A Bipolar Theory of Living Processes

We do not presume even to attempt to identify the electrical origin and development of each stage in the process of cell division. We do assume, however, that fundamentally, cell division is amenable to the law of bipolarity which, we believe, is the basis for all living processes. Among the facts which might be cited in support of the conceptions outlined above is the variation in the development of fresh water alge whereby cytoplasmic division may be

“Ву exposing the normal forms to lowered temperature, and in certain other ways, it was found that mitotie division may be so modified that although the chromosomes divide the daughternuclei do not separate normally and cytoplasmie division fails. Binucleate cells are thus produced, the two nuclei either remaining separate or fusing into one, which then grows to twice the normal size. In either of these cases the doubling of the nuclear mass is followed by growth of the cytosome to double the normal volume ; and by the continued division of such cells are produced giant filaments which may be reared to maturity, produce gametes of double the normal size, and conjugate to produce correspondingly enlarged zygotes.” (Wilson.) *

Another supporting fact is “the undoubted causal relation between nuclear volume and cytoplasmic growth, 1.е., the Karyoplasmic ratio of R. Hertwig,” to which Wilson alludes as “a fact of great theoretical interest.” Not only the primary cell divisions but also the permanent interrelations of the cells must be related to the bipolar conception. In the higher animals these intercellular relations are principally maintained in the ramifications of the nervous system; in the lower animals these interrelations are probably accomplished by means of the strands of protoplasm. Such connections are well known in many of the simplest plant forms. They exist also in certain animal tissues which are not provided with nerve fibers such as cartilage and bone.

“Plasmodesms or cell-bridges are of general occurrence in the epithelial tissues, where they were first observed in epidermal ‘spine-cells’ (fStachelzellen") and supposed to be spine-like processes from the membrane or cell-periphery (M. Schultze, 1864). Later studies by many observers (Ranvier, Renaut, Pfitzner, Schridde, Kromayer, Cajal, etc.) proved these structures to be protoplasmic inter-cellular bridges, and further showed that they are traversed by fibrillae, which may be followed from one cell to another and even through several cells. The plasma-bridges have since been found in the columnar epithelia generally. Further, it has been shown by a considerable number of observers that the germ-cells in both animals and plants may be connected with the surrounding somatic cells (follicle cells, ее.) by protoplasmic bridges. Plasma-bridges have also been described in the case of embryonic cells of many types and considerable evidence has been produced to show that they may here play an important part in maintaining the unity of the organism.

“The facts thus briefly reviewed have led some important modern writers to accept Heitzmann’s general conclusion almost in its entirety. A. Meyer, for example, expresses the opinion that both the plant and the animal individual is a continuous mass of protoplasm that forms a morphological unit whether it appear in the form of a single cell, a multinucleated cell, or a system of cells.” (Wilson.) 2 (Figs. 56-58.) It would seem probable that the universal presence of these conducting connections between cells would maintain electrical

Fic. 57.—Intercellular bridges (plasmodesms) in animal tissues (А, Flemming; B, Rio-Hortega; C, Ide). A, epithelium of the gilllamelle of salamander-larv:e, deeper layers in horizontal view; B, cells from the mucous membrane of а nasal polypus, fibrille traversing the inter-cellular bridges; C, human cancer-cells. (From Wilson: The Cell in Development and Heredity, New York, 1925, p. 105.) equilibrium among groups of cells and that, therefore, when they were broken by injury or chronie irritation this equilibrium would be destroyed with resultant comparative freedom of growth within one or another group of cells thus producing an unlimited growth among them (cancer). This would aecount for the apparent similarity between neoplasms in vegetable and in animal organisms and for the fact that in the production of each, chronic irritation plays an important rôle.

The symmetry of growth along axial planes would appear to be related to the fundamental polarity of living cells in every stage from the unicellular organisms to man. Fie. 58.—Connections between oocyte and follicle-cells in vertebrates (Retzius). A, in Chimaera; В, Raja; C, the rabbit; D, the pigeon; E, the domestic fowl. The foregoing from earlier stages of the odcyte; F, late stage in Lacerta. (From Wilson: The Cell in Development and Heredity, New York, 1925, p. 337.)

“Child has recently emphasized the general importance of ‘metabolic gradients’ as an expression (if not the actual cause) of functional polarity, which in his view may sometimes be merely a graded difference in the rate of metabolism in the direction of the axis (though it may often be more than this). In support of this he has proved experimentally, by a study of susceptibility to the action of poisons and narcotics, that such gradients undoubtedly exist in the direction of the main axes, both in organisms as a whole and in individual cells.

“Interesting possibilities for the further analysis of physiological polarity are opened by recent experiments. It has been shown that in hydroids the oral region is electronegative as compared with the basal; and also that axial differences of electrical potential similar in type, though different in detail, exist in other animals; and Lund has demonstrated that in the alga Fucus the polarity of the eggs shows a distinct orientation with respect to the electric field. Hyman and Bellamy emphasize the fact that in the various cases studied by them the electrical gradients closely correspond with the metabolic, levels of high metabolic rate being electronegative to those of lower.” (Wilson.) 3

That the nucleus is a primary agent in the constructive processes of cytoplasmic growth culminating in cell division is well recognized. Whatever may be the rôle of the chromosomes in the processes of reproduction, it is known that they bear direct relation to nuclear size. Wilson and others have called attention to the "remarkable contrast between nucleated and non-nucleated сеП- fragments in respect to synthetic processes. The earliest observations on this subject were the classical ones of Waller (1852) on the regeneration of nerve-fibers, which proved that this process only takes place when the axis-cylinders remain in connection with the nucleated cytosomes of the nerve-cells. When а nerve-fiber is severed the distal portion degenerates, while the proximal portion (still connected with the nerve-cell and its nucleus) may readily grow forth until the missing portion is restored. This observation, repeatedly confirmed by later observers, was not wholly decisive, but nevertheless gave the first clear indication of the necessity of the nucleus for growth, regeneration and differentiation.” 4

Moreover, if we are correct in our assumption that cell activities—funetion, growth, division—are due to a difference in potential between the nueleus and the cell body, it is certainly significant that, as has been observed by Conklin in studies of the life phenomena of Orepidula, “the largest nuclei appear in blastomeres that contain the largest amount of active protoplasm, irrespective of their total size." 5 (Fig. 59.) This is well shown in the accompanying cuts in which it will be seen that the largest nuclei are in the smallest cells and as we have already stated in another chapter, this variation in nuclear

Fig. 59.—Karyoplasmic relation in segmenting eggs of Crepidula after centrifuging. Direction of the centrifugal force shown by arrows (Conklin). Та each of these 2-cell and 4-cell stages the nuclear size is proportional to the amount of active protoplasm (stippled), not to that of the cell as a whole. (From Wilson: The Cell in Development and Heredity, New York, 1925, p. 732.) plasma ratio in its relation to the rapidity of growth and reproduction of cells is strikingly manifested in cancer cells. Wilson calls attention also to a fact which is of striking

“Tt is now rather generally accepted that the more primitive types of cells were very probably devoid of an individualized nucleus, ie. that the nuclear materials were originally scattered through the cell in the form of chromidia-like bodies which only at a later period became aggregated to form a nucleus of the ordinary massive or vesicular type. "This view finds its support in the present existence of various forms among the Protista in which the

nuclear material is actually thus distributed through the cell, and by the fact (if existing accounts are correct) that in some of these forms individualized nuclei may arise either by enlargement of the individual chromidia or by their aggregation into a granular mass, in the processes of gamete-formation, mitosis, or spore-formation (bacteria) .” 6 Such examples as these might be multiplied by anyone who will review the phenomena of growth and of cell division in the protozoa as they are described by Conklin, Hertwig and a multitude of other observers and cannot fail to be impressed by the extent to which each process may be explained by the bipolar theory.

It remains to offer a suggestion as to the possible manner in which the characteristics of the parent cells or of the parent ageregation of cells is impressed upon the new cell or on the descendent aggregation of cells. In accordance with the bipolar theory the nucleus of the original unicellular organism— the positive pole, was the prototype of the brain and central nervous system of the multicellular organisms. As we have noted above, the unicellular organism cannot function without the nucleus. It is the dynamic center, the control center, the organizing center. If the nucleus is removed from a unicellular organism it ceases to function and cannot reproduce itself. We must conceive therefore that in the nucleus of the ovum must reside those potential qualities which are to govern its later activities.

In the unfertilized ovum the nucleus is balanced by the cytoplasm, hence exerts no influence upon it. As soon, however, as the nucleus of the ovum is reinforced by the nuclear spermatazoón of the male a difference of potential is established which becomes at once effective in the initiation of the processes of cell division and differentiation by means of which the new individual is constructed. These premises, however, do not explain how the reinforced nucleus of the ovum can construct out of the apparently unorganized cytoplasm of the ovum and its own structure, a new individual resembling its parents. It must be presumed that there exists in the cytoplasm and in the nucleus the physical antecedents of the structures and parts of the fully developed organism. Just as in many unicellular organisms, projections from the nuclear structure appear to have definite neuromotor

functions, and parts of the cytoplasm are differentiated to perform the functions respectively of ingestion, digestion and elimination, so we may consider that in the cytoplasm and the nucleus may be found in miniature the organs and tissues of varying functions in the developed individual. Moreover, in the fertilized ovum are at once initiated electrical currents between the nucleus and the cytoplasm, by means of which the rudiments of the muscles, glands and viscera begin to be organized, electric currents which differ not at all from those which accomplish the further growth and development of the child after birth.

I can do no better in this connection than to quote again Wilson’s summary of the point of view of various investigators, with Wilson’s own comments thereupon. “To Bonnet and other preformationists the localization problem offered по difficulties. The Gordian knot was cut by the assumption that the embryo is from the first preformed in the egg. The final overthrow of this doctrine by Wolff and his successors excluded this easy solution, but the possibility still remained that the egg may contain prelocalized regions that are inevitably predestined for the parts to which they give rise—to a certain extent such predestination is indeed a matter of observation in the case of eggs that visibly display polarity and bilaterality. The questions here raised were discussed by Wilhelm His (74) in his interesting work Unsere Korperform. ‘It is clear, on the one hand, he says, ‘that every point in the embryonic region of the blastoderm (of the chick) must represent a later organ or part of an organ, and, on the other hand, that every organ developed from the blastoderm has its preformed germ (vorgebildete Anlage) їп a definitely located region of the germ-dise. . . . The material of the germ is already present in the flat germ-dise (in the chick), but is not yet morphologically marked off and hence not directly recognizable. But by following the development backwards we may determine the location of every such germ, even at a period when the morphological differentiation is incomplete or before it occurs; logically, indeed, we must extend this process back to the fertilized or even the unfertilized egg. According to this principle, the germ-dise contains the organ-germs spread out in a flat plate, and, conversely, every point of the germ-dise reappears in a later organ; I call this the principle of organ-forming germ-re-

gions? Ray Lankester (277) developed this conception as follows: ‘Though the substance of a cell may appear homogeneous under the most powerful microscope, it is quite possible, indeed certain, that it may contain, already formed and individualized, various kinds of physiological molecules. "The visible process of segregation is only the sequel of a differentiation already established and not visible. The egg-cytoplasm has a definite molecular organization directly handed down from the parent; cleavage sunders the various ‘physiological molecules’ and isolates them in particular cells. Whitman expresses a similar thought in his classical work on Clepsine: ‘While we cannot say that the embryo is predelineated, we can say that it is predetermined. The “histogenetic sundering” of embryonic elements begins with the cleavage, and every step in the process bears a definite and invariable relation to antecedent and subsequent steps. . . . The conclusion of Rabl (779) was similar: ‘In spite of ourselves the question is forced upon us whether we must not assume the existence, even in the unsegmented egg, of a quite definite and orderly grouping and distribution of the protoplasmic particles and molecules) Van Beneden (784) pointed out the close kinship of this conception to that of a theory of preformation: ‘If this were the case (ie., if the egg-axis coincided with the principal axis of the adult body), the old theory of evolution would not be as baseless as we think today.

The fact that in the ascidians, and probably in other bilateral animals, the median plane of the body of the future animal is marked out from the beginning of cleavage, fully justifies the hypothesis that the materials destined to form the right side of the body are situated in one of the lateral hemispheres of the egg, while the left hemisphere gives rise to all of the organs of the left half. “Later researches, both comparative and experimental, have brought forward a complete demonstration of the essential correctness of these conclusions. Many cases have now been made known in which the egg-substance is more or less definitely marked out— by the presence of pigment, specific types of granules and the like —into visibly different areas which give rise to particular parts of the embryo; and artificial destruction or removal of these areas is followed by the formation of an embryo from which the corresponding parts are absent or defective.” *

It is no more logical to conclude that every organ and tissue in the adult animal has not its physical, specific material in the nucleus and cytoplasm of the fertilized ovum than to say that the metal and wood and fabric in the automobile shop had no previous existence but appeared de novo in the finished motor car. We have stated in a preceding chapter the conception that within the brain and central nervous system of the individual have been laid down facilitated paths, each of which responds specifically to specific vibrations transmitted to it by certain stimuli. And these specific vibrations when conveyed to the muscles or glands or viscera excite in them specific vibrations resulting in specific action. In view of the theories outlined above and our own conception, is it not reasonable to assume that in the sex cells of the parents specific vibrations are developed for each of the infinitesimal differentiated portions from which the differentiated organism is to develop.

Thus each organ of a plant or of an animal may be regarded аз a separate species, and as such must furnish the specific energy to reproduce itself in the seed, just as if the organ were leading a separate existence and reproducing itself as simpler organisms do. Sex organs have the ability to affect profoundly certain other organs of the body, such as, for example, the brain, the thyroid, the adrenals, ete. If the diminutive cells of the sex organs can exert major influences on distant powerful organs, one would suppose the reverse ‘would be true, and that each organ must exert a distinct influence upon the cells of the sex organs.

The manner in which this influence may be developed may be hypothecated as follows:—The dynamic units forming protoplasm are the progenitors of the cells constructed on a similar pattern which form the organs which in turn form the larger organisms. Among the protoplasmic bipolar units a closer communication exists than among the organs of the larger animals and plants. And again the separation of the chemical constituents of the protoplasmic units is greater than the separation of the same chemical elements in non-living matter. Living matter is held together in electric systems which are more loosely bound together than in non-living matter, thus making possible the accumulation of charges—the development of internal strain until the dividing point is reached and identieal

new units are formed as the result of electric repulsion. These we believe are the pangens of De Vries which he considers to be the bearers of the hereditary characters. These protoplasmic units are constructed on the universal pattern of energy found in all the non-living and as we believe in all the living. Owing to their loose bonds these primary bipolar units are readily thrown out of equilibrium, and hence their size and character are readily changed. Such loosely balanced physieal units having such easily disturbed equilibrium could be easily modified by physical forces, especially by any specific electric vibratory force. Thus the dominant electrical units of the plant or animal might exert their characteristic influences on this sensitive physical unit through the mediation of specific electric forces. In growth and in repair, these specific electric forces would apparently build up all the characteristic structures including the protoplasm within the cells, this protoplasm containing the physical dynamic units. It may be argued that growth and repair come from the contiguous protoplasm and that cells do not play a réle, but in the case of divided nerves, or in the case of separation of the nucleus from its protoplasm, provided communication be maintained by a conducting strand, it is obvious that the specific energy that sustains and that repairs the detached protoplasm is furnished by the cell. It does not seem difficult to conceive that there may well be an analogous transference of specific energy from any parent plant or animal to the protoplasmic units—pangens (De Vries), physiological units (Spencer)—in the sex cells.

An important point in our conception is that what is inherited is the pattern of energy, not the chemical elements of the parent. The atoms and molecules of the chemical elements in the developing plant or animal are the building stones which are used by the energy of the primary physical unit. From identical atoms and molecules, therefore, this primary energy mechanism will build identical forms. If by the application of another form of specific energy than that by which it was created the physical structure of this labile unit is altered, a corresponding change in the related characteristics of the new individual will be carried on.

The demand for a physical basis for heredity was recognized by Darwin in his pangenesis theory, and by the conceptions of Weissman, Naegeli, De Vries and Spencer, respectively of “ancestral At 2 “Шор, ” *pangens," and “physiological units.” In all these conceptions, however, the purpose was to supply a form and structure as the fundamental basis of heredity. In our conception, on the other hand, specific energy is the fundamental basis, and form and structure are the products and tools of that specific energy. For if living organisms are energy transformers, it would seem logical that the primary róle should be played by energy and the secondary by form and structure. This hypothesis does not contradict Darwin’s theory of pangenesis, but it offers a possible explanation of that theory. Darwin’s theory assumes that the cells of the organism each соп- tribute a small gemmule larger than a chemical molecule but smaller than the smallest organism. ‘To Darwin’s mind it was logical to conceive that each cell must make a hereditary contribution to the sex cell. According to our conception instead of contributing a particle of material to the sex cells, the cells of the body construct the element in the sex cells at a distance by the specific energies of cells or cell groups, thus satisfying the logical requirements of Darwin’s pangenesis for the transportation of gemmules from somatic cell to sex cell. Darwin supposed that the gemmules thus transported to the sex cells would assimilate food and divide; but so could such a physical unit as we hypothecate. We may also logically assume that these specific energy units may remain dormant for generations, until other competing units, which dominate for a time, in turn allow them to dominate.

The physical balance which determines which type of primary unit will predominate may depend on the energy of the synergistic units. A unit may be represented in a sex cell and yet may not dominate the type of structure or of energy, just as in some lower forms a part of the animal may grow into a whole animal only when the dynamic preponderance of the whole animal is removed. Also it may be possible that the chance reinforcement of equal energy forms may give predominance in the medley of chances in breeding. Thus this energy theory of heredity indicates how pure breeds reinforce and mixed breeds coanterbalance and

hence reduce the whole energy effect, thus suggesting a physical It remains to consider by what mechanism the perpetuation of general form-species may be effected. It may be supposed that through long use—for eons of time—the dynamic units of force or energy transformation have become stabilized and static so that they cannot be modified by ontogenetic energies that are constantly developed in each generation. On the other hand, the energy units the activities of which created the ancestral forms, now have the power of creating the high spots only; e.g., although the human embryo passes through the stage of a fish, it only shows, as it were, a fish gesture. The form, the tail and fins, the shape of the head, the eyes, the scales, the psychic characteristics of a fish never appear. The fish stage and all the other stages of the ancestors are mere pale shadows of the past and do not represent. any organ of the bygone species that could perform an ancient function. At no time could a human embryo break off its progress toward becoming a human and become a fish. The phylogenetic remnants of species repeated in the human may not represent a complete roster of his predecessors, as many ancient forms may have been left behind altogether. In other words, the energy units of ancient forms could assimilate food, could grow and could reproduce only while the more labile hence more powerful energy transforming units of the more recent forms were gaining the ascendency; and when that ascendency was achieved the later units of energy outstripped the older units in acquiring and transforming energy and consequently the ancient units remained passive or regressed. The ancient or phylogenetic units, therefore, may serve no useful purpose, just as such vestiginous organs as the muscles of the ear of man do not serve any useful purpose; but have been outstripped by their competitors for food and energy.

The struggle for existence would bring about the elimination of such ancestral development, for if a human embryo developed into too much of a fish it could not become much of a man and hence would lose in the struggle for existence. The embryo may be said to make use of the energy principle of the ancestral forms just as the modern ocean liner makes use of the energy principle of Fulton’s steamboat, but does not continue the structure of the latter. It may be questioned whether the human embryo

organism requires the vestiginal muscles of the ear, requires the appendix vermiformis, or than the occasional horse requires the faint zebra stripes. The phylogenetic structures like the vestiginal may have historical value only. To suppose that the speed of growth of the human factors in the growth of the human embryo at the fish stage for example, is due to stimuli from the outside is to overlook the fact that outside stimuli are not available to the fetus in utero. That the human development predominates must mean rather that the energy units representing the human must outnumber greatly the energy units representing the fish. Та consequence the human elements which play upon each other and reciprocally increase each other’s speed of growth will overcome the energy units of the fish. Hence, the human organism can grow unhindered by the competing weaker fish units.