The Principles of Biology Vol. I/Chapter II.4
CHAPTER IV.
WASTE AND REPAIR.
§ 62. Throughout the vegetal kingdom, the processes of Waste and Repair are comparatively insignificant in their amounts. Though all parts of plants save the leaves, or other parts which are green, give out carbonic acid; yet this carbonic acid, assuming it to indicate consumption of tissue, or rather of the protoplasm contained in the tissue, indicates but a small consumption. Of course if there is little waste there can be but little repair—that is, little of the interstitial repair which restores the integrity of parts worn by functional activity. Nor, indeed, is there displayed by plants in any considerable degree, if at all, that other species of repair which consists in the restoration of lost or injured organs. Torn leaves and the shoots that are shortened by the pruner, do not reproduce their missing parts; and though when the branch of a tree is cut off close to the trunk, the place is in course of years covered over, it is not by any reparative action in the wounded surface but by the lateral growth of the adjacent bark. Hence, without saying that Waste and Repair do not go on at all in plants, we may fitly pass them over as of no importance.
There are but slight indications of waste in those lower orders of animals which, by their comparative inactivity, show themselves least removed from vegetal life. Actiniæ kept in an aquarium, do not appreciably diminish in bulk from prolonged abstinence. Even fish, though much more active than most other aquatic creatures, appear to undergo but little loss of substance when kept unfed during considerable periods. Reptiles, too, maintaining no great temperature, and passing their lives mostly in a state of torpor, suffer but little diminution of mass by waste. When, however, we turn to those higher orders of animals which are active and hot-blooded, we see that waste is rapid: producing, when unchecked, a notable decrease in bulk and weight, ending very shortly in death. Besides finding that waste is inconsiderable in creatures which produce but little insensible and sensible motion, and that it becomes conspicuous in creatures which produce much insensible and sensible motion; we find that in the same creatures there is most waste when most motion is generated. This is clearly proved by hybernating animals. "Valentin found that the waking marmot excreted in the average 75 times more carbonic acid, and inhaled 41 times more oxygen than the same animal in the most complete state of hybernation. The stages between waking and most profound hybernation yielded intermediate figures. A waking hedgehog yielded about 20.5 times more carbonic acid, and consumed 18.4 times more oxygen than one in the state of hybernation."[1] If we take these quantities of absorbed oxygen and excreted carbonic acid, as indicating something like the relative amounts of consumed organic substance, we see that there is a striking contrast between the waste accompanying the ordinary state of activity, and the waste accompanying complete quiescence and reduced temperature. This difference is still more definitely shown by the fact, that the mean daily loss from starvation in rabbits and guinea-pigs, bears to that from hybernation, the proportion of 18.3:1. Among men and domestic animals, the relation between degree of waste and amount of expended energy, though one respecting which there is little doubt, is less distinctly demonstrable; since waste is not allowed to go on uninterfered with. We have, however, in the lingering lives of invalids who are able to take scarcely any nutriment but are kept warm and still, an illustration of the extent to which waste diminishes as the expenditure of energy declines.
Besides the connexion between the waste of the organism as a whole and the production of sensible and insensible motion by the organism as a whole, there is a traceable connexion between the waste of special parts and the activities of such special parts. Experiments have shown that "the starving pigeon daily consumes in the average 40 times more muscular substance that the marmot in the state of torpor, and only 11 times more fat, 33 times more of the tissue of the alimentary canal, 18.3 times more liver, 15 times more lung, 5 times more skin." That is to say, in the hybernating animal the parts least consumed are the almost totally quiescent motor-organs, and the part most consumed is the hydro-carbonaceous deposit serving as a store of energy; whereas in the pigeon, similarly unsupplied with food but awake and active, the greatest loss takes place in the motor-organs. The relation between special activity and special waste, is illustrated, too, in the daily experiences of all: not indeed in the amount of decrease of the active parts in bulk or weight, for this we have no means of ascertaining; but in the diminished ability of such parts to perform their functions. That legs exerted for many hours in walking and arms long strained in rowing, lose their powers—that eyes become enfeebled by reading or writing without intermission—that concentrated attention, unbroken by rest, so prostrates the brain as to incapacitate it for thinking; are familiar truths. And though we have no direct evidence to this effect, there is little danger in concluding that muscles exercised until they ache or become stiff, and nerves of sense rendered weary or obtuse by work, are organs so much wasted by action as to be partially incompetent.
Repair is everywhere and always making up for waste. Though the two processes vary in their relative rates both are constantly going on. Though during the active, waking state of an animal waste is in excess of repair, yet repair is in progress; and though during sleep repair is in excess of waste, yet some waste is necessitated by the carrying on of certain never-ceasing functions. The organs of these never-ceasing functions furnish, indeed, the most conclusive proofs of the simultaneity of repair and waste. Day and night the heart never stops beating, but only varies in the rapidity and vigour of its beats; and hence the loss of substance which its contractions from moment to moment entail, must from moment to moment be made good. Day and night the lungs dilate and collapse; and the muscles which make them do this must therefore be kept in a state of integrity by a repair which keeps pace with waste, or which alternately falls behind and gets in advance of it to a very slight extent.
On a survey of the facts we see, as we might expect to see, that the progress of repair is most rapid when activity is most reduced. Assuming that the organs which absorb and circulate nutriment are in proper order, the restoration of the body to a state of integrity, after the disintegration consequent on expenditure of energy, is proportionate to the diminution in expenditure of energy. Thus we all know that those who are in health, feel the greatest return of vigour after profound sleep—after complete cessation of motion. We know that a night during which the quiescence, bodily and mental, has been less decided, is usually not followed by that spontaneous overflow of energy which indicates a high state of efficiency throughout the organism. We know, again, that long-continued recumbency, even with wakefulness (providing the wakefulness is not the result of disorder), is followed by a certain renewal of strength; though a renewal less than that which would have followed the greater inactivity of slumber. We know, too, that when exhausted by labour, sitting brings a partial return of vigour. And we also know that after the violent exertion of running, a lapse into the less violent exertion of walking, results in a gradual disappearance of that prostration which the running produced. This series of illustrations conclusively proves that the rebuilding of the organism is ever making up for the pulling down of it caused by action; and that the effect of this rebuilding becomes more manifest, in proportion as the pulling down is less rapid. From each digested meal there is every few hours absorbed into the mass of prepared nutriment circulating through the body, a fresh supply of the needful organic compounds; and from the blood, thus occasionally re-enriched, the organs through which it passes are ever taking up materials to replace the materials used up in the discharge of functions. During activity the reintegration falls in arrear of the disintegration; until, as a consequence, there presently comes a general state of functional languor; ending, at length, in a quiescence which permits the reintegration to exceed the disintegration, and restore the parts to their state of integrity. Here, as wherever there are antagonistic actions, we see rhythmical divergences on opposite sides of the medium state—changes which equilibrate each other by their alternate excesses. (First Principles, §§ 85, 173.)
Illustrations are not wanting of special repair that is similarly ever in progress, and similarly has intervals during which it falls below waste and rises above it. Every one knows that a muscle, or a set of muscles, continuously strained, as by holding out a weight at arm's length, soon loses its power; and that it recovers its power more or less fully after a short rest. The several organs of the special sensations yield us like experiences. Strong tastes, powerful odours, loud sounds, temporarily unfit the nerves impressed by them for appreciating faint tastes, odours, or sounds; but these incapacities are remedied by brief intervals of repose. Vision still better illustrates this simultaneity of waste and repair. Looking at the Sun so affects the eyes that, for a short time, they cannot perceive the things around with the usual clearness. After gazing at a bright light of a particular colour, we see, on turning the eyes to adjacent objects, an image of the complementary colour; showing that the retina has, for the moment, lost the power to feel small amounts of those rays which have strongly affected it. Such inabilities disappear in a few seconds or a few minutes, according to circumstances. And here, indeed, we are introduced to a conclusive proof that special repair is ever neutralizing special waste. For the rapidity with which the eyes recover their sensitiveness, varies with the reparative power of the individual. In youth the visual apparatus is so quickly restored to its state of integrity, that many of these photogenes, as they are called, cannot be perceived. When sitting on the far side of a room, and gazing out of the window against a light sky, a person who is debilitated by disease or advancing years, perceives, on transferring the gaze to the adjacent wall, a momentary negative image of the window—the sash-bars appearing light and the squares dark; but a young and healthy person has no such experience. With a rich blood and vigorous circulation, the repair of the visual nerves after impressions of moderate intensity, is nearly instantaneous.
Function carried to excess may produce waste so great that repair cannot make up for it during the ordinary daily periods of rest; and there may result incapacities of the over-taxed organs, lasting for considerable periods. We know that eyes strained by long-continued minute work lose their power for months or years: perhaps suffering an injury from which they never wholly recover. Brains, too, are often so unduly worked that permanent relaxation fails to restore them to vigour. Even of the motor organs the like holds. The most frequent cause of what is called "wasting palsy," or atrophy of the muscles, is habitual excess of exertion: the proof being that the disease occurs most frequently among those engaged in laborious handicrafts, and usually attacks first the muscles which have been most worked.
There has yet to be noticed another kind of repair—that, namely, by which injured or lost parts are restored. Among the Hydrozoa it is common for any portion of the body to reproduce the rest; even though the rest to be so reproduced is the greater part of the whole. In the more highly-organized Actinozoa the half of an individual will grow into a complete individual. Some of the lower Annelids, as the Nais, may be cut into thirty or forty pieces and each piece will eventually become a perfect animal. As we ascend to higher forms we find this reparative power much diminished, though still considerable. The reproduction of a lost claw by a lobster or crab, is a familiar instance. Some of the inferior Vertebrata also, as lizards, can develop new limbs or new tails, in place of those which have been cut off; and can even do this several times over, though with decreasing completeness. The highest animals, however, thus repair themselves to but a very small extent. Mammals and birds do it only in the healing of wounds; and very often but imperfectly even in this. For in muscular and glandular organs the tissues destroyed are not properly reproduced, but are replaced by tissue of an irregular kind which serves to hold the parts together. So that the power of reproducing lost parts is greatest where the organization is lowest; and almost disappears where the organization is highest. And though we cannot say that in the intermediate stages there is a constant inverse relation between reparative power and degree of organization; yet we may say that there is some approach to such a relation.
§ 63. There is an obvious and complete harmony between the first of the above inductions and the deduction which follows immediately from first principles. We have already seen ( § 23) "that whatever amount of power an organism expends in any shape, is the correlate and equivalent of a power that was taken into it from without." Motion, sensible or insensible, generated by an organism, is insensible motion which was absorbed in producing certain chemical compounds appropriated by the organism under the form of food. As much energy as was required to raise the elements of these complex atoms to their state of unstable equilibrium, is given out in their falls to a state of stable equilibrium; and having fallen to a state of stable equilibrium they can give out no further energy, but have to be got rid of as inert and useless. It is an inevitable corollary "from the persistence of force, that each portion of mechanical or other energy which an organism exerts, implies the transformation of as much organic matter as contained this energy in a latent state;" and that this organic matter in yielding up its latent energy, loses its value for the purposes of life, and becomes waste matter needing to be excreted. The loss of these complex unstable substances must hence be proportionate to the quantity of expended force. Here, then, is the rationale of certain general facts lately indicated. Plants do not waste to any considerable degree, for the obvious reason that the sensible and insensible motions they generate are inconsiderable. Between the small waste, small activity, and low temperature of the inferior animals, the relation is similarly one admitting of a priori establishment. Conversely, the rapid waste of energetic, hot-blooded animals might be foreseen with equal certainty. And not less manifestly necessary is the variation in waste which, in the same organism, attends the variation in the heat and mechanical motion produced.
Between the activity of a special part and the waste of that part, a like relation may be deductively inferred; though it cannot be inferred that this relation is equally definite. Were the activity of every organ quite independent of the activities of other organs, we might expect to trace out this relation distinctly; but since increased activity in any organ or group of organs, as the muscles, necessarily entails increased activity in other organs, as in the heart, lungs, and nervous system, it is clear that special waste and general waste are too much entangled to admit of a definite relation being established between special waste and special activity. We may fairly say, however, that this relation is quite as manifest as we can reasonably anticipate.
§ 64. Deductive interpretation of the phenomena of Repair, is by no means so easy. The tendency displayed by an animal organism, as well as by each of its organs, to return to a state of integrity by the assimilation of new matter, when it has undergone the waste consequent on activity, is a tendency which is not manifestly deducible from first principles; though it appears to be in harmony with them. If in the blood there existed ready-formed units exactly like in kind to those of which each organ consists, the sorting of these units, ending in the union of each kind with already existing groups of the same kind, would be merely a good example of Segregation (First Principles, § 163). It would be analogous to the process by which, from a mixed solution of salts, there are, after an interval, deposited separate masses of these salts in the shape of different crystals. But as already said ( § 54), though the selective assimilation by which the repair of organs is effected, may result in part from an action of this kind, the facts cannot be thus wholly accounted for; since organs are in part made up of units which do not exist as such in the circulating fluids. We must suppose that, as suggested in § 54, groups of compound units have a certain power of moulding adjacent fit materials into units of their own form. Let us see whether there is not reason to think such a power exists.
"The poison of small-pox or of scarlatina," remarks Mr. (now Sir James) Paget, "being once added to the blood, presently affects the composition of the whole: the disease pursues its course, and, if recovery ensue, the blood will seem to have returned to its previous condition: yet it is not as it was before; for now the same poison may be added to it with impunity." ... "The change once effected, may be maintained through life. And herein seems to be a proof of the assimilative force in the blood: for there seems no other mode of explaining these cases than by admitting that the altered particles have the power of assimilating to themselves all those by which they are being replaced: in other words, all the blood that is formed after such a disease deviates from the natural composition, so far as to acquire the peculiarity engendered by the disease: it is formed according to the altered model." Now if the compound molecules of the blood, or of an organism considered in the aggregate, have the power of moulding into their own type the matters which they absorb as nutriment; and if they have the power when their type has been changed by disease, of moulding materials afterwards received into the modified type; may we not reasonably suspect that the more or less specialized molecules of each organ have, in like manner, the power of moulding the materials which the blood brings to them into similarly specialized molecules? The one conclusion seems to be a corollary from the other. Such a power cannot be claimed for the component units of the blood without being conceded to the component units of every tissue. Indeed the assertion of this power is little more than an assertion of the fact that organs composed of specialized units are capable of resuming their structural integrity after they have been wasted by function. For if they do this, they must do it by forming from the materials brought to them, certain specialized units like in kind to those of which they are composed; and to say that they do this, is to say that their component units have the power of moulding fit materials into other units of the same order.
§ 65. What must we say of the ability an organism has to re-complete itself when one of its parts has been cut off? Is it of the same order as the ability of an injured crystal to re-complete itself. In either case new matter is so deposited as to restore the original outline. And if in the case of the crystal we say that the whole aggregate exerts over its parts a force which constrains the newly-integrated molecules to take a certain definite form, we seem obliged, in the case of the organism, to assume an analogous force. If when the leg of a lizard has been amputated there presently buds out the germ of a new one, which, passing through phases of development like those of the original leg, eventually assumes a like shape and structure, we assert only what we see, when we assert that the entire organism, or the adjacent part of it, exercises such power over the forming limb as makes it a repetition of its predecessor. If a leg is reproduced, where there was a leg, and a tail where there was a tail, there seems no alternative but to conclude that the forces around it control the formative processes going on in each part. And on contemplating these facts in connexion with various kindred ones, there is suggested the hypothesis, that the form of each species of organism is determined by a peculiarity in the constitution of its units—that these have a special structure in which they tend to arrange themselves; just as have the simpler units of inorganic matter. Let us glance at the evidences which more especially thrust this conclusion upon us.
A fragment of a Begonia-leaf imbedded in fit soil and kept at an appropriate temperature, will develop a young Begonia; and so small is the fragment which is thus capable of originating a complete plant, that something like a hundred plants may be produced from a single leaf. The friend to whom I owe this observation, tells me that various succulent plants have like powers of multiplication. Illustrating a similar power among animals, we have the often-cited experiments of Trembley on the common polype. Each of the four pieces into which one of these creatures was cut, grew into a perfect individual. In each of these, again, bisection and tri-section were followed by like results. And so with their segments, similarly produced, until as many as fifty polypes had resulted from the original one. Bodies when cut off regenerated heads; heads regenerated bodies; and when a polype had been divided into as many pieces as was practicable, nearly every piece survived and became a complete animal. What, now, is the implication? We cannot say that in each portion of a Begonia-leaf, and in every fragment of a Hydra's body, there exists a ready-formed model of the entire organism. Even were there warrant for the doctrine that the germ of every organism contains the perfect organism in miniature, it still could not be contended that each considerable part of the perfect organism resulting from such a germ, contains another such miniature. Indeed the one hypothesis negatives the other. The implication seems, therefore, to be that the living particles composing one of these fragments, have an innate tendency to arrange themselves into the shape of the organism to which they belong. We must infer that the active units composing a plant or animal of any species have an intrinsic aptitude to aggregate into the form of that species. It seems difficult to conceive that this can be so; but we see that it is so. Groups of units taken from an organism (providing they are of a certain bulk and not much differentiated into special structures) have this power of re-arranging themselves. Manifestly, too, if we are thus to interpret the reproduction of an organism from one of its amorphous fragments, we must thus interpret the reproduction of any minor portion of an organism by the remainder. When in place of its lost claw a lobster puts forth a cellular mass which, while increasing in bulk, assumes the form and structure of the original claw, we cannot avoid ascribing this result to a play of forces like that which moulds the materials contained in a piece of Begonia-leaf into the shape of a young Begonia.
§ 66. As we shall have frequent occasion hereafter to refer to these units which possess the property of arranging themselves into the special structures of the organisms to which they belong; it will be well here to ask by what name they may be most fitly called.
On the one hand, it cannot be in those chemical compounds characterizing organic bodies that this specific property dwells. It cannot be that the molecules of albumin, or fibrin, or gelatine, or other proteid, possess this power of aggregating into these specific shapes; for in such case there would be nothing to account for the unlikenesses of different organisms. If the proclivities of proteid molecules determined the forms of the organisms built up of them or by them, the occurrence of such endlessly varied forms would be inexplicable. Hence what we may call the chemical units are clearly not the possessors of this property.
On the other hand, this property cannot reside in what may be roughly distinguished as the morphological units. The germ of every organism is a minute portion of encased protoplasm commonly called a cell. It is by multiplication of cells that all the early developmental changes are effected. The various tissues which successively arise in the unfolding organism, are primarily cellular; and in many of them the formation of cells continues to be, throughout life, the process by which repair is carried on. But though cells are so generally the ultimate visible components of organisms, that they may with some show of reason be called the morphological units; yet we cannot say that this tendency to aggregate into special forms dwells in them. In many cases a fibrous tissue arises out of a nucleated blastema, without cell-formation; and in such cases cells cannot be regarded as units possessing the structural proclivity. But the conclusive proof that the morphological units are not the building factors in an organism composed of them, is yielded by their independent homologues the so-called unicellular organisms. For each of these displays the power to assume its specific structure. Clearly, if the ability of a multicellular organism to assume its specific structure resulted from the cooperation of its component cells, then a single cell, or the independent homologue of a single cell, having no other to cooperate with, could exhibit no structural traits. Not only, however, do single-celled organisms exhibit structural traits, but these, even among the simplest, are so distinct as to originate classification into orders, genera, and species; and they are so constant as to remain the same from generation to generation.
If, then, this organic polarity (as we might figuratively call this proclivity towards a specific structural arrangement) can be possessed neither by the chemical units nor the morphological units, we must conceive it as possessed by certain intermediate units, which we may term physiological. There seems no alternative but to suppose that the chemical units combine into units immensely more complex than themselves, complex as they are; and that in each organism the physiological units produced by this further compounding of highly compound molecules, have a more or less distinctive character. We must conclude that in each case some difference of composition in the units, or of arrangement in their components, leading to some difference in their mutual play of forces, produces a difference in the form which the aggregate of them assumes.
The facts contained in this chapter form but a small part of the evidence which thrusts this assumption upon us. We shall hereafter find various reasons for inferring that such physiological units exist, and that to their specific properties, more or less unlike in each plant and animal, various organic phenomena are due.
- ↑ In connexion with this matter I add here a statement made by Prof. Foster which it is difficult to understand: "Indeed it has been observed that a dormouse actually gained in weight during a hybernating period; it discharged during this period neither urine nor fæces, and the gain in weight was the excess of oxygen taken in over the carbonic acid given out." (Text-book of Physiology, 6th ed., Part II, page 859.)