Even on the other hypotheses, those of the formation of discrete suns and planets by the aggregation of meteoric dust, and the compensatory dispersal of such dust by radiation pressure, apparently insurmountable difficulties arise. All such hypotheses as we have indicated assume material substance and modes of energy-transformation similar to those that we study in laboratory processes, and all such hypotheses involve the notion of the degradation of energy. So long as we suppose that all cosmic processes are transformations of extended systems of material substances we must assume that energy is dissipated at every stage of the transformation, and whenever we assume this we admit that the processes are irreversible ones, and that the material universe as a whole tends towards a condition of inertia. Yet this, we see, cannot be true, for the universe teems with diversity. Is the progress towards the ultimate state of inertia an asymptotic one, as Ward suggests? This does not help us, since all that the suggestion does is to misapply a mathematical device of service only in the treatment of the problems for which it was developed. Somewhere or other, it has been said, the second law of thermodynamics must be evaded in our universe.
How can it be evaded? That movement or progress which we call inorganic is a movement of energy-transformations in one direction—towards their cessation. It is a movement which we can easily reverse in imagination. A cigarette consumed by a smoker represents the downfall of energy: the cellulose and oils of the tobacco burn with the liberation of heat, and the formation of water, carbon dioxide, and some soot; and this is what happens when potential energy contained in an organised substance becomes converted into kinetic energy. Now, the opposite process can clearly be conceived—it can even be pictured. If we make a kinematographic record of the smoking of the cigarette and then reverse the direction of motion of the film, we shall see the particles of soot recombining to form the substance of the cigarette, and we can imagine the concomitant combination of the water, carbon dioxide, and other substances formed during the combustion with the absorption of kinetic energy. This is not a mere analogy, for the same reversal of ordinary chemical happening occurs whenever a green plant builds up starch from the water and carbon dioxide of the atmosphere and it also occurs whenever a chemical synthesis of an “organic” compound, like that of urea by Wöhler, or that of the sugars by Fischer, is brought about in the laboratory. In all such syntheses the experimenter reverses the direction of inorganic chemical happening. He may cause endothermic chemical reactions, reactions accompanied by the absorption of available energy, to take place, and in these kinetic energy becomes transformed into potential energy. All the syntheses of organic compounds so complacently instanced by mechanistic biologists and chemists as indicative of the lack of distinction between the organic and the inorganic point to no such conclusion. Sugar is built up in the cells of the green plant from the inorganic compounds, water, and carbon dioxide, and is therefore a compound prepared by life—that of the plant organism. But sugar may also be built up in the laboratory from inorganic compounds, which may further have been synthesised by the chemist from their elements. Does this destroy the distinction between compounds formed by the agency of the organism and those formed by inorganic agencies? Obviously it does not, for in the green plant the sugar was formed as the result of the vital agency of the living chlorophyllian cell, while in the laboratory it was built up because of the intelligence of the experimenter. Apart from this intelligence or vital agency, the series of chemical transformations beginning with the elements carbon, oxygen, and hydrogen, and ending with the substance sugar, would not have occurred. We have no right to say, therefore, that such syntheses destroy the distinction between the organic and the inorganic. What they do indicate is the distinction between the tendency expressed by the second law of thermo-dynamics (inorganic processes), and those that occur as the result of direction conferred upon processes taken as a whole, either by the vital agency of the living cell, or by the intelligence of man (vital processes).
The direction, therefore, that may be conferred on a series of physico-chemical processes is what we must understand by the “vital impetus” of Bergson, or the “entelechy” of Driesch.
It must be admitted that it is difficult to describe more precisely than we have done above what is meant by these terms. It is with very much the same embarrassment that is experienced by the physicist when he has to apply the concepts of mass and inertia, in their eighteenth-century meaning, to his description of an universe in terms of electro-magnetic theory, that we seek to describe the modern concept of entelechy. Yet the physicist has had to make this step forward, and the same adventure awaits the biologist if the speculative side of his science is to make further progress, and if he is disinclined to make his science an appendage of physics and chemistry. Entelechy does not correspond to the eighteenth-century notion of a “vital force,” or to the “soul” of Descartes, as the writer of a book on evolutionary biology seems to suggest. It is a concept which is forced upon us mainly because of the failure of mechanistic hypotheses of the organism. If our physical analysis of the behaviour of the developing embryo, or the evolving race or stock, or the activities of the organism in the midst of an ever-changing environment, or even the reactions of the functioning gland, fail, then we seem to be forced to postulate an elemental agency in nature manifesting itself in the phenomena of the organism, but not in those of inorganic nature. This argument per ignorantium possesses little force to many minds: it makes little appeal to the thinker, or the critic, or the general reader, but it is almost impossible to over-estimate the appeal which it makes to the investigator, as his experience of the phenomena of the organism increases, and as he feels more and more the difficulty of describing in terms of the concepts of physics the activities of the living animal.
We may, however, attempt to illustrate mainly by analogy what is meant by Driesch’s entelechia, a more precise concept than is Bergson’s élan vital. We return to the consideration of the behaviour of the embryo at the close of the process of segmentation. The organism at this stage consists of a number of cells organically in continuity with each other, either by actual protoplasmic filaments or by the apposition of parts of their surfaces, thus constituting “semi-permeable” membranes. These cells are all similar to each other, both structurally and functionally. It does not matter that modern speculations on heredity describe them as unlike in that each contains a different part of the original germ-plasm which had been disintegrated in the process of the division of the ovum and the first few blastomeres; and it does not matter that these hypotheses are compelled to assume that a part of the original germ-plasm remains intact, being destined to form the gonads of the adult animal. These are hypotheses invented to account for the differentiation of the embryo in terms of eighteenth-century physics and chemistry, and they have yet to be supported by experiment before we can accept them as a description of what is to be observed in the processes of nuclear division and segmentation. Further, it is certainly the case that any one cell of the early embryo can give rise to any part of the larva. The segmented embryo is therefore a system of parts, all of which are potentially similar to each other. But actually each of these parts has a different fate in the process of the development of the larva, and this fate depends on what is the fate of the adjacent cells. There is also a plan or design in the development of the embryo—that is, a very definite structure results from this process—and each of the cells shares in the evolution of this design. The system of cells is therefore an harmonious equipotential system. The cells themselves are not the ultimate parts of this system, for each of them is an aggregate of a very great number of substances which are physico-chemically characterised—at least our methods of analysis seem to show that each cell is a mixture of a number of chemical compounds, but we must never forget that it is the dead cell which we thus subject to analysis, and not a living organism. Let us call these supposed chemical constituents of the living cells the elements of the system; then at the beginning of the process of development the latter is composed of elements which are not definitely arranged but which are distributed in an “homogeneous” manner very like the distribution which is effected on shuffling a pack of cards. But as differentiation proceeds, the elements of this system become unequally distributed, and the diversity becomes greater and greater, attaining its maximum when the definitive tissues and organs of the adult become established, just as at the close of a game of bridge the cards acquire a particular arrangement indicative of a very definite plan which was present in the minds of the players shortly after the game began.
Mechanistic biology would seek to explain this transformation of a homogeneous system of elements into a heterogeneous and specific arrangement by the interaction of the elements with each other, and by the reaction of the environment. But, given a homogeneous arrangement of elements capable of interacting with each other, then only one final phase can be supposed to be produced. A mixture of sulphur, carbon dust, copper and iron filings raised suddenly to a high temperature will only interact in one way, and the final phase of the system will depend on the composition of the mixture, on the temperature, and on the conduction of heat into the mixture in the initial stage of heating. A mixture of chloroform and water shaken up in a bottle is at first a “homogeneous” mixture of the particles of the two substances, but under the influence of gravity the liquids separate from each other and form two distinct layers, each of which will contain in solution some of the other liquid. A homogeneous mixture of different substances therefore becomes a heterogeneous arrangement in the inorganic system, as in the organic one, but while we can predict the former one we cannot predict the latter. We can express the result of the combination of the elements of the inorganic mixture as something that depends on chemical and physical potentials, but this is quite impossible in the case of the development of the embryonic system. It is not only that our knowledge of the developmental process is imperfect: the distinction between the two processes of differentiation is a fundamental one. A change in the conditions under which the inorganic system differentiates leads of necessity to a different final phase, but a change in the conditions under which the embryo develops need have no such effect. If some unforeseen occurrence takes place—some artificial interference with the process of segmentation, which could never have been experienced in the racial history of the organism—a regulation by the parts of the embryo occurs, and the final phase of development may be the same as if no interference had been experienced. That which is operating in the development of the embryo is something that is permitting, or suspending, or arranging physico-chemical reactions.
Let us think of the developing embryo merely as an aggregation of substances contained in an inorganic medium: the segmented frog’s egg floating on the water at the surface of a pond is an example. As an inorganic system its fate is determined. Autolysis of the substances in the cells will occur and the proteids will break down with the formation of amido-bodies, while other chemical changes, strictly predictable if our knowledge of organic chemistry were more complete than it is, would also occur. Putrefactive and fermentative bacteria will attack the proteids, fats, and carbohydrates, and in the end our aggregation of chemical substances will become an aggregation of much simpler compounds—water, carbon dioxide, marsh gas, sulphuretted hydrogen, phosphoretted hydrogen, ammonia, nitrates, etc., all of which will dissolve in the water of the pond, or will diffuse into the adjacent atmosphere. But in the living embryo this is not what occurs: an entirely different, and much more complex, arrangement of the chemical substances originally present in the segmented egg, or at least a physical and chemical re-arrangement, is brought about. The entelechy of the developing embryo prevents some reactions from occurring and directs the energy which is potential in the system towards the performance of other reactions.
Two analogies, suggested by Driesch, will perhaps make the rôle of entelechy more clear. A workman, a heap of bricks, some mortar, some food, and some oxygen constitute a system in the physico-chemical sense. From his heap of bricks and mortar the workman may build one of several different kinds of small house, or he may perhaps construct several walls without any definite arrangement, or he may merely convert one “disorderly” heap of bricks and mortar into another “disorderly” heap. In the same way a man, a case of movable types, some food, and some oxygen constitute another system. The initial phase of this system consists of the compositor, his food, and some fifty-odd boxes of types, each of which contains a large number of similar elements. A final phase of the system may be the arrangement of the types to form an epic poem, or a series of dramatic criticisms, or a meaningless jumble of correctly spelt words. In all these cases the same amount of energy was expended: the bricklayer used up the same quantity of food and oxygen and excreted the same quantities of water, carbon dioxide, and urea, whether he made a house, or a small chimney, or a heap of bricks without architectural arrangement. The system of bricks and mortar acquired during the process of differentiation a gradually increasing complexity; while in the case of the type-setting the diversity of arrangement acquired in the final phases may be of a very high order. Yet the intelligent mind of the worker remained in either case unchanged.
Let us consider further a man walking along the ties, or sleepers, of a railway track. The ties are at variable distances apart, so that the steps of the walker must vary in length, being sometimes closer together, sometimes further apart. The mean step has a definite length and requires the expenditure of a certain amount of energy, and the condition that the man takes sometimes a long step and sometimes a short one does not require that the energy expended on the steps should be more than if every one of them were of the mean length, for the additional energy that is required for the long steps is saved from the short ones. That which operates here is the power of regulation exercised by the walker regarded as a mechanism. There is no purely inorganic process precisely similar to this. It might be thought that the governor of a steam engine did very much the same thing, admitting more steam into the cylinder when the load on the engine increases, and vice versa. But the governor is a mechanism designed to compensate for variations that are given in advance. In the case of the man walking on the railway track, entelechy operates by suspending energetic happening (the muscular contractions of the short steps) when necessary, and allowing it to proceed when necessary. Entelechy itself, whatever it may be, need not be affected by these regulations.
The organism is therefore an aggregation of chemical substances arranged in a typical manner. These substances possess energy in the potential form, capable of undergoing transformation so that they may give rise to other chemical substances—secretions, for instance—or to energy in the kinetic form, that is, the movements of muscles. In the resting organism these transformations do not take place: the energy remains potential, so that chemical happening is suspended. In the unfertilised ovum, for instance, nothing happens although all the potentialities of segmentation are contained in the cell. If reactions did occur in consequence of the chemical potentials contained in the substances of the cells, the progress of these would be such as to lead to the formation of substances in which potential energy was minimal, and in which the original energy of the cell would be represented by the un-co-ordinated kinetic energy of the molecules resulting from the breakdown of the substances undergoing the chemical changes. This is not what happens in the differentiation of the ovum: the developing cell forms new substances from those of its inorganic medium similar to the substances of which it is already composed, and then these substances become arranged to produce the specific form of the organism into which the ovum is about to develop.