CHAPTER XII

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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