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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.
|13 - Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Or||13-09 - MORPHOLOGY, between members of the same class, between parts of the same individual||10||
We have seen that the members of the same class, independently of their habits of life, resemble each other in the general plan of their organisation.
This resemblance is often expressed by the term `unity of type;' or by saying that the several parts and organs in the different species of the class are homologous.
The whole subject is included under the general name of Morphology.
This is the most interesting department of natural history, and may be said to be its very soul.
What can be more curious than that the hand of a man, formed for grasping, that of a mole for digging, the leg of the horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include the same bones, in the same relative positions? Geoffroy St Hilaire has insisted strongly on the high importance of relative connexion in homologous organs: the parts may change to almost any extent in form and size, and yet they always remain connected together in the same order.
We never find, for instance, the bones of the arm and forearm, or of the thigh and leg, transposed.
Hence the same names can be given to the homologous bones in widely different animals.
We see the same great law in the construction of the mouths of insects: what can be more different than the immensely long spiral proboscis of a sphinx-moth, the curious folded one of a bee or bug, and the great jaws of a beetle? yet all these organs, serving for such different purposes, are formed by infinitely numerous modifications of an upper lip, mandibles, and two pairs of maxillae.
Analogous laws govern the construction of the mouths and limbs of crustaceans.
So it is with the flowers of plants.
Nothing can be more hopeless than to attempt to explain this similarity of pattern in members of the same class, by utility or by the doctrine of final causes.
The hopelessness of the attempt has been expressly admitted by Owen in his most interesting work on the `Nature of Limbs.' On the ordinary view of the independent creation of each being, we can only say that so it is; that it has so pleased the Creator to construct each animal and plant.
The explanation is manifest on the theory of the natural selection of successive slight modifications, each modification being profitable in some way to the modified form, but often affecting by correlation of growth other parts of the organisation.
In changes of this nature, there will be little or no tendency to modify the original pattern, or to transpose parts.
The bones of a limb might be shortened and widened to any extent, and become gradually enveloped in thick membrane, so as to serve as a fin; or a webbed foot might have all its bones, or certain bones, lengthened to any extent, and the membrane connecting them increased to any extent, so as to serve as a wing: yet in all this great amount of modification there will be no tendency to alter the framework of bones or the relative connexion of the several parts.
If we suppose that the ancient progenitor, the archetype as it may be called, of all mammals, had its limbs constructed on the existing general pattern, for whatever purpose they served, we can at once perceive the plain signification of the homologous construction of the limbs throughout the whole class.
So with the mouths of insects, we have only to suppose that their common progenitor had an upper lip, mandibles, and two pair of maxillae, these parts being perhaps very simple in form; and then natural selection will account for the infinite diversity in structure and function of the mouths of insects.
Nevertheless, it is conceivable that the general pattern of an organ might become so much obscured as to be finally lost, by the atrophy and ultimately by the complete abortion of certain parts, by the soldering together of other parts, and by the doubling or multiplication of others, variations which we know to be within the limits of possibility.
In the paddles of the extinct gigantic sea-lizards, and in the mouths of certain suctorial crustaceans, the general pattern seems to have been thus to a certain extent obscured.
There is another and equally curious branch of the present subject; namely, the comparison not of the same part in different members of a class, but of the different parts or organs in the same individual.
Most physiologists believe that the bones of the skull are homologous with that is correspond in number and in relative connexion with the elemental parts of a certain number of vertebrae.
The anterior and posterior limbs in each member of the vertebrate and articulate classes are plainly homologous.
We see the same law in comparing the wonderfully complex jaws and legs in crustaceans.
It is familiar to almost every one, that in a flower the relative position of the sepals, petals, stamens, and pistils, as well as their intimate structure, are intelligible in the view that they consist of metamorphosed leaves, arranged in a spire.
In monstrous plants, we often get direct evidence of the possibility of one organ being transformed into another; and we can actually see in embryonic crustaceans and in many other animals, and in flowers, that organs which when mature become extremely different, are at an early stage of growth exactly alike.
How inexplicable are these facts on the ordinary view of creation! Why should the brain be enclosed in a box composed of such numerous and such extraordinarily shaped pieces of bone? As Owen has remarked, the benefit derived from the yielding of the separate pieces in the act of parturition of mammals, will by no means explain the same construction in the skulls of birds.
Why should similar bones have been created in the formation of the wing and leg of a bat, used as they are for such totally different purposes? Why should one crustacean, which has an extremely complex mouth formed of many parts, consequently always have fewer legs; or conversely, those with many legs have simpler mouths? Why should the sepals, petals, stamens, and pistils in any individual flower, though fitted for such widely different purposes, be all constructed on the same pattern ?
On the theory of natural selection, we can satisfactorily answer these questions.
In the vertebrata, we see a series of internal vertebrae bearing certain processes and appendages; in the articulata, we see the body divided into a series of segments, bearing external appendages; and in flowering plants, we see a series of successive spiral whorls of leaves.
An indefinite repetition of the same part or organ is the common characteristic (as Owen has observed) of all low or little-modified forms; therefore we may readily believe that the unknown progenitor of the vertebrata possessed many vertebrae; the unknown progenitor of the articulata, many segments; and the unknown progenitor of flowering plants, many spiral whorls of leaves.
We have formerly seen that parts many times repeated are eminently liable to vary in number and structure; consequently it is quite probable that natural selection, during a long-continued course of modification, should have seized on a certain number of the primordially similar elements, many times repeated, and have adapted them to the most diverse purposes.
And as the whole amount of modification will have been effected by slight successive steps, we need not wonder at discovering in such parts or organs, a certain degree of fundamental resemblance, retained by the strong principle of inheritance.
In the great class of molluscs, though we can homologise the parts of one species with those of another and distinct species, we can indicate but few serial homologies; that is, we are seldom enabled to say that one part or organ is homologous with another in the same individual.
And we can understand this fact; for in molluscs, even in the lowest members of the class, we do not find nearly so much indefinite repetition of any one part, as we find in the other great classes of the animal and vegetable kingdoms.
Naturalists frequently speak of the skull as formed of metamorphosed vertebrae: the jaws of crabs as metamorphosed legs; the stamens and pistils of flowers as metamorphosed leaves; but it would in these cases probably be more correct, as Professor Huxley has remarked, to speak of both skull and vertebrae, both jaws and legs, &c., as having been metamorphosed, not one from the other, but from some common element.
Naturalists, however, use such language only in a metaphorical sense: they are far from meaning that during a long course of descent, primordial organs of any kind vertebrae in the one case and legs in the other have actually been modified into skulls or jaws.
Yet so strong is the appearance of a modification of this nature having occurred, that naturalists can hardly avoid employing language having this plain signification.
On my view these terms may be used literally; and the wonderful fact of the jaws, for instance, of a crab retaining numerous characters, which they would probably have retained through inheritance, if they had really been metamorphosed during a long course of descent from true legs, or from some simple appendage, is explained.
|13 - Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Or||13-08 - Extinction separates and defines groups||10||
Extinction, as we have seen in the fourth chapter, has played an important part in defining and widening the intervals between the several groups in each class.
We may thus account even for the distinctness of whole classes from each other for instance, of birds from all other vertebrate animals by the belief that many ancient forms of life have been utterly lost, through which the early progenitors of birds were formerly connected with the early progenitors of the other vertebrate classes.
|13 - Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Or||13-07 - Affinities, general, complex and radiating||10||
Mr. Waterhouse has remarked that, when a member belonging to one group of animals exhibits an affinity to a quite distinct group, this affinity in most cases is general and not special: thus, according to Mr. Waterhouse, of all Rodents, the bizcacha is most nearly related to Marsupials; but in the points in which it approaches this order, its relations are general, and not to any one marsupial species more than to another.
As the points of affinity of the bizcacha to Marsupials are believed to be real and not merely adaptive, they are due on my theory to inheritance in common.
Therefore we must suppose either that all Rodents, including the bizcacha, branched off from some very ancient Marsupial, which will have had a character in some degree intermediate with respect to all existing Marsupials; or that both Rodents and Marsupials branched off from a common progenitor, and that both groups have since undergone much modification in divergent directions.
On either view we may suppose that the bizcacha has retained, by inheritance, more of the character of its ancient progenitor than have other Rodents; and therefore it will not be specially related to any one existing Marsupial, but indirectly to all or nearly all Marsupials, from having partially retained the character of their common progenitor, or of an early member of the group.
On the other hand, of all Marsupials, as Mr. Waterhouse has remarked, the phascolomys resembles most nearly, not any one species, but the general order of Rodents.
In this case, however, it may be strongly suspected that the resemblance is only analogical, owing to the phascolomys having become adapted to habits like those of a Rodent.