It is half a century since the Austrian monk, Gregor Mendel, published in a provincial journal the results of his now famous breeding experiments with garden peas. They lay unnoticed until 1900, when three other breeders whose work had led them to similar conclusions, almost simultaneously discovered the work of Mendel and gave it to the world.

Breeding along the lines marked out by Mendel at once became the most popular method of attack, among those who were studying heredity. It became an extremely complicated subject, which can not be grasped without extended study, but its fundamentals can be briefly summarized.

Inherited differences in individuals, it will be admitted, are due to differences in their germ-plasms. It is convenient to think of these differences in germ-plasms (that is, differences in heredity) as being due to the presence in the germ-plasm of certain hypothetical units, which are usually referred to as factors. The factor, nowadays, is the ultimate unit of Mendelian research. Each of these factors is considered to be nearly or quite constant,—that is, it undergoes little, or no change from generation to generation. It is ordinarily resistant to "contamination" by other factors with which it may come in contact in the cell. The first fundamental principle of Mendelism, then, is the existence of relatively constant units, the Mendelian factors, as the basis for transmission of all the traits that go to make up an animal or plant.

Experimental breeding gives reason to believe that each factor has one or more alternatives, which may take its place in the mechanism of heredity, thereby changing the visible character of the individual plant or animal in which it occurs. To put the matter a little differently, one germ-cell differs from another in having alternatives present in place of some of the factors of the latter. A given germ-cell can never have more than one of the possible alternatives of each factor. These alternatives of a factor are called its allelomorphs.

Now a mature germ-cell has a single system of these factors: but when two germ-cells unite, there result from that union two kinds of cells—namely, immature germ-cells and body-cells; and both these kinds of cells contain a double system of factors, because of course they have received a single entire system from each parent. This is the second of the fundamental principles of Mendelism: that the factors are single in the mature germ-cell, but in duplicate in the body-cell (and also in the immature germ-cell).

In every cell with a double system of factors, there are necessarily present two representatives from each set of allelomorphs, but these may or may not be alike—or in technical language the individual may be homozygous, or heterozygous, as regards the given set of alternative factors. Looking at it from another angle, there is a single visible character in the plant or animal, but it is produced by a double factor, in the germ-plasm.

When the immature germ-cell, with its double system of factors, matures, it throws out half the factors, retaining only a single system: and the allelomorphic factors which then segregate into different cells are, as has been said above, ordinarily uninfluenced by their stay together.

But the allelomorphic factors are not the only ones which are segregated into different germ-cells, at the maturation of the cell; for the factors which are not alternative are likewise distributed, more or less independently of each other, so that it is largely a matter of chance whether factors which enter a cross in the same germ-cell, segregate into the same germ-cell or different ones, in the next generation. This is the next fundamental principle of Mendelism, usually comprehended under the term "segregation," although, as has been pointed out, it is really a double process, the segregation of alternative factors being a different thing from the segregation of non-alternative factors.

From this fact of segregation, it follows that as many kinds of germ-cells can be formed by an individual, as there are possible combinations of factors, on taking one alternative from each pair of allelomorphs present. In practice, this means that the possible number of different germ-cells is almost infinitely great, as would perhaps be suspected by anyone who has tried to find two living things that are just alike.

Fig. 46.—Many different lines of study have made it seem probable that much, although not all, of the heredity of an animal or plant is carried in the nucleus of the germ-cell and that in this nucleus it is further located in little rods or threads which can be easily stained so as to become visible, and which have the name of chromosomes. In the above illustration four different views of the nucleus of the germ-cell of an earthworm are shown, with the chromosomes in different stages; in section 19 each chromosome is doubled up like a hairpin. Study of the fruit-fly Drosophila has made it seem probable not only that the hypothetical factors of heredity are located in the chromosomes, but that each factor has a perfectly definite location in its chromosome; and T. H. Morgan and his associates have worked out an ingenious method of measuring the distance from either end, at which the factor lies. Photomicrograph after Foot and Strobell.

Such is the essence of Mendelism; and the reader is probably ready to admit that it is not a simple matter, even when reduced to the simplest terms. To sum up, the principal features at the base of the hypothetical structure are these:

1. There exist relatively constant units in the germ-plasm.

2. There are two very distinct relationships which these units may show to each other. Two (or more) unit factors may be alternatives in the mechanism of inheritance, indicating that one is a variation (or loss) of the other; or they may be independent of each other in the mechanism of inheritance.

3. The mature germ-cell contains a single system of independent factors (one representative from each set of alternates).

The immature germ-cells, and body-cells, have double systems of independent factors (two from each set of alternatives).

4. The double system arises simply from the union of two single systems (i. e., two germ-cells), without union or even contamination of the factors involved.

In the formation of a single system (mature germ-cells) from a double (immature germ-cells), pairs of alternates separate, passing into different germ-cells. Factors not alternates may or may not separate—the distribution is largely a matter of chance.

Such are the fundamental principles of Mendelism; but on them was early grafted a theoretical structure due mainly to the German zoÖlogist, August Weismann. To understand his part in the story, we must advert to that much mooted and too often misunderstood problem furnished by the chromosomes. (See Fig. 46.) These little rods of easily stained material, which are found in every cell of the body, were picked out by Professor Weismann as the probable carriers of heredity. With remarkable acuteness, he predicted their behavior at cell-division, the intricate nature of which is usually the despair of every beginner in biology. When Mendelian breeding, in the early years of this century, showed temporary pairing and subsequent separation of units in the germ-cell, it was soon realized that the observed facts of breeding fitted to a nicety the observed facts (predicted by Weismann) of chromosome-behavior; for at each cell-division the chromosomes, too, pair and separate again. The observed behavior of transmitted characters in animals and plants followed, in so many cases, the observed behavior of the chromosomes, that many students found it almost impossible to believe that there was no connection between the two, and Dr. Weismann's prediction, that the chromosomes are the carriers of heredity, came to be looked on as a fact, by many biologists.

But when so much of Professor Weismann's system was accepted, other parts of it went along, including a hypothetical system of "determiners" in the chromosome, which were believed to determine the development of characters in the organism. Every trait of an animal or plant, it was supposed, must be represented in the germ-plasm by its own determiner; one trait, one determiner. Did a notch in the ear run through a pedigree? Then it must be due to a determiner for a notch in the ear in the germ-plasm. Was mathematical ability hereditary? Then there must be a determiner, the expression of which was mathematical ability.

For a while, this hypothesis was of service in the development of genetics; some students even began to forget that it was a hypothesis, and to talk as if it were a fact. But the exhaustive tests of experimental breeding of plants and animals have long caused most of the advanced students of genetics to drop this simple hypothesis.

In its place stands the factorial hypothesis, evolved by workers in America, England, and France at about the same time. As explained in Chapter V, this hypothesis carries the assumption that every visible character is due to the effects of not one but many factors in the germ-cell.

In addition to these fundamentals, there are numerous extensions and corollaries, some of them of a highly speculative nature. The reader who is interested in pursuing the subject farther must turn to one of the text-books on Mendelism.

In plant-breeding a good deal of progress has been made in the exact study of Mendelian heredity; in animal breeding, somewhat less; in human heredity, very little. The reason is obvious: that experiments can not be made in man, and students must depend on the results of such matings as they can find; that only a very few offspring result from each mating; and that generations are so long that no one observer can have more than a few under his eyes. These difficulties make Mendelian research in man a very slow and uncertain matter.

Altogether, it is probable that something like a hundred characters in man have been pointed out as inherited in Mendelian fashion. A large part of these are pathological conditions or rare abnormalities.

But the present writers can not accept most of these cases. It has been pointed out in Chapter V that there are good reasons for doubting that feeble-mindedness is inherited in a simple Mendelian fashion, although it is widely accepted as such. We can not help feeling that in most cases heredity in man is being made to appear much simpler than it really is; and that particularly in mental characters, analysis of traits has by no means reached the bottom.

If we were asked to make out a list of characters, as to the Mendelian inheritance of which there could be little doubt, we would hardly be able to go farther than the following:

The sex-linked characters (one kind of color-blindness, hemophilia, one kind of night-blindness, atrophy of the optic nerve, and a few other rare abnormalities).

Albinism. This appears to be a recessive, but probably involves multiple allelomorphs in man, as in other animals.

Brachydactyly, apparently a dominant. This is so much cited in text-books on Mendelism that the student might think it is a common character. As a fact, it is extremely rare, being found in only a few families. The similar trait of orthodactyly or symphalangism, which likewise appears to be a good Mendelian dominant, seems to exist in only one family. Traits like these, which are easily defined and occur very rarely, make up a large part of the cases of probably Mendelian heredity. They are little more than curiosities, their rarity and abnormal nature depriving them of evolutionary significance other than to demonstrate that Mendelian heredity does operate in man.

White blaze in the hair or, as it might better be called to show its resemblance to the trait found in other mammals, piebaldism. A rather rare dominant.[204]

Huntington's Chorea, which usually appears to be a good dominant, although the last investigators (Muncey and Davenport) found some unconformable cases.

A few abnormalities, such as a premature graying of the hair (one family cited by K. Pearson) are well enough attested to be admitted. Many others, such as baldness, are probably Mendelian but not yet sufficiently supported by evidence.

None of these characters, it will be observed, is of much significance eugenically. If the exact manner of inheritance of some of the more important mental and physical traits were known, it would be of value. But it is not a prerequisite for eugenic action. Enough is known for a working program.

To sum up: the features in the modern view of heredity, which the reader must keep in mind, are the following:

1. That the various characters which make up the physical constitution of any individual plant or animal are due to the action (concurrently with the environment, of course) of what are called, for convenience, factors, separable hypothetical units in the germ-plasm, capable of independent transmission.

2. That each visible character is due to the coÖperative action of an indefinitely large number of factors; conversely, that each of these factors affects an indefinitely large number of characters.