LAWS OF DEVELOPMENT OF ORGANIC TRIBES AND OF INDIVIDUALS. PHYLOGENY AND
ONTOGENY.
Laws of the Development of Mankind: Differentiation and
Perfecting.—Mechanical Cause of these two Fundamental
Laws.—Progress without Differentiation, and
Differentiation without Progress.—Origin of Rudimentary
Organs by Non-use and Discontinuance of
Habit.—Ontogenesis, or Individual Development of
Organisms.—Its General Importance.—Ontogeny, or the
Individual History of Development of Vertebrate Animals,
including Man.—The Fructification of the Egg.—Formation
of the three Germ Layers.—History of the Development of
the Central Nervous System, of the Extremities, of the
Branchial Arches, and of the Tail of Vertebrate
Animals.—Causal Connection and Parallelism of
Ontogenesis and Phylogenesis, that is of the Development
of Individuals and Tribes.—Causal Connection of the
Parallelism of Phylogenesis and of Systematic
Development.—Parallelism of the three Organic Series of
Development.
If man wishes to understand his position in nature, and to comprehend as
natural facts his relations to the phenomena of the world cognisable by
him, it is absolutely necessary that he should compare human with
extra-human phenomena, and, above all, with animal phenomena. We have
already seen that the exceedingly important physiological laws of
Inheritance and Adaptation apply to the human organism in the same
manner as to the animal and vegetable kingdoms, and in both cases
interact with one another. Consequently, natural selection in the
struggle for life acts so as to transform human society, just as it
modifies animals and plants, and in both cases constantly produces new
forms. The comparison of the phenomena of human and animal
transformation is especially interesting in connection with the laws of
divergence and progress, the two fundamental laws which, at the end of
the last chapter, we proved to be direct and necessary consequences of
natural selection in the struggle for life.
A comparative survey of the history of nations, or what is called
“universal history,” will readily yield to us, as the first and most
general result, evidence of a continually _increasing variety_ of human
activities, both in the life of individuals and in that of families and
states. This differentiation or separation, this constantly increasing
divergence of human character and the form of human life, is caused by
the ever advancing and more complete division of labour among
individuals. While the most ancient and lowest stages of human
civilization show us throughout the same rude and simple conditions, we
see in every succeeding period of history, among different nations, a
greater variety of customs, practices, and institutions. The increasing
division of labour necessitates an increasing variety of forms
corresponding to it. This is expressed even in the formation of the
human face. Among the lowest tribes of nations, most of the individuals
resemble one another so much that European travellers often cannot
distinguish them at all. With increasing civilization the physiognomy of
individuals becomes differentiated, and finally, among the most highly
civilized nations, the English and Germans, the divergence in the
characters of the face is so great that we very rarely mistake one face
for another.
The second great fundamental law which is obvious in the history of
nations is the great law of progress or perfecting. Taken as a whole,
the history of man is the history of his _progressive development_. It
is true that everywhere and at all times we may notice individual
retrogressions, or observe that crooked roads towards progress have been
taken, which lead only towards one-sided and external perfecting, and
thus deviate more and more from the higher goal of internal and enduring
perfecting. However, on the whole, the movement of development of all
mankind is and remains a progressive one, inasmuch as man continually
removes himself further from his ape-like ancestors, and continually
approaches nearer to his own ideal.
Now, if we wish to know what causes actually determine these two great
laws of development in man, namely, the law of divergence and the law of
progress, we must compare them with the corresponding laws of
development in animals, and on a close examination we shall inevitably
come to the conclusion that the phenomena, as well as their causes, are
exactly the same in the two cases. The course of development in man,
just as in that of animals, being directed by the two fundamental laws
of differentiation and perfecting, is determined solely by purely
mechanical causes, and is solely the necessary consequence of natural
selection in the struggle for life.
Perhaps in the preceding discussion the question has presented itself to
some—“Are not these two laws identical? Is not progress in all cases
necessarily connected with divergence?” This question has often been
answered in the affirmative, and Carl Ernst Bär, for example, one of the
greatest investigators in the domain of the history of development, has
set forth the following proposition as one of the principal laws in the
ontogenesis of the animal body:—“The degree of development (or
perfecting) depends on the stage of separation (or differentiation) of
the parts.”(20) Correct as this proposition may be on the whole, yet it
is not universally true. In many individual cases it can be proved that
divergence and progress by no means always coincide. _Every progress is
not a differentiation, and every differentiation is not a progress._
Naturalists, guided by purely anatomical considerations, had already set
forth the law relating to progress in organization, that the perfecting
of an organism certainly depends, for the most part, upon the division
of labour among the individual organs and parts of the body, but that
there are also other organic transformations which determine a progress
in organization. One, in particular, which has been generally
recognized, is the _numerical diminution of identical parts_. If, for
example, we compare the lower articulated animals of the crustacean
group, which possess numerous pairs of legs, with spiders which never
have more than four pairs of legs, and with insects which always possess
only three pairs of legs, we find this law, for which a great number of
examples could be adduced, confirmed. The numerical diminution of pairs
of legs is a progress in the organization of articulated animals. In
like manner the numerical diminution of corresponding vertebral joints
in the trunk of vertebrate animals is a progress in their organization.
Fishes and amphibious animals with a very large number of identical
vertebral joints are, for this very reason, less perfect and lower than
birds and mammals, in which the vertebral joints, as a whole, are not
only very much more differentiated, but in which the number of
corresponding vertebræ is also much smaller. Further, according to the
same law of numerical diminution, flowers with numerous stamens are more
imperfect than the flowers of kindred plants with a smaller number of
stamens, etc. If therefore originally a great number of homogeneous
parts exist in an organic body, and if, in the course of very many
generations, this number be gradually decreased, this transformation
will be an example of perfecting.
Another law of progress, which is quite independent of differentiation,
nay, even appears to a certain extent opposed to it, is the law of
_centralization_. In general the whole organism is the more perfect the
more it is organized as a unit, the more the parts are subordinate to
the whole, and the more the functions and their organs are centralized.
Thus, for example, the system of blood-vessels is most perfect where a
centralized heart exists. In like manner, the dense mass of marrow which
forms the spinal cord of vertebrate animals, and the ventral cord of the
higher articulated animals, is more perfect than the decentralized chain
of ganglia of the lower articulated animals, and the scattered system of
ganglia in the molluscs. Considering the difficulty of explaining these
complicated laws of progress in detail, I cannot here enter upon a
closer discussion of them, and must refer to Bronn’s excellent
“Morphologischen Studien,” and to my “General Morphology” (Gen. Morph.
i. 370, 550; ii. 257-266).
Just as we have become acquainted with phenomena of progress, quite
independent of divergence, so we shall, on the other hand, very often
meet with divergencies which are not perfecting, but which are rather
the contrary, that is retrogressions or degenerations. It is easy to see
that the changes which every species of animal and plant experiences
cannot always be improvements. But rather many phenomena of
differentiation, which are of direct advantage to the organism itself,
are yet, in a wider sense, detrimental, inasmuch as they lessen its
general capabilities. Frequently a relapse to simpler conditions of life
takes place, and by adaptation to them a divergence in a retrograde
direction. If, for instance, organisms which have hitherto lived
independently accustom themselves to a parasitical life, they thereby
degenerate or retrograde. Such animals, which hitherto had possessed a
well-developed nervous system and quick organs of sense, as well as the
power of moving freely, lose these when they accustom themselves to a
parasitical mode of life; they consequently retrograde more or less.
There the differentiation viewed by itself is a degeneration, although
it is advantageous to the parasitical organism. In the struggle for life
such an animal, which has accustomed itself to live at the expense of
others, by retaining its eyes and apparatus of motion, which are of no
more use to it, would only expend so much material uselessly; and when
it loses these organs, then a great quantity of nourishment which was
employed for the maintenance of these parts, benefits other parts. In
the struggle for life between the different parasites, therefore, those
which make least pretensions will have advantage over the others, and
this favours their degeneration.
Just as this is found to be the case with the whole organism, so it is
also with the parts of the body of an individual organism. A
differentiation of parts, which leads to a partial degeneration, and
finally even to the loss of individual organs, is, when looked at by
itself, a degeneration, but yet may be advantageous to the organism in
the struggle for life. It is easier to fight when useless baggage is
thrown aside. Hence we meet everywhere, in the more highly-developed
animal and vegetable bodies, processes of divergence, the essence of
which is that they cause the degeneration, and finally the loss, of
particular parts. And at this point the most important and instructive
of all the series of phenomena bearing upon the history of organisms
presents itself to us, namely, that of _rudimentary or degenerate
organs_.
It will be remembered that even in my first chapter I considered this
exceedingly remarkable series of phenomena, from a theoretical point of
view, as one of the most important and most striking proofs of the truth
of the doctrine of descent. We designated as rudimentary organs those
parts of the body which are arranged for a definite purpose and yet are
without function. Let me remind the reader of the eyes of those animals
which live in the dark in caves and underground, and which consequently
never can use them. In these animals we find real eyes hidden under the
skin, frequently developed exactly as are the eyes of animals which
really see; and yet these eyes never perform any function, indeed
cannot, simply for the reason that they are covered by an opaque
membrane, and consequently no ray of light falls upon them (compare
above, p. 13). In the ancestors of these animals, which lived in open
daylight, the eyes were well developed, covered by a transparent horny
capsule (cornea), and actually served the purpose of seeing. But as the
animals gradually accustomed themselves to an underground mode of life,
and withdrew from the daylight and no longer used their eyes, these
became degenerated.
Very clear examples of rudimentary organs, moreover, are the wings of
animals which cannot fly; for example, the wings of the running birds,
like the ostrich, emeu, cassowary, etc., the legs of which have become
exceedingly developed. These birds having lost the habit of flying, have
consequently lost the use of their wings; however, the wings are still
there, although in a crippled form. We very frequently find such
crippled wings in the class of insects, most members of which can fly.
From reasons derived from comparative anatomy and other circumstances,
we can with certainty draw the inference that all insects now living
(all dragon-flies, grasshoppers, beetles, bees, bugs, flies,
butterflies, etc.) have originated from a single common parental form,
from a primary insect which possessed two well-developed pairs of wings,
and three pairs of legs. Yet there are very many insects in which either
one or both pairs of wings have become more or less degenerated, and
many in which they have even completely disappeared. For example, in the
whole order of flies, or Diptera, the hinder pair of wings—in the
bee-parasites, or Strepsiptera, on the other hand, the fore pair of
wings—have become degenerated or entirely disappeared. Moreover, in
every order of insects we find individual genera, or species, in which
the wings have more or less degenerated or disappeared. The latter is
the case especially in parasites. The females have frequently no wings,
whereas the males have; for instance, in the case of glow-worms
(Lampyris), Strepsiptera, etc. This partial or complete degeneration of
the wings of insects has evidently arisen from natural selection in the
struggle for life. For we find insects without wings living under
circumstances where flying would be useless, or even decidedly injurious
to them. If, for example, insects living on islands fly about much, it
may easily happen that when flying they are blown into the sea by the
wind, and if (as is always the case) the power of flying is differently
developed in different individuals, then those which fly badly have an
advantage over those which fly well; they are less easily blown into the
sea, and remain longer in life than the individuals of the same species
which fly well. In the course of many generations, by the action of
natural selection, this circumstance must necessarily lead to a complete
suppression of the wings. If this conclusion had been arrived at on
purely theoretical grounds, we might be pleased to find its truth
established by facts. For upon isolated islands the proportion of
wingless insects to those possessing wings is surprisingly large, much
larger than among the insects inhabiting continents. Thus, for example,
according to Wollaston, of the 550 species of beetles which inhabit the
island of Madeira, 220 are wingless, or possess such imperfect wings
that they can no longer fly; and of the 29 genera which belong to that
island exclusively, no less than 23 contain such species only. It is
evident that this remarkable circumstance does not need to be explained
by the special wisdom of the Creator, but is sufficiently accounted for
by natural selection, because in this case the hereditary disuse of the
wings, the discontinuance of flying in the presence of dangerous winds,
has been very advantageous in the struggle for life. In other wingless
insects the want of wings has been advantageous for other reasons.
Viewed by itself, the loss of wings is a degeneration, but in these
special conditions of life it is advantageous to the organism in the
struggle for life.
Among other rudimentary organs I may here, by way of example, further
mention the lungs of serpents and serpent-like lizards. All vertebrate
animals possessing lungs, such as amphibious animals, reptiles, birds,
and mammals, have a pair of lungs, a right and a left one. But in cases
where the body is exceedingly thin and elongated, as in serpents and
serpent-like lizards, there is no room for the one lung by the side of
the other, and it is an evident advantage to the mechanism of
respiration if only one lung is developed. A single large lung here
accomplishes more than two small ones side by side would do; and
consequently, in these animals, we invariably find only the right or
only the left lung fully developed. The other is completely aborted,
although existing as a useless rudiment. In like manner, in all birds
the right ovary is aborted and without function; only the left one is
developed, and yields all the eggs.
I mentioned in the first chapter that man also possesses such useless
and superfluous rudimentary organs, and I specified as such the muscles
which move the ears. Another of them is the rudiment of the tail which
man possesses in his 3—5 tail vertebræ, and which, in the human embryo,
stands out prominently during the first two months of its development
(compare Plates II. and III.). It afterwards becomes completely hidden.
The rudimentary little tail of man is an irrefutable proof of the fact
that he is descended from tailed ancestors. In woman the tail is
generally by one vertebra longer than in man. There still exist
rudimentary muscles in the human tail which formerly moved it.
Another case of human rudimentary organs, only belonging to the male,
and which obtains in like manner in all male mammals, is furnished by
the mammary glands on the breast, which, as a rule, are active only in
the female sex. However, cases of different mammals are known,
especially of men, sheep, and goats, in which the mammary glands were
fully developed in the male sex, and yielded milk as food for their
offspring. I have already mentioned before (p. 12) that the rudimentary
auricular muscles in man can still be employed to move their ears, by
some persons who have perseveringly practised them. In fact, rudimentary
organs are frequently very differently developed in different
individuals of the same species; in some they are tolerably large, in
others very small. This circumstance is very important for their
explanation, as is also the other circumstance that generally in
embryos, or in a very early period of life, they are much larger and
stronger in proportion to the rest of the body than they are in fully
developed and fully grown organisms. This can, in particular, be easily
pointed out in the rudimentary sexual organs of plants (stamens and
pistil), which I have already mentioned. They are proportionately much
larger in the young flower-bud than in the mature flower.
I have remarked (p. 15) that rudimentary or suppressed organs were the
strongest supports of the monistic or mechanical conception of the
universe. If its opponents, the dualists and teleologists, understood
the immense significance of rudimentary organs, it would put them into a
state of despair. Their ludicrous attempts to explain that rudimentary
organs were given to organisms by the Creator “for the sake of
symmetry,” or “as a formal provision,” or “in consideration of his
general plan of creation,” sufficiently prove the utter impotence of
their perverse conception of the universe. I must here repeat that, even
if we knew absolutely nothing of the other phenomena of development, we
should be obliged to believe in the truth of the Theory of Descent,
solely on the ground of the existence of rudimentary organs. Not one of
its opponents has been able to throw even a feeble glimmer of an
acceptable explanation upon these exceedingly remarkable and important
phenomena. There is scarcely any highly developed animal or vegetable
form which has not some rudimentary organs, and in most cases it can be
shown that they are the products of natural selection, and that they
have become suppressed by disuse. It is the reverse of the process of
formation in which new organs arise from adaptation to certain
conditions of life, and by the use of parts as yet incompletely
developed. It is true our opponents usually maintain that the origin of
altogether new parts is completely inexplicable by the Theory of
Descent. However, I distinctly assert that to those who possess a
knowledge of comparative anatomy and physiology this matter does not
present the slightest difficulty. Every one who is familiar with
comparative anatomy and the history of development will find as little
difficulty about the origin of completely new organs as about the utter
disappearance of rudimentary organs. The disappearance of the latter,
viewed by itself, is the converse of the origin of the former. Both
processes are particular phenomena of differentiation, which, like all
others, can be explained quite simply and mechanically by the action of
natural selection in the struggle for life.
The infinitely important study of rudimentary organs and their origin,
the comparison of their palæontological and embryological development,
now naturally leads us to the consideration of one of the most important
and instructive of all biological phenomena, namely, the parallelism
which the phenomena of progress and divergence present to us in three
different series. When, in the last chapter, we spoke of perfecting and
division of labour, we understood by those words progress and
separation, and those changes effected by them, which in the long and
slow course of the earth’s history have led to a continual variation of
the flora and fauna, to the origin of new and to the disappearance of
ancient species of animals and plants. Now, if we follow the origin, the
development, and the life of every single organic individual, we meet
with exactly the same phenomena of progress and differentiation. The
individual development, or the _ontogenesis_ of every single organism,
from the egg to the complete form is nothing but a growth attended by a
series of diverging and progressive changes. This applies equally to
animals, plants, and protista. If, for example, we consider the ontogeny
of any mammal, of man, of an ape, or of a pouched animal, or if we
follow the individual development of any other vertebrate animal of
another class, we everywhere find essentially the same phenomena. Every
one of these animals develops itself originally out of a single cell,
the egg. This cell increases by self-division, and forms a number of
cells, and by the growth of this accumulation of cells, by the divergent
development of originally identical cells, by the division of labour
among them, and by their perfecting, there arises the perfect organism,
the complicated composition of which excites our admiration.
It seems to me here indispensable to draw attention more closely to
those infinitely important and interesting processes which accompany
_ontogenesis, or the individual development of organisms_, and
especially to that of vertebrate animals, man included. I wish
especially to recommend these exceedingly remarkable and instructive
phenomena to the reader’s most careful consideration, first, because
they are among the strongest supports of the Theory of Descent, and
secondly, because, considering their immense general importance, they
have hitherto been properly considered only by a few privileged persons.
We cannot indeed but be astonished when we consider the deep ignorance
which still prevails, in the widest circles, about the facts of the
individual development of man and organisms in general. These facts, the
universal importance of which cannot be estimated too highly, were
established, in their most important outlines, even more than a hundred
years ago, in 1759, by the great German naturalist Caspar Friedriech
Wolff, in his classical “Theoria Generationis.” But, just as Lamarck’s
Theory of Descent, founded in 1809, lay dormant for half a century, and
was only awakened to new and imperishable life in 1859, by Darwin, in
like manner Wolff’s Theory of Epigenesis remained unknown for nearly
half a century; and it was only after Oken, in 1806 had published his
history of the development of the intestinal tube, and after Meckel, in
1812, had translated Wolff’s work (written in Latin) on the same subject
into German, that Wolff’s theory of epigenesis became more generally
known, and formed the foundation of all subsequent investigations of the
history of individual development. The study of ontogenesis now received
a great stimulus, and soon there appeared the classical investigations
of the two friends, Christian Pander (1817) and Carl Ernst Bär (1819).
Bär, in his remarkable “Entwickelungsgeschichte der Thiere,”(20) worked
out the ontogeny of vertebrate animals in all its important facts. He
carried out a series of such excellent observations, and illustrated
them by such profound philosophical reflections, that his work became
the foundation for a thorough understanding of this important group of
animals, to which, of course, man also belongs. The facts of embryology
alone would be sufficient to solve the question of man’s position in
nature, which is the highest of all problems. Look attentively at and
compare the eight figures which are represented on the adjoining Plates
II. and III., and it will be seen that the philosophical importance of
embryology cannot be too highly estimated.
We may well ask, What do our so-called “educated” circles, who think so
much of the high civilization of the 19th century, know of these most
important biological facts, of these indispensable foundations for
understanding their own organism? How much do our speculative
philosophers and theologians know about them, who fancy they can arrive
at an understanding of the human organism by mere guesswork or divine
inspiration? What indeed do the majority of naturalists, not excepting
the majority of the so-called “zoologists” (including the
entomologists!), know about them?
The answer to this question tells much to the shame of the persons above
indicated, and we must confess, willingly or unwillingly, that these
invaluable facts of human ontogeny are, even at the present day, utterly
unknown to most people, or are in no way valued as they deserve to be.
It is in the face of such a condition of things as this that we see
clearly upon what a wrong and one-sided road the much vaunted culture of
the 19th century still moves. Ignorance and superstition are the
foundations upon which most men construct their conception of their own
organism and its relation to the totality of things; and these palpable
facts of the history of development, which might throw the light of
truth upon them, are ignored. It is true these facts are not calculated
to excite approval among those who assume a thorough difference between
man and the rest of nature, and who will not acknowledge the animal
origin of the human race. That origin must be a very unpleasant truth to
members of the ruling and privileged castes in those nations among which
there exists an hereditary division of social classes, in consequence of
false ideas about the laws of inheritance. It is well known that, even
in our day, in many civilized countries the idea of hereditary grades of
rank goes so far, that, for example, the aristocracy imagine themselves
to be of a nature totally different from that of ordinary citizens, and
nobles who commit a disgraceful offence are punished by being expelled
from the caste of nobles, and thrust down among the pariahs of “vulgar
citizens.” What are these nobles to think of the noble blood which flows
in their privileged veins, when they learn that all human embryos, those
of nobles as well as commoners, during the first two months of
development, are scarcely distinguishable from the tailed embryos of
dogs and other mammals?
As the object of these pages is solely to further the general knowledge
of natural truths, and to spread, in wider circles, a natural conception
of the relations of man to the rest of nature, I shall be justified if I
do not pay any regard to the widely-spread prejudice in favour of an
exceptional and privileged position for man in creation, and simply give
here the embryological facts from which the reader will be able to draw
conclusions affirming the groundlessness of those prejudices. I wish all
the more to entreat him to reflect carefully upon these facts of
ontogeny, as it is my firm conviction that a general knowledge of them
can only promote the intellectual advance, and thereby the mental
perfecting, of the human race.
Amidst all the infinitely rich and interesting material which lies
before us in the ontogeny of vertebrate animals, that is, in the history
of their individual development, I shall here confine myself to showing
some of those facts which are of the greatest importance to the Theory
of Descent in general, as well as in its special application to man. Man
is at the beginning of his individual existence a simple egg, a single
little cell, just the same as every animal organism which originates by
sexual generation. The human egg is essentially the same as that of all
other mammals, and cannot be distinguished from the egg of the higher
mammals. The egg represented in Fig. 5 might be that of a man or an ape
as well as of a dog, a horse, or any other mammal. Not only the form and
structure, but even the size of the egg in most mammals is the same as
in man, namely, about the 120th part of an inch in diameter, so that the
egg under favorable circumstances, with the naked eye, can just be
perceived as a small speck. The differences which really exist between
the eggs of different mammals and that of man do not consist in the
form, but in the chemical mixture, in the molecular composition of the
albuminous combination of carbon, of which the egg essentially consists.
These minute individual differences of all eggs, which depend upon
indirect or potential adaptation (and especially upon the law of
individual adaptation), are indeed not directly perceptible to the
exceedingly imperfect senses of man, but are cognisable through indirect
means, as the primary causes of the difference of all individuals.
[Illustration: FIG. 5.—The human egg a hundred times enlarged. _a._ The
kernel speck, or nucleolus (the so-called germinal spot of the egg).
_b._ Kernel, or nucleus (the so-called germinal vesicle of the egg).
_c._ Cell-substance, or protoplasm (so-called yolk of the egg). _d._
Cell-membrane (the yolk-membrane of the egg; in mammals, on account of
its transparency, called zona pellucida). The eggs of other mammals are
of the same form.]
The human egg is, like that of all other mammals, a small globular
bladder, which contains all the constituent parts of a simple organic
cell (Fig. 5). The most essential parts of it are the mucous
cell-substance, or the protoplasma (_c_), which in an egg is called the
“yolk,” and the cell-kernel, or nucleus (_b_), surrounded by it, which
is here called by the special name of the “germinal vesicle.” The latter
is a delicate, clear, glassy globule of albumen, of about 1-600th part
of an inch in diameter, and surrounds, a still smaller, sharply-marked,
rounded granule (_a_), the kernel-speck, or the nucleolus of the cell
(in the egg it is called the “germinal spot”). The outside of the
globular egg-cell of a mammal is surrounded by a thick pellucid
membrane, the cell-membrane or yolk-membrane, which here bears the
special name of zona pellucida (_d_). The eggs of many lower animals
(for example of many Medusæ) differ from this in being _naked_ cells, as
the outer covering, or cell-membrane, is wanting.
As soon as the egg (ovulum) of the mammal has attained its full
maturity, it leaves the ovary of the female, in which it originates, and
passes into the oviduct, and through this narrow passage into the wider
pouch or womb (uterus). If, meanwhile, the egg is fructified by the male
seed (sperm), it develops itself in this pouch into an embryo, and does
not leave it until perfectly developed and capable of coming into the
world at birth as a young mammal.
The variations of form and transformations which the fructified egg must
go through within the uterus before it assumes the form of the mammal
are exceedingly remarkable, and proceed from the beginning in man, in
precisely the same way as in the other mammals. At first the fructified
egg of the mammal acts as a single-celled organism, which is about to
propagate independently and increase itself; for example, an Amœba
(compare Fig. 2, p. 188). In point of fact the simple egg-cell becomes
two, by the process of cell-division which I have previously described.
There arise from the single germinal spot (the small kernel-speck of the
original simple egg-cell) two new kernel-specks, and then in like
manner, out of the germinal vesicle (the nucleus), two new cell-kernels.
Then, and not until then, does the globular protoplasma first separate
itself by an equatorial furrow into two halves, in such a manner that
each half encloses one of the two kernels, together with its
kernel-speck. Thus the simple egg-cell, within the original cellular
membrane, has become two naked cells, each possessing its own kernel
(Fig. 6).
[Illustration: FIG. 6.—First commencement of the development of a
mammal’s egg, the so-called “yolk-cleavage” (propagation of the egg-cell
by repeated self-division). _A._ The egg, by the formation of the first
furrow, falls into two cells. _B._ These by division fall into four
cells. _C._ These latter have fallen into eight cells. _D._ By continued
division a globular mass of numerous cells has arisen.]
The same process of cell-division now repeats itself several times in
succession. In this way, from two cells (Fig. 6 _A_) there arise four
(Fig. 6 _B_); from four, eight (Fig. 6 _C_); from eight, sixteen; from
these, thirty-two, etc. Each time the division of the kernel-speck
precedes that of the kernel; this, again, precedes that of the
cell-substance, or protoplasma. As the division of the latter always
commences with the formation of a superficial annular _furrow_, or
cleft, the whole process is usually called the _furrowing of the egg_,
or yolk-cleavage, and the products of it, that is, the cells arising
from the continued halving, are called the _cleavage spheres_. However,
the whole process is nothing more than a simple, oft-repeated _division
of cells_, and the products of it are actual, naked _cells_. Finally,
through the continued division or “furrowing” of the mammal’s egg there
arises a mulberry-shaped ball, which is composed of a great number of
small spheres, naked cells, containing kernels (Fig. 6 _D_). These cells
are the materials out of which the body of the young mammal is
constructed. Every one of us has once been such a simple mulberry-shaped
ball, composed only of small equi-formal cells.
The further development of the globular lump of cells, which now
represents the young body of the mammal, consists first in its changing
into a globular bladder, as fluid accumulates within it. This bladder is
called the germ-bladder (vesicula blastodermica). Its wall is at first
composed of merely equi-formal cells. But soon, at one point on the
wall, arises a disc-shaped thickening, as the cells here increase
rapidly, and this thickening is now the foundation of the actual body of
the germ or embryo, while the other parts of the germ-bladder serve only
for its nutrition. The thickened disc, or foundation of the embryo, soon
assumes an oblong, and then a fiddle-shaped form, in consequence of its
right and left walls becoming convex (Fig. 7, p. 304). At this stage of
development in the first form of their germ or embryo, not only all
mammals, including man, but even all vertebrate animals in
general—birds, reptiles, amphibious animals, and fishes—can either not
be distinguished from one another at all, or only by very unessential
differences, such as the arrangement of the egg-coverings. In all the
whole body consists of nothing but a quite simple, oblong, oval, or
violin-shaped thin disc, which is composed of three closely connected
membranes or plates, lying one above another. Each of the three plates
or layers of the germ consists simply of cells all exactly like one
another; but each layer has a different function in the building up of
the vertebrate animal body. Out of the upper or outer germ-layer arises
solely the outer skin (epidermis), together with the central parts of
the nervous system (spinal marrow and brain); out of the lower or inner
layer arises only the inner delicate skin (epithelium) which lines the
whole intestinal tube from the mouth to the anus, together with all the
glands connected with it (lung, liver, salivary glands, etc.); out of
the middle germ-layer lying between the two others arise all the other
organs, muscles, bones, blood-vessels. Now, the processes by which the
various and exceedingly complicated parts of the fully-formed body of
vertebrate animals arise out of such simple material—out of the three
germ-layers composed only of cells—are, in the first place, the
repeated division, and consequently the increase of cells; in the second
place, the division of labour or differentiation of these cells; and
thirdly, the union of the variously developed or differentiated cells,
for the formation of the different organs. Thus arises the gradual
progress or perfecting which can be traced step by step in the
development of the embryonic body. The simple embryonic cells, which are
to constitute the body of the vertebrate animal, stand in the same
relation to each other as citizens who wish to found a state. Some take
to one occupation, others to another, and work together for the good of
the whole. By this division of labour, or differentiation, and the
perfecting (the organic progress) which is connected with it, it becomes
possible for the whole state to accomplish undertakings which would have
been impossible to the single individual. The whole body of the
vertebrate animal, like every other many-celled organism, is a
republican state of cells, and consequently it can accomplish organic
functions which the individual cell, as a solitary individual (for
example, an Amœba, or a single-celled plant), could never perform.
No sensible person supposes that carefully devised institutions, which
have been established for the good of the whole, as well as for the
individual, in every human state, are the results of the action of a
personal and supernatural Creator, acting for a definite purpose. On the
contrary, every one knows that these useful institutions of organization
in the state are the consequences of the co-operation of the individual
citizens and their common government, as well as of adaptation to the
conditions of existence of the outer world. Just in the same way we must
judge of the many-celled organism. In it also all the useful
arrangements are solely the natural and necessary result of the
co-operation, differentiation, and perfecting of the individual
citizens—the cells—and by no means the artificial arrangements of a
Creator acting for a definite purpose. If we rightly consider this
comparison, and pursue it further, we can distinctly see the perversity
of that dualistic conception of nature which discovers the action of a
creative plan of construction in the various adaptations of the
organization of living things.
Let us pursue the individual development of the vertebrate animal body a
few stages further, and see what is next done by the citizens of this
embryonic organism. In the central line of the violin-shaped disc, which
is composed of the three cellular germ-layers, there arises a straight
delicate furrow, the so-called “primitive streak,” by which the
violin-shaped body is divided into two equal lateral halves—a right and
a left part or “antimer.” On both sides of that streak or furrow, the
upper or external germ-layer rises in the form of a longitudinal fold,
and both folds then grow together over the furrow in the central line,
and thus form a cylindrical tube. This tube is called the marrow-tube,
or medullary canal, because it is the foundation of the central nervous
system, the _spinal marrow_ (medulla spinalis). At first it is pointed
both in front and behind, and it remains so for life in the lowest
vertebrate animal, the brainless, skull-less Lancelet (Amphioxus). But
in all other vertebrate animals, which we distinguish from the latter as
skulled animals, or Craniota, a difference between the fore and hinder
end of the marrow tube soon becomes visible, the fore end becoming
dilated, and changing into a roundish bladder, the foundation of the
_brain_.
In all Craniota, that is, in all vertebrate animals possessing skull and
brain, the brain, which is at first only the bladder-shaped dilatation
of the anterior end of the spinal marrow, divides into five bladders
lying one behind the other, four superficial, transverse in-nippings
being formed. These _five brain-bladders_, out of which afterwards arise
all the different parts of the intricately constructed brain, can be
seen in their original condition in the embryo represented in Fig. 7. It
is just the same whether we examine the embryo of a dog, a fowl, a
lizard, or any other higher vertebrate animal. For the embryos of the
different skulled animals (at least the three higher classes of them,
the reptiles, birds and mammals) cannot be in any way distinguished at
the stage represented in Fig. 7. The whole form of the body is as yet
exceedingly simple, being merely a thin, leaf-like disc. Face, legs,
intestines, etc., are as yet completely wanting. But the five bladders
are already quite distinct from one another.
[Illustration: FIG. 7.—Embryo of a mammal or bird, in which the five
brain-bladders have just commenced to develop. _v._ Fore brain. _z._
Twixt brain. _m._ Mid brain. _h._ Hind brain. _n._ After brain. _p._
Spinal-marrow. _a._ Eye-bladders. _w._ Primitive vertebræ. _d._
Spinal-axis or notochord.]
The _first_ bladder, the _fore brain_ (_a_), is in so far the most
important that it principally forms the hemispheres of the so-called
larger brain (cerebrum), that part which is the seat of the higher
mental activities. The more these activities are developed in the series
of vertebrate animals, the more do the two lateral halves of the fore
brain, or the hemispheres, grow at the expense of the other bladders,
and overlap them in front and from above. In man, where they are most
strongly developed, agreeing with his higher mental activity, they
eventually almost entirely cover the other parts from above (compare
Plates II. and III.) The _second_ bladder, the _twixt brain_ (_z_),
forms that portion of the brain which is called the _centre of sight_,
and stands in the closest relation to the eyes (_a_), which grow right
and left out of the fore brain in the shape of two bladders, and later
lie at the bottom of the twixt brain. The _third_ bladder, the _mid
brain_ (_m_), for the most part vanishes in the formation of the
so-called _four bulbs_, a bossy portion of the brain, which is strongly
developed in reptiles and birds (Fig. _E_, _F_, Plate II.), whereas in
mammals it recedes much more (Fig. _G_, _H_, Plate III.). The _fourth_
bladder, the _hind brain_ (_h_), forms the so-called _little
hemispheres_, together with the middle part of the _small brain_
(cerebellum), a part of the brain as to the function of which the most
contradictory conjectures are entertained, but which seems principally
to regulate the co-ordination of movements. Lastly, the _fifth_ bladder,
the _after brain_ (_n_), develops into that very important part of the
central nervous system which is called the _prolonged marrow_ (medulla
oblongata). It is the central organ of the respiratory movements, and of
other important functions, and an injury to it immediately causes death,
whereas the large hemispheres of the fore brain (or the organ of the
“soul,” in a restricted sense) can be removed bit by bit, and even
completely destroyed, without causing the death of the vertebrate
animal—only its higher mental activities disappearing in consequence.
These five brain bladders, in all vertebrate animals which possess a
brain at all, are originally arranged in the same manner and develop
gradually in the different groups so differently, that it is afterwards
very difficult to recognize the corresponding parts in the
fully-developed brains. In the early stage of development which is
represented in Fig. 7, it seems as yet quite impossible to distinguish
the embryos of the different mammals, birds, and reptiles, from one
another. But if we compare the much more developed embryos on Plates II.
and III. with one another, we can clearly see an inequality in their
development, and especially it will be perceived that the brain of the
two mammals (_G_ and _H_) already strongly differ from that of birds
(_F_) and of reptiles (_E_). In the two latter the mid brain
predominates, but in the former the fore brain. Even at this stage the
brain of the bird (_F_) is scarcely distinguishable from that of the
tortoise (_E_), and in like manner the brain of the dog (_G_) is as yet
almost the same as that of man (_H_). If, on the other hand, we compare
the brains of these four vertebrate animals in a fully developed
condition, we find them so very different in all anatomical particulars,
that we cannot doubt for a moment as to which animal each brain belongs.
I have here explained the original equality, the gradual commencement,
and the ever increasing separation or differentiation of the embryos in
the different vertebrate animals, taking the brain as a special example,
just because this organ of the soul’s activity is of special interest.
But I might as well have discussed in its stead the heart, or the liver,
or the limbs, in short, any other part of the body, since the same
wonder of creation is here ever repeated, namely, this, that all parts
are originally the same in the different vertebrate animals, and that
the variations by which the different classes, orders, families, genera,
etc., differ and deviate from one another, are only gradually developed.
There are certainly few parts of the body which are so differently
constructed as the _limbs or extremities_ of the vertebrate animals.
Now, I wish the reader to compare in Fig. _A-H_ on Plates II. and III.,
the four extremities (_bv_) of the embryos with one another, and he will
scarcely be able to perceive any important differences between the human
arm (_H bv_), the wing of a bird (_F bv_), the slim foreleg of a dog (_G
bv_), and the plump foreleg of the tortoise (_E bv_). In comparing the
hinder extremities (_bh_) in these figures he will find it equally
difficult to distinguish the leg of a man (_H bh_), of a bird (_F bh_),
the hind-leg of a dog (_G bh_), and that of a tortoise (_E bh_). The
fore as well as the hinder extremities are as yet short, broad
lumps, at the ends of which the foundations of the five toes are placed,
connected as yet by a membrane. At a still earlier stage (Fig. _A-D_)
the five toes are not marked out at all, and it is quite impossible to
distinguish even the fore and hinder extremities from one another. The
latter, as well as the former, are nothing but simple roundish
processes, which have grown out of the side of the trunk. At the very
early stage represented in Fig. 7 they are completely wanting, and the
whole embryo is a simple trunk without a trace of limbs.
[Illustration: Pl. II. Germs or Embryos of four Vertebrates.
_v._ Fore-brain. _z._ Twixt-brain. _m._ Mid-brain. _h._ Hind-brain. _n._
After-brain. _w._ Spine. _r._ Spinal-cord.]
[Illustration: Pl. III. Germs or Embryos of four Vertebrates.
_na._ Nose. _a._ Eyes. _o._ Ear. _k{1} k{2} k{3}._ Gill-arches.
_s._ Tail. _bv._ Fore-leg. _bh._ Hind-leg.]
I wish especially to draw attention in Plates II. and III., which
represents embryos in early stages of development (Fig. _A-D_)—and in
which we are not able to recognize a trace of the full-grown animal—to
an exceedingly important formation, which originally is common to all
vertebrate animals, but which at a later period is transformed into the
most different organs. Every one surely knows the _gill-arches_ of fish,
those arched bones which lie behind one another, to the number of three
or four, on each side of the neck, and which support the gills, the
respiratory organs of the fish (double rows of red leaves, which are
popularly called “fishes’ ears.”) Now, these gill-arches originally
exist exactly the same in man (_D_), in dogs (_C_), in fowls (_B_), and
in tortoises (_A_), as well as in all other vertebrate animals. (In Fig.
_A-D_ the three gill-arches of the right side of the neck are marked
_k{1} k{2} k{3}_). Now, it is only in fishes that these remain in
their original form, and develop into respiratory organs. In the other
vertebrate animals they are partly employed in the formation of the face
(especially the jaw apparatus), and partly in the formation of the organ
of hearing.
Finally, when comparing the embryos on Plates II. and III., we must not
fail to give attention again to the _human tail_ (_s_), an organ which,
in the original condition, man shares with all other vertebrate animals.
The discovery of tailed men was long anxiously expected by many monistic
philosophers, in order to establish a closer relationship between man
and the other mammals. And in like manner their dualistic opponents
often maintained with pride that the complete want of a tail formed one
of the most important bodily distinctions between men and animals,
though they did not bear in mind the many tailless animals which really
exist. Now, man in the first months of development possesses a real tail
as well as his nearest kindred, the tailless apes (orang-outang,
chimpanzee, gorilla), and vertebrate animals in general. But whereas, in
most of them—for example, the dog (_C_, _G_)—in the course of
development it always grows longer, in man (Fig. _D_, _H_) and in
tailless mammals, at a certain period of development, it degenerates and
finally completely disappears. However, even in fully developed men, the
remnant of the tail is seen in the three, four, or five tail vertebræ
(vertebræ coccygeæ) as an aborted or rudimentary organ, which forms the
hinder or lower end of the vertebral column (p. 289).
Most persons even now refuse to acknowledge the most important deduction
of the Theory of Descent, that is, the palæontological development of
man from ape-like, and through them from still lower, mammals, and
consider such a transformation of organic form as impossible. But, I
ask, are the phenomena of the individual development of man, the
fundamental features of which I have here given, in any way less
wonderful? Is it not in the highest degree remarkable that all
vertebrate animals of the most different classes—fishes, amphibious
animals, reptiles, birds, and mammals—in the first periods of their
embryonic development cannot be distinguished at all, and even much
later, at a time when reptiles and birds are already distinctly
different from mammals, that the dog and the man are almost identical?
Verily, if we compare those two series of development with one another,
and ask ourselves which of the two is the more wonderful, it must be
confessed that _ontogeny_, or the short and quick history of development
of the _individual_, is much more mysterious than _phylogeny_, or the
long and slow history of development of the _tribe_. For one and the
same grand change of form is accomplished by the latter in the course of
many thousands of years, and by the former in the course of a few
months. Evidently this most rapid and astonishing transformation of the
individual in ontogenesis, which we can actually point out at any moment
by direct observation, is in itself much more wonderful and astonishing
than the corresponding, but much slower and gradual transformation which
the long chain of ancestors of the same individual has gone through in
phylogenesis.
The two series of organic development, the ontogenesis of the individual
and the phylogenesis of the tribe to which it belongs, stand in the
closest causal connection with each other. I have endeavoured, in the
second volume of the “General Morphology,”(4) to establish this theory
in detail, as I consider it exceedingly important. As I have there
shown, _ontogenesis, or the development of the individual, is a short
and quick repetition_ (recapitulation) _of phylogenesis, or the
development of the tribe to which it belongs, determined by the laws of
inheritance and adaptation_; by tribe I mean the ancestors which form
the chain of progenitors of the individual concerned. (Gen. Morph. ii.
110-147, 371.)
In this intimate connection of ontogeny and phylogeny, I see one of the
most important and irrefutable proofs of the Theory of Descent. No one
can explain these phenomena unless he has recourse to the laws of
Inheritance and Adaptation; by these alone are they explicable. These
laws, which we have previously explained, are _the laws of abbreviated,
of homochronic, and of homotopic inheritance_, and here deserve renewed
consideration. As so high and complicated an organism as that of man, or
the organism of every other mammal, rises upwards from a simple cellular
state, and as it progresses in its differentiation and perfecting it
passes through the same series of transformations which its animal
progenitors have passed through, during immense spaces of time,
inconceivable ages ago. I have already pointed out this extremely
important parallelism of the development of individuals and tribes (p.
10). Certain very early and low stages in the development of man, and
the other vertebrate animals in general, correspond completely in many
points of structure with conditions which last for life in the lower
fishes. The next phase which follows upon this presents us with a change
of the fish-like being into a kind of amphibious animal. At a later
period the mammal, with its special characteristics, develops out of the
amphibian, and we can clearly see, in the successive stages of its later
development, a series of steps of progressive transformation which
evidently correspond with the differences of different mammalian orders
and families. Now, it is precisely in the same succession that we also
see the ancestors of man, and of the higher mammals, appear one after
the other in the earth’s history; first fishes, then amphibians, later
the lower, and at last the higher mammals. Here, therefore, the
embryonic development of the individual is completely parallel to the
palæontological development of the whole tribe to which it belongs, and
this exceedingly interesting and important phenomenon can be explained
only by the interaction of the laws of Inheritance and Adaptation.
The example last mentioned, of the parallelism of the palæontological
and of the individual developmental series, now directs our attention to
a third developmental series, which stands in the closest relations to
these two, and which likewise runs, on the whole, parallel to them. I
mean that series of development of forms which constitutes the object of
investigation in _comparative anatomy_, and which I will briefly call
the _systematic developmental series of species_. By this we understand
the chain of the different, but related and connected forms, which exist
_side by side_ at any one period of the earth’s history; as for example,
at the present moment. While comparative anatomy compares the different
forms of fully-developed organisms with one another, it endeavours to
discover the common prototypes which underlie, as it were, the manifold
forms of kindred genera, classes, etc., and which are more or less
concealed by their particular differentiation. It endeavours to make out
the series of progressive steps which are indicated in the different
degrees of perfection of the divergent branches of the tribe. To make
use again of the same particular instance, comparative anatomy shows us
how the individual organs and systems of organs in the tribe of
vertebrate animals—in the different classes, families, and species of
it—have unequally developed, differentiated, and perfected themselves.
It shows us how far the succession of classes of vertebrate animals,
from the Fishes upwards, through the Amphibia to the Mammals, and here
again, from the lower to the higher orders of Mammals, forms a
progressive series or ladder. This attempt to establish a connected
anatomical developmental series we may discover in the works of the
great comparative anatomists of all ages—in the works of Goethe,
Meckel, Cuvier, Johannes Müller, Gegenbaur, and Huxley.
The developmental series of mature forms, which comparative anatomy
points out in the different diverging and ascending steps of the organic
system, and which we call the systematic developmental series, is
parallel to the palæontological developmental series, because it deals
with the _result_ of palæontological development, and it is parallel to
the individual developmental series, because this is parallel to the
palæontological series. If two parallels are parallel to a third, they
must be parallel to one another.
The varied differentiation, and the unequal degree of perfecting which
comparative anatomy points out in the developmental series of the
System, is chiefly determined by the ever increasing variety of
conditions of existence to which the different groups adapt themselves
in the struggle for life, and by the different degrees of rapidity and
completeness with which this adaptation has been effected. Conservative
groups which have retained their inherited peculiarities most
tenaciously remain, in consequence, at the lowest and rudest stage of
development. Those groups progressing most rapidly and variously, and
which have adapted themselves to changed conditions of existence most
readily have attained the highest degree of perfection. The further the
organic world developed in the course of the earth’s history, the
greater must the gap between the lower conservative and the higher
progressive groups have become, as in fact may be seen too in the
history of nations. In this way also is explained the historical fact,
that the most perfect animal and vegetable groups have developed
themselves in a comparatively short time to a considerable height, while
the lowest or most conservative groups have remained stationary
throughout all ages in their original simple stage, or have progressed,
but very slowly and gradually. The series of man’s progenitors clearly
shows this state of things. The sharks of the present day are still very
like the primary fish, which are among the most ancient vertebrate
progenitors of man, and the lowest amphibians of the present day (the
gilled salamanders and salamanders) are very like the amphibians which
first developed themselves out of fishes. So, too, the later ancestors
of man, the Monotremata and Marsupials, the most ancient mammals, are at
the same time the most imperfect animals of the class which still exist.
The laws of inheritance and adaptation known to us are completely
sufficient to explain this exceedingly important and interesting
phenomenon, which may be briefly designated as the _parallelism of
individual, of palæontological, and of systematic development_. No
opponent of the Theory of Descent has been able to give an explanation
of this extremely wonderful fact, whereas it is perfectly explained,
according to the Theory of Descent, by the laws of Inheritance and
Adaptation.
If we examine this parallelism of the three organic series of
development more accurately, we have to add the following special
qualifications. _Ontogeny_, or the history of the individual development
of every organism (embryology and metamorphology), presents us with a
simple _unbranching_ or graduated chain of forms; and so it is with that
_portion of phylogeny_ which comprises the palæontological history of
development of the _direct ancestors only_ of an individual organism.
But _the whole of phylogeny_—which meets us in the _natural system_ of
every organic tribe or phylum, and which is concerned with the
investigation of the palæontological development _of all_ the branches
of this tribe—forms a _branching_ or tree-shaped developmental series,
a veritable pedigree. If we examine and compare the branches of this
pedigree, and place them together according to the degree of their
differentiation and perfection, we obtain the tree-shaped, branching,
_systematic developmental series of comparative anatomy_. Strictly
speaking, therefore, the latter is parallel to _the whole of phylogeny_,
and consequently is only partially parallel to ontogeny; for ontogeny
itself is parallel only to _a portion_ of phylogeny.
All the phenomena of organic development above discussed, especially the
threefold genealogical parallelism, and the laws of differentiation and
progress, which are evident in each of these three series of organic
development, and, further, the whole history of rudimentary organs, are
exceedingly important proofs of the truth of the Theory of Descent. For
by it alone can they be explained, whereas its opponents cannot even
offer a shadow of an explanation of them. Without the Doctrine of
Filiation, the fact of organic development in general cannot be
understood. We should therefore, for this reason alone, be forced to
accept Lamarck’s Theory of Descent, even if we did not possess Darwin’s
Theory of Selection.