Popular Science Monthly/Volume 76/May 1910/Heredity




MAY, 1910


By Professor W. E. CASTLE


THE conservation movement now in progress has for its end to preserve for future generations of men the natural resources of the earth. But it goes without saying that the movement is useless unless there are to be future generations of men capable of utilizing those resources. Thoughtful persons are beginning to wonder whether this is assured. Man is the product of two sets of agencies which we summarize in the terms heredity and environment. The question has often been asked which of these is the more important, but with this we need not concern ourselves. Both are indispensable. Seed and soil combined assure a harvest, but if either is lacking no harvest can be expected.

The public is awakening to the importance of providing mankind with a proper environment through the agencies of sanitation, education and good government, and this is well. This assures a suitable soil in which a crop of healthy human beings may develop. But what of the seed? This question has not yet been seriously considered. Only in England has it been more than suggested. There Francis Galton and his associates in the eugenics movement have started an inquiry as to why it is that the average physical condition of the English nation is declining although more and more attention is constantly being given to improving the environment. Likewise in Germany statistics show a steadily declining proportion of the young men fit for military service. There is a suspicion in the minds of many, that these nations are producing the new generation of citizens chiefly from inferior family and racial stocks. If this is so the remedy is obvious, though how easy of application remains to be ascertained. Would a farmer expect to have full harvests if each year he saved seed from the poorest yielding plants, or could he hope to secure the best results from his herds by selling or butchering the best stock and keeping only the scrubs? Obviously not, and no more can the civilized nations maintain their present standards of manhood if they follow a like practise.

But before any serious attempt can be made to improve the human race considered as an assemblage of animals possessed of certain desirable physical and intellectual attributes, it is obvious that we must know something about heredity in general, and how in particular each of the desired physical and intellectual attributes is produced. Considerations such as these lend general interest to the study of heredity, a subject which has always been of great practical concern to farmers, and of much theoretical interest to scientists. It is my purpose to review briefly some of the problems which the study of heredity presents, and some of the results obtained from their consideration.

"Like father like son" is a homely proverb which shows how general the recognition is that children resemble their parents. Resemblances to grandparents or ancestors even more remote are also of frequent occurrence, and it is convenient to use the term heredity as including all such resemblances, whether to near or to remote ancestors. The phenomenon of heredity is of course not restricted to human society. Heredity has for the stockman and plant-breeder a well-recognized commercial value, because by a knowledge of its laws he is enabled to produce in greater number or with greater certainty animals or plants of a particular type. Indeed, much of our present knowledge of heredity has been derived from a study of the domesticated animals or of the cultivated plants, and from the same sources we may expect to continue to draw, for here alone have we an unobstructed field for observation and experiment, the indispensable tools of scientific research. Just as the sciences of anatomy, embryology, physiology and pathology progressed but slowly so long as the phenomena of the human body alone were considered, but advanced by leaps and bounds when comparative studies on other animals were undertaken, so concerning heredity in man we have learned and can expect to learn but little from the study of man alone, but much from a study of other animals and of plants and from a comparison of the phenomena in the two cases.

Every new individual arises out of material derived exclusively from its parents. This is the basis of heredity. But it does not follow that the new individual will resemble its parents merely. It may resemble remote ancestors more strongly than either parent. For it represents a combination of materials or of qualities derived from the two parents and it is possible that neither parent may manifest all the peculiarities which it transmits to the offspring. For the parent is made up of two distinct parts, its own body and the reproductive substance contained within that body, and the two may not be identical in character.

The reproductive substance has been called by Weismann the germ-plasm. He it was who first clearly recognized the fact that the germ-plasm is distinct from the body which contains it, and that the influences which modify the character of the one do not of necessity modify the character of the other. Thus he was able to show experimentally that mutilations of the body, as loss of the tail in mice, are not inherited, and to establish with a considerable degree of certainty the principle that characters acquired by the body as a result of use, disuse or other agencies are not inherited, because they have not affected the constitution of the germ-plasm carried within the body.

Weismann's two principles are of fundamental importance to a right understanding of heredity. They are: (1) That the germ-plasm is independent of the body containing it, or, as Weismann put it, that the germ-plasm is continuous from generation to generation, whereas the body dies, and (2) that acquired characters are not inherited.

The hottest biological discussions of the last twenty years have been waged over these two principles and the contest is by no means ended, but year by year the correctness of Weismann's contentions is more generally admitted.

Common experiences support both principles. Thus the independence or continuity of the germ-plasm has been shown from time prehistoric in the practise of castration upon the domesticated animals or upon man. The germ-plasm is localized in particular organs of the body, the reproductive glands. If these are removed reproduction becomes impossible, though all other functions of the individual persist. Further, it is possible to show experimentally that the germ-plasm

PSM V76 D423 Three week old guinea pig.png

Fig. 1. A young black Guinea-pig, about three weeks old. The ovaries taken from an animal like this were transplanted into the albino shown in Fig. 2.

PSM V76 D424 Albino female guinea pig.png

Fig. 2. An Albino female Guinea-fig. Its ovaries were removed and in their place were introduced ovaries from a young black guinea-pig. Compare Fig. 1.

transplanted from one individual into another retains the character which it originally had, quite unaffected by the changed body with which it is associated. This Dr. John C. Phillips and the writer have recently shown in the following way. The ovaries were removed from a young black guinea-pig, Fig. 1, and these were transplanted into the body of a white guinea-pig, previously castrated, Fig. 2. The white guinea-pig was now mated with another white guinea-pig, Fig. 3. Normal white guinea-pigs produce only white offspring when mated with each other, but these two have now produced in three successive litters six young, all black. Three of these are shown in Fig. 4. Evidently the germ-plasm of the black guinea-pig retained its original character even after transplantation into the body of a white one.

In order better to understand the processes of heredity we should be familiar with what takes place when a new individual is formed. The new individual, whether an animal or a plant, has its beginning in the union of two bits of germ-plasm, an egg cell furnished by the mother and a sperm cell furnished by the father. Whether the union of the germ-plasm takes place within the maternal body or not is quite immaterial; among a great many animals it does not.

The new individual, it will be observed, is dual in origin, and to its dying day it retains a dual nature. For the maternal and paternal contributions of germ-plasm retain a certain distinctness as we shall see, and may in part separate from each other at reproduction.

Each germ-cell (egg or sperm), so far as its contribution to heredity is concerned, stands for a complete organism of its species, bears the potentialities of a complete organism, and under appropriate conditions can develop into such an organism. For this idea we have strong experimental evidence. It has long been known that the eggs of certain species of animals can develop without fertilization, i. e., without having united with a sperm or male sex-cell. In such cases there can be no question that the potentialities of an entire organism are contained in the egg, for without any outside help the egg develops into a complete individual of the species. In recent years it has been shown that the eggs of many species in which fertilization normally occurs may by artificial means be made to develop without having united with a sperm. This is true of the eggs of sea-urchins, star-fishes, and of certain worms and mollusks. Such eggs artificially stimulated to development produce entire individuals, similar to those produced by fertilization, but possibly less vigorous.

On the other hand, a sperm cell may be made to develop, if it is allowed to penetrate into a fragment of an egg, even a fragment which lacks the important cell-nucleus. In such cases the entire nuclear material of the embryo is furnished by the sperm, yet the embryo so produced is complete, lacking no essential part, and similar except in size and vigor to normal embryos produced by fertilization.

Accordingly the evidence is fairly complete that each germ-cell (egg or sperm), considered as the vehicle of heredity, represents a complete organism, and that an individual produced by the union of two such germ-cells represents twice over each heritable trait of the species. In other words, the germ-cell is single, the individual is double.

This fundamental principle of the singleness of the germ in contrast

PSM V76 D425 Albino male guinea pig.png
Fig. 3. An Albino male Guinea-pig, with which was mated the albino shown in Fig. 2.
PSM V76 D426 Group of three young produced by the albino pair.png
Fig. 4. A Group of three Young, produced by the pair of albinos shown in Figs. 2 and 3.

to the doubleness (duality) of the individual receives the fullest confirmation from experimental breeding.

If we mate a pure-bred black guinea-pig with a white one, the young are all black pigmented. This result seems to violate the principle previously stated that both parents contribute equally in heredity; in reality, however, that principle is not violated. The white parent has contributed its own character to the offspring, but that contribution is unseen in them simply because black hides it. The white will reappear among the grandchildren. In Fig. 5 we see a mother guinea-pig having a jet black coat. Beside her are four young of the same color as herself. The father too was black. In a word this black race breeds true. A female of this race was mated with the albino male shown in Fig. 6. Albinos have white hair and pink (unpigmented) eyes, the red eye color being due to the blood which shows through; they breed true among themselves, but the result is very different when they are mated with black individuals. Two children of the albino male and the black female are shown in Fig. 7. They are intensely black pigmented, as are all the young produced by this cross. Two of them when grown to maturity and mated with each other, produced a litter of four young, shown in Fig. 8. Three are black pigmented like the parents, but one is an albino similar in all respects to the albino grandsire. Here we notice the reappearance of the albino character after skipping a generation. The albino grandsire really made a hereditary contribution as regards the character hair color, but it did not show in the children, because black also was present in the children, and black obscured or dominated the white.

Applying our principle of single germ, dual individual to this case, we see that the facts observed are fully in harmony with it. The original cross brought together the characters B (black) and W (white) into an individual (or zygote as we call it, a joining together) B W, which showed only black. Two such individuals, a male and a female were now mated together. In the formation of germ-cells by these individuals there is a return to the single condition, B separates from W and passes into a different germ-cell. Accordingly, the mother forms eggs, B and W, respectively, and the father forms sperms of a like character. Now a new individual arises from a union of an egg with a sperm. Apparently either sort of sperm may unite with either sort of egg which it chances to meet. So there are formed in the next generation three sorts of zygotes (individuals), viz., B B, B W, and WW, instead of B W alone as in the previous generation. The chances for the occurrence of these three sorts of unions are 1 BB, 2 BW, and 1 WW. Any individual containing the character B will be black; accordingly the BWs as well as the BBs will be black and there should be three blacks to one white. These are in fact the observed proportions. The white individual should transmit no other character, because it contains only W. Such is indeed the observed fact. Any two white individuals mated together will produce only white offspring. But, if our reasoning is correct, two thirds of the black individuals of this generation (viz., the BWs) should transmit white as well as black, while the remaining one third, BB, should transmit only black. Experiment justifies both these conclusions. If we mate the black animals
PSM V76 D428 Black female guinea pig and her young.png
Fig. 5. A black female Guinea-pig and her Young.
PSM V76 D428 Albino guinea pig father of black young.png
Fig. 6. An Albino Guinea-pig, father cf black young, like those seen in Fig. 5.

of this generation, one by one, with albinos, we find that on the average two out of three of them will produce white offspring as well as black ones, while the third one produces only black offspring.

The scientific law which governs the inheritance of albinism, and of other characters transmitted in a similar fashion, is known as Mendel's law. It applies, apparently, to all cases of color-inheritance, as well as to the inheritance of characters of many other sorts. Through its operation new combinations of the peculiar characters of individuals or of races can be obtained in the course of one or two generations. Thus when a guinea-pig showing the two coat-characters seen in Fig. 9, dark and smooth coat, is mated with one showing the combination, white and rough, Fig. 10, young are produced showing a wholly new combination, dark and rough, Fig. 11. And if these young are at maturity bred together, a fourth combination, white and smooth, appears among their young, the grandchildren. See Fig. 12. Other grandchildren manifest the combinations seen, respectively, in the parents and in the grandparents. By selection any one of these combinations may be obtained in a pure race.

Oftentimes a new combination of characters obtained through
PSM V76 D429 Offspring of a black and albino guinea pig.png
Fig. 7. Two of the grown-up young of a Black and of an Albino Guinea-pig.
PSM V76 D429 Second generation offspring of a black and albino guinea pig strain.png
Fig. 8. A Group of four Young, produced by the animals shown in Fig. 7.

crosses coincides with a lost racial combination. Then the phenomenon is called reversion or atavism. Thus when yellow rabbits are crossed with black ones, gray offspring are obtained similar to wild rabbits in coloration. There is no longer anything mysterious about the process; it is simple recombination of different unit-characters formerly associated together in the same race, but since isolated in some of the derived races.

Very different in nature, apparently, from the Mendelian inheritance of unit-characters is the result obtained when races of animals are crossed differing in size or in the proportions of their parts. In such cases the children are intermediate in character, and the grandparental conditions do not reappear among the grandchildren. The result may be described as a blend apparently permanent. Fig. 13 shows the skulls of three rabbits, all adult, father, mother and son. The skull of the son is shown between that of his parents, the mother's skull being at the right. Size and proportions of parts are clearly intermediate in the son. No grandchildren were obtained like either grandparent in size. The color of the coat in this same family of rabbits clearly followed Mendel's law, although the size characters blended. The practical result is that one may at will produce a race
PSM V76 D430 A dark smooth haired guinea pig.png
Fig. 9. A dark smooth Guinea-pig.

of rabbits of any desired size within the known limits of variation in size among rabbits, and with any of the conceivable combinations of color factors. Size variation is apparently continuous and its inheritance blending, color variation is discontinuous and its inheritance Mendelian.

Notwithstanding the seemingly radical difference between these two types of inheritance, it is possible that they may, after all, prove to have

PSM V76 D430 A white rough haired guinea pig.png
Fig. 10. A white rough Guinea-pig.
PSM V76 D431 Dark rough haired guinea pig.png
Fig. 11. A dark rough Guinea-pig, the new combination of characters obtained when animals are mated like those shown in Figs'. 9 and 10, respectively.

a common basis. Blending inheritance may possibly be only a complex sort of Mendelian inheritance, in which many independent factors are simultaneously concerned. The question is one of much theoretical interest. Its solution awaits further investigation.

PSM V76 D431 White smooth haired guinea pig.png
Fig. 12. A white smooth Guinea-pig, a second new combination of characters, but obtained first among the grandchildren of such animals as are shown in Figs. 9 and 10. Other grandchildren are like the respective grandparents (Figs. 9 and 10) or the parents (Fig. 11).
PSM V76 D432 Skulls of three rabbits.png
Fig. 13. Skulls of three Rabbits, mother (3 and 3a), father (1 and la) and son (2 and 2a). From Publication No. 114, Carnegie Institution of Washington, by permission.

What has already been accomplished in the study of heredity gives us a hopeful outlook for the future. We are gaining a fuller knowledge of its processes, and a knowledge of processes is a first step toward their control.

  1. From a lecture delivered before Section F, American Association for the Advancement of Science, December 31, 1909.