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Mendelism - Third Edition
by Reginald Crundall Punnett
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One of the species of Euralia occurs in two very distinct forms (Pl. VI.), which were previously regarded as separate species under the names E. wahlbergi and E. mima. These two forms respectively resemble Amauris dominicanus and A. echeria. For purposes of argument we will assume A. echeria to be the more recent form of the two. On the modern Darwinian view certain individuals of A. dominicanus gradually diverged from the dominicanus type and eventually reached the echeria type, though why this should have happened does not appear to be clear. At the same time those specimens which tended to vary in the direction of A. echeria in places where this species was more abundant than A. dominicanus were encouraged by natural selection, and under its guiding hand the form mima eventually arose from wahlbergi.

According to Mendelian views, on the other hand, {146} A. echeria arose suddenly from A. dominicanus (or vice versa), and similarly mima arose suddenly from wahlbergi. If mima occurred where A. echeria was common and A. dominicanus was rare, its resemblance to the more plentiful distasteful form would give it the advantage over wahlbergi and allow it to establish itself in place of the latter. On the modern Darwinian view natural selection gradually shapes wahlbergi into the mima form owing to the presence of A. echeria; on the Mendelian view natural selection merely conserves the mima form when once it has arisen. Now this case of mimicry is one of especial interest, because we have experimental evidence that the relation between mima and wahlbergi is a simple Mendelian one, though at present it is uncertain which is the dominant and which the recessive form. The two have been proved to occur in families bred from the same female without the occurrence of any intermediates, and the fact that the two segregate cleanly is strong evidence in favour of the Mendelian view. On this view the genera Amauris and Euralia contain a similar set of pattern factors, and the conditions, whatever they may be, which bring about mutation in the former lead to the production of a similar mutation in the latter. Of the different forms of Euralia produced in any region that one has the best chance of survival, through the operation of natural selection, which resembles the most plentiful Amauris form. Mimetic resemblance is a true phenomenon, but natural selection plays the part of a conservative, not of a formative agent.



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It is interesting to recall that in earlier years Darwin was inclined to ascribe more importance to "sports" as opposed to continuous minute variation, and to consider that they might play a not inconsiderable part in the formation of new varieties in nature. This view, however, he gave up later, because he thought that the relatively rare sport or mutation would rapidly disappear through the swamping effects of crossing with the more abundant normal form, and so, even though favoured by natural selection, would never succeed in establishing itself. Mendel's discovery has eliminated this difficulty. For suppose that the sport differed from the normal in the loss of a factor and were recessive. When mated with the normal this character would seem to disappear, though, of course, half of the gametes of its progeny would bear it. By continual crossing with normals a small proportion of heterozygotes would eventually be scattered among the population, and as soon as any two of these mated together the recessive sport would appear in one quarter of their offspring.

A suggestive contribution to this subject was recently made by G. H. Hardy. Considering the distribution of a single factor in a mixed population consisting of the heterozygous and the two homozygous forms he showed that such a population breeding at random rapidly fell into a {148} stable condition with regard to the proportion of these three forms, whatever may have been the proportion of the three forms to start with. Let us suppose for instance, that the population consists of p homozygotes of one kind, r homozygotes of the other kind, and 2 q heterozygotes. Hardy pointed out that, other things being equal, such a population would be in equilibrium for this particular factor so long as the condition q^2 = pr was fulfilled. If the condition is fulfilled to start with, the population remains in equilibrium. If the condition is not fulfilled to start with, Hardy showed that a position of equilibrium becomes established after a single generation, and that this position is thereafter maintained. The proportions of the three classes which satisfy the equation q^2 = pr are exceedingly numerous, and populations in which they existed in the proportions shown in the appended table would remain in stable equilibrium generation after generation:—

p. 2q. r. 1 2 1 1 4 4 1 6 9 1 8 16 1 20,000 100,000,000 1 2n n^2

This, of course, assumes that all three classes are equally fertile, and that no form of selection is taking place to the {149} benefit of one class more than of another. Moreover, it makes no difference whether p represents the homozygous dominants or whether it stands for the recessives. A population containing a very small proportion of dominants and one containing a similar proportion of recessives are equally stable. The term dominant is in some respects apt to be misleading, for a dominant character cannot in virtue of its dominance establish itself at the expense of a recessive one. Brown eyes in man are dominant to blue, but there is no reason to suppose that as years go on the population of these islands will become increasingly brown eyed. Given equality of conditions both are on an equal footing. If, however, either dominant or recessive be favoured by selection the conditions are altered, and it can be shown that even a small advantage possessed by the one will rapidly lead to the elimination of the other. Even with but a 5 per cent selection advantage in its favour it can be shown that a rare sport will oust the normal form in a few hundred generations. In this way we are freed from a difficulty inherent in the older view that varieties arose through a long-continued process involving the accumulation of very slight variations. On that view the establishing of a new type was of necessity a very long and tedious business, involving many thousands of generations. For this reason the biologist has been accustomed to demand a very large supply of time, often a great deal more than the physicist is {150} disposed to grant, and this has sometimes led him to expostulate with the latter for cutting off the supply. On the newer views, however, this difficulty need not arise, for we realise that the origin and establishing of a new form may be a very much more rapid process than has hitherto been deemed possible.

One last question with regard to evolution. How far does Mendelism help us in connection with the problem of the origin of species? Among the plants and animals with which we have dealt we have been able to show that distinct differences, often considerable, in colour, size, and structure, may be interpreted in terms of Mendelian factors. It is not unlikely that most of the various characters which the systematist uses to mark off one species from another, the so-called specific characters, are of this nature. They serve as convenient labels, but are not essential to the conception of species. A systematist who defined the wild sweet pea could hardly fail to include in his definition such characters as the procumbent habit, the tendrils, the form of the pollen, the shape of the flower, and its purple colour. Yet all these and other characters have been proved to depend upon the presence of definite factors which can be removed by appropriate crossing. By this means we can produce a small plant a few inches in height with an erect habit of growth, without tendrils, with round instead of oblong pollen, and with colourless deformed flowers quite different {151} in appearance from those of the wild form. Such a plant would breed perfectly true, and a botanist to whom it was presented, if ignorant of its origin, might easily relegate it to a different genus. Nevertheless, though so widely divergent in structure, such a plant must yet be regarded as belonging to the species Lathyrus odoratus. For it still remains fertile with the many different varieties of sweet pea. It is not visible attributes that constitute the essential difference between one species and another. The essential difference, whatever it may be, is that underlying the phenomenon of sterility. The visible attributes are those made use of by the systematist in cataloguing the different forms of animal and plant life, for he has no other choice. But it must not be forgotten that they are often misleading. Until they were bred together Euralia wahlbergi and E. mima were regarded as perfectly valid species, and there is little doubt that numbers of recognised species will eventually fall to the ground in the same way as soon as we are in a position to apply the test of breeding. Mendelism has helped us to realise that specific characters may be but incidental to a species—that the true criterion of what constitutes a species is sterility, and that particular form of sterility which prevents two healthy gametes on uniting from producing a zygote with normal powers of growth and reproduction. For there are forms of sterility which are purely mechanical. The pollen of Mirabilis jalapa cannot fertilise M. {152} longiflora, because the pollen tubes of the former are not long enough to penetrate down to the ovules of the latter. Hybrids can nevertheless be obtained from the reciprocal cross. Nor should we expect offspring from a St. Bernard and a toy terrier without recourse to artificial fertilisation. Or sterility may be due to pathological causes which prevent the gametes from meeting one another in a healthy state. But in most cases it is probable that the sterility is due to some other cause. It is not inconceivable that definite differences in chemical composition render the protoplasm of one species toxic to the gametes of the other, and if this is so it is not impossible that we may some day be able to express these differences in terms of Mendelian factors. The very nature of the case makes it one of extreme difficulty for experimental investigation. At any rate, we realise more clearly than before that the problem of species is not one that can be resolved by the study of morphology or of systematics. It is a problem in physiology.

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CHAPTER XIV

ECONOMICAL

Since heredity lies at the basis of the breeder's work, it is evident that any contribution to a more exact knowledge of this subject must prove of service to him, and there is no doubt that he will be able to profit by Mendelian knowledge in the conduct of his operations. Indeed, as we shall see later, these ideas have already led to striking results in the raising of new and more profitable varieties. In the first place, heredity is a question of individuals. Identity of appearance is no sure guide to reproductive qualities. Two individuals similarly bred and indistinguishable in outward form may nevertheless behave entirely differently when bred from. Take, for instance, the family of sweet peas shown on Plate IV. The F2 generation here consists of seven distinct types, three sorts of purples, three sorts of reds, and whites. Let us suppose that our object is to obtain a true breeding strain of the pale purple picotee form. Now from the proportions in which they come we know that the dilute colour is due to the absence of the factor which intensifies the colour. Consequently the picotee cannot throw the {154} two deeper shades of red or purple. But it may be heterozygous for the purpling factor when it will throw the dilute red (tinged white), or it may be heterozygous for either or both of the two colour factors (cf. p. 44), in which case it will throw whites. Of the picotees which come in such a family, therefore, some will give picotees, tinged whites, and whites, others will give picotees and tinged whites only, others will give picotees and whites only, while others, again, and these the least numerous, will give nothing but picotees. The new variety is already fixed in a certain definite proportion of the plants; in this particular instance in 1 out of every 27. All that remains to be done is to pick out these plants. Since all the picotees look alike, whatever their breeding capacity, the only way to do this is to save the seed from a number of such plants individually, and to raise a further generation. Some of them will be found to breed true. The variety is then established, and may at once be put on the market with full confidence that it will hereafter throw none of the other forms. The all-important thing is to save and sow the seed of separate individuals separately. However alike they look, the seed from different individuals must on no account be mixed. Provided that due care is taken in this respect no long and tedious process of selection is required for the fixation of any given variety. Every possible variety arising from a cross appears in the F2 generation if only a sufficient {155} number is raised, and of all these different varieties a certain proportion of each is already fixed. Heredity is a question of individuals, and the recognition of this will save the breeder much labour, and enable him to fix his varieties in the shortest possible time.

Such cases as these of the sweet pea throw a fresh light upon another of the breeder's conceptions, that of purity of type. Hitherto the criterion of a "pure-bred" thing, whether plant or animal, has been its pedigree, and the individual was regarded as more or less pure bred for a given quality according as it could show a longer or shorter list of ancestors possessing this quality. To-day we realise that this is not essential. The pure-bred picotee appears in our F2 family though its parent was a purple bicolor, and its remoter ancestors whites for generations. So also from the cross between pure strains of black and albino rabbits we may obtain in the F2 generation animals of the wild agouti colour which breed as true to type as the pure wild rabbit of irreproachable pedigree. The true test of the pure breeding thing lies not in its ancestry but in the nature of the gametes which have gone to its making. Whenever two similarly constituted gametes unite, whatever the nature of the parents from which they arose, the resulting individual is homozygous in all respects and must consequently breed true. In deciding questions of purity it is to the gamete, and not to ancestry, that our appeal must henceforth be made. {156}

Improvement is after all the keynote to the breeder's operations. He is aiming at the production of a strain which shall combine the greatest number of desirable properties with the least number of undesirable ones. This good quality he must take from one strain, that from another, and that again from a third, while at the same time avoiding all the poor qualities that these different strains possess. It is evident that the Mendelian conception of characters based upon definite factors which are transmitted on a definite scheme must prove of the greatest service to him. For once these factors have been determined, their distribution is brought under control, and they can be associated together or dissociated at the breeder's will. The chief labour involved is that necessary for the determination of the factors upon which the various characters depend. For it often happens that what appears to be a simple character turns out when analysed to depend upon the simultaneous presence of several distinct factors. Thus the Malay fowl breeds true to the walnut comb, as does also the Leghorn to the single comb, and when pure strains are crossed all the offspring have walnut combs. At first sight it would be not unnatural to regard the difference as dependent upon the presence or absence of a single factor. Yet, as we have already seen, two other types of comb, the pea and the rose, make their appearance in the F_2 generation. Analysis shows that the difference between the walnut {157} and the single is a difference of two factors, and it is not until this has been determined that we can proceed with certainty to transfer the walnut character to a single-combed breed. Moreover, in his process of analysis the breeder must be prepared to encounter the various phenomena that we have described under the headings of interaction of factors, coupling and repulsion, and the recognition of these phenomena will naturally influence his procedure. Or again, his experiments may show him that one of the characters he wants, like the blue of the Andalusian fowl, is dependent upon the heterozygous nature of the individual which exhibits it, and if such is the case he will be wise to refrain from any futile attempt at fixing it. If it is essential it must be built up again in each generation, and he will recognise that the most economical way of doing this is to cross the two pure strains so that all the offspring may possess the desired character. The labour of analysis is often an intricate and tedious business. But once done it is done once for all. As soon as the various factors are determined, upon which the various characters of the individual depend, as soon as the material to be made use of has been properly analysed, the production and fixation of the required combinations becomes a matter of simple detail.

An excellent example of the practical application of Mendelian principles is afforded by the experiments which Professor Biffen has recently carried out in Cambridge. {158} Taken as a whole English wheats compare favourably with foreign ones in respect of their cropping power. On the other hand, they have two serious defects. They are liable to suffer from the attacks of the fungus which causes rust, and they do not bake into a good loaf. This last property depends upon the amount of gluten present, and it is the greater proportion of this which gives to the "hard" foreign wheat its quality of causing the loaf to rise well when baked. For some time it was held that "hard" wheat with a high glutinous content could not be grown in the English climate, and undoubtedly most of the hard varieties imported for trial deteriorated greatly in a very short time. Professor Biffen managed to obtain a hard wheat which kept its qualities when grown in England. But in spite of the superior quality of its grain from the baker's point of view its cropping capacity was too low for it to be grown profitably in competition with English wheats. Like the latter, it was also subject to rust. Among the many varieties which Professor Biffen collected and grew for observation he managed to find one which was completely immune to the attacks of the rust fungus, though in other respects it had no desirable quality to recommend it. Now as the result of an elaborate series of investigations he was able to show that the qualities of heavy cropping capacity, "hardness" of grain, and immunity to rust can all be expressed in terms of Mendelian factors. Having once analysed his material {159} the rest was comparatively simple, and in a few years he has been able to build up a strain of wheat which combines the cropping capacity of the best English varieties with the hardness of the foreign kinds, and at the same time is completely immune to rust. This wheat has already been shown to keep its qualities unchanged for several years, and there is little doubt that when it comes to be grown in quantity it will exert an appreciable influence on wheat-growing in Great Britain.



It may be objected that it is often with small differences rather than with the larger and more striking ones that the breeder is mainly concerned. It does not matter much to him whether the colour of a pea flower is purple or pink or white. But it does matter whether the plant bears rather larger seeds than usual, or rather more of them. Even a small difference when multiplied by the {160} size of the crop will effect a considerable difference in the profit. It is the general experience of seedsmen and others that differences of this nature are often capable of being developed up to a certain point by a process of careful selection each generation. At first sight this appears to be something very like the gradual accumulation of minute variations through the continuous application of a selective process. Some recent experiments by Professor Johannsen of Copenhagen set the matter in a different light. One of his investigations deals with the inheritance of the weight of beans, but as an account of these experiments would involve us in the consideration of a large amount of detail we may take a simple imaginary case to illustrate the nature of the conclusions at which he arrived. If we weigh a number of seeds collected from a patch of plants such as Johannsen's beans we should find that they varied considerably in size. The majority would probably not diverge very greatly from the general average, and as we approached the high or low extreme we should find a constantly decreasing number of individuals with these weights. Let us suppose that the weight of our seed varied between 4 and 20 grains, that the greatest number of seeds were of the mean weight, viz. 12 grains, and that as we passed to either extreme at 4 and 20 the number became regularly less. The weight relation of such a collection of seeds can be expressed by the accompanying curve (Fig. 30). Now if we select for {161} sowing only that seed which weighs over 12 grains, we shall find that in the next generation the average weight of the seed is raised and the curve becomes somewhat shifted to the right as in the dotted line of Fig. 30. By continually selecting we can shift our curve a little more to the right, i.e. we can increase the average weight of the seeds until at last we come to a limit beyond which further selection has no effect. This phenomenon has been long known, and it was customary to regard these variations as of a continuous nature, i.e. as all chance fluctuations in a homogeneous mass, and the effect of selection was supposed to afford evidence that small continuous variations could be increased by this process. But Johannsen's results point to another interpretation. Instead of our material being homogeneous it is probably a mixture of several strains each with its own average weight about {162} which the varying conditions of the environment cause it to fluctuate. Each of these strains is termed a PURE LINE. If we imagine that there are three such pure lines in our imaginary case, with average weights 10, 12, 14 grains respectively, and if the range of fluctuation of each of these pure lines is 12 grains, then our curve must be represented as made up of the three components

A fluctuating between 4 and 16 with a mean of 10 B " " 6 " 18 " " 12 C " " 8 " 20 " " 14



as is shown in Fig. 31. A seed that weighs 12 grains may belong to any of these three strains. It may be an average seed of B, or a rather large seed of A, or a rather small seed of C. If it belongs to B its offspring will average 12 grains, if to A they will average 10 grains, and if to C they will average 14 grains. Seeds of similar weight may give a different result because they happen to be fluctuations of different pure lines. But within the pure line any seed, large or small, produces the average result for that line. Thus a seed of line C which weighs 20 grains will give practically the same result as one that weighs 10 grains.

On this view we can understand why selection of the largest seed raises the average weight in the next generation. We are picking out more of C and less of A and B, and as this process is repeated the proportion of C gradually increases and we get the appearance of selection {163} acting on a continuously varying homogeneous material and producing a permanent effect. This is because the interval between the average weight of the different pure lines is small compared with the environmental fluctuations. None the less it is there, and the secret of separating and fixing any of these pure lines is again to breed from the individual separately. As soon as the pure line is separated further selection becomes superfluous.

Since the publication of Darwin's famous work upon the effects of cross and self fertilisation, it has been generally accepted that the effect of a cross is commonly, though not always, to introduce fresh vigour into the offspring, though why this should be so we are quite at a loss to explain. Continued close inbreeding, on the contrary, eventually leads to deterioration, though, as in many self-fertilised plants, a considerable number of generations may elapse before it shows itself in any marked degree. The fine quality of many of the seedsman's choice varieties of vegetables probably depends upon the fact that they had resulted from a cross but a few generations back, and it is possible that they often oust the older kinds not because they started as something intrinsically better, but because the latter had gradually deteriorated through continuous self-fertilisation. Most breeders are fully alive to the beneficial results of a cross so far as vigour is concerned, but they often hesitate to embark upon it owing to what was held {164} to be the inevitably lengthy and laborious business of recovering the original variety and refixing it, even if in the process it was not altogether lost. That danger Mendelism has removed, and we now know that by working on these lines it is possible in three or four generations to recover the original variety in a fixed state with all the superadded vigour that follows from a cross.

Nor is the problem one that concerns self-fertilised plants only. Plants that are reproduced asexually often appear to deteriorate after a few generations unless a sexual generation is introduced. New varieties of potato, for example, are frequently put upon the market, and their excellent qualities give them a considerable vogue. Much is expected of them, but time after time they deteriorate in a disappointing way and are lost to sight. It is not improbable that we are here concerned with a case in which the plants lose their vigour after a few asexual generations of reproduction from tubers, and can only recover it with the stimulus that results from the interpolation of a sexual generation. Unfortunately this generally means that the variety is lost, for owing to the haphazard way in which new kinds of potatoes are reproduced it is probable that most cultivated varieties are complex heterozygotes. Were the potato plant subjected to careful analysis and the various factors determined upon which its variations depend, we should be in a position to remake continually any good potato without {165} running the risk of losing it altogether, as is now so often the case.

The application of Mendelian principles is likely to prove of more immediate service for plants than animals, for owing to the large numbers which can be rapidly raised from a single individual and the prevalence of self-fertilisation, the process of analysis is greatly simplified. Even apart from the circumstance that the two sexes may sometimes differ in their powers of transmission, the mere fact of their separation renders the analysis of their properties more difficult. And as the constitution of the individual is determined by the nature and quality of its offspring, it is not easy to obtain this knowledge where the offspring, as in most animals, are relatively few. Still, as has been abundantly shown, the same principles hold good here also, and there is no reason why the process of analysis, though more troublesome, should not be effectively carried out. At the same time, it affords the breeder a rational basis for some familiar but puzzling phenomena. The fact, for instance, that certain characters often "skip a generation" is simply the effect of dominance in F_1 and the reappearance of the recessive character in the following generation. "Reversion" and "atavism," again, are phenomena which are no longer mysterious, but can be simply expressed in Mendelian terms as we have already suggested in Chap. VI. The occasional appearance of a sport in a supposedly pure strain is {166} often due to the reappearance of a recessive character. Thus even in the most highly pedigreed strains of polled cattle such as the Aberdeen Angus, occasional individuals with horns appear. The polled character is dominant to the horned, and the occasional reappearance of the horned animal is due to the fact that some of the polled herd are heterozygous in this character. When two such individuals are mated, the chances are 1 in 4 that the offspring will be horned. Though the heterozygous individuals may be indistinguishable in appearance from the pure dominant, they can be readily separated by the breeding test. For when crossed by the recessive, in this case horned animals, the pure dominant gives only polled beasts, while the heterozygous individual gives equal numbers of polled and horned ones. In this particular instance it would probably be impracticable to test all the cows by crossing with a horned bull. For in each case it would be necessary to have several polled calves from each before they could with reasonable certainty be regarded as pure dominants. But to ensure that no horned calves should come, it is enough to use a bull which is pure for that character. This can easily be tested by crossing him with a dozen or so horned cows. If he gets no horned calves out of these he may be regarded as a pure dominant and thenceforward put to his own cows, whether horned or polled, with the certainty that all his calves will be polled. {167}

Or, again, suppose that a breeder has a chestnut mare and wishes to make certain of a bay foal from her. We know that bay is dominant to chestnut, and that if a homozygous bay stallion is used a bay foal must result. In his choice of a sire, therefore, the breeder must be guided by the previous record of the animal, and select one that has never given anything but bays when put to either bay or chestnut mares. In this way he will assure himself of a bay foal from his chestnut mare, whereas if the record of the sire shows that he has given chestnuts he will be heterozygous, and the chances of his getting a bay or a chestnut out of a chestnut mare are equal.

It is not impossible that the breeder may be unwilling to test his animals by crossing them with a different breed through fear that their purity may be thereby impaired, and that the influence of the previous cross may show itself in succeeding generations. He might hesitate, for instance, to test his polled cows by crossing them with a horned bull for fear of getting horned calves when the cows were afterwards put to a polled bull of their own breed. The belief in the power of a sire to influence subsequent generations, or telegony as it is sometimes called, is not uncommon even to-day. Nevertheless, carefully conducted experiments by more than one competent observer have failed to elicit a single shred of unequivocal evidence in favour of the view. Until we have evidence based upon experiments which are capable of {168} repetition, we may safely ignore telegony as a factor in heredity.

Heterozygous forms play a greater part in the breeding of animals than of plants, for many of the qualities sought after by the breeder are of this nature. Such is the blue of the Andalusian fowl, and, according to Professor Wilson, the roan of the Shorthorn is similar, being the heterozygous form produced by mating red with white. The characters of certain breeds of canaries and pigeons again appear to depend upon their heterozygous nature. Such forms cannot, of course, ever be bred true, and where several factors are concerned they may when bred together produce but a small proportion of offspring like themselves. As soon, however, as their constitution has been analysed and expressed in terms of Mendelian factors, pure strains can be built up which when crossed will give nothing but offspring of the desired heterozygous form.

The points with which the breeder is concerned are often fine ones, not very evident except to the practised eye. Between an ordinary Dutch rabbit and a winner, or between the comb of a Hamburgh that is fit to show and one that is not, the differences are not very apparent to the uninitiated. Whether Mendelism will assist the breeder in the production of these finer points is at present doubtful. It may be that these small differences are heritable, such as those that form the basis of Johannsen's pure lines. In this case the breeder's outlook is {169} hopeful. But it may be that the variations which he seeks to perpetuate are of the nature of fluctuations, dependent upon the earlier life conditions of the individual, and not upon the constitution of the gametes by which it was formed. If such is the case, he will get no help from the science of heredity, for we know of no evidence which might lead us to suppose that variations of this sort can ever become fixed and heritable.

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CHAPTER XV

MAN



Though the interest attaching to heredity in man is more widespread than in other animals, it is far more difficult to obtain evidence that is both complete and accurate. The species is one in which the differentiating characters separating individual from individual are very numerous, while the number of the offspring is comparatively few, and the generations are far between. For these reasons, even if it were possible, direct experimental work with man would be likely to prove both tedious and expensive. There is, however, another method besides the direct one from which something can be learned. This consists in collecting all the evidence possible, arranging it in the form of pedigrees, and comparing it with standard cases already worked out in animals and plants. In this way it has been possible to demonstrate in man the existence of several characters showing simple Mendelian inheritance. As few besides medical men have hitherto been concerned practically with heredity, such records as exist are, for the most part, records of deformity or of disease. So it happens that most of the {171} pedigrees at present available deal with characters which are usually classed as abnormal. In some of these the inheritance is clearly Mendelian. One of the cases which has been most fully worked out is that of a deformity known as brachydactyly. In brachydactylous people the {172} whole of the body is much stunted, and the fingers and toes appear to have two joints only instead of three (cf. Figs. 32 and 33). The inheritance of this peculiarity has been carefully investigated by Dr. Drinkwater, who collected all the data he was able to find among the members of a large family in which it occurred. The result is the pedigree shown on p. 173. It is assumed that all who are recorded as having offspring were married to normals. Examination of the pedigree brings out the facts (1) that all affected individuals have an affected parent; (2) that none of the unaffected individuals, though sprung from the affected, ever have descendants who are affected, and (3) that in families where both affected and unaffected {173} occur, the numbers of the two classes are, on the average, equal. (The sum of such families in the complete pedigree is thirty-nine affected and thirty-six normals.) It is obvious that these are the conditions which are fulfilled in a simple Mendelian case, and there is nothing in this pedigree to contradict the assertion that brachydactyly, whatever it may be due to, behaves as a simple dominant to the normal form, i.e. that it depends upon a factor which the normal does not contain. The recessive normals cannot transmit the affected condition whatever their ancestry. Once free they are always free, and can marry other normals with full confidence that none of their children will show the deformity.



{174}

The evidence available from pedigrees has revealed the simplest form of Mendelian inheritance in several human defects and diseases, among which may be mentioned presenile cataract of the eyes, an abnormal form of skin thickening in the palms of the hands and soles of the feet, known as tylosis, and epidermolysis bullosa, a disease in which the skin rises up into numerous bursting blisters.

Among the most interesting of all human pedigrees is one recently built up by Mr. Nettleship from the records of a night-blind family living near Monpelier in the south of France. In night-blind people the retina is insensitive to light which falls below a certain intensity, and such people are consequently blind in failing daylight or in moonlight. As the Monpelier case had excited interest for some time, the records are unusually complete. They commence with a certain Jean Nougaret, who was born in 1637, and suffered from night-blindness, and they end for the present with children who are to-day but a few years of age. Particulars are known of over 2000 of the descendants of Jean Nougaret. Through ten generations and nearly three centuries the affection has behaved as a Mendelian dominant, and there is no sign that long-continued marriage with folk of normal vision has produced any amelioration of the night-blind state. {175}



Besides cases such as these where a simple form of Mendelian inheritance is obviously indicated, there are others which are more difficult to read. Of some it may be said that on the whole the peculiarity behaves as though it were an ordinary dominant; but that exceptions occur in which affected children are born to unaffected parents. It is not impossible that the condition may, like colour in the sweet pea, depend upon the presence or absence of more than one factor. In none of these cases, however, are the data sufficient for determining with certainty whether this is so or not.

A group of cases of exceptional interest is that in which the incidence of disease is largely, if not absolutely, restricted to one sex, and so far as is hitherto known the burden is invariably borne by the male. In the inheritance of colour-blindness (p. 117) we have already discussed an instance in which the defect is rare, though not {176} unknown, in the female. Sex-limited inheritance of a similar nature is known for one or two ocular defects, and for several diseases of the nervous system. In the peculiarly male disease known as haemophilia the blood refuses to clot when shed, and there is nothing to prevent great loss from even a superficial scratch. In its general trend the inheritance of haemophilia is not unlike that of horns among sheep, and it is possible that we are here again dealing with a character which is dominant in one sex and recessive in the other. But the evidence so far collected points to a difference somewhere, for in haemophilic families the affected males, instead of being equal in number to the unaffected, show a considerable preponderance. The unfortunate nature of the defect, however, forces us to rely for our interpretation almost entirely upon the families produced by the unaffected females who can transmit it. Our knowledge of the offspring of "bleeding" males is as yet far too scanty, and until it is improved, or until we can find some parallel case in animals or plants, the precise scheme of inheritance for haemophilia must remain undecided.

Though by far the greater part of the human evidence relates to abnormal or diseased conditions, a start has been made in obtaining pedigrees of normal characters. From the ease with which it can be observed, it was natural that eye-colour should be early selected as a subject of investigation, and the work of Hurst and others {177} has clearly demonstrated the existence of one Mendelian factor in operation here. Eyes are of many colours, and the colour depends upon the pigment in the iris. Some eyes have pigment on both sides of the iris—on the side that faces the retina as well as on the side that looks out upon the world. Other eyes have pigment on the retinal side only. To this class belong the blues and clear greys; while the eyes with pigment in front of the iris also are brown, hazel, or green in various shades according to the amount of pigment present. In albino animals the pigment is entirely absent, and as the little blood-vessels are not obscured the iris takes on its characteristic pinkish-red appearance. The condition in which pigment is present in front of the iris is dominant to that in which it is absent. Greens, browns, or hazels mated together may, if heterozygous, give the recessive blue, but no individuals of the brown class are to be looked for among the offspring of blues mated together. The blues, however, may carry factors which are capable of modifying the brown. Just as the pale pink-tinged sweet pea (Pl. IV., 9) when mated with a suitable white gives only deep purples, so an eye with very little brown pigment mated with certain blues produces progeny of a deep brown, far darker than either parent. The blue may carry a factor which brings about intensification of the brown pigment. There are doubtless other factors which modify the brown when present, but we do not yet know enough of the {178} inheritance of the various shades to justify any statement other than that the heredity of the pigment in front of the iris behaves as though it were due to a Mendelian factor.

Even this fact is of considerable importance, for it at once suggests that the present systems of classification of eye-colours, to which some anthropologists attach considerable weight, are founded on a purely empirical and unsatisfactory basis. Intensity of colour is the criterion at present in vogue, and it is customary to arrange the eye-colours in a scale of increasing depth of shade, starting with pale greys and ending with the deepest browns. On this system the lighter greens are placed among the blues. But we now know that blues may differ from the deep browns in the absence of only a single factor, while, on the other hand, the difference between a blue and a green may be a difference dependent upon more than one factor. To what extent eye-colour may be valuable as a criterion of race it is at present impossible to say, but if it is ever to become so, it will only be after a searching Mendelian analysis has disclosed the factors upon which the numerous varieties depend.

A discussion of eye-colour suggests reflections of another kind. It is difficult to believe that the markedly different states of pigmentation which occur in the same species are not associated with deep-seated chemical differences influencing the character and bent of the individual. {179} May not these differences in pigmentation be coupled with and so become in some measure a guide to mental and temperamental characteristics? In the National Portrait Gallery in London the pictures of celebrated men and women are largely grouped according to the vocations in which they have succeeded. The observant will probably have noticed that there is a tendency for a given type of eye-colour to predominate in some of the larger groups. It is rare to find anything but a blue among the soldiers and sailors, while among the actors, preachers, and orators the dark eye is predominant, although for the population as a whole it is far scarcer than the light. The facts are suggestive, and it is not impossible that future research may reveal an intimate connection between peculiarities of pigmentation and peculiarities of mind.

The inheritance of mental characters is often elusive, for it is frequently difficult to appraise the effects of early environment in determining a man's bent. That ability can be transmitted there is no doubt, for this is borne out by general experience, as well as by the numerous cases of able families brought together by Galton and others. But when we come to inquire more precisely what it is that is transmitted we are baffled. A distinguished son follows in the footsteps of a distinguished father. Is this due to the inheritance of a particular mental aptitude, or is it an instance of general mental ability displayed in a field rendered attractive by early association? We have {180} at present very little definite evidence for supposing that what appear to be special forms of ability may be due to specific factors. Hurst, indeed, has brought forward some facts which suggest that musical sense sometimes behaves as a recessive character, and it is likely that the study of some clean-cut faculty such as the mathematical one would yield interesting results.

The analysis of mental characters will no doubt be very difficult, and possibly the best line of attack is to search for cases where they are associated with some physical feature such as pigmentation. If an association of this kind be found, and the pigmentation factors be determined, it is evident that we should thereby obtain an insight into the nature of the units upon which mental conditions depend. Nor must it be forgotten that mental qualities, such as quickness, generosity, instability, etc.,—qualities which we are accustomed to regard as convenient units in classifying the different minds with which we are daily brought into contact,—are not necessarily qualities that correspond to heritable units. Effective mental ability is largely a matter of temperament, and this in turn is quite possibly dependent upon the various secretions produced by the different tissues of the body. Similar nervous systems associated with different livers might conceivably result in individuals upon whose mental ability the world would pass a very different judgment. Indeed, it is not at all impossible {181} that a particular form of mental ability may depend for its manifestation, not so much upon an essential difference in the structure of the nervous system, as upon the production by another tissue of some specific poison which causes the nervous system to react in a definite way. We have mentioned these possibilities merely to indicate how complex the problem may turn out to be. Though there is no doubt that mental ability is inherited, what it is that is transmitted, whether factors involving the quality and structure of the nervous system itself, or factors involving the production of specific poisons by other tissues, or both together, is at present uncertain.

Little as is known to-day of heredity in man, that little is of extraordinary significance. The qualities of men and women, physical and mental, depend primarily upon the inherent properties of the gametes which went to their making. Within limits these qualities are elastic, and can be modified to a greater or lesser extent by influences brought to bear upon the growing zygote, provided always that the necessary basis is present upon which these influences can work. If the mathematical faculty has been carried in by the gamete, the education of the zygote will enable him to make the most of it. But if the basis is not there, no amount of education can transform that zygote into a mathematician. This is a matter of common experience. Neither is there any reason for supposing that the superior education of a {182} mathematical zygote will thereby increase the mathematical propensities of the gametes which live within him. For the gamete recks little of quaternions. It is true that there is progress of a kind in the world, and that this progress is largely due to improvements in education and hygiene. The people of to-day are better fitted to cope with their material surroundings than were the people of even a few thousand years ago. And as time goes on they are able more and more to control the workings of the world around them. But there is no reason for supposing that this is because the effects of education are inherited. Man stores knowledge as a bee stores honey or a squirrel stores nuts. With man, however, the hoard is of a more lasting nature. Each generation in using it sifts, adds, and rejects, and passes it on to the next a little better and a little fuller. When we speak of progress we generally mean that the hoard has been improved, and is of more service to man in his attempts to control his surroundings. Sometimes this hoarded knowledge is spoken of as the inheritance which a generation receives from those who have gone before. This is misleading. The handing on of such knowledge has nothing more to do with heredity in the biological sense than has the handing on from parent to offspring of a picture, or a title, or a pair of boots. All these things are but the transfer from zygote to zygote of something extrinsic to the species. Heredity, on the other hand, deals with the {183} transmission of something intrinsic from gamete to zygote and from zygote to gamete. It is the participation of the gamete in the process that is our criterion of what is and what is not heredity.

Better hygiene and better education, then, are good for the zygote, because they help him to make the fullest use of his inherent qualities. But the qualities themselves remain unchanged in so far as the gamete is concerned, since the gamete pays no heed to the intellectual development of the zygote in whom he happens to dwell. Nevertheless, upon the gamete depend those inherent faculties which enable the zygote to profit by his opportunities, and, unless the zygote has received them from the gamete, the advantages of education are of little worth. If we are bent upon producing a permanent betterment that shall be independent of external circumstances, if we wish the national stock to become inherently more vigorous in mind and body, more free from congenital physical defect and feeble mentality, better able to assimilate and act upon the stores of knowledge which have been accumulated through the centuries, then it is the gamete that we must consult. The saving grace is with the gamete, and with the gamete alone.

People generally look upon the human species as having two kinds of individuals, males and females, and it is for them that the sociologists and legislators frame their schemes. This, however, is but an imperfect view to {184} take of ourselves. In reality we are of four kinds, male zygotes and female zygotes, large gametes and small gametes, and heredity is the link that binds us together. If our lives were like those of the starfish or the sea-urchin, we should probably have realised this sooner. For the gametes of these animals live freely, and contract their marriages in the waters of the sea. With us it is different, because half of us must live within the other half or perish. Parasites upon the rest, levying a daily toll of nutriment upon their hosts, they are yet in some measure the arbiters of the destiny of those within whom they dwell. At the moment of union of two gametes is decided the character of another zygote, as well as the nature of the population of gametes which must make its home within him. The union once affected the inevitable sequence takes its course, and whether it be good, or whether it be evil, we, the zygotes, have no longer power to alter it. We are in the hands of the gamete; yet not entirely. For though we cannot influence their behaviour we can nevertheless control their unions if we choose to do so. By regulating their marriages, by encouraging the desirable to come together, and by keeping the undesirable apart we could go far towards ridding the world of the squalor and the misery that come through disease and weakness and vice. But before we can be prepared to act, except, perhaps, in the simplest cases, we must learn far more about them. At present we are woefully ignorant {185} of much, though we do know that full knowledge is largely a matter of time and means. One day we shall have it, and the day may be nearer than most suspect. Whether we make use of it will depend in great measure upon whether we are prepared to recognise facts, and to modify or even destroy some of the conventions which we have become accustomed to regard as the foundations of our social life. Whatever be the outcome, there can be little doubt that the future of our civilisation, perhaps even the possibility of a future at all, is wrapped up with the recognition we accord to those who live unseen and inarticulate within us—the fateful race of gametes so irrevocably bound to us by that closest of all ties, heredity.

* * * * * {187}

APPENDIX

As some readers may possibly care to repeat Mendel's experiments for themselves, a few words on the methods used in crossing may not be superfluous. The flower of the pea with its standard, wings, and median keel is too familiar to need description. Like most flowers it is hermaphrodite. Both male and female organs occur on the same flower, and are covered by the keel. The anthers, ten in number, are arranged in a circle round the pistil. As soon as they are ripe they burst and shed their pollen on the style. The pollen tubes then penetrate the stigma, pass down the style, and eventually reach the ovules in the lower part of the pistil. Fertilisation occurs here. Each ovule, which is reached by a pollen tube, swells up and becomes a seed. At the same time the fused carpels enclosing the ovules enlarge to form the pod. When this, the normal mode of fertilisation, takes place, the flower is said to be SELFED.

In crossing, it is necessary to emasculate a flower on the plant chosen to be the female parent. For this purpose a young flower must be taken in which the anthers have not yet burst. The keel is depressed, and the stamens bearing the anthers are removed at their base by a {188} pair of fine forceps. It will probably be found necessary to tear the keel slightly in order to do this. The pistil is then covered up again with the keel, and the flower is enclosed in a bag of waxed paper until the following day. The stigma is then again exposed and dusted with ripe pollen from a flower of the plant selected as the male parent. This done, the keel is replaced, and the flower again enclosed in its bag to protect it from the possible attentions of insects until it has set seed. The bag may be removed in about a week after fertilisation. It is perhaps hardly necessary to add that strict biological cleanliness must be exercised during the fertilising operations. This is readily attained by sterilising fingers and forceps with a little strong spirit before each operation, thereby ensuring the death of any foreign pollen grains which may be present.

The above method applies also to sweet peas, with these slight modifications. As the anthers ripen relatively sooner in this species, emasculation must be performed at a rather earlier stage. It is generally safe to choose a bud about three parts grown. The interval between emasculation and fertilisation must be rather longer. Two to three days is generally sufficient. Further, the sweet pea is visited by the leaf-cutter bee, Megachile, which, unlike the honey bee, is able to depress the keel and gather pollen. If the presence of this insect is suspected, it is desirable to guard against the risk of admixture of {189} foreign pollen by selecting for pollinating purposes a flower which has not quite opened. If the standard is not erected, it is unlikely to have been visited by Megachile. Lastly, it not infrequently happens that the little beetle Meligethes is found inside the keel. Such flowers should be rejected for crossing purposes.

* * * * *

{191}

INDEX

Abraxas grossulariata, 99 "Acquired" characters, 14 Adaptation, 143 Agouti mice, 50 Albino mice, 50 Albinos, nature of, 53 Amauris, 144 Analysis of types, 156 Ancestral Heredity, Law of, 13 Andalusian fowls, 70 Axil colour in sweet peas, 93

Bateson, W., 14, 29, 55, 116, 132, 141 Biffen, R. H., 157 Blue Andalusian fowls, 71 Brachydactyly, 171 Bryony, 120 Bush sweet peas, 63

Castle, 132 Cattle, horns in, 86, 166 Colour, nature of, in flowers, 48 Colour-blindness, 117 Combs of fowls, 33, 43 Correns, C., 29, 120 Coupling of characters in gametes, 93 Cuenot, 50, 119 "Cupid" sweet peas, 62 Currant moth, 99

Darwin, C., 10, 65, 147, 163 De Vries, H., 15, 29, 141 Discontinuity in variation, 14 Dominant characters, 18 Doncaster, L., 99 Drinkwater, H., 172 Dutch rabbits, 60

Eggs, 2 Environment, influence of, 137 Euralia, 144 Evolution, 10, 85, 139 Eye, in primulas, 55 Eye-colour, in man, 176

Factor, definition of, 31 Factors, interaction of, 42 Fertilisation, 3 Fertilisation, self- and cross-, 163 Fixation of varieties, 153 Fluctuations, 138 Fowls, coloured from whites, 49, 73

Galton, 13, 179 Gametes, nature of, 6 Gregory, R. P., 55, 93

Haemophilia, 176 Hardy, G. H., 147 Heterozygote, definition of, 28 Heterozygote, of intermediate form, 68 Hieracium, 27, 132 Himalayan rabbits, 60 Homostyle primulas, 56 Homozygote, definition of, 28 Hooded sweet peas, 89 Horses, bay and chestnut in, 167 Hurst, C. C., 62, 176, 180

Immunity in wheat, 158 Individuality, 135 Inhibition, factors for, 74, 108 Intermediates, 125 {192}

Johannsen, W., 160

Lop-eared rabbits, 132

Mendel, 8, 17, 26, 132 Mental characters, 180 Mice, inheritance of coat colour in, 50 Mimicry, 143 Mirabilis, 151 Morgan, T. H., 116 Mulattos, 129 Mutation, 83, 138

Naegeli, C., 26 Natural selection, 11, 140, 142, 149 Nettleship, E., 175 Night-blindness, 175

Pararge egeria, 132 Parkinson, J., 122 Pea comb, 33 Peas, coloured flowers in, 24 Peas, tall and dwarf, 18 Pigeons, 86 Pin-eye in primulas, 55 Pisum, 17 Primulas, 31, 55, 68, 93 Pollen, 3 Pollen of sweet peas, 92 Pomace fly, 115 Population, inheritance of characters in a, 147 Presence and Absence theory, 35 Pure lines, 162 Purity of gametes, 24 Purity of type, 155

Rabbits, 53, 60 Ratios, Mendelian— 3 : 1, 20 9 : 3 : 3 : 1, 25, 34 9 : 3 : 4, 51 9 : 7, 49 Ray, John, 143 Recessive characters, 19 Repulsion between factors, 90 Reversion, 59, 165 in rabbits, 59 in sweet peas, 62 in fowls, 65 in pigeons, 65 Rose comb, 33

Saunders, E. R., 54, 122 Seeds, nature of, 4 Segregation, 22 Selection, 162 Sheep, horns in, 76 Silky fowls, 30, 105 Single comb, 32 Species, nature of, 150 Species, origin of, 11 Speckled wood butterfly, 132 Spermatozoa, 3 Sports, 147 Staples-Browne, R., 66 Sterility, 151 Sterility in sweet peas, 93 Stocks, double, 122 Stocks, hoariness in, 54 Sweet pea, colour in, 44, 79 history of, 82 inheritance of hood in, 89 inheritance of size in, 62

Telegony, 167 Thrum-eye in primulas, 55 Toe, extra toe in poultry, 76 Tschermak, E., 29

Unit-character, definition of, 31

Variation, 14, 137, 139

Walnut comb, 33 Weismann, A., 13 Wheat, beard in, 74 experiments with, 157 White, dominant in poultry, 72 Wilson, J., 168

Yellow mice, 119

Zygotes, nature of, 5

* * * * *

Notes

* * * * *

[1] Cf. note on p. 171.

[2] It has been found convenient to denote the various generations resulting from a cross by the signs F1, F2, F3, etc. F1 on this system denotes the first filial generation, F2 the second filial generation produced by two parents belonging to the F1 generation, and so on.

[3] Hurst's original cross was between a Belgian hare and an albina Angora, which turned out to be a masked Dutch.

[4] The Spot is an almost white bird, the colour being confined to the tail and the characteristic spot on the head.

[5] The reader who searches florists' catalogues for these varieties will probably experience disappointment. The sweet pea has been much "improved" in the past few years, and it is unlikely that the modern seedsman would list such unfashionable forms.

[6] It is to be understood that wherever a given factor is present the plant may be homozygous or heterozygous for it without alteration in its colour.

[7] It should be mentioned that as the shape of the pollen coat, like that of the seed coat, is a maternal character, all the grains of any given plant are either long or else round. The two kinds do not occur together on the same plant.

[8] For the most recent discussion of this peculiar case the reader is referred to Professor Castle's paper in Science, December 16, 1910.

[9] Paradisus Terrestris, London, 1629, p. 261.



* * * * *



Transcriber's note:

Corrections made to printed original.

Page 36, "two sorts, RP and Rp": 'PR and Rp' in original.

Page 51, "9 contain both C and G": 'c and G' in original.

Page 184, "in the simplest cases": 'simples' in original.

Footnote 3, "turned out to be a masked Dutch": 'turned to out be' in original.

THE END

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