Essays · Herbert Spencer
TRANSCENDENTAL PHYSIOLOGY.
Similarly with respect to nutriment. There are _entozoa_ which, living in the insides of other animals, and being constantly bathed by nutritive fluids, absorb a sufficiency through their outer surfaces; and so have no need of stomachs, and do not possess them. But all other animals, inhabiting media that are not in themselves nutritive, but only contain masses of food here and there, must have appliances by which these masses of food may be utilized. Evidently mere external contact of a solid organism with a solid portion of nutriment, could not result in the absorption of it in any moderate time, if at all. To effect absorption, there must be both a solvent or macerating action, and an extended surface fit for containing and imbibing the dissolved products: there must be a digestive cavity. Thus, given the ordinary conditions of animal life, and the possession of stomachs by all creatures living under these conditions may be deductively known.
Carrying out the train of reasoning still further, we may infer the existence of a vascular system or something equivalent to it, in all creatures of any size and activity. In a comparatively small inert animal, such as the hydra, which consists of little more than a sac having a double wall--an outer layer of cells forming the skin, and an inner layer forming the digestive and absorbent surface--there is no need for a special apparatus to diffuse through the body the aliment taken up; for the body is little more than a wrapper to the food it encloses. But where the bulk is considerable, or where the activity is such as to involve much waste and repair, or where both these characteristics exist, there is a necessity for a system of blood-vessels. It is not enough that there be adequately extensive surfaces for absorption and aeration; for in the absence of any means of conveyance, the absorbed elements can be of little or no use to the organism at large. Evidently there must be channels of communication. When, as in the _Medusæ_, we find these channels of communication consisting simply of branched canals opening out of the stomach and spreading through the disk, we may know, _a priori_, that such creatures are comparatively inactive; seeing that the nutritive liquid thus partially distributed throughout their bodies is crude and dilute, and that there is no efficient appliance for keeping it in motion. Conversely, when we meet with a creature of considerable size which displays much vivacity, we may know, _a priori_, that it must have an apparatus for the unceasing supply of concentrated nutriment, and of oxygen, to every organ--a pulsating vascular system.
It is manifest, then, that setting out from certain known fundamental conditions to vital activity, we may deduce from them sundry of the chief characteristics of organized bodies. Doubtless these known fundamental conditions have been inductively established. But what we wish to show is that, given these inductively-established primary facts in physiology, we may with safety draw certain general deductions from them. And, indeed, the legitimacy of such deductions, though not formally acknowledged, is practically recognized in the convictions of every physiologist, as may be readily proved. Thus, were a physiologist to find a creature exhibiting complex and variously co-ordinated movements, and yet having no nervous system; he would be less astonished at the breach of his empirical generalization that all such creatures have nervous systems, than at the disproof of his unconscious deduction that all creatures exhibiting complex and variously co-ordinated movements must have an "internuncial" apparatus by which the co-ordination may be effected. Or were he to find a creature having blood rapidly circulated and rapidly aerated, but yet showing a low temperature, the proof so afforded that active change of matter is not, as he had inferred from chemical data, the cause of animal heat, would stagger him more than would the exception to a constantly-observed relation. Clearly, then, the _a priori_ method already plays a part in physiological reasoning. If not ostensibly employed as a means of reaching new truths, it is at least privately appealed to for confirmation of truths reached _a posteriori_.
In an essay on "Progress: its Law and Cause," elsewhere published,[8] we have endeavoured to show that the transformation of the homogeneous into the heterogeneous, in which all progress, organic or other, essentially consists, is consequent on the production of many effects by one cause--many changes by one force. Having pointed out that this is a law of all things, we proceeded to show deductively that the multiform evolutions of the homogeneous into the heterogeneous--astronomic, geologic, ethnologic, social, &c.,--were explicable as consequences. And though in the case of organic evolution, lack of data disabled us from specifically tracing out the progressive complication as due to the multiplication of effects; yet, we found sundry indirect evidences that it was so. Now in so far as this conclusion, that organic evolution results from the decomposition of each expended force into several forces, was inferred from the general law previously pointed out, it was an example of deductive physiology. The particular was concluded from the universal. We here propose in the first place to show, that there is another general truth closely connected with the above; and in common with it underlying explanations of all progress, and therefore the progress of organisms--a truth which may indeed be considered as taking precedence of it in respect of time, if not in respect of generality. This truth is, that _the condition of homogeneity is a condition of unstable equilibrium_.
The phrase _unstable equilibrium_ is one used in mechanics to express a balance of forces of such kind, that the interference of any further force, however minute, will destroy the arrangement previously existing, and bring about a different arrangement. Thus, a stick poised on its lower end is in unstable equilibrium: however exactly it may be placed in a perpendicular position, as soon as it is left to itself it begins, at first imperceptibly and then visibly, to lean on one side, and with increasing rapidity falls into another position. Conversely, a stick suspended from its upper end is in stable equilibrium: however much disturbed, it will return to the same position. Our meaning is, then, that the state of homogeneity, like the state of the stick poised on its lower end, is one that cannot be maintained; and that hence results the first step in its gravitation towards the heterogeneous. Let us take a few illustrations.
Of mechanical ones the most familiar is that of the scales. If accurately made and not clogged by dirt or rust, a pair of scales cannot be perfectly balanced: eventually one scale will descend and the other ascend--they will assume a heterogeneous relation. Again, if we sprinkle over the surface of a liquid a number of equal-sized particles, having an attraction for one another, they will, no matter how uniformly distributed, by and by concentrate irregularly into groups. Were it possible to bring a mass of water into a state of perfect homogeneity--a state of complete quiescence, and exactly equal density throughout--yet the radiation of heat from neighbouring bodies, by affecting differently its different parts, would soon produce inequalities of density and consequent currents; and would so render it to that extent heterogeneous. Take a piece of red-hot matter, and however evenly heated it may at first be, it will quickly cease to be so: the exterior, cooling faster than the interior, will become different in temperature from it. And the lapse into heterogeneity of temperature, so obvious in this extreme case, is ever taking place more or less in all cases. The actions of chemical forces supply other illustrations. Expose a fragment of metal to air or water, and in course of time it will be coated with a film of oxide, carbonate, or other compound: its outer parts will become unlike its inner parts. Thus, every homogeneous aggregate of matter tends to lose its balance in some way or other--either mechanically, chemically, thermally or electrically; and the rapidity with which it lapses into a non-homogeneous state is simply a question of time and circumstances. Social bodies illustrate the law with like constancy. Endow the members of a community with equal properties, positions, powers, and they will forthwith begin to slide into inequalities. Be it in a representative assembly, a railway board, or a private partnership, the homogeneity, though it may continue in name, inevitably disappears in reality.
The instability thus variously illustrated becomes still more manifest if we consider its rationale. It is consequent on the fact that the several parts of any homogeneous mass are necessarily exposed to different forces--forces which differ either in their kinds or amounts; and being exposed to different forces they are of necessity differently modified. The relations of outside and inside, and of comparative nearness to neighbouring sources of influence, imply the reception of influences which are unlike in quantity or quality or both; and it follows that unlike changes will be wrought in the parts dissimilarly acted upon. The unstable equilibrium of any homogeneous aggregate can thus be shown both inductively and deductively. And now let us consider the bearing of this general truth on the evolution of organisms. The germ of a plant or animal is one of these homogeneous aggregates--relatively homogeneous if not absolutely so--whose equilibrium is unstable. But it has not simply the ordinary instability of homogeneous aggregates: it has something more. For it consists of units which are themselves specially characterized by instability. The constituent molecules of organic matter are distinguished by the feebleness of the affinities which hold their component elements together. They are extremely sensitive to heat, light, electricity, and the chemical actions of foreign elements; that is, they are peculiarly liable to be modified by disturbing forces. Hence then it follows, _a priori_, that a homogeneous aggregate of these unstable molecules will have an excessive tendency to lose its equilibrium. It will have a quite special liability to lapse into a non-homogeneous state. It will rapidly gravitate towards heretogeneity.
Moreover, the process must repeat itself in each of the subordinate groups of organic units which are differentiated by the modifying forces. Each of these subordinate groups, like the original group, must gradually, in obedience to the influences acting on it, lose its balance of parts--must pass from a uniform into a multiform state. And so on continuously. Thus, starting from the general laws of things, and the known chemical attributes of organic matter, we may conclude deductively that the homogeneous germs of organisms have a peculiar proclivity towards a non-homogeneous state; which may be either the state we call decomposition, or the state we call organization. * * * * * At present we have reached a conclusion only of the most general nature. We merely learn that _some_ kind of heterogeneity is inevitable; but as yet there is nothing to tell us _what_ kind. Besides that _orderly_ heterogeneity which distinguishes organisms, there is the _disorderly_ or _chaotic_ heterogeneity, into which a loose mass of inorganic matter lapses; and at present no reason has been given why the homogeneous germ of a plant or animal should not lapse into the disorderly instead of the orderly heterogeneity. But by pursuing still further the line of argument hitherto followed we shall find a reason.
We have seen that the instability of homogeneous aggregates in general, and of organic ones in particular, is consequent on the various ways and degrees in which their constituent parts are exposed to the disturbing forces brought to bear on them: their parts are differently acted upon, and therefore become different. Manifestly, then, a rationale of the special changes which a germ undergoes, must be sought in the particular relations which its several parts bear to each other and to their environment. However it may be masked, we may suspect the fundamental principle of organization to be, that the many like units forming a germ acquire those kinds and degrees of unlikeness which their respective positions entail. Take a mass of unorganized but organizable matter--either the body of one of the lowest living forms, or the germ of one of the higher. Consider its circumstances. It is immersed in water or air; or it is contained within a parent organism. Wherever placed, however, its outer and inner parts stand differently related to surrounding existences--nutriment, oxygen, and the various stimuli. But this is not all. Whether it lies quiescent at the bottom of the water, whether it moves through the water preserving some definite attitude, or whether it is in the inside of an adult; it equally results that certain parts of its surface are more directly exposed to surrounding agencies than other parts--in some cases more exposed to light, heat, or oxygen, and in others to the maternal tissues and their contents. The destruction of its original equilibrium is therefore certain. It may take place in one of two ways. Either the disturbing forces may be such as to overbalance the affinities of the organic elements, in which case there results that chaotic heterogeneity known as decomposition; or, as is ordinarily the case, such changes are induced as do not destroy the organic compounds, but only modify them: the parts most exposed to the modifying forces being most modified. Hence result those first differentiations which constitute incipient organization. From the point of view thus reached, suppose we look at a few cases: neglecting for the present all consideration of the tendency to assume the inherited type.
Note first what appear to be exceptions, as the _Amoeba_. In this creature and its allies, the substance of the jelly-like body remains throughout life unorganized--undergoes no permanent differentiations. But this fact, which seems directly opposed to our inference, is really one of the most significant evidences of its truth. For what is the peculiarity of the Rhizopods, exemplified by the _Amoeba_? They undergo perpetual and irregular changes of shape--they show no persistent relations of parts. What lately formed a portion of the interior is now protruded, and, as a temporary limb, is attached to some object it happens to touch. What is now a part of the surface will presently be drawn, along with the atom of nutriment sticking to it, into the centre of the mass. Thus there is an unceasing interchange of places; and the relations of inner and outer have no settled existence. But by the hypothesis, it is only in virtue of their unlike positions with respect to modifying forces, that the originally-like units of a living mass become unlike. We must not therefore expect any established differentiation of parts in creatures which exhibit no established differences of position in their parts.
This negative evidence is borne out by abundant positive evidence. When we turn from these ever-changing specks of living jelly to organisms having unchanging distributions of substance, we find differences of tissue corresponding to differences of relative position. In all the higher _Protozoa_, as also in the _Protophyta_, we meet with a fundamental differentiation into cell-membrane and cell-contents, answering to that fundamental contrast of conditions implied by the words outside and inside. And on passing from what are roughly classed as unicellular organisms to the lowest of those which consist of aggregated cells, we equally observe the connexion between structural differences and differences of circumstance. In the sponge, permeated throughout by currents of sea-water, the absence of definite organization corresponds with the absence of definite unlikeness of conditions. In the _Thalassicolla_ of Professor Huxley--a transparent, colourless body, found floating passively at the surface of the sea, and consisting essentially of "a mass of cells united by jelly"--there is displayed a rude structure obviously subordinated to the primary relations of centre and surface: in all of its many and important varieties, the parts exhibit a more or less concentric arrangement.
After this primary modification, by which the outer tissues are differentiated from the inner, the next in order of constancy and importance is that by which some part of the outer tissues is differentiated from the rest; and this corresponds with the almost universal fact that some part of the outer tissues is more directly exposed to certain environing influences than the rest. Here, as before, the apparent exceptions are extremely significant. Some of the lowest vegetable organisms, as the _Hematococci_ and _Protococci_, evenly imbedded in a mass of mucus, or dispersed through the Arctic snow, display no differentiations of surface: the several parts of the surface being subjected to no definite contrasts of conditions. The _Thalassicolla_ above mentioned, unfixed, and rolled about by the waves, presents all its sides successively to the same agencies; and all its sides are alike. A ciliated sphere like the _Volvox_ has no parts of its periphery unlike other parts; and it is not to be expected that it should have; seeing that as it revolves in all directions, it does not, in traversing the water, permanently expose any part to special conditions. But when we come to creatures that are either fixed, or while moving, severally preserve a definite attitude, we no longer find uniformity of surface. The gemmule of a Zoophyte, which during its locomotive stage is distinguishable only into outer and inner tissues, no sooner takes root than its upper end begins to assume a different structure from its lower. The free-swimming embryo of an aquatic annelid, being ovate and not ciliated all over, moves with one end foremost; and its differentiations proceed in conformity with this contrast of circumstances.
The principle thus displayed in the humbler forms of life, is traceable during the development of the higher; though being here soon masked by the assumption of the hereditary type, it cannot be traced far. Thus the "mulberry-mass" into which a fertilized ovum of a vertebrate animal first resolves itself, soon begins to exhibit a difference between the outer and inner parts answering to the difference of circumstances. The peripheral cells, after reaching a more complete development than the central ones, coalesce into a membrane enclosing the rest; and then the cells lying next to these outer ones become aggregated with them, and increase the thickness of the germinal membrane, while the central cells liquefy. Again, one part of the germinal membrane presently becomes distinguishable as the germinal spot; and without asserting that the cause of this is to be found in the unlike relations which the respective parts of the germinal membrane bear to environing influences, it is clear that we have in these unlike relations an element of disturbance tending to destroy the original homogeneity of the germinal membrane. Further, the germinal membrane by and by divides into two layers, internal and external; the one in contact with the liquefied interior part or yelk, the other exposed to the surrounding fluids: this contrast of circumstances being in obvious correspondence with the contrast of structures which follows it. Once more, the subsequent appearance of the vascular layer between these mucous and serous layers, as they have been named, admits of a like interpretation. And in this and the various complications which now begin to show themselves, we may see coming into play that general law of the multiplication of effects flowing from one cause, to which the increase of heterogeneity was elsewhere ascribed.[9]
Confining our remarks, as we do, to the most general facts of development, we think that some light is thus thrown on them. That the unstable equilibrium of a homogeneous germ must be destroyed by the unlike exposure of its several units to surrounding influences, is an _a priori_ conclusion. And it seems also to be an _a priori_ conclusion, that the several units thus differently acted upon, must either be decomposed, or must undergo such modifications of nature as may enable them to live in the respective circumstances they are thrown into: in other words--_they must either die or become adapted to their conditions_. Indeed, we might infer as much without going through the foregoing train of reasoning. The superficial organic units (be they the outer cells of a "mulberry-mass," or be they the outer molecules of an individual cell) must assume the function which their position necessitates; and assuming this function, must acquire such character as performance of it involves. The layer of organic units lying in contact with the yelk must be those through which the yelk is absorbed; and so must be adapted to the absorbent office. On this condition only does the process of organization appear possible. We might almost say that just as some race of animals, which multiplies and spreads into divers regions of the earth, becomes differentiated into several races through the adaptation of each to its conditions of life; so, the originally homogeneous population of cells arising in a fertilized germ-cell, becomes divided into several populations of cells that grow unlike in virtue of the unlikeness of their circumstances.
Moreover, it is to be remarked in further proof of our position, that it finds its clearest and most abundant illustrations where the conditions of the case are the simplest and most general--where the phenomena are the least involved: we mean in the production of individual cells. The structures which presently arise round nuclei in a blastema, and which have in some way been determined by those nuclei as centres of influence, evidently conform to the law; for the parts of the blastema in contact with the nuclei are differently conditioned from the parts not in contact with them. Again, the formation of a membrane round each of the masses of granules into which the endochrome of an alga-cell breaks up, is an instance of analogous kind. And should the recently-asserted fact that cells may arise round vacuoles in a mass of organizable substance, be confirmed, another good example will be furnished; for such portions of substance as bound these vacant spaces are subject to influences unlike those to which other portions of the substance are subject. If then we can most clearly trace this law of modification in these primordial processes, as well as in those more complex but analogous ones exhibited in the early changes of an ovum, we have strong reason for thinking that the law is fundamental.
But, as already more than once hinted, this principle, understood in the simple form here presented, supplies no key to the detailed phenomena of organic development. It fails entirely to explain generic and specific peculiarities; and leaves us equally in the dark respecting those more important distinctions by which families and orders are marked out. Why two ova, similarly exposed in the same pool, should become the one a fish, and the other a reptile, it cannot tell us. That from two different eggs placed under the same hen, should respectively come forth a duckling and a chicken, is a fact not to be accounted for on the hypothesis above developed. Here we are obliged to fall back upon the unexplained principle of hereditary transmission. The capacity possessed by an unorganized germ of unfolding into a complex adult which repeats ancestral traits in minute details, and that even when it has been placed in conditions unlike those of its ancestors, is a capacity impossible for us to understand. That a microscopic portion of seemingly structureless matter should embody an influence of such kind, that the resulting man will in fifty years after become gouty or insane, is a truth which would be incredible were it not daily illustrated.
But though the _manner_ in which hereditary likeness, in all its complications, is conveyed, is a mystery passing comprehension, it is quite conceivable that it is conveyed in subordination to the law of adaptation above explained; and we are not without reasons for thinking that it is so. Various facts show that acquired peculiarities resulting from the adaptation of constitution to conditions, are transmissible to offspring. Such acquired peculiarities consist of differences of structure or composition in one or more of the tissues. That is to say, of the aggregate of similar organic units composing a germ, the group going to the formation of a particular tissue, will take on the special character which the adaptation of that tissue to new circumstances had produced in the parents. We know this to be a general law of organic modifications. Further, it is the _only_ law of organic modifications of which we have any evidence.[10] It is not impossible then that it is the universal law; comprehending not simply those minor modifications which offspring inherit from recent ancestry, but comprehending also those larger modifications distinctive of species, genus, order, class, which they inherit from antecedent races of organisms. And thus it _may be_ that the law of adaptation is the sole law; presiding not only over the differentiation of any race of organisms into several races, but also over the differentiation of the race of organic units composing a germ, into the many races of organic units composing an adult. So understood, the process gone through by every unfolding organism will consist, partly in the direct adaptation of its elements to their several circumstances, and partly in the assumption of characters resulting from analogous adaptations of the elements of all ancestral organisms.
Ideas discussed in this chapter
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