UNITY OF TYPE IN THE GREAT CLASSES; AND MORPHOLOGICAL STRUCTURES
Unity of Type.
 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 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.
 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.
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: 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."
 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.
 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, 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. 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; 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.
 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.
 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.
 Origin, Ed. i. p. 438, vi. p. 602.
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. 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 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; 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: 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.
 Origin, Ed. i. p. 439, vi. p. 604.
 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.
 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.
 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. 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.
 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 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.
 Origin, Ed. i. p. 449, vi. p. 617.
In the economy, we will say of a feline animal, 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. 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 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.
 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.
 Origin, Ed. i. p. 447, vi. p. 613.
 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, 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.
 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.
 [[In the margin is written]] Aborted organs show, perhaps, something about period [[at]] 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 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, 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.
 See p. 42, note 5.[Note 160]
 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. 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 chapter I showed that there was in this respect a marked difference in natural and artificial selection, man not regularly exercising or adapting his varieties to new ends, whereas selection by nature presupposes such exercise and adaptation in each selected and changed part. The foregoing facts show and presuppose that slight variations occur at various periods of life after birth; the facts of monstrosity, on the other hand, show that many changes take place before birth, for instance, all such cases as extra fingers, hare-lip and all sudden and great alterations in structure; and these when inherited reappear during the embryonic period in the offspring. I will only add that at a period even anterior to embryonic life, namely, during the egg state, varieties appear in size and colour (as with the Hertfordshire duck with blackish eggs) which reappear in the egg; in plants also the capsule and membranes of the seed are very variable and inheritable.
 Origin, Ed. i. p. 444, vi. p. 610.
 In Var. under Dom., Ed. ii. vol. I. p. 295, such eggs are said to be laid early in each season by the black Labrador duck. In the next sentence in the text the author does not distinguish the characters of the vegetable capsule from those of the ovum.
If then the two following propositions are admitted (and I think the first can hardly be doubted), viz. that variation of structure takes place at all times of life, though no doubt far less in amount and seldomer in quite mature life (and then generally taking the form of disease); and secondly, that these variations tend to reappear at a corresponding period of life, which seems at least probable, then we might a priori have expected that in any selected breed the young animal would not partake in a corresponding degree the peculiarities characterising the full-grown parent; though it would in a lesser degree. For during the thousand or ten thousand selections of slight increments in the length of the limbs of individuals necessary to produce a long-limbed breed, we might expect that such increments would take place in different individuals (as we do not certainly know at what period they do take place), some earlier and some later in the embryonic state, and some during early youth; and these increments would reappear in their offspring only at corresponding periods. Hence, the entire length of limb in the new long-limbed breed would only be acquired at the latest period of life, when that one which was latest of the thousand primary increments of length supervened. Consequently, the foetus of the new breed during the earlier part of its existence would remain much less changed in the proportions of its limbs; and the earlier the period the less would the change be.
 This seems to me to be more strongly stated here than in the Origin, Ed. i.
Whatever may be thought of the facts on which this reasoning is grounded, it shows how the embryos and young of different species might come to remain less changed than their mature parents; and practically we find that the young of our domestic animals, though differing, differ less than their full-grown parents. Thus if we look at the young puppies of the greyhound and bulldog--(the two most obviously modified of the breeds of dog)--we find their puppies at the age of six days with legs and noses (the latter measured from the eyes to the tip) of the same length; though in the proportional thicknesses and general appearance of these parts there is a great difference. So it is with cattle, though the young calves of different breeds are easily recognisable, yet they do not differ so much in their proportions as the full-grown animals. We see this clearly in the fact that it shows the highest skill to select the best forms early in life, either in horses, cattle or poultry; no one would attempt it only a few hours after birth; and it requires great discrimination to judge with accuracy even during their full youth, and the best judges are sometimes deceived. This shows that the ultimate proportions of the body are not acquired till near mature age. If I had collected sufficient facts to firmly establish the proposition that in artificially selected breeds the embryonic and young animals are not changed in a corresponding degree with their mature parents, I might have omitted all the foregoing reasoning and the attempts to explain how this happens; for we might safely have transferred the proposition to the breeds or species naturally selected; and the ultimate effect would necessarily have been that in a number of races or species descended from a common stock and forming several genera and families the embryos would have resembled each other more closely than full-grown animals. Whatever may have been the form or habits of the parent-stock of the Vertebrata, in whatever course the arteries ran and branched, the selection of variations, supervening after the first formation of the arteries in the embryo, would not tend from variations supervening at corresponding periods to alter their course at that period: hence, the similar course of the arteries in the mammal, bird, reptile and fish, must be looked at as a most ancient record of the embryonic structure of the common parent-stock of these four great classes.
 Origin, Ed. i. p. 444, vi. p. 611.
A long course of selection might cause a form to become more simple, as well as more complicated; thus the adaptation of a crustaceous animal to live attached during its whole life to the body of a fish, might permit with advantage great simplification of structure, and on this view the singular fact of an embryo being more complex than its parent is at once explained.
 Origin, Ed. i. p. 441, vi. p. 607.
On the graduated complexity in each great class.
I may take this opportunity of remarking that naturalists have observed that in most of the great classes a series exists from very complicated to very simple beings; thus in Fish, what a range there is between the sand-eel and shark,--in the Articulata, between the common crab and the Daphnia,--between the Aphis and butterfly, and between a mite and a spider. Now the observation just made, namely, that selection might tend to simplify, as well as to complicate, explains this; for we can see that during the endless geologico-geographical changes, and consequent isolation of species, a station occupied in other districts by less complicated animals might be left unfilled, and be occupied by a degraded form of a higher or more complicated class; and it would by no means follow that, when the two regions became united, the degraded organism would give way to the aboriginally lower organism. According to our theory, there is obviously no power tending constantly to exalt species, except the mutual struggle between the different individuals and classes; but from the strong and general hereditary tendency we might expect to find some tendency to progressive complication in the successive production of new organic forms.
 Compare Origin, Ed. i. p. 419, vi. p. 575.
 [[Note in original.]] Scarcely possible to distinguish between non-development and retrograde development.
Modification by selection of the forms of immature animals.
I have above remarked that the feline form is quite of secondary importance to the embryo and to the kitten. Of course, during any great and prolonged change of structure in the mature animal, it might, and often would be, indispensable that the form of the embryo should be changed; and this could be effected, owing to the hereditary tendency at corresponding ages, by selection, equally well as in mature age: thus if the embryo tended to become, or to remain, either over its whole body or in certain parts, too bulky, the female parent would die or suffer more during parturition; and as in the case of the calves with large hinder quarters, the peculiarity must be either eliminated or the species become extinct. Where an embryonic form has to seek its own food, its structure and adaptation is just as important to the species as that of the full-grown animal; and as we have seen that a peculiarity appearing in a caterpillar (or in a child, as shown by the hereditariness of peculiarities in the milk-teeth) reappears in its offspring, so we can at once see that our common principle of the selection of slight accidental variations would modify and adapt a caterpillar to a new or changing condition, precisely as in the full-grown butterfly. Hence probably it is that caterpillars of different species of the Lepidoptera differ more than those embryos, at a corresponding early period of life, do which remain inactive in the womb of their parents. The parent during successive ages continuing to be adapted by selection for some one object, and the larva for quite another one, we need not wonder at the difference becoming wonderfully great between them; even as great as that between the fixed rock-barnacle and its free, crab-like offspring, which is furnished with eyes and well-articulated, locomotive limbs.
 See p. 42, where the same illustration is used.
 Var. under Dom., Ed. ii. vol. I. p. 452.
 Origin, Ed. i. p. 441, vi. p. 607.
Importance of embryology in classification.
We are now prepared to perceive why the study of embryonic forms is of such acknowledged importance in classification. For we have seen that a variation, supervening at any time, may aid in the modification and adaptation of the full-grown being; but for the modification of the embryo, only the variations which supervene at a very early period can be seized on and perpetuated by selection: hence there will be less power and less tendency (for the structure of the embryo is mostly unimportant) to modify the young: and hence we might expect to find at this period similarities preserved between different groups of species which had been obscured and quite lost in the full-grown animals. I conceive on the view of separate creations it would be impossible to offer any explanation of the affinities of organic beings thus being plainest and of the greatest importance at that period of life when their structure is not adapted to the final part they have to play in the economy of nature.
 Origin, Ed. i. p. 449, vi. p. 617.
Order in time in which the great classes have first appeared.
It follows strictly from the above reasoning only that the embryos of (for instance) existing vertebrata resemble more closely the embryo of the parent-stock of this great class than do full-grown existing vertebrata resemble their full-grown parent-stock. But it may be argued with much probability that in the earliest and simplest condition of things the parent and embryo must have resembled each other, and that the passage of any animal through embryonic states in its growth is entirely due to subsequent variations affecting only the more mature periods of life. If so, the embryos of the existing vertebrata will shadow forth the full-grown structure of some of those forms of this great class which existed at the earlier periods of the earth's history: and accordingly, animals with a fish-like structure ought to have preceded birds and mammals; and of fish, that higher organized division with the vertebræ extending into one division of the tail ought to have preceded the equal-tailed, because the embryos of the latter have an unequal tail; and of Crustacea, entomostraca ought to have preceded the ordinary crabs and barnacles--polypes ought to have preceded jelly-fish, and infusorial animalcules to have existed before both. This order of precedence in time in some of these cases is believed to hold good; but I think our evidence is so exceedingly incomplete regarding the number and kinds of organisms which have existed during all, especially the earlier, periods of the earth's history, that I should put no stress on this accordance, even if it held truer than it probably does in our present state of knowledge.
 Origin, Ed. i. p. 449, vi. p. 618.