That which we may distinguish as transverse integration, is well illustrated among the Annulosa in the development of the nervous system. Leaving out those simple forms which do not present distinct ganglia, it is to be observed that the lower annulose animals, in common with the larvæ of the higher, are severally characterized by a double chain of ganglia running from end to end of the body; while in the more advanced annulose animals this double chain becomes a single chain. Mr. Newport has described the course of this concentration in insects; and by Rathke it has been traced in crustaceans. In the early stages of the Astacus fluviatilis, or common cray-fish, there is a pair of separate ganglia to each ring. Of the fourteen pairs belonging to the head and thorax, the three pairs in advance of the mouth consolidate into one mass to form the brain, or cephalic ganglion. Meanwhile out of the remainder, the first six pairs severally unite in the median line, while the rest remain more or less separate. Of these six double ganglia thus formed, the anterior four coalesce into one mass; the remaining two coalesce into another mass; and then these two masses coalesce into one. Here we see longitudinal and transverse integration going on simultaneously; and in the highest crustaceans they are both carried still further. The Vertebrata exhibit this transverse integration in the development of the generative system. The lowest of the mammalia—the Monotremata—in common with birds, have oviducts which towards their lower extremities are dilated into cavities severally performing in an imperfect way the function of a uterus. "In the Marsupialia, there is a closer approximation of the two lateral sets of organs on the median line; for the oviducts converge towards one another and meet (without coalescing) on the median line; so that their uterine dilatations are in contact with each other, forming a true 'double uterus.' … As we ascend the series of 'placental' mammals, we find the lateral coalescence becoming gradually more and more complete. … In many of the Rodentia, the uterus still remains completely divided into two lateral halves; whilst in others, these coalesce at their lower portion, forming a rudiment of the true 'body' of the uterus in the Human subject. This part increases at the expense of the lateral 'cornua' in the higher Herbivora and Carnivora; but even in the lower Quadrumana, the uterus is somewhat cleft at its summit."[6] And this process of transverse integration, which is still more striking when observed in its details, is accompanied by parallel though less important changes in the opposite sex. Once more; in the increasing commissural connexion of the cerebral hemispheres, which, though separate in the lower vertebrata, become gradually more united in the higher, we have another instance. And further ones of a different order, but of like general implication, are supplied by the vascular system.
Now it seems to us that the various kinds of integration here exemplified, which are commonly set down as so many independent phenomena, ought to be generalized, and included in the formula describing the process of development. The fact that in an adult crab, many pairs of ganglia originally separate have become fused into a single mass, is a fact only second in significance to the differentiation of its alimentary canal into stomach and intestine. That in the higher Annulosa, a single heart replaces the string of rudimentary hearts constituting the dorsal blood-vessel in the lower Annulosa, (reaching in one species to the number of one hundred and sixty), is a truth as much needing to be comprised in the history of evolution, as is the formation of a respiratory surface by a branched expansion of the skin. A right conception of the genesis of a vertebral column, includes not only the differentiations from which result the chorda dorsalis and the vertebral segments imbedded in it; but quite as much it includes the coalescence of numerous vertebral processes with their respective vertebral bodies. The changes in virtue of which several things become one, demand recognition equally with those in virtue of which one thing becomes several. Evidently, then, the current statement which ascribes the developmental progress to differentiations alone, is incomplete. Adequately to express the facts, we must say that the transition from the homogeneous to the heterogeneous is carried on by differentiations and accompanying integrations.
It may not be amiss here to ask—What is the meaning of these integrations? The evidence seems to show that they are in some way dependent on community of function. The eight segments which coalesce to make the head of a centipede, jointly protect the cephalic ganglion, and afford a solid fulcrum for the jaws, &c. The many bones which unite to form a vertebral skull have like uses. In the consolidation of the several pieces which constitute a mammalian pelvis, and in the anchylosis of from ten to nineteen vertebræ in the sacrum of a bird, we have kindred instances of the integration of parts which transfer the weight of the body to the legs. The more or less extensive fusion of the tibia with the fibula and the radius with the ulna in the ungulated mammals, whose habits require only partial rotations of the limbs, is a fact of like meaning. And all the instances lately given—the concentration of ganglia, the replacement of many pulsating blood-sacs by fewer and finally by one, the fusion of two uteri into a single uterus—have the same implication. Whether, as in some cases, the integration is merely a consequence of the growth which eventually brings into contact adjacent parts performing similar duties; or whether, as in other cases, there is an actual approximation of these parts before their union; or whether, as in yet other cases, the integration is of that indirect kind which arises when, out of a number of like organs, one, or a group, discharges an ever-increasing share of the common function, and so grows while the rest dwindle and disappear;—the general fact remains the same, that there is a tendency to the unification of parts having similar duties.
The tendency, however, acts under limiting conditions; and recognition of them will explain some apparent exceptions. In the human fœtus, as in the lower vertebrata, the eyes are placed one on each side of the head. During evolution they become relatively nearer, and at birth are in front; though they are still, in the European infant as in the adult Mongol, proportionately further apart than they afterwards become. But this approximation shows no signs of further increase. Two reasons suggest themselves. One is that the two eyes have not quite the same function, since they are directed to slightly-different aspects of each object looked at; and, since the resulting binocular vision has an advantage over monocular vision, there results a check upon further approach towards identity of function and unity of structure. The other reason is that the interposed structures do not admit of any nearer approach. For the orbits of the eyes to be brought closer together, would imply a decrease in the olfactory chambers; and as these are probably not larger than is demanded by their present functional activity, no decrease can take place. Again, if we trace up the external organs of smell through fishes,[7] reptiles, ungulate mammals and unguiculate mammals, to man, we perceive a general tendency to coalescence in the median line; and on comparing the savage with the civilized, or the infant with the adult, we see this approach of the nostrils carried furthest in the most perfect of the species. But since the septum which divides them has the function both of an evaporating surface for the lachrymal secretion, and of a ramifying surface for a nerve ancillary to that of smell, it does not disappear entirely: the integration remains incomplete. These and other like instances do not however militate against the hypothesis. They merely show that the tendency is sometimes antagonized by other tendencies. Bearing in mind which qualification, we may say, that as differentiation of parts is connected with difference of function, so there appears to be a connexion between integration of parts and sameness of function.
Closely related to the general truth that the evolution of all organisms is carried on by combined differentiations and integrations, is another general truth, which physiologists appear not to have recognized. When we look at the organic world as a whole, we may observe that, on passing from lower to higher forms, we pass to forms which are not only characterized by a greater differentiation of parts, but are at the same time more completely differentiated from the surrounding medium. This truth may be contemplated under various aspects.
In the first place it is illustrated in structure. The advance from the homogeneous to the heterogeneous itself involves an