HEREDITY WITH VARIATION.
That an offspring always inherits from its parents many of their characteristics is well known; that it always varies, more or less, from them is also equally well known. Heredity and variation are twin forces that play upon every creature, holding it rigidly true to the parental type or compelling more or less divergence therefrom, according to the strength of the one or other power; so that every creature is the resultant of the activities of these two great parallel forces. Variation is coextensive with heredity, and every living creature gives evidence of the existence of variations.
Examples of Variations. No two leaves on a plant are exactly alike; no two children of the same parents give a perfect resemblance; no two individuals of the same species are molded in precisely the same pattern; of the thousands and thousands of faces that we observe in a city in the course of a year, each has some distinctive peculiarity.
The trained eye of the gardener recognizes each hyacinth among hundreds of bulbs; of the shepherd, each sheep in his flock; of the Laplander, each reindeer crowded in his herd like ants on the anthill. In a flock of 1,000 sheep each mother can even recognize a variation in the voice of her own lamb, all alike to us.
Every part of an animal is subject to variations, not only in bodily structure, but also in habits and instincts, and these variations are large in amount, numerous and diverse in character. Many observations, experiments and measurements that have been made at various times attest the truth of this assertion. Not only do variations take place in animals and plants under domestication, but also in the wild state.
Illustrations of Heredity. Mental heredity can be illustrated by studying the genealogies of such persons as Aristotle, Goethe, Darwin, Coleridge, Milton, etc. Probably the Bach family, of Germany, supply one of the best illustrations of the inheritance of intellectual character that we know of. The record of this family begins in 1550, lasting through eight generations to 1800. For about two centuries it gave to the world musicians and singers of high rank. The founder was Weit Bach, a baker of Presburg, who sought recreation from his routine work in song and music. For nearly two hundred years his descendants, who were very numerous in Franconia, Thuringia, and Saxony, retained a musical talent, being all church singers and organists.
When the members of the family had become very numerous and widely separated from one another, they decided to meet at a stated place once a year. Often more than a hundred persons—men, women and children—bearing the name of Bach were thus brought together. This family reunion continued until nearly the middle of the eighteenth century. In this family of musicians twenty-nine became eminent.
Inheritance of moral character is well established. Heredity, in its relation to crime and pauperism, has been thoroughly investigated by Mr. Dugdale in his most instructive little work entitled The Jukes. In this work the descendants of one vicious and neglected girl are traced through a large number of generations. It reveals that a large proportion of the descendants of this woman became licentious, for, in the course of six generations, fifty-two per cent. of the females became harlots and twenty-three per cent. of the children were illegitimate. It shows also that there were seven times more paupers among the women than among the average women of the State, and nine times more paupers among the male descendants than among the average men of the State.
The inheritance of physical peculiarities is so obvious as to need no illustration. Among the ancients the Romans stereotyped its truth by the use of such expressions as the labiones, or thick-lipped; the nasones, or big-nosed; the capitones, or big-headed; and the buccones, or swollen-cheeked, etc. In more recent times we read of the Austrian lip and the Bourbon nose.
Questions of heredity and variation are cytological ones—that is, questions of the anatomy, physiology, physiological chemistry, and pathology of cells. The most important part of a cell, as far as these questions are concerned, is the nucleus. The nucleus is the physical basis of all the heritages of an organism, from the simplest to the most complex. The nuclear threads may, therefore, very appropriately be termed the hereditary threads, or, collectively, the hereditary mass; and the physiological units in them the hereditary units. The nucleus is of fundamental importance in the reproduction or multiplication of both unicellular and multicellular animals and plants.
In unicellular creatures multiplication may take place by fission and by conjugation. Both of these processes can be studied by observation of the infusorians. Maupas’s beautiful investigations on these unicellular animals have demonstrated that multiplication by fission may proceed to a prodigious extent for many generations, but that a time comes when the process fails, and the species will become exhausted and die out unless there is a rejuvenation of it by conjugation of individuals. In conjugation two individual infusoria come in apposition with each other, the nucleus in each undergoes subdivision. They reciprocally exchange part of their nuclear contents so that each infusorian comes to contain hereditary threads of two distinct individuals. From these rejuvenated (or fertilized) individuals multitudes of others may be derived by fission until exhaustion again takes place.
Multiplication in multicellular creatures may be accomplished by budding (which is allied to fission), and is exemplified in the plant, hydra, the queen bee (parthenogenesis), etc., and by fertilization (which is allied to conjugation). A knowledge of the phenomena of fertilization of the ovum by the spermatozoid is essential to any understanding of the problems of heredity and variation in mankind. The nuclear threads of the ovum are its hereditary threads—the groups of maternal hereditary units; likewise, the nucleus of the spermatozoid contains the paternal groups of hereditary units.
Fertilization. In fertilization, the spermatozoid (a nucleated flagellate cell) penetrates the ovum (a nucleated, encysted cell), its protoplasm mixes with that of the ovum, and its nuclear threads come into relation with the nuclear threads of the ovum; so that the fertilized ovum (a new creature, a veritable microcosm) is still a nucleated cell, but one in which the nucleus is compound, is hermaphroditic, in that it contains maternal and paternal threads—that is, maternal and paternal hereditary units which constitute its hereditary mass.
It will be convenient to speak of the maternal and paternal hereditary units in the fertilized ovum as ancestral hereditary units.
This hermaphroditic cell passes through complex phases, illustrated by embryology, to the adult. In doing so this hermaphroditic cell (mother cell) first divides into two smaller cells (daughter cells). The mother cell divides in such a way (by mitosis) that one-half of its nucleus and part of its protoplasm goes to one daughter cell and the other half of the nucleus, with the remainder of the protoplasm, goes to the other daughter cell. It is an interesting fact that although the amount of protoplasm which goes from the mother cell to the two daughter cells may be unequal at times, yet the amount of the nucleus in one daughter cell is always exactly equal[2] to that in the other; so that each daughter cell contains maternal and paternal hereditary masses of equal quantity and quality, being in fact one-half that of the fertilized ovum. In consequence of the fact that each hereditary unit in the nucleus of the daughter cell can absorb nutriment and grow, it comes about that the nucleus of each daughter cell attains to the size of that of the mother cell. The enveloping protoplasm of the nuclei also grows to a greater or less extent, so that the cells as a whole grow. These two daughter cells go through the same process and form other daughter cells, and so on through all the mitoses of development, until all the myriads of cells of a living organism are produced, each of which contains maternal and paternal hereditary masses of equal quality and quantity, and also of the same character as that of the fertilized ovum whence they are all derived.
Apart from their activity in absorbing nutriment and growing, the great majority of the hereditary units in the nuclei of the forming cells remain latent. But some of the hereditary units in each cell produced are active. They multiply, grow and migrate out of the nucleus, and get among the units of the enveloping protoplasm. During this activity they undergo physical and chemical changes and effect corresponding differentiations in the protoplasm of the cell. Thus, through many mitoses and many differentiations of the protoplasm of cells, we finally derive from the fertilized ovum all the cells that constitute the adult body, such as muscle cells, glandular cells (as liver cells, kidney cells, etc.), nerve cells, skeletal cells (as bone, cartilage, and connective-tissue cells), and the ova and spermatozoids. According to this theory, the nucleus of each cell in the adult animal or plant is pure hereditary mass, exactly like that in the nucleus of the fertilized ovum; but the protoplasm of each adult cell that envelopes the nucleus may differ greatly in different cell groups, as in muscle, nerve, cartilage, and the like. Of course, the protoplasm of those cells that develop into the ova and the spermatozoids has differentiated along such lines as to become like that of the ovum and spermatozoid, the junction of which formed the fertilized ovum. These statements hold true for plants and the lower animal organisms, although they cannot be verified for the higher animals. More than likely the pure hereditary masses are present in the body (somatic) cells of the higher organism in latent conditions, but are unable ever to be developed owing to the greater specialization of these cells. It is thus seen that all the cells of many animals and plants can perform their own special functions and at the same time contain all of the hereditary units of the complex organism in a state ready to develop under favoring conditions.
Since the ova and spermatozoids are cells specially differentiated for the purpose of propagating the species by sexual generation, and since their conjugation produces the germ of a new creature, they may very appropriately be spoken of as germ cells. Since all the other cells of the adult form the great bulk of the body that envelopes and protects the germ cells, they may be termed the body cells, or somatic cells.
Suppose that ova, containing maternal and paternal groups of hereditary units, are fertilized by similarly complex spermatozoids, and the process is repeated generation after generation. There will come a time when the fertilized ovum will have a highly complex nucleus composed of many different ancestral groups of hereditary units.
One often hears the expression that a child is a chip of the old block; but this is only a very partial truth, for a child is preëminently a composite chip of many old blocks.
The complex nucleus of the fertilized ovum may be compared to a modern Italian building which has been constructed of material—a column here, a cornice there, a lintel yonder—gathered from different classic buildings of varying antiquity. In view of the increasing number of ancestral groups of hereditary units that must have accumulated in the nuclei of ova in the course of time, there must necessarily, for mechanical reasons, have arrived a period when these nuclei could receive no more of them by fertilization, unless natural selection should develop some saving device; hence we have, possibly, an explanation of the phenomena of maturation in ova (the reducing process of Weismann and Hertwig). Here the ovum, prior to fertilization, undergoes mitosis twice in succession, by which the polar bodies are formed and the hereditary mass is diminished by one-half (the mature germ cells having only one-half the number of nuclear threads that the body cells possess). A homologous process takes place in the maturation of spermatozoids. Fertilization increases the amount of the hereditary mass in the ovum to the original quantity, and thus restores the number of nuclear threads to the specific number. All the body cells derived from the fertilized ovum possess, also, the specific number of threads.
This union of two distinct hereditary masses is called amphimixis as well as fertilization.
Maturation and amphimixis or fertilization are the source of many variations in the body, good and bad, beautiful and ugly, geniuses and monstrosities; because, in the commingling of distinct hereditary masses, there is a struggle for existence between the hereditary units and a survival of the fittest.
In this struggle some of the hereditary units are strengthened so that heritages may be augmented; some mix so that there may be a blending of characteristics; some are mutually exclusive; some are prepotent; some are neutralized; some are destroyed; some lie dormant (latent) for varying lengths of time, and some are so altered as to produce much modified forms; and thus the possibilities of combination, reactions and modifications of the hereditary units, and therefore heritages, are almost endless.
The augmentation of heritages in the fertilized ovum is well displayed, for instance, where fleet horses are bred with fleet ones, until, by careful selection, generation after generation, a progeny may be secured much more swift than the original stock from whence they were derived. In the same way good milch cows have been produced.
The mixing of hereditary units and blending of heritages is shown in the color of the skin, as where a mulatto child is born to a negress by a white father; mutually exclusive heritages are well illustrated in the color of the eyes, as where a child has either the blue eyes of one parent or the black of the other, but never any blending of the colors; this may also be illustrated where the white game bird and black one are crossed, the young being either white or black, but never blended. Prepotency is illustrated where the silky variety of fan-tailed pigeon is mated with any other small-sized variety of pigeon, for the silkiness is invariably transmitted. A most interesting case of prepotency in mankind, mentioned by Ribot, is that of Lislet-Geoffrey, an engineer in Mauritius. He was the son of a very stupid negress and an educated white man. In physical constitution he was as much a negro as his mother; he had the woolly hair, the features, the complexion, and the peculiar odor of his race. He was so thoroughly a white man as regards intellectual development that he succeeded in vanquishing the prejudices of race, so strong in the French colonies, and in being admitted into the most aristocratic houses. At the time of his death he was Corresponding Member of the Academy of Sciences.
In this, it will be observed, we have prepotency in the mother’s physical constitution, and in the father’s intellectual characteristics.
The struggle of heritages in the impregnated ovum may lead to such structural changes of the nucleus, and therefore of the cell, as to develop the most marked variations—such variations as the biologists call sports.
In the latter part of the eighteenth century the farmers of Massachusetts had flocks of ordinary sheep on their farms. These sheep were continually jumping fences and getting on neighboring farms. They were the source of many disputes and much irritation between neighboring farmers. Finally, one of the sheep had a lamb which, when grown, displayed well-marked peculiarities (a sport). It had a longer body than the ordinary sheep and shorter legs, which were bowed. It was noticed that this sheep could not get over the fences. The cute Yankee farmer, noticing this valuable peculiarity, carefully preserved this peculiar sheep, and from it was ultimately derived, by careful selective breeding, a special variety known as the Ancon sheep.
The germinal variations resulting from the mixing of two separate hereditary masses by impregnation find their expression in the most varied qualities of the minds and bodies of developing children. If the variations are not especially marked, they are looked upon as normal and attract no special attention.
But if the variations are so pronounced as to compel attention, and at the same time it is known that they are useful, they are spoken of as talents, or, on the other hand, if they are harmful or useless, they are designated as pathological or monstrosities.
These are truly what the biologists call sports; and to those classes of sports that occur as specially gifted in human culture, in the varied fields of science, art, or literature, we assign such a person as Shakespeare, and call the remarkable variations embodied in him genius. On the other hand, such variations as lead to certain forms of pigmentary degeneration of the retina, and to Daltonism, to dyschromatopsia and achromatopsia, to certain supernumerary glands, polydactylism, and such like, which are either useless or harmful, we designate as pathological cases or monstrosities.
Hereditary units (the carriers of heritages) may be latent—that is, they may appear late in life, or in the offspring, or, still again, in remote descendants; in the latter cases the heritages are spoken of as reversional or atavistic. Latent hereditary units may very usefully be compared to dormant seeds buried in the ground. It is stated that buried seeds may lie dormant for many years, so that when a plot of ground is plowed deeply and upturned, plants that have not been seen there within the memory of one will often make their appearance and flourish. The hereditary units are veritable living seeds, that, under certain and often unknown stimuli, grow and unfold their heritages as do the buried seeds.
Latent heritages are well illustrated by a study of secondary sexual characters as developed at puberty.
Among our barnyard fowls, the hens often, when they have atrophy or degeneration of the ovaries, although up to this time they have laid eggs for years, stop this function, put aside the plumage and appearance proper to their sex, and don more or less completely the garments of the rooster. Thus females have latent in them many secondary sexual characters of the male. For similar reasons the male develops, occasionally, female characters.
This latency is illustrated again in deer. In most species of the deer tribe the males alone possess antlers, yet it is a well-known circumstance that in females with degenerations of the ovaries rudimentary horns that are never shed appear. A study of congenital color-blindness illustrates beautifully latent heritages, showing how the females of one generation may be free from the malady and the males of the next afflicted.
A study of the regeneration of lost parts in various animals and plants illustrates well the latency of many hereditary units. A cutting made from the willow and planted sends out roots and finally reaches the dimensions of the adult tree. Here the body cells of the stem evidently contained, in a latent condition, many hereditary units in their nuclei, which became active through the special stimulus of being planted as a cutting. Adult plants can be raised from the cuttings of many other plants.
If the garden worm is cut in two, the head-part will reproduce the tail-portion. If the little fresh-water polyp, Hydra, be cut into a number of pieces, each segment will reproduce a perfect animal. Many lizards, after losing their tails by violence, manufacture new tails through the agency of the latent hereditary units contained in the body cells of the stump left. If the tentacle or “horn” of a snail, which contains an eye with a perfect lens and retina, be cut off, the animal can reproduce another one with a perfect eye, and this can be repeated a number of times. Often newts, when fighting with one another, or lobsters when fighting, lose a leg or a claw. These highly organized animals have the power of creating new limbs, making bones, ligaments, muscles, nerves, cuticle, and so on. All of this is done through the hereditary masses in the nuclei of the body cells at the site of the injury.
A study of the phenomena of polymorphism in hydroids and insects will beautifully and most interestingly illustrate latent hereditary units.
Much that is speculative and fanciful is included under the subject of atavism, and the safest plan for pathologists and biologists, in considering any abnormality, is to remember a golden rule of Gegenbaur, that only those structures are reversional which are taxonomically not far distant or phylogenetically not very old. Embryology is also a very important check in considering such subjects.
In mankind supernumerary limbs and digits, microcephalia and micrencephalia, have been looked upon as reversions to the simian type.
Lombroso, in contrasting the criminal with normal man, looks upon his homo delinquens as an illustration of atavism, contrasting with homo sapiens. But, as Ziegler, in his Pathology, well observes, many writers have gone too far in this respect, and have characterized as atavistic formations various acquired pathological formations and fresh variations of germ cells.
I think one can safely say that supernumerary ribs and those supernumerary nipples and mammary glands along the line of the deep epigastric and internal mammary arteries are truly atavistic structures; also certain muscles normally belonging to those mammalia which come near to man in the scale of relationship, and which appear in man as muscular variations, are reversional.
Children are often born with pigmented hairy patches on their bodies known as moles; sometimes these hairy moles are only of the size of a split pea, in other cases they are several square inches in area, while in rare cases almost all of the trunk may be thus covered. Although many similar pathological cases are often but marked variations called sports, yet the illustrations mentioned are undoubtedly reversional. Of the multitudinous illustrations of atavism that could be mentioned I wish to refer to but one more case.
The conjunctiva is a modification of skin, and frequently proclaims its ancestry by reverting to its original form. It is by no means a very rare event to see a patient having a patch of hair-covered skin growing upon the ocular conjunctiva. While a clinical assistant at the Royal Ophthalmic Hospital in London, we saw one such case, and Dr. Treacher Collins, the pathologist of that eye hospital, has stated that about twelve cases are seen there annually of this pathological condition, which is atavistic, according to Sutton, although it seemingly violates Gegenbaur’s rule about phylogenetic remoteness, and may be looked upon by some as a pathological illustration of a sport.
In speaking of inheritance, we should carefully discriminate between heredity and pseudo-heredity. Physicians constantly write of tuberculosis, lepra, smallpox, and syphilis as hereditary; but it is incorrect and misleading to do so. When a person has syphilis, say, from the earliest existence—that is, from the fertilized ovum by transmission of a syphilitic microbe through the germ cells of the parents—this should be designated by its proper name as congenital bacterial infection. This is totally different from the hereditary qualities that flow from the structural equilibrium following the commingling and struggle for existence of multitudes of hereditary units.
The one set of hereditary qualities is purely germinal, while the other is germinal profoundly modified by the presence of an infecting microbe. Of course, to the extent that any toxines that are secreted by the bacteria may cause permanent structural changes in the germ-cells, to that extent may the germinal characteristics be transmitted and become hereditary.
Many instances of infection of the child in utero have been reported in cases of endocarditis, scarlet fever, and smallpox; and there can no longer be any doubt, from experimental investigation and recent observation, that pneumococci, typhoid bacilli, anthrax bacilli, and pus cocci are able to pass to the fœtus through the placenta. But the diseases that develop in this way can be called hereditary with even less semblance of correctness than in the case of the fertilized ovum that is invaded with a microbe.
All of these cases are illustrations of pseudo-hereditary transmission, and should, for the sake of clearness and accuracy, be spoken of as prenatal infections.
So far as the problems of heredity and variation are concerned, we may say that the life cycle begins and ends with the germ cell. Insects lay their eggs in old age; among plants the annuals flower but to die; in higher creatures the cessation of the procreative power often marks the beginning of bodily decline.
Bearing in mind that the human body consists of two great classes of cells, germ cells and somatic cells, the following scheme will be found very useful in discussing heredity with variation—viz.:
| Germ-Cell | { | Stable (Heredity) | ||||||
| { | Unstable (Variation) | { | Blastogenetic | { | Temperature. | } | Environment | |
| { | Chemical substances in solution in the fluids that bathe the germ-cells, as food, drugs, poisons, etc. | } | ||||||
| { | Somatogenetic | { | Habitat | } | ||||
| { | Temperature | } | ||||||
| { | Climate | } | ||||||
| { | Air | } | ||||||
| { | Food | } | ||||||
| { | Soil | } | ||||||
| { | Water | } | ||||||
| { | “Use” | } | ||||||
| { | “Disuse” | } | ||||||
Just in proportion as fertilized germ cells during the mitoses of ontogeny give origin, among the somatic cells, to other germ cells that are structurally, and therefore physiologically, like themselves, just to that extent do we have heredity; on the other hand, just to the degree that the new germ cells which are produced are unstable, to that degree also do we meet with variations.
ENVIRONMENT.
In zoölogy the environment of an organism means the sum-total of the conditions of life that surround and affect it, such as food, air, water, climate, etc.
We have already stated that as far as evolution is concerned the structures of fundamental importance in an organism are the Germ-Cells; therefore, for our purposes, we will define environment to be the sum-total of the conditions that directly or indirectly influence in any way the germ-cells, by which variations in them may be produced, or through which stability may be maintained.
There are two great classes of environmental factors that bring about variations in the germ-cells. One of these classes acts directly on the germ-cells, and is therefore called blastogenetic; the other acts indirectly through the body cells, and is therefore designated somatogenetic. Many of the blastogenetic factors bringing about structural changes in the delicate mechanism of germ-cells are entirely unknown, and are therefore designated as fortuitous. Many other causes, such as poisons in solution in the fluids that bathe them, can readily enough be appreciated.
Blastogenetic Factors. It has been demonstrated that various chemical substances, such as chloroform, morphia, chloral, etc., have a pronounced influence upon the vital activities of cells. It is well known that microscopic unicellular plants constitute the essential part of yeast. These little cells have the power of causing fermentation in solutions of grape sugar by which alcohol and carbon dioxide are formed, the latter being a gas and escaping as bubbles. If chloroform or ether be added to the solution of sugar, before adding the yeast, no fermentation takes place, for the yeast-cells are paralyzed. But when the yeast is separated from the chloroform solution and rinsed with distilled water, it soon regains the power of causing fermentation in pure solutions of sugar.
Ova and Spermatozoids are subject to the action of drugs in a similar manner. If actively motile spermatozoids of a sea-urchin be placed in a one-half of one per cent. solution of chloral in sea water, it will be found that after five minutes their action will be completely arrested. These motions can soon be restored if the chloral solution be sufficiently diluted with pure sea water. These temporarily paralyzed spermatozoids, when completely recovered, will unite as quickly with ova as fresh spermatozoids. When spermatozoids are kept for half an hour in the chloral solution, a more decided paralysis will be observed, which persists for some time after the removal of the poisonous agent. A few minutes elapse before some of the spermatozoids exhibit feeble movements which finally become active. Even when placed near ova, it is some time before they fertilize them, although several may attach themselves to the egg’s surface. But, finally, fertilization does take place by the penetration of one spermatozoid, and the egg normally develops.
In like manner, if ova are subjected to chloral solution of varying strength, they also are influenced in a marked degree; for, when fertilized, they develop in an abnormal manner. Ordinarily, normal ova are fertilized by one spermatozoid. If fertilized by two or more, they become diseased, and develop pathologically. The chloral solution favors this fertilization by several spermatozoids. The stronger the solution of chloral, the larger the number of spermatozoids that fertilize the ovum. Experiment and observation show that the behavior of the nuclear hereditary mass is modified, during mitosis, by the chloral and other solutions. It is thus seen that the germ-cells of the lower animals can be profoundly modified by various substances.
Equally true is it that the man or woman who makes use of such drugs as alcohol, opium, chloral, and such like, in an intemperate manner, contains these poisons in solution in the blood, circulating to every part of the body, and thus bathing and profoundly influencing the germ-cells. In consequence of this fact an acquired and habitual intemperance will seldom fail to leave its impress upon one or more of the offspring, either like the original vice or one very closely allied to it. Intemperate people not only profoundly impair the health, the intelligence, and the morals of their offspring, by poisoning these delicate germ-cells, but they also transmit the fatal tendency to crave for the very substances that have acted as poisons on these germ-cells before and after fertilization. And one of the saddest features of this great medical truth is that the hereditary units which are concerned in transmitting these grave abnormal tendencies may lie dormant in the germs of one generation, to become active in those of the next; so that children of intemperate parents may lead honorable and temperate lives, and take every pains to rear, in turn, their own children in a wholesome and refining atmosphere, and yet these children of good environment may become intemperate through heredity, so that the sins of the grandparents may be visited, not on the children, but on the grandchildren.
These profound truths should lead all, and especially law-makers, to remember that “the man who inherits from his parents an impulsive or easily tempted nature and an inert will and judgment, and commits a crime under the influence of strong emotion, can no more be placed in the same category of responsibility with a man of more favorable constitution and temperament than can a man who steals a loaf under the pangs of starvation with a merchant who commits a forgery to afford him the means of prolonging a guilty career.”[3]
Not only do certain known poisons circulating in the blood, or other fluid that may bathe the germ-cells of living creatures, profoundly affect the germ-cells, but many other substances probably have great influence upon them. Certainly, the amount and character of the food have a very decided influence on them, as will be understood from the following facts.
According to Yung, who has experimented very extensively upon tadpoles, all tadpoles pass through a bisexual (hermaphroditic) stage, as is the case probably with most animals. During this tadpole phase external influences, and, more particularly, food, determine their fate as regards sex. In Yung’s experiments it was found that when tadpoles were left to themselves, the percentage of females was in the majority, the average being probably about 57 per cent. females and 43 per cent. males. In experimenting with three broods, those fed on beef gave 78 per cent. females; those fed on fish gave 80 per cent. females; and those fed on the highly nutritious flesh of frogs gave 92 per cent. females.
In Mrs. Treat’s interesting experiments on moths and butterflies, it was observed that if caterpillars were confined and starved before they entered the chrysalis state, the resultant moths or butterflies were males, but others of the same brood that had been highly nourished came out females.
The study of bees illustrates the same conclusions. It is well known that in a beehive there are three kinds of inmates, as the queen, the drones, and the workers,—the last-mentioned being females whose reproductive organs are imperfectly developed. It is believed that the eggs that give rise to queens and workers are fertilized and developed normally. But it is a very curious fact that the eggs which develop into drones do so without fertilization (parthenogenesis). What factor or factors decide the destiny of the two former, determining whether a given ovum will develop into a queen, and thus be the possible mother of a new generation, or stay at the lower grade of a working, non-fertile female? These factors are the quality and quantity of the food. An abundance of what is called royal food causes the development of the larva in such a way that the queen with her reproductive organs is formed. If a larva on the road to develop into a worker (non-fertile female) “receive by chance some crumbs from the royal superfluity,” it is found that the reproductive organs may develop to such an extent that workers partially fertile may be formed. A worker larva may, by this royal food, be intentionally reared into a queen bee.
It is thus seen how profoundly the germ-cells, in their growth, may be affected and made to vary by such a blastogenetic factor as food.
Somatogenetic Factors. As to somatogenetic factors—granting that structural changes in the body (body-cells) of an animal or plant can profoundly influence in some way the germ-cells, and that, therefore, acquired characters can be transmitted—they are many and well defined. Some of them are the habitat of an animal or plant, the temperature, climate, air, food, soil, water, structures in use or disuse (so-called “Use” and “Disuse”), etc.
The following brief descriptions will enable the reader to understand that change in the surroundings (environment) of a living creature may cause its body (body-cells) to vary.
A certain species of snail was introduced into Lexington, Virginia, a few years ago from Europe. In its new habitat it varied very much. One hundred and twenty-five varieties have been discovered there, sixty-seven of which are new and unknown in Europe, the native home of the species.
The common ringed snake, when living in its natural habitat, deposits eggs in the sand, which are hatched by the heat of the sun; but when this snake is confined in a cage in which no sand is strewn, it gives birth to little living snakes.
In experimenting on moths it has been found that the variations of temperature to which the pupæ, and probably also the larvæ, are subjected, tend to bring about very pronounced differences in the moths. Cold has a tendency to develop a darker hue in the perfect insect.
English dogs when taken to hot climates, like that of India, are known to degenerate in a few generations. It is well known how climate affects the hairiness of animals. When greyhounds are taken to the uplands of Mexico they are unable to course on account of the rarity of the air.
In 1870 a number of pupæ of a certain species of moth (Saturnia) were taken from Texas to Switzerland. After passing the winter there, the pupæ emerged from their cocoons as moths, and resembled the Texan species entirely. The young of these moths were fed on the leaves of a plant different from that the moths in Texas feed on, and they developed into moths so different in form and color-pattern from their parents that entomologists classified them as a distinct species.
We have seen how certain foods affect the germ-cells and act as blastogenetic factors; the preceding case and the following show how certain foods act as somatogenetic factors and modify the body-cells. If the bullfinch be fed on hemp seed, its color is changed to black; if the canary be fed on cayenne, its plumage becomes darker; if the common green Amazonian parrot be fed on the fat of siluroid fishes, it assumes a beautiful variegation of red and yellow.
The character of the soil has a marked influence in inducing somatogenetic variations. In France an experimenter collected seed from the wild radish and sowed one lot in heavy soil in the country, while another lot was sown by him in the dry, light soil near the Museum of Natural History in Paris. The radish “roots” grown in these two places presented marked differences in color and form. Those grown in Paris were either of a rose or white color and elongated; while those from the country were violet, dark-brown or nearly black in color, and more rounded than the former.
In the summer of 1847 Professor Buckman gathered seed from wild parsnips, and sowed them in the spring of 1848 under changed conditions of life. Most of the plants grown from these seeds were like the wild parsnips, but some of them developed the light-green color and hairless, smooth appearance characteristic of the cultivated plant. The roots also were found to be more fleshy than those of the wild variety.
Peas and squashes, when grown in different soils, often show remarkable variations.
There is one species of shrimp that inhabits brackish water, and another that lives in water which is much more salt. These crustaceans differ from one another in the character of the spines they bear and in the form of the tail-lobes. They have been regarded as distinct species, and yet either of them can be transformed into the other in the course of a few generations, by gradually altering the saline conditions of the water.
For a long while the siredon and amblystoma were regarded as being distinct genera of amphibians. Siredon was looked upon as a permanent gill-breather, while amblystoma passes through a metamorphosis and becomes a permanent lung-breather. It is now known that the former can change into the latter. If there is plenty of water the siredon remains indefinitely a gill-breather and reproduces freely; but when the water dries up it changes into the lung-breathing amblystoma. These two cases illustrate very well the power of environment to modify the development of organic forms.
As to “use” and “disuse”: It can readily be observed that exercise increases the size of muscles; that by steady application the capacity for thinking can be developed; that the oarsman’s constant use of his hands leads to the hardening and thickening of the cuticle; that the arm of the blacksmith and the legs of the mountaineer are much enlarged, etc.
When an organ is exercised properly, there is an increased blood supply to it, and, consequently, stimulated nutrition and growth in various parts, such as in the muscular, nervous or other tissues.
When an organ is disused there is diminished blood supply, and, consequently, diminished growth and functional capacity. In man it is known that certain activities, such as coal-heaving, shoemaking, etc., produce recognizable effects upon the muscular system, the skeleton, and other parts of the body.
The peculiar habits of a tribe, such as tree-climbing among those natives of the interior of New Guinea, who build their houses in the upper limbs of lofty trees, modify the body in ways that are readily recognizable.
After considering many facts in connection with the brains of rabbits, Darwin announced that this most complicated and important organ in an animal is subject to the law of decrease in size from disuse. We have very interesting illustrations of the effects of “use” and “disuse” in causing somatogenetic variations, in the differences between domestic ducks and the wild ones from which they have been undoubtedly derived. The wild duck, which must constantly be on the alert for enemies, and uses its wings so much more extensively and its legs comparatively less than the domestic duck, is a much more intelligent fowl than the stupid, well-protected domestic one. The wings of the wild duck are stronger and its legs shorter than those of the barnyard duck. It has been shown that in the wild duck the brain is nearly twice as heavy in proportion to the body as it is in the comparatively imbecile domestic duck.
Many other useful illustrations of disuse, such as the cattle and goats in India, that have dependent ears; also cats in China, and horses in parts of Russia, whose ears are dependent, could be referred to. Use and disuse are included among the factors of environment, because by those terms we mean certain groups of body-cells that are functionally active or inactive; for body-cells on any theory of modified pangenesis constitute an exceedingly important environment of the germ-cells.
The surrounding conditions (environment) of an animal or plant having the power to cause variations in the living creature by affecting its germ-cells or its body-cells, the environment may be spoken of as blastogenetic and somatogenetic.
Whether it is a fact or not that somatic variations can induce corresponding variations in the germ-cells, and thus be transmitted by heredity, it is certainly true that all heritages must come through the germ-cells. For this reason, it is clearly seen that so far as evolution is concerned the germ-cells are the factors of fundamental importance in organisms. Therefore, we may repeat that environment is the sum-total of the conditions of life that affect the germ-cells directly or indirectly.
ACQUIRED CHARACTERS.
All heritages, then, are derived directly through the germ cells. Can there be any heritages indirectly from the somatic cells through the germ-cells, as has hitherto been assumed? In other words, can acquired characteristics be transmitted to the offspring? This question has given origin to the battle royal that is still going on between opposing schools of biology. The contending parties have appealed to such biological evidence as is furnished by a study of use-inheritance, reflex and instinctive actions in animals, etc., and to such experimental evidence as the induction of traumatic epilepsy in guinea pigs, a change in the shape of the ear by cutting the cervical sympathetic nerve, protrusion of the eyeball through injury to the restiform body of the brain, and such like, noting the effects on the offspring, and have drawn very different conclusions.
As to the transmission or non-transmission of acquired characters, some have maintained that only germinal variations are transmitted (because they believe the germ cells are insulated from the body cells, and therefore from somatic influences). For instance, Ziegler, in his work on General Pathology, says: “If a disease, such as nearsightedness, is the product of a special inherited predisposition, plus the effect of harmful influences which have acted upon the body during life, only that part can be transmitted which was received by inheritance, but not that part which was derived from external influences.” In other words, there is no transmission of acquired character. In this belief it will be observed that he follows Weismann.
On the contrary, other investigators, like Darwin and Spencer, teach that somatic variations—the plus element in Ziegler’s illustration of nearsightedness—do influence the germ-cells (through some such agency as Darwin’s theory of pangenesis suggests), and that, therefore, acquired characters can be transmitted. The question is one of fundamental importance, and yet no crucial experiment has been devised or fact observed which can compel the correct answer. The evidence seems to favor the view that acquired characters can be transmitted.
The theories as to the transmission or non-transmission of acquired characters may be better understood by reference to schemes No. 1, 2 and 3. Scheme No. 1 represents the theory of Pangenesis, which teaches that reproductive cells are not formed from pre-existing reproductive cells, but by the body cells themselves. Darwin taught that all the cells of the body, such as skeletal-cells, muscle-cells, nerve-cells, and so on, are continually giving off infinitely small cell germs or gemmules, which have the power of growing and forming cells exactly like themselves. These gemmules have a great affinity for one another, and, circulating in the blood in countless numbers, they finally come together in the reproductive glands and form the reproductive cells. On this theory the fact of the transmission of acquired characters can readily be appreciated, and it can easily be understood how the parent molds the child. Suppose, for instance, that the parent, by exercise, has become a skillful athlete. In him certain muscles have become greatly developed and strengthened. During all the time of the exercise of these muscles, the modifying muscle cells have been continually giving off to the blood modified gemmules, which collect in the reproductive cells and make it possible for the offspring to develop into an athlete because the modified gemmules develop into modified muscles like those of the athlete.
Scheme 1 shows the absence of any arrow like those shown in schemes 2 and 3, directly connecting germ-cell with germ-cell; this means that in this theory there is no continuity of the germ-cells. But arrows are seen extending from the various body-cells (skeletal, glandular, etc.) to the germ-cell; this means that the germ-cells are formed by influences or gemmules emanating from the various body cells.
Scheme 2 teaches that a germ-cell (when fertilized, of course) can produce many cells, some of which differentiate, finally, into skeletal cells, some into glandular, some into muscle and nerve cells, and some into new germ-cells; so that an animal or plant, I, is formed. In like manner a germ-cell of animal, I, can give rise to the germ and body cells of animal, II, and so on indefinitely. This scheme shows that there is a direct continuity of the germ-cells; and it also shows that the germ-cells are entirely insulated, as it were, from the body-cells (skeletal, glandular, etc.), inasmuch as no influences (arrows) extend from the body-cells to the germ-cells. This means that the transmission of acquired characters, bodily, mental, moral, etc., is impossible. It means, in other words, that none of the advantages gained by a parent in the course of his life can be handed on to his offspring by heredity. There are many biologists and pathologists who teach this theory as the correct one.
The majority of biologists accept the theory illustrated in Scheme 3. This is the theory of modified Pangenesis, which teaches that there is a direct continuity of the germ-cells and that these germ-cells are not insulated from the body-cells, but that the latter, when modified as the result of experience, can send off influences that correspondingly modify the germ-cells; so that the latter, when developing into a new individual, may cause the same body variations that exist in the parent. In short, this scheme illustrates not only that there is a germinal inheritance, but also an inheritance of acquired characters. In this Scheme 3, the oblique arrows show that germ-cells produce other germ cells; the perpendicular arrows show that the germ-cells are modified by influences that proceed from the body cells.
Germinal characteristics are transmitted with vastly greater amplitude and swiftness than merely body (acquired) characteristics. If, for instance, a man were born with that physical constitution that makes with ease a first-class pianist out of him, his sons may easily, through heredity, be first-class pianists. But if a man be born without such a congenital tendency and has by constant labor and practice so developed the muscles of his forearm, his nerves, his brain, etc., that he becomes a very good pianist (acquired characters); and, further, if his male descendants for thousands of generations, in succession, have become very good pianists by constant practice, we may expect that the sons of these last generations may obtain a congenital tendency to become first-class pianists quite easily. The constant improvement, by practice, of groups of body-cells (muscle-cells, nerve-cells, etc.) for generations, has, in each generation, tended to so correspondingly modify the germ-cells that they have acquired the power to develop into men who may become very good pianists with very little practice. This illustrates that there may be a continuous summation of feeble germ-cell variations that have been induced by prolonged influences emanating from somatic variations, so that, in the course of many generations, robust acquired characters may ultimately be translated into strong congenital characters (Scheme 3).
Scheme 1. Illustrating the theory of Pangenesis. Here the germ-cell (a) develops into the body-cells, e, e, e, e, of animal I, as indicated by the oblique arrows, but not into any germ-cells, as indicated by absence of arrow between germ-cell (a) and germ-cells (b). The germ-cells (b) in animal I are formed by the aggregation of infinite numbers of gemmules from the various groups of body-cells, e, e, e, e, as indicated by the perpendicular arrows. The germ-cell (a) transmits germinal heritages to the body-cells e, e, e, e; these body-cells transmit the heritages to the germ-cells (b) by means of the gemmules. If the body-cells are modified in any way, correspondingly modified gemmules are sent to the germ-cells (b), and these germ-cells are modified and thus transmit acquired characters to animal II, and so on.
Scheme 2. Illustrating the theory of Continuity of the Germ-Cells; pure germinal inheritance; and the non-transmissibility of acquired characters. The germ-cells are insulated from the body cells. The germ-cell (a) develops into the body-cells, e, e, e, e, and the germ-cells (b), in animal I. The body-cells, e, e, e, e, do not influence in any way the germ-cells (b), as indicated by the absence of perpendicular arrows. The germ-cells (b) get all their heritages from the antecedent germ-cell (a), as indicated by the oblique arrow from (a) to (b). All heritages are purely through the germ-cells. The same with the animals II and III. Germ-cells (a), (b), (c), (d), are connected together by obliquely placed arrows, indicating the continuity of the germ-cells.
Scheme 3. Illustrating the theory of Continuity with Modified Pangenesis. A germ-cell (a) develops into the body-cells, e, e, e, e, and the germ-cells (b) of animal I. The germ-cells (b) get their heritages directly from the germ-cell (a), as indicated by the long, obliquely-situated arrow (continuity of the germ-cells). The germ-cells (b) are, moreover, modified by influences extending from the body-cells, e, e, e, e, as indicated by the perpendicular arrows. A modified germ-cell (b) can develop into a modified animal II, and the body-cells of this animal can influence and modify the germ-cells (c); and so on, indefinitely. The perpendicular arrows indicate that acquired characters are transmitted, and that, too, through the germ-cells.
Professor Morgan, of England, has advanced the ingenious theory, which may reconcile the above-mentioned antagonistic views, that somatic variations, in the direction of adaptation, pave the way for germinal variations, so that, while somatic modifications as such are not inherited, they are yet the favoring conditions under which germinal variations are preserved by the great principle of natural selection. If this is true, as we think it is, then we can safely state that each man in his totality is the resultant of two great factors—heredity and environment, the latter including not only food, water, climate, occupation, etc., but also the character of the civilization, the state of morals in society, the ideals and examples most frequently seen, etc., etc.
Heredity brings down to him the streams of tendency of former generations, often of a healthy and beneficent character, but also often surcharged with lust and passion, and reeking with disease.
Environment is the coöperating and, to us, vitally important factor, inasmuch as it may supplement and thus reënforce the hereditary tendencies, whether good or bad; or it may even tend to turn them into new channels, correcting the evil or vitiating the good.
Man is not simply a creature of the present, but profoundly a product of the past. Bodily structure, moral and intellectual tendencies, disease, vices, and virtues are all in the marvelous stream of heritage that comes to him from the past. “Diseases that no facts in the individual life can account for point gaunt fingers of blame from one generation to another. Not a murderer is hung, not a daughter starts on the downward way, but a great company, like those who were present at the stoning of Stephen, stand by inaugurating and consenting to the ruin.”[4]
Truly has it been said that the past is at work in the present, its powers reaching down through the ages, to all the race, largely molding every human life, touching and influencing every individual’s thought and will, and, more than any other force, coloring history.
Studies in heredity illustrate most luridly that the continuity of the human race is a terrible but remorseless reality.
If the ignorance and the perverted pleasures of one generation may produce the vices and the crimes and the diseases of another, a question of tremendous import arises: Is heredity as potent in the direction of virtue and health as of vice and disease? At the first look one is almost tempted to answer Nay! for the most striking examples of heredity seem to be in the direction of evil. But this is perfectly natural. Decay is always more rapid than growth. A cherry rots much more quickly than it ripens. Vice and disease spread much more quickly and widely than virtue and health. But all history and all social and medical science teach that vice and disease carry within themselves the seeds of decay, and virtue and health the seeds of endurance and growth.
Through the great Darwinian principle of natural selection, or survival of the fittest, vice and disease will become less and less predominant, and virtue and hygienic constitutions more and more disseminated.
As influencing a man’s life and character,[5] which is the stronger factor, heredity or environment? Fatalism or choice? In our opinion, as the result of long study and reading, where we have an average man of “mens sana in corpore sano,” environment will be the stronger factor whether for good or for evil—that is, in men in general, who have no organic defect, such as insanity or idiocy, and allied affections, the stronger force is environment; but in those having such defect, heredity is the controlling power, and, we may add, the destroying power.
It must be recalled, though, that the average man with a “sound mind in a sound body,” in his development to his present estate, has become possessed of a vast aggregate of diverse heritages, of varying age, strength and dignity. Some of them are so old and strong that they seem to be cast in unyielding molds, while others are so weak and recent that they fluctuate with every passing circumstance. The most dignified and important of all his heritages is that of rational volition. It is the play of this volition upon many of his other heritages that gives him the power of selecting, to a limited extent, his environment.
Every man is born into the world with a certain physical constitution, and, therefore, with a given temperament; with certain passions; with the power of judgment; and with a certain strength of will. If the power of his will be not equal to the strength of his passions, the latter will surely predominate and will display him as the slave of heredity. If he has such an organization of his nervous system that his volition is superior to his passions, he will be none the less the servant of heredity, though a being now possessed of the power of Free-Will.
Man is, to a far greater degree than is ordinarily realized, the servant of heredity. It seems to us an incontrovertible fact that every living creature, at any given moment, is swayed infinitely more by the totality of its heritages than by its environment. No one can possibly deny this so far as plants and most animals are concerned. Nor, if one look below the surface, can it be denied of the higher animals and of man. Happily, the average man, with his present constitution, has his diverse heritages so proportioned that we may repeat that his life and character (in customs, morals, and religion) are vastly more influenced by environment than by heredity.
The standards for estimating the life and character of men, namely, human customs, morals, and religions, are such recent acquisitions, geologically speaking, that they have, as yet, very slightly if at all influenced the germ-cells. They are acquired (somatic) characteristics, and not congenital (germinal) qualities. They are preëminently the creations of environment. If the infants of a Catholic family which is descended from a long line of Catholic ancestors were to be placed and retained in a purely Mohammedan environment, heredity would carry no Christian customs, morals or religion into that environment, but, on the contrary, the Mohammedan surroundings would instill new customs, different ethical ideas, and a different religion. This illustrates how very feebly indeed are germ-cells correspondingly impressed by pure acquired characters. It is almost certain that the translation of somatic changes into germinal changes is appallingly slow. As far, then, as social customs, morals, and religion are concerned, the average man is, in our opinion, infinitely more the creature of nurture than of nature. But, as far as his temperament, his emotional nature, his judgment, his strength of will, in short, his physical and therefore his mental constitution, are concerned, he is almost absolutely the creature of heredity. The equilibrium of qualities or heritages in the average man, resident in a given, stable community, is in harmony with the average customs, ethical ideas, and religious beliefs of that community. But in all stable communities there are men whose resultant of heritages, some in one direction and some in another, places them out of harmony with the average of their social environment, and they are looked upon, some as idiots, some as geniuses, some as criminals, and others as saints, and so on. So that again we may say that a man’s character in a community is the resultant of an hereditary physical constitution, and his environment. Some men may inherit such a physical constitution that in spite of the best environment they are much debased below the average man; others may possess such heritages that, notwithstanding adverse circumstances, they reach a level of character much above the average man. And there are all gradations between the two extremes.