The Principles of Biology, Volume 1 (of 2). Spencer Herbert
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While then it is certain, a priori, that there cannot be growth in the absence of such substances as those of which an organism consists; and while it is equally certain that the amount of growth must primarily be governed by the supply of these substances; it is not less certain that extra supply will not produce extra growth, beyond a point very soon reached. Deduction shows to be necessary, as induction makes familiar, the truths that the value of food for purposes of growth depends not on the quantity of the various organizable materials it contains, but on the quantity of the material most needed; that given a right proportion of materials, the pre-existing structure of the organism limits their availability; and that the higher the structure, the sooner is this limit reached.
§ 46. But why should the growth of every organism be finally arrested? Though the rate of increase may, in each case, be necessarily restricted within a narrow range of variation – though the increment that is possible in a given time, cannot exceed a certain amount; yet why should the increments decrease and finally become insensible? Why should not all organisms, when supplied with sufficient materials, continue to grow as long as they live? To find an answer to this question we must revert to the nature and functions of organic matter.
In the first three chapters of Part I, it was shown that plants and animals mainly consist of substances in states of unstable equilibrium – substances which have been raised to this unstable equilibrium by the expenditure of the forces we know as solar radiations, and which give out these forces in other forms on falling into states of stable equilibrium. Leaving out the water, which serves as a vehicle for these materials and a medium for their changes; and excluding those mineral matters that play either passive or subsidiary parts; organisms are built up of compounds which are stores of force. Thus complex colloids and crystalloids which, as united together, form organized bodies, are the same colloids and crystalloids which give out, on their decomposition, the forces expended by organized bodies. Thus these nitrogenous and carbonaceous substances, being at once the materials for organic growth and the sources of organic energy, it results that as much of them as is used up for the genesis of energy is taken away from the means of growth, and as much as is economized by diminishing the genesis of energy, is available for growth. Given that limited quantity of nutritive matter which the pre-existing structure of an organism enables it to absorb; and it is a necessary corollary from the persistence of force, that the matter accumulated as growth cannot exceed that surplus which remains undecomposed after the production of the required amounts of sensible and insensible motion. This, which would be rigorously true under all conditions if exactly the same substances were used in exactly the same proportions for the production of force and for the formation of tissue, requires, however, to be taken with the qualification that some of the force-evolving substances are not constituents of tissue; and that thus there may be a genesis of force which is not at the expense of potential growth. But since organisms (or at least animal organisms, with which we are here chiefly concerned) have a certain power of selective absorption, which, partially in an individual and more completely in a race, adapts the proportions of the substances absorbed to the needs of the system; then if a certain habitual expenditure of force leads to a certain habitual absorption of force-evolving matters that are not available for growth; and if, were there less need for such matters, the ability to absorb matters available for growth would be increased to an equivalent extent; it follows that the antagonism described does, in the long run, hold even without this qualification. Hence, growth is substantially equivalent to the absorbed nutriment, minus the nutriment used up in action.
This, however, is no answer to the question – why has individual growth a limit? – why do the increments of growth bear decreasing ratios to the mass and finally come to an end? The question is involved. There are more causes than one why the excess of absorbed nutriment over expended nutriment must, other things equal, become less as the size of the animal becomes greater. In similarly-shaped bodies the masses, and therefore the weights, vary as the cubes of the dimensions; whereas the powers of bearing the stresses imposed by the weights vary as the squares of the dimensions. Suppose a creature which a year ago was one foot high, has now become two feet high, while it is unchanged in proportions and structure; what are the necessary concomitant changes? It is eight times as heavy; that is to say, it has to resist eight times the strain which gravitation puts upon certain of its parts; and when there occurs sudden arrest of motion or sudden genesis of motion, eight times the strain is put upon the muscles employed. Meanwhile the muscles and bones have severally increased their abilities to bear strains in proportion to the areas of their transverse sections, and hence have severally only four times the tenacity they had. This relative decrease in the power of bearing stress does not imply a relative decrease in the power of generating energy and moving the body; for in the case supposed the muscles have not only increased four times in their transverse sections but have become twice as long, and will therefore generate an amount of energy proportionate to their bulk. The implication is simply that each muscle has only half the power to withstand those shocks and strains which the creature's movements entail; and that consequently the creature must be either less able to bear these, or must have muscles and bones having relatively greater transverse dimensions: the result being that greater cost of nutrition is inevitably caused and therefore a correlative tendency to limit growth. This necessity will be seen still more clearly if we leave out the motor apparatus, and consider only the forces required and the means of supplying them. For since, in similar bodies, the areas vary as the squares of the dimensions, and the masses vary as the cubes; it follows that the absorbing surface has become four times as great, while the weight to be moved by the matter absorbed has become eight times as great. If then, a year ago, the absorbing surface could take up twice as much nutriment as was needed for expenditure, thus leaving one-half for growth, it is now able only just to meet expenditure, and can provide nothing for growth. However great the excess of assimilation over waste may be during the early life of an active organism, we see that because a series of numbers increasing as the cubes, overtakes a series increasing as the squares, even though starting from a much smaller number, there must be reached, if the organism lives long enough, a point at which the surplus assimilation is brought down to nothing – a point at which expenditure balances nutrition – a state of moving equilibrium. The only way in which the difficulty can be met is by gradual re-organization of the alimentary system; and, in the first place, this entails direct cost upon the organism, and, in the second place, indirect cost from the carrying of greater weight: both tending towards limitation. There are two other varying relations between degrees of growth and amounts of expended force; one of which conspires with the last, while the other conflicts with it. Consider, in the first place, the cost at which nutriment is distributed through the body and effete matters removed from it. Each increment of growth being added at the periphery of the organism, the force expended in the transfer of matter must increase in a rapid progression – a progression more rapid than that of the mass. But as the dynamic expense of distribution is small compared with the dynamic value of the materials distributed, this item in the calculation is unimportant. Now consider, in the second place, the changing proportion between production and loss of heat. In similar organisms the quantities of heat generated by similar actions going on throughout their substance, must increase as the masses, or as the cubes of the dimensions. Meanwhile, the surfaces from which loss of heat takes place, increase only as the squares of the dimensions. Though the loss of heat does not therefore increase only as the squares of the dimensions, it certainly increases at a smaller rate than the cubes. And to the extent that augmentation of mass results in a greater retention of heat, it effects an economization of force. This advantage is not, however, so important as at first appears. Organic heat is a concomitant of organic action, and is so abundantly produced during action that the loss of it is then usually of no consequence: