# The Principles of Biology Vol. I/Chapter III.13

CHAPTER XIII.
THE CO-OPERATION OF THE FACTORS.
§ 169. Thus the phenomena of Organic Evolution may be interpreted in the same way as the phenomena of all other Evolution. Fully to see this, it will be needful for us to contemplate in their ensemble, the several processes separately described in the four preceding chapters.
If the forces acting on any aggregate remain the same, the changes produced by them will presently reach a limit, at which the outer forces are balanced by the inner forces; and thereafter no further metamorphosis will take place. Hence, that there may be continuous changes of structure in organisms, there must be continuous changes in the incident forces. This condition to the evolution of animal and vegetal forms, we find to be fully satisfied. The astronomic, geologic, and meteorologic changes that have been slowly but incessantly going on, and have been increasing in the complexity of their combinations, have been perpetually altering the circumstances of organisms; and organisms, becoming more numerous in their kinds and higher in their kinds, have been perpetually altering one another's circumstances. Thus, for those progressive modifications upon modifications which organic evolution implies, we find a sufficient cause. The increasing inner changes for which we thus find a cause in the perpetual outer changes, conform, so far as we can trace them, to the universal law of the instability of the ​homogeneous. In organisms, as in all other things, the exposure of different parts to different kinds and amounts of incident forces, has necessitated their differentiation; and, for the like reason, aggregates of individuals have been lapsing into varieties, and species, and genera, and orders. Further, in each type of organism, as in the aggregate of types, the multiplication of effects has continually aided this transition from a more homogeneous to a more heterogeneous state. And yet again, that increasing segregation and concomitant increasing definiteness, associated with the growing heterogeneity of organisms, has been aided by the continual destruction of those which expose themselves to aggregates of external actions markedly incongruous with the aggregates of their internal actions, and the survival of those subject only to comparatively small incongruities. Finally, we have found that each change of structure, superposed on preceding changes, has been a re-equilibration necessitated by the disturbance of a preceding equilibrium. The maintenance of life being the maintenance of a balanced combination of functions, it follows that individuals and species that have continued to live, are individuals and species in which the balance of functions has not been overthrown. Hence survival through successive changes of conditions, implies successive adjustments of the balance to the new conditions.
The actions that are here specified in succession, are in reality simultaneous; and they must be so conceived before organic evolution can be rightly understood. Some aid towards so conceiving them will be given by the annexed table, representing the co-operation of the factors.
§ 170. Respecting this co-operation, it remains only to point out the respective shares of the factors in producing the total result; and the way in which the proportions of their respective shares vary as evolution progresses.
| | | | | | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | on each species: affecting | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | its individuals, | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | immediately through their functions; | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | which, partially in the first generation, and completely in the course of generations, are directly equilibrated with the changed agencies. |
| | | Astronomic changes Geologic changes Meteorologic changes | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | alter the incidence of inorganic forces. | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | on each species: affecting | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | its individuals, | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | immediately through their functions; | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | which have their direct equilibration with the changed agencies, aided by indirect equilibration, through the more frequent survival of those in which the direct equilibration is most rapid. |
| | | Astronomic changes Geologic changes Meteorologic changes | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | alter the incidence of inorganic forces. | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | on each species: affecting | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | its individuals, | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | mediately through the aggregate of individuals; | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | positively—by aiding the multiplication of those whose moving equilibria happen to be most congruous with the changed agencies: thus, in the course of generations, indirect equilibrating certain individua with them. |
| Enemies Competitors Co-operators Prey | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | varying in number | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | alter the incidence of organic forces. | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | on each species: affecting | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | its individuals, | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | mediately through the aggregate of individuals; | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | negatively—by killing those whose moving equilibria are most incongruous with the changed agencies: thus, in the course of generations, indirectly equilibrating each of its surviving individuals with them. |
| Enemies Competitors Co-operators Prey | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | varying in number | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | alter the incidence of organic forces. | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | on each species: affecting | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | its aggregate of individuals, | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | by acting on it in some parts of the habitat more than in others; and thus differentiating the species into local varieties. | by acting on it in some parts of the habitat more than in others; and thus differentiating the species into local varieties. | by acting on it in some parts of the habitat more than in others; and thus differentiating the species into local varieties. |
| Enemies Competitors Co-operators Prey | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | varying in kind | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | alter the incidence of organic forces. | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | on each species: affecting | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | its aggregate of individuals, | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | by acting differently on slightly-unlike individuals in the same locality; | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | and thus causing differentiations of the species into varieties, irrespective of locality. |
| Enemies Competitors Co-operators Prey | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | varying in kind | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | alter the incidence of organic forces. | $\scriptstyle {\left.{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right\}\,$ | on each species: affecting | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | its aggregate of individuals, | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | by acting differently on slightly-unlike individuals in the same locality; | $\scriptstyle {\left\{{\begin{matrix}\ \\\\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \\\ \ \end{matrix}}\right.}$ | and thus causing modification of the species as a whole, by abstracting a certain class of its units. |
​At first, changes in the amounts and combinations of inorganic forces, astronomic, geologic, and meteorologic, were the only causes of the successive modifications; and these changes have continued to be causes. But as, through the diffusion of organisms and consequent differential actions of inorganic forces, there arose unlikenesses among them, producing varieties, species, genera, orders, classes, the actions of organisms on one another became new sources of organic modifications. And as fast as types have multiplied and become more complex, so fast have the mutual actions of organisms come to be more influential factors in their respective evolutions: eventually becoming the chief factors.
Passing from the external causes of change to the internal processes of change entailed by them, we see that these, too, have varied in their proportions: that which was originally the most important and almost the sole process, becoming gradually less important, if not at last the least important. Always there must have been, and always there must continue to be, a survival of the fittest; natural selection must have been in operation at the outset, and can never cease to operate. While yet organisms had small abilities to coordinate their actions, and adjust them to environing actions, natural selection worked almost alone in moulding and remoulding organisms into fitness for their changing environments; and natural selection has remained almost the sole agency by which plants and inferior orders of animals have been modified and developed. The equilibration of organisms that are almost passive, is necessarily effected indirectly, by the action of incident forces on the species as a whole. But along with the evolution of organisms having some activity, there grows up a kind of equilibration which is in part direct. In proportion as the activity increases direct equilibration plays a more important part. Until, when the nervo-muscular apparatus becomes greatly developed, and the power of varying the actions to fit the varying requirements becomes considerable, the share taken by direct equilibration rises into co-ordinate importance or greater importance. As fast as essential faculties multiply, and as ​fast as the number of organs which co-operate in any given function increases, indirect equilibration through natural selection becomes less and less capable of producing specific adaptations; and remains capable only of maintaining the general fitness of constitution to conditions. The production of adaptations by direct equilibration then takes the first place: indirect equilibration serving to facilitate it. Until at length, among the civilized human races, the equilibration becomes mainly direct: the action of natural selection being limited to the destruction of those who are constitutionally too feeble to live, even with external aid. As the preservation of incapables is secured by our social arrangements; and as very few save incarcerated criminals are prevented by their inferiorities from leaving the average number of offspring; it results that survival of the fittest can scarcely at all act in such way as to produce specialities of nature, either bodily or mental. Here the specialities of nature, chiefly mental, which we see produced, and which are so rapidly produced that a few centuries show a considerable change, must be ascribed almost wholly to direct equilibration.