoophyta (sponges, medusae, corals, siphonophorae,
ctenophora).
b. Most worms.
c. Most molluscs.
d. Many echinoderms (viviparous species and some others).
e. Some of the lower articulata (both crustacea and tracheata).
f. Many tunicata.
g. Cyclostoma, the oldest fishes, amphibia, mammals (not including
man).

2. Partial Segmentation. Meroblastic ova. Gastrula with separate
food-yelk. Merogastrula.

2.3. Discoid Segmentation. Discoblastic ova. Discoid gastrula.
c. Cephalopods or cuttlefish.
e. Many articulata, wood-lice, scorpions, etc.
g. Primitive fishes, bony fishes, reptiles, birds, monotremes.

2.4. Superficial Segmentation. Periblastic ova. Spherical-gastrula.
e. The great majority of the articulata (crustaceans, myriapods,
arachnids, insects).


CHAPTER 1.9. THE GASTRULATION OF THE VERTEBRATE.*

(* Cf. Balfour's Manual of Comparative Embryology volume 2; Theodore
Morgan's The Development of the Frog's Egg.)

The remarkable processes of gastrulation, ovum-segmentation, and
formation of germinal layers present a most conspicuous variety. There
is to-day only the lowest of the vertebrates, the amphioxus, that
exhibits the original form of those processes, or the palingenetic
gastrulation which we have considered in the preceding chapter, and
which culminates in the formation of the archigastrula (Figure 1.38).
In all other extant vertebrates these fundamental processes have been
more or less modified by adaptation to the conditions of embryonic
development (especially by changes in the food-yelk); they exhibit
various cenogenetic types of the formation of germinal layers.
However, the different classes vary considerably from each other. In
order to grasp the unity that underlies the manifold differences in
these phenomena and their historical connection, it is necessary to
bear in mind always the unity of the vertebrate-stem. This
"phylogenetic unity," which I developed in my General Morphology in
1866, is now generally admitted. All impartial zoologists agree to-day
that all the vertebrates, from the amphioxus and the fishes to the ape
and man, descend from a common ancestor, "the primitive vertebrate."
Hence the embryonic processes, by which each individual vertebrate is
developed, must also be capable of being reduced to one common type of
embryonic development; and this primitive type is most certainly
exhibited to-day by the amphioxus.

It must, therefore, be our next task to make a comparative study of
the various forms of vertebrate gastrulation, and trace them backwards
to that of the lancelet. Broadly speaking, they fall first into two
groups: the older cyclostoma, the earliest fishes, most of the
amphibia, and the viviparous mammals, have holoblastic ova--that is to
say, ova with total, unequal segmentation; while the younger
cyclostoma, most of the fishes, the cephalopods, reptiles, birds, and
monotremes, have meroblastic ova, or ova with partial discoid
segmentation. A closer study of them shows, however, that these two
groups do not present a natural unity, and that the historical
relations between their several divisions are very complicated. In
order to understand them properly, we must first consider the various
modifications of gastrulation in these classes. We may begin with that
of the amphibia.

The most suitable and most available objects of study in this class
are the eggs of our indigenous amphibia, the tailless frogs and toads,
and the tailed salamander. In spring they are to be found in clusters
in every pond, and careful examination of the ova with a lens is
sufficient to show at least the external features of the segmentation.
In order to understand the whole process rightly and follow the
formation of the germinal layers and the gastrula, the ova of the frog
and salamander must be carefully hardened; then the thinnest possible
sections must be made of the hardened ova with the microtome, and the
tinted sections must be very closely compared under a powerful
microscope.

The ova of the frog or toad are globular in shape, about the twelfth
of an inch in diameter, and are clustered in jelly-like masses, which
are lumped together in the case of the frog, but form long strings in
the case of the toad. When we examine the opaque, grey, brown, or
blackish ova closely, we find that the upper half is darker than the
lower. The middle of the upper half is in many species black, while
the middle of the lower half is white.* (* The colouring of the eggs
of the amphibia is caused by the accumulation of dark-colouring matter
at the animal pole of the ovum. In consequence of this, the animal
cells of the ectoderm are darker than the vegetal cells of the
entoderm. We find the reverse of this in the case of most animals, the
protoplasm of the entoderm cells being usually darker and
coarser-grained.) In this way we get a definite axis of the ovum with
two poles. To give a clear idea of the segmentation of this ovum, it
is best to compare it with a globe, on the surface of which are marked
the various parallels of longitude and latitude. The superficial
dividing lines between the different cells, which come from the
repeated segmentation of the ovum, look like deep furrows on the
surface, and hence the whole process has been given the name of
furcation. In reality, however, this "furcation," which was formerly
regarded as a very mysterious process, is nothing but the familiar,
repeated cell-segmentation. Hence also the segmentation-cells which
result from it are real cells.

(FIGURE 1.40. The cleavage of the frog's ovum (magnified ten times). A
stem-cell. B the first two segmentation-cells. C four cells. D eight
cells (4 animal and 4 vegetative). E twelve cells (8 animal and 4
vegetative). F sixteen cells (8 animal and 8 vegetative). G
twenty-four cells (16 animal and 8 vegetative). H thirty-two cells. I
forty-eight cells. K sixty-four cells. L ninety-six cells. M 160 cells
(128 animal and 32 vegetative).

(FIGURES 1.41 TO 1.44. Four vertical sections of the fertilised ovum
of the toad, in four successive stages of development. The letters
have the same meaning throughout: F segmentation-cavity. D covering of
same (D dorsal half of the embryo, P ventral half). P yelk-stopper
(white round field at the lower pole). Z yelk-cells of the entoderm
(Remak's "glandular embryo"). N primitive gut cavity (progaster or
Rusconian alimentary cavity). The primitive mouth (prostoma) is closed
by the yelk-stopper, P. s partition between the primitive gut cavity
(N) and the segmentation cavity (F). k k apostrophe, section of the
large circular lip-border of the primitive mouth (the Rusconian anus).
The line of dots between k and k apostrophe indicates the earlier
connection of the yelk-stopper (P) with the central mass of the
yelk-cells (Z). In Figure 1.44 the ovum has turned 90 degrees, so that
the back of the embryo is uppermost and the ventral side down. (From
Stricker.)).

The unequal segmentation which we observe in the ovum of the amphibia
has the special feature of beginning at the upper and darker pole (the
north pole of the terrestrial globe in our illustration), and slowly
advancing towards the lower and brighter pole (the south pole). Also
the upper and darker hemisphere remains in this position throughout
the course of the segmentation, and its cells multiply much more
briskly. Hence the cells of the lower hemisphere are found to be
larger and less numerous. The cleavage of the stem-cell (Figure 1.40
A) begins with the formation of a complete furrow, which starts from
the north pole and reaches to the south (B). An hour later a second
furrow arises in the same way, and this cuts the first at a right
angle (Figure 1.40 C). The ovum is thus divided into four equal parts.
Each of these four "segmentation cells" has an upper and darker and a
lower, brighter half. A few hours later a third furrow appears,
vertically to the first two (Figure 1.40 D). The globular germ now
consists of eight cells, four smaller ones above (northern) and four
larger ones below (southern). Next, each of the four upper ones
divides into two halves by a cleavage beginning from the north pole,
so that we now have eight above and four below (Figure 1.40 E). Later,
the four new longitudinal divisions extend gradually to the lower
cells, and the number rises from twelve to sixteen (F). Then a second
circular furrow appears, parallel to the first, and nearer to the
north pole, so that we may compare it to the north polar circle. In
this way we get twenty-four segmentation-cells--sixteen upper,
smaller, and darker ones, and eight smaller and brighter ones below
(G). Soon, however, the latter also sub-divide into sixteen, a third
or "meridian of latitude" appearing, this time in the southern
hemisphere: this makes thirty-two cells altogether (H). Then eight new
longitudinal lines are formed at the north pole, and these proceed to
divide, first the darker cells above and afterwards the lighter
southern cells, and finally reach the south pole. In this way we get
in succession forty, forty-eight, fifty-six, and at last sixty-four
cells (I, K). In the meantime, the two hemispheres differ more and
more from each other. Whereas the sluggish lower hemisphere long
remains at thirty-two cells, the lively northern hemisphere briskly
sub-divides twice, producing first sixty-four and then 128 cells (L,
M). Thus we reach a stage in which we count on the surface of the ovum
128 small cells in the upper half and thirty-two large ones in the
lower half, or 160 altogether. The dissimilarity of the two halves
increases: while the northern breaks up into a great number of small
cells, the southern consists of a much smaller number of larger cells.
Finally, the dark cells of the upper half grow almost over the surface
of the ovum, leaving only a small circular spot at the south pole,
where the large and clear cells of the lower half are visible. This
white region at the south pole corresponds, as we shall see
afterwards, to the primitive mouth of the gastrula. The whole mass of
the inner and larger and clearer cells (including the white polar
region) belongs to the entoderm or ventral layer. The outer envelope
of dark smaller cells forms the ectoderm or skin-layer.

In the meantime, a large cavity, full of fluid, has been formed within
the globular body--the segmentation-cavity or embryonic cavity
(blastocoel, Figures 1.41 to 1.44 F). It extends considerably as the
cleavage proceeds, and afterwards assumes an almost semi-circular form
(Figure 1.41 F). The frog-embryo now represents a modified embryonic
vesicle or blastula, with hollow animal half and solid vegetal half.

Now a second, narrower but longer, cavity arises by a process of
folding at the lower pole, and by the falling away from each other of
the white entoderm-cells (Figures 1.41 to 1.44 N). This is the
primitive gut-cavity or the gastric cavity of the gastrula, progaster
or archenteron. It was first observed in the ovum of the amphibia by
Rusconi, and so called the Rusconian cavity. The reason of its
peculiar narrowness here is that it is, for the most part, full of
yelk-cells of the entoderm. These also stop up the whole of the wide
opening of the primitive mouth, and form what is known as the
"yelk-stopper," which is seen freely at the white round spot at the
south pole (P). Around it the ectoderm is much thicker, and forms the
border of the primitive mouth, the most important part of the embryo
(Figure 1.44 k, k apostrophe). Soon the primitive gut-cavity stretches
further and further at the expense of the segmentation-cavity (F),
until at last the latter disappears altogether. The two cavities are
only separated by a thin partition (Figure 1.43 s). With the formation
of the primitive gut our frog-embryo has reached the gastrula stage,
though it is clear that this cenogenetic amphibian gastrula is very
different from the real palingenetic gastrula we have considered
(Figures 1.30 to 1.36).

In the growth of this hooded gastrula we cannot sharply mark off the
various stages which we distinguish successively in the bell-gastrula
as morula and gastrula. Nevertheless, it is not difficult to reduce
the whole cenogenetic or disturbed development of this amphigastrula
to the true palingenetic formation of the archigastrula of the
amphioxus.

(FIGURE 1.45. Blastula of the water-salamander (Triton). fh
segmentation-cavity, dz yelk-cells, rz border-zone. (From Hertwig.)

FIGURE 1.46. Embryonic vesicle of triton (blastula), outer view, with
the transverse fold of the primitive mouth (u). (From Hertwig.)

FIGURE 1.47. Sagittal section of a hooded-embryo (depula) of triton
(blastula at the commencement of gastrulation). ak outer germinal
layer, ik inner germinal layer, fh segmentation-cavity, ud primitive
gut, u primitive mouth, dl and vl dorsal and ventral lips of the
mouth, dz yelk-cells. (From Hertwig.))

This reduction becomes easier if, after considering the gastrulation
of the tailless amphibia (frogs and toads), we glance for a moment at
that of the tailed amphibia, the salamanders. In some of the latter,
that have only recently been carefully studied, and that are
phylogenetically older, the process is much simpler and clearer than
is the case with the former and longer known. Our common
water-salamander (Triton taeniatus) is a particularly good subject for
observation. Its nutritive yelk is much smaller and its formative yelk
less obscured with black pigment-cells than in the case of the frog;
and its gastrulation has better retained the original palingenetic
character. It was first described by Scott and Osborn (1879), and
Oscar Hertwig especially made a careful study of it (1881), and
rightly pointed out its great importance in helping us to understand
the vertebrate development. Its globular blastula (Figure 1.45)
consists of loosely-aggregated, yelk-filled entodermic cells or
yelk-cells (dz) in the lower vegetal half; the upper, animal half
encloses the hemispherical segmentation-cavity (fh), the curved roof
of which is formed of two or three strata of small ectodermic cells.
At the point where the latter pass into the former (at the equator of
the globular vesicle) we have the border zone (rz). The folding which
leads to the formation of the gastrula takes place at a spot in this
border zone, the primitive mouth (Figure 1.46 u).

Unequal segmentation takes place in some of the cyclostoma and in the
oldest fishes in just the same way as in most of the amphibia. Among
the cyclostoma ("round-mouthed") the familiar lampreys are
particularly interesting. In respect of organisation and development
they are half-way between the acrania (lancelet) and the lowest real
fishes (Selachii); hence I divided the group of the cyclostoma in 1886
from the real fishes with which they were formerly associated, and
formed of them a special class of vertebrates. The ovum-segmentation
in our common river-lamprey (Petromyzon fluviatilis) was described by
Max Schultze in 1856, and afterwards by Scott (1882) and Goette
(1890).

Unequal total segmentation follows the same lines in the oldest
fishes, the selachii and ganoids, which are directly descended from
the cyclostoma. The primitive fishes (Selachii), which we must regard
as the ancestral group of the true fishes, were generally considered,
until a short time ago, to be discoblastic. It was not until the
beginning of the twentieth century that Bashford Dean made the
important discovery in Japan that one of the oldest living fishes of
the shark type (Cestracion japonicus) has the same total unequal
segmentation as the amphiblastic plated fishes (ganoides).* (*
Bashford Dean, Holoblastic Cleavage in the Egg of a Shark, Cestracion
japonicus Macleay. Annotationes zoologicae japonenses, volume 4 Tokio
1901.) This is particularly interesting in connection with our
subject, because the few remaining survivors of this division, which
was so numerous in paleozoic times, exhibit three different types of
gastrulation. The oldest and most conservative forms of the modern
ganoids are the scaly sturgeons (Sturiones), plated fishes of great
evolutionary importance, the eggs of which are eaten as caviar; their
cleavage is not essentially different from that of the lampreys and
the amphibia. On the other hand, the most modern of the plated fishes,
the beautifully scaled bony pike of the North American rivers
(Lepidosteus), approaches the osseous fishes, and is discoblastic like
them. A third genus (Amia) is midway between the sturgeons and the
latter.

(FIGURE 1.48. Sagittal section of the gastrula of the water-salamander
(Triton). (From Hertwig.) Letters as in Figure 1.47; except--p
yelk-stopper, mk beginning of the middle germinal layer.)

The group of the lung-fishes (Dipneusta or Dipnoi) is closely
connected with the older ganoids. In respect of their whole
organisation they are midway between the gill-breathing fishes and the
lung-breathing amphibia; they share with the former the shape of the
body and limbs, and with the latter the form of the heart and lungs.
Of the older dipnoi (Paladipneusta) we have now only one specimen, the
remarkable Ceratodus of East Australia; its amphiblastic gastrulation
has been recently explained by Richard Semon (cf. Chapter 2.21). That
of the two modern dipneusta, of which Protopterus is found in Africa
and Lepidosiren in America, is not materially different. (Cf. Figure
1.51.)

(FIGURE 1.49. Ovum-segmentation of the lamprey (Petromyzon
fluviatalis), in four successive stages. The small cells of the upper
(animal) hemisphere divide much more quickly than the cells of the
lower (vegetal) hemisphere.

FIGURE 1.50. Gastrulation of the lamprey (Petromyzon fluviatilis). A
blastula, with wide embryonic cavity (blastocoel, bl), g incipient
invagination. B depula, with advanced invagination, from the primitive
mouth (g). C gastrula, with complete primitive gut: the embryonic
cavity has almost disappeared in consequence of invagination.)

All these amphiblastic vertebrates, Petromyzon and Cestracion,
Accipenser and Ceratodus, and also the salamanders and batrachia,
belong to the old, conservative groups of our stem. Their unequal
ovum-segmentation and gastrulation have many peculiarities in detail,
but can always be reduced with comparative ease to the original
cleavage and gastrulation of the lowest vertebrate, the amphioxus; and
this is little removed, as we have seen, from the very simple
archigastrula of the Sagitta and Monoxenia (see Figures 1.29 to 1.36).
All these and many other classes of animals generally agree in the
circumstance that in segmentation their ovum divides into a large
number of cells by repeated cleavage. All such ova have been called,
after Remak, "whole-cleaving" (holoblasta), because their division
into cells is complete or total.

(FIGURE 1.51. Gastrulation of ceratodus (from Semon). A and C stage
with four cells, B and D with sixteen cells. A and B are seen from
above, C and D sideways. E stage with thirty-two
cells; F blastula; G gastrula in longitudinal section. fh
segmentation-cavity. gh primitive gut or gastric cavity.)

In a great many other classes of animals this is not the case, as we
find (in the vertebrate stem) among the birds, reptiles, and most of
the fishes; among the insects and most of the spiders and crabs (of
the articulates); and the cephalopods (of the molluscs). In all these
animals the mature ovum, and the stem-cell that arises from it in
fertilisation, consist of two different and separate parts, which we
have called formative yelk and nutritive yelk. The formative yelk
alone consists of living protoplasm, and is the active, evolutionary,
and nucleated part of the ovum; this alone divides in segmentation,
and produces the numerous cells which make up the embryo. On the other
hand, the nutritive yelk is merely a passive part of the contents of
the ovum, a subordinate element which contains nutritive material
(albumin, fat, etc.), and so represents in a sense the provision-store
of the developing embryo. The latter takes a quantity of food out of
this store, and finally consumes it all. Hence the nutritive yelk is
of great indirect importance in embryonic development, though it has
no direct share in it. It either does not divide at all, or only later
on, and does not generally consist of cells. It is sometimes large and
sometimes small, but generally many times larger than the formative
yelk; and hence it is that it was formerly thought the more important
of the two. As the respective significance of these two parts of the
ovum is often wrongly described, it must be borne in mind that the
nutritive yelk is only a secondary addition to the primary cell, it is
an inner enclosure, not an external appendage. All ova that have this
independent nutritive yelk are called, after Remak,
"partially-cleaving" (meroblasta). Their segmentation is incomplete or
partial.

(FIGURE 1.52. Ovum of a deep-sea bony fish. b protoplasm of the
stem-cell, k nucleus of same, d clear globule of albumin, the
nutritive yelk, f fat-globule of same, c outer membrane of the ovum,
or ovolemma.)

There are many difficulties in the way of understanding this partial
segmentation and the gastrula that arises from it. We have only
recently succeeded, by means of comparative research, in overcoming
these difficulties, and reducing this cenogenetic form of gastrulation
to the original palingenetic type. This is comparatively easy in the
small meroblastic ova which contain little nutritive yelk--for
instance, in the marine ova of a bony fish, the development of which I
observed in 1875 at Ajaccio in Corsica. I found them joined together
in lumps of jelly, floating on the surface of the sea; and, as the
little ovula were completely transparent, I could easily follow the
development of the germ step by step. These ovula are glossy and
colourless globules of little more than the 50th of an inch. Inside a
structureless, thin, but firm membrane (ovolemma, Figure 1.52 c) we
find a large, quite clear, and transparent globule of albumin (d). At
both poles of its axis this globule has a pit-like depression. In the
pit at the upper, animal pole (which is turned downwards in the
floating ovum) there is a bi-convex lens composed of protoplasm, and
this encloses the nucleus (k); this is the formative yelk of the
stem-cell, or the germinal disk (b). The small fat-globule (f) and the
large albumin-globule (d) together form the nutritive yelk. Only the
formative yelk undergoes cleavage, the nutritive yelk not dividing at
all at first.

The segmentation of the lens-shaped formative yelk (b) proceeds quite
independently of the nutritive yelk, and in perfect geometrical order.

When the mulberry-like cluster of cells has been formed, the
border-cells of the lens separate from the rest and travel into the
yelk and the border-layer. From this the blastula is developed; the
regular bi-convex lens being converted into a disk, like a
watch-glass, with thick borders. This lies on the upper and less
curved polar surface of the nutritive yelk like the watch glass on the
yelk. Fluid gathers between the outer layer and the border, and the
segmentation-cavity is formed. The gastrula is then formed by
invagination, or a kind of turning-up of the edge of the blastoderm.
In this process the segmentation-cavity disappears.

The space underneath the entoderm corresponds to the primitive
gut-cavity, and is filled with the decreasing food-yelk (n). Thus the
formation of the gastrula of our fish is complete. In contrast to the
two chief forms of gastrula we considered previously, we give the name
of discoid gastrula (discogastrula, Figure 1.54) to this third
principal type.

Very similar to the discoid gastrulation of the bony fishes is that of
the hags or myxinoida, the remarkable cyclostomes that live
parasitically in the body-cavity of fishes, and are distinguished by
several notable peculiarities from their nearest relatives, the
lampreys. While the amphiblastic ova of the latter are small and
develop like those of the amphibia, the cucumber-shaped ova of the hag
are about an inch long, and form a discoid gastrula. Up to the present
it has only been observed in one species (Bdellostoma Stouti), by Dean
and Doflein (1898).

It is clear that the important features which distinguish the discoid
gastrula from the other chief forms we have considered are determined
by the large food-yelk. This takes no direct part in the building of
the germinal layers, and completely fills the primitive gut-cavity of
the gastrula, even protruding at the mouth-opening. If we imagine the
original bell-gastrula (Figures 1.30 to 1.36) trying to swallow a ball
of food which is much bigger than itself, it would spread out round it
in discoid shape in the attempt, just as we find to be the case here
(Figure 1.54). Hence we may derive the discoid gastrula from the
original bell-gastrula, through the intermediate stage of the hooded
gastrula. It has arisen through the accumulation of a store of
food-stuff at the vegetal pole, a "nutritive yelk" being thus formed
in contrast to the "formative yelk." Nevertheless, the gastrula is
formed here, as in the previous cases, by the folding or invagination
of the blastula. We can, therefore, reduce this cenogenetic form of
the discoid segmentation to the palingenetic form of the primitive
cleavage.

(FIGURE 1.53. Ovum-segmentation of a bony fish. A first cleavage of
the stem-cell (cytula), B division of same into four
segmentation-cells (only two visible), C the germinal disk divides
into the blastoderm (b) and the periblast (p). d nutritive yelk, f
fat-globule, c ovolemma, z space between the ovolemma and the ovum,
filled with a clear fluid.)

This reduction is tolerably easy and confident in the case of the
small ovum of our deep-sea bony fish, but it becomes difficult and
uncertain in the case of the large ova that we find in the majority of
the other fishes and in all the reptiles and birds. In these cases the
food-yelk is, in the first place, comparatively colossal, the
formative yelk being almost invisible beside it; and, in the second
place, the food-yelk contains a quantity of different elements, which
are known as "yelk-granules, yelk-globules, yelk-plates, yelk-flakes,
yelk-vesicles," and so on. Frequently these definite elements in the
yelk have been described as real cells, and it has been wrongly stated
that a portion of the embryonic body is built up from these cells.
This is by no means the case. In every case, however large it is--and
even when cell-nuclei travel into it during the cleavage of the
border--the nutritive yelk remains a dead accumulation of food, which
is taken into the gut during embryonic development and consumed by the
embryo. The latter develops solely from the living formative yelk of
the stem-cell. This is equally true of the ova of our small bony
fishes and of the colossal ova of the primitive fishes, reptiles, and
birds.

(FIGURE 1.54. Discoid gastrula (discogastrula) of a bony fish. e
ectoderm, i entoderm, w border-swelling or primitive mouth, n
albuminous globule of the nutritive yelk, f fat-globule of same, c
external membrane (ovolemma), d partition between entoderm and
ectoderm (earlier the segmentation-cavity).)

The gastrulation of the primitive fishes or selachii (sharks and rays)
has been carefully studied of late years by Ruckert, Rabl, and H.E.
Ziegler in particular, and is very important in the sense that this
group is the oldest among living fishes, and their gastrulation can be
derived directly from that of the cyclostoma by the accumulation of a
large quantity of food-yelk. The oldest sharks (Cestracion) still have
the unequal segmentation inherited from the cyclostoma. But while in
this case, as in the case of the amphibia, the small ovum completely
divides into cells in segmentation, this is no longer so in the great
majority of the selachii (or Elasmobranchii). In these the
contractility of the active protoplasm no longer suffices to break up
the huge mass of the passive deutoplasm completely into cells; this is
only possible in the upper or dorsal part, but not in the lower or
ventral section. Hence we find in the primitive fishes a blastula with
a small eccentric segmentation-cavity (Figure 1.55 b), the wall of
which varies greatly in composition. The circular border of the
germinal disk which connects the roof and floor of the
segmentation-cavity corresponds to the border-zone at the equator of
the amphibian ovum. In the middle of its hinder border we have the
beginning of the invagination of the primitive gut (Figure 1.56 ud);
it extends gradually from this spot (which corresponds to the
Rusconian anus of the amphibia) forward and around, so that the
primitive mouth becomes first crescent-shaped and then circular, and,
as it opens wider, surrounds the ball of the larger food-yelk.

Essentially different from the wide-mouthed discoid gastrula of most
of the selachii is the narrow-mouthed discoid gastrula (or
epigastrula) of the amniotes, the reptiles, birds, and monotremes;
between the two--as an intermediate stage--we have the amphigastrula
of the amphibia. The latter has developed from the amphigastrula of
the ganoids and dipneusts, whereas the discoid amniote gastrula has
been evolved from the amphibian gastrula by the addition of food-yelk.
This change of gastrulation is still found in the remarkable ophidia
(Gymnophiona, Coecilia, or Peromela), serpent-like amphibia that live
in moist soil in the tropics, and in many respects represent the
transition from the gill-breathing amphibia to the lung-breathing
reptiles. Their embryonic development has been explained by the fine
studies of the brothers Sarasin of Ichthyophis glutinosa at Ceylon
(1887), and those of August Brauer of the Hypogeophis rostrata in the
Seychelles (1897). It is only by the historical and comparative study
of these that we can understand the difficult and obscure gastrulation
of the amniotes.

The bird's egg is particularly important for our purpose, because most
of the chief studies of the development of the vertebrates are based
on observations of the hen's egg during hatching. The mammal ovum is
much more difficult to obtain and study, and for this practical and
obvious reason very rarely thoroughly investigated. But we can get
hens' eggs in any quantity at any time, and, by means of artificial
incubation, follow the development of the embryo step by step. The
bird's egg differs considerably from the tiny mammal ovum in size, a
large quantity of food-yelk accumulating within the original yelk or
the protoplasm of the ovum. This is the yellow ball which we commonly
call the yolk of the egg. In order to understand the bird's egg
aright--for it is very often quite wrongly explained--we must examine
it in its original condition, and follow it from the very beginning of
its development in the bird's ovary. We then see that the original
ovum is a quite small, naked, and simple cell with a nucleus, not
differing in either size or shape from the original ovum of the
mammals and other animals (cf. Figure 1.13 E). As in the case of all
the craniota (animals with a skull), the original or primitive ovum
(protovum) is covered with a continuous layer of small cells. This
membrane is the follicle, from which the ovum afterwards issues.
Immediately underneath it the structureless yelk-membrane is secreted
from the yelk.

(FIGURE 1.55. Longitudinal section through the blastula of a shark
(Pristiuris). (From Ruckert.) (Looked at from the left; to the right
is the hinder end, H, to the left the fore end, V.) B
segmentation-cavity, kz cells of the germinal membrane, dk
yelk-nuclei.

FIGURE 1.56. Longitudinal section of the blastula of a shark
(Pristiurus) at the beginning of gastrulation. (From Ruckert.) (Seen
from the left.) V fore end, H hind end, B segmentation-cavity, ud
first trace of the primitive gut, dk yelk-nuclei, fd fine-grained
yelk, gd coarse-grained yelk.)

The small primitive ovum of the bird begins very early to take up into
itself a quantity of food-stuff through the yelk-membrane, and work it
up into the "yellow yelk." In this way the ovum enters on its second
stage (the metovum), which is many times larger than the first, but
still only a single enlarged cell. Through the accumulation of the
store of yellow yelk within the ball of protoplasm the nucleus it
contains (the germinal vesicle) is forced to the surface of the ball.
Here it is surrounded by a small quantity of protoplasm, and with this
forms the lens-shaped formative yelk (Figure 1.15 b). This is seen on
the yellow yelk-ball, at a certain point of the surface, as a small
round white spot--the "tread" (cicatricula). From this point a
thread-like column of white nutritive yelk (d), which contains no
yellow yelk-granules, and is softer than the yellow food-yelk,
proceeds to the middle of the yellow yelk-ball, and forms there a
small central globule of white yelk (Figure 1.15 d). The whole of this
white yelk is not sharply separated from the yellow yelk, which shows
a slight trace of concentric layers in the hard-boiled egg (Figure
1.15 c). We also find in the hen's egg, when we break the shell and
take out the yelk, a round small white disk at its surface which
corresponds to the tread. But this small white "germinal disk" is now
further developed, and is really the gastrula of the chick. The body
of the chick is formed from it alone. The whole white and yellow
yelk-mass is without any significance for the formation of the embryo,
it being merely used as food by the developing chick. The clear,
glarous mass of albumin that surrounds the yellow yelk of the bird's
egg, and also the hard chalky shell, are only formed within the
oviduct round the impregnated ovum.

When the fertilisation of the bird's ovum has taken place within the
mother's body, we find in the lens-shaped stem-cell the progress of
flat, discoid segmentation (Figure 1.57). First two equal
segmentation-cells (A) are formed from the ovum. These divide into
four (B), then into eight, sixteen (C), thirty-two, sixty-four, and so
on. The cleavage of the cells is always preceded by a division of
their nuclei. The cleavage surfaces between the segmentation-cells
appear at the free surface of the tread as clefts. The first two
divisions are vertical to each other, in the form of a cross (B). Then
there are two more divisions, which cut the former at an angle of
forty-five degrees. The tread, which thus becomes the germinal disk,
now has the appearance of an eight-rayed star. A circular cleavage
next taking place round the middle, the eight triangular cells divide
into sixteen, of which eight are in the middle and eight distributed
around (C). Afterwards circular clefts and radial clefts, directed
towards the centre, alternate more or less irregularly (D, E). In most
of the amniotes the formation of concentric and radial clefts is
irregular from the very first; and so also in the hen's egg. But the
final outcome of the cleavage-process is once more the formation of a
large number of small cells of a similar nature. As in the case of the
fish-ovum, these segmentation-cells form a round, lens-shaped disk,
which corresponds to the morula, and is embedded in a small depression
of the white yelk. Between the lens-shaped disk of the morula-cells
and the underlying white yelk a small cavity is now formed by the
accumulation of fluid, as in the fishes. Thus we get the peculiar and
not easily recognisable blastula of the bird (Figure 1.58). The small
segmentation-cavity (fh) is very flat and much compressed. The upper
or dorsal wall (dw) is formed of a single layer of clear, distinctly
separated cells; this corresponds to the upper or animal hemisphere of
the triton-blastula (Figure 1.45). The lower or ventral wall of the
flat dividing space (vw) is made up of larger and darker
segmentation-cells; it corresponds to the lower or vegetal hemisphere
of the blastula of the water-salamander (Figure 1.45 dz). The nuclei
of the yelk-cells, which are in this case especially numerous at the
edge of the lens-shaped blastula, travel into the white yelk, increase
by cleavage, and contribute even to the further growth of the germinal
disk by furnishing it with food-stuff.

(FIGURE 1.57. Diagram of discoid segmentation in the bird's ovum
(magnified about ten times). Only the formative yelk (the tread) is
shown in these six figures (A to F), because cleavage only takes place
in this. The much larger food-yelk, which does not share in the
cleavage, is left out and merely indicated by the dark ring without.)

The invagination or the folding inwards of the bird-blastula takes
place in this case also at the hinder pole of the subsequent chief
axis, in the middle of the hind border of the round germinal disk
(Figure 1.59 s). At this spot we have the most brisk cleavage of the
cells; hence the cells are more numerous and smaller here than in the
fore-half of the germinal disk. The border-swelling or thick edge of
the disk is less clear but whiter behind, and is more sharply
separated from contiguous parts. In the middle of its hind border
there is a white, crescent-shaped groove--Koller's sickle-groove (Fig
1.59 s); a small projecting process in the centre of it is called the
sickle-knob (sk). This important cleft is the primitive mouth, which
was described for a long time as the "primitive groove." If we make a
vertical section through this part, we see that a flat and broad cleft
stretches under the germinal disk forwards from the primitive mouth;
this is the primitive gut (Figure 1.60 ud). Its roof or dorsal wall is
formed by the folded upper part of the blastula, and its floor or
ventral wall by the white yelk (wd), in which a number of yelk-nuclei
(dk) are distributed. There is a brisk multiplication of these at the
edge of the germinal disk, especially in the neighbourhood of the
sickle-shaped primitive mouth.

We learn from sections through later stages of this discoid
bird-gastrula that the primitive gut-cavity, extending forward from
the primitive mouth as a flat pouch, undermines the whole region of
the round flat lens-shaped blastula (Figure 1.61 ud). At the same
time, the segmentation-cavity gradually disappears altogether, the
folded inner germinal layer (ik) placing itself from underneath on the
overlying outer germinal layer (ak). The typical process of
invagination, though greatly disguised, can thus be clearly seen in
this case, as Goette and Rauber, and more recently Duval (Figure
1.61), have shown.

(FIGURE 1.58. Vertical section of the blastula of a hen
(discoblastula). fh segmentation-cavity, dw dorsal wall of same, vw
ventral wall, passing directly into the white yelk (wd) (From Duval.)

FIGURE 1.59. The germinal disk of the hen's ovum at the beginning of
gastrulation; A before
incubation, B in the first hour of incubation. (From Koller.) ks
germinal-disk, V its fore and H its hind border; es embryonic shield,
s sickle-groove, sk sickle knob, d yelk.

FIGURE 1.60. Longitudinal section of the germinal disk of a siskin
(discogastrula). (From Duval.) ud primitive gut, vl, hl fore and hind
lips of the primitive mouth (or sickle-edge); ak outer germinal layer,
ik inner germinal layer, dk yelk-nuclei, wd white yelk.

FIGURE 1.61. Longitudinal section of the discoid gastrula of the
nightingale. (From Duval.) ud primitive gut, vl, hl fore and hind lips
of the primitive mouth; ak, ik outer and inner germinal layers; vr
fore-border of the discogastrula.)

The older embryologists (Pander, Baer, Remak), and, in recent times
especially, His, Kolliker, and others, said that the two primary
germinal layers of the hen's ovum--the oldest and most frequent
subject of observation!--arose by horizontal cleavage of a simple
germinal disk. In opposition to this accepted view, I affirmed in my
Gastraea Theory (1873) that the discoid bird-gastrula, like that of
all other vertebrates, is formed by folding (or invagination), and
that this typical process is merely altered in a peculiar way and
disguised by the immense accumulation of food-yelk and the flat
spreading of the discoid blastula at one part of its surface. I
endeavoured to establish this view by the derivation of the
vertebrates from one source, and especially by proving that the birds
descend from the reptiles, and these from the amphibia. If this is
correct, the discoid gastrula of the amniotes must have been formed by
the folding-in of a hollow blastula, as has been shown by Remak and
Rusconi of the discoid gastrula of the amphibia, their direct
ancestors. The accurate and extremely careful observations of the
authors I have mentioned (Goette, Rauber, and Duval) have decisively
proved this recently for the birds; and the same has been done for the
reptiles by the fine studies of Kupffer, Beneke, Wenkebach, and
others. In the shield-shaped germinal disk of the lizard (Figure
1.62), the crocodile, the tortoise, and other reptiles, we find in the
middle of the hind border (at the same spot as the sickle groove in
the bird) a transverse furrow (u), which leads into a flat,
pouch-like, blind sac, the primitive gut. The fore (dorsal) and hind
(ventral) lips of the transverse furrow correspond exactly to the lips
of the primitive mouth (or sickle-groove) in the birds.

(FIGURE 1.62. Germinal disk of the lizard (Lacerta agilis). (From
Kupffer.) u primitive mouth, s sickle, es embryonic shield, hf and df
light and dark germinative area.)

The gastrulation of the mammals must be derived from this special
embryonic development of the reptiles and birds. This latest and most
advanced class of the vertebrates has, as we shall see afterwards,
evolved at a comparatively recent date from an older group of
reptiles; and all these amniotes must have come originally from a
common stem-form. Hence the distinctive embryonic process of the
mammal must have arisen by cenogenetic modifications from the older
form of gastrulation of the reptiles and birds. Until we admit this
thesis we cannot understand the formation of the germinal layers in
the mammal, and therefore in man.

I first advanced this fundamental principle in my essay On the
Gastrulation of Mammals (1877), and sought to show in this way that I
assumed a gradual degeneration of the food-yelk and the yelk-sac on
the way from the proreptiles to the mammals. "The cenogenetic process
of adaptation," I said, "which has occasioned the atrophy of the
rudimentary yelk-sac of the mammal, is perfectly clear. It is due to
the fact that the young of the mammal, whose ancestors were certainly
oviparous, now remain a long time in the womb. As the great store of
food-yelk, which the oviparous ancestors gave to the egg, became
superfluous in their descendants owing to the long carrying in the
womb, and the maternal blood in the wall of the uterus made itself the
chief source of nourishment, the now useless yelk-sac was bound to
atrophy by embryonic adaptation."

My opinion met with little approval at the time; it was vehemently
attacked by Kolliker, Hensen, and His in particular. However, it has
been gradually accepted, and has recently been firmly established by a
large number of excellent studies of mammal gastrulation, especially
by Edward Van Beneden's studies of the rabbit and bat, Selenka's on
the marsupials and rodents, Heape's and Lieberkuhn's on the mole,
Kupffer and Keibel's on the rodents, Bonnet's on the ruminants, etc.
From the general comparative point of view, Carl Rabl in his theory of
the mesoderm, Oscar Hertwig in the latest edition of his Manual
(1902), and Hubrecht in his Studies in Mammalian Embryology (1891),
have supported the opinion, and sought to derive the peculiarly
modified gastrulation of the mammal from that of the reptile.

(FIGURE 1.63. Ovum of the opossum (Didelphys) divided into four. (From
Selenka.) b the four segmentation-cells, r directive body, c
unnucleated coagulated matter, p, albumin-membrane.)

In the meantime (1884) the studies of Wilhelm Haacke and Caldwell
provided a proof of the long-suspected and very interesting fact, that
the lowest mammals, the monotremes, LAY EGGS, like the birds and
reptiles, and are not viviparous like the other mammals. Although the
gastrulation of the monotremes was not really known until studied by
Richard Semon in 1894, there could be little doubt, in view of the
great size of their food-yelk, that their ovum-segmentation was
discoid, and led to the formation of a sickle-mouthed discogastrula,
as in the case of the reptiles and birds. Hence I had, in 1875 (in my
essay on The Gastrula and Ovum-segmentation of Animals), counted the
monotremes among the discoblastic vertebrates. This hypothesis was
established as a fact nineteen years afterwards by the careful
observations of Semon; he gave in the second volume of his great work,
Zoological Journeys in Australia (1894), the first description and
correct explanation of the discoid gastrulation of the monotremes. The
fertilised ova of the two living monotremes (Echidna and
Ornithorhynchus) are balls of one-fifth of an inch in diameter,
enclosed in a stiff shell; but they grow considerably during
development, so that when laid the egg is three times as large. The
structure of the plentiful yelk, and especially the relation of the
yellow and the white yelk, are just the same as in the reptiles and
birds. As with these, partial cleavage takes place at a spot on the
surface at which the small formative yelk and the nucleus it encloses
are found. First is formed a lens-shaped circular germinal disk. This
is made up of several strata of cells, but it spreads over the
yelk-ball, and thus becomes a one-layered blastula. If we then imagine
the yelk it contains to be dissolved and replaced by a clear liquid,
we have the characteristic blastula of the higher mammals. In these
the gastrulation proceeds in two phases, as Semon rightly observes:
firstly, formation of the entoderm by cleavage at the centre and
further growth at the edge; secondly, invagination. In the monotremes
more primitive conditions have been retained better than in the
reptiles and birds. In the latter, before the commencement of the
gastrula-folding, we have, at least at the periphery, a two-layered
embryo forming from the cleavage. But in the monotremes the formation
of the cenogenetic entoderm does not precede the invagination; hence
in this case the construction of the germinal layers is less modified
than in the other amniota.

The marsupials, a second sub-class, come next to the oviparous
monotremes, the oldest of the mammals. But as in their case the
food-yelk is already atrophied, and the little ovum develops within
the mother's body, the partial cleavage has been reconverted into
total. One section of the marsupials still show points of agreement
with the monotremes, while another section of them, according to the
splendid investigations of Selenka, form a connecting-link between
these and the placentals.

(FIGURE 1.64. Blastula of the opossum (Didelphys). (From Selenka.) a
animal pole of the blastula, v vegetal pole, en mother-cell of the
entoderm, ex ectodermic cells, s spermia, ib unnucleated yelk-balls
(remainder of the food-yelk), p albumin membrane.)

The fertilised ovum of the opossum (Didelphys) divides, according to
Selenka, first into two, then four, then eight equal cells; hence the
segmentation is at first equal or homogeneous. But in the course of
the cleavage a larger cell, distinguished by its less clear plasm and
its containing more yelk-granules (the mother cell of the entoderm,
Figure 1.64 en), separates from the others; the latter multiply more
rapidly than the former. As, further, a quantity of fluid gathers in
the morula, we get a round blastula, the wall of which is of varying
thickness, like that of the amphioxus (Figure 1.38 E) and the amphibia
(Figure 1.45). The upper or animal hemisphere is formed of a large
number of small cells; the lower or vegetal hemisphere of a small
number of large cells. One of the latter, distinguished by its size
(Figure 1.64 en), lies at the vegetal pole of the blastula-axis, at
the point where the primitive mouth afterwards appears. This is the
mother-cell of the entoderm; it now begins to multiply by cleavage,
and the daughter-cells (Figure 1.65 i) spread out from this spot over
the inner surface of the blastula, though at first only over the
vegetal hemisphere. The less clear entodermic cells (i) are
distinguished at first by their rounder shape and darker nuclei from
the higher, clearer, and longer entodermic cells (e), afterwards both
are greatly flattened, the inner blastodermic cells more than the
outer.

(FIGURE 1.65. Blastula of the opossum (Didelphys) at the beginning of
gastrulation. (From Selenka.) e ectoderm, i entoderm; a animal pole, u
primitive mouth at the vegetal pole, f segmentation-cavity, d
unnucleated yelk-balls (relics of the reduced food-yelk), c nucleated
curd (without yelk-granules).

FIGURE 1.66. Oval gastrula of the opossum (Didelphys), about eight
hours old. (From Selenka) (external view).)

The unnucleated yelk-balls and curd (Figure 1.65 d) that we find in
the fluid of the blastula in these marsupials are very remarkable;
they are the relics of the atrophied food-yelk, which was developed in
their ancestors, the monotremes, and in the reptiles.

In the further course of the gastrulation of the opossum the oval
shape of the gastrula (Figure 1.66) gradually changes into globular, a
larger quantity of fluid accumulating in the vesicle. At the same
time, the entoderm spreads further and further over the inner surface
of the ectoderm (e). A globular vesicle is formed, the wall of which
consists of two thin simple strata of cells; the cells of the outer
germinal layer are rounder, and those of the inner layer flatter. In
the region of the primitive mouth (p) the cells are less flattened,
and multiply briskly. From this point--from the hind (ventral) lip of
the primitive mouth, which extends in a central cleft, the primitive
groove--the construction of the mesoderm proceeds.

Gastrulation is still more modified and curtailed cenogenetically in
the placentals than in the marsupials. It was first accurately known
to us by the distinguished investigations of Edward Van Beneden in
1875, the first object of study being the ovum of the rabbit. But as
man also belongs to this sub-class, and as his as yet unstudied
gastrulation cannot be materially different from that of the other
placentals, it merits the closest attention. We have, in the first
place, the peculiar feature that the two first segmentation-cells that
proceed from the cleavage of the fertilised ovum (Figure 1.68) are of
different sizes and natures; the difference is sometimes greater,
sometimes less (Figure 1.69). One of these first daughter-cells of the
ovum is a little larger, clearer, and more transparent than the other.
Further, the smaller cell takes a colour in carmine, osmium, etc.,
more strongly than the larger. By repeated cleavage of it a morula is
formed, and from this a blastula, which changes in a very
characteristic way into the greatly modified gastrula. When the number
of the segmentation-cells in the mammal embryo has reached ninety-six
(in the rabbit, about seventy hours after impregnation) the foetus
assumes a form very like the archigastrula (Figure 1.72). The
spherical embryo consists of a central mass of thirty-two soft, round
cells with dark nuclei, which are flattened into polygonal shape by
mutual pressure, and colour dark-brown with osmic acid (Figure 1.72
i). This dark central group of cells is surrounded by a lighter
spherical membrane, consisting of sixty-four cube-shaped, small, and
fine-grained cells which lie close together in a single stratum, and
only colour slightly in osmic acid (Figure 1.72 e). The authors who
regard this embryonic form as the primary gastrula of the placental
conceive the outer layer as the ectoderm and the inner as the
entoderm. The entodermic membrane is only interrupted at one spot,
one, two, or three of the ectodermic cells being loose there. These
form the yelk-stopper, and fill up the mouth of the gastrula (a). The
central primitive gut-cavity (d) is full of entodermic cells. The
uni-axial type of the mammal gastrula is accentuated in this way.
However, opinions still differ considerably as to the real nature of
this "provisional gastrula" of the placental and its relation to the
blastula into which it is converted.

As the gastrulation proceeds a large spherical blastula is formed from
this peculiar solid amphigastrula of the placental, as we saw in the
case of the marsupial. The accumulation of fluid in the solid gastrula
(Figure 1.73 A) leads to the formation of an eccentric cavity, the
group of the darker entodermic cells (hy) remaining directly attached
at one spot with the round enveloping stratum of the lighter
ectodermic cells (ep). This spot corresponds to the original primitive
mouth (prostoma or blastoporus). From this important spot the inner
germinal layer spreads all round on the inner surface of the outer
layer, the cell-stratum of which forms the wall of the hollow sphere;
the extension proceeds from the vegetal towards the animal pole.

(FIGURE 1.67. Longitudinal section through the oval gastrula of the
opossum (Figure 1.69). (From Selenka.) p primitive mouth, e ectoderm,
i entoderm, d yelk remains in the primitive gut-cavity (u).)

The cenogenetic gastrulation of the placental has been greatly
modified by secondary adaptation in the various groups of this most
advanced and youngest sub-class of the mammals. Thus, for instance, we
find in many of the rodents (guinea-pigs, mice, etc.) APPARENTLY a
temporary inversion of the two germinal layers. This is due to a
folding of the blastodermic wall by what is called the "girder," a
plug-shaped growth of Rauber's "roof-layer." It is a thin layer of
flat epithelial cells, that is freed from the surface of the
blastoderm in some of the rodents; it has no more significance in
connection with the general course of placental gastrulation than the
conspicuous departure from the usual globular shape in the blastula of
some of the ungulates. In some pigs and ruminants it grows into a
thread-like, long and thin tube.

(FIGURE 1.68. Stem-cell of the mammal ovum (from the rabbit). k
stem-nucleus, n nuclear corpuscle, p protoplasm of the stem-cell, z
modified zona pellucida, h outer albuminous membrane, s dead
sperm-cells.

FIGURE 1.69. Incipient cleavage of the mammal ovum (from the rabbit).
The stem-cell has divided into two unequal cells, one lighter (e) and
one darker (i). z zona pellucida, h outer albuminous membrane, s dead
sperm-cell.

FIGURE 1.70. The first four segmentation-cells of the mammal ovum
(from the rabbit). e the two larger (and lighter) cells, i the two
smaller (and darker) cells, z zona pellucida, h outer albuminous
membrane.

FIGURE 1.71. Mammal ovum with eight segmentation-cells (from the
rabbit). e four larger and lighter cells, i four smaller and darker
cells, z zona pellucida, h outer albuminous membrane.)

Thus the gastrulation of the placentals, which diverges most from that
of the amphioxus, the primitive form, is reduced to the original type,
the invagination of a modified blastula. Its chief peculiarity is that
the folded part of the blastoderm does not form a completely closed
(only open at the primitive mouth) blind sac, as is usual; but this
blind sac has a wide opening at the ventral curve (opposite to the
dorsal mouth); and through this opening the primitive gut communicates
from the first with the embryonic cavity of the blastula. The folded
crest-shaped entoderm grows with a free circular border on the inner
surface of the entoderm towards the vegetal pole; when it has reached
this, and the inner surface of the blastula is completely grown over,
the primitive gut is closed. This remarkable direct transition of the
primitive gut-cavity into the segmentation-cavity is explained simply
by the assumption that in most of the mammals the yelk-mass, which is
still possessed by the oldest forms of the class (the monotremes) and
their ancestors (the reptiles), is atrophied. This proves the
essential unity of gastrulation in all the vertebrates, in spite of
the striking differences in the various classes.

In order to complete our consideration of the important processes of
segmentation and gastrulation, we will, in conclusion, cast a brief
glance at the fourth chief type--superficial segmentation. In the
vertebrates this form is not found at all. But it plays the chief part
in the large stem of the articulates--the insects, spiders, myriapods,
and crabs. The distinctive form of gastrula that comes of it is the
"vesicular gastrula" (Perigastrula).

In the ova which undergo this superficial cleavage the formative yelk
is sharply divided from the nutritive yelk, as in the preceding cases
of the ova of birds, reptiles, fishes, etc.; the formative yelk alone
undergoes cleavage. But while in the ova with discoid gastrulation the
formative yelk is not in the centre, but at one pole of the uni-axial
ovum, and the food-yelk gathered at the other pole, in the ova with
superficial cleavage we find the formative yelk spread over the whole
surface of the ovum; it encloses spherically the food-yelk, which is
accumulated in the middle of the ova. As the segmentation only affects
the former and not the latter, it is bound to be entirely
"superficial"; the store of food in the middle is quite untouched by
it. As a rule, it proceeds in regular geometrical progression. In the
end the whole of the formative yelk divides into a number of small and
homogeneous cells, which lie close together in a single stratum on the
entire surface of the ovum, and form a superficial blastoderm. This
blastoderm is a simple, completely closed vesicle, the internal cavity
of which is entirely full of food-yelk. This real blastula only
differs from that of the primitive ova in its chemical composition. In
the latter the content is water or a watery jelly; in the former it is
a thick mixture, rich in food-yelk, of albuminous and fatty
substances. As this quantity of food-yelk fills the centre of the ovum
before cleavage begins, there is no difference in this respect between
the morula and the blastula. The two stages rather agree in this.

When the blastula is fully formed, we have again in this case the
important folding or invagination that determines gastrulation. The
space between the skin-layer and the gut-layer (the remainder of the
segmentation-cavity) remains full of food-yelk, which is gradually
used up. This is the only material difference between our vesicular
gastrula (perigastrula) and the original form of the bell-gastrula
(archigastrula). Clearly the one has been developed from the other in
the course of time, owing to the accumulation of food-yelk in the
centre of the ovum.* (* On the reduction of all forms of gastrulation
to the original palingenetic form see especially the lucid treatment
of the subject in Arnold Lang's Manual of Comparative Anatomy (1888),
Part 1.)

We must count it an important advance that we are thus in a position
to reduce all the various embryonic phenomena in the different groups
of animals to these four principal forms of segmentation and
gastrulation. Of these four forms we must regard one only as the
original palingenetic, and the other three as cenogenetic and
derivative. The unequal, the discoid, and the superficial segmentation
have all clearly arisen by secondary adaptation from the primary
segmentation; and the chief cause of their development has been the
gradual formation of the food-yelk, and the increasing antithesis
between animal and vegetal halves of the ovum, or between ectoderm
(skin-layer) and entoderm (gut-layer).

(FIGURE 1.72. Gastrula of the placental mammal (epigastrula from the
rabbit), longitudinal section through the axis. e ectodermic cells
(sixty-four, lighter and smaller), i entodermic cells (thirty-two,
darker and larger), d central entodermic cell, filling the primitive
gut-cavity, o peripheral entodermic cell, stopping up the opening of
the primitive mouth (yelk-stopper in the Rusconian anus).)

(FIGURE 1.73. Gastrula of the rabbit. A as a solid, spherical cluster
of cells, B changing into the embryonic vesicle, bp primitive mouth,
ep ectoderm, hy entoderm.)

The numbers of careful studies of animal gastrulation that have been
made in the last few decades have completely established the views I
have expounded, and which I first advanced in the years 1872 to 1876.
For a time they were greatly disputed by many embryologists. Some said
that the original embryonic form of the metazoa was not the gastrula,
but the "planula"--a double-walled vesicle with closed cavity and
without mouth-aperture; the latter was supposed to pierce through
gradually. It was afterwards shown that this planula (found in several
sponges, etc.) was a later evolution from the gastrula. It was also
shown that what is called delamination--the rise of the two primary
germinal layers by the folding of the surface of the blastoderm (for
instance, in the Geryonidae and other medusae)--was a secondary
formation, due to cenogenetic variations from the original
invagination of the blastula. The same may be said of what is called
"immigration," in which certain cells or groups of cells are detached
from the simple layer of the blastoderm, and travel into the interior
of the blastula; they attach themselves to the inner wall of the
blastula, and form a second internal epithelial layer--that is to say,
the entoderm. In these and many other controversies of modern
embryology the first requisite for clear and natural explanation is a
careful and discriminative distinction between palingenetic
(hereditary) and cenogenetic (adaptive) processes. If this is properly
attended to, we find evidence everywhere of the biogenetic law.


CHAPTER 1.10. THE COELOM THEORY.

The two "primary germinal layers" which the gastraea theory has shown
to be the first foundation in the construction of the body are found
in this simplest form throughout life only in animals of the lowest
grade--in the gastraeads, olynthus (the stem-form of the sponges),
hydra, and similar very simple animals. In all the other animals new
strata of cells are formed subsequently between these two primary
body-layers, and these are generally comprehended under the title of
the middle layer, or mesoderm. As a rule, the various products of this
middle layer afterwards constitute the great bulk of the animal frame,
while the original entoderm, or internal germinal layer, is restricted
to the clothing of the alimentary canal and its glandular appendages;
and, on the other hand, the ectoderm, or external germinal layer,
furnishes the outer clothing of the body, the skin and nervous system.

In some large groups of the lower animals, such as the sponges,
corals, and flat-worms, the middle germinal layer remains a single
connected mass, and most of the body is developed from it; these have
been called the three-layered metazoa, in opposition to the
two-layered animals described. Like the two-layered animals, they have
no body-cavity--that is to say, no cavity distinct from the alimentary
system. On the other hand, all the higher animals have this real
body-cavity (coeloma), and so are called coelomaria. In all these we
can distinguish four secondary germinal layers, which develop from the
two primary layers. To the same class belong all true vermalia
(excepting the platodes), and also the higher typical animal stems
that have been evolved from them--molluscs, echinoderms, articulates,
tunicates, and vertebrates.

(FIGURES 1.74 AND 1.75. Diagram of the four secondary germinal layers,
transverse section through the metazoic embryo: Figure 1.74 of an
annelid, Figure 1.75 of a vermalian. a primitive gut, dd ventral
glandular layer, df ventral fibre-layer, hm skin-fibre-layer, hs
skin-sense-layer, u beginning of the rudimentary kidneys, n beginning
of the nerve-plates.)

The body-cavity (coeloma) is therefore a new acquisition of the animal
body, much younger than the alimentary system, and of great
importance. I first pointed out this fundamental significance of the
coelom in my Monograph on the Sponges (1872), in the section which
draws a distinction between the body-cavity and the gut-cavity, and
which follows immediately on the germ-layer theory and the ancestral
tree of the animal kingdom (the first sketch of the gastraea theory).
Up to that time these two principal cavities of the animal body had
been confused, or very imperfectly distinguished; chiefly because
Leuckart, the founder of the coelenterata group (1848), has attributed
a body-cavity, but not a gut-cavity, to these lowest metazoa. In
reality, the truth is just the other way about.

The ventral cavity, the original organ of nutrition in the
multicellular animal-body, is the oldest and most important organ of
all the metazoa, and, together with the primitive mouth, is formed in
every case in the gastrula as the primitive gut; it is only at a much
later stage that the body-cavity, which is entirely wanting in the
coelenterata, is developed in some of the metazoa between the ventral
and the body wall. The two cavities are entirely different in content
and purport. The alimentary cavity (enteron) serves the purpose of
digestion; it contains water and food taken from without, as well as
the pulp (chymus) formed from this by digestion. On the other hand,
the body-cavity, quite distinct from the gut and closed externally,
has nothing to do with digestion; it encloses the gut itself and its
glandular appendages, and also contains the sexual products and a
certain amount of blood or lymph, a fluid that is transuded through
the ventral wall.

As soon as the body-cavity appears, the ventral wall is found to be
separated from the enclosing body-wall, but the two continue to be
directly connected at various points. We can also then always
distinguish a number of different layers of tissue in both walls--at
least two in each. These tissue-layers are formed originally from four
different simple cell-layers, which are the much-discussed four
secondary germinal layers. The outermost of these, the
skin-sense-layer (Figures 1.74 and 1.75 hs), and the innermost, the
gut-gland-layer (dd), remain at first simple epithelia or
covering-layers. The one covers the outer surface of the body, the
other the inner surface of the ventral wall; hence they are called
confining or limiting layers. Between them are the two middle-layers,
or mesoblasts, which enclose the body-cavity.

(FIGURE 1.76. Coelomula of sagitta (gastrula with a couple of
coelom-pouches. (From Kowalevsky.) bl.p primitive mouth, al primitive
gut, pv coelom-folds, m permanent mouth.)

The four secondary germinal layers are so distributed in the structure
of the body in all the coelomaria (or all metazoa that have a
body-cavity) that the outer two, joined fast together, constitute the
body-wall, and the inner two the ventral wall; the two walls are
separated by the cavity of the coelom. Each of the walls is made up of
a limiting layer and a middle layer. The two limiting layers chiefly
give rise to epithelia, or covering-tissues, and glands and nerves,
while the middle layers form the great bulk of the fibrous tissue,
muscles, and connective matter. Hence the latter have also been called
fibrous or muscular layers. The outer middle layer, which lies on the
inner side of the skin-sense-layer, is the skin fibre-layer; the inner
middle layer, which attaches from without to the ventral glandular
layer, is the ventral fibre layer. The former is usually called
briefly the parietal, and the latter the visceral layer or mesoderm.
Of the many different names that have been given to the four secondary
germinal layers, the following are those most in use to-day:--

1.  Skin-sense-layer (outer limiting layer) and 2. Skin-fibre-layer
(outer middle layer).

I. Neural layer (neuroblast) and II. Parietal layer (myoblast). The
two secondary germinal layers of the body-wall: 1. Epithelial. 2.
Fibrous.

3. Gut-fibre-layer (inner middle layer) and 4. Gut-gland-layer (inner
limiting layer).

III. Visceral layer (gonoblast) and IV. Enteral layer (enteroblast).
The two secondary germinal layers of the gut-wall: 3. Fibrous. 4.
Epithelial.

The first scientist to recognise and clearly distinguish the four
secondary germinal layers was Baer. It is true that he was not quite
clear as to their origin and further significance, and made several
mistakes in detail in explaining them. But, on the whole, their great
importance did not escape him. However, in later years his view had to
be given up in consequence of more accurate observations. Remak then
propounded a three-layer theory, which was generally accepted. These
theories of cleavage, however, began to give way thirty years ago,
when Kowalevsky (1871) showed that in the case of Sagitta (a very
clear and typical subject of gastrulation) the two middle germinal
layers and the two limiting layers arise not by cleavage, but by
folding--by a secondary invagination of the primary inner germ-layer.
This invagination or folding proceeds from the primitive mouth, at the
two sides of which (right and left) a couple of pouches are formed. As
these coelom-pouches or coelom-sacs detach themselves from the
primitive gut, a double body-cavity is formed (Figures 1.74 to 1.76).

(FIGURE 1.77. Coelomula of sagitta, in section. (From Hertwig.) D
dorsal side, V ventral side, ik inner germinal layer, mv visceral
mesoblast, lh body-cavity, mp parietal mesoblast, ak outer germinal
layer.)

The same kind of coelom-formation as in sagitta was afterwards found
by Kowalevsky in brachiopods and other invertebrates, and in the
lowest vertebrate--the amphioxus. Further instances were discovered by
two English embryologists, to whom we owe very considerable advance in
ontogeny--E. Ray-Lankester and F. Balfour. On the strength of these
and other studies, as well as most extensive research of their own,
the brothers Oscar and Richard Hertwig constructed in 1881 the Coelom
Theory. In order to appreciate fully the great merit of this
illuminating and helpful theory, one must remember what a chaos of
contradictory views was then represented by the "problem of the
mesoderm," or the much-disputed "question of the origin of the middle
germinal layer." The coelom theory brought some light and order into
this infinite confusion by establishing the following points: 1. The
body-cavity originates in the great majority of animals (especially in
all the vertebrates) in the same way as in sagitta: a couple of
pouches or sacs are formed by folding inwards at the primitive mouth,
between the two primary germinal layers; as these pouches detach from
the primitive gut, a pair of coelom-sacs (right and left) are formed;
the coalescence of these produces a simple body-cavity. 2. When these
coelom-embryos develop, not as a pair of hollow pouches, but as solid
layers of cells (in the shape of a pair of mesodermal streaks)--as
happens in the higher vertebrates--we have a secondary (cenogenetic)
modification of the primary (palingenetic) structure; the two walls of
the pouches, inner and outer, have been pressed together by the
expansion of the large food-yelk. 3. Hence the mesoderm consists from
the first of TWO genetically distinct layers, which do not originate
by the cleavage of a primary simple middle layer (as Remak supposed).
4. These two middle layers have, in all vertebrates, and the great
majority of the invertebrates, the same radical significance for the
construction of the animal body; the inner middle layer, or the
visceral mesoderm, (gut-fibre layer), attaches itself to the original
entoderm, and forms the fibrous, muscular, and connective part of the
visceral wall; the outer middle layer, or the parietal mesoderm
(skin-fibre-layer), attaches itself to the original ectoderm and forms
the fibrous, muscular, and connective part of the body-wall. 5. It is
only at the point of origination, the primitive mouth and its
vicinity, that the four secondary germinal layers are directly
connected; from this point the two middle layers advance forward
separately between the two primary germinal layers, to which they
severally attach themselves. 6. The further separation or
differentiation of the four secondary germinal layers and their
division into the various tissues and organs take place especially in
the later fore-part or head of the embryo, and extend backwards from
there towards the primitive mouth.

(FIGURE 1.78. Section of a young sagitta. (From Hertwig.) dh visceral
cavity, ik and ak inner and outer limiting layers, mv and mp inner and
outer middle layers, lk body-cavity, dm and vm dorsal and visceral
mesentery.)

All animals in which the body-cavity demonstrably arises in this way
from the primitive gut (vertebrates, tunicates, echinoderms,
articulates, and a part of the vermalia) were comprised by the
Hertwigs under the title of enterocoela, and were contrasted with the
other groups of the pseudocoela (with false body-cavity) and the
coelenterata (with no body-cavity). However, this radical distinction
and the views as to classification which it occasioned have been shown
to be untenable. Further, the absolute differences in tissue-formation
which the Hertwigs set up between the enterocoela and pseudocoela
cannot be sustained in this connection. For these and other reasons
their coelom-theory has been much criticised and partly abandoned.
Nevertheless, it has rendered a great and lasting service in the
solution of the difficult problem of the mesoderm, and a material part
of it will certainly be retained. I consider it an especial merit of
the theory that it has established the identity of the development of
the two middle layers in all the vertebrates, and has traced them as
cenogenetic modifications back to the original palingenetic form of
development that we still find in the amphioxus. Carl Rabl comes to
the same conclusion in his able Theory of the Mesoderm, and so do
Ray-Lankester, Rauber, Kupffer, Ruckert, Selenka, Hatschek, and
others. There is a general agreement in these and many other recent
writers that all the different forms of coelom-construction, like
those of gastrulation, follow one and the same strict hereditary law
in the vast vertebrate stem; in spite of their apparent differences,
they are all only cenogenetic modifications of one palingenetic type,
and this original type has been preserved for us down to the present
day by the invaluable amphioxus.

(FIGURES 1.79 AND 1.80. Transverse section of amphioxus-larvae. (From
Hatschek.) Figure 1.79 at the commencement of coelom formation (still
without segments), Figure 1.80 at the stage with four primitive
segments. ak, ik, mk outer, inner, and middle germinal layer, hp horn
plate, mp medullary plate, ch chorda, asterisk and asterisk,
disposition of the coelom-pouches, lh body-cavity.)

But before we go into the regular coelomation of the amphioxus, we
will glance at that of the arrow-worm (Sagitta), a remarkable deep-sea
worm that is interesting in many ways for comparative anatomy and
ontogeny. On the one hand, the transparency of the body and the
embryo, and, on the other hand, the typical simplicity of its
embryonic development, make the sagitta a most instructive object in
connection with various problems. The class of the chaetogatha, which
is only represented by the cognate genera of Sagitta and Spadella, is
in another respect also a most remarkable branch of the extensive
vermalia stem. It was therefore very gratifying that Oscar Hertwig
(1880) fully explained the anatomy, classification, and evolution of
the chaetognatha in his careful monograph.

The spherical blastula that arises from the impregnated ovum of the
sagitta is converted by a folding at one pole into a typical
archigastrula, entirely similar to that of the Monoxenia which I
described (Chapter 1.8, Figure 1.29). This oval, uni-axial cup-larva
(circular in section) becomes bilateral (or tri-axial) by the growth
of a couple of coelom-pouches from the primitive gut (Figures 1.76 and
1.77). To the right and left a sac-shaped fold appears towards the top
pole (where the permanent mouth, m, afterwards arises). The two sacs
are at first separated by a couple of folds of the entoderm (Figure
1.76 pv), and are still connected with the primitive gut by wide
apertures; they also communicate for a short time with the dorsal side
(Figure 1.77 d). Soon, however, the coelom-pouches completely separate
from each other and from the primitive gut; at the same time they
enlarge so much that they close round the primitive gut (Figure 1.78).
But in the middle line of the dorsal and ventral sides the pouches
remain separated, their approaching walls joining here to form a thin
vertical partition, the mesentery (dm and vm). Thus Sagitta has
throughout life a double body-cavity (Figure 1.78 lk), and the gut is
fastened to the body-wall both above and below by a mesentery--below
by the ventral mesentery (vm), and above by the dorsal mesentery (dm).
The inner layer of the two coelom-pouches (mv) attaches itself to the
entoderm (ik), and forms with it the visceral wall. The outer layer
(mp) attaches itself to the ectoderm (ak), and forms with it the outer
body-wall. Thus we have in Sagitta a perfectly clear and simple
illustration of the original coelomation of the enterocoela. This
palingenetic fact is the more important, as the greater part of the
two body-cavities in Sagitta changes afterwards into sexual
glands--the fore or female part into a pair of ovaries, and the hind
or male part into a pair of testicles.

Coelomation takes place with equal clearness and transparency in the
case of the amphioxus, the lowest vertebrate, and its nearest
relatives, the invertebrate tunicates, the sea-squirts. However, in
these two stems, which we class together as Chordonia, this important
process is more complex, as two other processes are associated with
it--the development of the chorda from the entoderm and the separation
of the medullary plate or nervous centre from the ectoderm. Here again
the skulless amphioxus has preserved to our own time by tenacious
heredity the chief phenomena in their original form, while it has been
more or less modified by embryonic adaptation in all the other
vertebrates (with skulls). Hence we must once more thoroughly
understand the palingenetic embryonic features of the lancelet before
we go on to consider the cenogenetic forms of the craniota.

(FIGURES 1.81 AND 1.82. Transverse section of amphioxus embryo. Figure
1.81 at the stage with five somites, Figure 1.82 at the stage with
eleven somites. (From Hatschek.) ak outer germinal layer, mp medullary
plate, n nerve-tube, ik inner germinal layer, dh visceral cavity, lh
body-cavity, mk middle germinal layer (mk1 parietal, mk2 visceral), us
primitive segment, ch chorda.)

The coelomation of the amphioxus, which was first observed by
Kowalevsky in 1867, has been very carefully studied since by Hatschek
(1881). According to him, there are first formed on the bilateral
gastrula we have already considered (Figures 1.36 and 1.37) three
parallel longitudinal folds--one single ectodermal fold in the central
line of the dorsal surface, and a pair of entodermic folds at the two
sides of the former. The broad ectodermal fold that first appears in
the middle line of the flattened dorsal surface, and forms a shallow
longitudinal groove, is the beginning of the central nervous system,
the medullary tube. Thus the primary outer germinal layer divides into
two parts, the middle medullary plate (Figure 1.81 mp) and the
horny-plate (ak), the beginning of the outer skin or epidermis. As the
parallel borders of the concave medullary plate fold towards each
other and grow underneath the horny-plate, a cylindrical tube is
formed, the medullary tube (Figure 1.82 n); this quickly detaches
itself altogether from the horny-plate. At each side of the medullary
tube, between it and the alimentary tube (Figures 1.79 to 1.82 dh),
the two parallel longitudinal folds grow out of the dorsal wall of the
alimentary tube, and these form the two coelom-pouches (Figures 1.80
and 1.81 lh). This part of the entoderm, which thus represents the
first structure of the middle germinal layer, is shown darker than the
rest of the inner germinal layer in Figures 1.79 to 1.82. The edges of
the folds meet, and thus form closed tubes (Figure 1.81 in section).

During this interesting process the outline of a third very important
organ, the chorda or axial rod, is being formed between the two
coelom-pouches. This first foundation of the skeleton, a solid
cylindrical cartilaginous rod, is formed in the middle line of the
dorsal primitive gut-wall, from the entodermal cell-streak that
remains here between the two coelom-pouches (Figures 1.79 to 1.82 ch).
The chorda appears at first in the shape of a flat longitudinal fold
or a shallow groove (Figures 1.80 and 1.81); it does not become a
solid cylindrical cord until after separation from the primitive gut
(Figure 1.82). Hence we might say that the dorsal wall of the
primitive gut forms three parallel longitudinal folds at this
important period--one single fold and a pair of folds. The single
middle fold becomes the chorda, and lies immediately below the groove
of the ectoderm, which becomes the medullary tube; the pair of folds
to the right and left lie at the sides between the former and the
latter, and form the coelom-pouches. The part of the primitive gut
that remains after the cutting off of these three dorsal primitive
organs is the permanent gut; its entoderm is the gut-gland-layer or
enteric layer.

(FIGURES 1.83 AND 1.84. Chordula of the amphioxus. Figure 1.83 median
longitudinal section (seen from the left). Figure 1.84 transverse
section. (From Hatschek.) In Figure 1.83 the coelom-pouches are
omitted, in order to show the chordula more clearly. Figure 1.84 is
rather diagrammatic. h horny-plate, m medullary tube, n wall of same
(n apostrophe, dorsal, n double apostrophe, ventral), ch chorda, np
neuroporus, ne canalis neurentericus, d gut-cavity, r gut dorsal wall,
b gut ventral wall, z yelk-cells in the latter, u primitive mouth, o
mouth-pit, p promesoblasts (primitive or polar cells of the mesoderm),
w parietal layer, v visceral layer of the mesoderm, c coelom, f rest
of the segmentation-cavity.

FIGURES 1.85 AND 1.86. Chordula of the amphibia (the ringed adder).
(From Goette.) Figure 85 median longitudinal section (seen from the
left), Figure 1.86 transverse section (slightly diagrammatic).
Lettering as in Figures 1.83 and 1.84.

FIGURES 1.87 AND 1.88. Diagrammatic vertical section of
coelomula-embryos of vertebrates. (From Hertwig.) Figure 1.87,
vertical section THROUGH the primitive mouth, Figure 1.88, vertical
section BEFORE the primitive mouth. u primitive mouth, ud primitive
gut. d yelk, dk yelk-nuclei, dh gut-cavity, lh body-cavity, mp
medullary plate, ch chorda plate, ak and ik outer and inner germinal
layers, pb parietal and vb visceral mesoblast.

FIGURES 1.89 AND 1.90. Transverse section of coelomula embryos of
triton. (From Hertwig.) Figure 1.89, section THROUGH the primitive
mouth. Figure 1.90, section in front of the primitive mouth, u
primitive mouth. dh gut-cavity, dz yelk-cells, dp yelk-stopper, ak
outer and ik inner germinal layer, pb parietal and vb visceral middle
layer, m medullary plate, ch chorda.)

I give the name of chordula or chorda-larva to the embryonic stage of
the vertebrate organism which is represented by the amphioxus larva at
this period (Figures 1.83 and 1.84, in the third period of development
according to Hatschek). (Strabo and Plinius give the name of cordula
or cordyla to young fish larvae.) I ascribe the utmost phylogenetic
significance to it, as it is found in all the chorda-animals
(tunicates as well as vertebrates) in essentially the same form.
Although the accumulation of food-yelk greatly modifies the form of
the chordula in the higher vertebrates, it remains the same in its
main features throughout. In all cases the nerve-tube (m) lies on the
dorsal side of the bilateral, worm-like body, the gut-tube (d) on the
ventral side, the chorda (ch) between the two, on the long axis, and
the coelom pouches (c) at each side. In every case these primitive
organs develop in the same way from the germinal layers, and the same
organs always arise from them in the mature chorda-animal. Hence we
may conclude, according to the laws of the theory of descent, that all
these chordonia or chordata (tunicates and vertebrates) descend from
an ancient common ancestral form, which we may call Chordaea. We
should regard this long-extinct Chordaea, if it were still in
existence, as a special class of unarticulated worm (chordaria). It is
especially noteworthy that neither the dorsal nerve-tube nor the
ventral gut-tube, nor even the chorda that lies between them, shows
any trace of articulation or segmentation; even the two coelom-sacs
are not segmented at first (though in the amphioxus they quickly
divide into a series of parts by transverse folding). These
ontogenetic facts are of the greatest importance for the purpose of
learning those ancestral forms of the vertebrates which we have to
seek in the group of the unarticulated vermalia. The coelom-pouches
were originally sexual glands in these ancient chordonia.

(FIGURE 1.91. A, B, C. Vertical section of the dorsal part of three
triton-embryos. (From Hertwig.) In Figure A the medullary swellings
(the parallel borders of the medullary plate) begin to rise; in Figure
B they grow towards each other; in Figure C they join and form the
medullary tube. mp medullary plate, mf medullary folds, n nerve-tube,
ch chorda, lh body-cavity, mk1 and mk2 parietal and visceral
mesoblasts, uv primitive-segment cavities, ak ectoderm, ik entoderm,
dz yelk-cells, dh gut-cavity.)

From the evolutionary point of view the coelom-pouches are, in any
case, older than the chorda; since they also develop in the same way
as in the chordonia in a number of invertebrates which have no chorda
(for instance, Sagitta, Figures 1.76 to 1.78). Moreover, in the
amphioxus the first outline of the chorda appears later than that of
the coelom-sacs. Hence we must, according to the biogenetic law,
postulate a special intermediate form between the gastrula and the
chordula, which we will call coelomula, an unarticulated, worm-like
body with primitive gut, primitive mouth, and a double body-cavity,
but no chorda. This embryonic form, the bilateral coelomula (Figure
1.81), may in turn be regarded as the ontogenetic reproduction
(maintained by heredity) of an ancient ancestral form of the
coelomaria, the Coelomaea (cf. Chapter 2.20).

In Sagitta and other worm-like animals the two coelom-pouches
(presumably gonads or sex-glands) are separated by a complete median
partition, the dorsal and ventral mesentery (Figure 1.78 dm and vm);
but in the vertebrates only the upper part of this vertical partition
is maintained, and forms the dorsal mesentery. This mesentery
afterwards takes the form of a thin membrane, which fastens the
visceral tube to the chorda (or the vertebral column). At the under
side of the visceral tube the coelom-sacs blend together, their inner
or median walls breaking down and disappearing. The body-cavity then
forms a single simple hollow, in which the gut is quite free, or only
attached to the dorsal wall by means of the mesentery.

The development of the body-cavity and the formation of the chordula
in the higher vertebrates is, like that of the gastrula, chiefly
modified by the pressure of the food-yelk on the embryonic structures,
which forces its hinder part into a discoid expansion. These
cenogenetic modifications seem to be so great that until twenty years
ago these important processes were totally misunderstood. It was
generally believed that the body-cavity in man and the higher
vertebrates was due to the division of a simple middle layer, and that
the latter arose by cleavage from one or both of the primary germinal
layers. The truth was brought to light at last by the comparative
embryological research of the Hertwigs. They showed in their Coelom
Theory (1881) that all vertebrates are true enterocoela, and that in
every case a pair of coelom-pouches are developed from the primitive
gut by folding. The cenogenetic chordula-forms of the craniotes must
therefore be derived from the palingenetic embryology of the amphioxus
in the same way as I had previously proved for their gastrula-forms.

The chief difference between the coelomation of the acrania
(amphioxus) and the other vertebrates (with skulls--craniotes) is that
the two coelom-folds of the primitive gut in the former are from the
first hollow vesicles, filled with fluid, but in the latter are empty
pouches, the layers of which (inner and outer) close with each other.
In common parlance we still call a pouch or pocket by that name,
whether it is full or empty. It is different in ontogeny; in some of
our embryological literature ordinary logic does not count for very
much. In many of the manuals and large treatises on this science it is
proved that vesicles, pouches, or sacs deserve that name only when
they are inflated and filled with a clear fluid. When they are not so
filled (for instance, when the primitive gut of the gastrula is filled
with yelk, or when the walls of the empty coelom-pouches are pressed
together), these vesicles must not be cavities any longer, but "solid
structures."

The accumulation of food-yelk in the ventral wall of the primitive gut
(Figures 1.85 and 1.86) is the simple cause that converts the
sac-shaped coelom-pouches of the acrania into the leaf-shaped
coelom-streaks of the craniotes. To convince ourselves of this we need
only compare, with Hertwig, the palingenetic coelomula of the
amphioxus (Figures 1.80 and 1.81) with the corresponding cenogenetic
form of the amphibia (Figures 1.89 to 1.90), and construct the simple
diagram that connects the two (Figures 1.87 and 1.88). If we imagine
the ventral half of the primitive gut-wall in the amphioxus embryo
(Figures 1.79 to 1.84) distended with food-yelk, the vesicular
coelom-pouches (lh) must be pressed together by this, and forced to
extend in the shape of a thin double plate between the gut-wall and
body-wall (Figures 1.86 and 1.87). This expansion follows a downward
and forward direction. They are not directly connected with these two
walls. The real unbroken connection between the two middle layers and
the primary germ-layers is found right at the back, in the region of
the primitive mouth (Figure 1.87 u). At this important spot we have
the source of embryonic development (blastocrene), or "zone of
growth," from which the coelomation (and also the gastrulation)
originally proceeds.

(FIGURE 1.92. Transverse section of the chordula-embryo of a bird
(from a hen's egg at the close of the first day of incubation). (From
Kolliker,) h horn-plate (ectoderm), m medullary plate, Rf dorsal folds
of same, Pv medullary furrow, ch chorda, uwp median (inner) part of
the middle layer (median wall of the coelom-pouches), sp lateral
(outer) part of same, or lateral plates, uwh structure of the
body-cavity, dd gut-gland-layer.)

Hertwig even succeeded in showing, in the coelomula-embryo of the
water salamander (Triton), between the first structures of the two
middle layers, the relic of the body-cavity, which is represented in
the diagrammatic transitional form (Figures 1.87 and 1.88). In
sections both through the primitive mouth itself (Figure 1.89) and in
front of it (Figure 1.90) the two middle layers (pb and vb) diverge
from each other, and disclose the two body-cavities as narrow clefts.
At the primitive-mouth itself (Figure 1.90 u) we can penetrate into
them from without. It is only here at the border of the primitive
mouth that we can show the direct transition of the two middle layers
into the two limiting layers or primary germinal layers.

The structure of the chorda also shows the same features in these
coelomula-embryos of the amphibia (Figure 1.91) as in the amphioxus
(Figures 1.79 to 1.82). It arises from the entodermic cell-streak,
which forms the middle dorsal-line of the primitive gut, and occupies
the space between the flat coelom-pouches (Figure 1.91 A). While the
nervous centre is formed here in the middle line of the back and
separated from the ectoderm as "medullary tube," there takes place at
the same time, directly underneath, the severance of the chorda from
the entoderm (Figure 1.91 A, B, C). Under the chorda is formed (out of
the ventral entodermic half of the gastrula) the permanent gut or
visceral cavity (enteron) (Figure 1.91 B, dh). This is done by the
coalescence, under the chorda in the median line, of the two dorsal
side-borders of the gut-gland-layer (ik), which were previously
separated by the chorda-plate (Figure 1.91 A, ch); these now alone
form the clothing of the visceral cavity (dh) (enteroderm, Figure 1.91
C). All these important modifications take place at first in the fore
or head-part of the embryo, and spread backwards from there; here at
the hinder end, the region of the primitive mouth, the important
border of the mouth (or properistoma) remains for a long time the
source of development or the zone of fresh construction, in the
further building-up of the organism. One has only to compare carefully
the illustrations given (Figures 1.85 to 1.91) to see that, as a fact,
the cenogenetic coelomation of the amphibia can be deduced directly
from the palingenetic form of the acrania (Figures 1.79 to 1.84).

(FIGURE 1.93. Transverse section of the vertebrate-embryo of a bird
(from a hen's egg on the second day of incubation). (From Kolliker.) h
horn-plate, mr medullary tube, ch chorda, uw primitive segments, uwh
primitive-segment cavity (median relic of the coelom), sp lateral
coelom-cleft, hpl skin-fibre-layer, df gut-fibre-layer, ung
primitive-kidney passage, ao primitive aorta, dd gut-gland-layer.)

The same principle holds good for the amniotes, the reptiles, birds,
and mammals, although in this case the processes of coelomation are
more modified and more difficult to identify on account of the
colossal accumulation of food-yelk and the corresponding notable
flattening of the germinal disk. However, as the whole group of the
amniotes has been developed at a comparatively late date from the
class of the amphibia, their coelomation must also be directly
traceable to that of the latter. This is really possible as a matter
of fact; even the older illustrations showed an essential identity of
features. Thus forty years ago Kolliker gave, in the first edition of
his Human Embryology (1861), some sections of the chicken-embryo, the
features of which could at once be reduced to those already described
and explained in the sense of Hertwig's coelom-theory. A section
through the embryo in the hatched hen's egg towards the close of the
first day of incubation shows in the middle of the dorsal surface a
broad ectodermic medullary groove (Figure 1.92 Rf), and underneath the
middle of the chorda (ch) and at each side of it a couple of broad
mesodermic layers (sp). These enclose a narrow space or cleft (uwh),
which is nothing else than the structure of the body-cavity. The two
layers that enclose it--the upper parietal layer (hpl) and the lower
visceral layer (df)--are pressed together from without, but clearly
distinguishable. This is even clearer a little later, when the
medullary furrow is closed into the nerve-tube (Figure 1.93 mr).

Special importance attaches to the fact that here again the four
secondary germinal layers are already sharply distinct, and easily
separated from each other. There is only one very restricted area in
which they are connected, and actually pass into each other; this is
the region of the primitive mouth, which is contracted in the amniotes
into a dorsal longitudinal cleft, the primitive groove. Its two
lateral lip-borders form the primitive streak, which has long been
recognised as the most important embryonic source and starting-point
of further processes. Sections through this primitive streak (Figures
1.94 and 1.95) show that the two primary germinal layers grow at an
early stage (in the discoid gastrula of the chick, a few hours after
incubation) into the primitive streak (x), and that the two middle
layers extend outward from this thickened axial plate (y) to the right
and left between the former. The plates of the coelom-layers, the
parietal skin-fibre-layer (m) and the visceral gut-fibre-layer (f),
are seen to be still pressed close together, and only diverge later to
form the body-cavity. Between the inner borders of the two flat
coelom-pouches lies the chorda (Figure 1.95 x), which here again
develops from the middle line of the dorsal wall of the primitive gut.

(FIGURES 1.94 AND 1.95. Transverse section of the primitive-streak
(primitive mouth) of the chick. Figure 1.94 a few hours after the
commencement of incubation, Figure 1.95 a little later. (From
Waldeyer.) h horn-plate, n nerve-plate, m skin-fibre-layer, f
gut-fibre-layer, d gut-gland-layer, y primitive streak or axial plate,
in which all four germinal layers meet, x structure of the chorda, u
region of the later primitive kidneys.)

Coelomation takes place in the vertebrates in just the same way as in
the birds and reptiles. This was to be expected, as the characteristic
gastrulation of the mammal has descended from that of the reptiles. In
both cases a discoid gastrula with primitive streak arises from the
segmented ovum, a two-layered germinal disk with long and small hinder
primitive mouth. Here again the two primary germinal layers are only
directly connected (Figure 1.96 pr) along the primitive streak (at the
folding-point of the blastula), and from this spot (the border of the
primitive mouth) the middle germinal layers (mk) grow out to right and
left between the preceding. In the fine illustration of the coelomula
of the rabbit which Van Beneden has given us (Figure 1.96) one can
clearly see that each of the four secondary germinal layers consists
of a single stratum of cells.

Finally, we must point out, as a fact of the utmost importance for our
anthropogeny and of great general interest, that the four-layered
coelomula of man has just the same construction as that of the rabbit
(Figure 1.96). A vertical section that Count Spee made through the
primitive mouth or streak of a very young human germinal disk (Figure
1.97) clearly shows that here again the four secondary germ-layers are
inseparably connected only at the primitive streak, and that here also
the two flattened coelom-pouches (mk) extend outwards to right and
left from the primitive mouth between the outer and inner germinal
layers. In this case, too, the middle germinal layer consists from the
first of two separate strata of cells, the parietal (mp) and visceral
(mv) mesoblasts.

(FIGURE 1.96. Transverse section of the primitive groove (or primitive
mouth) of a rabbit. (From Van Beneden.) pr primitive mouth, ul lips of
same (primitive lips), ak and ik outer and inner germinal layers, mk
middle germinal layer, mp parietal layer, mv visceral layer of the
mesoderm.

FIGURE 1.97. Transverse section of the primitive mouth (or groove) of
a human embryo (at the
coelomula stage). (From Count Spee.) pr primitive mouth, ul lips of
same (primitive folds), ak and ik outer and inner germinal layers, mk
middle layer, mp parietal layer, mv visceral layer of the mesoblasts.)

These concordant results of the best recent investigations (which have
been confirmed by the observations of a number of scientists I have
not enumerated) prove the unity of the vertebrate-stem in point of
coelomation, no less than of gastrulation. In both respects the
invaluable amphioxus--the sole survivor of the acrania--is found to be
the original model that has preserved for us in palingenetic form by a
tenacious heredity these most important embryonic processes. From this
primary model of construction we can cenogenetically deduce all the
embryonic forms of the other vertebrates, the craniota, by secondary
modifications. My thesis of the universal formation of the gastrula by
folding of the blastula has now been clearly proved for all the
vertebrates; so also has been Hertwig's thesis of the origin of the
middle germinal layers by the folding of a couple of coelom-pouches
which appear at the border of the primitive mouth. Just as the
gastraea-theory explains the origin and identity of the two primary
layers, so the coelom-theory explains those of the four secondary
layers. The point of origin is always the properistoma, the border of
the original primitive mouth of the gastrula, at which the two primary
layers pass directly into each other.

Moreover, the coelomula is important as the immediate source of the
chordula, the embryonic reproduction of the ancient, typical,
unarticulated, worm-like form, which has an axial chorda between the
dorsal nerve-tube and the ventral gut-tube. This instructive chordula
(Figures 1.83 to 1.86) provides a valuable support of our phylogeny;
it indicates the important moment in our stem-history at which the
stem of the chordonia (tunicates and vertebrates) parted for ever from
the divergent stems of the other metazoa (articulates, echinoderms,
and molluscs).

I may express here my opinion, in the form of a chordaea-theory, that
the characteristic chordula-larva of the chordonia has in reality this
great significance--it is the typical reproduction (preserved by
heredity) of the ancient common stem-form of all the vertebrates and
tunicates, the long-extinct Chordaea. We will return in Chapter 2.20
to these worm-like ancestors, which stand out as luminous points in
the obscure stem-history of the invertebrate ancestors of our race.


CHAPTER 1.11. THE VERTEBRATE CHARACTER OF MAN.

We have now secured a number of firm standing-places in the
labyrinthian course of our individual development by our study of the
important embryonic forms which we have called the cytula, morula,
blastula, gastrula, coelomula, and chordula. But we have still in
front of us the difficult task of deriving the complicated frame of
the human body, with all its different parts, organs, members, etc.,
from the simple form of the chordula. We have previously considered
the origin of this four-layered embryonic form from the two-layered
gastrula. The two primary germinal layers, which form the entire body
of the gastrula, and the two middle layers of the coelomula that
develop between them, are the four simple cell-strata, or epithelia,
which alone go to the formation of the complex body of man and the
higher animals. It is so difficult to understand this construction
that we will first seek a companion who may help us out of many
difficulties.

This helpful associate is the science of comparative anatomy. Its task
is, by comparing the fully-developed bodily forms in the various
groups of animals, to learn the general laws of organisation according
to which the body is constructed; at the same time, it has to
determine the affinities of the various groups by critical
appreciation of the degrees of difference between them. Formerly, this
work was conceived in a teleological sense, and it was sought to find
traces of the plan of the Creator in the actual purposive organisation
of animals. But comparative anatomy has gone much deeper since the
establishment of the theory of descent; its philosophic aim now is to
explain the variety of organic forms by adaptation, and their
similarity by heredity. At the same time, it has to recognise in the
shades of difference in form the degree of blood-relationship, and
make an effort to construct the ancestral tree of the animal world. In
this way, comparative anatomy enters into the closest relations with
comparative embryology on the one hand, and with the science of
classification on the other.

Now, when we ask what position man occupies among the other organisms
according to the latest teaching of comparative anatomy and
classification, and how man's place in the zoological system is
determined by comparison of the mature bodily forms, we get a very
definite and significant reply; and this reply gives us extremely
important conclusions that enable us to understand the embryonic
development and its evolutionary purport. Since Cuvier and Baer, since
the immense progress that was effected in the early decades of the
nineteenth century by these two great zoologists, the opinion has
generally prevailed that the whole animal kingdom may be distributed
in a small number of great divisions or types. They are called types
because a certain typical or characteristic structure is constantly
preserved within each of these large sections. Since we applied the
theory of descent to this doctrine of types, we have learned that this
common type is an outcome of heredity; all the animals of one type are
blood-relatives, or members of one stem, and can be traced to a common
ancestral form. Cuvier and Baer set up four of these types: the
vertebrates, articulates, molluscs, and radiates. The first three of
these are still retained, and may be conceived as natural phylogenetic
unities, as stems or phyla in the sense of the theory of descent. It
is quite otherwise with the fourth type--the radiata. These animals,
little known as yet at the beginning of the nineteenth century, were
made to form a sort of lumber-room, into which were cast all the lower
animals that did not belong to the other three types. As we obtained a
closer acquaintance with them in the course of the last sixty years,
it was found that we must distinguish among them from four to eight
different types. In this way the total number of animal stems or phyla
has been raised to eight or twelve (cf. Chapter 2.20).

These twelve stems of the animal kingdom are, however, by no means
co-ordinate and independent types, but have definite relations, partly
of subordination, to each other, and a very different phylogenetic
meaning. Hence they must not be arranged simply in a row one after the
other, as was generally done until thirty years ago, and is still done
in some manuals. We must distribute them in three subordinate
principal groups of very different value, and arrange the various
stems phylogenetically on the principles which I laid down in my
Monograph on the Sponges, and developed in the Study of the Gastraea
Theory. We have first to distinguish the unicellular animals
(protozoa) from the multicellular tissue-forming (metazoa). Only the
latter exhibit the important processes of segmentation and
gastrulation; and they alone have a primitive gut, and form germinal
layers and tissues.

The metazoa, the tissue-animals or gut-animals, then sub-divide into
two main sections, according as a body-cavity is or is not developed
between the primary germinal layers. We may call these the coelenteria
and coelomaria, the former are often also called zoophytes or
coelenterata, and the latter bilaterals. This division is the more
important as the coelenteria (without coelom) have no blood and
blood-vessels, nor an anus. The coelomaria (with body-cavity) have
generally an anus, and blood and blood-vessels. There are four stems
belonging to the coelenteria: the gastraeads ("primitive-gut
animals"), sponges, cnidaria, and platodes. Of the coelomaria we can
distinguish six stems: the vermalia at the bottom represent the common
stem-group (derived from the platodes) of these, the other five
typical stems of the coelomaria--the molluscs, echinoderms,
articulates, tunicates, and vertebrates--being evolved from them.

Man is, in his whole structure, a true vertebrate, and develops from
an impregnated ovum in just the same characteristic way as the other
vertebrates. There can no longer be the slightest doubt about this
fundamental fact, nor of the fact that all the vertebrates form a
natural phylogenetic unity, a single stem. The whole of the members of
this stem, from the amphioxus and the cyclostoma to the apes and man,
have the same characteristic disposition, connection, and development
of the central organs, and arise in the same way from the common
embryonic form of the chordula. Without going into the difficult
question of the origin of this stem, we must emphasise the fact that
the vertebrate stem has no direct affinity whatever to five of the
other ten stems; these five isolated phyla are the sponges, cnidaria,
molluscs, articulates, and echinoderms. On the other hand, there are
important and, to an extent, close phylogenetic relations to the other
five stems--the protozoa (through the amoebae), the gastraeads
(through the blastula and gastrula), the platodes and vermalia
(through the coelomula), and the tunicates (through the chordula).

How we are to explain these phylogenetic relations in the present
state of our knowledge, and what place is assigned to the vertebrates
in the animal ancestral tree, will be considered later (Chapter 2.20).
For the present our task is to make plainer the vertebrate character
of man, and especially to point out the chief peculiarities of
organisation by which the vertebrate stem is profoundly separated from
the other eleven stems of the animal kingdom. Only after these
comparative-anatomical considerations shall we be in a position to
attack the difficult question of our embryology. The development of
even the simplest and lowest vertebrate from the simple chordula
(Figures 1.83 to 1.86) is so complicated and difficult to follow that
it is necessary to understand the organic features of the fully-formed
vertebrate in order to grasp the course of its embryonic evolution.
But it is equally necessary to confine our attention, in this general
anatomic description of the vertebrate-body, to the essential facts,
and pass by all the unessential. Hence, in giving now an ideal
anatomic description of the chief features of the vertebrate and its
internal organisation, I omit all the subordinate points, and restrict
myself to the most important characteristics.

Much, of course, will seem to the reader to be essential that is only
of subordinate and secondary interest, or even not essential at all,
in the light of comparative anatomy and embryology. For instance, the
skull and vertebral column and the extremities are non-essential in
this sense. It is true that these parts are very important
PHYSIOLOGICALLY; but for the MORPHOLOGICAL conception of the
vertebrate they are not essential, because they are only found in the
higher, not the lower, vertebrates. The lowest vertebrates have
neither skull nor vertebrae, and no extremities or limbs. Even the
human embryo passes through a stage in which it has no skull or
vertebrae; the trunk is quite simple, and there is yet no trace of
arms and legs. At this stage of development man, like every other
higher vertebrate, is essentially similar to the simplest vertebrate
form, which we now find in only one living specimen. This one lowest
vertebrate that merits the closest study--undoubtedly the most
interesting of all the vertebrates after man--is the famous lancelet
or amphioxus, to which we have already often referred. As we are going
to study it more closely later on (Chapters 2.16 and 2.17), I will
only make one or two passing observations on it here.

The amphioxus lives buried in the sand of the sea, is about one or two
inches in length, and has, when fully developed, the shape of a very
simple, longish, lancet-like leaf; hence its name of the lancelet. The
narrow body is compressed on both sides, almost equally pointed at the
fore and hind ends, without any trace of external appendages or
articulation of the body into head, neck, breast, abdomen, etc. Its
whole shape is so simple that its first discoverer thought it was a
naked snail. It was not until much later--half a century ago--that the
tiny creature was studied more carefully, and was found to be a true
vertebrate. More recent investigations have shown that it is of the
greatest importance in connection with the comparative anatomy and
ontogeny of the vertebrates, and therefore with human phylogeny. The
amphioxus reveals the great secret of the origin of the vertebrates
from the invertebrate vermalia, and in its development and structure
connects directly with certain lower tunicates, the ascidia.

When we make a number of sections of the body of the amphioxus,
firstly vertical longitudinal sections through the whole body from end
to end, and secondly transverse sections from right to left, we get
anatomic pictures of the utmost instructiveness (cf. Figures 1.98 to
1.102). In the main they correspond to the ideal which we form, with
the aid of comparative anatomy and ontogeny, of the primitive type or
build of the vertebrate--the long-extinct form to which the whole stem
owes its origin. As we take the phylogenetic unity of the vertebrate
stem to be beyond dispute, and assume a common origin from a primitive
stem-form for all the vertebrates, from amphioxus to man, we are
justified in forming a definite morphological idea of this primitive
vertebrate (Prospondylus or Vertebraea). We need only imagine a few
slight and unessential changes in the real sections of the amphioxus
in order to have this ideal anatomic figure or diagram of the
primitive vertebrate form, as we see in Figures 1.98 to 1.102. The
amphioxus departs so little from this primitive form that we may, in a
certain sense, describe it as a modified "primitive vertebrate."* (*
The ideal figure of the vertebrate as given in Figures 1.98 to 1.102
is a hypothetical scheme or diagram, that has been chiefly constructed
on the lines of the amphioxus, but with a certain attention to the
comparative anatomy and ontogeny of the ascidia and appendicularia on
the one hand, and of the cyclostoma and selachii on the other. This
diagram has no pretension whatever to be an "exact picture," but
merely an attempt to reconstruct hypothetically the unknown and long
extinct vertebrate stem-form, an ideal "archetype.")

The outer form of our hypothetical primitive vertebrate was at all
events very simple, and probably more or less similar to that of the
lancelet. The bilateral or bilateral-symmetrical body is stretched out
lengthways and compressed at the sides (Figures 1.98 to 1.100), oval
in section (Figures 1.101 and 1.102). There are no external
articulation and no external appendages, in the shape of limbs, legs,
or fins. On the other hand, the division of the body into two
sections, head and trunk, was probably clearer in Prospondylus than it
is in its little-changed ancestor, the amphioxus. In both animals the
fore or head-half of the body contains different organs from the
trunk, and different on the dorsal from on the ventral side. As this
important division is found even in the sea-squirt, the remarkable
invertebrate stem-relative of the vertebrates, we may assume that it
was also found in the prochordonia, the common ancestors of both
stems. It is also very pronounced in the young larvae of the
cyclostoma; this fact is particularly interesting, as this
palingenetic larva-form is in other respects also an important
connecting-link between the higher vertebrates and the acrania.

(FIGURES 1.98 TO 1.102. The ideal primitive vertebrate (prospondylus).
Diagram. Figure 1.98 side-view (from the left). Figure 1.99 back-view.
Figure 1.100 front view. Figure 1.101 transverse section through the
head (to the left through the gill-pouches, to the right through the
gill-clefts). Figure 1.102 transverse section of the trunk (to the
right a pro-renal canal is affected). a aorta, af anus, au eye, b
lateral furrow (primitive renal process), c coeloma (body-cavity), d
small intestine, e parietal eye (epiphysis), f fin border of the skin,
g auditory vesicle, gh brain, h heart, i muscular cavity (dorsal
coelom-pouch), k gill-grut, ka gill-artery, kg gill-arch, ks
gill-folds, l liver, ma stomach, md mouth, ms muscles, na nose (smell
pit), n renal canals, u apertures of same, o outer skin, p gullet, r
spinal marrow, a sexual glands (gonads), t corium, u kidney-openings
(pores of the lateral furrow), v visceral vein (chief vein). x chorda,
y hypophysis (urinary appendage), z gullet-groove or gill-groove
(hypobranchial groove).)

The head of the acrania, or the anterior half of the body (both of the
real amphioxus and the ideal prospondylus), contains the branchial
(gill) gut and heart in the ventral section and the brain and
sense-organs in the dorsal section. The trunk, or posterior half of
the body, contains the hepatic (liver) gut and sexual-glands in the
ventral part, and the spinal marrow and most of the muscles in the
dorsal part.

In the longitudinal section of the ideal vertebrate (Figure 1.98) we
have in the middle of the body a thin and flexible, but stiff,
cylindrical rod, pointed at both ends (ch). It goes the whole length
through the middle of the body, and forms, as the central skeletal
axis, the original structure of the later vertebral column. This is
the axial rod, or chorda dorsalis, also called chorda vertebralis,
vertebral cord, axial cord, dorsal cord, notochorda, or, briefly,
chorda. This solid, but flexible and elastic, axial rod consists of a
cartilaginous mass of cells, and forms the inner axial skeleton or
central frame of the body; it is only found in vertebrates and
tunicates, not in any other animals. As the first structure of the
spinal column it has the same radical significance in all vertebrates,
from the amphioxus to man. But it is only in the amphioxus and the
cyclostoma that the axial rod retains its simplest form throughout
life. In man and all the higher vertebrates it is found only in the
earlier embryonic period, and is afterwards replaced by the
articulated vertebral column.

The axial rod or chorda is the real solid chief axis of the vertebrate
body, and at the same time corresponds to the ideal long-axis, and
serves to direct us with some confidence in the orientation of the
principal organs. We therefore take the vertebrate-body in its
original, natural disposition, in which the long-axis lies
horizontally, the dorsal side upward and the ventral side downward
(Figure 1.98). When we make a vertical section through the whole
length of this long axis, the body divides into two equal and
symmetrical halves, right and left. In each half we have ORIGINALLY
the same organs in the same disposition and connection; only their
disposal in relation to the vertical plane of section, or median
plane, is exactly reversed: the left half is the reflection of the
right. We call the two halves antimera (opposed-parts). In the
vertical plane of section that divides the two halves the sagittal
("arrow") axis, or "dorsoventral axis," goes from the back to the
belly, corresponding to the sagittal seam of the skull. But when we
make a horizontal longitudinal section through the chorda, the whole
body divides into a dorsal and a ventral half. The line of section
that passes through the body from right to left is the transverse,
frontal, or lateral axis.

The two halves of the vertebrate body that are separated by this
horizontal transverse axis and by the chorda have quite different
characters. The dorsal half is mainly the animal part of the body, and
contains the greater part of what are called the animal organs, the
nervous system, muscular system, osseous system, etc.--the instruments
of movement and sensation. The ventral half is essentially the
vegetative half of the body, and contains the greater part of the
vertebrate's vegetal organs, the visceral and vascular systems, sexual
system, etc.--the instruments of nutrition and reproduction. Hence in
the construction of the dorsal half it is chiefly the outer, and in
the construction of the ventral half chiefly the inner, germinal layer
that is engaged. Each of the two halves develops in the shape of a
tube, and encloses a cavity in which another tube is found. The dorsal
half contains the narrow spinal-column cavity or vertebral canal ABOVE
the chorda, in which lies the tube-shaped central nervous system, the
medullary tube. The ventral half contains the much more spacious
visceral cavity or body-cavity UNDERNEATH the chorda, in which we find
the alimentary canal and all its appendages.

The medullary tube, as the central nervous system or psychic organ of
the vertebrate is called in its first stage, consists, in man and all
the higher vertebrates, of two different parts: the large brain,
contained in the skull, and the long spinal cord which stretches from
there over the whole dorsal part of the trunk. Even in the primitive
vertebrate this composition is plainly indicated. The fore half of the
body, which corresponds to the head, encloses a knob-shaped vesicle,
the brain (gh); this is prolonged backwards into the thin cylindrical
tube of the spinal marrow (r). Hence we find here this very important
psychic organ, which accomplishes sensation, will, and thought, in the
vertebrates, in its simplest form. The thick wall of the nerve-tube,
which runs through the long axis of the body immediately over the
axial rod, encloses a narrow central canal filled with fluid (Figures
1.98 to 1.102 r). We still find the medullary tube in this very simple
form for a time in the embryo of all the vertebrates, and it retains
this form in the amphioxus throughout life; only in the latter case
the cylindrical medullary tube barely indicates the separation of
brain and spinal cord. The lancelet's medullary tube runs nearly the
whole length of the body, above the chorda, in the shape of a long
thin tube of almost equal diameter throughout, and there is only a
slight swelling of it right at the front to represent the rudiment of
a cerebral lobe. It is probable that this peculiarity of the amphioxus
is connected with the partial atrophy of its head, as the ascidian
larvae on the one hand and the young cyclostoma on the other clearly
show a division of the vesicular brain, or head marrow, from the
thinner, tubular spinal marrow.

Probably we must trace to the same phylogenetic cause the defective
nature of the sense organs of the amphioxus, which we will describe
later (Chapter 2.16). Prospondylus, on the other hand, probably had
three pairs of sense-organs, though of a simple character, a pair of,
or a single olfactory depression, right in front (Figures 1.98 and
1.99, na), a pair of eyes (au) in the lateral walls of the brain, and
a pair of simple auscultory vesicles (g) behind. There was also,
perhaps, a single parietal or "pineal" eye at the top of the skull
(epiphysis, e).

In the vertical median plane (or middle plane, dividing the bilateral
body into right and left halves) we have in the acrania, underneath
the chorda, the mesentery and visceral tube, and above it the
medullary tube; and above the latter a membranous partition of the two
halves of the body. With this partition is connected the mass of
connective tissue which acts as a sheath both for the medullary tube
and the underlying chorda, and is, therefore, called the chord-sheath
(perichorda); it originates from the dorsal and median part of the
coelom-pouches, which we shall call the skeleton plate or "sclerotom"
in the craniote embryo. In the latter the chief part of the
skeleton--the vertebral column and skull--develops from this
chord-sheath; in the acrania it retains its simple form as a soft
connective matter, from which are formed the membranous partitions
between the various muscular plates or myotomes (Figures 1.98 and 1.99
ms).

To the right and left of the cord-sheath, at each side of the
medullary tube and the underlying axial rod, we find in all the
vertebrates the large masses of muscle that constitute the musculature
of the trunk and effect its movements. Although these are very
elaborately differentiated and connected in the developed vertebrate
(corresponding to the various parts of the bony skeleton), in our
ideal primitive vertebrate we can distinguish only two pairs of these
principal muscles, which run the whole length of the body parallel to
the chorda. These are the upper (dorsal) and lower (ventral) lateral
muscles of the trunk. The upper (dorsal) muscles, or the original
dorsal muscles (Figure 1.102 ms), form the thick mass of flesh on the
back. The lower (ventral) muscles, or the original muscles of the
belly, form the fleshy wall of the abdomen. Both sets are segmented,
and consist of a double row of muscular plates (Figures 1.98 and 1.99
ms); the number of these myotomes determines the number of joints in
the trunk, or metamera. The myotomes are also developed from the thick
wall of the coelom-pouches (Figure 1.102 i).

Outside this muscular tube we have the external envelope of the
vertebrate body, which is known as the corium or cutis. This strong
and thick envelope consists, in its deeper strata, chiefly of fat and
loose connective tissue, and in its upper layers of cutaneous muscles
and firmer connective tissue. It covers the whole surface of the
fleshy body, and is of considerable thickness in all the craniota. But
in the acrania the corium is merely a thin plate of connective tissue,
an insignificant "corium-plate" (lamella corii, Figures 1.98 to 1.102
t).

Immediately above the corium is the outer skin (epidermis, o), the
general covering of the whole outer surface. In the higher vertebrates
the hairs, nails, feathers, claws, scales, etc., grow out of this
epidermis. It consists, with all its appendages and products, of
simple cells, and has no blood-vessels. Its cells are connected with
the terminations of the sensory nerves. Originally, the outer skin is
a perfectly simple covering of the outer surface of the body, composed
only of homogeneous cells--a permanent horn-plate. In this simplest
form, as a one-layered epithelium, we find it, at first, in all the
vertebrates, and throughout life in the acrania. It afterwards grows
thicker in the higher vertebrates, and divides into two strata--an
outer, firmer corneous (horn) layer and an inner, softer mucus-layer;
also a number of external and internal appendages grow out of it:
outwardly, the hairs, nails, claws, etc., and inwardly, the
sweat-glands, fat-glands, etc.

It is probable that in our primitive vertebrate the skin was raised in
the middle line of the body in the shape of a vertical fin border (f).
A similar fringe, going round the greater part of the body, is found
to-day in the amphioxus and the cyclostoma; we also find one in the
tail of fish-larvae and tadpoles.

Now that we have considered the external parts of the vertebrate and
the animal organs, which mainly lie in the dorsal half, above the
chorda, we turn to the vegetal organs, which lie for the most part in
the ventral half, below the axial rod. Here we find a large
body-cavity or visceral cavity in all the craniota. The spacious
cavity that encloses the greater part of the viscera corresponds to
only a part of the original coeloma, which we considered in Chapter
1.10; hence it nay be called the metacoeloma. As a rule, it is still
briefly called the coeloma; formerly it was known in anatomy as the
pleuroperitoneal cavity. In man and the other mammals (but only in
these) this coeloma divides, when fully developed, into two different
cavities, which are separated by a transverse partition--the muscular
diaphragm. The fore or pectoral cavity (pleura-cavity) contains the
oesophagus (gullet), heart, and lungs; the hind or peritoneal or
abdominal cavity contains the stomach, small and large intestines,
liver, pancreas, kidneys, etc. But in the vertebrate embryo, before
the diaphragm is developed, the two cavities form a single continuous
body-cavity, and we find it thus in all the lower vertebrates
throughout life. This body-cavity is clothed with a delicate layer of
cells, the coelom-epithelium. In the acrania the coelom is segmented
both dorsally and ventrally, as their muscular pouches and primitive
genital organs plainly show (Figure 1.102).

The chief of the viscera in the body-cavity is the alimentary canal,
the organ that represents the whole body in the gastrula. In all the
vertebrates it is a long tube, enclosed in the body-cavity and more or
less differentiated in length, and has two apertures--a mouth for
taking in food (Figures 1.98 and 1.100 md) and an anus for the
ejection of unusable matter or excrements (af). With the alimentary
canal a number of glands are connected which are of great importance
for the vertebrate body, and which all grow out of the canal. Glands
of this kind are the salivary glands, the lungs, the liver, and many
smaller glands. Nearly all these glands are wanting in the acrania;
probably there were merely a couple of simple hepatic tubes (Figures
1.98 and 1.100 l) in the vertebrate stem-form. The wall of the
alimentary canal and all its appendages consists of two different
layers; the inner, cellular clothing is the gut-gland-layer, and the
outer, fibrous envelope consists of the gut-fibre-layer; it is mainly
composed of muscular fibres which accomplish the digestive movements
of the canal, and of connective-tissue fibres that form a firm
envelope. We have a continuation of it in the mesentery, a thin,
bandage-like layer, by means of which the alimentary canal is fastened
to the ventral side of the chorda, originally the dorsal partition of
the two coelom-pouches. The alimentary canal is variously modified in
the vertebrates both as a whole and in its several sections, though
the original structure is always the same, and is very simple. As a
rule, it is longer (often several times longer) than the body, and
therefore folded and winding within the body-cavity, especially at the
lower end. In man and the higher vertebrates it is divided into
several sections, often separated by valves--the mouth, pharynx,
oesophagus, stomach, small and large intestine, and rectum. All these
parts develop from a very simple structure, which originally
(throughout life in the amphioxus) runs from end to end under the
chorda in the shape of a straight cylindrical canal.

As the alimentary canal may be regarded morphologically as the oldest
and most important organ in the body, it is interesting to understand
its essential features in the vertebrate more fully, and distinguish
them from unessential features. In this connection we must
particularly note that the alimentary canal of every vertebrate shows
a very characteristic division into two sections--a fore and a hind
chamber. The fore chamber is the head-gut or branchial gut (Figures
1.98 to 1.100 p, k), and is chiefly occupied with respiration. The
hind section is the trunk-gut or hepatic gut, which accomplishes
digestion (ma, d). In all vertebrates there are formed, at an early
stage, to the right and left in the fore-part of the head-gut, certain
special clefts that have an intimate connection with the original
respiratory apparatus of the vertebrate--the branchial (gill) clefts
(ks). All the lower vertebrates, the lancelets, lampreys, and fishes,
are constantly taking in water at the mouth, and letting it out again
by the lateral clefts of the gullet. This water serves for breathing.
The oxygen contained in it is inspired by the blood-canals, which
spread out on the parts between the gill-clefts, the gill-arches (kg).
These very characteristic branchial clefts and arches are found in the
embryo of man and all the higher vertebrates at an early stage of
development, just as we find them throughout life in the lower
vertebrates. However, these clefts and arches never act as respiratory
organs in the mammals, birds, and reptiles, but gradually develop into
quite different parts. Still, the fact that they are found at first in
the same form as in the fishes is one of the most interesting proofs
of the descent of these three higher classes from the fishes.

Not less interesting and important is an organ that develops from the
ventral wall in all vertebrates--the gill-groove or hypobranchial
groove. In the acrania and the ascidiae it consists throughout life of
a glandular ciliated groove, which runs down from the mouth in the
ventral middle line of the gill-gut, and takes small particles of food
to the stomach (Figure 1.101 z). But in the craniota the thyroid gland
(thyreoidea) is developed from it, the gland that lies in front of the
larynx, and which, when pathologically enlarged, forms goitre
(struma).

From the head-gut we get not only the gills, the organs of
water-breathing in the lower vertebrates, but also the lungs, the
organs of atmospheric breathing in the five higher classes. In these
cases a vesicular fold appears in the gullet of the embryo at an early
stage, and gradually takes the shape of two spacious sacs, which are
afterwards filled with air. These sacs are the two air-breathing
lungs, which take the place of the water-breathing gills. But the
vesicular invagination, from which the lungs arise, is merely the
familiar air-filled vesicle, which we call the floating-bladder of the
fish, and which alters its specific weight, acting as hydrostatic
organ or floating apparatus. This structure is not found in the lowest
vertebrate classes--the acrania and cyclostoma. We shall see more of
it in Volume 2.

The second chief section of the vertebrate-gut, the trunk or
liver-gut, which accomplishes digestion, is of very simple
construction in the acrania. It consists of two different chambers.
The first chamber, immediately behind the gill-gut, is the expanded
stomach (ma); the second, narrower and longer chamber, is the straight
small intestine (d): it issues behind on the ventral side by the anus
(af). Near the limit of the two chambers in the visceral cavity we
find the liver, in the shape of a simple tube or blind sac (l); in the
amphioxus it is single; in the prospondylus it was probably double
(Figures 1.98 and 1.100 l).

Closely related morphologically and physiologically to the alimentary
canal is the vascular system of the vertebrate, the chief sections of
which develop from the fibrous gut-layer. It consists of two different
but directly connected parts, the system of blood-vessels and that of
lymph-vessels. In the passages of the one we find red blood, and in
the other colourless lymph. To the lymphatic system belong, first of
all, the lymphatic canals proper or absorbent veins, which are
distributed among all the organs, and absorb the used-up juices from
the tissues, and conduct them into the venous blood; but besides these
there are the chyle-vessels, which absorb the white chyle, the milky
fluid prepared by the alimentary canal from the food, and conduct this
also to the blood.

The blood-vessel system of the vertebrate has a very elaborate
construction, but seems to have had a very simple form in the
primitive vertebrate, as we find it to-day permanently in the annelids
(for instance, earth-worms) and the amphioxus. We accordingly
distinguish first of all as essential, original parts of it two large
single blood-canals, which lie in the fibrous wall of the gut, and run
along the alimentary canal in the median plane of the body, one above
and the other underneath the canal. These principal canals give out
numerous branches to all parts of the body, and pass into each other
by arches before and behind; we will call them the primitive artery
and the primitive vein. The first corresponds to the dorsal vessel,
the second to the ventral vessel, of the worms. The primitive or
principal artery, usually called the aorta (Figure 1.98 a), lies above
the gut in the middle line of its dorsal side, and conducts oxidised
or arterial blood from the gills to the body. The primitive or
principal vein (Figure 1.100 v) lies below the gut, in the middle line
of its ventral side, and is therefore also called the vena
subintestinalis; it conducts carbonised or venous blood back from the
body to the gills. At the branchial section of the gut in front the
two canals are connected by a number of branches, which rise in arches
between the gill-clefts. These "branchial vascular arches" (kg) run
along the gill-arches, and have a direct share in the work of
respiration. The anterior continuation of the principal vein which
runs on the ventral wall of the gill-gut, and gives off these vascular
arches upwards, is the branchial artery (ka). At the border of the two
sections of the ventral vessel it enlarges into a contractile
spindle-shaped tube (Figures 1.98 and 1.100 h). This is the first
outline of the heart, which afterwards becomes a four-chambered pump
in the higher vertebrates and man. There is no heart in the amphioxus,
probably owing to degeneration. In prospondylus the ventral gill-heart
probably had the simple form in which we still find it in the ascidia
and the embryos of the craniota (Figures 1.98 and 1.100 h).

The kidneys, which act as organs of excretion or urinary organs in all
vertebrates, have a very different and elaborate construction in the
various sections of this stem; we will consider them further in
Chapter 2.29. Here I need only mention that in our hypothetical
primitive vertebrate they probably had the same form as in the actual
amphioxus--the primitive kidneys (protonephra). These are originally
made up of a double row of little canals, which directly convey the
used-up juices or the urine out of the body-cavity (Figure 1.102 n).
The inner aperture of these pronephridial canals opens with a ciliated
funnel into the body-cavity; the external aperture opens in lateral
grooves of the epidermis, a couple of longitudinal grooves in the
lateral surface of the outer skin (Figure 1.102 b). The pronephridial
duct is formed by the closing of this groove to the right and left at
the sides. In all the craniota it develops at an early stage in the
horny plate; in the amphioxus it seems to be converted into a wide
cavity, the atrium, or peribranchial space.

Next to the kidneys we have the sexual organs of the vertebrate. In
most of the members of this stem the two are united in a single
urogenital system; it is only in a few groups that the urinary and
sexual organs are separated (in the amphioxus, the cyclostoma, and
some sections of the fish-class). In man and all the higher
vertebrates the sexual apparatus is made up of various parts, which we
will consider in Chapter 2.29. But in the two lowest classes of our
stem, the acrania and cyclostoma, they consist merely of simple sexual
glands or gonads, the ovaries of the female sex and the testicles
(spermaria) of the male; the former provide the ova, the latter the
sperm. In the craniota we always find only one pair of gonads; in the
amphioxus several pairs, arranged in succession. They must have had
the same form in our hypothetical prospondylus (Figures 1.98 and 1.100
s). These segmental pairs of gonads are the original ventral halves of
the coelom-pouches.

The organs which we have now enumerated in this general survey, and of
which we have noted the characteristic disposition, are those parts of
the organism that are found in all vertebrates without exception in
the same relation to each other, however much they may be modified. We
have chiefly had in view the transverse section of the body (Figures
1.101 and 1.102), because in this we see most clearly the distinctive
arrangement of them. But to complete our picture we must also consider
the segmentation or metamera-formation of them, which has yet been
hardly noticed, and which is seen best in the longitudinal section. In
man and all the more advanced vertebrates the body is made up of a
series or chain of similar members, which succeed each other in the
long axis of the body--the segments or metamera of the organism. In
man these homogeneous parts number thirty-three in the trunk, but they
run to several hundred in many of the vertebrates (such as serpents or
eels). As this internal articulation or metamerism is mainly found in
the vertebral column and the surrounding muscles, the sections or
metamera were formerly called pro-vertebrae. As a fact, the
articulation is by no means chiefly determined and caused by the
skeleton, but by the muscular system and the segmental arrangement of
the kidneys and gonads. However, the composition from these
pro-vertebrae or internal metamera is usually, and rightly, put
forward as a prominent character of the vertebrate, and the manifold
division or differentiation of them is of great importance in the
various groups of the vertebrates. But as far as our present task--the
derivation of the simple body of the primitive vertebrate from the
chordula--is concerned, the articulate parts or metamera are of
secondary interest, and we need not go into them just now.

(FIGURE 1.103 A, B, C, D. Instances of redundant mammary glands and
nipples (hypermastism). A a
pair of small redundant breasts (with two nipples on the left) above
the large normal ones; from a 45-year-old Berlin woman, who had had
children 17 times (twins twice). (From Hansemann.) B the highest
number: ten nipples (all giving milk), three pairs above, one pair
below, the large normal breasts; from a 22-year-old servant at
Warschau. (From Neugebaur.) C three pairs of nipples: two pairs on the
normal glands and one pair above; from a 19-year-old Japanese girl. D
four pairs of nipples: one pair above the normal and two pairs of
small accessory nipples underneath; from a 22-year-old Bavarian
soldier. (From Wiedersheim.))

The characteristic composition of the vertebrate body develops from
the embryonic structure in the same way in man as in all the other
vertebrates. As all competent experts now admit the monophyletic
origin of the vertebrates on the strength of this significant
agreement, and this "common descent of all the vertebrates from one
original stem-form" is admitted as an historical fact, we have found
the answer to "the question of questions." We may, moreover, point out
that this answer is just as certain and precise in the case of the
origin of man from the mammals. This advanced vertebrate class is also
monophyletic, or has evolved from one common stem-group of lower
vertebrates (reptiles, and, earlier still, amphibia). This follows
from the fact that the mammals are clearly distinguished from the
other classes of the stem, not merely in one striking particular, but
in a whole group of distinctive characters.

It is only in the mammals that we find the skin covered with hair, the
breast-cavity separated from the abdominal cavity by a complete
diaphragm, and the larynx provided with an epiglottis. The mammals
alone have three small auscultory bones in the tympanic cavity--a
feature that is connected with the characteristic modification of
their maxillary joint. Their red blood-cells have no nucleus, whereas
this is retained in all other vertebrates. Finally, it is only in the
mammals that we find the remarkable function of the breast structure
which has given its name to the whole class--the feeding of the young
by the mother's milk. The mammary glands which serve this purpose are
interesting in so many ways that we may devote a few lines to them
here.

As is well known, the lower mammals, especially those which beget a
number of young at a time, have several mammary glands at the breast.
Hedgehogs and sows have five pairs, mice four or five pairs, dogs and
squirrels four pairs, cats and bears three pairs, most of the
ruminants and many of the rodents two pairs, each provided with a teat
or nipple (mastos). In the various genera of the half-apes (lemurs)
the number varies a good deal. On the other hand, the bats and apes,
which only beget one young at a time as a rule, have only one pair of
mammary glands, and these are found at the breast, as in man.

These variations in the number or structure of the mammary apparatus
(mammarium) have become doubly interesting in the light of recent
research in comparative anatomy. It has been shown that in man and the
apes we often find redundant mammary glands (hyper-mastism) and
corresponding teats (hyper-thelism) in both sexes. Figure 1.103 shows
four cases of this kind--A, B, and C of three women, and D of a man.
They prove that all the above-mentioned numbers may be found
occasionally in man. Figure 1.103 A shows the breast of a Berlin woman
who had had children seventeen times, and who has a pair of small
accessory breasts (with two nipples on the left one) above the two
normal breasts; this is a common occurrence, and the small soft pad
above the breast is not infrequently represented in ancient statues of
Venus. In Figure 1.103 C we have the same phenomenon in a Japanese
girl of nineteen, who has two nipples on each breast besides (three
pairs altogether). Figure 1.103 D is a man of twenty-two with four
pairs of nipples (as in the dog), a small pair above and two small
pairs beneath the large normal teats. The maximum number of five pairs
(as in the sow and hedgehog) was found in a Polish servant of
twenty-two who had had several children; milk was given by each
nipple; there were three pairs of redundant nipples above and one pair
underneath the normal and very large breasts (Figure 1.103 B).

A number of recent investigations (especially among recruits) have
shown that these things are not uncommon in the male as well as the
female sex. They can only be explained by evolution, which attributes
them to atavism and latent heredity. The earlier ancestors of all the
primates (including man) were lower placentals, which had, like the
hedgehog (one of the oldest forms of the living placentals), several
mammary glands (five or more pairs) in the abdominal skin. In the apes
and man only a couple of them are normally developed, but from time to
time we get a development of the atrophied structures. Special notice
should be taken of the arrangement of these accessory mammae; they
form, as is clearly seen in Figure 1.103 B and D, two long rows, which
diverge forward (towards the arm-pit), and converge behind in the
middle line (towards the loins). The milk-glands of the polymastic
lower placentals are arranged in similar lines.

The phylogenetic explanation of polymastism, as given in comparative
anatomy, has lately found considerable support in ontogeny. Hans
Strahl, E. Schmitt, and others, have found that there are always in
the human embryo at the sixth week (when it is three-fifths of an inch
long) the microscopic traces of five pairs of mammary glands, and that
they are arranged at regular distances in two lateral and divergent
lines, which correspond to the mammary lines. Only one pair of
them--the central pair--are normally developed, the others atrophying.
Hence there is for a time in the human embryo a normal hyperthelism,
and this can only be explained by the descent of man from lower
primates (lemurs) with several pairs.

But the milk-gland of the mammal has a great morphological interest
from another point of view. This organ for feeding the young in man
and the higher mammals is, as is known, found in both sexes. However,
it is usually active only in the female sex, and yields the valuable
"mother's milk"; in the male sex it is small and inactive, a real
rudimentary organ of no physiological interest. Nevertheless, in
certain cases we find the breast as fully developed in man as in
woman, and it may give milk for feeding the young.

(FIGURE 1.104. A Greek gynecomast.)

We have a striking instance of this gynecomastism (large milk-giving
breasts in a male) in Figure 1.104. I owe the photograph (taken from
life) to the kindness of Dr. Ornstein, of Athens, a German physician,
who has rendered service by a number of anthropological observations,
(for instance, in several cases of tailed men). The gynecomast in
question is a Greek recruit in his twentieth year, who has both
normally developed male organs and very pronounced female breasts. It
is noteworthy that the other features of his structure are in accord
with the softer forms of the female sex. It reminds us of the marble
statues of hermaphrodites which the ancient Greek and Roman sculptors
often produced. But the man would only be a real hermaphrodite if he
had ovaries internally besides the (externally visible) testicles.

I observed a very similar case during my stay in Ceylon (at
Belligemma) in 1881. A young Cinghalese in his twenty-fifth year was
brought to me as a curious hermaphrodite, half-man and half-woman. His
large breasts gave plenty of milk; he was employed as "male nurse" to
suckle a new-born infant whose mother had died at birth. The outline
of his body was softer and more feminine than in the Greek shown in
Figure 1.104. As the Cinghalese are small of stature and of graceful
build, and as the men often resemble the women in clothing (upper part
of the body naked, female dress on the lower part) and the dressing of
the hair (with a comb), I first took the beardless youth to be a
woman. The illusion was greater, as in this remarkable case
gynecomastism was associated with cryptorchism--that is to say, the
testicles had kept to their original place in the visceral cavity, and
had not travelled in the normal way down into the scrotum. (Cf.
Chapter 2.29.) Hence the latter was very small, soft, and empty.
Moreover, one could feel nothing of the testicles in the inguinal
canal. On the other hand, the male organ was very small, but normally
developed. It was clear that this apparent hermaphrodite also was a
real male.

Another case of practical gynecomastism has been described by
Alexander von Humboldt. In a South American forest he found a solitary
settler whose wife had died in child-birth. The man had laid the
new-born child on his own breast in despair; and the continuous
stimulus of the child's sucking movements had revived the activity of
the mammary glands. It is possible that nervous suggestion had some
share in it. Similar cases have been often observed in recent years,
even among other male mammals (such as sheep and goats).

The great scientific interest of these facts is in their bearing on
the question of heredity. The stem-history of the mammarium rests
partly on its embryology (Chapter 2.24.) and partly on the facts of
comparative anatomy and physiology. As in the lower and higher mammals
(the monotremes, and most of the marsupials) the whole lactiferous
apparatus is only found in the female; and as there are traces of it
in the male only in a few younger marsupials, there can be no doubt
that these important organs were originally found only in the female
mammal, and that they were acquired by these through a special
adaptation to habits of life.

Later, these female organs were communicated to both sexes by
heredity; and they have been maintained in all persons of either sex,
although they are not physiologically active in the males. This normal
permanence of the female lactiferous organs in BOTH sexes of the
higher mammals and man is independent of any selection, and is a fine
instance of the much-disputed "inheritance of acquired characters."


CHAPTER 1.12. EMBRYONIC SHIELD AND GERMINATIVE AREA.

The three higher classes of vertebrates which we call the
amniotes--the mammals, birds, and reptiles--are notably distinguished
by a number of peculiarities of their development from the five lower
classes of the stem--the animals without an amnion (the anamnia). All
the amniotes have a distinctive embryonic membrane known as the amnion
(or "water-membrane"), and a special embryonic appendage--the
allantois. They have, further, a large yelk-sac, which is filled with
food-yelk in the reptiles and birds, and with a corresponding clear
fluid in the mammals. In consequence of these later-acquired
structures, the original features of the development of the amniotes
are so much altered that it is very difficult to reduce them to the
palingenetic embryonic processes of the lower amnion-less vertebrates.
The gastraea theory shows us how to do this, by representing the
embryology of the lowest vertebrate, the skull-less amphioxus, as the
original form, and deducing from it, through a series of gradual
modifications, the gastrulation and coelomation of the craniota.

It was somewhat fatal to the true conception of the chief embryonic
processes of the vertebrate that all the older embryologists, from
Malpighi (1687) and Wolff (1750) to Baer (1828) and Remak (1850),
always started from the investigation of the hen's egg, and
transferred to man and the other vertebrates the impressions they
gathered from this. This classical object of embryological research
is, as we have seen, a source of dangerous errors. The large round
food-yelk of the bird's egg causes, in the first place, a flat discoid
expansion of the small gastrula, and then so distinctive a development
of this thin round embryonic disk that the controversy as to its
significance occupies a large part of embryological literature.

(FIGURE 1.105. Severance of the discoid mammal embryo from the
yelk-sac, in transverse section (diagrammatic). A The germinal disk
(h, hf) lies flat on one side of the branchial-gut vesicle (kb). B In
the middle of the germinal disk we find the medullary groove (mr), and
underneath it the chorda (ch). C The gut-fibre-layer (df) has been
enclosed by the gut-gland-layer (dd). D The skin-fibre-layer (hf) and
gut-fibre-layer (df) divide at the periphery; the gut (d) begins to
separate from the yelk-sac or umbilical vesicle (nb). E The medullary
tube (mr) is closed; the body-cavity (c) begins to form. F The
provertebrae (w) begin to grow round the medullary tube (mr) and the
chorda (ch): the gut (d) is cut off from the umbilical vesicle (nb). H
The vertebrae (w) have grown round the medullary tube (mr) and chorda;
the body-cavity is closed, and the umbilical vesicle has disappeared.
The amnion and serous membrane are omitted. The letters have the same
meaning throughout: h horn-plate, mr medullary tube, hf
skin-fibre-layer, w provertebrae, ch chorda, c body-cavity or coeloma,
df gut-fibre-layer, dd gut-gland-layer, d gut-cavity, nb umbilical
vesicle.)

One of the most unfortunate errors that this led to was the idea of an
original antithesis of germ and yelk. The latter was regarded as a
foreign body, extrinsic to the real germ, whereas it is properly a
part of it, an embryonic organ of nutrition. Many authors said there
was no trace of the embryo until a later stage, and outside the yelk;
sometimes the two-layered embryonic disk itself, at other times only
the central portion of it (as distinguished from the germinative area,
which we will describe presently), was taken to be the first outline
of the embryo. In the light of the gastraea theory it is hardly
necessary to dwell on the defects of this earlier view and the
erroneous conclusions drawn from it. In reality, the first
segmentation-cell, and even the stem-cell itself and all that issues
therefrom, belong to the embryo. As the large original yelk-mass in
the undivided egg of the bird only represents an inclosure in the
greatly enlarged ovum, so the later contents of its embryonic yelk-sac
(whether yet segmented or not) are only a part of the entoderm which
forms the primitive gut. This is clearly shown by the ova of the
amphibia and cyclostoma, which explain the transition from the
yelk-less ova of the amphioxus to the large yelk-filled ova of the
reptiles and birds.

It is precisely in the study of these difficult features that we see
the incalculable value of phylogenetic considerations in explaining
complex ontogenetic facts, and the need of separating cenogenetic
phenomena from palingenetic. This is particularly clear as regards the
comparative embryology of the vertebrates, because here the
phylogenetic unity of the stem has been already established by the
well-known facts of paleontology and comparative anatomy. If this
unity of the stem, on the basis of the amphioxus, were always borne in
mind, we should not have these errors constantly recurring.

In many cases the cenogenetic relation of the embryo to the food-yelk
has until now given rise to a quite wrong idea of the first and most
important embryonic processes in the higher vertebrates, and has
occasioned a number of false theories in connection with them. Until
thirty years ago the embryology of the higher vertebrates always
started from the position that the first structure of the embryo is a
flat, leaf-shaped disk; it was for this reason that the cell-layers
that compose this germinal disk (also called germinative area) are
called "germinal layers." This flat germinal disk, which is round at
first and then oval, and which is often described as the tread or
cicatricula in the laid hen's egg, is found at a certain part of the
surface of the large globular food-yelk. I am convinced that it is
nothing else than the discoid, flattened gastrula of the birds. At the
beginning of germination the flat embryonic disk curves outwards, and
separates on the inner side from the underlying large yelk-ball. In
this way the flat layers are converted into tubes, their edges folding
and joining together (Figure 1.105). As the embryo grows at the
expense of the food-yelk, the latter becomes smaller and smaller; it
is completely surrounded by the germinal layers. Later still, the
remainder of the food-yelk only forms a small round sac, the yelk-sac
or umbilical vesicle (Figure 1.105 nb). This is enclosed by the
visceral layer, is connected by a thin stalk, the yelk-duct, with the
central part of the gut-tube, and is finally, in most of the
vertebrates, entirely absorbed by this (H). The point at which this
takes place, and where the gut finally closes, is the visceral navel.
In the mammals, in which the remainder of the yelk-sac remains without
and atrophies, the yelk-duct at length penetrates the outer ventral
wall. At birth the umbilical cord proceeds from here, and the point of
closure remains throughout life in the skin as the navel.

As the older embryology of the higher vertebrates was mainly based on
the chick, and regarded the antithesis of embryo (or formative-yelk)
and food-yelk (or yelk-sac) as original, it had also to look upon the
flat leaf-shaped structure of the germinal disk as the primitive
embryonic form, and emphasise the fact that hollow grooves were formed
of these flat layers by folding, and closed tubes by the joining
together of their edges.

This idea, which dominated the whole treatment of the embryology of
the higher vertebrates until thirty years ago, was totally false. The
gastraea theory, which has its chief application here, teaches us that
it is the very reverse of the truth. The cup-shaped gastrula, in the
body-wall of which the two primary germinal layers appear from the
first as closed tubes, is the original embryonic form of all the
vertebrates, and all the multicellular invertebrates; and the flat
germinal disk with its superficially expanded germinal layers is a
later, secondary form, due to the cenogenetic formation of the large
food-yelk and the gradual spread of the germ-layers over its surface.
Hence the actual folding of the germinal layers and their conversion
into tubes is not an original and primary, but a much later and
tertiary, evolutionary process. In the phylogeny of the vertebrate
embryonic process we may distinguish the following three stages:--

   A. First Stage: Primary (palingenetic) embryonic process.

The germinal layers form from the first closed tubes, the one-layered
blastula being converted into the two-layered gastrula by
invagination. No food-yelk. (Amphioxus.)

B. Second Stage: Secondary (cenogenetic) embryonic process.

The germinal layers spread out leaf-wise, food-yelk gathering in the
ventral entoderm, and a large yelk-sac being formed from the middle of
the gut-tube. (Amphibia.)

C. Third Stage: Tertiary (cenogenetic) embryonic process.

The germinal layers form a flat germinal disk, the borders of which
join together and form closed tubes, separating from the central
yelk-sac. (Amniotes.)

As this theory, a logical conclusion from the gastraea theory, has
been fully substantiated by the comparative study of gastrulation in
the last few decades, we must exactly reverse the hitherto prevalent
mode of treatment. The yelk-sac is not to be treated, as was done
formerly, as if it were originally antithetic to the embryo, but as an
essential part of it, a part of its visceral tube. The primitive gut
of the gastrula has, on this view, been divided into two parts in the
higher animals as a result of the cenogenetic formation of the
food-yelk--the permanent gut (metagaster), or permanent alimentary
canal, and the yelk-sac (lecithoma), or umbilical vesicle. This is
very clearly shown by the comparative ontogeny of the fishes and
amphibia. In these cases the whole yelk undergoes cleavage at first,
and forms a yelk-gland, composed of yelk-cells, in the ventral wall of
the primitive gut. But it afterwards becomes so large that a part of
the yelk does not divide, and is used up in the yelk-sac that is cut
off outside.

(FIGURE 1.106. The visceral embryonic vesicle (blastocystis or
gastrocystis) of a rabbit (the "blastula" or vesicula blastodermica of
other writers), a outer envelope (ovolemma), b skin-layer or ectoderm,
forming the entire wall of the yelk-vesicle, c groups of dark cells,
representing the visceral layer or entoderm.

FIGURE 1.107. The same in section. Letters as above. d cavity of the
vesicle. (From Bischoff.))

When we make a comparative study of the embryology of the amphioxus,
the frog, the chick, and the rabbit, there cannot, in my opinion, be
any further doubt as to the truth of this position, which I have held
for thirty years. Hence in the light of the gastraea theory we must
regard the features of the amphioxus as the only and real primitive
structure among all the vertebrates, departing very little from the
palingenetic embryonic form. In the cyclostoma and the frog these
features are, on the whole, not much altered cenogenetically, but they
are very much so in the chick, and most of all in the rabbit. In the
bell-gastrula of the amphioxus and in the hooded gastrula of the
lamprey and the frog the germinal layers are found to be closed tubes
or vesicles from the first. On the other hand, the chick-embryo (in
the new laid, but not yet hatched, egg) is a flat circular disk, and
it was not easy to recognise this as a real gastrula. Rauber and
Goette have, however, achieved this. As the discoid gastrula grows
round the large globular yelk, and the permanent gut then separates
from the outlying yelk-sac, we find all the processes which we have
shown (diagrammatically) in Figure 1.108--processes that were hitherto
regarded as principal acts, whereas they are merely secondary.

The oldest, oviparous mammals, the monotremes, behave in the same way
as the reptiles and birds. But the corresponding embryonic processes
in the viviparous mammals, the marsupials and placentals, are very
elaborate and distinctive. They were formerly quite misinterpreted; it
was not until the publication of the studies of Edward van Beneden
(1875) and the later research of Selenka, Kuppfer, Rabl, and others,
that light was thrown on them, and we were in a position to bring them
into line with the principles of the gastraea theory and trace them to
the embryonic forms of the lower vertebrates. Although there is no
independent food-yelk, apart from the formative yelk, in the mammal
ovum, and although its segmentation is total on that account,
nevertheless a large yelk-sac is formed in their embryos, and the
"embryo proper" spreads leaf-wise over its surface, as in the reptiles
and birds, which have a large food-yelk and partial segmentation. In
the mammals, as well as in the latter, the flat, leaf-shaped germinal
disk separates from the yelk-sac, and its edges join together and form
tubes.

How can we explain this curious anomaly? Only as a result of very
characteristic and peculiar cenogenetic modifications of the embryonic
process, the real causes of which must be sought in the change in the
rearing of the young on the part of the viviparous mammals. These are
clearly connected with the fact that the ancestors of the viviparous
mammals were oviparous amniotes like the present monotremes, and only
gradually became viviparous. This can no longer be questioned now that
it has been shown (1884) that the monotremes, the lowest and oldest of
the mammals, still lay eggs, and that these develop like the ova of
the reptiles and birds. Their nearest descendants, the marsupials,
formed the habit of retaining the eggs, and developing them in the
oviduct; the latter was thus converted into a womb (uterus). A
nutritive fluid that was secreted from its wall, and passed through
the wall of the blastula, now served to feed the embryo, and took the
place of the food-yelk. In this way the original food-yelk of the
monotremes gradually atrophied, and at last disappeared so completely
that the partial ovum-segmentation of their descendants, the rest of
the mammals, once more became total. From the discogastrula of the
former was evolved the distinctive epigastrula of the latter.

It is only by this phylogenetic explanation that we can understand the
formation and development of the peculiar, and hitherto totally
misunderstood, blastula of the mammal. The vesicular condition of the
mammal embryo was discovered 200 years ago (1677) by Regner de Graaf.
He found in the uterus of a rabbit four days after impregnation small,
round, loose, transparent vesicles, with a double envelope. However,
Graaf's discovery passed without recognition. It was not until 1827
that these vesicles were rediscovered by Baer, and then more closely
studied in 1842 by Bischoff in the rabbit (Figures 1.106 and 1.107).
They are found in the womb of the rabbit, the dog, and other small
mammals, a few days after copulation. The mature ova of the mammal,
when they have left the ovary, are fertilised either here or in the
oviduct immediately afterwards by the invading sperm-cells.* (* In man
and the other mammals the fertilisation of the ova probably takes
place, as a rule, in the oviduct; here the ova, which issue from the
female ovary in the shape of the Graafian follicle, and enter the
inner aperture of the oviduct, encounter the mobile sperm-cells of the
male seed, which pass into the uterus at copulation, and from this
into the external aperture of the oviduct. Impregnation rarely takes
place in the ovary or in the womb.) (As to the womb and oviduct see
Chapter 2.29.) The cleavage and formation of the gastrula take place
in the oviduct. Either here in the oviduct or after the mammal
gastrula has passed into the uterus it is converted into the globular
vesicle which is shown externally in Figure 1.106, and in section in
Figure 1.107. The thick, outer, structureless envelope that encloses
it is the original ovolemma or zona pellucida, modified, and clothed
with a layer of albumin that has been deposited on the outside. From
this stage the envelope is called the external membrane, the primary
chorion or prochorion (a). The real wall of the vesicle enclosed by it
consists of a simple layer of ectodermic cells (b), which are
flattened by mutual pressure, and generally hexagonal; a light nucleus
shines through their fine-grained protoplasm (Figure 1.108). At one
part (c) inside this hollow ball we find a circular disc, formed of
darker, softer, and rounder cells, the dark-grained entodermic cells
(Figure 1.109).

(FIGURE 1.108. Four entodermic cells from the embryonic vesicle of the
rabbit.

FIGURE 1.109. Two entodermic cells from the embryonic vesicle of the
rabbit.)

The characteristic embryonic form that the developing mammal now
exhibits has up to the present usually been called the "blastula"
(Bischoff), "sac-shaped embryo" (Baer), "vesicular embryo" (vesicula
blastodermica, or, briefly, blastosphaera). The wall of the hollow
vesicle, which consists of a single layer of cells, was called the
"blastoderm," and was supposed to be equivalent to the cell-layer of
the same name that forms the wall of the real blastula of the
amphioxus and many of the invertebrates (such as Monoxenia, Figure
1.29 F, G). Formerly this real blastula was generally believed to be
equivalent to the embryonic vesicle of the mammal. However, this is by
no means the case. What is called the "blastula" of the mammal and the
real blastula of the amphioxus and many of the invertebrates are
totally different embryonic structures. The latter (blastula) is
palingenetic, and precedes the formation of the gastrula. The former
(blastodermic vesicle) is cenogenetic, and follows gastrulation. The
globular wall of the blastula is a real blastoderm, and consists of
homogeneous (blastodermic) cells; it is not yet differentiated into
the two primary germinal layers. But the globular wall of the mammal
vesicle is the differentiated ectoderm, and at one point in it we find
a circular disk of quite different cells--the entoderm. The round
cavity, filled with fluid, inside the real blastula is the
segmentation-cavity. But the similar cavity within the mammal vesicle
is the yelk-sac cavity, which is connected with the incipient
gut-cavity. This primitive gut-cavity passes directly into the
segmentation-cavity in the mammals, in consequence of the peculiar
cenogenetic changes in their gastrulation, which we have considered
previously (Chapter 1.9). For these reasons it is very necessary to
recognise the secondary embryonic vesicle in the mammal (gastrocystis
or blastocystis) as a characteristic structure peculiar to this class,
and distinguish it carefully from the primary blastula of the
amphioxus and the invertebrates.

(FIGURE 1.110. Ovum of a rabbit from the uterus, one sixth of an inch
in diameter. The embryonic vesicle (b) has withdrawn a little from the
smooth ovolemma (a). In the middle of the ovolemma we see the round
germinal disk (blastodiscus, c), at the edge of which (at d) the inner
layer of the embryonic vesicle is already beginning to expand.
(Figures 1.110 to 1.114 from Bischoff.)

FIGURE 1.111. The same ovum, seen in profile. Letters as in Figure
1.110.

FIGURE 1.112. Ovum of a rabbit from the uterus, one-fourth of an inch
in diameter. The blastoderm is already for the most part two-layered
(b). The ovolemma, or outer envelope, is tufted (a).

FIGURE 1.113. The same ovum, seen in profile. Letters as in Figure
1.112.

FIGURE 1.114. Ovum of a rabbit from the uterus, one-third of an inch
in diameter. The embryonic vesicle is now nearly everywhere
two-layered (k) only remaining one-layered below (at d).

FIGURE 1.115. Round germinative area of the rabbit, divided into the
central light area (area pellucida) and the peripheral dark area (area
opaca). The light area seems darker on account of the dark ground
appearing through it.)

The small, circular, whitish, and opaque spot which the gastric disk
(Figure 1.106) forms at a certain part of the surface of the clear and
transparent embryonic vesicle has long been known to science, and
compared to the germinal disk of the birds and reptiles. Sometimes it
has been called the germinal disk, sometimes the germinal spot, and
usually the germinative area. From the area the further development of
the embryo proceeds. However, the larger part of the embryonic vesicle
of the mammal is not directly used for building up the later body, but
for the construction of the temporary umbilical vesicle. The embryo
separates from this in proportion as it grows at its expense; the two
are only connected by the yelk-duct (the stalk of the yelk-sac), and
this maintains the direct communication between the cavity of the
umbilical vesicle and the forming visceral cavity (Figure 1.105).

The germinative area or gastric disk of the animal consists at first
(like the germinal disk of birds and reptiles) merely of the two
primary germinal layers, the ectoderm and entoderm. But soon there
appears in the middle of the circular disk between the two a third
stratum of cells, the rudiment of the middle layer or fibrous layer
(mesoderm). This middle germinal layer consists from the first, as we
have seen in Chapter 1.10, of two separate epithelial plates, the two
layers of the coelom-pouches (parietal and visceral). However, in all
the amniotes (on account of the large formation of yelk) these thin
middle plates are so firmly pressed together that they seem to
represent a single layer. It is thus peculiar to the amniotes that the
middle of the germinative area is composed of four germinal layers,
the two limiting (or primary) layers and the middle layers between
them (Figures 1.96 and 1.97). These four secondary germinal layers can
be clearly distinguished as soon as what is called the sickle-groove
(or "embryonic sickle") is seen at the hind border of the germinative
area. At the borders, however, the germinative area of the mammal only
consists of two layers. The rest of the wall of the embryonic vesicle
consists at first (but only for a short time in most of the mammals)
of a single layer, the outer germinal layer.

(FIGURE 1.116. Oval area, with the opaque whitish border of the dark
area without.)

From this stage, however, the whole wall of the embryonic vesicle
becomes two-layered. The middle of the germinative area is much
thickened by the growth of the cells of the middle layers, and the
inner layer expands at the same time, and increases at the border of
the disk all round. Lying close on the outer layer throughout, it
grows over its inner surface at all points, covers first the upper and
then the lower hemisphere, and at last closes in the middle of the
inner layer (Figures 1.110 to 1.114). The wall of the embryonic
vesicle now consists throughout of two layers of cells, the ectoderm
without and the entoderm within. It is only in the centre of the
circular area, which becomes thicker and thicker through the growth of
the middle layers, that it is made up of all four layers. At the same
time, small structureless tufts or warts are deposited on the surface
of the outer ovolemma or prochorion, which has been raised above the
embryonic vesicle (Figures 1.112 to 1.114 a).

(FIGURE 1.117. Oval germinal disk of the rabbit, magnified about ten
times. As the delicate, half-transparent disk lies on a black ground,
the pellucid area looks like a dark ring, and the opaque area (lying
outside it) like a white ring. The oval shield in the centre also
looks whitish, and in its axis we see the dark medullary groove. (From
Bischoff.))

We may now disregard both the outer ovolemma and the greater part of
the vesicle, and concentrate our attention on the germinative area and
the four-layered embryonic disk. It is here alone that we find the
important changes which lead to the differentiation of the first
organs. It is immaterial whether we examine the germinative area of
the mammal (the rabbit, for instance) or the germinal disk of a bird
or a reptile (such as a lizard or tortoise). The embryonic processes
we are now going to consider are essentially the same in all members
of the three higher classes of vertebrates which we call the amniotes.
Man is found to agree in this respect with the rabbit, dog, ox, etc.;
and in all these animals the germinative area undergoes essentially
the same changes as in the birds and reptiles. They are most
frequently and accurately studied in the chick, because we can have
incubated hens' eggs in any quantity at any stage of development.
Moreover, the round germinal disk of the chick passes immediately
after the beginning of incubation (within a few hours) from the
two-layered to the four-layered stage, the two-layered mesoderm
developing from the median primitive groove between the ectoderm and
entoderm (Figures 1.82 to 1.95).

The first change in the round germinal disk of the chick is that the
cells at its edges multiply more briskly, and form darker nuclei in
their protoplasm. This gives rise to a dark ring, more or less sharply
set off from the lighter centre of the germinal disk (Figure 1.115).
From this point the latter takes the name of the "light area" (area
pellucida), and the darker ring is called the "dark area" (area
opaca). (In a strong light, as in Figures 1.115 to 1.117, the light
area seems dark, because the dark ground is seen through it; and the
dark area seems whiter). The circular shape of the area now changes
into elliptic, and then immediately into oval (Figures 1.116 and
1.117). One end seems to be broader and blunter, the other narrower
and more pointed; the former corresponds to the anterior and the
latter to the posterior section of the subsequent body. At the same
time, we can already trace the characteristic bilateral form of the
body, the antithesis of right and left, before and behind. This will
be made clearer by the "primitive streak," which appears at the
posterior end.

(FIGURE 1.118. Pear-shaped germinal shield of the rabbit (eight days
old), magnified twenty times. rf medullary groove. pr primitive groove
(primitive mouth). (From Kolliker.)

FIGURE 1.119. Median longitudinal section of the gastrula of four
vertebrates. (From Rabl.) A discogastrula of a shark (Pristiurus). B
amphigastrula of a sturgeon (Accipenser). C amphigastrula of an
amphibium (Triton). D epigastrula of an amniote (diagram). a ventral,
b dorsal lip of the primitive mouth.)

At an early stage an opaque spot is seen in the middle of the clear
germinative area, and this also passes from a circular to an oval
shape. At first this shield-shaped marking is very delicate and barely
perceptible; but it soon becomes clearer, and now stands out as an
oval shield, surrounded by two rings or areas (Figure 1.117). The
inner and brighter ring is the remainder of the pellucid area, and the
dark outer ring the remainder of the opaque area; the opaque
shield-like spot itself is the first rudiment of the dorsal part of
the embryo. We give it briefly the name of embryonic shield or dorsal
shield. In most works this embryonic shield is described as "the first
rudiment or trace of the embryo," or "primitive embryo." But this is
wrong, though it rests on the authority of Baer and Bischoff. As a
matter of fact, we already have the embryo in the stem-cell, the
gastrula, and all the subsequent stages. The embryonic shield is
simply the first rudiment of the dorsal part, which is the earliest to
develop. As the older names of "embryonic rudiment" and "germinative
area" are used in many different senses--and this has led to a fatal
confusion in embryonic literature--we must explain very clearly the
real significance of these important embryonic parts of the amniote.
It will be useful to do so in a series of formal principles:--

1. The so-called "first trace of the embryo" in the amniotes, or the
embryonic shield, in the centre of the pellucid area, consists merely
of an early differentiation and formation of the middle dorsal parts.

2. Hence the best name for it is "the dorsal shield," as I proposed
long ago.

3. The germinative area, in which the first embryonic blood-vessels
appear at an early stage, is not opposed as an external area to the
"embryo proper," but is a part of it.

4. In the same way, the yelk-sac or the umbilical vesicle is not a
foreign external appendage of the embryo, but an outlying part of its
primitive gut.

5. The dorsal shield gradually separates from the germinative area and
the yelk-sac, its edges growing downwards and folding together to form
ventral plates.

6. The yelk-sac and vessels of the germinative area, which soon spread
over its whole surface, are, therefore, real embryonic organs, or
temporary parts of the embryo, and have a transitory importance in
connection with the nutrition of the growing later body; the latter
may be called the "permanent body" in contrast to them.

The relation of these cenogenetic features of the amniotes to the
palingenetic structures of the older non-amniotic vertebrates may be
expressed in the following theses: The original gastrula, which
completely passes into the embryonic body in the acrania, cyclostoma,
and amphibia, is early divided into two parts in the amniotes--the
embryonic shield, which represents the dorsal outline of the permanent
body; and the temporary embryonic organs of the germinative area and
its blood-vessels, which soon grow over the whole of the yelk-sac. The
differences which we find in the various classes of the vertebrate
stem in these important particulars can only be fully understood when
we bear in mind their phylogenetic relations on the one hand, and, on
the other, the cenogenetic modifications of structure that have been
brought about by changes in the rearing of the young and the variation
in the mass of the food-yelk.

We have already described in Chapter 1.9 the changes which this
increase and decrease of the nutritive yelk causes in the form of the
gastrula, and especially in the situation and shape of the primitive
mouth. The primitive mouth or prostoma is originally a simple round
aperture at the lower pole of the long axis; its dorsal lip is above
and ventral lip below. In the amphioxus this primitive mouth is a
little eccentric, or shifted to the dorsal side (Figure 1.39). The
aperture increases with the growth of the food-yelk in the cyclostoma
and ganoids; in the sturgeon it lies almost on the equator of the
round ovum, the ventral lip (a) in front and the dorsal lip (b) behind
(Figure 1.119 b). In the wide-mouthed, circular discoid gastrula of
the selachii or primitive fishes, which spreads quite flat on the
large food-yelk, the anterior semi-circle of the border of the disk is
the ventral, and the posterior semicircle the dorsal lip (Figure 1.119
A). The amphiblastic amphibia are directly connected with their
earlier fish-ancestors, the dipneusts and ganoids, and further the
oldest selachii (Cestracion); they have retained their total unequal
segmentation, and their small primitive mouth (Figure 1.119 C, ab),
blocked up by the yelk-stopper, lies at the limit of the dorsal and
ventral surface of the embryo (at the lower pole of its equatorial
axis), and there again has an upper dorsal and a lower ventral lip (a,
b). The formation of a large food-yelk followed again in the
stem-forms of the amniotes, the protamniotes or proreptilia, descended
from the amphibia (Figure 1.119 D). But here the accumulation of the
food-yelk took place only in the ventral wall of the primitive-gut, so
that the narrow primitive mouth lying behind was forced upwards, and
came to lie on the back of the discoid "epigastrula" in the shape of
the "primitive groove"; thus (in contrast to the case of the selachii,
Figure 1.119 A) the dorsal lip (b) had to be in front, and the ventral
lip (a) behind (Figure 1.119 D). This feature was transmitted to all
the amniotes, whether they retained the large food-yelk (reptiles,
birds, and monotremes), or lost it by atrophy (the viviparous
mammals).

This phylogenetic explanation of gastrulation and coelomation, and the
comparative study of them in the various vertebrates, throw a clear
and full light on many ontogenetic phenomena, as to which the most
obscure and confused opinions were prevalent thirty years ago. In this
we see especially the high scientific value of the biogenetic law and
the careful separation of palingenetic from cenogenetic processes. To
the opponents of this law the real explanation of these remarkable
phenomena is impossible. Here, and in every other part of embryology,
the true key to the solution lies in phylogeny.


CHAPTER 1.13. DORSAL BODY AND VENTRAL BODY.

The earliest stages of the human embryo are, for the reasons already
given, either quite unknown or only imperfectly known to us. But as
the subsequent embryonic forms in man behave and develop just as they
do in all the other mammals, there cannot be the slightest doubt that
the preceding stages also are similar. We have been able to see in the
coelomula of the human embryo (Figure 1.97), by transverse sections
through its primitive mouth, that its two coelom-pouches are developed
in just the same way as in the rabbit (Figure 1.96); moreover, the
peculiar course of the gastrulation is just the same.

(FIGURE 1.120. Embryonic vesicle of a seven-days-old rabbit with oval
embryonic shield (ag).
A seen from above, B from the side. (From Kolliker.) ag dorsal shield
or embryonic spot. In B the upper half of the vesicle is made up of
the two primary germinal layers, the lower (up to ge) only from the
outer layer.)

The germinative area forms in the human embryo in the same way as in
the other mammals, and in the middle part of this we have the
embryonic shield, the purport of which we considered in Chapter 1.12.
The next changes in the embryonic disk, or the "embryonic spot," take
place in corresponding fashion. These are the changes we are now going
to consider more closely.

The chief part of the oval embryonic shield is at first the narrow
hinder end; it is in the middle line of this that the primitive streak
appears (Figure 1.121 ps). The narrow longitudinal groove in it--the
so-called "primitive groove"--is, as we have seen, the primitive mouth
of the gastrula. In the gastrula-embryos of the mammals, which are
much modified cenogenetically, this cleft-shaped prostoma is
lengthened so much that it soon traverses the whole of the hinder half
of the dorsal shield; as we find in a rabbit embryo of six to eight
days (Figure 1.122 pr). The two swollen parallel borders that limit
this median furrow are the side lips of the primitive mouth, right and
left. In this way the bilateral-symmetrical type of the vertebrate
becomes pronounced. The subsequent head of the amniote is developed
from the broader and rounder fore-half of the dorsal shield.

In this fore-half of the dorsal shield a median furrow quickly makes
its appearance (Figure 1.123 rf). This is the broader dorsal furrow or
medullary groove, the first beginning of the central nervous system.
The two parallel dorsal or medullary swellings that enclose it grow
together over it afterwards, and form the medullary tube. As is seen
in transverse sections, it is formed only of the outer germinal layer
(Figures 1.95 and 1.136). The lips of the primitive mouth, however,
lie, as we know, at the important point where the outer layer bends
over the inner, and from which the two coelom pouches grow between the
primary germinal layers.

(FIGURE 1.121. Oval embryonic shield of the rabbit (A of six days
eighteen hours, B of eight days). (From Kolliker.) ps primitive
streak, pr primitive groove, arg area germinalis, sw sickle-shaped
germinal growth.

FIGURE 1.122. Dorsal shield (ag) and germinative area of a
rabbit-embryo of eight days. (From Kolliker.) pr primitive groove, rf
dorsal furrow.

FIGURE 1.123. Embryonic shield of a rabbit of eight days. (From Van
Beneden.) pr primitive groove, cn canalis neurentericus, nk nodus
neurentericus (or "Hensen's ganglion"), kf head-process (chorda).

FIGURE 1.124. Longitudinal section of the coelomula of amphioxus (from
the left). i entoderm, d primitive gut, cn medullary duct, n nerve
tube, m mesoderm, s first primitive segment, c coelom-pouches. (From
Hatschek.))

Thus the median primitive furrow (pr) in the hind-half and the median
medullary furrow (rf) in the fore-half of the oval shield are totally
different structures, although the latter seems to a superficial
observer to be merely the forward continuation of the former. Hence
they were formerly always confused. This error was the more pardonable
as immediately afterwards the two grooves do actually pass into each
other in a very remarkable way. The point of transition is the
remarkable neurenteric canal (Figure 1.124 cn). But the direct
connection which is thus established does not last long; the two are
soon definitely separated by a partition.

The enigmatic neurenteric canal is a very old embryonic organ, and of
great phylogenetic interest, because it arises in the same way in all
the chordonia (both tunicates and vertebrates). In every case it
touches or embraces like an arch the posterior end of the chorda,
which has been developed here in front out of the middle line of the
primitive gut (between the two coelom-folds of the sickle groove)
("head-process," Figure 1.123 kf). These very ancient and strictly
hereditary structures, which have no physiological significance
to-day, deserve (as "rudimentary organs") our closest attention. The
tenacity with which the useless neurenteric canal has been transmitted
down to man through the whole series of vertebrates is of equal
interest for the theory of descent in general, and the phylogeny of
the chordonia in particular.

The connection which the neurenteric canal (Figure 1.123 cn)
establishes between the dorsal nerve-tube (n) and the ventral gut-tube
(d) is seen very plainly in the amphioxus in a longitudinal section of
the coelomula, as soon as the primitive mouth is completely closed at
its hinder end. The medullary tube has still at this stage an opening
at the forward end, the neuroporus (Figure 1.83 np). This opening also
is afterwards closed. There are then two completely closed canals over
each other--the medullary tube above and the gastric tube below, the
two being separated by the chorda. The same features as in the acrania
are exhibited by the related tunicates, the ascidiae.

Again, we find the neurenteric canal in just the same form and
situation in the amphibia. A longitudinal section of a young tadpole
(Figure 1.125) shows how we may penetrate from the still open
primitive mouth (x) either into the wide primitive gut-cavity (al) or
the narrow overlying nerve-tube. A little later, when the primitive
mouth is closed, the narrow neurenteric canal (Figure 1.126 ne)
represents the arched connection between the dorsal medullary canal
(mc) and the ventral gastric canal.

(FIGURE 1.125. Longitudinal section of the chordula of a frog. (From
Balfour.) nc nerve-tube, x canalis neurentericus, al alimentary canal,
yk yelk-cells, m mesoderm.

FIGURE 1.126. Longitudinal section of a frog-embryo. (From Goette.) m
mouth, l liver, an anus, ne canalis neurentericus, mc medullary-tube,
pn pineal body (epiphysis), ch chorda.

FIGURES 1.127 AND 1.128. Dorsal shield of the chick. (From Balfour.)
The medullary furrow (me), which is not yet visible in Figure 1.130,
encloses with its hinder end the fore end of the primitive groove (pr)
in Figure 1.131.)

In the amniotes this original curved form of the neurenteric canal
cannot be found at first, because here the primitive mouth travels
completely over to the dorsal surface of the gastrula, and is
converted into the longitudinal furrow we call the primitive groove.
Hence the primitive groove (Figure 1.128 pr), examined from above,
appears to be the straight continuation of the fore-lying and younger
medullary furrow (me). The divergent hind legs of the latter embrace
the anterior end of the former. Afterwards we have the complete
closing of the primitive mouth, the dorsal swellings joining to form
the medullary tube and growing over it. The neurenteric canal then
leads directly, in the shape of a narrow arch-shaped tube (Figure
1.129 ne), from the medullary tube (sp) to the gastric tube (pag).
Directly in front of it is the latter end of the chorda (cli).

While these important processes are taking place in the axial part of
the dorsal shield, its external form also is changing. The oval form
(Figure 1.117) becomes like the sole of a shoe or sandal, lyre-shaped
or finger-biscuit shaped (Figure 1.130). The middle third does not
grow in width as quickly as the posterior, and still less than the
anterior third; thus the shape of the permanent body becomes somewhat
narrow at the waist. At the same time, the oval form of the
germinative area returns to a circular shape, and the inner pellucid
area separates more clearly from the opaque outer area (Figure 1.131
a). The completion of the circle in the area marks the limit of the
formation of blood-vessels in the mesoderm.

(FIGURE 1.129. Longitudinal section of the hinder end of a chick.
(From Balfour.) sp medullary tube, connected with the terminal gut
(pag) by the neurenteric canal (ne), ch chorda, pr neurenteric (or
Hensen's) ganglion, al allantois, ep ectoderm, hy entoderm, so
parietal layer, sp visceral layer, an anus-pit, am amnion.)

The characteristic sandal-shape of the dorsal shield, which is
determined by the narrowness of the middle part, and which is compared
to a violin, lyre, or shoe-sole, persists for a long time in all the
amniotes. All mammals, birds, and reptiles have substantially the same
construction at this stage, and even for a longer or shorter period
after the division of the primitive segments into the coelom-folds has
begun (Figure 1.132). The human embryonic shield assumes the
sandal-form in the second week of development; towards the end of the
week our sole-shaped embryo has a length of about one-twelfth of an
inch (Figure 1.133).

The complete bilateral symmetry of the vertebrate body is very early
indicated in the oval form of the embryonic shield (Figure 1.117) by
the median primitive streak; in the sandal-form it is even more
pronounced (Figures 1.131 to 1.135). In the lateral parts of the
embryonic shield a darker central and a lighter peripheral zone become
more obvious; the former is called the stem-zone (Figure 1.134 stz),
and the latter the parietal zone (pz); from the first we get the
dorsal and from the second the ventral half of the body-wall. The
stem-zone of the amniote embryo would be called more appropriately the
dorsal zone or dorsal shield; from it develops the whole of the dorsal
half of the later body (or permanent body)--that is to say, the dorsal
body (episoma). Again, it would be better to call the "parietal zone"
the ventral zone or ventral shield; from it develop the ventral
"lateral plates," which afterwards separate from the embryonic vesicle
and form the ventral body (hyposoma)--that is to say, the ventral half
of the permanent body, together with the body-cavity and the gastric
canal that it encloses.

(FIGURE 1.130. Germinal area or germinal disk of the rabbit, with
sole-shaped embryonic shield, magnified about ten times. The clear
circular field (d) is the opaque area. The pellucid area (c) is
lyre-shaped, like the embryonic shield itself (b). In its axis is seen
the dorsal furrow or medullary furrow (a). (From Bischoff.))

The sole-shaped germinal shields of all the amniotes are still, at the
stage of construction which Figure 1.134 illustrates in the rabbit and
Figure 1.135 in the opossum, so like each other that we can either not
distinguish them at all or only by means of quite subordinate
peculiarities in the size of the various parts. Moreover, the human
sandal-shaped embryo cannot at this stage be distinguished from those
of other mammals, and it particularly resembles that of the rabbit. On
the other hand, the outer form of these flat sandal-shaped embryos is
very different from the corresponding form of the lower animals,
especially the acrania (amphioxus). Nevertheless, the body is just the
same in the essential features of its structure as that we find in the
chordula of the latter (Figures 1.83 to 1.86), and in the embryonic
forms which immediately develop from it. The striking external
difference is here again due to the fact that in the palingenetic
embryos of the amphioxus (Figures 1.83 and 1.84) and the amphibia
(Figures 1.85 and 1.86) the gut-wall and body-wall form closed tubes
from the first, whereas in the cenogenetic embryos of the amniotes
they are forced to expand leaf-wise on the surface owing to the great
extension of the food-yelk.

(FIGURE 1.131. Embryo of the opossum, sixty hours old, one-sixth of an
inch in diameter. (From Selenka) b the globular embryonic vesicle, a
the round germinative area, b limit of the ventral plates, r dorsal
shield, v its fore part, u the first primitive segment, ch chorda, chr
its fore-end, pr primitive groove (or mouth).

FIGURE 1.132. Sandal-shaped embryonic shield of a rabbit of eight
days, with the fore part of the germinative area (ao opaque, ap
pellucid area). (From Kolliker.) rf dorsal furrow, in the middle of
the medullary plate, h, pr primitive groove (mouth), stz dorsal (stem)
zone, pz ventral (parietal) zone. In the narrow middle part the first
three primitive segments may be seen.)

It is all the more notable that the early separation of dorsal and
ventral halves takes place in the same rigidly hereditary fashion in
all the vertebrates. In both the acrania and the craniota the dorsal
body is about this period separated from the ventral body. In the
middle part of the body this division has already taken place by the
construction of the chorda between the dorsal nerve-tube and the
ventral canal. But in the outer or lateral part of the body it is only
brought about by the division of the coelom-pouches into two
sections--a dorsal episomite (dorsal segment or provertebra) and a
ventral hyposomite (or ventral segment) by a frontal constriction. In
the amphioxus each of the former makes a muscular pouch, and each of
the latter a sex-pouch or gonad.

These important processes of differentiation in the mesoderm, which we
will consider more closely in the next chapter, proceed step by step
with interesting changes in the ectoderm, while the entoderm changes
little at first. We can study these processes best in transverse
sections, made vertically to the surface through the sole-shaped
embryonic shield. Such a transverse section of a chick embryo, at the
end of the first day of incubation, shows the gut-gland layer as a
very simple epithelium, which is spread like a leaf over the outer
surface of the food-yelk (Figure 1.92). The chorda (ch) has separated
from the dorsal middle line of the entoderm; to the right and left of
it are the two halves of the mesoderm, or the two coelom-folds. A
narrow cleft in the latter indicates the body-cavity (uwh); this
separates the two plates of the coelom-pouches, the lower (visceral)
and upper (parietal). The broad dorsal furrow (Rf) formed by the
medullary plate (m) is still wide open, but is divided from the
lateral horn-plate (h) by the parallel medullary swellings, which
eventually close.

(FIGURE 1.133. Human embryo at the sandal-stage, one-twelfth of an
inch long, from the end of the second week, magnified twenty-five
times. (From Count Spee.)

FIGURE 1.134. Sandal-shaped embryonic shield of a rabbit of nine days.
(From Kolliker.) (Back view from above.) stz stem-zone or dorsal
shield (with eight pairs of primitive segments), pz parietal or
ventral zone, ap pellucid area, af amnion-fold, h heart, ph
pericardial cavity, vo omphalo-mesenteric vein, ab eye-vesicles, vh
fore brain, mh middle brain, hh hind brain, uw primitive segments (or
vertebrae).)

During these processes important changes are taking place in the outer
germinal layer (the "skin-sense layer"). The continued rise and growth
of the dorsal swellings causes their higher parts to bend together at
their free borders, approach nearer and nearer (Figure 1.136 w), and
finally unite. Thus in the end we get from the open dorsal furrow, the
upper cleft of which becomes narrower and narrower, a closed
cylindrical tube (Figure 1.137 mr). This tube is of the utmost
importance; it is the beginning of the central nervous system, the
brain and spinal marrow, the medullary tube. This embryonic fact was
formerly looked upon as very mysterious. We shall see presently that
in the light of the theory of descent it is a thoroughly natural
process. The phylogenetic explanation of it is that the central
nervous system is the organ by means of which all intercourse with the
outer world, all psychic action and sense-perception, are
accomplished; hence it was bound to develop originally from the outer
and upper surface of the body, or from the outer skin. The medullary
tube afterwards separates completely from the outer germinal layer,
and is surrounded by the middle parts of the provertebrae and forced
inwards (Figure 1.146). The remaining portion of the skin-sense layer
(Figure 1.93 h) is now called the horn-plate or horn-layer, because
from it is developed the whole of the outer skin or epidermis, with
all its horny appendages (nails, hair, etc.).

(FIGURE 1.135. Sandal-shaped embryonic shield of an opossum
(Didelphys), three days old. (From Selenka.) (Back view from above.)
stz stem-zone or dorsal shield (with eight pairs of primitive
segments), pz parietal or ventral zone, ap pellucid area, ao opaque
area, hh halves of the heart, v fore-end, h hind-end. In the median
line we see the chorda (ch) through the transparent medullary tube
(m). u primitive segment, pr primitive streak (or primitive mouth).)

A totally different organ, the prorenal (primitive kidney) duct (ung),
is found to be developed at an early stage from the ectoderm. This is
originally a quite simple, tube-shaped, lengthy duct, or straight
canal, which runs from front to rear at each side of the provertebrae
(on the outer side, Figure 1.93 ung). It originates, it seems, out of
the horn-plate at the side of the medullary tube, in the gap that we
find between the provertebral and the lateral plates. The prorenal
duct is visible in this gap even at the time of the severance of the
medullary tube from the horn-plate. Other observers think that the
first trace of it does not come from the skin-sense layer, but the
skin-fibre layer.

The inner germinal layer, or the gut-fibre layer (Figure 1.93 dd),
remains unchanged during these processes. A little later, however, it
shows a quite flat, groove-like depression in the middle line of the
embryonic shield, directly under the chorda. This depression is called
the gastric groove or furrow. This at once indicates the future lot of
this germinal layer. As this ventral groove gradually deepens, and its
lower edges bend towards each other, it is formed into a closed tube,
the alimentary canal, in the same way as the medullary groove grows
into the medullary tube. The gut-fibre layer (Figure 1.137 f), which
lies on the gut-gland layer (d), naturally follows it in its folding.
Moreover, the incipient gut-wall consists from the first of two
layers, internally the gut-gland layer and externally the gut-fibre
layer.

The formation of the alimentary canal resembles that of the medullary
tube to this extent--in both cases a straight groove or furrow arises
first of all in the middle line of a flat layer. The edges of this
furrow then bend towards each other, and join to form a tube (Figure
1.137). But the two processes are really very different. The medullary
tube closes in its whole length, and forms a cylindrical tube, whereas
the alimentary canal remains open in the middle, and its cavity
continues for a long time in connection with the cavity of the
embryonic vesicle. The open connection between the two cavities is
only closed at a very late stage, by the construction of the navel.
The closing of the medullary tube is effected from both sides, the
edges of the groove joining together from right and left. But the
closing of the alimentary canal is not only effected from right and
left, but also from front and rear, the edges of the ventral groove
growing together from every side towards the navel. Throughout the
three higher classes of vertebrates the whole of this process of the
construction of the gut is closely connected with the formation of the
navel, or with the separation of the embryo from the yelk-sac or
umbilical vesicle.

In order to get a clear idea of this, we must understand carefully the
relation of the embryonic shield to the germinative area and the
embryonic vesicle. This is done best by a comparison of the five
stages which are shown in longitudinal section in Figures 1.138 to
1.142. The embryonic shield (c), which at first projects very slightly
over the surface of the germinative area, soon begins to rise higher
above it, and to separate from the embryonic vesicle. At this point
the embryonic shield, looked at from the dorsal surface, shows still
the original simple sandal-shape (Figures 1.133 to 1.135). We do not
yet see any trace of articulation into head, neck, trunk, etc., or
limbs. But the embryonic shield has increased greatly in thickness,
especially in the anterior part. It now has the appearance of a thick,
oval swelling, strongly curved over the surface of the germinative
area. It begins to sever completely from the embryonic vesicle, with
which it is connected at the ventral surface. As this severance
proceeds, the back bends more and more; in proportion as the embryo
grows the embryonic vesicle decreases, and at last it merely hangs as
a small vesicle from the belly of the embryo (Figure 1.142 ds). In
consequence of the growth-movements which cause this severance, a
groove-shaped depression is formed at the surface of the vesicle, the
limiting furrow, which surrounds the vesicle in the shape of a pit,
and a circular mound or dam (Figure 1.139 ks) is formed at the outside
of this pit by the elevation of the contiguous parts of the germinal
vesicle.

(FIGURE 1.136. Transverse section of the embryonic disk of a chick at
the end of the first day of incubation, magnified about twenty times.
The edges of the medullary plate (m), the medullary swellings (w),
which separate the medullary from the horn-plate (h), are bending
towards each other. At each side of the chorda (ch) the primitive
segment plates (u) have separated from the lateral plates (sp). A
gut-gland layer. (From Remak.))

In order to understand clearly this important process, we may compare
the embryo to a fortress with its surrounding rampart and trench. The
ditch consists of the outer part of the germinative area, and comes to
an end at the point where the area passes into the vesicle. The
important fold of the middle germinal layer that brings about the
formation of the body-cavity spreads beyond the borders of the embryo
over the whole germinative area. At first this middle layer reaches as
far as the germinative area; the whole of the rest of the embryonic
vesicle consists in the beginning only of the two original limiting
layers, the outer and inner germinal layers. Hence, as far as the
germinative area extends the germinal layer splits into the two plates
we have already recognised in it, the outer skin-fibre layer and the
inner gut-fibre layer. These two plates diverge considerably, a clear
fluid gathering between them (Figure 1.140 am). The inner plate, the
gut-fibre layer, remains on the inner layer of the embryonic vesicle
(on the gut-gland layer). The outer plate, the skin-fibre layer, lies
close on the outer layer of the germinative area, or the skin-sense
layer, and separates together with this from the embryonic vesicle.
From these two united outer plates is formed a continuous membrane.
This is the circular mound that rises higher and higher round the
whole embryo, and at last joins above it (Figures 1.139 to 1.142 am).
To return to our illustration of the fortress, we must imagine the
circular rampart to be extraordinarily high and towering far above the
fortress. Its edges bend over like the combs of an overhanging wall of
rock that would enclose the fortress; they form a deep hollow, and at
last join together above. In the end the fortress lies entirely within
the hollow that has been formed by the growth of the edges of this
large rampart.

(FIGURE 1.137. Three diagrammatic transverse sections of the embryonic
disk of the higher vertebrate, to show the origin of the tubular
organs from the bending germinal layers. In Figure A the medullary
tube (n) and the alimentary canal (a) are still open grooves. In
Figure B the medullary tube (n) and the dorsal wall are closed, but
the alimentary canal (a) and the ventral wall are open; the prorenal
ducts (u) are cut off from the horn-plate (h) and internally connected
with segmental prorenal canals. In Figure C both the medullary tube
and the dorsal wall above and the alimentary canal and ventral wall
below are closed. All the open grooves have become closed tubes; the
primitive kidneys are directed inwards. The letters have the same
meaning in all three figures: h skin-sense layer, n medullary tube, u
prorenal ducts, x axial rod, s primitive-vertebra, r dorsal wall, b
ventral wall, c body-cavity or coeloma, f gut-fibre layer, t primitive
artery (aorta), v primitive vein (subintestinal vein), d gut-fibre
layer, a alimentary canal.)

As the two outer layers of the germinative area thus rise in a fold
about the embryo, and join above it, they come at last to form a
spacious sac-like membrane about it. This envelope takes the name of
the germinative membrane, or water-membrane, or amnion (Figure 1.142
am). The embryo floats in a watery fluid, which fills the space
between the embryo and the amnion, and is called the amniotic fluid
(Figures 1.141 and 1.142 ah). We will deal with this remarkable
formation and with the allantois later on (Chapter 1.15). In front of
the allantois the yelk-sac or umbilical vesicle (ds), the remainder of
the original embryonic vesicle, starts from the open belly of the
embryo (Figure 1.138 kh). In more advanced embryos, in which the
gastric wall and the ventral wall are nearly closed, it hangs out of
the navel-opening in the shape of a small vesicle with a stalk
(Figures 1.141 and 1.142 ds). The more the embryo grows, the smaller
becomes the vitelline (yelk) sac. At first the embryo looks like a
small appendage of the large embryonic vesicle. Afterwards it is the
yelk-sac, or the remainder of the embryonic vesicle, that seems a
small pouch-like appendage of the embryo (Figure 1.142 ds). It ceases
to have any significance in the end. The very wide opening, through
which the gastric cavity at first communicates with the umbilical
vesicle, becomes narrower and narrower, and at last disappears
altogether. The navel, the small pit-like depression that we find in
the developed man in the middle of the abdominal wall, is the spot at
which the remainder of the embryonic vesicle (the umbilical vesicle)
originally entered into the ventral cavity, and joined on to the
growing gut.

(FIGURES 1.138 TO 1.142. Five diagrammatic longitudinal sections of
the maturing mammal embryo and its envelopes. In Figures 1.138 to
1.141 the longitudinal section passes through the sagittal or middle
plane of the body, dividing the right and left halves; in Figure 1.142
the embryo is seen from the left side. In Figure 1.138 the tufted it
prochorion (dd apostrophe) encloses the germinal vesicle, the wall of
which consists of the two primary layers. Between the outer (a) and
inner (i) layer the middle layer (m) has been developed in the region
of the germinative area. In Figure 1.139 the embryo (e) begins to
separate from the embryonic vesicle (ds), while the wall of the
amnion-fold rises about it (in front as head-sheath, ks, behind as
tail-sheath, ss). In Figure 1.140 the edges of the amniotic fold (am)
rise together over the back of the embryo, and form the amniotic
cavity (ah); as the embryo separates more completely from the
embryonic vesicle (ds) the alimentary canal (dd) is formed, from the
hinder end of which the allantois grows (al). In Figure 1.141 the
allantois is larger; the yelk-sac (ds) smaller. In Figure 1.142 the
embryo shows the gill-clefts and the outline of the two legs; the
chorion has formed branching villi (tufts.) In all four figures e =
embryo, a outer germinal layer, m middle germinal layer, i inner
germinal layer, am amnion (ks head-sheath, ss tail-sheath), ah
amniotic cavity, as amniotic sheath of the umbilical cord, kh
embryonic vesicle, ds yelk-sac (umbilical vesicle), dg vitelline duct,
df gut-fibre layer, dd gut-gland layer, al allantois, vl = hh place of
heart, d vitelline membrane (ovolemma or prochorion), d apostrophe
tufts or villi of same, sh serous membrane (serolemma), sz tufts of
same, ch chorion, chz tufts or villi, st terminal vein, r pericoelom
or serocoelom (the space, filled with fluid, between the amnion and
chorion). (From Kolliker.))

The origin of the navel coincides with the complete closing of the
external ventral wall. In the amniotes the ventral wall originates in
the same way as the dorsal wall. Both are formed substantially from
the skin-fibre layer, and externally covered with the horn-plate, the
border section of the skin-sense layer. Both come into existence by
the conversion of the four flat germinal layers of the embryonic
shield into a double tube by folding from opposite directions; above,
at the back, we have the vertebral canal which encloses the medullary
tube, and below, at the belly, the wall of the body-cavity which
contains the alimentary canal (Figure 1.137).

We will consider the formation of the dorsal wall first, and that of
the ventral wall afterwards (Figures 1.143 to 1.147). In the middle of
the dorsal surface of the embryo there is originally, as we already
know, the medullary (mr) tube directly underneath the horn-plate (h),
from the middle part of which it has been developed. Later, however,
the provertebral plates (uw) grow over from the right and left between
these originally connected parts (Figures 1.145 and 1.146). The upper
and inner edges of the two provertebral plates push between the
horn-plate and medullary tube, force them away from each other, and
finally join between them in a seam that corresponds to the middle
line of the back. The coalescence of these two dorsal plates and the
closing in the middle of the dorsal wall take place in the same way as
the medullary tube, which is henceforth enclosed by the vertebral
tube. Thus is formed the dorsal wall, and the medullary tube takes up
a position inside the body. In the same way the provertebral mass
grows afterwards round the chorda, and forms the vertebral column.
Below this the inner and outer edge of the provertebral plate splits
on each side into two horizontal plates, of which the upper pushes
between the chorda and medullary tube, and the lower between the
chorda and gastric tube. As the plates meet from both sides above and
below the chorda, they completely enclose it, and so form the tubular,
outer chord-sheath, the sheath from which the vertebral column is
formed (perichorda, Figure 1.137 C, s; Figures 1.145 uwh, 1.146).

(FIGURES 1.143 TO 1.146. Transverse sections of embryos (of chicks).
Figure 1.143 of the second, Figure 1.144 of the third, Figure 1.145 of
the fourth, and Figure 1.146 of the fifth day of incubation. Figures
1.143 to 1.145 from Kolliker, magnified about 100 times; Figure 1.146
from Remak, magnified about twenty times. h horn-plate, mr medullary
tube, ung prorenal duct, un prorenal vesicles, hp skin-fibre layer, m
= mu = mp muscle-plate, uw provertebral plate (wh cutaneous rudiment
of the body of the vertebra, wb of the arch of the vertebra, wq the
rib or transverse continuation), uwh provertebral cavity, ch axial rod
or chorda, sh chorda-sheath, bh ventral wall, g hind and v fore root
of the spinal nerves, a = af = am amniotic fold, p body-cavity or
coeloma, df gut-fibre layer, ao primitive aortas, sa secondary aorta,
vc cardinal veins, d = dd gut-gland layer, dr gastric groove. In
Figure 1.143 the larger part of the right half, in Figure 1.144 the
larger part of the left half, of the section is omitted. Of the
yelk-sac or remainder of the embryonic vesicle only a small piece of
the wall is indicated below.)

We find in the construction of the ventral wall precisely the same
processes as in the formation of the dorsal wall (Figure 1.137 B,
Figure 1.144 hp, Figure 1.146 bh). It is formed on the flat embryonic
shield of the amniotes from the upper plates of the parietal zone. The
right and left parietal plates bend downwards towards each other, and
grow round the gut in the same way as the gut itself closes. The outer
part of the lateral plates forms the ventral wall or the lower wall of
the body, the two lateral plates bending considerably on the inner
side of the amniotic fold, and growing towards each other from right
and left. While the alimentary canal is closing, the body-wall also
closes on all sides. Hence the ventral wall, which encloses the whole
ventral cavity below, consists of two parts, two lateral plates that
bend towards each other. These approach each other all along, and at
last meet at the navel. We ought, therefore, really to distinguish two
navels, an inner and an outer one. The internal or intestinal navel is
the definitive point of the closing of the gut wall, which puts an end
to the open communication between the ventral cavity and the cavity of
the yelk-sac (Figure 1.105). The external navel in the skin is the
definitive point of the closing of the ventral wall; this is visible
in the developed body as a small depression.

(FIGURE 1.147. Median longitudinal section of the embryo of a chick
(fifth day of incubation), seen from the right side (head to the
right, tail to the left). Dorsal body dark, with convex outline. d
gut, o mouth, a anus, l lungs, h liver, g mesentery, v auricle of the
heart, k ventricle of the heart, b arch of the arteries, t aorta, c
yelk-sac, m vitelline (yelk) duct, u allantois, r pedicle (stalk) of
the allantois, n amnion, w amniotic cavity, s serous membrane. (From
Baer.))

With the formation of the internal navel and the closing of the
alimentary canal is connected the formation of two cavities, which we
call the capital and the pelvic sections of the visceral cavity. As
the embryonic shield lies flat on the wall of the embryonic vesicle at
first, and only gradually separates from it, its fore and hind ends
are independent in the beginning; on the other hand, the middle part
of the ventral surface is connected with the yelk-sac by means of the
vitelline or umbilical duct (Figure 1.147 m). This leads to a notable
curving of the dorsal surface; the head-end bends downwards towards
the breast and the tail-end towards the belly. We see this very
clearly in the excellent old diagrammatic illustration given by Baer
(Figure 1.147), a median longitudinal section of the embryo of the
chick, in which the dorsal body or episoma is deeply shaded. The
embryo seems to be trying to roll up, like a hedgehog protecting
itself from its pursuers. This pronounced curve of the back is due to
the more rapid growth of the convex dorsal surface, and is directly
connected with the severance of the embryo from the yelk-sac. The
further bending of the embryo leads to the formation of the
"head-cavity" of the gut (Figure 1.148 above D) and a similar one at
the tail, known as its "pelvic cavity."

As a result of these processes the embryo attains a shape that may be
compared to a wooden shoe, or, better still, to an overturned canoe.
Imagine a canoe or boat with both ends rounded and a small covering
before and behind; if this canoe is turned upside down, so that the
curved keel is uppermost, we have a fair picture of the canoe-shaped
embryo (Figure 1.147). The upturned convex keel corresponds to the
middle line of the back; the small chamber underneath the fore-deck
represents the capital cavity, and the small chamber under the
rear-deck the pelvic chamber of the gut (cf. Figure 1.140).

The embryo now, as it were, presses into the outer surface of the
embryonic vesicle with its free ends, while it moves away from it with
its middle part. As a result of this change the yelk-sac becomes
henceforth only a pouch-like outer appendage at the middle of the
ventral wall. The ventral appendage, growing smaller and smaller, is
afterwards called the umbilical (navel) vesicle. The cavity of the
yelk-sac or umbilical vesicle communicates with the corresponding
visceral cavity by a wide opening, which gradually contracts into a
narrow and long canal, the vitelline (yelk) duct (ductus vitellinus,
Figure 1.147 m). Hence, if we were to imagine ourselves in the cavity
of the yelk-sac, we could get from it through the yelk-duct into the
middle and still wide open part of the alimentary canal. If we were to
go forward from there into the head-part of the embryo, we should
reach the capital cavity of the gut, the fore-end of which is closed
up.

The reader will ask: "Where are the mouth and the anus?" These are not
at first present in the embryo. The whole of the primitive gut-cavity
is completely closed, and is merely connected in the middle by the
vitelline duct with the equally closed cavity of the embryonic vesicle
(Figure 1.140). The two later apertures of the alimentary canal--the
anus and the mouth--are secondary constructions, formed from the outer
skin. In the horn-plate, at the spot where the mouth is found
subsequently, a pit-like depression is formed, and this grows deeper
and deeper, pushing towards the blind fore-end of the capital cavity;
this is the mouth-pit. In the same way, at the spot in the outer skin
where the anus is afterwards situated a pit-shaped depression appears,
grows deeper and deeper, and approaches the blind hind-end of the
pelvic cavity; this is the anus-pit. In the end these pits touch with
their deepest and innermost points the two blind ends of the primitive
alimentary canal, so that they are now only separated from them by
thin membranous partitions. This membrane finally disappears, and
henceforth the alimentary canal opens in front at the mouth and in the
rear by the anus (Figures 1.141 and 1.147). Hence at first, if we
penetrate into these pits from without, we find a partition cutting
them off from the cavity of the alimentary canal, which gradually
disappears. The formation of mouth and anus is secondary in all the
vertebrates.

(FIGURE 1.148. Longitudinal section of the fore half of a chick-embryo
at the end of the first day of incubation (seen from the left side). k
head-plates, ch chorda. Above it is the blind fore-end of the ventral
tube (m); below it the capital cavity of the gut. d gut-gland layer,
df gut-fibre layer, h horn plate, hh cavity of the heart, hk
heart-capsule, ks head-sheath, kk head-capsule. (From Remak.))

During the important processes which lead to the formation of the
navel, and of the intestinal wall and ventral wall, we find a number
of other interesting changes taking place in the embryonic shield of
the amniotes. These relate chiefly to the prorenal ducts and the first
blood-vessels. The prorenal (primitive kidney) ducts, which at first
lie quite flat under the horn-plate or epiderm (Figure 1.93 ung), soon
back towards each other in consequence of special growth movements
(Figures 1.143 to 1.145 ung). They depart more and more from their
point of origin, and approach the gut-gland layer. In the end they lie
deep in the interior, on either side of the mesentery, underneath the
chorda, (Figure 1.145 ung). At the same time, the two primitive aortas
change their position (cf. Figures 1.138 to 1.145 ao); they travel
inwards underneath the chorda, and there coalesce at last to form a
single secondary aorta, which is found under the rudimentary vertebral
column (Figure 1.145 ao). The cardinal veins, the first venous
blood-vessels, also back towards each other, and eventually unite
immediately above the rudimentary kidneys (Figures 1.145 vc, 152 cav).
In the same spot, at the inner side of the fore-kidneys, we soon see
the first trace of the sexual organs. The most important part of this
apparatus (apart from all its appendages) is the ovary in the female
and the testicle in the male. Both develop from a small part of the
cell-lining of the body-cavity, at the spot where the skin-fibre layer
and gut-fibre layer touch. The connection of this embryonic gland with
the prorenal ducts, which lie close to it and assume most important
relations to it, is only secondary.

(FIGURE 1.149. Longitudinal section of a human embryo of the fourth
week, one-fifth of an inch long, magnified fifteen times. Showing:
bend of skull, yelk-sac, umbilical cord, terminal gut, rudimentary
kidneys, mesoderm, head-gut (with gill-clefts), primitive lungs,
liver, stomach, pancreas, mesentery, primitive kidneys, allantoic
duct, rectum. (From Kollmann.)

FIGURE 1.150. Transverse section of a human embryo of fourteen days.
mr medullary tube, ch chorda. vu umbilical vein, mt myotome, mp middle
plate, ug prorenal duct, lh body-cavity, e ectoderm, bh ventral skin,
hf skin-fibre layer, df gut-fibre layer. (From Kollmann.)

FIGURE 1.151. Transverse section of a shark-embryo (or young
selachius). mr medullary tube, ch chorda, a aorta, d gut, vp principal
(or subintestinal) vein, mt myotome, mm muscular mass of the
provertebra, mp middle plate, ug prorenal duct, lh body-cavity, e
ectoderm of the rudimentary extremities, mz mesenchymic cells, z point
where the myotome and nephrotome separate. (From H.E. Ziegler.)

FIGURE 1.152. Transverse section of a duck-embryo with twenty-four
primitive segments. (From Balfour.) From a dorsal lateral joint of the
medullary tube (spc) the spinal ganglia (spg) grow out between it and
the horn-plate. ch chorda, ao double aorta, hy gut-gland layer, sp
gut-fibre layer, with blood-vessels in section, ms muscle plate, in
the dorsal wall of the myocoel (episomite). Below the cardinal vein
(cav) is the prorenal duct (wd) and a segmental prorenal canal (st).
The skin-fibre layer of the body-wall (so) is continued in the
amniotic fold (am). Between the four secondary germinal layers and the
structures formed from them there is formed embryonic connective
matter with stellate cells and vascular structures (Hertwig's
"mesenchym").)


CHAPTER 1.14. THE ARTICULATION OF THE BODY.*

(* The term articulation is used in this chapter to denote both
"segmentation" and "articulation" in the ordinary sense.--Translator.)

The vertebrate stem, to which our race belongs as one of the latest
and most advanced outcomes of the natural development of life, is
rightly placed at the head of the animal kingdom. This privilege must
be accorded to it, not only because man does in point of fact soar far
above all other animals, and has been lifted to the position of "lord
of creation"; but also because the vertebrate organism far surpasses
all the other animal-stems in size, in complexity of structure, and in
the advanced character of its functions. From the point of view of
both anatomy and physiology, the vertebrate stem outstrips all the
other, or invertebrate, animals.

There is only one among the twelve stems of the animal kingdom that
can in many respects be compared with the vertebrates, and reaches an
equal, if not a greater, importance in many points. This is the stem
of the articulates, composed of three classes: 1, the annelids
(earth-worms, leeches, and cognate forms); 2, the crustacea (crabs,
etc.); 3, the tracheata (spiders, insects, etc.). The stem of the
articulates is superior not only to the vertebrates, but to all other
animal-stems, in variety of forms, number of species, elaborateness of
individuals, and general importance in the economy of nature.

When we have thus declared the vertebrates and the articulates to be
the most important and most advanced of the twelve stems of the animal
kingdom, the question arises whether this special position is accorded
to them on the ground of a peculiarity of organisation that is common
to the two. The answer is that this is really the case; it is their
segmental or transverse articulation, which we may briefly call
metamerism. In all the vertebrates and articulates the developed
individual consists of a series of successive members (segments or
metamera = "parts"); in the embryo these are called primitive segments
or somites. In each of these segments we have a certain group of
organs reproduced in the same arrangement, so that we may regard each
segment as an individual unity, or a special "individual" subordinated
to the entire personality.

The similarity of their segmentation, and the consequent physiological
advance in the two stems of the vertebrates and articulates, has led
to the assumption of a direct affinity between them, and an attempt to
derive the former directly from the latter. The annelids were supposed
to be the direct ancestors, not only of the crustacea and tracheata,
but also of the vertebrates. We shall see later (Chapter 2.20) that
this annelid theory of the vertebrates is entirely wrong, and ignores
the most important differences in the organisation of the two stems.
The internal articulation of the vertebrates is just as profoundly
different from the external metamerism of the articulates as are their
skeletal structure, nervous system, vascular system, and so on. The
articulation has been developed in a totally different way in the two
stems. The unarticulated chordula (Figures 1.83 to 1.86), which we
have recognised as one of the chief palingenetic embryonic forms of
the vertebrate group, and from which we have inferred the existence of
a corresponding ancestral form for all the vertebrates and tunicates,
is quite unthinkable as the stem-form of the articulates.

All articulated animals came originally from unarticulated ones. This
phylogenetic principle is as firmly established as the ontogenetic
fact that every articulated animal-form develops from an unarticulated
embryo. But the organisation of the embryo is totally different in the
two stems. The chordula-embryo of all the vertebrates is characterised
by the dorsal medullary tube, the neurenteric canal, which passes at
the primitive mouth into the alimentary canal, and the axial chorda
between the two. None of the articulates, either annelids or
arthropods (crustacea and tracheata), show any trace of this type of
organisation. Moreover, the development of the chief systems of organs
proceeds in the opposite way in the two stems. Hence the segmentation
must have arisen independently in each. This is not at all surprising;
we find analogous cases in the stalk-articulation of the higher plants
and in several groups of other animal stems.

The characteristic internal articulation of the vertebrates and its
importance in the organisation of the stem are best seen in the study
of the skeleton. Its chief and central part, the cartilaginous or bony
vertebral column, affords an obvious instance of vertebrate
metamerism; it consists of a series of cartilaginous or bony pieces,
which have long been known as vertebrae (or spondyli). Each vertebra
is directly connected with a special section of the muscular system,
the nervous system, the vascular system, etc. Thus most of the "animal
organs" take part in this vertebration. But we saw, when we were
considering our own vertebrate character (in Chapter 1.11), that the
same internal articulation is also found in the lowest primitive
vertebrates, the acrania, although here the whole skeleton consists
merely of the simple chorda, and is not at all articulated. Hence the
articulation does not proceed primarily from the skeleton, but from
the muscular system, and is clearly determined by the more advanced
swimming-movements of the primitive chordonia-ancestors.

(FIGURES 1.153 TO 1.155. Sole-shaped embryonic disk of the chick, in
three successive stages of development, looked at from the dorsal
surface, magnified about twenty times, somewhat diagrammatic. Figure
1.153 with six pairs of somites. Brain a simple vesicle (hb).
Medullary furrow still wide open from x; greatly widened at z. mp
medullary plates, sp lateral plates, y limit of gullet-cavity (sh) and
fore-gut (vd). Figure 1.154 with ten pairs of somites. Brain divided
into three vesicles: v fore-brain, m middle-brain, h hind-brain, c
heart, dv vitelline-veins. Medullary furrow still wide open behind
(z). mp medullary plates. Figure 1.155 with sixteen pairs of somites.
Brain divided into five vesicles: v fore-brain, z intermediate-brain,
m middle-brain, h hind-brain, n after-brain, a optic vesicles, g
auditory vesicles, c heart, dv vitelline veins, mp medullary plate, uw
primitive vertebra.)

It is, therefore, wrong to describe the first rudimentary segments in
the vertebrate embryo as primitive vertebrae or provertebrae; the fact
that they have been so called for some time has led to much error and
misunderstanding. Hence we shall give the name of "somites" or
primitive segments to these so-called "primitive vertebrae." If the
latter name is retained at all, it should only be used of the
sclerotom--i.e., the small part of the somites from which the later
vertebra does actually develop.

Articulation begins in all vertebrates at a very early embryonic
stage, and this indicates the considerable phylogenetic age of the
process. When the chordula (Figures 1.83 to 1.86) has completed its
characteristic composition, often even a little earlier, we find in
the amniotes, in the middle of the sole-shaped embryonic shield,
several pairs of dark square spots, symmetrically distributed on both
sides of the chorda (Figures 1.131 to 1.135). Transverse sections
(Figure 1.93 uw) show that they belong to the stem-zone (episoma) of
the mesoderm, and are separated from the parietal zone (hyposoma) by
the lateral folds; in section they are still quadrangular, almost
square, so that they look something like dice. These pairs of "cubes"
of the mesoderm are the first traces of the primitive segments or
somites, the so-called "protovertebrae." (Figures 1.153 to 1.155 uw).

(FIGURE 1.156. Embryo of the amphioxus, sixteen hours old, seen from
the back. (From Hatschek.) d primitive gut, u primitive mouth, p polar
cells of the mesoderm, c coelom-pouches, m their first segment, n
medullary tube, i entoderm, e ectoderm, s first segment-fold.

FIGURE 1.157. Embryo of the amphioxus, twenty hours old, with five
somites. (Right view; for left view see Figure 1.124.) (From
Hatschek.) V fore end, H hind end. ak, mk, ik outer, middle, and inner
germinal layers; dh alimentary canal, n neural tube, cn canalis
neurentericus, ush coelom-pouches (or primitive-segment cavities), us1
first (and foremost) primitive segment.)

Among the mammals the embryos of the marsupials have three pairs of
somites (Figure 1.131) after sixty hours, and eight pairs after
seventy-two hours (Figure 1.135). They develop more slowly in the
embryo of the rabbit; this has three somites on the eighth day (Figure
1.132), and eight somites a day later (Figure 1.134). In the incubated
hen's egg the first somites make their appearance thirty hours after
incubation begins (Figure 1.153). At the end of the second day the
number has risen to sixteen or eighteen (Figure 1.155). The
articulation of the stem-zone, to which the somites owe their origin,
thus proceeds briskly from front to rear, new transverse constrictions
of the "protovertebral plates" forming continuously and successively.
The first segment, which is almost half-way down in the embryonic
shield of the amniote, is the foremost of all; from this first somite
is formed the first cervical vertebra with its muscles and skeletal
parts. It follows from this, firstly, that the multiplication of the
primitive segments proceeds backwards from the front, with a constant
lengthening of the hinder end of the body; and, secondly, that at the
beginning of segmentation nearly the whole of the anterior half of the
sole-shaped embryonic shield of the amniote belongs to the later head,
while the whole of the rest of the body is formed from its hinder
half. We are reminded that in the amphioxus (and in our hypothetic
primitive vertebrate, Figures 1.98 to 1.102) nearly the whole of the
fore half corresponds to the head, and the hind half to the trunk.

The number of the metamera, and of the embryonic somites or primitive
segments from which they develop, varies considerably in the
vertebrates, according as the hind part of the body is short or is
lengthened by a tail. In the developed man the trunk (including the
rudimentary tail) consists of thirty-three metamera, the solid centre
of which is formed by that number of vertebrae in the vertebral column
(seven cervical, twelve dorsal, five lumbar, five sacral, and four
caudal). To these we must add at least nine head-vertebrae, which
originally (in all the craniota) constitute the skull. Thus the total
number of the primitive segments of the human body is raised to at
least forty-two; it would reach forty-five to forty-eight if
(according to recent investigations) the number of the original
segments of the skull is put at twelve to fifteen. In the tailless or
anthropoid apes the number of metamera is much the same as in man,
only differing by one or two; but it is much larger in the long-tailed
apes and most of the other mammals. In long serpents and fishes it
reaches several hundred (sometimes 400).

(FIGURES 1.158 TO 1.160. Embryo of the amphioxus, twenty four hours
old, with eight somites. (From Hatschek.) Figures 1.158 and 1.159
lateral view (from left). Figure 1.160 seen from back. In Figure 1.158
only the outlines of the eight primitive segments are indicated, in
Figure 1.159 their cavities and muscular walls. V fore end, H hind
end, d gut, du under and dd upper wall of the gut, ne canalis
neurentericus, nv ventral, nd dorsal wall of the neural tube, np
neuroporus, dv fore pouch of the gut, ch chorda, mf mesodermic fold,
pm polar cells of the mesoderm (ms), e ectoderm.)

In order to understand properly the real nature and origin of
articulation in the human body and that of the higher vertebrates, it
is necessary to compare it with that of the lower vertebrates, and
bear in mind always the genetic connection of all the members of the
stem. In this the simple development of the invaluable amphioxus once
more furnishes the key to the complex and cenogenetically modified
embryonic processes of the craniota. The articulation of the amphioxus
begins at an early stage--earlier than in the craniotes. The two
coelom-pouches have hardly grown out of the primitive gut (Figure
1.156 c) when the blind fore part of it (farthest away from the
primitive mouth, u) begins to separate by a transverse fold (s): this
is the first primitive segment. Immediately afterwards the hind part
of the coelom-pouches begins to divide into a series of pieces by new
transverse folds (Figure 1.157). The foremost of these primitive
segments (us1) is the first and oldest; in Figures 1.124 and 1.157
there are already five formed. They separate so rapidly, one behind
the other, that eight pairs are formed within twenty-four hours of the
beginning of development, and seventeen pairs twenty-four hours later.
The number increases as the embryo grows and extends backwards, and
new cells are formed constantly (at the primitive mouth) from the two
primitive mesodermic cells (Figures 1.159 to 1.160).

(FIGURES 1.161 AND 1.162. Transverse section of shark-embryos (through
the region of the kidneys). (From Wijhe and Hertwig.) In Figure 1.162
the dorsal segment-cavities (h) are already separated from the
body-cavity (lh), but they are connected a little earlier (Figure
1.161), nr neural tube, ch chorda, sch subchordal string, ao aorta, sk
skeletal-plate, mp muscle-plate, cp cutis-plate, w connection of
latter (growth-zone), vn primitive kidneys, ug prorenal duct, uk
prorenal canals, us point where they are cut off, tr prorenal funnel,
mk middle germ-layer (mk1 parietal, mk2 visceral), ik inner germ-layer
(gut-gland layer).)

This typical articulation of the two coelom-sacs begins very early in
the lancelet, before they are yet severed from the primitive gut, so
that at first each segment-cavity (us) still communicates by a narrow
opening with the gut, like an intestinal gland. But this opening soon
closes by complete severance, proceeding regularly backwards. The
closed segments then extend more, so that their upper half grows
upwards like a fold between the ectoderm (ak) and neural tube (n), and
the lower half between the ectoderm and alimentary canal (ch; Figure
1.82 d, left half of the figure). Afterwards the two halves completely
separate, a lateral longitudinal fold cutting between them (mk, right
half of Figure 1.82). The dorsal segments (sd) provide the muscles of
the trunk the whole length of the body (1.159): this cavity afterwards
disappears. On the other hand, the ventral parts give rise, from their
uppermost section, to the pronephridia or primitive-kidney canals, and
from the lower to the segmental rudiments of the sexual glands or
gonads. The partitions of the muscular dorsal pieces (myotomes)
remain, and determine the permanent articulation of the vertebrate
organism. But the partitions of the large ventral pieces (gonotomes)
become thinner, and afterwards disappear in part, so that their
cavities run together to form the metacoel, or the simple permanent
body-cavity.

The articulation proceeds in substantially the same way in the other
vertebrates, the craniota, starting from the coelom-pouches. But
whereas in the former case there is first a transverse division of the
coelom-sacs (by vertical folds) and then the dorso-ventral division,
the procedure is reversed in the craniota; in their case each of the
long coelom-pouches first divides into a dorsal (primitive segment
plates) and a ventral (lateral plates) section by a lateral
longitudinal fold. Only the former are then broken up into primitive
segments by the subsequent vertical folds; while the latter (segmented
for a time in the amphioxus) remain undivided, and, by the divergence
of their parietal and visceral plates, form a body-cavity that is
unified from the first. In this case, again, it is clear that we must
regard the features of the younger craniota as cenogenetically
modified processes that can be traced palingenetically to the older
acrania.

We have an interesting intermediate stage between the acrania and the
fishes in these and many other respects in the cyclostoma (the hag and
the lamprey, cf. Chapter 2.21).

(FIGURE 1.163. Frontal (or horizontal-longitudinal) section of a
triton-embryo with three pairs of primitive segments. ch chorda, us
primitive segments, ush their cavity, ak horn plate.)

Among the fishes the selachii, or primitive fishes, yield the most
important information on these and many other phylogenetic questions
(Figures 1.161 and 1.162). The careful studies of Ruckert, Van Wijhe,
H.E. Ziegler, and others, have given us most valuable results. The
products of the middle germinal layer are partly clear in these cases
at the period when the dorsal primitive segment cavities (or myocoels,
h) are still connected with the ventral body-cavity (lh; Figure
1.161). In Figure 1.162, a somewhat older embryo, these cavities are
separated. The outer or lateral wall of the dorsal segment yields the
cutis-plate (cp), the foundation of the connective corium. From its
inner or median wall are developed the muscle-plate (mp, the rudiment
of the trunk-muscles) and the skeletal plate, the formative matter of
the vertebral column (sk).

In the amphibia, also, especially the water-salamander (Triton), we
can observe very clearly the articulation of the coelom-pouches and
the rise of the primitive segments from their dorsal half (cf. Figure
1.91, A, B, C). A horizontal longitudinal section of the
salamander-embryo (Figure 1.163) shows very clearly the series of
pairs of these vesicular dorsal segments, which have been cut off on
each side from the ventral side-plates, and lie to the right and left
of the chorda.

(FIGURE 1.164. The third cervical vertebra (human).

FIGURE 1.165. The sixth dorsal vertebra (human).

FIGURE 1.166. The second lumbar vertebra (human).)

The metamerism of the amniotes agrees in all essential points with
that of the three lower classes of vertebrates we have considered; but
it varies considerably in detail, in consequence of cenogenetic
disturbances that are due in the first place (like the degeneration of
the coelom-pouches) to the large development of the food-yelk. As the
pressure of this seems to force the two middle layers together from
the start, and as the solid structure of the mesoderm apparently
belies the original hollow character of the sacs, the two sections of
the mesoderm, which are at that time divided by the lateral fold--the
dorsal segment-plates and ventral side-plates--have the appearance at
first of solid layers of cells (Figures 1.94 to 1.97). And when the
articulation of the somites begins in the sole-shaped embryonic
shield, and a couple of protovertebrae are developed in succession,
constantly increasing in number towards the rear, these cube-shaped
somites (formerly called protovertebrae, or primitive vertebrae) have
the appearance of solid dice, made up of mesodermic cells (Figure
1.93). Nevertheless, there is for a time a ventral cavity, or
provertebral cavity, even in these solid "protovertebrae" (Figure
1.143 uwh). This vesicular condition of the provertebra is of the
greatest phylogenetic interest; we must, according to the coelom
theory, regard it as an hereditary reproduction of the hollow dorsal
somites of the amphioxus (Figures 1.156 to 1.160) and the lower
vertebrates (Figures 1.161 to 1.163). This rudimentary "provertebral
cavity" has no physiological significance whatever in the
amniote-embryo; it soon disappears, being filled up with cells of the
muscular plate.

(FIGURE 1.167. Head of a shark embryo (Pristiurus), one-third of an
inch long, magnified twenty times. (From Parker.) Seen from the
ventral side.)

The innermost median part of the primitive segment plates, which lies
immediately on the chorda (Figure 1.145 ch) and the medullary tube
(m), forms the vertebral column in all the higher vertebrates (it is
wanting in the lowest); hence it may be called the skeleton plate. In
each of the provertebrae it is called the "sclerotome" (in opposition
to the outlying muscular plate, the "myotome"). From the phylogenetic
point of view the myotomes are much older than the sclerotomes. The
lower or ventral part of each sclerotome (the inner and lower edge of
the cube-shaped provertebra) divides into two plates, which grow round
the chorda, and thus form the foundation of the body of the vertebra
(wh). The upper plate presses between the chorda and the medullary
tube, the lower between the chorda and the alimentary canal (Figure
1.137 C). As the plates of two opposite provertebral pieces unite from
the right and left, a circular sheath is formed round this part of the
chorda. From this develops the BODY of a vertebra--that is to say, the
massive lower or ventral half of the bony ring, which is called the
"vertebra" proper and surrounds the medullary tube (Figures 1.164 to
1.166). The upper or dorsal half of this bony ring, the vertebral arch
(Figure 1.145 wb), arises in just the same way from the upper part of
the skeletal plate, and therefore from the inner and upper edge of the
cube-shaped primitive vertebra. As the upper edges of two opposing
somites grow together over the medullary tube from right and left, the
vertebra-arch becomes closed.

The whole of the secondary vertebra, which is thus formed from the
union of the skeletal plates of two provertebral pieces and encloses a
part of the chorda in its body, consists at first of a rather soft
mass of cells; this afterwards passes into a firmer, cartilaginous
stage, and finally into a third, permanent, bony stage. These three
stages can generally be distinguished in the greater part of the
skeleton of the higher vertebrates; at first most parts of the
skeleton are soft, tender, and membranous; they then become
cartilaginous in the course of their development, and finally bony.

(FIGURES 1.168 AND 1.169. Head of a chick embryo, of the third day.
Figure 1.168 from the front, Figure 1.169 from the right. n
rudimentary nose (olfactory pit), l rudimentary eye (optic pit,
lens-cavity), g rudimentary ear (auditory pit), v fore-brain, gl
eye-cleft. Of the three pairs of gill-arches the first has passed into
a process of the upper jaw (o) and of the lower jaw (u). (From
Kolliker.))

At the head part of the embryo in the amniotes there is not generally
a cleavage of the middle germinal layer into provertebral and lateral
plates, but the dorsal and ventral somites are blended from the first,
and form what are called the "head-plates" (Figure 1.148 k). From
these are formed the skull, the bony case of the brain, and the
muscles and corium of the body. The skull develops in the same way as
the membranous vertebral column. The right and left halves of the head
curve over the cerebral vesicle, enclose the foremost part of the
chorda below, and thus finally form a simple, soft, membranous capsule
about the brain. This is afterwards converted into a cartilaginous
primitive skull, such as we find permanently in many of the fishes.
Much later this cartilaginous skull becomes the permanent bony skull
with its various parts. The bony skull in man and all the other
amniotes is more highly differentiated and modified than that of the
lower vertebrates, the amphibia and fishes. But as the one has arisen
phylogenetically from the other, we must assume that in the former no
less than the latter the skull was originally formed from the
sclerotomes of a number of (at least nine) head-somites.

While the articulation of the vertebrate body is always obvious in the
episoma or dorsal body, and is clearly expressed in the segmentation
of the muscular plates and vertebrae, it is more latent in the
hyposoma or ventral body. Nevertheless, the hyposomites of the vegetal
half of the body are not less important than the episomites of the
animal half. The segmentation in the ventral cavity affects the
following principal systems of organs: 1, the gonads or sex-glands
(gonotomes); 2, the nephridia or kidneys (nephrotomes); and 3, the
head-gut with its gill-clefts (branchiotomes).

(FIGURE 1.170. Head of a dog embryo, seen from the front. a the two
lateral halves of the foremost cerebral vesicle, b rudimentary eye, c
middle cerebral vesicle, de first pair of gill-arches (e upper-jaw
process, d lower-jaw process), f, f apostrophe, f double apostrophe,
second, third, and fourth pairs of gill-arches, g h i k heart (g
right, h left auricle; i left, k right ventricle), l origin of the
aorta with three pairs of arches, which go to the gill-arches. (From
Bischoff.))

The metamerism of the hyposoma is less conspicuous because in all the
craniotes the cavities of the ventral segments, in the walls of which
the sexual products are developed, have long since coalesced, and
formed a single large body-cavity, owing to the disappearance of the
partition. This cenogenetic process is so old that the cavity seems to
be unsegmented from the first in all the craniotes, and the rudiment
of the gonads also is almost always unsegmented. It is the more
interesting to learn that, according to the important discovery of
Ruckert, this sexual structure is at first segmental even in the
actual selachii, and the several gonotomes only blend into a simple
sexual gland on either side secondarily.

(FIGURE 1.171. Human embryo of the fourth week (twenty-six days old),
one-fourth of an inch in length magnified twenty times, showing: point
of development of the hind-leg, umbilical cord (underneath it the
tail, bent upwards), trigeminal nerve V Trigeminus, optic-muscle nerve
III Oculo-motorius, rolling muscle nerve IV Trochlearis, rudiment of
ear (labyrinthic vesicles), pneumogastric nerve X Vagus, terminal
nerve XI Accessorius, hypoglossal nerve XII Hypoglossus, first spinal
nerve, point of development of arm (or fore-leg), true spinal nerve.
(From Moll.) The rudiments of the cerebral nerves and the roots of the
spinal nerves are especially marked. Underneath the four gill-arches
(left side) is the heart (with auricle, V and ventricle, K), under
this again the liver (L).)

Amphioxus, the sole surviving representative of the acrania, once more
yields us most interesting information; in this case the sexual glands
remain segmented throughout life. The sexually mature lancelet has, on
the right and left of the gut, a series of metamerous sacs, which are
filled with ova in the female and sperm in the male. These segmental
gonads are originally nothing else than the real gonotomes, separate
body-cavities, formed from the hyposomites of the trunk.

The gonads are the most important segmental organs of the hyposoma, in
the sense that they are phylogenetically the oldest. We find sexual
glands (as pouch-like appendages of the gastro-canal system) in most
of the lower animals, even in the medusae, etc., which have no
kidneys. The latter appear first (as a pair of excretory tubes) in the
platodes (turbellaria), and have probably been inherited from these by
the articulates (annelids) on the one hand and the unarticulated
prochordonia on the other, and from these passed to the articulated
vertebrates. The oldest form of the kidney system in this stem are the
segmental pronephridia or prorenal canals, in the same arrangement as
Boveri found them in the amphioxus. They are small canals that lie in
the frontal plane, on each side of the chorda, between the episoma and
hyposoma (Figure 1.102 n); their internal funnel-shaped opening leads
into the various body-cavities, their outer opening is the lateral
furrow of the epidermis. Originally they must have had a double
function, the carrying away of the urine from the episomites and the
release of the sexual cells from the hyposomites.

The recent investigations of Ruckert and Van Wijhe on the mesodermic
segments of the trunk and the excretory system of the selachii show
that these "primitive fishes" are closely related to the amphioxus in
this further respect. The transverse section of the shark-embryo in
Figure 1.161 shows this very clearly.

In other higher vertebrates, also, the kidneys develop (though very
differently formed later on) from similar structures, which have been
secondarily derived from the segmental pronephridia of the acrania.
The parts of the mesoderm at which the first traces of them are found
are usually called the middle or mesenteric plates. As the first
traces of the gonads make their appearance in the lining of these
middle plates nearer inward (or the middle) from the inner funnels of
the nephro-canals, it is better to count this part of the mesoderm
with the hyposoma.

The chief and oldest organ of the vertebrate hyposoma, the alimentary
canal, is generally described as an unsegmented organ. But we could
just as well say that it is the oldest of all the segmented organs of
the vertebrate; the double row of the coelom-pouches grows out of the
dorsal wall of the gut, on either side of the chorda. In the brief
period during which these segmental coelom-pouches are still openly
connected with the gut, they look just like a double chain of
segmented visceral glands. But apart from this, we have originally in
all vertebrates an important articulation of the fore-gut, that is
wanting in the lower gut, the segmentation of the branchial (gill)
gut.

(FIGURE 1.172. Transverse section of the shoulder and fore-limb (wing)
of a chick-embryo of the fourth day, magnified about twenty times.
Beside the medullary tube we can see on each side three clear streaks
in the dark dorsal wall, which advance into the rudimentary fore-limb
or wing (e). The uppermost of them is the muscular plate; the middle
is the hind and the lowest the fore root of a spinal nerve. Under the
chorda in the middle is the single aorta, at each side of it a
cardinal vein, and below these the primitive kidneys. The gut is
almost closed. The ventral wall advances into the amnion, which
encloses the embryo. (From Remak.)

FIGURE 1.173. Transverse section of the pelvic region and hind legs of
a chick-embryo of the fourth day, magnified about forty times. h
horn-plate, w medullary tube, n canal of the tube, u primitive
kidneys, x chorda, e hind legs, b allantoic canal in the ventral wall,
t aorta, v cardinal veins, a gut, d gut-gland layer, f gut-fibre
layer, g embryonic epithelium, r dorsal muscles, c body-cavity or
coeloma. (From Waldeyer.))

The gill-clefts, which originally in the older acrania pierced the
wall of the fore-gut, and the gill-arches that separated them, were
presumably also segmental, and distributed among the various metamera
of the chain, like the gonads in the after-gut and the nephridia. In
the amphioxus, too, they are still segmentally formed. Probably there
was a division of labour of the hyposomites in the older (and long
extinct) acrania, in such wise that those of the fore-gut took over
the function of breathing and those of the after-gut that of
reproduction. The former developed into gill-pouches, the latter into
sex-pouches. There may have been primitive kidneys in both. Though the
gills have lost their function in the higher animals, certain parts of
them have been generally maintained in the embryo by a tenacious
heredity. At a very early stage we notice in the embryo of man and the
other amniotes, at each side of the head, the remarkable and important
structures which we call the gill-arches and gill-clefts (Figures
1.167 to 1.170 f). They belong to the characteristic and inalienable
organs of the amniote-embryo, and are found always in the same spot
and with the same arrangement and structure. There are formed to the
right and left in the lateral wall of the fore-gut cavity, in its
foremost part, first a pair and then several pairs of sac-shaped
inlets, that pierce the whole thickness of the lateral wall of the
head. They are thus converted into clefts, through which one can
penetrate freely from without into the gullet. The wall thickens
between these branchial folds, and changes into an arch-like or
sickle-shaped piece--the gill, or gullet-arch. In this the muscles and
skeletal parts of the branchial gut separate; a blood-vessel arch
rises afterwards on their inner side (Figure 1.98 ka). The number of
the branchial arches and the clefts that alternate with them is four
or five on each side in the higher vertebrates (Figure 1.170 d, f, f
apostrophe, f double apostrophe). In some of the fishes (selachii) and
in the cyclostoma we find six or seven of them permanently.

These remarkable structures had originally the function of respiratory
organs--gills. In the fishes the water that serves for breathing, and
is taken in at the mouth, still always passes out by the branchial
clefts at the sides of the gullet. In the higher vertebrates they
afterwards disappear. The branchial arches are converted partly into
the jaws, partly into the bones of the tongue and the ear. From the
first gill-cleft is formed the tympanic cavity of the ear.

There are few parts of the vertebrate organism that, like the outer
covering or integument of the body, are not subject to metamerism. The
outer skin (epidermis) is unsegmented from the first, and proceeds
from the continuous horny plate. Moreover, the underlying cutis is
also not metamerous, although it develops from the segmental structure
of the cutis-plates (Figures 1.161 and 1.162 cp). The vertebrates are
strikingly and profoundly different from the articulates in these
respects also.

Further, most of the vertebrates still have a number of unarticulated
organs, which have arisen locally, by adaptation of particular parts
of the body to certain special functions. Of this character are the
sense-organs in the episoma, and the limbs, the heart, the spleen, and
the large visceral glands--lungs, liver, pancreas, etc.--in the
hyposoma. The heart is originally only a local spindle-shaped
enlargement of the large ventral blood-vessel or principal vein, at
the point where the subintestinal passes into the branchial artery, at
the limit of the head and trunk (Figures 1.170 and 1.171). The three
higher sense-organs--nose, eye, and ear--were originally developed in
the same form in all the craniotes, as three pairs of small
depressions in the skin at the side of the head.

The organ of smell, the nose, has the appearance of a pair of small
pits above the mouth-aperture, in front of the head (Figure 1.169 n).
The organ of sight, the eye, is found at the side of the head, also in
the shape of a depression (Figures 1.169 l and 1.170 b), to which
corresponds a large outgrowth of the foremost cerebral vesicle on each
side. Farther behind, at each side of the head, there is a third
depression, the first trace of the organ of hearing (Figure 1.169 g).
As yet we can see nothing of the later elaborate structure of these
organs, nor of the characteristic build of the face.

(FIGURE 1.174. Development of the lizard's legs (Lacerta agilis), with
special relation to their blood-vessels. 1, 3, 5, 7, 9, 11 right
fore-leg; 13, 15 left fore-leg; 2, 4, 6, 8, 10, 12 right hind-leg; 14,
16 left hind-leg; SRV lateral veins of the trunk, VU umbilical vein.
(From F. Hochstetter.))

When the human embryo has reached this stage of development, it can
still scarcely be distinguished from that of any other higher
vertebrate. All the chief parts of the body are now laid down: the
head with the primitive skull, the rudiments of the three higher
sense-organs and the five cerebral vesicles, and the gill-arches and
clefts; the trunk with the spinal cord, the rudiment of the vertebral
column, the chain of metamera, the heart and chief blood-vessels, and
the kidneys. At this stage man is a higher vertebrate, but shows no
essential morphological difference from the embryos of the mammals,
the birds, the reptiles, etc. This is an ontogenetic fact of the
utmost significance. From it we can gather the most important
phylogenetic conclusions.

There is still no trace of the limbs. Although head and trunk are
separated and all the principal internal organs are laid down, there
is no indication whatever of the "extremities" at this stage; they are
formed later on. Here again we have a fact of the utmost interest. It
proves that the older vertebrates had no feet, as we find to be the
case in the lowest living vertebrates (amphioxus and the cyclostoma).
The descendants of these ancient footless vertebrates only acquired
extremities--two fore-legs and two hind-legs--at a much later stage of
development. These were at first all alike, though they afterwards
vary considerably in structure--becoming fins (of breast and belly) in
the fishes, wings and legs in the birds, fore and hind legs in the
creeping animals, arms and legs in the apes and man. All these parts
develop from the same simple original structure, which forms
secondarily from the trunk-wall (Figures 1.172 and 1.173). They have
always the appearance of two pairs of small buds, which represent at
first simple roundish knobs or plates. Gradually each of these plates
becomes a large projection, in which we can distinguish a small inner
part and a broader outer part. The latter is the rudiment of the foot
or hand, the former that of the leg or arm. The similarity of the
original rudiment of the limbs in different groups of vertebrates is
very striking.

(FIGURE 1.175. Human embryo, five weeks old, half an inch long, seen
from the right, magnified ten times. (From Russel Bardeen and Harmon
Lewis.) In the undissected head we see the eye, mouth, and ear. In the
trunk the skin and part of the muscles have been removed, so that the
cartilaginous vertebral column is free; the dorsal root of a spinal
nerve goes out from each vertebra (towards the skin of the back). In
the middle of the lower half of the figure part of the ribs and
intercostal muscles are visible. The skin and muscles have also been
removed from the right limbs; the internal rudiments of the five
fingers of the hand, and five toes of the foot, are clearly seen
within the fin-shaped plate, and also the strong network of nerves
that goes from the spinal cord to the extremities. The tail projects
under the foot, and to the right of it is the first part of the
umbilical cord.)

How the five fingers or toes with their blood-vessels gradually
differentiate within the simple fin-like structure of the limbs can be
seen in the instance of the lizard in Figure 1.174. They are formed in
just the same way in man: in the human embryo of five weeks the five
fingers can clearly be distinguished within the fin-plate (Figure
1.175).

The careful study and comparison of human embryos with those of other
vertebrates at this stage of development is very instructive, and
reveals more mysteries to the impartial student than all the religions
in the world put together. For instance, if we compare attentively the
three successive stages of development that are represented, in twenty
different amniotes we find a remarkable likeness. When we see that as
a fact twenty different amniotes of such divergent characters develop
from the same embryonic form, we can easily understand that they may
all descend from a common ancestor.

(FIGURES 1.176 TO 1.178. Embryos of the bat (Vespertilio murinus) at
three different stages. (From Oscar Schultze.) Figure 1.176:
Rudimentary limbs (v fore-leg, h hind-leg). l lenticular depression, r
olfactory pit, ok upper jaw, uk lower jaw, k2, k3, k4 first, second
and third gill-arches, a amnion, n umbilical vessel, d yelk-sac.
Figure 1.177: Rudiment of flying membrane, membranous fold between
fore and hind leg. n umbilical vessel, o ear-opening, f flying
membrane. Figure 1.178: The flying membrane developed and stretched
across the fingers of the hands, which cover the face.)

In the first stage of development, in which the head with the five
cerebral vesicles is already clearly indicated, but there are no
limbs, the embryos of all the vertebrates, from the fish to man, are
only incidentally or not at all different from each other. In the
second stage, which shows the limbs, we begin to see differences
between the embryos of the lower and higher vertebrates; but the human
embryo is still hardly distinguishable from that of the higher
mammals. In the third stage, in which the gill-arches have disappeared
and the face is formed, the differences become more pronounced. These
are facts of a significance that cannot be exaggerated.* (* Because
they show how the most diverse structures may be developed from a
common form. As we actually see this in the case of the embryos, we
have a right to assume it in that of the stem-forms. Nevertheless,
this resemblance, however great, is never a real identity. Even the
embryos of the different individuals of one species are usually not
really identical. If the reader can consult the complete edition of
this work at a library, he will find six plates illustrating these
twenty embryos.)

If there is an intimate causal connection between the processes of
embryology and stem-history, as we must assume in virtue of the laws
of heredity, several important phylogenetic conclusions follow at once
from these ontogenetic facts. The profound and remarkable similarity
in the embryonic development of man and the other vertebrates can only
be explained when we admit their descent from a common ancestor. As a
fact, this common descent is now accepted by all competent scientists;
they have substituted the natural evolution for the supernatural
creation of organisms.


CHAPTER 1.15. FOETAL MEMBRANES AND CIRCULATION.

Among the many interesting phenomena that we have encountered in the
course of human embryology, there is an especial importance in the
fact that the development of the human body follows from the beginning
just the same lines as that of the other viviparous mammals. As a
fact, all the embryonic peculiarities that distinguish the mammals
from other animals are found also in man; even the ovum with its
distinctive membrane (zona pellucida, Figure 1.14) shows the same
typical structure in all mammals (apart from the older oviparous
monotremes). It has long since been deduced from the structure of the
developed man that his natural place in the animal kingdom is among
the mammals. Linne (1735) placed him in this class with the apes, in
one and the same order (primates), in his Systema Naturae. This
position is fully confirmed by comparative embryology. We see that man
entirely resembles the higher mammals, and most of all the apes, in
embryonic development as well as in anatomic structure. And if we seek
to understand this ontogenetic agreement in the light of the
biogenetic law, we find that it proves clearly and necessarily the
descent of man from a series of other mammals, and proximately from
the primates. The common origin of man and the other mammals from a
single ancient stem-form can no longer be questioned; nor can the
immediate blood-relationship of man and the ape.

(FIGURE 1.179. Human embryos from the second to the fifteenth week,
natural size, seen from the left, the curved back turned towards the
right. (Mostly from Ecker.) II of fourteen days. III of three weeks.
IV of four weeks. V of five weeks. VI of six weeks. VII of seven
weeks. VIII of eight weeks. XII of twelve weeks. XV of fifteen weeks.)

The essential agreement in the whole bodily form and inner structure
is still visible in the embryo of man and the other mammals at the
late stage of development at which the mammal-body can be recognised
as such. But at a somewhat earlier stage, in which the limbs,
gill-arches, sense-organs, etc., are already outlined, we cannot yet
recognise the mammal embryos as such, or distinguish them from those
of birds and reptiles. When we consider still earlier stages of
development, we are unable to discover any essential difference in
bodily structure between the embryos of these higher vertebrates and
those of the lower, the amphibia and fishes. If, in fine, we go back
to the construction of the body out of the four germinal layers, we
are astonished to perceive that these four layers are the same in all
vertebrates, and everywhere take a similar part in the building-up of
the fundamental organs of the body. If we inquire as to the origin of
these four secondary layers, we learn that they always arise in the
same way from the two primary layers; and the latter have the same
significance in all the metazoa (i.e., all animals except the
unicellulars). Finally, we see that the cells which make up the
primary germinal layers owe their origin in every case to the repeated
cleavage of a single simple cell, the stem-cell or fertilised ovum.

(FIGURE 1.180. Very young human embryo of the fourth week, one-fourth
of an inch long (taken from the womb of a suicide eight hours after
death). (From Rabl.) n nasal pits, a eye, u lower jaw, z arch of hyoid
bone, k3 and k4 third and fourth gill-arch, h heart; s primitive
segments, vg fore-limb (arm), hg hind-limb (leg), between the two the
ventral pedicle.)

It is impossible to lay too much stress on this remarkable agreement
in the chief embryonic features in man and the other animals. We shall
make use of it later on for our monophyletic theory of descent--the
hypothesis of a common descent of man and all the metazoa from the
gastraea. The first rudiments of the principal parts of the body,
especially the oldest organ, the alimentary canal, are the same
everywhere; they have always the same extremely simple form. All the
peculiarities that distinguish the various groups of animals from each
other only appear gradually in the course of embryonic development;
and the closer the relation of the various groups, the later they are
found. We may formulate this phenomenon in a definite law, which may
in a sense be regarded as an appendix to our biogenetic law. This is
the law of the ontogenetic connection of related animal forms. It
runs: The closer the relation of two fully-developed animals in
respect of their whole bodily structure, and the nearer they are
connected in the classification of the animal kingdom, the longer do
their embryonic forms retain their identity, and the longer is it
impossible (or only possible on the ground of subordinate features) to
distinguish between their embryos. This law applies to all animals
whose embryonic development is, in the main, an hereditary summary of
their ancestral history, or in which the original form of development
has been faithfully preserved by heredity. When, on the other hand, it
has been altered by cenogenesis, or disturbance of development, we
find a limitation of the law, which increases in proportion to the
introduction of new features by adaptation (cf. Chapter 1.1). Thus the
apparent exceptions to the law can always be traced to cenogenesis.

(FIGURE 1.181. Human embryo of the middle of the fifth week, one-third
of an inch long. (From Rabl.) Letters as in Figure 1.180, except sk
curve of skull, ok upper jaw, hb neck-indentation.)

When we apply to man this law of the ontogenetic connection of related
forms, and run rapidly over the earliest stages of human development
with an eye to it, we notice first of all the structural identity of
the ovum in man and the other mammals at the very beginning (Figures
1.1 and 1.14). The human ovum possesses all the distinctive features
of the ovum of the viviparous mammals, especially the characteristic
formation of its membrane (zona pellucida), which clearly
distinguishes it from the ovum of all other animals. When the human
foetus has attained the age of fourteen days, it forms a round vesicle
(or "embryonic vesicle") about a quarter of an inch in diameter. A
thicker part of its border forms a simple sole-shaped embryonic shield
one-twelfth of an inch long (Figure 1.133). On its dorsal side we find
in the middle line the straight medullary furrow, bordered by the two
parallel dorsal or medullary swellings. Behind, it passes by the
neurenteric canal into the primitive gut or primitive groove. From
this the folding of the two coelom-pouches proceeds in the same way as
in the other mammals (cf. Figures 1.96 and 1.97). In the middle of the
sole-shaped embryonic shield the first primitive segments immediately
begin to make their appearance. At this age the human embryo cannot be
distinguished from that of other mammals, such as the hare or dog.

A week later (or after the twenty-first day) the human embryo has
doubled its length; it is now about one-fifth of an inch long, and,
when seen from the side, shows the characteristic bend of the back,
the swelling of the head-end, the first outline of the three higher
sense-organs, and the rudiments of the gill-clefts, which pierce the
sides of the neck (Figure 1.179, III). The allantois has grown out of
the gut behind. The embryo is already entirely enclosed in the amnion,
and is only connected in the middle of the belly by the vitelline duct
with the embryonic vesicle, which changes into the yelk-sac. There are
no extremities or limbs at this stage, no trace of arms or legs. The
head-end has been strongly differentiated from the tail-end; and the
first outlines of the cerebral vesicles in front, and the heart below,
under the fore-arm, are already more or less clearly seen. There is as
yet no real face. Moreover, we seek in vain at this stage a special
character that may distinguish the human embryo from that of other
mammals.

(FIGURE 1.182. Median longitudinal section of the tail of a human
embryo, two-thirds of an inch long. (From Ross Granville Harrison.)
Med medullary tube, Ca.fil caudal filament, ch chorda, ao caudal
artery, V.c.i caudal vein, an anus, S.ug sinus urogenitalis.)

A week later (after the fourth week, on the twenty-eighth to thirtieth
day of development) the human embryo has reached a length of about
one-third of an inch (Figure 1.179 IV). We can now clearly distinguish
the head with its various parts; inside it the five primitive cerebral
vesicles (fore-brain, middle-brain, intermediate-brain, hind-brain,
and after-brain); under the head the gill-arches, which divide the
gill-clefts; at the sides of the head the rudiments of the eyes, a
couple of pits in the outer skin, with a pair of corresponding simple
vesicles growing out of the lateral wall of the fore-brain (Figures
1.180, 1.181 a). Far behind the eyes, over the last gill-arches, we
see a vesicular rudiment of the auscultory organ. The rudimentary
limbs are now clearly outlined--four simple buds of the shape of round
plates, a pair of fore (vg) and a pair of hind legs (hg), the former a
little larger than the latter. The large head bends over the trunk,
almost at a right angle. The latter is still connected in the middle
of its ventral side with the embryonic vesicle; but the embryo has
still further severed itself from it, so that it already hangs out as
the yelk-sac. The hind part of the body is also very much curved, so
that the pointed tail-end is directed towards the head. The head and
face-part are sunk entirely on the still open breast. The bend soon
increases so much that the tail almost touches the forehead (Figure
1.179 V.; Figure 1.181). We may then distinguish three or four special
curves on the round dorsal surface--namely, a skull-curve in the
region of the second cerebral vesicle, a neck-curve at the beginning
of the spinal cord, and a tail-curve at the fore-end. This pronounced
curve is only shared by man and the higher classes of vertebrates (the
amniotes); it is much slighter, or not found at all, in the lower
vertebrates. At this age (four weeks) man has a considerable tail,
twice as long as his legs. A vertical longitudinal section through the
middle plane of this tail (Figure 1.182) shows that the hinder end of
the spinal marrow extends to the point of the tail, as also does the
underlying chorda (ch), the terminal continuation of the vertebral
column. Of the latter, the rudiments of the seven coccygeal (or
lowest) vertebrae are visible--thirty-two indicates the third and
thirty-six the seventh of these. Under the vertebral column we see the
hindmost ends of the two large blood-vessels of the tail, the
principal artery (aorta caudalis or arteria sacralis media, Ao), and
the principal vein (vena caudalis or sacralis media). Underneath is
the opening of the anus (an) and the urogenital sinus (S.ug). From
this anatomic structure of the human tail it is perfectly clear that
it is the rudiment of an ape-tail, the last hereditary relic of a long
hairy tail, which has been handed down from our tertiary primate
ancestors to the present day.

(FIGURE 1.183. Human embryo, four weeks old, opened on the ventral
side. Ventral and dorsal walls are cut away, so as to show the
contents of the pectoral and abdominal cavities. All the appendages
are also removed (amnion, allantois, yelk-sac), and the middle part of
the gut. n eye, 3 nose, 4 upper jaw, 5 lower jaw, 6 second, 6 double
apostrophe, third gill-arch, ov heart (o right, o apostrophe, left
auricle; v right, v apostrophe, left ventricle), b origin of the
aorta, f liver (u umbilical vein), e gut (with vitelline artery, cut
off at a apostrophe), j apostrophe, vitelline vein, m primitive
kidneys, t rudimentary sexual glands, r terminal gut (cut off at the
mesentery z), n umbilical artery, u umbilical vein, 9 fore-leg, 9
apostrophe, hind-leg. (From Coste.)

FIGURE 1.184. Human embryo, five weeks old, opened from the ventral
side (as in Figure 1.183). Breast and belly-wall and liver are
removed. 3 outer nasal process, 4 upper jaw, 5 lower jaw, z tongue, v
right, v apostrophe, left ventricle of heart, o apostrophe, left
auricle, b origin of aorta, b apostrophe, b double apostrophe, b
triple apostrophe, first, second, and third aorta-arches, c, c
apostrophe, c double apostrophe, vena cava, ae lungs (y pulmonary
artery), e stomach, m primitive kidneys (j left vitelline vein, s
cystic vein, a right vitelline artery, n umbilical artery, u umbilical
vein), x vitelline duct, i rectum, 8 tail, 9 fore-leg, 9 apostrophe,
hind-leg. (From Coste.))

It sometimes happens that we find even external relics of this tail
growing. According to the illustrated works of Surgeon-General
Bernhard Ornstein, of Greece, these tailed men are not uncommon; it is
not impossible that they gave rise to the ancient fables of the
satyrs. A great number of such cases are given by Max Bartels in his
essay on "Tailed Men" (1884, in the Archiv fur Anthropologie, Band
15), and critically examined. These atavistic human tails are often
mobile; sometimes they contain only muscles and fat, sometimes also
rudiments of caudal vertebrae. They have a length of eight to ten
inches and more. Granville Harrison has very carefully studied one of
these cases of "pigtail," which he removed by operation from a six
months old child in 1901. The tail moved briskly when the child cried
or was excited, and was drawn up when at rest.

(FIGURE 1.185. The head of Miss Julia Pastrana. (From a photograph by
Hintze.)

FIGURE 1.186. Human ovum of twelve to thirteen days (?). (From Allen
Thomson.) 1. Not opened, natural size. 2. Opened and magnified. Within
the outer chorion the tiny curved foetus lies on the large embryonic
vesicle, to the left above.

FIGURE 1.187. Human ovum of ten days. (From Allen Thomson.) Natural
size, opened; the small foetus in the right half, above.

FIGURE 1.188. Human foetus of ten days, taken from the preceding ovum,
magnified ten times, a yelk-sac, b neck (the medullary groove already
closed), c head (with open medullary groove), d hind part (with open
medullary groove), e a shred of the amnion.

FIGURE 1.189. Human ovum of twenty to twenty-two days. (From Allen
Thomson.) Natural size, opened. The chorion forms a spacious vesicle,
to the inner wall of which the small foetus (to the right above) is
attached by a short umbilical cord.

FIGURE 1.190. Human foetus of twenty to twenty-two days, taken from
the preceding ovum, magnified. a amnion, b yelk-sac, c lower-jaw
process of the first gill-arch, d upper-jaw process of same, e second
gill-arch (two smaller ones behind). Three gill-clefts are clearly
seen. f rudimentary fore-leg, g auditory vesicle, h eye, i heart.)

In the opinion of some travellers and anthropologists, the atavistic
tail-formation is hereditary in certain isolated tribes (especially in
south-eastern Asia and the archipelago), so that we might speak of a
special race or "species" of tailed men (Homo caudatus). Bartels has
"no doubt that these tailed men will be discovered in the advance of
our geographical and ethnographical knowledge of the lands in
question" (Archiv fur Anthropologie, Band 15 page 129).

When we open a human embryo of one month (Figure 1.183), we find the
alimentary canal formed in the body-cavity, and for the most part cut
off from the embryonic vesicle. There are both mouth and anus
apertures. But the mouth-cavity is not yet separated from the nasal
cavity, and the face not yet shaped. The heart shows all its four
sections; it is very large, and almost fills the whole of the pectoral
cavity (Figure 1.183 ov). Behind it are the very small rudimentary
lungs. The primitive kidneys (m) are very large; they fill the greater
part of the abdominal cavity, and extend from the liver (f) to the
pelvic gut. Thus at the end of the first month all the chief organs
are already outlined. But there are at this stage no features by which
the human embryo materially differs from that of the dog, the hare,
the ox, or the horse--in a word, of any other higher mammal. All these
embryos have the same, or at least a very similar, form; they can at
the most be distinguished from the human embryo by the total size of
the body or some other insignificant difference in size. Thus, for
instance, in man the head is larger in proportion to the trunk than in
the ox. The tail is rather longer in the dog than in man. These are
all negligible differences. On the other hand, the whole internal
organisation and the form and arrangement of the various organs are
essentially the same in the human embryo of four weeks as in the
embryos of the other mammals at corresponding stages.

(FIGURE 1.191. Human embryo of sixteen to eighteen days. (From Coste.)
Magnified. The embryo is surrounded by the amnion, (a), and lies free
with this in the opened embryonic vesicle. The belly is drawn up by
the large yelk-sac (d), and fastened to the inner wall of the
embryonic membrane by the short and thick pedicle (b). Hence the
normal convex curve of the back (Figure 1.190) is here changed into an
abnormal concave surface. h heart, m parietal mesoderm. The spots on
the outer wall of the serolemma are the roots of the branching
chorion-villi, which are free at the border.

FIGURE 1.192. Human embryo of the fourth week, one-third of an inch
long, lying in the dissected chorion.

FIGURE 1.193. Human embryo of the fourth week, with its membranes,
like Figure 1.192, but a little older. The yelk-sac is rather smaller,
the amnion and chorion larger.)

It is otherwise in the second month of human development. Figure 1.179
represents a human embryo of six weeks (VI), one of seven weeks (VII),
and one of eight weeks (VIII), at natural size. The differences which
mark off the human embryo from that of the dog and the lower mammals
now begin to be more pronounced. We can see important differences at
the sixth, and still more at the eighth week, especially in the
formation of the head. The size of the various sections of the brain
is greater in man, and the tail is shorter. Other differences between
man and the lower mammals are found in the relative size of the
internal organs. But even at this stage the human embryo differs very
little from that of the nearest related mammals--the apes, especially
the anthropomorphic apes. The features by means of which we
distinguish between them are not clear until later on. Even at a much
more advanced stage of development, when we can distinguish the human
foetus from that of the ungulates at a glance, it still closely
resembles that of the higher apes. At last we get the distinctive
features, and we can distinguish the human embryo confidently at the
first glance from that of all other mammals during the last four
months of foetal life--from the sixth to the ninth month of pregnancy.
Then we begin to find also the differences between the various races
of men, especially in regard to the formation of the skull and the
face. (Cf. Chapter 2.23.)

(FIGURE 1.194. Human embryo with its membranes, six weeks old. The
outer envelope of the whole ovum is the chorion, thickly covered with
its branching villi, a product of the serous membrane. The embryo is
enclosed in the delicate amnion-sac. The yelk-sac is reduced to a
small pear-shaped umbilical vesicle; its thin pedicle, the long
vitelline duct, is enclosed in the umbilical cord. In the latter,
behind the vitelline duct, is the much shorter pedicle of the
allantois, the inner lamina of which (the gut-gland layer) forms a
large vesicle in most of the mammals, while the outer lamina is
attached to the inner wall of the outer embryonic coat, and forms the
placenta there. (Half diagrammatic.))

The striking resemblance that persists so long between the embryo of
man and of the higher apes disappears much earlier in the lower apes.
It naturally remains longest in the large anthropomorphic apes
(gorilla, chimpanzee, orang, and gibbon). The physiognomic similarity
of these animals, which we find so great in their earlier years,
lessens with the increase of age. On the other hand, it remains
throughout life in the remarkable long-nosed ape of Borneo (Nasalis
larvatus). Its finely-shaped nose would be regarded with envy by many
a man who has too little of that organ. If we compare the face of the
long-nosed ape with that of abnormally ape-like human beings (such as
the famous Miss Julia Pastrana, Figure 1.185), it will be admitted to
represent a higher stage of development. There are still people among
us who look especially to the face for the "image of God in man." The
long-nosed ape would have more claim to this than some of the
stumpy-nosed human individuals one meets.

This progressive divergence of the human from the animal form, which
is based on the law of the ontogenetic connection between related
forms, is found in the structure of the internal organs as well as in
external form. It is also expressed in the construction of the
envelopes and appendages that we find surrounding the foetus
externally, and that we will now consider more closely. Two of these
appendages--the amnion and the allantois--are only found in the three
higher classes of vertebrates, while the third, the yelk-sac, is found
in most of the vertebrates. This is a circumstance of great
importance, and it gives us valuable data for constructing man's
genealogical tree.

(FIGURE 1.195. Diagram of the embryonic organs of the mammal (foetal
membranes and appendages). (From Turner.) E, M, H outer, middle, and
inner germ layer of the embryonic shield, which is figured in median
longitudinal section, seen from the left. am amnion. AC amniotic
cavity, UV yelk-sac or umbilical vesicle, ALC allantois, al pericoelom
or serocoelom (inter-amniotic cavity), sz serolemma (or serous
membrane), pc prochorion (with villi).)

As regards the external membrane that encloses the ovum in the mammal
womb, we find it just the same in man as in the higher mammals. The
ovum is, the reader will remember, first surrounded by the transparent
structureless ovolemma or zona pellucida (Figures 1.1 and 1.14). But
very soon, even in the first week of development, this is replaced by
the permanent chorion. This is formed from the external layer of the
amnion, the serolemma, or "serous membrane," the formation of which we
shall consider presently; it surrounds the foetus and its appendages
as a broad, completely closed sac; the space between the two, filled
with clear watery fluid, is the serocoelom, or interamniotic cavity
("extra-embryonic body-cavity"). But the smooth surface of the sac is
quickly covered with numbers of tiny tufts, which are really hollow
outgrowths like the fingers of a glove (Figures 1.186, 1.191 and 1.198
chz). They ramify and push into the corresponding depressions that are
formed by the tubular glands of the mucous membrane of the maternal
womb. Thus, the ovum secures its permanent seat (Figures 1.186 to
1.194).

In human ova of eight to twelve days this external membrane, the
chorion, is already covered with small tufts or villi, and forms a
ball or spheroid of one-fourth to one-third of an inch in diameter
(Figures 1.186 to 1.188). As a large quantity of fluid gathers inside
it, the chorion expands more and more, so that the embryo only
occupies a small part of the space within the vesicle. The villi of
the chorion grow larger and more numerous. They branch out more and
more. At first the villi cover the whole surface, but they afterwards
disappear from the greater part of it; they then develop with
proportionately greater vigour at a spot where the placenta is formed
from the allantois.

When we open the chorion of a human embryo of three weeks, we find on
the ventral side of the foetus a large round sac, filled with fluid.
This is the yelk-sac, or "umbilical vesicle," the origin of which we
have considered previously. The larger the embryo becomes the smaller
we find the yelk-sac. In the end we find the remainder of it in the
shape of a small pear-shaped vesicle, fastened to a long thin stalk
(or pedicle), and hanging from the open belly of the foetus (Figure
1.194). This pedicle is the vitelline duct, and is separated from the
body at the closing of the navel.

Behind the yelk-sac a second appendage, of much greater importance, is
formed at an early stage at the belly of the mammal embryo. This is
the allantois or "primitive urinary sac," an important embryonic
organ, only found in the three higher classes of vertebrates. In all
the amniotes the allantois quickly appears at the hinder end of the
alimentary canal, growing out of the cavity of the pelvic gut (Figure
1.147 r, u, Figure 1.195 ALC}.

(FIGURE 1.196. Diagrammatic frontal section of the pregnant human
womb. (From Longet.) The embryo hangs by the umbilical cord, which
encloses the pedicle of the allantois (al). nb umbilical vessel, am
amnion, ch chorion, ds decidua serotina, dv decidua vera, dr decidua
reflexa, z villi of the placenta, c cervix uteri, u uterus.)

The further development of the allantois varies considerably in the
three sub-classes of the mammals. The two lower sub-classes,
monotremes and marsupials, retain the simpler structure of their
ancestors, the reptiles. The wall of the allantois and the enveloping
serolemma remains smooth and without villi, as in the birds. But in
the third sub-class of the mammals the serolemma forms, by
invagination at its outer surface, a number of hollow tufts or villi,
from which it takes the name of the chorion or mallochorion. The
gut-fibre layer of the allantois, richly supplied with branches of the
umbilical vessel, presses into these tufts of the primary chorion, and
forms the "secondary chorion." Its embryonic blood-vessels are closely
correlated to the contiguous maternal blood-vessels of the environing
womb, and thus is formed the important nutritive apparatus of the
embryo which we call the placenta.

The pedicle of the allantois, which connects the embryo with the
placenta and conducts the strong umbilical vessels from the former to
the latter, is covered by the amnion, and, with this amniotic sheath
and the pedicle of the yelk-sac, forms what is called the umbilical
cord (Figure 1.196 al). As the large and blood-filled vascular network
of the foetal allantois attaches itself closely to the mucous lining
of the maternal womb, and the partition between the blood-vessels of
mother and child becomes much thinner, we get that remarkable
nutritive apparatus of the foetal body which is characteristic of the
placentalia (or choriata). We shall return afterwards to the closer
consideration of this (cf. Chapter 2.23).

In the various orders of mammals the placenta undergoes many
modifications, and these are in part of great evolutionary importance
and useful in classification. There is only one of these that need be
specially mentioned--the important fact, established by Selenka in
1890, that the distinctive human placentation is confined to the
anthropoids. In this most advanced group of the mammals the allantois
is very small, soon loses its cavity, and then, in common with the
amnion, undergoes certain peculiar changes. The umbilical cord
develops in this case from what is called the "ventral pedicle." Until
very recently this was regarded as a structure peculiar to man. We now
know from Selenka that the much-discussed ventral pedicle is merely
the pedicle of the allantois, combined with the pedicle of the amnion
and the rudimentary pedicle of the yelk-sac. It has just the same
structure in the orang and gibbon (Figure 1.197) and very probably in
the chimpanzee and gorilla, as in man; it is, therefore, not a
DISPROOF, but a striking fresh proof, of the blood-relationship of man
and the anthropoid apes.

(FIGURE 1.197. Male embryo of the Siamang-gibbon (Hylobates siamanga)
of Sumatra, two-thirds natural size; to the left the dissected uterus,
of which only the dorsal half is given. The embryo has been taken out,
and the limbs folded together; it is still connected by the umbilical
cord with the centre of the circular placenta which is attached to the
inside of the womb. This embryo takes the head-position in the womb,
and this is normal in man also.)

We find only in the anthropoid apes--the gibbon and orang of Asia and
the chimpanzee and gorilla of Africa--the peculiar and elaborate
formation of the placenta that characterises man (Figure 1.198). In
this case there is at an early stage an intimate blending of the
chorion of the embryo and the part of the mucous lining of the womb to
which it attaches. The villi of the chorion with the blood-vessels
they contain grow so completely into the tissue of the uterus, which
is rich in blood, that it becomes impossible to separate them, and
they form together a sort of cake. This comes away as the "afterbirth"
at parturition; at the same time, the part of the mucous lining of the
womb that has united inseparably with the chorion is torn away; hence
it is called the decidua ("falling-away membrane"), and also the
"sieve-membrane," because it is perforated like a sieve. We find a
decidua of this kind in most of the higher placentals; but it is only
in man and the anthropoid apes that it divides into three parts--the
outer, inner, and placental decidua. The external or true decidua
(Figure 1.196 du, Figure 1.199 g) is the part of the mucous lining of
the womb that clothes the inner surface of the uterine cavity wherever
it is not connected with the placenta. The placental or spongy decidua
(placentalis or serotina, Figure 1.196 ds, Figure 1.199 d) is really
the placenta itself, or the maternal part of it (placenta
uterina)--namely, that part of the mucous lining of the womb which
unites intimately with the chorion-villi of the foetal placenta. The
internal or false decidua (interna or reflexa, Figure 1.196 dr, Figure
1.199 f) is that part of the mucous lining of the womb which encloses
the remaining surface of the ovum, the smooth chorion (chorion laeve),
in the shape of a special thin membrane. The origin of these three
different deciduous membranes, in regard to which quite erroneous
views (still retained in their names) formerly prevailed, is now quite
clear, The external decidua vera is the specially modified and
subsequently detachable superficial stratum of the original mucous
lining of the womb. The placental decidua serotina is that part of the
preceding which is completely transformed by the ingrowth of the
chorion-villi, and is used for constructing the placenta. The inner
decidua reflexa is formed by the rise of a circular fold of the mucous
lining (at the border of the decidua vera and serotina), which grows
over the foetus (like the anmnion) to the end.

The peculiar anatomic features that characterise the human foetal
membranes are found in just the same way in the higher apes. Until
recently it was thought that the human embryo was distinguished by its
peculiar construction of a solid allantois and a special ventral
pedicle, and that the umbilical cord developed from this in a
different way than in the other mammals. The opponents of the
unwelcome "ape-theory" laid great stress on this, and thought they had
at last discovered an important indication that separated man from all
the other placentals. But the remarkable discoveries published by the
distinguished zoologist Selenka in 1890 proved that man shares these
peculiarities of placentation with the anthropoid apes, though they
are not found in the other apes. Thus the very feature which was
advanced by our critics as a disproof became a most important piece of
evidence in favour of our pithecoid origin.)

(FIGURE 1.198. Frontal section of the pregnant human womb, showing:
end of the decidua, uterine cavity, chorion (laeve), amniotic cavity,
foetal placenta, oviduct, spongy decidua serotina, umbilical vesicle,
amnion, decidua reflexa, decidua vera, muscular wall of the uterus,
mouth of the uterus. (From Turner.) The embryo (a month old) hangs in
the middle of the amniotic cavity by the ventral pedicle or umbilical
cord, which connects it with the placenta (above).

FIGURE 1.199. Human foetus, twelve weeks old, with its membranes.
Natural size. The umbilical cord goes from its navel to the placenta.
b amnion, c chorion, d placenta, d apostrophe, relics of villi on
smooth chorion, f internal or reflex decidua, g external or true
decidua. (From B. Schultze.)

FIGURE 1.200. Mature human foetus (at the end of pregnancy, in its
natural position, taken out of the uterine cavity). On the inner
surface of the latter (to the left) is the placenta, which is
connected by the umbilical cord with the child's navel. (From Bernhard
Schultze.))

Of the three vesicular appendages of the amniote embryo which we have
now described the amnion has no blood-vessels at any moment of its
existence. But the other two vesicles, the yelk-sac and the allantois,
are equipped with large blood-vessels, and these effect the
nourishment of the embryonic body. We may take the opportunity to make
a few general observations on the first circulation in the embryo and
its central organ, the heart. The first blood-vessels, the heart, and
the first blood itself, are formed from the gut-fibre layer. Hence it
was called by earlier embryologists the "vascular layer." In a sense
the term is quite correct. But it must not be understood as if all the
blood-vessels in the body came from this layer, or as if the whole of
this layer were taken up only with the formation of blood-vessels.
Neither of these suppositions is true. Blood-vessels may be formed
independently in other parts, especially in the various products of
the skin-fibre layer.

The first blood-vessels of the mammal embryo have been considered by
us previously, and we shall study the development of the heart in the
second volume.

(FIGURE 1.201. Vitelline vessels in the germinative area of a
chick-embryo, at the close of the third day of incubation. (From
Balfour.) The detached germinative area is seen from the ventral side:
the arteries are dark, the veins light. H heart, AA aorta-arches, Ao
aorta, R.Of.A right omphalo-mesenteric artery, S.T sinus terminalis,
L.Of and R.Of right and left omphalo-mesenteric veins, S.V sinus
venosus, D.C ductus Cuvieri, S.Ca.V and V.Ca fore and hind cardinal
veins.)

In every vertebrate it lies at first in the ventral wall of the
fore-gut, or in the ventral (or cardiac) mesentery, by which it is
connected for a time with the wall of the body. But it soon severs
itself from the place of its origin, and lies freely in a cavity--the
cardiac cavity. For a short time it is still connected with the former
by the thin plate of the mesocardium. Afterwards it lies quite free in
the cardiac cavity, and is only directly connected with the gut-wall
by the vessels which issue from it.

The fore-end of the spindle-shaped tube, which soon bends into an
S-shape (Figure 1.202), divides into a right and left branch. These
tubes are bent upwards arch-wise, and represent the first arches of
the aorta. They rise in the wall of the fore-gut, which they enclose
in a sense, and then unite above, in the upper wall of the fore
gut-cavity, to form a large single artery, that runs backward
immediately under the chorda, and is called the aorta (Figure 1.201
Ao). The first pair of aorta-arches rise on the inner wall of the
first pair of gill-arches, and so lie between the first gill-arch (k)
and the fore-gut (d), just as we find them throughout life in the
fishes. The single aorta, which results from the conjunction of these
two first vascular arches, divides again immediately into two parallel
branches, which run backwards on either side of the chorda. These are
the primitive aortas which we have already mentioned; they are also
called the posterior vertebral arteries. These two arteries now give
off at each side, behind, at right angles, four or five branches, and
these pass from the embryonic body to the germinative area, they are
called omphalo-mesenteric or vitelline arteries. They represent the
first beginning of a foetal circulation. Thus, the first blood-vessels
pass over the embryonic body and reach as far as the edge of the
germinative area. At first they are confined to the dark or "vascular"
area. But they afterwards extend over the whole surface of the
embryonic vesicle. In the end, the whole of the yelk-sac is covered
with a vascular net-work. These vessels have to gather food from the
contents of the yelk-sac and convey it to the embryonic body. This is
done by the veins, which pass first from the germinative area, and
afterwards from the yelk-sac, to the farther end of the heart. They
are called vitelline, or, frequently, omphalo-mesenteric, veins.

These vessels naturally atrophy with the degeneration of the umbilical
vesicle, and the vitelline circulation is replaced by a second, that
of the allantois. Large blood-vessels are developed in the wall of the
urinary sac or the allantois, as before, from the gut-fibre layer.
These vessels grow larger and larger, and are very closely connected
with the vessels that develop in the body of the embryo itself. Thus,
the secondary, allantoic circulation gradually takes the place of the
original vitelline circulation. When the allantois has attached itself
to the inner wall of the chorion and been converted into the placenta,
its blood-vessels alone effect the nourishment of the embryo. They are
called umbilical vessels, and are originally double--a pair of
umbilical arteries and a pair of umbilical veins. The two umbilical
veins (Figure 1.183 u), which convey blood from the placenta to the
heart, open it first into the united vitelline veins. The latter then
disappear, and the right umbilical vein goes with them, so that
henceforth a single large vein, the left umbilical vein, conducts all
the blood from the placenta to the heart of the embryo. The two
arteries of the allantois, or the umbilical arteries (Figures 1.183 n
and 1.184 n), are merely the ultimate terminations of the primitive
aortas, which are strongly developed afterwards. This umbilical
circulation is retained until the nine months of embryonic life are
over, and the human embryo enters into the world as the independent
individual. The umbilical cord (Figure 1.196 al), in which these large
blood-vessels pass from the embryo to the placenta, comes away,
together with the latter, in the after-birth, and with the use of the
lungs begins an entirely new form of circulation, which is confined to
the body of the infant.

(FIGURE 1.202. Boat-shaped embryo of the dog, from the ventral side,
magnified about ten times. In front under the forehead we can see the
first pair of gill-arches; underneath is the S-shaped heart, at the
sides of which are the auditory vesicles. The heart divides behind
into the two vitelline veins, which expand in the germinative area
(which is torn off all round). On the floor of the open belly lie,
between the protovertebrae, the primitive aortas, from which five
pairs of vitelline arteries are given off. (From Bischoff.))

There is a great phylogenetic significance in the perfect agreement
which we find between man and the anthropoid apes in these important
features of embryonic circulation, and the special construction of the
placenta and the umbilical cord. We must infer from it a close
blood-relationship of man and the anthropomorphic apes--a common
descent of them from one and the same extinct group of lower apes.
Huxley's "pithecometra-principle" applies to these ontogenetic
features as much as to any other morphological relations: "The
differences in construction of any part of the body are less between
man and the anthropoid apes than between the latter and the lower
apes."

This important Huxleian law, the chief consequence of which is "the
descent of man from the ape," has lately been confirmed in an
interesting and unexpected way from the side of the experimental
physiology of the blood. The experiments of Hans Friedenthal at Berlin
have shown that human blood, mixed with the blood of lower apes, has a
poisonous effect on the latter; the serum of the one destroys the
blood-cells of the other. But this does not happen when human blood is
mixed with that of the anthropoid ape. As we know from many other
experiments that the mixture of two different kinds of blood is only
possible without injury in the case of two closely related animals of
the same family, we have another proof of the close
blood-relationship, in the literal sense of the word, of man and the
anthropoid ape.

(FIGURE 1.203. Lar or white-handed gibbon (Hylobates lar or
albimanus), from the Indian mainland (From Brehm.)

FIGURE 1.204. Young orang (Satyrus orang), asleep.)

The existing anthropoid apes are only a small remnant of a large
family of eastern apes (or Catarrhinae), from which man was evolved
about the end of the Tertiary period. They fall into two geographical
groups--the Asiatic and the African anthropoids. In each group we can
distinguish two genera. The oldest of these four genera is the gibbon
Hylobates, Figure 1.203); there are from eight to twelve species of it
in the East Indies. I made observations of four of them during my
voyage in the East Indies (1901), and had a specimen of the ash-grey
gibbon (Hylobates leuciscus) living for several months in the garden
of my house in Java. I have described the interesting habits of this
ape (regarded by the Malays as the wild descendant of men who had lost
their way) in my Malayischen Reisebriefen (chapter 11).
Psychologically, he showed a good deal of resemblance to the children
of my Malay hosts, with whom he played and formed a very close
friendship.

(FIGURE 1.205. Wild orang (Dyssatyrus auritius). (From R. Fick and
Leutemann.))

The second, larger and stronger, genus of Asiatic anthropoid ape is
the orang (Satyrus); he is now found only in the islands of Borneo and
Sumatra. Selenka, who has published a very thorough Study of the
Development and Cranial Structure of the Anthropoid Apes (1899),
distinguishes ten races of the orang, which may, however, also be
regarded as "local varieties or species." They fall into two
sub-genera or genera: one group, Dissatyrus (orang-bentang, Figure
1.205), is distinguished for the strength of its limbs, and the
formation of very peculiar and salient cheek-pads in the elderly male;
these are wanting in the other group, the ordinary orang-outang
(Eusatyrus).

(FIGURE 1.206. The bald-headed chimpanzee (Anthropithecus calvus).
Female. This fresh species, described by Frank Beddard in 1897 as
Troglodytes calvus, differs considerably from the ordinary A. niger
Figure 1.207) in the structure of the head, the colouring, and the
absence of hair in parts.)

Several species have lately been distinguished in the two genera of
the black African anthropoid apes (chimpanzee and gorilla). In the
genus Anthropithecus (or Anthropopithecus, formerly Troglodytes), the
bald-headed chimpanzee, A. calvus (Figure 1.206), and the gorilla-like
A. mafuca differ very strikingly from the ordinary Anthropithecus
niger (Figure 1.207), not only in the size and proportion of many
parts of the body, but also in the peculiar shape of the head,
especially the ears and lips, and in the hair and colour. The
controversy that still continues as to whether these different forms
of chimpanzee and orang are "merely local varieties" or "true species"
is an idle one; as in all such disputes of classifiers there is an
utter absence of clear ideas as to what a species really is.

Of the largest and most famous of all the anthropoid apes, the
gorilla, Paschen has lately discovered a giant-form in the interior of
the Cameroons, which seems to differ from the ordinary species
(Gorilla gina Figure 1.208), not only by its unusual size and
strength, but also by a special formation of the skull. This giant
gorilla (Gorilla gigas, Figure 1.209) is six feet eight inches long;
the span of its great arms is about nine feet; its powerful chest is
twice as broad as that of a strong man.

(FIGURE 1.207. Female chimpanzee (Anthropithecus niger). (From Brehm.)

FIGURE 1.208. Female gorilla. (From Brehm.)

FIGURE 1.209. Male giant-gorilla (Gorilla gigas), from Yaunde, in the
interior of the Cameroons. killed by H. Paschen, stuffed by Umlauff.)

The whole structure of this huge anthropoid ape is not merely very
similar to that of man, but it is substantially the same. "The same
200 bones, arranged in the same way, form our internal skeleton; the
same 300 muscles effect our movements; the same hair covers our skin;
the same groups of ganglionic cells compose the ingenious mechanism of
our brain; the same four-chambered heart is the central pump of our
circulation." The really existing differences in the shape and size of
the various parts are explained by differences in their growth, due to
adaptation to different habits of life and unequal use of the various
organs. This of itself proves morphologically the descent of man from
the ape. We will return to the point in Chapter 2.23. But I wanted to
point already to this important solution of "the question of
questions," because that agreement in the formation of the embryonic
membranes and in foetal circulation which I have described affords a
particularly weighty proof of it. It is the more instructive as even
cenogenetic structures may in certain circumstances have a high
phylogenetic value. In conjunction with the other facts, it affords a
striking confirmation of our biogenetic law.





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