The Principles of Biology Vol. I/Chapter III.10
CHAPTER X.
INTERNAL FACTORS.
§ 153. We saw at the outset ( §§ 10- 16), that organic matter is built up of molecules so unstable, that the slightest variation in their conditions destroys their equilibrium, and causes them either to assume altered structures or to decompose. But a substance which is beyond all others changeable by the actions and reactions of the forces liberated from instant to instant within its own mass, must be a substance which is beyond all others changeable by the forces acting on it from without. If their composition fits organic aggregates for undergoing with special facility and rapidity those re-distributions of matter and motion whence result individual organization and life; then their composition must make them similarly apt to undergo those permanent re-distributions of matter and motion which are expressed by changes of structure, in correspondence with permanent re-distributions of matter and motion in their environments.
In First Principles, when considering the phenomena of Evolution at large, the leading characters and causes of those changes which constitute organic evolution were briefly traced. Under each of the derivative laws of force to which the passage from an incoherent, indefinite homogeneity to a coherent, definite heterogeneity, conforms, were given illustrations drawn from the metamorphoses of living bodies. Here it will be needful to contemplate the several resulting processes as going on at once, in both individuals and species.
§ 154. Our postulate being that organic evolution in general commenced with homogeneous organic matter, we have first to remember that the state of homogeneity is an unstable state (First Principles, § 149). In any aggregate "the relations of outside and inside, and of comparative nearness to neighbouring sources of influence, imply the reception of influences that are unlike in quantity, or quality, or both; and it follows that unlike changes will be produced in the parts thus dissimilarly acted upon." Further, "if any given whole, instead of being absolutely uniform throughout, consists of parts distinguishable from one another—if each of these parts, while somewhat unlike other parts, is uniform within itself; then, each of them being in unstable equilibrium, it follows that while the changes set up within it must render it multiform, they must at the same time render the whole more multiform than before;" and hence, "whether that state with which we commence be or be not one of perfect homogeneity, the process must equally be towards a relative heterogeneity." This loss of homogeneity which the special instability of organic aggregates fits them to display more promptly and variously than any other aggregates, must be shown in more numerous ways in proportion as the incident forces are more numerous. Every differentiation of structure being a result of some difference in the relations of the parts to the agencies acting on them, it follows that the more multiplied and more unlike the agencies, the more varied must be the differentiations wrought. Hence the change from a state of homogeneity to a state of heterogeneity, will be marked in proportion as the environing actions to which the organism is supposes it is only are complex. This transition from a uniform to a multiform state, must continue through successive individuals. Given a series of organisms, each of which is developed from a portion of a preceding organism, and the question is whether, after exposure of the series for a million years to changed incident forces, one of its members will be the same as though the incident forces had only just changed. To say that it will, is implicitly to deny the persistence of force. In relation to any cause of divergence, the whole series of such organisms may be considered as fused together into a continuously-existing organism; and when so considered, it becomes manifest that a continuously-acting cause will go on working a continuously-increasing effect, until some counteracting cause prevents any further effect.
But now if any primordial organic aggregate must, in itself and through its descendants, gravitate from uniformity to multiformity, in obedience to the more or less multiform forces acting on it; what must happen if these multiform forces are themselves undergoing slow variations and complications? Clearly the process, ever-advancing towards a temporary limit but ever having its limit removed, must go on unceasingly. On those structural changes wrought in the once homogeneous aggregate by an original set of incident forces, will be superposed further changes wrought by a modified set of incident forces; and so on throughout all time. Omitting for the present those circumstances which check and qualify its consequences, the instability of the homogeneous must be recognized as an ever-acting cause of organic evolution, as of all other evolution.
While it follows that every organism, considered as an individual and as one of a series, tends thus to pass into a more heterogeneous state; it also follows that every species, considered as an aggregate of individuals, tends to do the like. Throughout the area it inhabits, the conditions can never be absolutely uniform: its members must, in different parts of the area, be exposed to different sets of incident forces. Still more decided must this difference of exposure be when its members spread into other habitats. Those expansive and repressive energies which set to each species a limit that perpetually oscillates from side to side of a certain mean, are, as we lately saw, frequently changed by new combinations of the external factors—astronomic, geologic, meteorologic, and organic. Hence there from time to time arise lines of diminished resistance, along which the species flows into new localities. Such portions of the species as thus migrate, are subject to circumstances unlike its previous average circumstances. And from multiformity of the circumstances, must come multiformity of the species.
Thus the law of the instability of the homogeneous has here a three-fold corollary. As interpreted in connexion with the ever-progressing, ever-complicating changes in external factors, it involves the conclusion that there is a prevailing tendency towards greater heterogeneity in all kinds of organisms, considered both individually and in successive generations; as well as in each assemblage of organisms constituting a species; and, by consequence, in each genus, order, and class.
§ 155. When considering the causes of evolution in general, we further saw (First Principles, § 156), that the multiplication of effects aids continually to increase that heterogeneity into which homogeneity inevitably lapses. It was pointed out that since "the several parts of an aggregate are differently modified by any incident force;" and since "by the reactions of the differently modified parts the incident force itself must be divided into differently modified parts;" it follows that "each differentiated division of the aggregate thus becomes a centre from which a differentiated division of the original force is again diffused. And since unlike forces must produce unlike results, each of these differentiated forces must produce, throughout the aggregate, a further series of differentiations." To this it was added that, in proportion as the heterogeneity increases, the complications arising from this multiplication of effects grow more marked; because the more strongly contrasted the parts of an aggregate become, the more different must be their reactions on incident forces, and the more unlike must be the secondary effects which these initiate; and because every increase in the number of unlike parts adds to the number of such differentiated incident forces, and such secondary effects.
How this multiplication of effects conspires, with the instability of the homogeneous, to work an increasing multiformity of structure in an organism, was shown at the time; and the foregoing pages contain further incidental illustrations. In § 69 it was pointed out that a change in one function must produce ever-complicating perturbations in other functions; and that, eventually, all parts of the organism must be modified in their states. Suppose that the head of a bison becomes much heavier, what must be the indirect results? The muscles of the neck are put to greater exertions; and its vertebræ have to bear additional tensions and pressures, caused both by the increased weight of the head, and by the stronger contractions of the muscles that support and move it. These muscles also affect their special attachments: several of the dorsal spines suffer augmented strains; and the vertebræ to which they are fixed are more severely taxed. Further, this heavier head and the more massive neck it necessitates, require a stronger fulcrum: the whole thoracic arch, and the fore-limbs which support it, are subject to greater continuous stress and more violent occasional shocks. And the required strengthening of the fore-quarters cannot take place without the centre of gravity being changed, and the hind limbs being differently reacted upon during locomotion. Any one who compares the outline of the bison with that of its congener, the ox, will see how profoundly a heavier head affects the entire osseous and muscular systems. Besides this multiplication of mechanical effects, there is a multiplication of physiological effects. The vascular apparatus is modified throughout its whole structure by each considerable modification in the proportions of the body. Increase in the size of any organ implies a quantitative, and often a qualitative, reaction on the blood; and thus alters the nutrition of all other organs. Such physiological correlations are exemplified in the many differences which accompany difference of sex. That the minor sexual peculiarities are brought about by the physiological actions and reactions, is shown both by the fact that they are commonly but faintly marked until the fundamentally distinctive organs are developed, and that when the development of these is prevented, the minor sexual peculiarities do not arise. No further proof is, I think, needed, that in any individual organism or its descendants, a new external action must, besides the primary internal change which it works, work many secondary changes, as well as tertiary changes still more multiplied. That tendency towards greater heterogeneity which is given to an organism by disturbing its environment, is helped by the tendency which every modification has to produce other modifications—modifications which must become more numerous in proportion as the organism becomes more complex. Lastly, among the indirect and involved manifestations of this tendency, we must not omit the innumerable small irregularities of structure which result from the crossing of dissimilarly-modified individuals. It was shown ( §§ 89, 90) that what are called "spontaneous variations," are interpretable as results of miscellaneously compounding the changes wrought in different lines of ancestors by different conditions of life. These still more complex and multitudinous effects so produced, are further illustrations of the multiplication of effects.
Equally in the aggregate of individuals constituting a species, does multiplication of effects become the continual cause of increasing multiformity. The lapse of a species into divergent varieties, initiates fresh combinations of forces tending to work further divergences. The new varieties compete with the parent species in new ways; and so add new elements to its circumstances. They modify somewhat the conditions of other species existing in their habitat, or in the habitat they have invaded; and the modifications wrought in such other species become additional sources of influence. The Flora and Fauna of every region are united by their entangled relations into a whole, of which no part can be affected without affecting the rest. Hence, each differentiation in a local assemblage of species, becomes the cause of further differentiations.
§ 156. One of the universal principles to which we saw that the re-distribution of matter and motion conforms, is that in any aggregate made up of mixed units, incident forces produce segregation—separate unlike units and bring together like units; and it was shown that the increasing integration and definiteness which characterizes each part of an evolving organic aggregate, as of every other aggregate, results from this (First Principles, § 166). It remains here to say that while the actions and reactions between organisms and their changing environments, add to the heterogeneity of organic structures, they also give to the heterogeneity this growing distinctness. At first sight the reverse might be inferred. It might be argued that any new set of effects wrought in an organism by some new set of external forces, must tend more or less to obliterate the effects previously wrought—must produce confusion or indefiniteness. A little consideration, however, will dissipate this impression.
Doubtless the condition under which alone increasing definiteness of structure can be acquired by any part of an organism, either in an individual or in successive generations, is that such part shall be exposed to some set of tolerably-constant forces; and doubtless, continual change of circumstances interferes with this. But the interference can never be considerable. For the pre-existing structure of an organism prevents it from living under any new conditions except such as are congruous with the fundamental characters of its organization—such as subject its essential organs to actions substantially the same as before. Great changes must kill it. Hence, it can continuously expose itself and its descendants, only to those moderate changes which do not destroy the general harmony between the aggregate of incident forces and the aggregate of its functions. That is, it must remain under influences calculated to make greater the definiteness of the chief differentiations already produced. If, for example, we set out with an animal in which a rudimentary vertebral column with its attached muscular system has been established; it is clear that the mechanical arrangements have become thereby so far determined, that subsequent modifications are extremely likely, if not certain, to be consistent with the production of movement by the actions of muscles on a flexible central axis. Hence, there will continue a general similarity in the play of forces to which the flexible central axis is subject; and so, notwithstanding the metamorphoses which the vertebrate type undergoes, there will be a maintenance of conditions favourable to increasing definiteness and integration of the vertebral column. Moreover, this maintenance of such conditions becomes secure in proportion as organization advances. Each further complexity of structure, implying some further complexity in the relations between an organism and its environment, must tend to specialize the actions and reactions between it and its environment—must tend to increase the stringency with which it is restrained within such environments as admit of those special actions and reactions for which its structure fits it; that is, must further guarantee the continuance of those actions and reactions to which its essential organs respond, and therefore the continuance of the segregating process.
How in each species, considered as an aggregate of individuals, there must arise stronger and stronger contrasts among those divergent varieties which result from the instability of the homogeneous and the multiplication of effects, need only be briefly indicated. It has already been shown (First Principles, § 166), that in conformity to the universal law that mixed units are segregated by like incident forces, there are produced increasingly-definite distinctions among varieties, wherever there occur definitely-distinguished sets of conditions to which the varieties are respectively subject.
§ 157. Probably in the minds of some, the reading of this chapter has been accompanied by a running commentary, to the effect that the argument proves too much. The apparent implication is, that the passage from an indefinite, incoherent homogeneity to a definite, coherent heterogeneity in organic aggregates, must have been going on universally; whereas we find that in many cases there has been persistence without progression. This apparent implication, however, is not a real one.
For though every environment on the Earth's surface undergoes changes; and though usually the organisms which each environment contains, cannot escape certain resulting new influences; yet occasionally such new influences are escaped, by the survival of species in the unchanged parts of their habitats, or by their spread into neighbouring habitats which the change has rendered like their original habitats, or by both. Any alteration in the temperature of a climate or its degree of humidity, is unlikely to affect simultaneously the whole area occupied by a species; and further, it can scarcely fail to happen that the addition or subtraction of heat or moisture, will give to a part of some adjacent area, a climate like that to which the species has been habituated. If, again, the circumstances of a species are modified by the intrusion of some foreign kind of plant or animal, it follows that since the intruders will probably not spread throughout its whole habitat, the species will, in one or more localities, remain unaffected by them. Especially among marine creatures, must there frequently occur cases in which modifying causes are continually eluded. Comparatively uniform as are the physical conditions to which the sea exposes its inhabitants, it becomes possible for such of them as live on widely-diffused food, to be widely distributed; and wide distribution generally prevents the members of a species from being all subject to the same cause. Our commonest cirriped, for instance, subsisting on minute creatures everywhere dispersed through the water; needing only to have some firm surface on which to build up its shell; and in scarcely any danger from surrounding animals; is able to exist on shores so widely remote from one another, that nearly every change in the incident forces must fall within narrower areas than that which the species occupies. Nearly always, therefore, a portion of the species will survive unmodified. Its easily-transported germs will take possession of such new habitats as have been rendered fitter by the change that has unfitted some parts of its original habitat. Hence, on successive occasions, while some parts of the species are slightly transformed, another part may continually escape transformation by migrating hither and thither, where the simple conditions needed for its existence recur in nearly the same combinations as before. And it will so become possible for it to survive, with insignificant structural changes, throughout long geologic periods.
§ 158. The results to which we find ourselves led, are these.
In subordination to the different amounts and kinds of forces to which its different parts are exposed, every individual organic aggregate, like all other aggregates, tends to pass from its original indistinct simplicity towards a more distinct complexity. Unless we deny the persistence of force, we must admit that the lapse of an organism's structure from an indefinitely homogeneous to a definitely heterogeneous state, must be cumulative in successive generations, if the forces causing it continue to act. And for the like reasons, the increasing assemblage of individuals arising from a common stock, is also liable to lose its original uniformity; and, in successive generations, to grow more pronounced in its multiformity.
These changes, which would go to but a comparatively small extent were organisms exposed to constant external conditions, are kept up by the continual changes in external conditions, produced by astronomic, geologic, meteorologic, and organic agencies: the average result being, that on previous complications wrought by previous incident forces, new complications are continually superposed by new incident forces. And hence simultaneously arises increasing heterogeneity in the structures of individuals, in the structures of species, and in the structures of the Earth's Flora and Fauna.
But while, in very many or in most cases, the ever-changing incidence of forces is ever adding to the complexity of organisms, and to the complexity of the organic world as a whole; it does this only where its action cannot be eluded. And since, by migration, it is possible for a species to keep itself under conditions that are tolerably constant, there must be a proportion of cases in which greater heterogeneity of structure is not to be expected.
To show, however, that there must arise a certain average tendency to the production of greater heterogeneity is not sufficient. Aggregates might be rendered more heterogeneous by changing incident forces, without having given to them that kind of heterogeneity required for carrying on life. Hence it remains now to inquire how the production and maintenance of this kind of heterogeneity is insured.