Still it may be fairly objected that however the attributes of the two parents are variously mingled in their offspring, they must in all of them fall between the extremes displayed in the parents. In no characteristic could one of the young exceed both parents, were there no cause of "spontaneous variation" but the one alleged. Evidently, then, there is a cause yet unfound.
§ 89. Thus far we have contemplated the process under its simplest aspect. While we have assumed the two parents to be somewhat unlike, we have assumed that each parent has a homogeneous constitution—is built up of physiological units which are exactly alike. But in no case can such a homogeneity exist. Each parent had parents who were more or less contrasted—each parent inherited at least two orders of physiological units not quite identical. Here then we have a further cause of variation. The sperm-cells or germ-cells which any organism produces, will differ from each other not quantitatively only but qualitatively. Of the slightly-unlike physiological units bequeathed to it, the reproductive cells it casts off cannot habitually contain the same proportions; and we may expect the proportions to vary not slightly but greatly. Just as, during the evolution of an organism, the physiological units derived from the two parents tend to segregate, and produce likeness to the male parent in this part and to the female parent in that; so, during the formation of reproductive cells, there will arise in one a predominance of the physiological units derived from the father, and in another a predominance of the physiological units derived from the mother. Thus, then, every fertilized germ, besides containing different amounts of the two parental influences, will contain different kinds of influences—this having received a marked impress from one grandparent, and that from another. Without further exposition the reader will see how this cause of complication, running back through each line of ancestry, must produce in every germ numerous minute differences among the units.
Here, then, we have a clue to the multiplied variations, and sometimes extreme variations, that arise in races which have once begun to vary. Amid countless different combinations of units derived from parents, and through them from ancestors, immediate and remote—amid the various conflicts in their slightly-different organic polarities, opposing and conspiring with one another in all ways and degrees; there will from time to time arise special proportions causing special deviations. From the general law of probabilities it may be concluded that while these involved influences, derived from many progenitors, must, on the average of cases, obscure and partially neutralize one another; there must occasionally result such combinations of them as will produce considerable divergences from average structures; and, at rare intervals, such combinations as will produce very marked divergences. There is thus a correspondence between the inferable results and the results as habitually witnessed.
§ 90. Still there remains a difficulty. It may be said that admitting functional change to be the initiator of variation—granting that the physiological units of an organism long subject to new conditions, will tend to become modified in such way as to cause change of structure in offspring; yet there will still be no cause of the supposed heterogeneity among the physiological units of different individuals. There seems validity in the objection, that as all the members of a species whose circumstances have been altered will be affected in the same manner, the results, when they begin to show themselves in descendants, will show themselves in the same manner: not multiform variations will arise, but deviations all in one direction.
The reply is simple. The members of a species thus circumstanced will not be similarly affected. In the absence of absolute uniformity among them, the functional changes caused in them will be more or less dissimilar. Just as men of slightly-unlike dispositions behave in quite opposite ways under the same circumstances; or just as men of slightly-unlike constitutions get diverse disorders from the same cause, and are diversely acted on by the same medicine; so, the insensibly-differentiated members of a species whose conditions have been changed, may at once begin to undergo various kinds of functional changes. As we have already seen, small initial contrasts may lead to large terminal contrasts. The intenser cold of the climate into which a species has migrated, may cause in one individual increased consumption of food to balance the greater loss of heat; while in another individual the requirement may be met by a thicker growth of fur. Or, when meeting with the new foods which a new region furnishes, accident may determine one member of the species to begin with one kind and another member with another kind; and hence may arise established habits in these respective members and their descendants. Now when the functional divergences thus set up in sundry families of a species have lasted long enough to affect their constitutions, and to modify somewhat the physiological units thrown off in their reproductive cells, the divergences produced by these in offspring will be of divers kinds. And the original homogeneity of constitution having been thus destroyed, variation may go on with increasing facility. There will result a heterogeneous mixture of modifications of structure caused by modifications of function; and of still more numerous correlated modifications, indirectly so caused. By natural selection of the most divergent forms, the unlikenesses of parents will be rendered more marked, and the limits of variation wider. Until at length the divergences of constitutions and modes of life, become great enough to lead to segregation of the varieties.
§ 91. That variations must occur, and that they must ever tend, both directly and indirectly, towards adaptive modifications, are conclusions deducible from first principles; apart from any detailed interpretations like the above. That the state of homogeneity is an unstable state we have found to be a universal truth. Each species must pass from the uniform into the more or less multiform, unless the incidence of external forces is exactly the same for all its members, which it never can be. Through the process of differentiation and integration, which of necessity brings together, or keeps together, like individuals, and separates unlike ones from them, there must nevertheless be maintained a tolerably uniform species, so long as there continues a tolerably uniform set of conditions in which it may exist. But if the conditions change, either absolutely by some disturbance of the habitat or relatively by spread of the species into other habitats, then the divergent individuals that result must be segregated by the divergent sets of conditions into distinct varieties (First Principles, § 166). When, instead of contemplating a species in the aggregate, we confine our attention to a single member and its descendants, we see it to be a corollary from the general law of equilibration that the moving equilibrium constituted by the vital actions in each member of this family, must remain constant so long as the external actions to which they correspond remain constant; and that if the external actions are changed, the disturbed balance of internal changes, if not overthrown, cannot cease undergoing modification until the internal changes are again in equilibrium with the external actions: corresponding structural alterations having arisen.
On passing from these derivative laws to the ultimate law, we see that Variation is necessitated by the persistence of force. The members of a species inhabiting any area cannot be subject to like sets of forces over the whole of that area. And if, in different parts of the area, different kinds or amounts or combinations of forces act on them, they cannot but become different in themselves and in their progeny. To say otherwise, is to say that differences in the forces will not produce differences in the effects; which is to deny the persistence of force.
CHAPTER X.
GENESIS, HEREDITY, AND VARIATION.
§ 92. A question raised, and hypothetically answered, in [§§ 78] and [79], was there postponed until we had dealt with the topics of Heredity and Variation. Let us now resume the consideration of this question, in connexion with sundry others which the facts suggest.