Mendelism Part 1

Mendelism.

by Reginald Crundall Punnett.

PREFACE

A few years ago I published a short sketch of Mendel's discovery in heredity, and of some of the recent experiments which had arisen from it.

Since then progress in these studies has been rapid, and the present account, though bearing the same t.i.tle, has been completely rewritten. A number of ill.u.s.trations have been added, and here I may acknowledge my indebtedness to Miss Wheldale for the two coloured plates of sweet peas, to the Hon. Walter Rothschild for the b.u.t.terflies figured on Plate VI., to Professor Wood for photographs of sheep, and to Dr. Drinkwater for the figures of human hands. To my former publishers also, Messrs. Bowes and Bowes, I wish to express my thanks for the courtesy with which they acquiesced in my desire that the present edition should be published elsewhere.

As the book is intended to appeal to a wide audience, I have not attempted to give more experimental instances than were necessary to ill.u.s.trate the story, nor have I burdened it with bibliographical reference. The reader who desires further information may be referred to Mr. Bateson's indispensable Volume on _Mendel's {vi} Principles of Heredity_ (Cambridge, 1909), where a full account of these matters is readily accessible. Neither have I alluded to recent cytological work in so far as it may bear upon our problems. Many of the facts connected with the division of the chromosomes are striking and suggestive, but while so much difference of opinion exists as to their interpretation they are hardly suited for popular treatment.

In choosing typical examples to ill.u.s.trate the growth of our ideas it was natural that I should give the preference to those with which I was most familiar. For this reason the book is in some measure a record of the work accomplished by the Cambridge School of Genetics, and it is not unfair to say that under the leadership of William Bateson the contributions of this school have been second to none. But it should not be forgotten that workers in other European countries, and especially in America, have ama.s.sed a large and valuable body of evidence with which it is impossible to deal in a small volume of this scope.

It is not long since the English language was enriched by two new words--Eugenics and Genetics--and their similarity of origin has sometimes led to confusion between them on the part of those who are innocent of Greek. Genetics is the term applied to the experimental study of heredity and variation in animals and plants, and the main concern of its students is the establishing of law and order among the phenomena {vii} there encountered. Eugenics, on the other hand, deals with the improvement of the human race under existing conditions of law and sentiment. The Eugenist has to take into account the religious and social beliefs and prejudices of mankind. Other issues are involved besides the purely biological one, though as time goes on it is coming to be more clearly recognised that the Eugenic ideal is sharply circ.u.mscribed by the facts of heredity and variation, and by the laws which govern the transmission of qualities in living things. What these facts, what these laws are, in so far as we at present know them, I have endeavoured to indicate in the following pages; for I feel convinced that if the Eugenist is to achieve anything solid it is upon them that he must primarily build. Little enough material, it is true, exists at present, but that we now see to be largely a question of time and means. Whatever be the outcome, whatever the form of the structure which is eventually to emerge, we owe it first of all to Mendel that the foundations can be well and truly laid.

R. C. P.

CAMBRIDGE, _March, 1911_.

For although it be a more new and difficult way, to find out the nature of things, by the things themselves; then by reading of Books, to take our knowledge upon trust from the opinions of Philosophers: yet must it needs be confessed, that the former is much more open, and lesse fraudulent, especially in the Secrets relating to _Natural Philosophy_.

WILLIAM HARVEY, _Anatomical Exercitations_, 1653.

CHAPTER I

THE PROBLEM

A curious thing in the history of human thought so far as literature reveals it to us is the strange lack of interest shown in one of the most interesting of all human relationships. Few if any of the more primitive peoples seem to have attempted to define the part played by either parent in the formation of the offspring, or to have a.s.signed peculiar powers of transmission to them, even in the vaguest way. For ages man must have been more or less consciously improving his domesticated races of animals and plants, yet it is not until the time of Aristotle that we have clear evidence of any hypothesis to account for these phenomena of heredity. The production of offspring by man was then held to be similar to the production of a crop from seed. The seed came from the man, the woman provided the soil. This remained the generally accepted view for many centuries, and it was not until the recognition of woman as more than a pa.s.sive agent that the physical basis of heredity became established. That recognition was effected by the microscope, for only with its advent was actual {2} observation of the minute s.e.xual cells made possible. After more than a hundred years of conflict lasting until the end of the eighteenth century, scientific men settled down to the view that each of the s.e.xes makes a definite material contribution to the offspring produced by their joint efforts. Among animals the female contributes the ovum and the male the spermatozoon; among plants the corresponding cells are the ovules and pollen grains.

As a general rule it may be stated that the reproductive cells produced by the female are relatively large and without the power of independent movement. In addition to the actual living substance which is to take part in the formation of a new individual, the ova are more or less heavily loaded with the yolk substance that is to provide for the nutrition of the developing embryo during the early stages of its existence. The size of the ova varies enormously in different animals. In birds and reptiles where the contents of the egg form the sole resources of the developing young they are very large in comparison with the size of the animal which lays them.

In mammals, on the other hand, where the young are parasitic upon the mother during the earlier stages of their growth, the eggs are minute and only contain the small amount of yolk that enables them to reach the stage at which they develop the processes for attaching themselves to the wall of the maternal uterus. But whatever the differences in the size and appearance of the ova produced by different {3} animals, they are all comparable in that each is a distinct and separate s.e.xual cell which, as a rule, is unable to develop into a new individual of its species unless it is fertilised by union with a s.e.xual cell produced by the male.

The male s.e.xual cells are always of microscopic size and are produced in the generative gland or testis in exceedingly large numbers. In addition to their minuter size they differ from the ova in their power of active movement. Animals present various mechanisms by which the s.e.xual elements may be brought into juxtaposition, but in all cases some distance must be traversed in a fluid or semifluid medium (frequently within the body of the female parent) before the necessary fusion can occur. To accomplish this latter end of its journey the spermatozoon is endowed with some form of motile apparatus, and this frequently takes the form of a long flagellum, or whip-like process, by the lashing of which the little creature propels itself much as a tadpole with its tail.

In plants as in animals the female cells or ovules are larger than the pollen grains, though the disparity in size is not nearly so marked. Still they are always relatively minute cells since the circ.u.mstances of their development as parasites upon the mother plant render it unnecessary for them to possess any great supply of food yolk. The ovules are found surrounded by maternal tissue in the ovary, but through the stigma and down the pistil a {4} potential pa.s.sage is left for the male cell. The majority of flowers are hermaphrodite, and in many cases they are also self-fertilising. The anthers burst and the contained pollen grains are then shed upon the stigma. When this happens, the pollen cell slips through a little hole in its coat and bores its way down the pistil to reach an ovule in the ovary. Complete fusion occurs, and the minute embryo of a new plant immediately results. But for some time it is incapable of leading a separate existence, and, like the embryo mammal, it lives as a parasite upon its parent. By the parent it is provided with a protective wrapping, the seed coat, and beneath this the little embryo swells until it reaches a certain size, when as a ripe seed it severs its connection with the maternal organism. It is important to realise that the seed of a plant is not a s.e.xual cell but a young individual which, except for the coat that it wears, belongs entirely to the next generation. It is with annual plants in some respects as with many b.u.t.terflies. During one summer they are initiated by the union of two s.e.xual cells and pa.s.s through certain stages of larval development--the b.u.t.terfly as a caterpillar, the plant as a parasite upon its mother. As the summer draws to a close each pa.s.ses into a resting-stage against the winter cold--the b.u.t.terfly as a pupa and the plant as a seed, with the difference that while the caterpillar provides its own coat, that of the plant is provided by its mother. With the advent of spring both b.u.t.terfly and {5} plant emerge, become mature, and themselves ripen germ cells which give rise to a new generation.

Whatever the details of development, one cardinal fact is clear. Except for the relatively rare instances of parthenogenesis a new individual, whether plant or animal, arises as the joint product of two s.e.xual cells derived from individuals of different s.e.xes. Such s.e.xual cells, whether ovules or ova, spermatozoa or pollen grains, are known by the general term of GAMETES, or marrying cells, and the individual formed by the fusion or yoking together of two gametes is spoken of as a ZYGOTE. Since a zygote arises from the yoking together of two separate gametes, the individual so formed must be regarded throughout its life as a double structure in which the components brought in by each of the gametes remain intimately fused in a form of partnership. But when the zygote in its turn comes to form gametes, the partnership is broken and the process is reversed. The component parts of the dual structure are resolved, with the formation of a set of single structures, the gametes.

The life cycle of a species from among the higher plants or animals may be regarded as falling into three periods: (1) a period of isolation in the form of gametes, each a living unit incapable of further development without intimate a.s.sociation with another produced by the opposite s.e.x; (2) a period of a.s.sociation in which two gametes become yoked together into a zygote and react upon one {6} another to give rise by a process of cell division to what we ordinarily term an individual with all its various attributes and properties; and (3) a period of dissociation when the single structured gametes separate out from that portion of the double structured zygote which const.i.tutes its generative gland. What is the relation between gamete and zygote, between zygote and gamete? how are the properties of the zygote represented in the gamete, and in what manner are they distributed from the one to the other?--these are questions which serve to indicate the nature of the problem underlying the process of heredity.

Owing to their peculiar power of growth and the relatively large size to which they attain, many of the properties of zygotes are appreciable by observation. The colour of an animal or of a flower, the shape of a seed, or the pattern on the wings of a moth are all zygotic properties, and all capable of direct estimation. It is otherwise with the properties of gametes. While the difference between a black and a white fowl is sufficiently obvious, no one by inspection can tell the difference between the egg that will hatch into a black and that which will hatch into a white. Nor from a ma.s.s of pollen grains can any one to-day pick out those that will produce white from those that will produce coloured flowers.

Nevertheless, we know that in spite of apparent similarity there must exist fundamental differences among the gametes, even {7} among those that spring from the same individual. At present our only way of appreciating those differences is to observe the properties of the zygotes which they form.

And as it takes two gametes to form a zygote, we are in the position of attempting to decide the properties of two unknowns from one known.

Fortunately the problem is not entirely one of simple mathematics. It can be attacked by the experimental method, and with what measure of success will appear in the following pages.

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CHAPTER II

HISTORICAL

To Gregor Mendel, monk and abbot, belongs the credit of founding the modern science of heredity. Through him there was brought into these problems an entirely new idea, an entirely fresh conception of the nature of living things. Born in 1822 of Austro-Silesian parentage, he early entered the monastery of Brunn, and there in the seclusion of the cloister garden he carried out with the common pea the series of experiments which has since become so famous. In 1865 after eight years' work he published the results of his experiments in the _Proceedings of the Natural History Society of Brunn_, in a brief paper of some forty pages. But brief as it is the importance of the results and the lucidity of the exposition will always give it high rank among the cla.s.sics of biological literature. For thirty-five years Mendel's paper remained unknown, and it was not until 1900 that it was simultaneously discovered by several distinguished botanists. The causes of this curious neglect are not altogether without interest. Hybridisation experiments before Mendel there had been in plenty.

The cla.s.sificatory work of {9} Linnaeus in the latter half of the eighteenth century had given a definite significance to the word species, and scientific men began to turn their attention to attempting to discover how species were related to one another. And one obvious way of attacking the problem was to cross different species together and see what happened.

This was largely done during the earlier half of the nineteenth century, though such work was almost entirely confined to the botanists. Apart from the fact that plants lend themselves to hybridisation work more readily than animals, there was probably another reason why zoologists neglected this form of investigation. The field of zoology is a wider one than that of botany, presenting a far greater variety of type and structure. Partly owing to their importance in the study of medicine, and partly owing to their smaller numbers, the anatomy of the vegetable was far better known than that of the animal kingdom. It is, therefore, not surprising that the earlier part of the nineteenth century found the zoologists, under the influence of Cuvier and his pupils, devoting their entire energies to describing the anatomy of the new forms of animal life which careful search at home and fresh voyages of discovery abroad were continually bringing to light. During this period the zoologist had little inclination or inducement to carry on those investigations in hybridisation which were occupying the attention of some botanists. Nor did the efforts of the botanists afford much {10} encouragement to such work, for in spite of the labour devoted to these experiments, the results offered but a confused tangle of facts, contributing in no apparent way to the solution of the problem for which they had been undertaken. After half a century of experimental hybridisation the determination of the relation of species and varieties to one another seemed as remote as ever. Then in 1859 came the _Origin of Species_, in which Darwin presented to the world a consistent theory to account for the manner in which one species might have arisen from another by a process of gradual evolution. Briefly put, that theory was as follows: In any species of plant or animal the reproductive capacity tends to outrun the available food supply, and the resulting compet.i.tion leads to an inevitable struggle for existence. Of all the individuals born, only a portion, and that often a very small one, can survive to produce offspring. According to Darwin's theory, the nature of the surviving portion is not determined by chance alone. No two individuals of a species are precisely alike, and among the variations that occur some enable their possessors to cope more successfully with the compet.i.tive conditions under which they exist. In comparison with their less favoured brethren they have a better chance of surviving in the struggle for existence and consequently of leaving offspring. The argument is completed by the further a.s.sumption of a principle of heredity, in virtue of which offspring tend to {11} resemble their parents more than other members of the species. Parents possessing a favourable variation tend to transmit that variation to their offspring, to some in greater, to others in less degree. Those possessing it in greater degree will again have a better chance of survival, and will transmit the favourable variation in even greater degree to some of their offspring. A compet.i.tive struggle for existence working in combination with certain principles of variation and heredity results in a slow and continuous transformation of species through the operation of a process which Darwin termed natural selection.

The coherence and simplicity of the theory, supported as it was by the great array of facts which Darwin had patiently marshalled together, rapidly gained the enthusiastic support of the great majority of biologists. The problem of the relation of species at last appeared to be solved, and for the next forty years zoologists and botanists were busily engaged in cla.s.sifying by the light of Darwin's theory the great ma.s.ses of anatomical facts which had already acc.u.mulated and in adding and cla.s.sifying fresh ones. The study of comparative anatomy and embryology received a new stimulus, for with the acceptance of the theory of descent with modification it became inc.u.mbent upon the biologist to demonstrate the manner in which animals and plants differing widely in structure and appearance could be conceivably related to one another. Thenceforward the energies of both {12} botanists and zoologists have been devoted to the construction of hypothetical pedigrees suggesting the various tracks of evolution by which one group of animals or plants may have arisen from another through a long continued process of natural selection. The result of such work on the whole may be said to have shown that the diverse forms under which living things exist to-day, and have existed in the past so far as palaeontology can tell us, are consistent with the view that they are all related by the community of descent which the accepted theory of evolution demands, though as to the exact course of descent for any particular group of animals there is often considerable diversity of opinion. It is obvious that all this work has little or nothing to do with the manner in which species are formed. Indeed, the effect of Darwin's _Origin of Species_ was to divert attention from the way in which species originate. At the time that it was put forward his explanation appeared so satisfying that biologists accepted the notions of variation and heredity there set forth and ceased to take any further interest in the work of the hybridisers. Had Mendel's paper appeared a dozen years earlier it is difficult to believe that it could have failed to attract the attention it deserved. Coming as it did a few years after the publication of Darwin's great work, it found men's minds set at rest on the problems that he raised and their thoughts and energies directed to other matters. {13}

Nevertheless one interesting and noteworthy attempt to give greater precision to the term heredity was made about this time. Francis Galton, a cousin of Darwin, working upon data relating to the breeding of Ba.s.set hounds, found that he could express on a definite statistical scheme the proportion in which the different colours appeared in successive generations. Every individual was conceived of as possessing a definite heritage which might be expressed as unity. Of this, was on the average derived from the two parents (_i.e._ from each parent), from the four grandparents, 1/8 from the eight great-grandparents, and so on. _The Law of Ancestral Heredity_, as it was termed, expresses with fair accuracy some of the statistical phenomena relating to the transmission of characters in a mixed population. But the problem of the way in which characters are distributed from gamete to zygote and from zygote to gamete remained as before. Heredity is essentially a physiological problem, and though statistics may be suggestive in the initiation of experiment, it is upon the basis of experimental fact that progress must ultimately rest. For this reason, in spite of its ingenuity and originality, Galton's theory and the subsequent statistical work that has been founded upon it failed to give us any deeper insight into the nature of the hereditary process.

While Galton was working in England the German zoologist August Weismann was elaborating the complicated {14} theory of heredity which eventually appeared in his work on _The Germplasm_ (1885), a book which will be remembered for one notable contribution to the subject. Until the publication of Weismann's work it had been generally accepted that the modifications brought about in the individual during its lifetime, through the varying conditions of nutrition and environment, could be transmitted to the offspring. In this biologists were but following Darwin, who held that the changes in the parent resulting from increased use or disuse of any part or organ were pa.s.sed on to the children. Weismann's theory involved the conception of a sharp cleavage between the general body tissues or somatoplasm and the reproductive glands or germplasm. The individual was merely a carrier for the essential germplasm whose properties had been determined long before he was capable of leading a separate existence. As this conception ran counter to the possibility of the inheritance of "acquired characters," Weismann challenged the evidence upon which it rested and showed that it broke down wherever it was critically examined. By thus compelling biologists to revise their ideas as to the inherited effects of use and disuse, Weismann rendered a valuable service to the study of genetics and did much to clear the way for subsequent research.

A further important step was taken in 1895, when Bateson once more drew attention to the problem of the origin {15} of species, and questioned whether the accepted ideas of variation and heredity were after all in consonance with the facts. Speaking generally, species do not grade gradually from one to the other, but the differences between them are sharp and specific. Whence comes this prevalence of discontinuity if the process by which they have arisen is one of acc.u.mulation of minute and almost imperceptible differences? Why are not intermediates of all sorts more abundantly produced in nature than is actually known to be the case?

Bateson saw that if we are ever to answer this question we must have more definite knowledge of the nature of variation and of the nature of the hereditary process by which these variations are transmitted. And the best way to obtain that knowledge was to let the dead alone and to return to the study of the living. It was true that the past record of experimental breeding had been mainly one of disappointment. It was true also that there was no tangible clue by which experiments might be directed in the present.

Nevertheless in this kind of work alone there seemed any promise of ultimate success.

A few years later appeared the first volume of de Vries' remarkable book on _The Mutation Theory_. From a prolonged study of the evening primrose (_Oenothera_) de Vries concluded that new varieties suddenly arose from older ones by sudden sharp steps or mutations, and not by any process involving the gradual acc.u.mulation of minute {16} differences. The number of striking cases from among widely different plants which he was able to bring forward went far to convincing biologists that discontinuity in variation was a more widespread phenomenon than had hitherto been suspected, and not a few began to question whether the account of the mode of evolution so generally accepted for forty years was after all the true account. Such in brief was the outlook in the central problem of biology at the time of the rediscovery of Mendel's work.

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CHAPTER III

MENDEL'S WORK

The task that Mendel set before himself was to gain some clear conception of the manner in which the definite and fixed varieties found within a species are related to one another, and he realised at the outset that the best chance of success lay in working with material of such a nature as to reduce the problem to its simplest terms. He decided that the plant with which he was to work must be normally self-fertilising and unlikely to be crossed through the interference of insects, while at the same time it must possess definite fixed varieties which bred true to type. In the common pea (_Pisum sativum_) he found the plant he sought. A hardy annual, prolific, easily worked, _Pisum_ has a further advantage in that the insects which normally visit flowers are unable to gather pollen from it and so to bring about cross fertilisation. At the same time it exists in a number of strains presenting well-marked and fixed differences. The flowers may be purple, or red, or white; the plants may be tall or dwarf; the ripe seeds may be yellow or green, round or wrinkled--such are a few of the characters in which the various races of peas differ from one another. {18}

In planning his crossing experiments Mendel adopted an att.i.tude which marked him off sharply from the earlier hybridisers. He realised that their failure to elucidate any general principle of heredity from the results of cross fertilisation was due to their not having concentrated upon particular characters or traced them carefully through a sequence of generations. That source of failure he was careful to avoid, and throughout his experiments he crossed plants presenting sharply contrasted characters, and devoted his efforts to observing the behaviour of these characters in successive generations. Thus in one series of experiments he concentrated his attention on the transmission of the characters tallness and dwarfness, neglecting in so far as these experiments were concerned any other characters in which the parent plants might differ from one another. For this purpose he chose two strains of peas, one of about 6 feet in height, and another of about 1 feet. Previous testing had shown that each strain bred true to its peculiar height. These two strains were artificially crossed[1] with one another, and it was found to make no difference which was used as the pollen parent and which was used as the ovule parent. In either case the result was the same. The result of crossing tall with dwarf was in every case nothing but talls, as tall or even a little taller than the tall parent. For this reason Mendel termed tallness the DOMINANT and {19} dwarfness the RECESSIVE character. The next stage was to collect and sow the seeds of these tall hybrids. Such seeds in the following year gave rise to a mixed generation consisting of talls and dwarfs _but no intermediates_. By raising a considerable number of such plants Mendel was able to establish the fact that the number of talls which occurred in this generation was almost exactly three times as great as the number of the dwarfs. As in the previous year, seed were carefully collected from this, the second hybrid generation, and in every case _the seeds from each individual plant were harvested separately and separately sown in the following year_. By this respect for the individuality of the different plants, however closely they resembled one another, Mendel found the clue that had eluded the efforts of all his predecessors. The seeds collected from the dwarf recessives bred true, giving nothing but dwarfs. And this was true for every dwarf tested. But with the talls it was quite otherwise.

Although indistinguishable in appearance, some of them bred true, while others behaved like the original tall hybrids, giving a generation consisting of talls and dwarfs in the proportion of three of {20} the former to one of the latter. Counting showed that the number of the talls which gave dwarfs was double that of the talls which bred true.

T D --------------------- P T(D) --------------------- F_1 +-----------+-----------+-------------+ T T(D) T(D) D --- F_2 +---+----+--+ +--+----+---+ T T T(D) T(D) D T T(D) T(D) D D --- F_3 T D --- F_4

If we denote a dwarf plant as D, a true breeding tall plant as T, and a tall which gives both talls and dwarfs in the ratio 3 : 1 as T(D), the result of these experiments may be briefly summarised in the foregoing scheme.[2]

Mendel experimented with other pairs of contrasted characters and found that in every instance they followed the same scheme of inheritance. Thus coloured flowers were dominant to white, in the ripe seeds yellow was dominant to green, and round shape was dominant to wrinkled, and so on. In every case where the inheritance of an alternative pair of characters was concerned the effect of the cross in successive generations was to produce three and only three different sorts of individuals, viz. dominants which bred true, dominants which gave both dominant and recessive offspring in the ratio 3 : 1, and recessives which always bred true. Having determined a general scheme of inheritance which experiment showed to hold good for each of the seven pairs of alternative characters with which he worked, Mendel set himself to providing a theoretical interpretation of this scheme which, as he clearly realised, must be in terms of germ cells. He {21} conceived of the gametes as bearers of something capable of giving rise to the characters of the plant, but he regarded any individual gamete as being able to carry one and one only of any alternative pair of characters. A given gamete could carry tallness _or_ dwarfness, but not both. The two were mutually exclusive so far as the gamete was concerned. It must be pure for one or the other of such a pair, and this conception of the purity of the gametes is the most essential part of Mendel's theory.

[Ill.u.s.tration: FIG. 1.