The Foundations of the Origin of Species Two Essays written in 1842 and 1844
CHAPTER VIII
UNITY OF TYPE IN THE GREAT CLASSES; AND MORPHOLOGICAL STRUCTURES
_Unity of Type_{454}.
{454} _Origin_, Ed. i. p. 434, vi. p. 595. Ch. VIII corresponds to a section of Ch. XIII in the _Origin_, Ed. i.
Scarcely anything is more wonderful or has been oftener insisted on than that the organic beings in each great class, though living in the most distant climes and at periods immensely remote, though fitted to widely different ends in the economy of nature, yet all in their internal structure evince an obvious uniformity. What, for instance, is more wonderful than that the hand to clasp, the foot or hoof to walk, the bat's wing to fly, the porpoise's fin{455} to swim, should all be built on the same plan? and that the bones in their position and number should be so similar that they can all be classed and called by the same names. Occasionally some of the bones are merely represented by an apparently useless, smooth style, or are soldered closely to other bones, but the unity of type is not by this destroyed, and hardly rendered less clear. We see in this fact some deep bond of union between the organic beings of the same great classes--to illustrate which is the object and foundation of the natural system. The perception of this bond, I may add, is the evident cause that naturalists make an ill-defined distinction between true and adaptive affinities.
{455} _Origin_, Ed. i. p. 434, vi. p. 596. In the _Origin_, Ed. i. these examples occur under the heading _Morphology_; the author does not there draw much distinction between this heading and that of _Unity of Type_.
_Morphology._
There is another allied or rather almost identical class of facts admitted by the least visionary naturalists and included under the name of Morphology. These facts show that in an individual organic being, several of its organs consist of some other organ metamorphosed{456}: thus the sepals, petals, stamens, pistils, &c. of every plant can be shown to be metamorphosed leaves; and thus not only can the number, position and transitional states of these several organs, but likewise their monstrous changes, be most lucidly explained. It is believed that the same laws hold good with the gemmiferous vesicles of Zoophytes. In the same manner the number and position of the extraordinarily complicated jaws and palpi of Crustacea and of insects, and likewise their differences in the different groups, all become simple, on the view of these parts, or rather legs and all metamorphosed appendages, being metamorphosed legs. The skulls, again, of the Vertebrata are composed of three metamorphosed vertebræ, and thus we can see a meaning in the number and strange complication of the bony case of the brain. In this latter instance, and in that of the jaws of the Crustacea, it is only necessary to see a series taken from the different groups of each class to admit the truth of these views. It is evident that when in each species of a group its organs consist of some other part metamorphosed, that there must also be a "unity of type" in such a group. And in the cases as that above given in which the foot, hand, wing and paddle are said to be constructed on a uniform type, if we could perceive in such parts or organs traces of an apparent change from some other use or function, we should strictly include such parts or organs in the department of morphology: thus if we could trace in the limbs of the Vertebrata, as we can in their ribs, traces of an apparent change from being processes of the vertebræ, it would be said that in each species of the Vertebrata the limbs were "metamorphosed spinal processes," and that in all the species throughout the class the limbs displayed a "unity of type{457}."
{456} See _Origin_, Ed. i. p. 436, vi. p. 599, where the parts of the flower, the jaws and palpi of Crustaceans and the vertebrate skull are given as examples.
{457} The author here brings _Unity of Type_ and _Morphology_ together.
These wonderful parts of the hoof, foot, hand, wing, paddle, both in living and extinct animals, being all constructed on the same framework, and again of the petals, stamina, germens, &c. being metamorphosed leaves, can by the creationist be viewed only as ultimate facts and incapable of explanation; whilst on our theory of descent these facts all necessary follow: for by this theory all the beings of any one class, say of the mammalia, are supposed to be descended from one parent-stock, and to have been altered by such slight steps as man effects by the selection of chance domestic variations. Now we can see according to this view that a foot might be selected with longer and longer bones, and wider connecting membranes, till it became a swimming organ, and so on till it became an organ by which to flap along the surface or to glide over it, and lastly to fly through the air: but in such changes there would be no tendency to alter the framework of the internal inherited structure. Parts might become lost (as the tail in dogs, or horns in cattle, or the pistils in plants), others might become united together (as in the feet of the Lincolnshire breed of pigs{458}, and in the stamens of many garden flowers); parts of a similar nature might become increased in number (as the vertebræ in the tails of pigs, &c., &c. and the fingers and toes in six-fingered races of men and in the Dorking fowls), but analogous differences are observed in nature and are not considered by naturalists to destroy the uniformity of the types. We can, however, conceive such changes to be carried to such length that the unity of type might be obscured and finally be undistinguishable, and the paddle of the Plesiosaurus has been advanced as an instance in which the uniformity of type can hardly be recognised{459}. If after long and gradual changes in the structure of the co-descendants from any parent stock, evidence (either from monstrosities or from a graduated series) could be still detected of the function, which certain parts or organs played in the parent stock, these parts or organs might be strictly determined by their former function with the term "metamorphosed" appended. Naturalists have used this term in the same metaphorical manner as they have been obliged to use the terms of affinity and relation; and when they affirm, for instance, that the jaws of a crab are metamorphosed legs, so that one crab has more legs and fewer jaws than another, they are far from meaning that the jaws, either during the life of the individual crab or of its progenitors, were really legs. By our theory this term assumes its literal meaning{460}; and this wonderful fact of the complex jaws of an animal retaining numerous characters, which they would probably have retained if they had really been metamorphosed during many successive generations from true legs, is simply explained.
{458} The solid-hoofed pigs mentioned in _Var. under Dom._, Ed. ii. vol. II. p. 424 are not _Lincolnshire pigs_. For other cases see Bateson, _Materials for the Study of Variation_, 1894, pp. 387-90.
{459} In the margin C. Bell is given as authority, apparently for the statement about Plesiosaurus. See _Origin_, Ed. i. p. 436, vi. p. 598, where the author speaks of the "general pattern" being obscured in "extinct gigantic sea lizards." In the same place the suctorial Entomostraca are added as examples of the difficulty of recognising the type.
{460} _Origin_, Ed. i. p. 438, vi. p. 602.
_Embryology_.
The unity of type in the great classes is shown in another and very striking manner, namely, in the stages through which the embryo passes in coming to maturity{461}. Thus, for instance, at one period of the embryo, the wings of the bat, the hand, hoof or foot of the quadruped, and the fin of the porpoise do not differ, but consist of a simple undivided bone. At a still earlier period the embryo of the fish, bird, reptile and mammal all strikingly resemble each other. Let it not be supposed this resemblance is only external; for on dissection, the arteries are found to branch out and run in a peculiar course, wholly unlike that in the full-grown mammal and bird, but much less unlike that in the full-grown fish, for they run as if to ærate blood by branchiæ{462} on the neck, of which even the slit-like orifices can be discerned. How wonderful it is that this structure should be present in the embryos of animals about to be developed into such different forms, and of which two great classes respire only in the air. Moreover, as the embryo of the mammal is matured in the parent's body, and that of the bird in an egg in the air, and that of the fish in an egg in the water, we cannot believe that this course of the arteries is related to any external conditions. In all shell-fish (Gasteropods) the embryo passes through a state analogous to that of the Pteropodous Mollusca: amongst insects again, even the most different ones, as the moth, fly and beetle, the crawling larvæ are all closely analogous: amongst the Radiata, the jelly-fish in its embryonic state resembles a polype, and in a still earlier state an infusorial animalcule--as does likewise the embryo of the polype. From the part of the embryo of a mammal, at one period, resembling a fish more than its parent form; from the larvæ of all orders of insects more resembling the simpler articulate animals than their parent insects{463}; and from such other cases as the embryo of the jelly-fish resembling a polype much nearer than the perfect jelly-fish; it has often been asserted that the higher animal in each class passes through the state of a lower animal; for instance, that the mammal amongst the vertebrata passes through the state of a fish{464}: but Müller denies this, and affirms that the young mammal is at no time a fish, as does Owen assert that the embryonic jelly-fish is at no time a polype, but that mammal and fish, jelly-fish and polype pass through the same state; the mammal and jelly-fish being only further developed or changed.
{461} _Origin_, Ed. i. p. 439, vi. p. 604.
{462} The uselessness of the branchial arches in mammalia is insisted on in the _Origin_, Ed. i. p. 440, vi. p. 606. Also the uselessness of the spots on the young blackbird and the stripes of the lion-whelp, cases which do not occur in the present Essay.
{463} In the _Origin_, Ed. i. pp. 442, 448, vi. pp. 608, 614 it is pointed out that in some cases the young form resembles the adult, _e.g._ in spiders; again, that in the Aphis there is no "worm-like stage" of development.
{464} In the _Origin_, Ed. i. p. 449, vi. p. 618, the author speaks doubtfully about the recapitulation theory.
As the embryo, in most cases, possesses a less complicated structure than that into which it is to be developed, it might have been thought that the resemblance of the embryo to less complicated forms in the same great class, was in some manner a necessary preparation for its higher development; but in fact the embryo, during its growth, may become less, as well as more, complicated{465}. Thus certain female Epizoic Crustaceans in their mature state have neither eyes nor any organs of locomotion; they consist of a mere sack, with a simple apparatus for digestion and procreation; and when once attached to the body of the fish, on which they prey, they never move again during their whole lives: in their embryonic condition, on the other hand, they are furnished with eyes, and with well articulated limbs, actively swim about and seek their proper object to become attached to. The larvæ, also, of some moths are as complicated and are more active than the wingless and limbless females, which never leave their pupa-case, never feed and never see the daylight.
{465} This corresponds to the _Origin_, Ed. i. p. 441, vi. p. 607, where, however, the example is taken from the Cirripedes.
_Attempt to explain the facts of embryology._
I think considerable light can be thrown by the theory of descent on these wonderful embryological facts which are common in a greater or less degree to the whole animal kingdom, and in some manner to the vegetable kingdom: on the fact, for instance, of the arteries in the embryonic mammal, bird, reptile and fish, running and branching in the same courses and nearly in the same manner with the arteries in the full-grown fish; on the fact I may add of the high importance to systematic naturalists{466} of the characters and resemblances in the embryonic state, in ascertaining the true position in the natural system of mature organic beings. The following are the considerations which throw light on these curious points.
{466} _Origin_, Ed. i. p. 449, vi. p. 617.
In the economy, we will say of a feline animal{467}, the feline structure of the embryo or of the sucking kitten is of quite secondary importance to it; hence, if a feline animal varied (assuming for the time the possibility of this) and if some place in the economy of nature favoured the selection of a longer-limbed variety, it would be quite unimportant to the production by natural selection of a long-limbed breed, whether the limbs of the embryo and kitten were elongated if they _became_ so _as soon_ as the animal had to provide food for itself. And if it were found after continued selection and the production of several new breeds from one parent-stock, that the successive variations had supervened, not very early in the youth or embryonic life of each breed (and we have just seen that it is quite unimportant whether it does so or not), then it obviously follows that the young or embryos of the several breeds will continue resembling each other more closely than their adult parents{468}. And again, if two of these breeds became each the parent-stock of several other breeds, forming two genera, the young and embryos of these would still retain a greater resemblance to the one original stock than when in an adult state. Therefore if it could be shown that the period of the slight successive variations does not always supervene at a very early period of life, the greater resemblance or closer unity in type of animals in the young than in the full-grown state would be explained. Before practically{469} endeavouring to discover in our domestic races whether the structure or form of the young has or has not changed in an exactly corresponding degree with the changes of full-grown animals, it will be well to show that it is at least quite _possible_ for the primary germinal vesicle to be impressed with a tendency to produce some change on the growing tissues which will not be fully effected till the animal is advanced in life.
{467} This corresponds to the _Origin_, Ed. i. pp. 443-4, vi. p. 610: the "feline animal" is not used to illustrate the generalisation, but is so used in the Essay of 1842, p. 42.
{468} _Origin_, Ed. i. p. 447, vi. p. 613.
{469} In the margin is written "Get young pigeons"; this was afterwards done, and the results are given in the _Origin_, Ed. i. p. 445, vi. p. 612.
From the following peculiarities of structure being inheritable and appearing only when the animal is full-grown--namely, general size, tallness (not consequent on the tallness of the infant), fatness either over the whole body, or local; change of colour in hair and its loss; deposition of bony matter on the legs of horses; blindness and deafness, that is changes of structure in the eye and ear; gout and consequent deposition of chalk-stones; and many other diseases{470}, as of the heart and brain, &c., &c.; from all such tendencies being I repeat inheritable, we clearly see that the germinal vesicle is impressed with some power which is wonderfully preserved during the production of infinitely numerous cells in the ever changing tissues, till the part ultimately to be affected is formed and the time of life arrived at. We see this clearly when we select cattle with any peculiarity of their horns, or poultry with any peculiarity of their second plumage, for such peculiarities cannot of course reappear till the animal is mature. Hence, it is certainly _possible_ that the germinal vesicle may be impressed with a tendency to produce a long-limbed animal, the full proportional length of whose limbs shall appear only when the animal is mature{471}.
{470} In the _Origin_, Ed. i. the corresponding passages are at pp. 8, 13, 443, vi. pp. 8, 15, 610. In the _Origin_, Ed. i. I have not found a passage so striking as that which occurs a few lines lower "that the germinal vesicle is impressed with some power which is wonderfully preserved, &c." In the _Origin_ this _preservation_ is rather taken for granted.
{471} Aborted organs show, perhaps, something about period which changes supervene in embryo.
In several of the cases just enumerated we know that the first cause of the peculiarity, when _not_ inherited, lies in the conditions to which the animal is exposed during mature life, thus to a certain extent general size and fatness, lameness in horses and in a lesser degree blindness, gout and some other diseases are certainly in some degree caused and accelerated by the habits of life, and these peculiarities when transmitted to the offspring of the affected person reappear at a nearly corresponding time of life. In medical works it is asserted generally that at whatever period an hereditary disease appears in the parent, it tends to reappear in the offspring at the same period. Again, we find that early maturity, the season of reproduction and longevity are transmitted to corresponding periods of life. Dr Holland has insisted much on children of the same family exhibiting certain diseases in similar and peculiar manners; my father has known three brothers{472} die in very old age in a _singular_ comatose state; now to make these latter cases strictly bear, the children of such families ought similarly to suffer at corresponding times of life; this is probably not the case, but such facts show that a tendency in a disease to appear at particular stages of life can be transmitted through the germinal vesicle to different individuals of the same family. It is then certainly possible that diseases affecting widely different periods of life can be transmitted. So little attention is paid to very young domestic animals that I do not know whether any case is on record of selected peculiarities in young animals, for instance, in the first plumage of birds, being transmitted to their young. If, however, we turn to silk-worms{473}, we find that the caterpillars and coccoons (which must correspond to a _very early_ period of the embryonic life of mammalia) vary, and that these varieties reappear in the offspring caterpillars and coccoons.
{472} See p. 42, note 5.{Note 160}
{473} The evidence is given in _Var. under Dom._, I. p. 316.
I think these facts are sufficient to render it probable that at whatever period of life any peculiarity (capable of being inherited) appears, whether caused by the action of external influences during mature life, or from an affection of the primary germinal vesicle, it _tends_ to reappear in the offspring at the corresponding period of life{474}. Hence (I may add) whatever effect training, that is the full employment or action of every newly selected slight variation, has in fully developing and increasing such variation, would only show itself in mature age, corresponding to the period of training; in the second