Mendelism - Third Edition
by Reginald Crundall Punnett
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Transcriber's note:

A few typographical errors have been corrected: they are listed at the end of the text.

Fig. 8 has been re-mastered to match the text (the Black boxes were shown as Albino and the heterozygous Albinos as Black).

Superscripted numbers are indicated by a carat character followed by the superscript. For example, 2^4 denotes 2 raised to the fourth power and 2^(10) denotes 2 raised to the tenth power.

Subscripted numbers are indicated by an underscore followed by the subscript. For example, in the expression "F_1" the 1 is a subscript.

Page numbers in this text file are enclosed in curly brackets. This enables the reader to use the index by searching for the page number. To find page 35, search for {35}.




Fellow of Gonville and Caius College Professor of Biology in the University Of Cambridge

THIRD EDITION Entirely Rewritten and Much Enlarged

New York The MacMillan Company 1911

All rights reserved

Copyright, 1911, by The MacMillan Company.

Set up and electrotyped. Published May, 1911.

Norwood Press J. S. Cushing Co.—Berwick & Smith Co. Norwood, Mass., U.S.A.



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 title, has been completely rewritten. A number of illustrations 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 butterflies 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 illustrate 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 illustrate 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 amassed 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 circumscribed 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.

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CHAPTER I PAGE The Problem 1

CHAPTER II Historical 8

CHAPTER III Mendel's Work 17

CHAPTER IV The Presence and Absence Theory 29

CHAPTER V Interaction of Factors 42

CHAPTER VI Reversion 59

CHAPTER VII Dominance 68

{x} CHAPTER VIII Wild Forms and Domestic Varieties 79

CHAPTER IX Repulsion and Coupling of Factors 88


CHAPTER XI Sex (continued) 115

CHAPTER XII Intermediates 125

CHAPTER XIII Variation and Evolution 135

CHAPTER XIV Economical 153




* * * * *





Gregor Mendel Frontispiece

I. Rabbits To face 60

II. Sweet Peas " 64

III. Sheep " 78

IV. Sweet Peas " 80

V. Fowls " 107

VI. Butterflies " 146



1. Scheme of Inheritance in simple Mendelian Case 21

2. Feathers of Silky and Common Fowl 30

3. Single and Double Primulas 31

4. Fowls' Combs 32

5. Diagram of Inheritance of Fowls' Combs 37

6. Fowls' Combs 39

7. Diagram of F_2 Generation resulting from Cross between two White Sweet Peas 46

8. Diagram illustrating 9 : 3 : 4 Ratio in Mice 52

9. Sections of Primulas 55

{xii} 10. Small and Large-eyed Primulas 56

11. Diagram illustrating Reversion in Pigeons 67

12. Primula sinensis x Primula stellata 68

13. Diagram illustrating Cross between Dominant and Recessive White Fowls 72

14. Bearded and Beardless Wheat 75

15. Feet of Fowls 76

16. Scheme of Inheritance of Horns in Sheep 76

17. Abraxas grossulariata and var. lacticolor 99

18. Scheme of Inheritance in Abraxas 102

19. Scheme of Inheritance of Silky Hen x Brown Leghorn Cock 105

20. Scheme of Inheritance of Brown Leghorn Hen x Silky Cock 106

21. Scheme of F_1 (ex Brown Leghorn x Silky Cock) crossed with pure Brown Leghorn 107

22. Scheme for Silky Hen x Brown Leghorn Cock 108

23. Scheme for Brown Leghorn Hen x Silky Cock 109

24. Diagram illustrating Nature of Offspring from Brown Leghorn Hen x F_1 Cock 110

25. Scheme to illustrate Heterozygous Nature of Brown Leghorn Hen 111

26. Scheme of Inheritance of Colour-blindness 117

27. Single and Double Stocks 122

28. F_2 Generation ex Silky Hen x Brown Leghorn Cock 127

29. Pedigree of Eurasian Family 130

30. Curve illustrating Influence of Selection 159

{xiii} 31. Curve illustrating Conception of pure Lines 162

32. Brachydactylous and Normal Hands 170

33. Radiograph of Brachydactylous Hand 170

34. Pedigree of Brachydactylous Family 173

35. Pedigree of Haemophilic Family 175

* * * * *


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.

* * * * *




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 assigned 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 passive 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 sexual 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 sexes 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 sexual cell which, as a rule, is unable to develop into a new individual of its species unless it is fertilised by union with a sexual cell produced by the male.

The male sexual 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 sexual 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 circumstances 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 passage 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 sexual 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 butterflies. During one summer they are initiated by the union of two sexual cells and pass through certain stages of larval development—the butterfly as a caterpillar, the plant as a parasite upon its mother. As the summer draws to a close each passes into a resting-stage against the winter cold—the butterfly 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 butterfly 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 sexual cells derived from individuals of different sexes. Such sexual 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 association with another produced by the opposite sex; (2) a period of association 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 constitutes 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 mass 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.

* * * * *




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 Bruenn, 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 Bruenn, 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 classics 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 classificatory 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 competition 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 competitive 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 assumption 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 competitive 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 classifying by the light of Darwin's theory the great masses of anatomical facts which had already accumulated and in adding and classifying 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 incumbent 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 Basset 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, 1/2 was on the average derived from the two parents (i.e. 1/4 from each parent), 1/4 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 passed 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 accumulation 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 accumulation 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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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 attitude 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 1-1/2 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 x D - P T(D) - F1 - - - T T(D) T(D) D - F2 - - T T T(D) T(D) D T T(D) T(D) D D - F3 T D - F4

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.

We may now proceed with the help of the accompanying scheme (Fig. 1) to deduce the results that should flow from Mendel's conception of the nature of the gametes, and to see how far they are in accordance with the facts. Since the original tall plant belonged to a strain which bred true, all the gametes produced by it must bear the tall character. Similarly all the gametes of the original dwarf plant must bear the dwarf character. A cross between these two means the union of {22} a gamete containing tallness with one bearing dwarfness. Owing to the completely dominant nature of the tall character, such a plant is in appearance indistinguishable from the pure tall, but it differs markedly from it in the nature of the gametes to which it gives rise. When the formation of the gametes occurs, the elements representing dwarfness and tallness SEGREGATE from one another, so that half of the gametes produced contain the one, and half contain the other of these two elements. For on hypothesis every gamete must be pure for one or other of these two characters. And this is true for the ovules as well as for the pollen grains. Such hybrid F_1 plants, therefore, must produce a series of ovules consisting of those bearing tallness and those bearing dwarfness, and must produce them in equal numbers. And similarly for the pollen grains. We may now calculate what should happen when such a series of pollen grains meets such a series of ovules, _i.e._ the nature of the generation that should be produced when the hybrid is allowed to fertilise itself. Let us suppose that there are 4x ovules so that 2x are "tall" and 2x are "dwarf." These are brought in contact with a mass of pollen grains of which half are "tall" and half are "dwarf." It is obvious that a "tall" ovule has an equal chance of being fertilised by a "tall" or a "dwarf" pollen grain. Hence of our 2x "tall" ovules, x will be fertilised by "tall" pollen grains and x will be fertilised by "dwarf" pollen grains. The former must give rise to tall {23} plants, and since the dwarf character has been entirely eliminated from them they must in the future breed true. The latter must also give rise to tall plants, but since they carry also the recessive dwarf character they must when bred from produce both tails and dwarfs. Each of the 2x dwarf ovules, again, has an equal chance of being fertilised by a "tall" or by a "dwarf" pollen grain. Hence x will give rise to tall plants carrying the recessive dwarf character, while x will produce plants from which the tall character has been eliminated, _i.e._ to pure recessive dwarfs. Consequently from the 4x ovules of the self-fertilised hybrid we ought to obtain 3x tall and x dwarf plants. And of the 3x talls x should breed true to tallness, while the remaining 2x, having been formed like the original hybrid by the union of a "tall" and a "dwarf" gamete, ought to behave like it when bred from and give talls and dwarfs in the ratio 3 : 1. Now this is precisely the result actually obtained by experiment (cf. p. 17), and the close accord of the experimental results with those deduced on the assumption of the purity of the gametes as enunciated by Mendel affords the strongest of arguments for regarding the nature of the gametes and their relation to the characters of the zygotes in the way that he has done.

It is possible to put the theory to a further test. The explanation of the 3 : 1 ratio of dominants and recessives in the F_2 generation is regarded as due to the F_1 individuals producing equal numbers of gametes bearing the {24} dominant and recessive elements respectively. If now the F_1 plant be crossed with the pure recessive, we are bringing together a series of gametes consisting of equal numbers of dominants and recessives with a series consisting solely of recessives. We ought from such a cross to obtain equal numbers of dominant and recessive individuals, and further, the dominants so produced ought all to give both dominants and recessives in the ratio 3 : 1 when they themselves are bred from. Both of these expectations were amply confirmed by experiment, and crossing with the recessive is now a recognised way of testing whether a plant or animal bearing a dominant character is a pure dominant, or an impure dominant which is carrying the recessive character. In the former case the offspring will be all of the dominant form, while in the latter they will consist on the average of equal numbers of dominants and recessives.

So far we have been concerned with the results obtained when two individuals differing in a single pair of characters are crossed together and with the interpretation of those results. But Mendel also used plants which differed in more than a single pair of differentiating characters. In such cases he found that each pair of characters followed the same definite rule, but that the inheritance of each pair was absolutely independent of the other. Thus, for example, when a tall plant bearing coloured flowers was crossed with a dwarf plant {25} bearing white flowers the resulting hybrid was a tall plant with coloured flowers. For coloured flowers are dominant to white, and tallness is dominant to dwarfness. In the succeeding generation there are plants with coloured flowers and plants with white flowers in the proportion of 3 : 1, and at the same time tall plants and dwarf plants in the same proportion. Hence the chances that a tall plant will have coloured flowers are three times as great as its chance of having white flowers. And this is also true for the dwarf plants. As the result of this cross, therefore, we should expect an F2 generation consisting of four classes, viz. coloured talls, white talls, coloured dwarfs, and white dwarfs, and we should further expect these four forms to appear in the ratio of 9 coloured talls, 3 white talls, 3 coloured dwarfs, and 1 white dwarf. For this is the only ratio which satisfies the conditions that the talls should be to the dwarfs as 3 : 1, and at the same time the coloured should be to the whites as 3 : 1. And these are the proportions that Mendel found to obtain actually in his experiments. Put in a more general form, it may be stated that when two individuals are crossed which differ in two pairs of differentiating characters the hybrids (F1) are all of the same form, exhibiting the dominant character of each of the two pairs, while the F2 generation produced by such hybrids consists on the average of 9 showing both dominants, 3 showing one dominant and one recessive, {26} 3 showing the other dominant and the other recessive, and 1 showing both recessive characters. And, as Mendel pointed out, the principle may be extended indefinitely. If, for example, the parents differ in three pair of characters A, B, and C, respectively dominant to a, b, and c, the F1 individuals will be all of the form ABC, while the F2 generation will consists of 27 ABC, 9 ABc, 9 AbC, 9 aBC, 3 Abc, 3 aBc, 3 abC, and 1 abc. When individuals differing in a number of alternative characters are crossed together, the hybrid generation, provided that the original parents were of pure strains, consists of plants of the same form; but when these are bred from a redistribution of the various characters occurs. That redistribution follows the same definite rule for each character, and if the constitution of the original parents be known, the nature of the F2 generation, i.e. the number of possible forms and the proportions in which they occur, can be readily calculated. Moreover, as Mendel showed, we can calculate also the chances of any given form breeding true. To this point, however, we shall return later.

Of Mendel's experiments with beans it is sufficient to say here that they corroborated his more ample work with peas. He is also known to have made experiments with many other plants, and a few of his results are incidentally given in his series of letters to Naegeli the botanist. To the breeding and crossing of bees he also devoted much {27} time and attention, but unhappily the record of these experiments appears to have been lost. The only other published work that we possess dealing with heredity is a brief paper on some crossing experiments with the hawkweeds (_Hieracium_), a genus that he chose for working with because of the enormous number of forms under which it naturally exists. By crossing together the more distinct varieties, he evidently hoped to produce some of these numerous wild forms, and so throw light upon their origin and nature. In this hope he was disappointed. Owing in part to the great technical difficulties attending the cross fertilisation of these flowers he succeeded in obtaining very few hybrids. Moreover, the behaviour of those which he did obtain was quite contrary to what he had found in the peas. Instead of giving a variety of forms in the F_2 generation, they bred true and continued to do so as long as they were kept under observation. More recent research has shown that this is due to a peculiar form of parthenogenesis (cf. p. 135), and not to any failure of the characters to separate clearly from one another in the gametes. Mendel, however, could not have known of this, and his inability to discover in _Hieracium_ any indication of the rule which he had found to hold good for both peas and beans must have been a source of considerable disappointment. Whether for this reason, or owing to the utter neglect of his work by the scientific world, Mendel gave up his experimental {28} researches during the latter part of his life. His closing years were shadowed with ill-health and embittered by a controversy with the Government on a question of the rights of his monastery. He died of Bright's disease in 1884.

Note.—Shortly after the discovery of Mendel's paper a need was felt for terms of a general nature to express the constitution of individuals in respect of inherited characters, and Bateson accordingly proposed the words homozygote and heterozygote. An individual is said to be homozygous for a given character when it has been formed by two gametes each bearing the character, and all the gametes of a homozygote bear the character in respect of which it is homozygous. When, however, the zygote is formed by two gametes of which one bears the given character while the other does not, it is said to be heterozygous for the character in question, and only half the gametes produced by such a heterozygote bear the character. An individual may be homozygous for one or more characters, and at the same time may be heterozygous for others.

* * * * *




It was fortunate for the development of biological science that the rediscovery of Mendel's work found a small group of biologists deeply interested in the problems of heredity, and themselves engaged in experimental breeding. To these men the extraordinary significance of the discovery was at once apparent. From their experiments, undertaken in ignorance of Mendel's paper, de Vries, Correns, and Tschermak were able to confirm his results in peas and other plants, while Bateson was the first to demonstrate their application to animals. Thenceforward the record has been one of steady progress, and the result of ten years' work has been to establish more and more firmly the fundamental nature of Mendel's discovery. The scheme of inheritance, which he was the first to enunciate, has been found to hold good for such diverse things as height, hairiness, and flower colour and flower form in plants, the shape of pollen grains, and the structure of fruits; while among animals the coat colour of mammals, the form of the feathers and of the comb in poultry, the waltzing habit of Japanese mice, and eye {30} colour in man are but a few examples of the diversity of characters which all follow the same law of transmission. And as time went on many cases which at first seemed to fall without the scheme have been gradually brought into line in the light of fuller knowledge. Some of these will be dealt with in the succeeding chapters of this book. Meanwhile we may concern ourselves with the single modification of Mendel's original views which has arisen out of more ample knowledge.

As we have already seen, Mendel considered that in the gamete there was either a definite something {31} corresponding to the dominant character or a definite something corresponding to the recessive character, and that these somethings whatever they were could not coexist in any single gamete. For these somethings we shall in future use the term FACTOR. The factor, then, is what corresponds in the gamete to the UNIT-CHARACTER that appears in some shape or other in the development of the zygote. Tallness in the pea is a unit-character, and the gametes in which it is represented are said to contain the factor for tallness. Beyond their existence in the gamete and their mode of transmission we make no suggestion as to the nature of these factors.


On Mendel's view there was a factor corresponding to the dominant character and another factor corresponding to the recessive character of each alternative pair of unit-characters, and the characters were alternative because no gamete could carry more than one of the two factors belonging to the alternative pair. On the other hand, Mendel supposed that it always carried either one or the other of such a pair. As experimental work proceeded, {33} it soon became clear that there were cases which could not be expressed in terms of this conception. The nature of the difficulty and the way in which it was met will perhaps be best understood by considering a set of experiments in which it occurred. Many of the different breeds of poultry are characterised by a particular form of comb, and in certain cases the inheritance of these has been carefully worked out. It was shown that the rose comb (Fig. 4, B) with its flattened papillated upper surface and backwardly projecting pike was dominant in the ordinary way to the deeply serrated high single comb (Fig. 4, C) which is characteristic of the Mediterranean races. Experiment also showed that the pea comb (Fig. 4, A), a form with a low central and two well-developed lateral ridges, such as is found in Indian game, behaves as a simple dominant to the single comb. The interesting question arose as to what would happen when the rose and the pea, two forms each dominant to the same third form, were mated together. It seemed reasonable to suppose that things which were alternative to the same thing would be alternative to one another—that either rose or pea would dominate in the hybrids, and that the F_2 generation would consist of dominants and recessives in the ratio 3 : 1. The result of the experiment was, however, very different. The cross rose x pea led to the production of a comb quite unlike either of them. This, the so-called walnut comb (Fig. 4, D), {34} from its resemblance to the half of a walnut, is a type of comb which is normally characteristic of the Malay fowl. Moreover, when these F_1 birds were bred together, a further unlooked-for result was obtained. As was expected, there appeared in the F_2 generation the three forms walnut, rose, and pea. But there also appeared a definite proportion of single-combed birds, and among many hundreds of chickens bred in this way the proportions in which the four forms walnut, rose, pea, and single appeared was 9 : 3 : 3 : 1.

Rose x Pea + Walnut x Walnut + - - Walnut Rose Pea Single (9) (3) (3) (1)

Now this, as Mendel showed, is the ratio found in an F_2 generation when the original parents differ in two pairs of alternative characters, and from the proportions in which the different forms of comb occur we must infer that the walnut contains both dominants, the rose and the pea one dominant each, while the single is pure for both recessive characters. This accorded with subsequent breeding experiments, for the singles bred perfectly true as soon as they had once made their appearance. So far the case is clear. The difficulty comes when we attempt to define these two pairs of characters. How are we to express the fact that while single behaves as a simple recessive to either pure rose, or to pure pea, it can yet appear in F_2 from a cross {35} between these two pure forms, though neither of them should, on Mendel's view, contain the single? An explanation which covers the facts in a simple way is that which has been termed the "Presence and Absence" theory. On this theory the dominant character of an alternative pair owes its dominance to the presence of a factor which is absent in the recessive. The tall pea is tall owing to the presence in it of the factor for tallness, but in the absence of this factor the pea remains a dwarf. All peas are dwarf, but the tall is a dwarf plus a factor which turns it into a tall. Instead of the characters of an alternative pair being due to two separate factors, we now regard them as the expression of the only two possible states of a single factor, viz. its presence or its absence. The conception will probably become clearer if we follow its application in detail to the case of the fowl's combs. In this case we are concerned with the transmission of the two factors, rose (R) and pea (P), the presence of each of which is alternative to its absence. The rose-combed bird contains the factor for rose but not that for pea, and the pea-combed bird contains the factor for pea but not that for rose. When both factors are present in a bird, as in the hybrid made by crossing rose with pea, the result is a walnut. For convenience of argument we may denote the presence of a given factor by a capital letter and its absence by the corresponding small letter. The use of the small letter is merely a symbolic way of intimating {36} that a particular factor is absent in a gamete or zygote. Represented thus the zygotic constitution of a pure rose-combed bird is RRpp; for it has been formed by the union of two gametes both of which contained R but not P. Similarly we may denote the pure pea-combed bird as rrPP. On crossing the rose with the pea union occurs between a gamete Rp and a gamete rP, resulting in the formation of a heterozygote of the constitution RrPp. The use of the small letters here informs us that such a zygote contains only a single dose of each of the factors R and P, although, of course, it is possible for a zygote, if made in a suitable way, to have a double dose of any factor. Now when such a bird comes to form gametes a separation takes place between the part of the zygotic cell containing R and the part which does not contain it (r). Half of its gametes, therefore, will contain R and the other half will be without it (r). Similarly half of its gametes will contain P and the other half will be without it (p). It is obvious that the chances of R being distributed to a gamete with or without P are equal. Hence the gametes containing R will be of two sorts, RP and Rp, and these will be produced in equal numbers. Similarly the gametes without R will also be of two sorts, rP and rp, and these, again, will be produced in equal numbers. Each of the hybrid walnut-combed birds, therefore, gives rise to a series consisting of equal numbers of gametes of the four different types RP, Rp, rP, and rp; and the breeding {37} together of such F_1 birds means the bringing together of two such series of gametes. When this happens an ovum of any one of the four types has an equal chance of being fertilised by a spermatozoon of any one of the four types. A convenient and simple method of demonstrating what happens under such circumstances is the method sometimes termed the "chessboard" method. For two series each consisting of four different types of gamete we require a square divided up into 16 parts. The four terms of the gametic series are first written horizontally across the four sets of four squares, so that the series is repeated four times. It is then written vertically four times, care being taken to keep to the same order. In this simple mechanical way all the possible combinations are represented and in their proper proportions.

+ -+ -+ -+ -+ RP RP RP RP RP Rp rP rp Walnut Walnut Walnut Walnut + -+ -+ -+ -+ Rp Rp Rp Rp RP Rp rP rp Walnut Rose Walnut Rose + -+ -+ -+ -+ rP rP rP rP RP Rp rP rp Walnut Walnut Pea Pea + -+ -+ -+ -+ rp rp rp rp RP Rp rP rp Walnut Rose Pea Single + -+ -+ -+ -+

FIG. 5.

Diagram to illustrate the nature of the F_2 generation from the cross of rose comb x pea comb.

Fig. 5 shows the result of applying this method to our series RP, Rp, rP, rp, and the 16 squares represent the different kinds of zygotes formed and the proportions in which they occur. As {38} the figure shows, 9 zygotes contain both R and P, having a double or a single dose of either or both of these factors. Such birds must be all walnut combed. Three out of the 16 zygotes contain R but not P, and these must be rose-combed birds. Three, again, contain P but not R and must be pea-combed birds. Finally one out of the 16 contains neither R nor P. It cannot be rose—it cannot be pea. It must, therefore, be something else. As a matter of fact it is single. Why it should be single and not something else follows from what we already know about the behaviour of these various forms of comb. For rose is dominant to single; therefore on the Presence and Absence theory a rose is a single plus a factor which turns the single into a rose. If we could remove the "rose" factor from a rose-combed bird the underlying single would come into view. Similarly a pea comb is a single plus a factor which turns the single into a pea, and a walnut is a single which possesses two additional modifying factors. Singleness, in fact, underlies all these combs, and if we write their zygotic constitution in full we must denote a walnut as RRPPSS, a rose as RRppSS, a pea as rrPPSS, and a single as rrppSS. The crossing of rose with pea results in a reshuffling of the factors concerned, and in accordance with the principle of segregation some zygotes are formed in which neither of the modifying factors R and P are present, and the single character can then become manifest. {39}

The Presence and Absence theory is to-day generally accepted by students of these matters. Not only does it afford a simple explanation of the remarkable fact that in all cases of Mendelian inheritance we should be able to express our unit-characters in terms of alternative pairs, but, as we shall have occasion to refer to later, it suggests a clue as to the course by which the various domesticated varieties of plants and animals have arisen from their wild prototypes.

Before leaving this topic we may draw attention to some experiments which offer a pretty confirmation of the view that the rose comb is a single to which a modifying factor for roseness has been added. It was argued that if we could find a type of comb in which the factor for singleness was absent, then on crossing such a comb with a rose we ought, if singleness really underlies rose, to obtain some single combs in F_2 from such a cross. Such a comb we had the good fortune to find in the Breda fowl, a breed largely used in Holland. This fowl is usually spoken of as combless, for the place of the comb is taken by a covering of short bristlelike feathers (Fig. 6, D). In reality it possesses the vestige of a comb in the form of two minute lateral knobs of comb tissue. Characteristic also of this breed is the high development of the horny nostrils, a feature probably correlated with the almost complete absence of comb. The first step in the experiment was to prove the absence of the factor for singleness in the Breda. {40} On crossing Breda with single the F_1 birds exhibit a large comb of the form of a double single comb in which the two portions are united anteriorly, but diverge from one another towards the back of the head (Fig. 6, C). The Breda contains an element of duplicity which is dominant to the simplicity of the ordinary single comb. But it cannot contain the factor for the single comb, because as soon as that is put into it by crossing with a single the comb {41} assumes a large size, and is totally distinct in appearance from its almost complete absence in the pure Breda. Now when the Breda is crossed with the rose duplicity is dominant to simplicity, and rose is dominant to lack of comb, and the F_1 generation consists of birds possessing duplex rose combs (Fig. 6, A and B). On breeding such birds together we obtain a generation consisting of Bredas, duplex roses, roses, duplex singles, and singles. From our previous experiment we know that the singles could not have come from the Breda, since a Breda comb to which the factor for single has been added no longer remains a Breda. Therefore it must have come from the rose, thus confirming our view that the rose is in reality a single comb which contains in addition a dominant modifying factor (R) whose presence turns it into a rose. We shall take it, therefore, that there is good experimental evidence for the Presence and Absence theory, and we shall express in terms of it the various cases which come up for discussion in succeeding chapters.

Rose x Breda - -+ Duplex x Duplex Rose Rose + - - - Duplex Rose Duplex Single Breda Rose Single (Duplex and Simplex)

* * * * *




We have now reached a point at which it is possible to formulate a definite conception of the living organism. A plant or animal is a living entity whose properties may in large measure be expressed in terms of unit-characters, and it is the possession of a greater or lesser number of such unit-characters renders it possible for us to draw sharp distinctions between one individual and another. These unit-characters are represented by definite factors in the gamete which in the process of heredity behave as indivisible entities, and are distributed according to a definite scheme. The factor for this or that unit-character is either present in the gamete or it is not present. It must be there in its entirety or completely absent. Such at any rate is the view to which recent experiment has led us. But as to the nature of these factors, the conditions under which they exist in the gamete, and the manner in which they produce their specific effects in the zygote, we are at present almost completely in the dark.

The case of the fowls' combs opens up the important question of the extent to which the various factors can {43} influence one another in the zygote. The rose and the pea factors are separate entities, and each when present alone produces a perfectly distinct and characteristic effect upon the single comb, turning it into a rose or a pea as the case may be. But when both are present in the same zygote their combined effect is to produce the walnut comb, a comb which is quite distinct from either and in no sense intermediate between them. The question of the influence of factors upon one another did not present itself to Mendel because he worked with characters which affected different parts of the plant. It was unlikely that the factor which led to the production of colour in the flower would affect the shape of the pod, or that the height of the plant would be influenced by the presence or absence of the factor that determined the shape of the ripe seed. But when several factors can modify the same structure it is reasonable to suppose that they will influence one another in the effects which their simultaneous presence has upon the zygote. By themselves the pea and the rose factors each produce a definite modification of the single comb, but when both are present in the zygote, whether as a single or double dose, the modification that results is quite different to that produced by either when present alone. Thus we are led to the conception of characters which depend for their manifestation on more than one factor in the zygote, and in the present chapter we may consider a few of the {44} phenomena which result from such interaction between separate and distinct factors.

White x White Red F1 Red White - F2 (9) (7)

One of the most interesting and instructive cases in which the interaction between separate factors has been demonstrated is a case in the sweet pea. All white sweet peas breed true to whiteness. And generally speaking the result of crossing different whites is to produce nothing but whites, whether in F_1 or in succeeding generations. But there are certain strains of white sweet peas which when crossed together produce only coloured flowers. The colour may be different in different cases, though for our present purpose we may take a case in which the colour is red. When such reds are allowed to self-fertilise themselves in the normal way and the seeds sown, the resulting F_2 generation consists of reds and whites, the former being rather more numerous than the latter in the proportion of 9 : 7. The raising of a further generation from the seeds of these F_2 plants shows that the whites always breed true to whiteness, but that different reds may behave differently. Some breed true, others give reds and whites in the ratio 3 : 1, while others, again, give reds and whites in the ratio 9 : 7. As in the case of the fowls' combs, this case may be interpreted in terms of the presence and absence of two factors. {45}

White White AAbb aaBB / / / / Ab Ab aB aB gametes of parents '———-' Red F_1 AaBb / / AB AB Female gametes Ab Ab Male gametes of F_1 aB aB of F_1 ab ab

Red in the sweet pea results from the interaction of two factors, and unless these are both present the red colour cannot appear. Each of the white parents carried one of the two factors whose interaction is necessary for the production of the red colour, and as a cross between them brings these two complementary factors together the F_1 plants must all be red. As this case is of considerable importance for the proper understanding of much that is to follow, and as it has been completely worked out, we shall consider it in some detail. Denoting these two colour factors by A and B respectively we may proceed to follow out the consequences of this cross. Since all the F_1 plants were red the constitution of the parental whites must have been AAbb and aaBB respectively, and their gametes consequently Ab and aB. The constitution of the F_1 plants must, therefore, be AaBb. Such a plant being heterozygous for two factors produces a series of gametes of the four kinds AB, Ab, aB, ab, and produces them in equal numbers (cf. p. 36). To obtain the various types of zygotes which are produced when such {46} a series of pollen grains meets a similar series of ovules we may make use of the same "chessboard" system which we have already adopted in the case of the fowls' combs.

+ + + + + AB.... AB.... AB.... AB.... AB.... Ab.... aB.... ab.... ...... ...... ...... ...... + + + + + Ab.... Ab Ab.... Ab AB.... Ab aB.... ab ...... ...... + + + + + aB.... aB.... aB aB AB.... Ab.... aB ab ...... ...... + + + + + ab.... ab ab ab AB.... Ab aB ab ...... + + + + +

FIG. 7.

Diagram to illustrate the nature of the F2 generation from the two white sweet peas which give a coloured F1.

An examination of this figure (Fig. 7) shows that 9 out of the 16 squares contain both A and B, while 7 contain either A or B alone, or neither. In other words, on this view of the nature of the two white sweet peas we should in the F_2 generation look for the appearance of coloured and white flowers in the ratio 9 : 7. And this, as we have already seen, is what was actually found by experiment. Further examination of the figure shows that the coloured plants are not all of the same constitution, but are of four kinds with respect to their zygotic constitution, viz. AABB, AABb, AaBB, and AaBb. Since AABB is homozygous for both A and B, all the gametes which it produces must contain both of these factors, and such a plant must therefore breed true to the red colour. A plant of the {47} constitution AABb is homozygous for the factor A, but heterozygous for B. All of its gametes will contain A, but only one-half of them will contain B, _i.e._ it produces equal numbers of gametes AB and Ab. Two such series of gametes coming together must give a generation consisting of x AABB, 2x AABb, and x AAbb, that is, reds and whites in the ratio 3 : 1. Lastly the red zygotes of the constitution AaBb have the same constitution as the original red made from the two whites, and must therefore when bred from give reds and whites in the ratio 9 : 7. The existence of all these three sorts of reds was demonstrated by experiment, and the proportions in which they were met with tallied with the theoretical explanation.

The theory was further tested by an examination into the properties of the various F2 whites which come from a coloured plant that has itself been produced by the mating of two whites. As Fig. 7 shows, these are, in respect of their constitution, of five different kinds, viz. AAbb, Aabb, aaBB, aaBb, and aabb. Since none of them produce anything but whites on self-fertilisation it was found necessary to test their properties in another way, and the method adopted was that of crossing them together. It is obvious that when this is done we should expect different results in different cases. Thus the cross between two whites of the constitution AAbb and aaBB should give nothing but coloured plants; for these two whites are of {48} the same constitution as the original two whites from which the experiment started. On the other hand, the cross between a white of the constitution aabb and any other white can never give anything but whites. For no white contains both A and B, or it would not be white, and a plant of the constitution aabb cannot supply the complementary factor necessary for the production of colour. Again, two whites of the constitution Aabb and aaBb when crossed should give both coloured and white flowers, the latter being three times as numerous as the former. Without going into further detail it may be stated that the results of a long series of crosses between the various F2 whites accorded closely with the theoretical explanation.

From the evidence afforded by this exhaustive set of experiments it is impossible to resist the deduction that the appearance of colour in the sweet pea depends upon the interaction of two factors which are independently transmitted according to the ordinary scheme of Mendelian inheritance. What these factors are is still an open question. Recent evidence of a chemical nature indicates that colour in a flower is due to the interaction of two definitive substances: (1) a colourless "chromogen," or colour basis; and (2) a ferment which behaves as an activator of the chromogen, and by inducing some process of oxidation, leads to the formation of a coloured substance. But whether these two bodies exist as such {49} in the gametes or whether in some other form we have as yet no means of deciding.

Since the elucidation of the nature of colour in the sweet pea phenomena of a similar kind have been witnessed in other plants, notably in stocks, snapdragons, and orchids. Nor is this class of phenomena confined to plants. In the course of a series of experiments upon the plumage colour of poultry, indications were obtained that different white breeds did not always owe their whiteness to the same cause. Crosses were accordingly made between the white Silky fowl and a pure white strain derived from the white Dorking. Each of these had been previously shown to behave as a simple recessive to colour. When the two were crossed only fully coloured birds resulted. From analogy with the case of the sweet pea it was anticipated that such F1 coloured birds when bred together would produce an F2 generation consisting of coloured and white birds in the ratio 9 : 7, and when the experiment was made this was actually shown to be the case. With the growth of knowledge it is probable that further striking parallels of this nature between the plant and animal worlds will be met with.

Before quitting the subject of these experiments attention may be drawn to the fact that the 9 : 7 ratio is in reality a 9 : 3 : 3 : 1 ratio in which the last three terms are indistinguishable owing to the special circumstances that neither factor can produce a visible effect without {50} the co-operation of the other. And we may further emphasise the fact that although the two factors thus interact upon one another they are nevertheless transmitted quite independently and in accordance with the ordinary Mendelian scheme.

Agouti x Agouti + -+ Agouti x Agouti + -+ -+ Agouti Black Albino (9) (3) (4)

One of the earliest sets of experiments demonstrating the interaction of separate factors was that made by the French zoologist Cuenot on the coat colours of mice. It was shown that in certain cases agouti, which is the colour of the ordinary wild grey mouse, behaves as a dominant to the albino variety, i.e. the F2 generation from such a cross consists of agoutis and albinos in the ratio 3 : 1. But in other cases the cross between albino and agouti gave a different result. In the F1 generation appeared only agoutis as before, but the F2 generation consisted of three distinct types, viz. agoutis, albinos, and blacks. Whence the sudden appearance of the new type? The answer is a simple one. The albino parent was really a black. But it lacked the factor without which the colour is unable to develop, and consequently it remained an albino. If we denote this factor by C, then the constitution of an albino must be cc, while that of a coloured animal may be CC or Cc, according as to whether it breeds true to colour or can {51} throw albinos. Agouti was previously known to be a simple dominant to black, i.e. an agouti is a black rabbit plus an additional greying factor which modifies the black into agouti. This factor we will denote by G, and we will use B for the black factor. Our original agouti and albino parents we may therefore regard as in constitution GGCCBB and ggccBB respectively. Both of the parents are homozygous for black. The gametes produced by the two parents are GCB, and gcB, and the constitution of the F1 animals must be GgCcBB. Being heterozygous for two factors they will produce four kinds of gametes in equal numbers, viz. GCB, GcB, gCB, and gcB. The results of the mating of two such similar series of gametes when the F1 animals are bred together we may determine by the usual "chessboard" method (Fig. 8). Out of the 16 squares 9 contain both C and G in addition to B. Such animals must be agoutis. Three squares contain C but not G. Such animals must be coloured, but as they do not contain the modifying agouti factor their colour will be black. The remaining four squares do not contain C, and in the absence of this colour-developing factor they must all be albinos. Theory demands that the three classes agouti, black, and albino should appear in F2 in the ratio 9 : 3 : 4; experiment has shown that these are the only classes that appear, and that the proportions in which they are produced accord closely with the theoretical expectation. Put briefly, then, the explanation {52} of this case is that all the animals are black, and that we are dealing with the presence and absence of two factors, a colour developer (C), and a colour modifier (G), both acting, as it were, upon a substratum of black. The F2 generation really consists of the four classes agoutis, blacks, albino agoutis, and albino blacks in the ratio 9 : 3 : 3 : 1. But since in the absence of the colour developer C the colour modifier G can produce no visible result, the last two classes of the ratio are indistinguishable, and our F2 generation comes to consist of three classes in the ratio 9 : 3 : 4, instead of four classes in the ratio 9 : 3 : 3 : 1.

+ -+ -+ -+ -+ GCB.... GCB.... GCB.... GCB.... GCB.... GcB.... gCB.... gcB.... ....... ....... ....... ....... .Agouti .Agouti .Agouti .Agouti + -+ -+ -+ -+ GcB.... GcB GcB.... GcB GCB.... GcB gCB.... gcB ....... ....... .Agouti Albino .Agouti Albino + -+ -+ -+ -+ gCB.... gCB.... gCB# gCB# GCB.... GcB.... gCB# gcB# ....... ....... ## ## .Agouti .Agouti #BLACK #BLACK + -+ -+ -+ -+ gcB.... gcB gcB# gcB GCB.... GcB gCB# gcB ....... ## .Agouti Albino #BLACK Albino + -+ -+ -+ -+

FIG. 8.

Diagram to illustrate the nature of the F_2 generation which may arise from the mating of agouti with albino in mice or rabbits.

This explanation was further tested by experiments with the albinos. In an F2 family of this nature there ought to be three kinds, viz. albinos homozygous for G (GGccBB), albinos heterozygous for G (GgccBB), and albinos without G (ggccBB). These albinos are, as it were, like photographic plates exposed but undeveloped. {53} Their potentialities may be quite different, although they all look alike, but this can only be tested by treating them with a colour developer. In the case of the mice and rabbits the potentiality for which we wish to test is the presence or absence of the factor G, and in order to develop the colour we must introduce the factor C. Our developer, therefore, must contain C but not G. In other words, it must be a homozygous black mouse or rabbit, ggCCBB. Since such an animal is pure for C it must, when mated with any of the albinos, produce only coloured offspring. And since it does not contain G the appearance of agoutis among its offspring must be attributed to the presence of G in the albino. Tested in this way the F2 albinos were proved, as was expected, to be of three kinds: (1) those which gave only agouti, i.e. which were homozygous for G; (2) those which gave agoutis and blacks in approximately equal numbers, i.e. which were heterozygous for G; and (3) those which gave only blacks, and therefore did not contain G.

Though albinos, whether mice, rabbits, rats, or other animals, breed true to albinism, and though albinism behaves as a simple recessive to colour, yet albinos may be of many different sorts. There are in fact just as many kinds of albinos as there are coloured forms—neither more nor less. And all these different kinds of albinos may breed together, transmitting the various colour factors according to the Mendelian scheme of inheritance, {54} and yet the visible result will be nothing but albinos. Under the mask of albinism is all the while occurring that segregation of the different colour factors which would result in all the varieties of coloured forms, if only the essential factor for colour development were present. But put in the developer by crossing with a pure coloured form and their variety of constitution can then at last become manifest.

So far we have dealt with cases in which the production of a character is dependent upon the interaction of two factors. But it may be that some characters require the simultaneous presence of a greater number of factors for their manifestation, and the experiments of Miss Saunders have shown that there is a character in stocks which is unable to appear except through the interaction of three distinct factors. Coloured stocks may be either hoary, with the leaves and stem covered by small hairs, or they may lack the hairy covering, in which case they are termed glabrous. Hoariness is dominant to glabrousness; that is to say, there is a definite factor which can turn the glabrous into a hoary plant when it is present. But in families where coloured and white stocks occur the white are always glabrous, while the coloured plants may or may not be hoary. Now colour in the stock as in the sweet pea has been proved to be dependent upon the interaction of two separate factors. Hence hoariness depends upon three separate factors, and a stock cannot be hoary unless {55} it contains the hoary factor in addition to the two colour factors. It requires the presence of all these three factors to produce the hoary character, though how this comes about we have not at present the least idea.

A somewhat different and less usual form of interaction between factors may be illustrated by a case in primulas recently worked out by Bateson and Gregory. Like the common primrose, the primula exhibits both pin-eyed and thrum-eyed varieties. In the former the style is long, and the centre of the eye is formed by the end of the stigma which more or less plugs up the opening of the corolla (cf. Fig. 9, A); in the latter the style is short and hidden by the four anthers which spring from higher up in the corolla and form the centre of the eye (cf. Fig. 9, B). The greater part of the "eye" is formed by the greenish-yellow patches on each petal just at the opening {56} of the corolla. In most primulas the eye is small, but there are some in which it is large and extends as a flush over a considerable part of the petals (Fig. 10). Experiments showed that these two pairs of characters behave in simple Mendelian fashion, short style ( = "thrum") being dominant to long style (= "pin") and small eye dominant to large. Besides the normal long and short styled forms, there occurs a third form, which has been termed homostyle. In this form the anthers are placed low down in the corolla tube as they are in the long-styled form, but the style remains short instead of reaching up to the corolla opening (Fig. 9, C). In the course of their experiments Bateson and Gregory crossed a large-eyed homostyle plant with a small-eyed thrum ( = short style). The F1 plants were all short styled with small eyes. {57} On self-fertilisation these gave an F2 generation consisting of four types, viz. short styled with small eyes, short styled with large eyes, long styled with small eyes, and homostyled with large eyes. The notable feature of this generation is the appearance of long-styled plants, which, however, occur only in association with the small eye. The proportions in which these four types appeared shows that the presence or absence of but two factors is concerned, and at the same time provides the key to the nature of the homostyled plants. These are potentially long styled, and the position of the anthers is that of normal long-styled plants, but owing to some interaction between the factors the style itself is unable to reach its full development unless the factor for the small eye is present. For this reason long-styled plants with the large eye are always of the homostyle form. What the connecting-link between these apparently unrelated structures may be we cannot yet picture to ourselves, any more than we can picture the relation between flower {58} colour and hairiness in stocks. It is evident, however, that the conception of the interaction of factors, besides clearing up much that is paradoxical in heredity, promises to indicate lines of research which may lead to valuable extensions in our knowledge of the way in which the various parts of the living organism are related to one another.

Short style } { Homo style small eye } x { large eye Short style small eye - - Short style Short style Long style Homo style small eye large eye ("pin") large eye (9) (3) (3) (1)

* * * * *




As soon as the idea was grasped that characters in plants and animals might be due to the interaction of complementary factors, it became evident that this threw clear light upon the hitherto puzzling phenomenon of reversion. We have already seen that in certain cases the cross between a black mouse or rabbit and an albino, each belonging to true breeding strains, might produce nothing but agoutis. In other words, the cross between the black and the white in certain instances results in a complete reversion to the wild grey form. Expressed in Mendelian terms, the production of the agouti was the necessary consequence of the meeting of the factors C and G in the same zygote. As soon as they are brought together, no matter in what way, the reversion is bound to occur. Reversion, therefore, in such cases we may regard as the bringing together of complementary factors which had somehow in the course of evolution become separated from one another. In the simplest cases, such as that of the black and the white rabbit, only two factors are concerned, and one of them is brought in from each of the {60} two parents. But in other cases the nature of the reversion may be more complicated owing to a larger number of factors being concerned, though the general principle remains the same. Careful breeding from the reversions will enable us in each case to determine the number and nature of the factors concerned, and in illustration of this we may take another example from rabbits. The Himalayan rabbit is a well-known breed. In appearance it is a white rabbit with pink eyes, but the ears, paws, and nose are black (Pl. I., 2). The Dutch rabbit is another well-known breed. Generally speaking, the anterior portion of the body is white, and the posterior part coloured. Anteriorly, however, the eyes are surrounded by coloured patches extending up to the ears, which are entirely coloured. At the same time the hind paws are white (cf. Pl. I., 1). Dutch rabbits exist in many varieties of colour, though in each one of these the distribution of colour and white shows the same relations. In the experiments about to be described a yellow Dutch rabbit was crossed with a Himalaya. The result was a reversion to the wild agouti colour (Pl. I., 3). Some of the F_1 individuals showed white patches, while others were self-coloured. On breeding from the F_1 animals a series of coloured forms resulted in F_2. These were agoutis, blacks, yellows, and sooty yellows, the so-called tortoise shells of the fancy (Pl. I., 4-7).


Yellow x Himalayan + -+ Agouti x Agouti + + + -+ + Agouti Yellow Black Tortoise Himalayan Shell (27) (9) (9) (3) (16)

In addition to these appeared Himalayans with either black points or with lighter brownish ones, and the proportions in which they came showed the Himalayan character to be a simple recessive. A certain number of the coloured forms exhibited the Dutch marking to a greater or less extent, but as its inheritance in this set of experiments is complicated and has not yet been worked out, we may for the present neglect it and confine our attention to the coloured types and to the Himalayans. The proportion in which the four coloured types appeared in F_2 was very nearly 9 agoutis, 3 blacks, 3 yellows, and 1 tortoiseshell. Evidently we are here dealing with two factors: (1) the grey factor (G), which modifies black into agouti, or tortoiseshell into yellow; and (2) an intensifying factor (I), which intensifies yellow into agouti and tortoiseshell into black. It may be mentioned here that other experiments confirmed the view that the yellow rabbit is a dilute agouti, and the tortoiseshell a dilute black. The Himalayan pattern behaves as a recessive to self-colour. It is a self-coloured black rabbit lacking a factor that allows the colour to develop except in the points. That factor we may denote {62} by X, and as far as it is concerned the Himalayan is constitutionally xx. The Himalayan contains the intensifying factor, for such pigment as it possesses in the points is full coloured. At the same time it is black, _i.e._ lacking in the factor G. With regard to these three factors, therefore, the constitution of the Himalayan is ggIIxx. The last character which we have to consider in this cross is the Dutch character. This was found by Hurst to behave as a recessive to self-colour (S), and for our present purpose we will regard it as differing from a self-coloured rabbit in the lack of this factor.[3] The Himalayan is really a self-coloured animal, which, however, is unable to show itself as a full black owing to its not possessing the factor X. The results of breeding experiments then suggest that we may denote the Himalayan by the formula ggIIxxSS and the yellow Dutch by GGiiXXss. Each lacks two of the factors upon the full complement of which the agouti colour depends. By crossing them the complete series GIXS is brought into the same zygote, and the result is a reversion to the colour of the wild rabbit.

Bush x Cupid Tall F1 + + -+ + + Tall Bush Cupid Cupid F2 (procumbent) (erect)

Most of the instances of reversion yet worked out are those in which colour characters are concerned. The sweet pea, however, supplies us with a good example of reversion in structural characters. A dwarf variety known as the "Cupid" has been extensively grown for {63} some years. In these little plants the internodes are very short and the stems are few in number, and attain to a length of only 9-10 inches. In course of growth they diverge from one another, and come to lie prostrate on the ground (Pl. II., 2). Curiously enough, although the whole plant is dwarfed in other respects, this does not seem to affect the size of the flower, which is that of a normal sweet pea. Another though less-known variety is the "Bush" sweet pea. Its name is derived from its habit of growth. The numerous stems do not diverge from one another, but all grow up side by side, giving the plant the appearance of a compact bush (Pl. II., 1). Under ordinary conditions it attains a height of 3-1/2-4 feet. A number of crosses were made between the Bush and Cupid varieties, with the somewhat unexpected result that in every instance the F1 plants showed complete reversion to the size and habit of the ordinary tall sweet pea (Pl. II., 3), which is the form of the wild plant as it occurs in Sicily to-day. The F2 generation from these reversionary talls consisted of four different types, viz. {64} talls, bushes, Cupids of the procumbent type like the original Cupid parent, and Cupids with the compact upright Bush habit (Pl. II., 4). These four types appeared in the ratio 9 : 3 : 3 : 1, and this, of course, provided the clue to the nature of the case. The characters concerned are (1) long internode of stem between the leaves which is dominant to short internode, and (2) the creeping procumbent habit which is dominant to the erect bush-like habit. Of these characters length of internode was carried by the Bush, and the procumbent habit by the original Cupid parent. The bringing of them together by the cross resulted in a procumbent plant with long internodes. This is the ordinary tall sweet pea of the wild Sicilian type, reversion here, again, being due to the bringing together of two complementary factors which had somehow become separated in the course of evolution.

To this interpretation it may be objected that the ordinary sweet pea is a plant of upright habit. This, however, is not true. It only appears so because the conventional way of growing it is to train it up sticks. In reality it is of procumbent habit, with divergent stems like the ordinary Cupid, a fact which can easily be observed by anyone who will watch them grow without the artificial aid of prepared supports.


The cases of reversion with which we have so far dealt have been cases in which the reversion occurs as an immediate result of a cross, _i.e._ in the F_1 generation. This is perhaps the commonest mode of reversion, but instances are known in which the reversion that occurs when two pure types are crossed does not appear until the F_2 generation. Such a case we have already met with in the fowls' combs. It will be remembered that the cross between pure pea and pure rose gave walnut combs in F_1, while in the F_2 generation a definite proportion, 1 in 16, of single combs appeared (cf. p. 32). Now the single comb is the form that is found in the wild jungle fowl, which is generally regarded as the ancestor of the domestic breeds. If this is so, we have a case of reversion in F_2; and this in the _absence_ of the two factors brought together by the rose-comb and pea-comb parents. Instead of the reversion being due to the bringing together of two complementary factors, we must regard it here as due to the association of two complementary absences. To this question, however, we shall revert later in discussing the origin of domesticated varieties.

Black Barb x White Fantail Black Barb x Spot[4] Dark x Dark Among the offspring one very similar to the wild blue rock.

Black White Barb x Fantail + + Black x Black (White Splashed) (White Splashed) + + + -+ -+ Black Black Blue Blue White (White Splashed) (White Splashed) / -/ (9) (3) (4)

There is one other instance of reversion to which we must allude. This is Darwin's famous case of the occasional appearance of pigeons reverting to the wild blue rock (_Columba livia_), when certain domesticated races are crossed together. As is well known, Darwin made use of this as an argument for regarding all the domesticated varieties as having arisen from the same wild species. The original experiment is somewhat complicated, and is shown in the accompanying scheme. Essentially it lay in {66} following the results flowing from crosses between blacks and whites. Experiments recently made by Staples-Browne have shown that this case of reversion also can be readily interpreted in Mendelian terms. In these experiments the cross was made between black barbs and white fantails. The F_1 birds were all black with some white splashes, evidently due to a separate factor introduced by the fantail. On breeding these blacks together they gave an F_2 generation, consisting of blacks (with or without white splashes), blues (with or without white splashes), and whites in the ratio 9 : 3 : 4. The factors concerned are colour (C), in the absence of {67} which a bird is white, and a black modifier (B), in the absence of which a coloured bird is blue. The original black barb contained both of these factors, being in constitution CCBB. The fantail, however, contained neither, and was constitutionally ccbb. The F_1 birds produced by crossing were in constitution CcBb, and being heterozygous for two factors produced in equal numbers the four sorts of gametes CB, Cb, cB, cb. The results of two such series of gametes being brought together are shown in the usual way in Fig. 11. A blue is a bird containing the colour factor but lacking the black modifier, _i.e._ of the constitution CCbb, or Ccbb, and such birds as the figure shows appear in the F_2 generation on the average three times out of sixteen. Reversion here comes about in F_2, when the redistribution of the factors leads to the formation of zygotes containing one of the two factors but not the other.

+ -+ -+ -+ -+ CB## CB## CB## CB## CB## Cb## cB## cb## ## ## ## ## #BLACK #BLACK #BLACK #BLACK + -+ -+ -+ -+ Cb## Cb..... Cb## Cb..... CB## Cb..... cB## cb..... ## ....... ## ....... #BLACK ...Blue #BLACK ...Blue + -+ -+ -+ -+ cB## cB## cB cB CB## Cb## cB cb ## ## #BLACK #BLACK + -+ -+ -+ -+ cb## cb..... cb cb CB## Cb..... cB cb ## ....... #BLACK ...Blue + -+ -+ -+ -+

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