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“Changes of any kind in the conditions of life, even extremely slight changes, often suffice to cause variability…. Animals and plants continue to be variable for an immense period after their first domestication; … In the course of time they can be habituated to certain changes, so as to become less variable; … There is good evidence that the power of changed conditions accumulates; so that two, three, or more generations must be exposed to new conditions before any effect is visible…. Some variations are induced by the direct action of the surrounding conditions on the whole organization, or on certain parts alone, and other variations are induced indirectly through the reproductive system being affected in the same manner as is so common with organic beings when removed from their natural conditions.”–(Animals and Plants under Domestication, vol. ii, 270.)

There are to be recognized two modes of this effect produced by changed conditions on the reproductive system, and consequently on offspring. Simple arrest of development is one. But beyond the variations of offspring arising from imperfectly developed reproductive systems in parents–variations which must be ordinarily in the nature of imperfections–there are others due to a changed balance of functions caused by changed conditions. The fact noted by Mr. Darwin in the above passage, “that the power of changed conditions accumulates; so that two, three, or more generations must be exposed to new conditions before any effect is visible,” implies that during these generations there is going on some change of constitution consequent on the changed proportions and relations of the functions. I will not dwell on the implication, which seems tolerably clear, that this change must consist of such modifications of organs as adapt them to their changed functions; and that if the influence of changed conditions “accumulates,” it must be through the inheritance of such modifications. Nor will I press the question–What is the nature of the effect registered in the reproductive elements, and which is subsequently manifested by variations?–Is it an effect entirely irrelevant to the new requirements of the variety?–Or is it an effect which makes the variety less fit for the new requirements?–Or is it an effect which makes it more fit for the new requirements? But not pressing these questions, it suffices to point out the necessary implication that changed functions of organs do, in some way or other, register themselves in changed proclivities of the reproductive elements. In face of these facts it cannot be denied that the modified action of a part produces an inheritable effect–be the nature of that effect what it may.

The second of the remarks above adverted to as made by Mr. Darwin, is contained in his sections dealing with correlated variations. In the Origin of Species, p. 114, he says–

“The whole organization is so tied together during its growth and development, that when slight variations in any one part occur, and are accumulated through natural selection, other parts become modified.”

And a parallel statement contained in Animals and Plants under Domestication, vol. ii, p. 320, runs thus–

“Correlated variation is an important subject for us; for when one part is modified through continued selection, either by man or under nature, other parts of the organization will be unavoidably modified. From this correlation it apparently follows that, with our domesticated animals and plants, varieties rarely or never differ from each other by some single character alone.”

By what process does a changed part modify other parts? By modifying their functions in some way or degree, seems the necessary answer. It is indeed, imaginable, that where the part changed is some dermal appendage which, becoming larger, has abstracted more of the needful material from the general stock, the effect may consist simply in diminishing the amount of this material available for other dermal appendages, leading to diminution of some or all of them, and may fail to affect in appreciable ways the rest of the organism: save perhaps the blood-vessels near the enlarged appendage. But where the part is an active one–a limb, or viscus, or any organ which constantly demands blood, produces waste matter, secretes, or absorbs–then all the other active organs become implicated in the change. The functions performed by them have to constitute a moving equilibrium; and the function of one cannot, by alteration of the structure performing it, be modified in degree or kind, without modifying the functions of the rest–some appreciably and others inappreciably, according to the directness or indirectness of their relations. Of such inter-dependent changes, the normal ones are naturally inconspicuous; but those which are partially or completely abnormal, sufficiently carry home the general truth. Thus, unusual cerebral excitement affects the excretion through the kidneys in quantity or quality or both. Strong emotions of disagreeable kinds check or arrest the flow of bile. A considerable obstacle to the circulation offered by some important structure in a diseased or disordered state, throwing more strain upon the heart, causes hypertrophy of its muscular walls; and this change which is, so far as concerns the primary evil, a remedial one, often entails mischiefs in other organs. “Apoplexy and palsy, in a scarcely credible number of cases, are directly dependent on hypertrophic enlargement of the heart.” And in other cases, asthma, dropsy, and epilepsy are caused. Now if a result of this inter-dependence as seen in the individual organism, is that a local modification of one part produces, by changing their functions, correlative modifications of other parts, then the question here to be put is–Are these correlative modifications, when of a kind falling within normal limits, inheritable or not. If they are inheritable, then the fact stated by Mr. Darwin that “when one part is modified through continued selection,” “other parts of the organization will be unavoidably modified” is perfectly intelligible: these entailed secondary modifications are transmitted pari passu with the successive modifications produced by selection. But what if they are not inheritable? Then these secondary modifications caused in the individual, not being transmitted to descendants, the descendants must commence life with organizations out of balance, and with each increment of change in the part affected by selection, their organizations must get more out of balance–must have a larger and larger amounts of re-organization to be made during their lives. Hence the constitution of the variety must become more and more unworkable.