Read Ebook: Mendelism Third Edition by Punnett Reginald Crundall

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I promised good faith, and immediately he burst into the frantic blasphemy of one who has escaped from supervision, cursing the priests, the lying filthy cheating immoral priests who had ruined his life, who had murdered his brother and the girl, whom he dared not murder back because they held the key of heaven and could ruin him in the next life too. "Thus are we tricked!" was his cry and he stood up and kicked at the azure ripples with his feet, till he had obscured them with a cloud of sand.

I too was moved. The story of Giuseppe, for all itis 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 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 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.

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: 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; a period of association in which two gametes become yoked together into a zygote and react upon one 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 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 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.

HISTORICAL

A further important step was taken in 1895, when Bateson once more drew attention to the problem of the origin 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.

MENDEL'S WORK

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, the result of these experiments may be briefly summarised in the foregoing scheme.

THE PRESENCE AND ABSENCE THEORY

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 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 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.

FIG. 5.

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 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.

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.

INTERACTION OF FACTORS

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.

FIG. 7.

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: a colourless "chromogen," or colour basis; and 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 in the gametes or whether in some other form we have as yet no means of deciding.

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 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.

FIG. 8.

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, 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 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.

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REVERSION

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.

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

FIG. 11.

DOMINANCE

In such cases as this it is obvious that we cannot speak of dominance. And with the disappearance of this phenomenon we lose one criterion for determining which of the two parent forms possesses the additional factor. Are we, for example, to regard the black Andalusian as a splashed white to which has been added a double dose of a colour-intensifying factor, or are we to consider the white splashed bird as a black which is unable to show its true pigmentation owing to the possession of some inhibiting factor which prevents the manifestation of the black. Either interpretation fits the facts equally well, and until further experiments have been devised and carried out it is not possible to decide which is the correct view.

Besides these comparatively rare cases where the heterozygote cannot be said to bear a closer resemblance to one parent more than to the other, there are cases in which it is often possible to draw a visible distinction between the heterozygote and the pure dominant. There are certain white breeds of poultry, notably the White Leghorn, in which the white behaves as a dominant to colour. But the heterozygous whites made by crossing the dominant white birds with a pure coloured form almost invariably show a few coloured feathers or "ticks" in their plumage. The dominance of white is not quite complete, and renders it possible to distinguish the pure from the impure dominant without recourse to breeding experiments.

FIG. 13.

WILD FORMS AND DOMESTIC VARIETIES

Again, the proportion in which the three classes of purples appeared was 9 bicolors, 3 deep purples, 4 picotees. We are, therefore, concerned here with the operation of two factors: a light wing factor, which renders the bicolor dominant to the dark winged form; and a factor for intense colour, which occurs in the bicolor and in the deep purple, but is lacking in the dilute picotee. And here it should be mentioned that these conclusions rest upon an exhaustive set of experiments involving the breeding of many thousands of plants. In this cross, therefore, we are concerned with the presence or absence of five factors, which we may denote as follows:--

On this notation our six coloured forms are:--

It will be noticed in this series that the various coloured forms can be expressed by the omission of one or more factors from the purple bicolor of the wild type. With the complete omission of each factor a new colour type results, and it is difficult to resist the inference that the various cultivated forms of the sweet pea have arisen from the wild by some process of this kind. Such a view tallies with what we know of the behaviour of the wild form when crossed by any of the garden varieties. Wherever such crossing has been made the form of the hybrid has been that of the wild, thus supporting the view that the wild contains a complete set of all the differentiating factors which are to be found in the sweet pea.

Moreover, this view is in harmony with such historical evidence as is to be gleaned from botanical literature, and from old seedsmen's catalogues. The wild sweet pea first reached England in 1699, having been sent from Sicily by the monk Franciscus Cupani as a present to a certain Dr. Uvedale in the county of Middlesex. Somewhat later we hear of two new varieties, the red bicolor, or Painted Lady, and the white, each of which may be regarded as having "sported" from the wild purple by the omission of the purple factor, or of one of the two colour factors. In 1793 we find a seedsman offering also what he called black and scarlet varieties. It is probable that these were our deep purple and Miss Hunt varieties, and that somewhere about this time the factor for the light wing was dropped out in certain plants. In 1860 we have evidence that the pale purple or Picotee, and with it doubtless the Tinged White, had come into existence. This time it was the factor for intense colour which had dropped out. And so the story goes on until the present day, and it is now possible to express by the same simple method the relation of the modern shades, of purple and reds, of blues and pinks, of hooded and wavy standards, to one another and to the original wild form. The constitution of many of these has now been worked out, and to-day it would be a simple though perhaps tedious task to denote all the different varieties by a series of letters indicating the factors which they contain, instead of by the present system of calling them after kings and queens, and famous generals, and ladies more or less well known.

Why these mutations arise: what leads to the surmised unequal division of the gametes: of this we know practically nothing. Nor until we can induce the production of mutations at will are we likely to understand the conditions which govern their formation. Nevertheless there are already hints scattered about the recent literature of experimental biology which lead us to hope that we may know more of these matters in the future.

In respect of the evolution of its now multitudinous varieties, the story of the sweet pea is clear and straightforward. These have all arisen from the wild by a process of continuous loss. Everything was there in the beginning, and as the wild plant parted with factor after factor there came into being the long series of derived forms. Exquisite as are the results of civilization, it is by the degradation of the wild that they have been brought about. How far are we justified in regarding this as a picture of the manner in which evolution works?

There are certainly other species in which we must suppose that this is the way that the various domesticated forms have arisen. Such, for example, is the case in the rabbit, where most of the colour varieties are recessive to the wild agouti form. Such also is the case in the rat, where the black and albino varieties and the various pattern forms are also recessive to the wild agouti type. And with the exception of a certain yellow variety to which we shall refer later, such is also the case with the many fancy varieties of mice.

On the whole, therefore, we must be prepared to admit that the evolution of domestic varieties may come about by a process of addition of factors in some cases and of subtraction in others. It may be that what we term additional factors fall into distinct categories from the rest. So far, experiment seems to show that they are either of the nature of melanic factors, or of inhibitory factors, or of reduplication factors as in the case of the fowls' combs. But while the data remain so scanty, speculation in these matters is too hazardous to be profitable.

REPULSION AND COUPLING OF FACTORS

Although different factors may act together to produce specific results in the zygote through their interaction, yet in all the cases we have hitherto considered the heredity of each of the different factors is entirely independent. The interaction of the factors affects the characters of the zygote, but makes no difference to the distribution of the separate factors, which is always in strict accordance with the ordinary Mendelian scheme. Each factor in this respect behaves as though the other were not present.

A few cases have been worked out in which the distribution of the different factors to the gametes is affected by their simultaneous presence in the zygote. And the influence which they are able to exert upon one another in such cases is of two kinds. They may repel one another, refusing, as it were, to enter into the same zygote, or they may attract one another, and, becoming linked together, pass into the same gamete, as it were by preference. For the moment we may consider these two sets of phenomena apart.

These four cases have already been found in the sweet pea, and similar phenomena have been met with by Gregory in primulas. To certain seemingly analogous cases in animals where sex is concerned we shall refer later.

Nor for the present can we suggest why certain factors should be linked together in the peculiar way that we have reason to suppose that they are during the process of the formation of the gametes. Nevertheless the phenomena are very definite, and it is not unlikely that a further study of them may throw important light on the architecture of the living cell.

As it is possible that some readers may care, in spite of its complexity, to enter rather more fully into the peculiar phenomenon of the coupling of characters, I have brought together some further data in this Appendix. In the case we have already considered, where the factors for blue colour and long pollen are concerned, we have been led to suppose that the gametes produced by the heterozygous plant are of the nature 7 BL : 1 Bl : 1 bL : 7 bl. Such a series of ovules fertilised by a similar series of pollen grains will give a generation of the following composition:--

In such a series the number of gametes containing A is equal to the number lacking A, and the same is true for B. Consequently the number of zygotes formed containing A is three times as great as the number of zygotes which do not contain A; and similarly for B. The proportion of dominants to recessives in each case is 3 : 1. It is only in the distribution of the characters with relation to one another that these cases differ from a simple Mendelian case.

As the study of these series presents another feature of some interest, we may consider it in a little more detail. In the accompanying table are set out the results produced by these different series of gametes. The series marked by an asterisk have already been demonstrated experimentally. The first term in the series, in which all the four kinds of gametes are produced in equal numbers is, of course, that of a simple Mendelian case where no coupling occurs.

SEX

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