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|13 - Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Or||13-10 - EMBRYOLOGY, laws of, explained by variations not supervening at an early age, and being inherited at a corresponding age||10||
It has already been casually remarked that certain organs in the individual, which when mature become widely different and serve for different purposes, are in the embryo exactly alike.
The embryos, also, of distinct animals within the same class are often strikingly similar: a better proof of this cannot be given, than a circumstance mentioned by Agassiz, namely, that having forgotten to ticket the embryo of some vertebrate animal, he cannot now tell whether it be that of a mammal, bird, or reptile.
The vermiform larvae of moths, flies, beetles, &c., resemble each other much more closely than do the mature insects; but in the case of larvae, the embryos are active, and have been adapted for special lines of life.
A trace of the law of embryonic resemblance, sometimes lasts till a rather late age: thus birds of the same genus, and of closely allied genera, often resemble each other in their first and second plumage; as we see in the spotted feathers in the thrush group.
In the cat tribe, most of the species are striped or spotted in lines; and stripes can be plainly distinguished in the whelp of the lion.
We occasionally though rarely see something of this kind in plants: thus the embryonic leaves of the ulex or furze, and the first leaves of the phyllodineous acaceas, are pinnate or divided like the ordinary leaves of the leguminosae.
The points of structure, in which the embryos of widely different animals of the same class resemble each other, often have no direct relation to their conditions of existence.
We cannot, for instance, suppose that in the embryos of the vertebrata the peculiar loop-like course of the arteries near the branchial slits are related to similar conditions, in the young mammal which is nourished in the womb of its mother, in the egg of the bird which is hatched in a nest, and in the spawn of a frog under water.
We have no more reason to believe in such a relation, than we have to believe that the same bones in the hand of a man, wing of a bat, and fin of a porpoise, are related to similar conditions of life.
No one will suppose that the stripes on the whelp of a lion, or the spots on the young blackbird, are of any use to these animals, or are related to the conditions to which they are exposed.
The case, however, is different when an animal during any part of its embryonic career is active, and has to provide for itself.
The period of activity may come on earlier or later in life; but whenever it comes on, the adaptation of the larva to its conditions of life is just as perfect and as beautiful as in the adult animal.
From such special adaptations, the similarity of the larvae or active embryos of allied animals is sometimes much obscured; and cases could be given of the larvae of two species, or of two groups of species, differing quite as much, or even more, from each other than do their adult parents.
In most cases, however, the larvae, though active, still obey more or less closely the law of common embryonic resemblance.
Cirripedes afford a good instance of this: even the illustrious Cuvier did not perceive that a barnacle was, as it certainly is, a crustacean; but a glance at the larva shows this to be the case in an unmistakeable manner.
So again the two main divisions of cirripedes, the pedunculated and sessile, which differ widely in external appearance, have larvae in all their several stages barely distinguishable.
The embryo in the course of development generally rises in organisation: I use this expression, though I am aware that it is hardly possible to define clearly what is meant by the organisation being higher or lower.
But no one probably will dispute that the butterfly is higher than the caterpillar.
In some cases, however, the mature animal is generally considered as lower in the scale than the larva, as with certain parasitic crustaceans.
To refer once again to cirripedes: the larvae in the first stage have three pairs of legs, a very simple single eye, and a probosciformed mouth, with which they feed largely, for they increase much in size.
In the second stage, answering to the chrysalis stage of butterflies, they have six pairs of beautifully constructed natatory legs, a pair of magnificent compound eyes, and extremely complex antennae; but they have a closed and imperfect mouth, and cannot feed: their function at this stage is, to search by their well-developed organs of sense, and to reach by their active powers of swimming, a proper place on which to become attached and to undergo their final metamorphosis.
When this is completed they are fixed for life: their legs are now converted into prehensile organs; they again obtain a well-constructed mouth; but they have no antennae, and their two eyes are now reconverted into a minute, single, and very simple eye-spot.
In this last and complete state, cirripedes may be considered as either more highly or more lowly organised than they were in the larval condition.
But in some genera the larvae become developed either into hermaphrodites having the ordinary structure, or into what I have called complemental males: and in the latter, the development has assuredly been retrograde; for the male is a mere sack, which lives for a short time, and is destitute of mouth, stomach, or other organ of importance, excepting for reproduction.
We are so much accustomed to see differences in structure between the embryo and the adult, and likewise a close similarity in the embryos of widely different animals within the same class, that we might be led to look at these facts as necessarily contingent in some manner on growth.
But there is no obvious reason why, for instance, the wing of a bat, or the fin of a porpoise, should not have been sketched out with all the parts in proper proportion, as soon as any structure became visible in the embryo.
And in some whole groups of animals and in certain members of other groups, the embryo does not at any period differ widely from the adult: thus Owen has remarked in regard to cuttle-fish, `there is no metamorphosis; the cephalopodic character is manifested long before the parts of the embryo are completed;' and again in spiders, `there is nothing worthy to be called a metamorphosis.' The larvae of insects, whether adapted to the most diverse and active habits, or quite inactive, being fed by their parents or placed in the midst of proper nutriment, yet nearly all pass through a similar worm-like stage of development; but in some few cases, as in that of Aphis, if we look to the admirable drawings by Professor Huxley of the development of this insect, we see no trace of the vermiform stage.
How, then, can we explain these several facts in embryology, namely the very general, but not universal difference in structure between the embryo and the adult; of parts in the same individual embryo, which ultimately become very unlike and serve for diverse purposes, being at this early period of growth alike; of embryos of different species within the same class, generally, but not universally, resembling each other; of the structure of the embryo not being closely related to its conditions of existence, except when the embryo becomes at any period of life active and has to provide for itself; of the embryo apparently having sometimes a higher organisation than the mature animal, into which it is developed.
I believe that all these facts can be explained, as follows, on the view of descent with modification.
It is commonly assumed, perhaps from monstrosities often affecting the embryo at a very early period, that slight variations necessarily appear at an equally early period.
But we have little evidence on this head indeed the evidence rather points the other way; for it is notorious that breeders of cattle, horses, and various fancy animals, cannot positively tell, until some time after the animal has been born, what its merits or form will ultimately turn out.
We see this plainly in our own children; we cannot always tell whether the child will be tall or short, or what its precise features will be.
The question is not, at what period of life any variation has been caused, but at what period it is fully displayed.
The cause may have acted, and I believe generally has acted, even before the embryo is formed; and the variation may be due to the male and female sexual elements having been affected by the conditions to which either parent, or their ancestors, have been exposed.
Nevertheless an effect thus caused at a very early period, even before the formation of the embryo, may appear late in life; as when an hereditary disease, which appears in old age alone, has been communicated to the offspring from the reproductive element of one parent.
Or again, as when the horns of cross-bred cattle have been affected by the shape of the horns of either parent.
For the welfare of a very young animal, as long as it remains in its mother's womb, or in the egg, or as long as it is nourished and protected by its parent, it must be quite unimportant whether most of its characters are fully acquired a little earlier or later in life.
It would not signify, for instance, to a bird which obtained its food best by having a long beak, whether or not it assumed a beak of this particular length, as long as it was fed by its parents.
Hence, I conclude, that it is quite possible, that each of the many successive modifications, by which each species has acquired its present structure, may have supervened at a not very early period of life; and some direct evidence from our domestic animals supports this view.
But in other cases it is quite possible that each successive modification, or most of them, may have appeared at an extremely early period.
I have stated in the first chapter, that there is some evidence to render it probable, that at whatever age any variation first appears in the parent, it tends to reappear at a corresponding age in the offspring.
Certain variations can only appear at corresponding ages, for instance, peculiarities in the caterpillar, cocoon, or imago states of the silk-moth; or, again, in the horns of almost full-grown cattle.
But further than this, variations which, for all that we can see, might have appeared earlier or later in life, tend to appear at a corresponding age in the offspring and parent.
I am far from meaning that this is invariably the case; and I could give a good many cases of variations (taking the word in the largest sense) which have supervened at an earlier age in the child than in the parent.
These two principles, if their truth be admitted, will, I believe, explain all the above specified leading facts in embryology.
But first let us look at a few analogous cases in domestic varieties.
Some authors who have written on Dogs, maintain that the greyhound and bulldog, though appearing so different, are really varieties most closely allied, and have probably descended from the same wild stock; hence I was curious to see how far their puppies differed from each other: I was told by breeders that they differed just as much as their parents, and this, judging by the eye, seemed almost to be the case; but on actually measuring the old dogs and their six-days old puppies, I found that the puppies had not nearly acquired their full amount of proportional difference.
So, again, I was told that the foals of cart and race-horses differed as much as the full-grown animals; and this surprised me greatly, as I think it probable that the difference between these two breeds has been wholly caused by selection under domestication; but having had careful measurements made of the dam and of a three-days old colt of a race and heavy cart-horse, I find that the colts have by no means acquired their full amount of proportional difference.
As the evidence appears to me conclusive, that the several domestic breeds of pigeon have descended from one wild species, I compared young pigeons of various breeds, within twelve hours after being hatched; I carefully measured the proportions (but will not here give details) of the beak, width of mouth, length of nostril and of eyelid, size of feet and length of leg, in the wild stock, in pouters, fantails, runts, barbs, dragons, carriers, and tumblers.
Now some of these birds, when mature, differ so extraordinarily in length and form of beak, that they would, I cannot doubt, be ranked in distinct genera, had they been natural productions.
But when the nestling birds of these several breeds were placed in a row, though most of them could be distinguished from each other, yet their proportional differences in the above specified several points were incomparably less than in the full-grown birds.
Some characteristic points of difference for instance, that of the width of mouth -- could hardly be detected in the young.
But there was one remarkable exception to this rule, for the young of the short-faced tumbler differed from the young of the wild rock-pigeon and of the other breeds, in all its proportions, almost exactly as much as in the adult state.
The two principles above given seem to me to explain these facts in regard to the later embryonic stages of our domestic varieties.
Fanciers select their horses, dogs, and pigeons, for breeding, when they are nearly grown up: they are indifferent whether the desired qualities and structures have been acquired earlier or later in life, if the full-grown animal possesses them.
And the cases just given, more especially that of pigeons, seem to show that the characteristic differences which give value to each breed, and which have been accumulated by man's selection, have not generally first appeared at an early period of life, and have been inherited by the offspring at a corresponding not early period.
But the case of the short-faced tumbler, which when twelve hours old had acquired its proper proportions, proves that this is not the universal rule; for here the characteristic differences must either have appeared at an earlier period than usual, or, if not so, the differences must have been inherited, not at the corresponding, but at an earlier age.
Now let us apply these facts and the above two principles which latter, though not proved true, can be shown to be in some degree probable to species in a state of nature.
Let us take a genus of birds, descended on my theory from some one parent-species, and of which the several new species have become modified through natural selection in accordance with their diverse habits.
Then, from the many slight successive steps of variation having supervened at a rather late age, and having been inherited at a corresponding age, the young of the new species of our supposed genus will manifestly tend to resemble each other much more closely than do the adults, just as we have seen in the case of pigeons.
We may extend this view to whole families or even classes.
The fore-limbs, for instance, which served as legs in the parent-species, may become, by a long course of modification, adapted in one descendant to act as hands, in another as paddles, in another as wings; and on the above two principles namely of each successive modification supervening at a rather late age, and being inherited at a corresponding late age the fore-limbs in the embryos of the several descendants of the parent-species will still resemble each other closely, for they will not have been modified.
But in each individual new species, the embryonic fore-limbs will differ greatly from the fore-limbs in the mature animal; the limbs in the latter having undergone much modification at a rather late period of life, and having thus been converted into hands, or paddles, or wings.
Whatever influence long-continued exercise or use on the one hand, and disuse on the other, may have in modifying an organ, such influence will mainly affect the mature animal, which has come to its full powers of activity and has to gain its own living; and the effects thus produced will be inherited at a corresponding mature age.
Whereas the young will remain unmodified, or be modified in a lesser degree, by the effects of use and disuse.
In certain cases the successive steps of variation might supervene, from causes of which we are wholly ignorant, at a very early period of life, or each step might be inherited at an earlier period than that at which it first appeared.
In either case (as with the short-faced tumbler) the young or embryo would closely resemble the mature parent-form.
We have seen that this is the rule of development in certain whole groups of animals, as with cuttle-fish and spiders, and with a few members of the great class of insects, as with Aphis.
With respect to the final cause of the young in these cases not undergoing any metamorphosis, or closely resembling their parents from their earliest age, we can see that this would result from the two following contingencies; firstly, from the young, during a course of modification carried on for many generations, having to provide for their own wants at a very early stage of development, and secondly, from their following exactly the same habits of life with their parents; for in this case, it would be indispensable for the existence of the species, that the child should be modified at a very early age in the same manner with its parents, in accordance with their similar habits.
Some further explanation, however, of the embryo not undergoing any metamorphosis is perhaps requisite.
If, on the other hand, it profited the young to follow habits of life in any degree different from those of their parent, and consequently to be constructed in a slightly different manner, then, on the principle of inheritance at corresponding ages, the active young or larvae might easily be rendered by natural selection different to any conceivable extent from their parents.
Such differences might, also, become correlated with successive stages of development; so that the larvae, in the first stage, might differ greatly from the larvae in the second stage, as we have seen to be the case with cirripedes.
The adult might become fitted for sites or habits, in which organs of locomotion or of the senses, &c., would be useless; and in this case the final metamorphosis would be said to be retrograde.
As all the organic beings, extinct and recent, which have ever lived on this earth have to be classed together, and as all have been connected by the finest gradations, the best, or indeed, if our collections were nearly perfect, the only possible arrangement, would be genealogical.
Descent being on my view the hidden bond of connexion which naturalists have been seeking under the term of the natural system.
On this view we can understand how it is that, in the eyes of most naturalists, the structure of the embryo is even more important for classification than that of the adult.
For the embryo is the animal in its less modified state; and in so far it reveals the structure of its progenitor.
In two groups of animal, however much they may at present differ from each other in structure and habits, if they pass through the same or similar embryonic stages, we may feel assured that they have both descended from the same or nearly similar parents, and are therefore in that degree closely related.
Thus, community in embryonic structure reveals community of descent.
It will reveal this community of descent, however much the structure of the adult may have been modified and obscured; we have seen, for instance, that cirripedes can at once be recognised by their larvae as belonging to the great class of crustaceans.
As the embryonic state of each species and group of species partially shows us the structure of their less modified ancient progenitors, we can clearly see why ancient and extinct forms of life should resemble the embryos of their descendants, our existing species.
Agassiz believes this to be a law of nature; but I am bound to confess that I only hope to see the law hereafter proved true.
It can be proved true in those cases alone in which the ancient state, now supposed to be represented in many embryos, has not been obliterated, either by the successive variations in a long course of modification having supervened at a very early age, or by the variations having been inherited at an earlier period than that at which they first appeared.
It should also be borne in mind, that the supposed law of resemblance of ancient forms of life to the embryonic stages of recent forms, may be true, but yet, owing to the geological record not extending far enough back in time, may remain for a long period, or for ever, incapable of demonstration.
Thus, as it seems to me, the leading facts in embryology, which are second in importance to none in natural history, are explained on the principle of slight modifications not appearing, in the many descendants from some one ancient progenitor, at a very early period in the life of each, though perhaps caused at the earliest, and being inherited at a corresponding not early period.
Embryology rises greatly in interest, when we thus look at the embryo as a picture, more or less obscured, of the common parent-form of each great class of animals.
|04 - Natural Selection||04-05 - Sexual Selection||10||
Inasmuch as peculiarities often appear under domestication in one sex and become hereditarily attached to that sex, so no doubt it will be under nature.
Thus it is rendered possible for the two sexes to be modified through natural selection in relation to different habits of life, as is sometimes the case; or for one sex to be modified in relation to the other sex, as commonly occurs.
This leads me to say a few words on what I have called Sexual Selection. This form of selection depends, not on a struggle for existence in relation to other organic beings or to external conditions, but on a struggle between the individuals of one sex, generally the males, for the possession of the other sex.
The result is not death to the unsuccessful competitor, but few or no offspring. Sexual selection is, therefore, less rigorous than natural selection.
Generally, the most vigorous males, those which are best fitted for their places in nature, will leave most progeny.
But in many cases, victory depends not so much on general vigor, as on having special weapons, confined to the male sex.
A hornless stag or spurless cock would have a poor chance of leaving numerous offspring.
Sexual selection, by always allowing the victor to breed, might surely give indomitable courage, length to the spur, and strength to the wing to strike in the spurred leg, in nearly the same manner as does the brutal cockfighter by the careful selection of his best cocks.
How low in the scale of nature the law of battle descends, I know not; male alligators have been described as fighting, bellowing, and whirling round, like Indians in a war-dance, for the possession of the females; male salmons have been observed fighting all day long; male stagbeetles sometimes bear wounds from the huge mandibles of other males; the males of certain hymenopterous insects have been frequently seen by that inimitable observer M. Fabre, fighting for a particular female who sits by, an apparently unconcerned beholder of the struggle, and then retires with the conqueror.
The war is, perhaps, severest between the males of polygamous animals, and these seem oftenest provided with special weapons.
The males of carnivorous animals are already well armed; though to them and to others, special means of defence may be given through means of sexual selection, as the mane of the lion, and the hooked jaw to the male salmon; for the shield may be as important for victory, as the sword or spear.
|13 - Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Or||13-12 - Summary||10||
In this chapter I have attempted to show, that the subordination of group to group in all organisms throughout all time; that the nature of the relationship, by which all living and extinct beings are united by complex, radiating, and circuitous lines of affinities into one grand system; the rules followed and the difficulties encountered by naturalists in their classifications; the value set upon characters, if constant and prevalent, whether of high vital importance, or of the most trifling importance, or, as in rudimentary organs, of no importance; the wide opposition in value between analogical or adaptive characters, and characters of true affinity; and other such rules; all naturally follow on the view of the common parentage of those forms which are considered by naturalists as allied, together with their modification through natural selection, with its contingencies of extinction and divergence of character.
In considering this view of classification, it should be borne in mind that the element of descent has been universally used in ranking together the sexes, ages, and acknowledged varieties of the same species, however different they may be in structure.
If we extend the use of this element of descent, the only certainly known cause of similarity in organic beings, we shall understand what is meant by the natural system: it is genealogical in its attempted arrangement, with the grades of acquired difference marked by the terms varieties, species, genera, families, orders, and classes.
On this same view of descent with modification, all the great facts in Morphology become intelligible, whether we look to the same pattern displayed in the homologous organs, to whatever purpose applied, of the different species of a class; or to the homologous parts constructed on the same pattern in each individual animal and plant.
On the principle of successive slight variations, not necessarily or generally supervening at a very early period of life, and being inherited at a corresponding period, we can understand the great leading facts in Embryology; namely, the resemblance in an individual embryo of the homologous parts, which when matured will become widely different from each other in structure and function; and the resemblance in different species of a class of the homologous parts or organs, though fitted in the adult members for purposes as different as possible.
Larvae are active embryos, which have become specially modified in relation to their habits of life, through the principle of modifications being inherited at corresponding ages.
On this same principle and bearing in mind, that when organs are reduced in size, either from disuse or selection, it will generally be at that period of life when the being has to provide for its own wants, and bearing in mind how strong is the principle of inheritance the occurrence of rudimentary organs and their final abortion, present to us no inexplicable difficulties; on the contrary, their presence might have been even anticipated.
The importance of embryological characters and of rudimentary organs in classification is intelligible, on the view that an arrangement is only so far natural as it is genealogical.
Finally, the several classes of facts which have been considered in this chapter, seem to me to proclaim so plainly, that the innumerable species, genera, and families of organic beings, with which this world is peopled, have all descended, each within its own class or group, from common parents, and have all been modified in the course of descent, that I should without hesitation adopt this view, even if it were unsupported by other facts or arguments.
|12 - Geographical Distribution -- continued||12-60 - Summary of the last and present chapters||10||
In these chapters I have endeavoured to show, that if we make due allowance for our ignorance of the full effects of all the changes of climate and of the level of the land, which have certainly occurred within the recent period, and of other similar changes which may have occurred within the same period; if we remember how profoundly ignorant we are with respect to the many and curious means of occasional transport, a subject which has hardly ever been properly experimentised on; if we bear in mind how often a species may have ranged continuously over a wide area, and then have become extinct in the intermediate tracts, I think the difficulties in believing that all the individuals of the same species, wherever located, have descended from the same parents, are not insuperable.
And we are led to this conclusion, which has been arrived at by many naturalists under the designation of single centres of creation, by some general considerations, more especially from the importance of barriers and from the analogical distribution of sub-genera, genera, and families.
With respect to the distinct species of the same genus, which on my theory must have spread from one parent-source; if we make the same allowances as before for our ignorance, and remember that some forms of life change most slowly, enormous periods of time being thus granted for their migration, I do not think that the difficulties are insuperable; though they often are in this case, and in that of the individuals of the same species, extremely grave.
As exemplifying the effects of climatal changes on distribution, I have attempted to show how important has been the influence of the modern Glacial period, which I am fully convinced simultaneously affected the whole world, or at least great meridional belts.
As showing how diversified are the means of occasional transport, I have discussed at some little length the means of dispersal of fresh-water productions.
If the difficulties be not insuperable in admitting that in the long course of time the individuals of the same species, and likewise of allied species, have proceeded from some one source; then I think all the grand leading facts of geographical distribution are explicable on the theory of migration (generally of the more dominant forms of life), together with subsequent modification and the multiplication of new forms.
We can thus understand the high importance of barriers, whether of land or water, which separate our several zoological and botanical provinces.