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Mendelism - Third Edition
by Reginald Crundall Punnett
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FIG. 11.

Diagram to illustrate the appearance of the reversionary blue pigeon in F_2 from the cross of black with white.

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{68}

CHAPTER VII

DOMINANCE



Sinensis x Stellata Intermediate F1 + -+ + -+ -+ Sinensis Inter. Inter. Stellata - F2 Sinensis Stellata - F{3} + -+ -+ -+ Sinensis Sin. Int. Int. Stell. Stellata - F{4}

In the cases which we have hitherto considered the presence of a factor produces its full effect whether it is introduced by both of the gametes which go to form the zygote, or by one of them alone. The heterozygous tall pea or the heterozygous rose-combed fowl cannot be distinguished from the homozygous form by mere inspection, however close. Breeding tests alone can decide which is the heterozygous and which the homozygous form. Though this is true for the majority of characters yet investigated, there are cases known in which the heterozygous form differs in appearance from either parent. Among plants such a case has been met with in the primula. The ordinary Chinese primula (_P. sinensis_) (Fig. 12) has large rather wavy petals much crenated at the edges. In the Star Primula (_P. stellata_) the flowers are much smaller, while the petals are flat and present only a terminal notch instead of the numerous crenations of _P. sinensis_. The heterozygote produced by crossing these forms is intermediate in size and appearance. When self-fertilised such plants behave in simple Mendelian fashion, {69} giving a generation consisting of _sinensis_, intermediates, and _stellata_ in the ratio 1 : 2 : 1. Subsequent breeding from these plants showed that both the _sinensis_ and _stellata_ which appeared in the F_2 generation bred true, while the intermediates always gave all three forms again in the same proportion. But though there is no dominance of the character of either parent in such a case as this, the Mendelian principle of segregation could hardly have a better illustration.

{70}

Blue x Blue + -+ + + Black Blue x Blue White + + + + -+ Black Black Blue Blue White White Black - x - White Blue (all)

Among birds a case of similar nature is that of the Blue Andalusian fowl. Fanciers have long recognised the difficulty of getting this variety to breed true. Of a slaty blue colour itself with darker hackles and with black lacing on the feathers of the breast, it always throws "wasters" of two kinds, viz. blacks, and whites splashed with black. Careful breeding from the blues shows that the three sorts are always produced in the same definite {71} proportions, viz., one black, two blues, one splashed white. This at once suggests that the black and the splashed white are the two homozygous forms, and that the blues are heterozygous, i.e., producing equal numbers of "black" and "white splashed" gametes. The view was tested by breeding the "wasters" together—black with black, and splashed white with splashed white—and it was found that each bred true to its respective type. But when the black and the splashed white were crossed they gave, as was expected, nothing but blues. In other words, we have the seeming paradox of the black and the splashed white producing twice as many blues as do the blues when bred together. The black and the splashed white "wasters" are in reality the pure breeds, while the "pure" Blue Andalusian is a mongrel which no amount of selection will ever be able to fix.

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, {72} 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 (such as the Brown Leghorn) 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.

+ + + + + CI CI CI CI CI Ci cI ci + + + + + Ci Ci.... Ci Ci.... CI Ci.... cI ci.... ...... ...... ...... ...... + + + + + cI cI cI cI CI Ci cI ci + + + + + ci ci.... ci ci CI Ci.... cI ci ...... ...... + + + + +

FIG. 13.

Diagram to illustrate the nature of the F_2 generation from the cross between dominant white and recessive white fowls.

This case of the dominant white fowl opens up another interesting problem in connection with dominance. By accepting the "Presence and Absence" hypothesis we are committed to the view that the dominant form possesses an extra factor as compared with the recessive. The natural way of looking at this case of the fowl is to regard white as the absence of colour. But were this so, colour should be dominant to white, which is not the case. We are therefore forced to suppose that the absence of colour in this instance is due to the presence of a factor whose {73} property is to inhibit the production of colour in what would otherwise be a pure coloured bird. On this view the dominant white fowl is a coloured bird plus a factor which inhibits the development of the colour. The view can be put to the test of experiment. We have already seen that there are other white fowls in which white is recessive to colour, and that the whiteness of such birds is due to the fact that they lack a factor for the development of colour. If we denote this factor by C and our postulated inhibitor factor in the dominant white bird by I, then we must write the constitution of the recessive white as ccii, and the dominant white as CCII. We may now work out the results we ought to obtain when a cross is made between these two pure white breeds. The constitution of the F_1 bird must be CcIi. Such birds being heterozygous for the inhibitor factor, should be whites showing some coloured "ticks." Being heterozygous for both of the two factors C and I, they will produce in equal numbers the four different sorts of gametes CI, Ci, cI, ci. The result of bringing two such similar series of gametes together is shown in Fig. 13. Out of the sixteen squares, twelve contain I; these will be white birds either with or without a few coloured ticks. Three contain C but not I: these must be coloured birds. One contains neither C nor I; this must be a white. From such a mating we ought, therefore, to obtain both white and coloured birds in the ratio 13 : 3. The results thus theoretically {74} deduced were found to accord with the actual facts of experiment. The F_1 birds were all "ticked" whites, and in the F_2 generation came white and coloured birds in the expected ratio. There seems, therefore, little reason to doubt that the dominant white is a coloured bird in which the absence of colour is due to the action of a colour-inhibiting factor, though as to the nature of that factor we can at present make no surmise. It is probable that other facts, which at first sight do not appear to be in agreement with the "Presence and Absence" hypothesis, will eventually be brought into line through the action of inhibitor factors. Such a case, for instance, is that of bearded and beardless wheats. Though the beard is obviously the additional character, the bearded condition is recessive to the beardless. Probably we ought to regard the beardless as a bearded wheat in which there is an inhibitor that stops the beard from growing. It is not unlikely that as time goes on we shall {75} find many more such cases of the action of inhibitor factors, and we must be prepared to find that the same visible effect may be produced either by the addition or by the omission of a factor. The dominant and recessive white poultry are indistinguishable in appearance. Yet the one contains a factor more and the other a factor less than the coloured bird.



{76}

A phenomenon sometimes termed irregularity of dominance has been investigated in a few cases. In certain breeds of poultry such as Dorkings there occurs an extra toe directed backwards like the hallux (cf. Fig. 15). In some families this character behaves as an ordinary dominant to the normal, giving the expected 3 : 1 ratio in F_2. But in other families similarly bred the proportions of birds with and without the extra toe appear to be unusual. It has been shown that in such a family some of the birds without the extra toe may nevertheless transmit the peculiarity when mated with birds belonging to strains in which the extra toe never occurs. Though the external appearance of the bird generally affords some indication of the nature of the gametes which it is carrying, this is not always the case. Nevertheless we have reason to suppose that the character segregates in the gametes, though the nature of these cannot always be decided from the appearance of the bird which bears them.



There are cases in which an apparent irregularity of dominance has been shown to depend upon another character, as in the experiments with sheep carried out by Professor Wood. In these experiments two breeds were crossed, of which one, the Dorset, is horned in both sexes, while the other, the Suffolk, is without horns in either sex. Whichever way the cross was made the resulting F1 generation was similar; the rams were horned, and {77} the ewes were hornless. In the F2 generation raised from these F1 animals both horned and hornless types appeared in both sexes but in very different proportions. While the horned rams were about three times as numerous as the hornless, this relation was reversed among the females, in which the horned formed only about one-quarter of the total. The simplest explanation of this interesting case is to suppose that the dominance of the horned character depends upon the sex of the animal—that it is dominant in the male but recessive in the female. A pretty experiment was devised for putting this view to the test. If it is true, equal numbers of gametes with and without the horned factor must be produced by the F1 ewes, while the factor should be lacking in all the gametes of the hornless F2 rams. A {78} hornless ram, therefore, put to a flock of F1 ewes should give rise to equal numbers of zygotes which are heterozygous for the horned character, and of zygotes in which it is completely absent. And since the heterozygous males are horned, while the heterozgyous females are hornless, we should expect from this mating equal numbers of horned and hornless rams, but only hornless ewes. The result of the experiment confirmed this expectation. Of the ram lambs 9 were horned and 8 were hornless, while all the 11 ewe lambs were completely destitute of horns.



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{79}

CHAPTER VIII

WILD FORMS AND DOMESTIC VARIETIES

In discussing the phenomena of reversion we have seen that in most cases such reversion occurs when the two varieties which are crossed each contain certain factors lacking in the other, of which the full complement is necessary for the production of the reversionary wild form. This at once suggests the idea that the various domestic forms of animals and plants have arisen by the omission from time to time of this factor or of that. In some cases we have clear evidence that this is the most natural interpretation of the relation between the cultivated and the wild forms. Probably the species in which it is most evident is the sweet pea (_Lathyrus odoratus_). We have already seen reason to suppose that as regards certain structural features the Bush variety is a wild lacking the factor for the procumbent habit, that the Cupid is a wild without the factor for the long inter-node, and that the Bush Cupid is a wild minus both these factors. Nor is the evidence less clear for the many colour varieties. In illustration we may consider in more detail a case in which the cross between two whites resulted {80} in a complete reversion to the purple colour characteristic of the wild Sicilian form (Pl. IV.). In this particular instance subsequent breeding from the purples resulted in the production of six different colour forms in addition to whites. The proportion of the coloured forms to the whites was 9 : 7 (cf. p. 44), but it is with the relation of the six coloured forms that we are concerned here. Of these six forms three were purples and three were reds. The three purple forms were (1) the wild bicolor purple with blue wings known in cultivation as the Purple Invincible (Pl. IV., 4); (2) a deep purple with purple wings (Pl. IV., 5); and (3) a very dilute purple known as the Picotee (Pl. IV., 6). Corresponding to these three purple forms were three reds: (1) a bicolor red known as Painted Lady (Pl. IV., 7); (2) a deep red with red wings known as Miss Hunt (Pl. IV., 8); and (3) a very pale red which we have termed Tinged White[5] (Pl. IV., 9). In the F_2 generation the total number of purples bore to the total number of reds the ratio 3 : 1, and this ratio was maintained for each of the corresponding classes. Purple, therefore, is dominant to red, and each of the three classes of red differs from its corresponding purple in not possessing the blue factor (B) which turns it into purple.



{81} 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: (1) a light wing factor, which renders the bicolor dominant to the dark winged form; and (2) 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:—

A colour base, R. A colour developer, C. A purple factor, B. A light wing factor, L. A factor for intense colour, I.

On this notation our six coloured forms are:—

(1) Purple bicolor CRBLI.[6] (2) Deep purple CRBlI. (3) Picotee CRBLi or CRBli. (4) Red bicolor ( = Painted Lady) CRbLI. (5) Deep red ( = Miss Hunt) CRblI. (6) Tinged white CRbLi or CRbli.

It will be noticed in this series that the various coloured {82} 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 {83} light wing (L) 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.

From what we know of the history of the various strains of sweet peas one thing stands out clearly. The new character does not arise from a pre-existing variety by any process of gradual selection, conscious or otherwise. It turns up suddenly complete in itself, and thereafter it can be associated by crossing with other existing characters to produce a gamut of new varieties. If, for example, the character of hooding in the standard (cf. Pl. II., 7) suddenly turned up in such a family as that shown on Plate IV. we should be able to get a hooded form corresponding to each of the forms with the erect {84} standard; in other words, the arrival of the new form would give us the possibility of fourteen varieties instead of seven. As we know, the hooded character already exists. It is recessive to the erect standard, and we have reason to suppose that it arose as a sudden sport by the omission of the factor in whose presence the standard assumes the erect shape characteristic of the wild flower. It is largely by keeping his eyes open and seizing upon such sports for crossing purposes that the horticulturist "improves" the plants with which he deals. How these sports or mutations come about we can now surmise. They must owe their origin to a disturbance in the processes of cell division through which the gametes originate. At some stage or other the normal equal distribution of the various factors is upset, and some of the gametes receive a factor less than others. From the union of two such gametes, provided that they are still capable of fertilisation, comes the zygote which in course of growth develops the new character.

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. {85}

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.

Nevertheless there are other cases in which we must suppose evolution to have proceeded by the interpolation of characters. In discussing reversion on crossing, we have already seen that this may not occur until the F_2 generation, as, for example, in the instance of the fowls' combs (cf. p. 65). The reversion to the single comb occurred as the result of the removal of the two factors {86} for rose and pea. These two domesticated varieties must be regarded as each possessing an additional factor in comparison with the wild single-combed bird. During the evolution of the fowl, these two factors must be conceived of as having been interpolated in some way. And the same holds good for the inhibitory factor on which, as we have seen, the dominant white character of certain poultry depends. In pigeons, too, if we regard the blue rock as the ancestor of the domesticated breeds, we must suppose that an additional melanic factor has arisen at some stage. For we have already seen that black is dominant to blue, and the characters of F_1, together with the greater number of blacks than blues in F_2, negatives the possibility that we are here dealing with an inhibitory factor. The hornless or polled condition of cattle, again, is dominant to the horned condition, and if, as seems reasonable, we regard the original ancestors of domestic cattle as having been horned, we have here again the interpolation of an inhibitory factor somewhere in the course of evolution.

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 {87} 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.

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{88}

CHAPTER IX

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. {89}

One of the best illustrations of repulsion between factors occurs in the sweet pea. We have already seen that the loss of the blue or purple factor (B) from the wild bicolor results in the formation of the red bicolor known as Painted Lady (Pl. IV., 7). Further, we have seen that the hooded standard is recessive to the ordinary erect standard. The omission of the factor for the erect standard (E) from the purple bicolor (Pl. II., 5) results in a hooded purple known as Duke of Westminster (Pl. II., 7). And here it should be mentioned that in the corresponding hooded forms the difference in colour between the wings and standard is not nearly so marked as in the forms with the erect standard, but the difference in structure appears to affect the colour, which becomes nearly uniform. This may be readily seen by comparing the picture of the purple bicolor on Plate II. with that of the Duke of Westminster flower.

Now when a Duke of Westminster is mated with a Painted Lady the factor for erect standard (E) is brought in by the red, and that for blue (B) by the Duke, and the offspring are consequently all purple bicolors. Purples so formed are all heterozygous for these two factors, and were the case a simple one, such as those which have already been discussed, we should expect the F_2 generation to consist of the four forms: erect purple, hooded purple, erect red, and hooded red in the ratio 9 : 3 : 3 : 1. Such, however, is not the case. The F_2 generation {90} actually consists of only three forms, viz. erect red, erect purple, and hooded purple, and the ratio in which these three forms occur is 1 : 2 : 1. No hooded red has been known to occur in such a family. Moreover further breeding shows that while the erect reds and the hooded purples always breed true, the erect purples in such families _never_ breed true, but always behave like the original F_1 plant, giving the three forms again in the ratio 1 : 2 : 1. Yet we know that there is no difficulty in getting purple bicolors to breed true from other families; and we know also that hooded red sweet peas exist in other strains.

Painted Lady x Duke of Westminster (erect red) (hooded purple) Purple Invincible (erect purple) + -+ -+ Painted Purple Invincible Duke of Lady Westminster (1) (2) (1)



EEbb eeBB Parents / / / / / / Eb Eb eB eB gametes ——————/ EeBb F2 / / Fem. gametes of F1 Eb —-> EEbb <—- Eb Male gametes of F1 Eb —-> EeBb EeBb eeBB <—- eB ——/ F2 generation

On the assumption that there exists a repulsion between the factors for erect standard and blue in a plant which is heterozygous for both, this peculiar case receives a simple explanation. The constitutions of the erect red and the hooded purple are EEbb and eeBB respectively and that of the F_1 erect purple is EeBb. Now let us suppose that in such a zygote there exists a repulsion {91} between E and B, such that when the plant forms gametes these two factors will not go into the same gamete. On this view it can only form two kinds of gametes, viz. Eb and eB, and these, of course, will be formed in equal numbers. Such a plant on self-fertilisation must give the zygotic series EEbb + 2 EeBb + eeBB, _i.e._ 1 erect red, 2 erect purples, and 1 hooded purple. And because the erect reds and the hooded purples are respectively homozygous for E and B, they must thenceforward breed true. The erect purples, on the other hand, being always formed by the union of a gamete Eb with a gamete eB, are always heterozygous for both of these factors. They can, consequently, never breed true, but must always give erect reds, erect purples, and hooded purples in the ratio 1 : 2 : 1. The experimental facts are readily explained on the assumption of repulsion between the two {92} factors B and E during the formation of the gametes in a plant which is heterozygous for both.

Other similar cases of factorial repulsion have been demonstrated in the sweet pea, and two of these are also concerned with the two factors with which we have just been dealing. Two distinct varieties of pollen grains occur in this species, viz. the ordinary oblong form and a rather smaller rounded grain. The former is dominant to the latter.[7] When a cross is made between a purple with round pollen and a red with long pollen the F_1 plant is a long pollened purple. But the F_2 generation consists of purples with round pollen, purples with long pollen, and reds with long pollen in the ratio 1 : 2 : 1. No red with round pollen appears in F_2 owing to repulsion between the factors for purple (B) and for long pollen (L). Similarly plants produced by crossing a red hooded long with a red round having an erect standard give in F_1 long pollened reds with an erect standard, and these in F_2 produce the three types, round pollened erect, long pollened erect, and long pollened hooded, in the ratio 1 : 2 : 1. The repulsion here is between the long pollen factor (L) and the factor for the erect standard (E).

{93}

Yet another similar case is known in which we are concerned with quite different factors. In some sweet peas the axils whence the leaves and flower stalks spring from the main stem are of a deep red colour. In others they are green. The dark pigmented axil is dominant to the light one. Again, in some sweet peas the anthers are sterile, setting no pollen, and this condition is recessive to the ordinary fertile condition. When a sterile plant with a dark axil is crossed by a fertile plant with a light axil, the F1 plants are all fertile with dark axils. But such plants in F2 give fertiles with light axils, fertiles with dark axils, and steriles with dark axils in the ratio 1 : 2 : 1. No light axilled steriles appear from such a cross owing to the repulsion between the factor for dark axil (D) and that for the fertile anther (F).

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.

Now all of these four cases present a common feature which probably has not escaped the attention of the reader. In all of them _the original cross was such as to introduce one of the repelling factors with each of the two parents_. If we denote our two factors by A and B, the crosses have always been of the nature AAbb x aaBB. Let us now consider what happens when both of the {94} factors, which in these cases repel one another, are introduced by one of the parents, and neither by the other parent. And in particular we will take the case in which we are concerned with purple and red flower colour, and with long and round pollen, _i.e._ with the factors B and L. When a purple long (BBLL) is crossed with a red round (bbll) the F_1 (BbLl) is a purple with long pollen, identical in appearance with that produced by crossing the long pollened red with the round pollened purple. But the nature of the F_2 generation is in some respects very different. The ratio of purples to reds and of longs to rounds is in each case 3 : 1, as before. But instead of an association between the red and the long pollen characters the reverse is the case. The long pollen character is now associated with purple and the round pollen with red. The association, however, is not quite complete, and the examination of a large quantity of similarly bred material shows that the purple longs are about twelve times as numerous as the purple rounds, while the red rounds are rather more than three times as many as the red longs. Now this peculiar result could be brought about if the gametic series produced by the F_1 plant consisted of 7 BL + 1 Bl + 1 bL + 7 bl out of every 16 gametes. Fertilization between two such similar series of 16 gametes would result in 256 plants, of which 177 would be purple longs, 15 purple rounds, 15 red longs, and 49 red rounds—a proportion of the four different kinds very close to {95} that actually found by experiment. It will be noticed that in the whole family the purples are to the reds as 3 : 1, and the longs are also three times as numerous as the rounds. The peculiarity of the case lies in the distribution of these two characters with regard to one another. In some way or other the factors for blue and for long pollen become linked together in the cell divisions that give rise to the gametes, but the linking is not complete. This holds good for all the four cases in which repulsion between the factors occurs when one of the two factors is introduced by each of the parents. _When both of the factors are brought into the cross by the same parent we get coupling between them instead of repulsion._ The phenomena of repulsion and coupling between separate factors are intimately related, though hitherto we have not been able to suggest why this should be so.

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.

APPENDIX TO CHAPTER IX

As it is possible that some readers may care, in spite of its complexity, to enter rather more fully into the peculiar phenomenon {96} 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:—

49 BBLL + 7 BBLl + 7 BbLL + 49 BbLl + 7 BBLl + 7 BbLL + BbLl + BbLl + 49 BbLl ————————————————-/ 177 purple, long

+ BBll + 7 Bbll + bbLL + 7 bbLl + 49 bbll + 7 Bbll + 7 bbLl ——————-/ —————-/ ——-/ 15 purple, 15 red, 49 red, round long round

and as this theoretical result fits closely with the actual figures obtained by experiment we have reason for supposing that the heterozygous plant produces a series of gametes in which the factors are coupled in this way. The intensity of the coupling, however, varies in different cases. Where we are dealing with another, viz. fertility (F) and the dark axil (D), the experimental numbers accord with the view that the gametic series is here 15 FD : 1 Fd : 1 fD : 15 fd. The coupling is in this instance more intense. In the case of the erect standard (E) and blueness (B) the coupling is even more intense, and the experimental evidence available at present points to the gametic series here being 63 Eb : 1 EB : 1 eB : 63 eb. There is evidence also for supposing that the intensity of the coupling may vary in different families for the same pair of factors. The coupling between blue and long pollen is generally on the 7 : 1 : 1 : 7 {97} basis, but in some cases it may be on the 15 : 1 : 1 : 15 basis. But though the intensity of the coupling may vary it varies in an orderly way. If A and B are the two factors concerned, the results obtained in F_2 are explicable on the assumption that the ratio of the four sorts of gametes produced is a term of the series—

3 AB + Ab + aB + 3 ab 7 AB + Ab + aB + 7 ab 15 AB + Ab + aB + 15 ab, etc., etc.

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, {98} 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.

+ -+ + -+ -+ No. of Distribution of No. of Gametes Factors in Gametic Zygotes Form of F_2 Generation. in Series produced. series. + -+ + -+ -+ AB. Ab. aB. ab. AB. Ab. aB. ab. 4 1: 1: 1: 1 16 9 3 3 1 8 3: 1: 1: 3 64 49 7 7 9 16 7: 1: 1: 7 256 177 15 15 49* 32 15: 1: 1: 15 1024 737 31 31 225* 64 31: 1: 1: 31 4096 3009 63 63 961 128 63: 1: 1: 63 16384 12161 127 127 3969* 2n (n-1): 1: 1:(n-1) 4n^2 3n^2-(2n-1) 2n-1 2n-1 n^2-(2n-1) + -+ + -+ -+

Now, as the table shows, it is possible to express the gametic series by a general formula (n + 1) AB + Ab + aB + (n - 1) ab, where 2n is the total number of the gametes in the series. A plant producing such a series of gametes gives rise to a family of zygotes in which 3n^2 - (2n - 1) show both of the dominant characters and n^2 - (2n - 1) show both of the recessive characters, while the number of the two classes which each show one of the two dominants is (2n - 1). When in such a series the coupling becomes closer the value of n increases, but in comparison with n^2 its value becomes less and less. The larger n becomes the more negligible is its value relatively to n^2. If, therefore, the coupling were very close, the series 3n^2 - (2n - 1) : (2n - 1) : (2n - 1) : n^2 - (2n - 1) would approximate more and more to the series 3n^2 : n^2, i.e. to a simple 3 : 1 ratio. Though the point is probably of more theoretical than practical interest, it is not impossible that some of the cases which have hitherto been regarded as following a simple 3 : 1 ratio will turn out on further analysis to belong to this more complicated scheme.

* * * * *

{99}

CHAPTER X

SEX



In their simplest expression the phenomena exhibited by Mendelian characters are sharp and clean cut. Clean cut and sharp also are the phenomena of sex. It was natural, therefore, that a comparison should have been early instituted between these two sets of phenomena. As a general rule, the cross between a male and a female results in the production of the two sexes in approximately equal numbers. The cross between a heterozygous dominant and a recessive also leads to equal numbers of recessives and of heterozygous dominants. Is it not, therefore, possible that one of the sexes is heterozygous for a factor which is lacking in the other, and that the presence or absence of this factor determines the sex of the zygote? The results of some recent experiments would appear to justify this interpretation, at any rate in particular cases. Of these, the simplest is that of the common currant moth (Abraxas grossulariata), of which there exists a pale variety (Fig. 17) known as lacticolor. The experiments of Doncaster and Raynor showed that the variety behaved as a simple recessive to the normal form. But the distribution of the dominants and {100} recessives with with regard to the sexes was peculiar. The original cross was between a lacticolor female and a normal male. All the F1 moths of both sexes were of the normal grossulariata type. The F1 insects were then paired together and gave a generation consisting of 3 normals : 1 lacticolor. But all the lacticolor were females, and all the males were of the normal pattern. It was, however, found possible to obtain the lacticolor male by mating a lacticolor female with the F1 male. The family resulting from this cross consisted of normal males and normal females, lacticolor males and lacticolor females, and the {101} four sorts were produced in approximately equal numbers. In such a family there was no special association of either of the two colour varieties with one sex rather than the other. But the reverse cross, F1 female by lacticolor male, gave a very different result. As in the previous cross such families contained equal numbers of the normal form and of the recessive variety. But all of the normal grossulariata were males, while all the lacticolor were females. Now this seemingly complex collection of facts is readily explained if we make the following three assumptions:—



(1) The grossulariata character (G) is dominant to the lacticolor character (g). This is obviously justified by the experiments, for, leaving the sex distribution out of account, we get the expected 3 : 1 ratio from F1 x F1, and also the expected ratio of equality when the heterozygote is crossed with the recessive.

(2) The female is heterozygous for a dominant factor (F) which is lacking in the male. The constitution of a female is consequently Ff, and of a male ff. This assumption is in harmony with the fact that the sexes are produced in approximately equal numbers.

(3) There exists repulsion between the factors G and F in a zygote which is heterozygous for them both. Such zygotes (FfGg) must always be females, and on this assumption will produce gametes Fg and fG in equal numbers. {102}



We may now construct a scheme for comparison with that on page 100 to show how these assumptions explain the experimental results. The original parents were lacticolor female and grossulariata male, which on our assumptions must be Ffgg and ffGG respectively in constitution. Since the female is always heterozygous for F, her gametes must be of two kinds, viz. Fg and fg, while those of the pure grossulariata male must be all fG. When an ovum Fg is fertilised by a spermatozoon fG, the resulting zygote, FfGg, is heterozygous for both F and G, and in appearance is a female grossulariata. The zygote resulting from the fertilisation of an ovum fg by a spermatozoon fG is heterozygous for G, but does not contain F, and therefore is a male grossulariata. Such a male being in constitution {103} ffGg must produce gametes of two kinds, fG and fg, in equal numbers. And since we are assuming repulsion between F and G, the F1 female being in constitution FfGg, must produce equal numbers of gametes Fg and fG. For on our assumption F and G cannot enter into the same gamete. The series of gametes produced by the F1 moths, therefore, are fG, fg by the male and Fg, fG by the female. The resulting F2 generation consequently consists of the four classes of zygotes Ffgg, FfGg, ffGg, and ffGG in equal numbers. In other words, the sexes are produced in equal numbers, the proportion of normal grossulariata to lacticolor is 3 : 1, and all of the lacticolor are females; that is to say, the results worked out on our assumptions accord with those actually produced by experiment. We may now turn to the results which should be obtained by crossing the F1 moths with the lacticolor variety. And first we will take the cross lacticolor female x F1 male. The gametes produced by the lacticolor female we have already seen to be Fg and fg, while those produced by the F1 male are fG and fg. The bringing together of these two series of gametes must result in equal numbers of the four kinds of zygotes FfGg, Ffgg, ffGg, and ffgg, i.e. of female grossulariata and lacticolor, and of male grossulariata and lacticolor in equal numbers. Here, again, the calculated results accord with those of experiment. Lastly, we may examine what should happen when the F1 female is crossed with the lacticolor {104} male. The F1 female, owing to the repulsion between F and G, produces only the two kinds of ova Fg and fG, and produces them in equal numbers. Since the lacticolor male can contain neither F nor G, all of its spermatozoa must be fg. The results of such a cross, therefore, should be to produce equal numbers of the two kinds of zygote Ffgg and ffGg, i.e. of lacticolor females and of grossulariata males. And this, as we have already seen, is the actual result of such a cross.

Before leaving the currant moth we may allude to an interesting discovery which arose out of these experiments. The lacticolor variety in Great Britain is a southern form and is not known to occur in Scotland. Matings were made between wild Scotch females and lacticolor males. The families resulting from such matings were precisely the same as those from lacticolor males and F1 females, viz. grossulariata males and lacticolor females only. We are, therefore, forced to regard the constitution of the wild grossulariata female as identical with that of the F1 female, i.e. as heterozygous for the grossulariata factor as well as for the factor for femaleness. Though from a region where lacticolor is unknown, the "pure" wild grossulariata female is nevertheless a permanent mongrel, but it can never reveal its true colours unless it is mated with a male which is either heterozygous for G or pure lacticolor. And as all the wild northern males are {105} pure for the grossulariata character this can never happen in a state of nature.



An essential feature of the case of the currant moth lies in the different results given by reciprocal crosses. Lacticolor female x grossulariata male gives grossulariata alone of both sexes. But grossulariata female x lacticolor male gives only grossulariata males and lacticolor females. Such a difference between reciprocal crosses has also been found in other animals, and the experimental results, though sometimes more complicated, are explicable on the same lines. An interesting case in which three factors are concerned has been recently worked out in poultry. The Silky breed of fowls is characterised among other peculiarities by a remarkable abundance of melanic pigment. The skin is dull black, while the comb and wattles are of a deep purple colour contrasting sharply with the white plumage (Pl. V., 3). Dissection shows that this black pigment is widely spread throughout the body, being especially marked in such membranes as the mesenteries, the periosteum, and the pia mater surrounding the brain. It also occurs in the connective tissues among the muscles. In the Brown Leghorn, on the other hand, this pigment is not found. Reciprocal crosses between these two breeds gave a remarkable difference in result. A cross between the Silky hen and the Brown Leghorn cock produced F1 birds in which both sexes exhibited only traces of the pigment. On casual observation they might have {106} passed for unpigmented birds, for with the exception of an occasional fleck of pigment their skin, comb and wattles were as clear as in the Brown Leghorn (Pl. V., 1 and 4). Dissection revealed the presence of a slight amount of internal pigment. Such birds bred together gave some offspring with the full pigmentation of the Silky, some without any pigment, and others showing different degrees of pigment. None of the F2 male birds, however, showed the full deep pigmentation of the Silky.



When, however, the cross was made the other way, viz. Brown Leghorn hen x Silky cock, the result was different. While the F_1 male birds were almost destitute of pigment as in the previous cross, the F_1 hens, on the other hand, were nearly as deeply pigmented as the pure Silky {107} (Pl. V., 2). The male Silky transmitted the pigmentation, but only to his daughters. Such birds bred together gave an F_2 generation containing chicks with the full deep pigment, chicks without pigment, and chicks with various grades of pigmentation, all the different kinds in both sexes.



In analysing this complicated case many other different crosses were made, but for the present it will be sufficient to mention but one of these, viz. that between the F_1 birds and the pure Brown Leghorn. The cross between the F_1 hen and the Brown Leghorn cock produced only birds with a slight amount of pigment and birds without pigment. And this was true for both the deeply pigmented and the slightly pigmented types of F_1 hen. But when the F_1 cock was mated to a Brown Leghorn hen, a definite proportion of the chicks, one in eight, was deeply pigmented, and _these deeply pigmented birds were always females_ (cf. Fig. 21). And in this respect all the F_1 males behaved alike, whether they were from the Silky hen or from the Silky cock. We have, therefore, the paradox that the F_1 hen, though herself deeply pigmented, cannot transmit this condition to any of her offspring when she is mated to the unpigmented Brown Leghorn, but that, when similarly mated, the F_1 cock can transmit this pigmented condition to a quarter of his female offspring though he himself is almost devoid of pigment.



{108}



Now all these apparently complicated results, as well as many others to which we have not alluded, can be expressed by the following simple scheme. There are three factors affecting pigment, viz. (1) a pigmentation factor (P); (2) a factor which inhibits the production of pigment (I); and (3) a factor for femaleness (F), for which the female birds are heterozygous, but which is not present in the males. Further, we make the assumptions (a) that there is repulsion between F and I in the female zygote (FfIi), and (b) that the male Brown Leghorn is homozygous for the inhibitor factor (I), but that the hen Brown Leghorn is always heterozygous for this factor just in the same way as the female of the currant moth is always heterozygous for the grossulariata factor. We may now proceed to show how this explanation fits the experimental facts which we have given.

The Silky is pure for the pigmentation factor, but does not contain the inhibitor factor. The Brown Leghorn, on the other hand, contains the inhibitor factor, but not the {109} pigmentation factor. In crossing a Silky hen with a Brown Leghorn cock we are mating two birds of the constitution FfPPii and ffppII, and all the F_1 birds are consequently heterozygous for both P and I. In such birds the pigment is almost but not completely suppressed, and as both sexes are of the same constitution with regard to these two factors they are both of similar appearance.



In the reciprocal cross, on the other hand, we are mating a Silky male (ffPPii) with a Brown Leghorn hen which on our assumption is heterozygous for the inhibitor factor (I), and in constitution therefore is FfppIi. Owing to the repulsion between F and I the gametes produced by such a bird are Fpi and fpI in equal numbers. All the gametes produced by the Silky cock are fPi. Hence the constitution of the F_1 male birds produced by this cross is ffPpIi as before, but the female birds must be all of the constitution FfPpii. The Silky cock transmits the fully pigmented condition to his daughters, because the gametes of the Brown Leghorn hen which contain the factor for femaleness do not contain the {110} inhibitory factor owing to the repulsion between these factors. The nature of the F_2 generation in each case is in harmony with the above scheme. As, however, it serves to illustrate certain points in connection with intermediate forms we shall postpone further consideration of it till we discuss these matters, and for the present shall limit ourselves to the explanation of the different behaviour of the F_1 males and females when crossed with the Brown Leghorn. And, first, the cross of Brown Leghorn female by F_1 male. The Brown Leghorn hen is on our hypothesis FfppIi, and produces gametes Fpi and fpI. The F_1 cock is on our hypothesis ffPpIi, and produces in equal numbers the four kinds of gametes fPI, fPi, fpI, fpi. The result of the meeting of these two series of gametes is given in Fig. 24. Of the eight different kinds of zygote formed only one contains P in the absence of I, and this is a female. The result, as we have already seen, is in accordance with the experimental facts.



On the other hand, the Brown Leghorn cock is on our hypothesis ffppII. All his gametes consequently contain the inhibitor factor, and when he is mated with an F_1 {111} hen all the zygotes produced must contain I. None of his offspring, therefore, can be fully pigmented, for this condition only occurs in the absence of the inhibitor factor among zygotes which are either homozygous or heterozygous for P.



The interpretation of this case turns upon the constitution of the Brown Leghorn hen, upon her heterozygous condition with regard to the two factors F and I, and upon the repulsion that occurs between them when the gametes are formed. Through an independent set of experiments this view of the nature of the Brown Leghorn hen has been confirmed in an interesting way. There are fowls which possess neither the factor for pigment nor the inhibitory factor, which are in constitution ppii. Such birds when crossed with the Silky give dark pigmented birds of both sexes in F_1, and the F_2 generation consists of pigmented and unpigmented in the ratio 3 : 1. Now a cock of such a strain crossed with a Brown Leghorn hen should give only completely unpigmented birds. But if, as we have supposed, the Brown Leghorn hen is producing gametes Fpi and fpI, the male birds produced by such a cross should be heterozygous for I, {112} _i.e._ in constitution ffppIi, while the hen birds, though identical in appearance so far as absence of pigmentation goes, should not contain this factor but should be constitutionally Ffppii. Crossed with the pure Silky, the F_1 birds of opposite sexes should give an entirely different result. For while the hens should give only deeply pigmented birds of both sexes, the cocks should give equal numbers of deeply pigmented and slightly pigmented birds (cf. Fig. 25). These were the results which the experiment actually gave, thus affording strong confirmation of the view which we have been led to take of the Brown Leghorn hen. Essentially the poultry case is that of the currant moth. It differs in that the factor which {113} repels femaleness produces no visible effect, and its presence or absence can only be determined by the introduction of a third factor, that for pigmentation.

This conception of the nature of the Brown Leghorn hen leads to a curious paradox. We have stated that the Silky cock transmits the pigmented condition, but transmits it to his daughters only. Apparently the case is one of unequal transmission by the father. Actually, as our analysis has shown, it is one of unequal transmission by the mother, the father's contribution to the offspring being identical for each sex. The mother transmits to the daughters her dominant quality of femaleness, but to balance this, as it were, she transmits to her sons another quality which her daughters do not receive. It is a matter of common experience among human families that in respect to particular qualities the sons tend to resemble their mothers more than the daughters do, and it is not improbable that such observations have a real foundation for which the clue may be provided by the Brown Leghorn hen.

Nor is this the only reflection that the Brown Leghorn suggests. Owing to the repulsion between the factors for femaleness and for pigment inhibition, it is impossible by any form of mating to make a hen which is homozygous for the inhibitor factor. She has bartered away for femaleness the possibility of ever receiving a double dose of this factor. We know that in some cases, as, for example, {114} that of the blue Andalusian fowl, the qualities of the individual are markedly different according as to whether he or she has received a single or a double dose of a given factor. It is not inconceivable that some of the qualities in which a man differs from a woman are founded upon a distinction of this nature. Certain qualities of intellect, for example, may depend upon the existence in the individual of a double dose of some factor which is repelled by femaleness. If this is so, and if woman is bent upon achieving the results which such qualities of intellect imply, it is not education or training that will help her. Her problem is to get the factor on which the quality depends into an ovum that carries also the factor for femaleness.

* * * * *

{115}

CHAPTER XI

SEX (continued)

The cases which we have considered in the last chapter belong to a group in which the peculiarities of inheritance are most easily explained by supposing that the female is heterozygous for some factor that is not found in the male. Femaleness is an additional character superposed upon a basis of maleness, and as we imagine that there is a separate factor for each the full constitutional formula for a female is FfMM, and for a male ffMM. Both sexes are homozygous for the male element, and the difference between them is due to the presence or absence of the female element F.

There are, however, other cases for which the explanation will not suffice, but can be best interpreted on the view that the male is heterozygous for a factor which is not found in the female. Such a case is that recently described by Morgan in America for the pomace fly (Drosophila ampelophila). Normally this little insect has a red eye, but white eyed individuals are known to occur as rare sports. Red eye is dominant to white. In their relation to sex the eye colours of the pomace fly {116} are inherited on the same lines as the grossulariata and lacticolor patterns of the currant moth, but with one essential difference. The factor which repels the red-eye factor is in this case to be found in the male, and here consequently it is the male which must be regarded as heterozygous for a sex factor that is lacking in the female.



In order to bring these cases and others into line an interesting suggestion has recently been put forward by Bateson. On this suggestion each sex is heterozygous for its own sex factor only, and does not contain the factor proper to the opposite sex. The male is of the constitution, Mmff and the female Ffmm. Each sex produces two sorts of gametes, Mf and mf in the case of the male, and Fm, fm in that of the female. But on this view a further supposition is necessary. If each of the two kinds of spermatozoa were capable of fertilising each of the two kinds of ova, we should get individuals of the constitution MmFf and mmff, as well as the normal males and females, Mmff and Ffmm. As the facts of ordinary bisexual reproduction afford us no grounds for assuming the existence of these two classes of individuals, whatever they may be, we must suppose that fertilisation. is productive only between the spermatozoa carrying M and the ova without F, or between the spermatozoa {117} without M and the ova containing F. In other words we must on this view suppose that fertilisations between certain forms of gametes, even if they can occur, are incapable of giving rise to zygotes with the capacity for further development. If we admit this supposition, the scheme just given will cover such cases as those of the currant moth and the fowl, equally as well as that of the pomace fly. In the former there is repulsion between either the grossulariata factor and F, or else between the pigment inhibitor factor and F, while in the latter there is repulsion between the factor for red eye and M.



Whatever the merits or demerits of such a scheme it certainly does offer an explanation of a peculiar form of sex limited inheritance in man. It has long been a matter of common knowledge that colour-blindness is much more common among men than among women, and also that unaffected women can transmit it to their sons. At first sight the case is not unlike that of the sheep, where the horned character is apparently dominant in the male but recessive in the female. The hypothesis that the colour-blind condition is due to the presence of an extra factor as compared with the normal, and that a single dose of it will produce {118} colour-blindness in the male but not in the female, will cover a good many of the observed facts (cf. Fig. 26). Moreover, it serves to explain the remarkable fact that all the sons of colour-blind women are also colour-blind. For a woman cannot be colour-blind unless she is homozygous for the colour-blind factor, in which case all her children must get a single dose of it even if she marries a normal male. And this is sufficient to produce colour-blindness in the male, though not in the female.

But there is one notable difference in this case as compared with that of the sheep. When crossed with pure hornless ewes the heterozygous horned ram transmits the horned character to half his male offspring (cf. p. 71). But the heterozygous colour-blind man does not behave altogether like a sheep, for he apparently does not transmit the colour-blind condition to any of his male offspring. If, however, we suppose that the colour-blind factor is repelled by the factor for maleness, the amended scheme will cover the observed facts. For, denoting the colour-blind factor by X, the gametes produced by the colour-blind male are of two sorts only, viz. Mfx and mfX. If he marries a normal woman (Ffmmxx), the spermatozoa Mfx unite with ova fmx to give normal males, while the spermatozoa mfX unite with ova Fmx to give females which are heterozygous for the colour-blind factor. These daughters are themselves normal, but transmit the condition to about half their sons. {119}

The attempt to discover a simple explanation of the nature of sex has led us to assume that certain combinations between gametes are incapable of giving rise to zygotes which can develop further. In the various cases hitherto considered there is no reason to suppose that anything of the sort occurs, or that the different gametes are otherwise than completely fertile one with another. One peculiar case, however, has been known for several years in which some of the gametes are apparently incapable of uniting to produce offspring. Yellow in the mouse is dominant to agouti, but hitherto a homozygous yellow has never been met with. The yellows from families where only yellows and agoutis occur produce, when bred together, yellows and agoutis in the ratio 2 : 1. If it were an ordinary Mendelian case the ratio should be 3 : 1, and one out of every three yellows so bred should be homozygous and give only yellows when crossed with agouti. But Cuenot and others have shown that all of the yellows are heterozygous, and when crossed with agoutis give both yellows and agoutis. We are led, therefore, to suppose that an ovum carrying the yellow factor is unproductive if fertilised by a spermatozoon which also bears this factor. In this way alone does it seem possible to explain the deficiency of yellows and the absence of homozygous ones in the families arising from the mating of yellows together. At present, however, it remains the only definite instance among animals in which we have {120} grounds for assuming that anything in the nature of unproductive fertilisation takes place.[8]

If we turn from animals to plants we find a more complicated state of affairs. Generally speaking, the higher plants are hermaphrodite, both ovules and pollen grains occurring on the same flower. Some plants, however like most animals, are of separate sexes, a single plant bearing only male or female flowers. In other plants the separate flowers are either male or female, though both are borne on the same individual. In others, again, the conditions are even more complex, for the same plant may bear flowers of three kinds, viz. male, female, and hermaphrodite. Or it may be that these three forms occur in the same species but in different individuals—female and hermaphrodites in one species; males, females, and hermaphrodites in another. One case, however, must be mentioned as it suggests a possibility which we have not hitherto encountered. In the common English bryony (Bryonia dioica) the sexes are separate, some plants having only male and others only female flowers. In another European species, B. alba, both male and female flowers occur on the same plant. Correns crossed these two species reciprocally, and also fertilised B. dioica by its own male with the following results:—

{121}

dioica [female] x dioica [male] gave [female] [female] and [male] [male] " x alba [male] " [female] [female] only alba [female] x dioica [male] " [female] [female] and [male] [male].

The point of chief interest lies in the striking difference shown by the reciprocal crosses between _dioica_ and _alba_. Males appear when _alba_ is used as the female parent but not when the female _dioica_ is crossed by male _alba_. It is possible to suggest more than one scheme to cover these facts, but we may confine ourselves here to that which seems most in accord with the general trend of other cases. We will suppose that in _dioica_ femaleness is dominant to maleness, and that the female is heterozygous for this additional factor. In this species, then, the female produces equal numbers of ovules with and without the female factor, while this factor is absent in all the pollen grains. _Alba_ [female] x _dioica_ [male] gives the same result as _dioica_ [female] x _dioica_ [male], and we must therefore suppose that alba produces male and female ovules in equal numbers. _Alba_ [male] x _dioica_ [female], however, gives nothing but females. Unless, therefore, we assume that there is selective fertilisation we must suppose that all the pollen grains of alba carry the female factor—in other words, that so far as the sex factors are concerned there is a difference between the ovules and pollen grains borne by the same plant. Unfortunately further investigation of this case is rendered impossible owing to the complete sterility of the F_1 plants. {122}



That the possibility of a difference between the ovules and pollen grains of the same individual must be taken into account in future work there is evidence from quite a different source. The double stock is an old horticultural favourite, and for centuries it has been known that of itself it sets no seed, but must be raised from special strains of the single variety. "You must understand withall," wrote John Parkinson of his gilloflowers,[9] "that those plants that beare double flowers, doe beare no seed at all ... but the onely way to have double flowers any yeare is to save the seedes of those plants of this kinde that beare single flowers, for from that seede will rise some that will beare single, and some double flowers." With regard to the nature of these double-throwing strains of singles, Miss Saunders has recently brought out some interesting facts. She crossed the double-throwing singles with pure singles belonging to strains in which doubles never occur. The cross was made both ways, and in both cases all the F1 plants were single. A distinction, however, appeared when a further generation was raised from the F1 plants. All the F1 plants from the pollen of the double-throwing single behaved like double-throwing singles, but of the F1 plants from the ovules of the double throwers some behaved as double throwers, and some as pure singles. We are led to infer, therefore, that the ovules and pollen grains {123} of the double throwers, though both produced by the same plant, differ in their relation to the factor (or factors) for doubleness. Doubleness is apparently carried by all the pollen grains of such plants, but only by some of the ovules. Though the nature of doubleness in stocks is not yet clearly understood, the facts discovered by Miss Saunders suggest strongly that the ovules and pollen grains of the same plant may differ in their transmitting properties, probably owing to some process of segregation in the growing plant which leads to an unequal distribution of some or other factors to the cells which give rise to the ovules as compared with those from which {124} the pollen grains eventually spring. Whether this may turn out to be the true account or not, the possibility must not be overlooked in future work.

Single - Single Double / Pollen of x Ovule Pollen x Ovule of pure single pure single Single Single Single - - Single Single Double Single Double - - Single Single Double Single Double

From all this it is clear enough that there is much to be done before the problem of sex is solved even so far as the biologist can ever expect to solve it. The possibilities are many, and many a fresh set of facts is needed before we can hope to decide among them. Yet the occasional glimpses of clear-cut and orderly phenomena, which Mendelian spectacles have already enabled us to catch, offer a fair hope that some day they may all be brought into focus, and assigned their proper places in a general scheme which shall embrace them all. Then, though not till then, will the problem of the nature of sex pass from the hands of the biologist into those of the physicist and the chemist.

* * * * *

{125}

CHAPTER XII

INTERMEDIATES

So far as we have gone we have found it possible to express the various characters of animals and plants in terms of definite factors which are carried by the gametes, and are distributed according to a definite scheme. Whatever may be the nature of these factors it is possible for purposes of analysis to treat them as indivisible entities which may or may not be present in any given gamete. When the factor is present it is present as a whole. The visible properties developed by a zygote in the course of its growth depend upon the nature and variety of the factors carried in by the two gametes which went to its making, and to a less degree upon whether each factor was brought in by both gametes or by one only. If the given factor is brought in by one gamete only, the resulting heterozygote may be more or less intermediate between the homozygous form with a double dose of the factor and the homozygous form which is entirely destitute of the factor. Cases in point are those of the primula flowers and the Andalusian fowls. Nevertheless these intermediates produce only pure gametes, as is {126} shown by the fact that the pure parental types appear in a certain proportion of their offspring. In such cases as these there is but a single type of intermediate, and the simple ratio in which this and the two homozygous forms appear renders the interpretation obvious. But the nature of the F2 generation may be much more complex, and, where we are dealing with factors which interact upon one another, may even present the appearance of a series of intermediate forms grading from the condition found in one of the original parents to that which occurred in the other. As an illustration we may consider the cross between the Brown Leghorn and Silky fowls which we have already dealt with in connection with the inheritance of sex. The offspring of a Silky hen mated with a Brown Leghorn are in both sexes birds with but a trace of the Silky pigmentation. But when such birds are bred together they produce a generation consisting of chicks as deeply pigmented as the original Silky parent, chicks devoid of pigment like the Brown Leghorn, and chicks in which the pigmentation shows itself in a variety of intermediate stages. Indeed from a hundred chicks bred in this way it would be possible to pick out a number of individuals and arrange them in an apparently continuous series of gradually increasing pigmentation, with the completely unpigmented at one end and the most deeply pigmented at the other. Nevertheless, the case is one in which complete segregation of the different factors takes {127} place, place, and the apparently continuous series of intermediates is the result of the interaction of the different factors upon one another. The constitution of the F1 [male] is a ffPpIi, and such a bird produces in equal numbers the four sorts of gametes fPI, fPi, fpI, fpi. The constitution of the F1 [female] in this case is FfPpIi. Owing to the repulsion between F and I she produces the four kinds of gametes FPi, Fpi, fPI, fpi, and produces them in equal numbers. The result of bringing two such series of gametes together is shown in Fig. 28. Out of the sixteen types of zygote formed one (FfPPii) is homozygous for the pigmentation factor, and does not contain the inhibitor factor. Such a bird is as deeply pigmented as the pure Silky parent. Two, again, contain a single dose of P in the absence of I. These are nearly as dark as the pure Silky. Four zygotes are destitute of P, though they may or may not contain I. These birds are completely devoid of pigment like the Brown Leghorn. The remaining nine zygotes show {128} various combinations of the two factors P and I, being either PPIi, PPII, PpII, or PpIi, and in each of these cases the pigment is more or less intense according to the constitution of the bird. Thus a bird of the constitution PPIi approaches in pigmentation a bird of the constitution Ppii, while a bird of the constitution PpII has but little more pigment than the unpigmented bird. In this way we have seven distinct grades of pigmentation, and the series is further complicated by the fact that these various grades exhibit a rather different amount of pigmentation according as they occur in a male or a female bird, for, generally speaking, the female of a given grade exhibits rather more pigment than the corresponding male. The examination of a number of birds bred in this way might quite well suggest that in this case we were dealing with a character which could break up, as it were, to give a continuous series of intergrading forms between the two extremes. With the constant handling of large numbers it becomes possible to recognise most of the different grades, though even so it is possible to make mistakes. Nevertheless, as breeding tests have amply shown, we are dealing with but two interacting factors which segregate cleanly from one another according to the strict Mendelian rule. The approach to continuity in variation exhibited by the F2 generation depends upon the fact that these two factors interact upon one another, and to different degrees according as the zygote is for one {129} or other or both of them in a homozygous or a heterozygous state. Moreover, certain of these intermediates will breed true to an intermediate condition of the pigmentation. A male of the constitution ffPPII when bred with females of the constitution FfPPIi will produce only males like itself and females like the maternal parent. We have dealt with this case in some detail, because the existence of families showing a series of intermediate stages between two characters has sometimes been brought forward in opposition to the view that the characters of organisms depend upon specific factors which are transmitted according to the Mendelian rule. But, as this case from poultry shows clearly, neither the existence of such a continuous series of intermediates, nor the fact that some of them may breed true to the intermediate condition, are incompatible with the Mendelian principle of segregation.



In connection with intermediates a more cogent objection to the Mendelian view is the case of the first cross between two definite varieties thenceforward breeding true. The case that will naturally occur to the reader is that of the mulatto, which results from the cross between the negro and the white. According to general opinion, these mulattos, of intermediate pigmentation, continue to produce mulattos. Unfortunately this interesting case has never been critically investigated, and the statement that the mulatto breeds true rests almost entirely upon {130} information that is general and often vague. It may be that the inheritance of skin pigmentation in this instance is a genuine exception to the normal rule, but at the same time it must not be forgotten that it may be one in which several interacting factors are concerned, and that the pure white and the pure black are the result of combinations which from their rarity are apt to be overlooked. But until we are in possession of accurate information it is impossible to pronounce definitely upon the nature of the inheritance in this case.



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On the other hand, from the cross between the darkly pigmented Eastern races and the white segregation seems to occur in subsequent generations. Families are to be found in which one parent is a pure white, while the other has arisen from the cross between the dark and light in the first or some subsequent generation. Such families may contain children indistinguishable from pure blonds as well as children of very dark and of intermediate shades. As an example, I may give the following pedigree, which was kindly communicated to me by an Anglo-Indian friend (Fig. 29). The family had resided in England for several generations, so that in this case there was no question of a further admixture of black. Most noticeable is the family produced by a very dark lady who had married a white man. Some of the children were intermediate in colour, but two were fair whites and two were dark as dark Hindus. This sharp segregation or splitting out of blacks and whites in addition to intermediates strongly suggests that the nature of the inheritance is Mendelian, though it may be complicated by the existence of several factors which may also react upon one another. Nor must it be forgotten that in so far as these different factors are concerned the whites themselves may differ in constitution without showing any trace of it in their appearance. Before the case can be regarded as settled all these different possibilities will have to be definitely tested. With the dark Eastern races as with the negro we cannot {132} hope to come to any conclusion until we have evidence collected by critical and competent observers.

Though for the present we must regard the case of the negro as not proven, there are nevertheless two others in which the heredity would appear not to follow the Mendelian rule. Castle in America crossed the lop-eared rabbit with the normal form, and found that the F_1 animals were intermediate with respect to their ears. And subsequent experiment showed that, on the whole, they bred true to this intermediate condition. The other case relates to Lepidoptera. The speckled wood butterfly (_Pararge egeria_) has a southern form which differs from the northern one in the greater brightness and depth of its yellow-brown markings. The northern form is generally distinguished as var. _egeriades_. Bateson crossed the southern form from the south of France with the paler British form, and found that the offspring were more or less intermediate in colour, and that in subsequent generations the parental types did not recur. These cases at present stand alone. It is possible that further research may reveal complications which mask or interfere with an underlying process of segregation. Or it may be that segregation does not occur owing to some definite physiological reason which at present we do not understand.

And here it is impossible not to recall Mendel's own experiences with the Hawkweeds (Hieracium). This {133} genus of plants exhibits an extraordinary profusion of forms differing from one another sometimes in a single feature, sometimes in several. The question as to how far these numerous forms were to be classified as distinct species, how far as varieties, and how far as products of chance hybridisation, was even at that time a source of keen controversy among botanists. There is little doubt that Mendel undertook his experiments on the Hawkweeds in the hope that the conception of unit-characters so brilliantly demonstrated for the pea would serve to explain the great profusion of forms among the Hieraciums. Owing to the minute size of their florets, these plants offer very considerable technical difficulties in the way of cross fertilisation. By dint of great perseverance and labour, however, Mendel succeeded in obtaining a few crosses between different forms. These hybrids were reared and a further generation produced from them, and, no doubt somewhat to Mendel's chagrin, every one of them proved to breed true. There was a complete absence of that segregation of characters which he had shown to exist in peas and beans, and had probably looked forward with some confidence to finding in Hieracium. More than thirty years passed before the matter was cleared up. To-day we know that the peculiar behaviour of the hybrid Hieraciums is due to the fact that they normally produce seed by a peculiar process of parthenogenesis. It is possible to take an unopened flower and to shear off with a {134} razor all the male organs together with the stigmata through which the pollen reaches the ovules. The flower, nevertheless, sets perfectly good seed. But the cells from which the seeds develop are not of the same nature as the normal ovules of a plant. They are not gametes but retain the double structure of the maternal cells. They are rather to be regarded as of the nature of buds which early become detached from the parent stock to lead an independent existence, and, like buds, they reproduce exactly the maternal characteristics. The discovery of the true nature of this case was only rendered possible by the development of the study of cytology, and it was not given to Mendel to live long enough to learn why his hybrid Hieraciums all bred true.

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CHAPTER XIII

VARIATION AND EVOLUTION

Through the facts of heredity we have reached a new conception of the individual. Hitherto we have been accustomed to distinguish between the members of a family of rabbits like that illustrated on Plate I. by assigning to each an individuality, and by making use of certain external features, such as the coat colour or the markings, as convenient outward signs to express our idea that the individuality of these different animals is different. Apart from this, our notions as to what constituted the individuality in each case were at best but vague. Mendelian analysis has placed in our hands a more precise method of estimating and expressing the variations that are to be found between one individual and another. Instead of looking at the individual as a whole, which is in some vague way endowed with an individuality marking it off from its fellows, we now regard it as an organism built up of definite characters superimposed on a basis beyond which for the moment our analysis will not take us. We have begun to realise that each individual has a definite architecture, and that this architecture depends {136} primarily upon the number and variety of the factors that existed in the two gametes that went to its building. Now most species exhibit considerable variation and exist in a number, often very large, of more or less well-defined varieties. How far can this great variety be explained in terms of a comparatively small number of factors if the number of possible forms depends upon the number of the factors which may be present or absent?

In the simple case where the homozygous and heterozygous conditions are indistinguishable in appearance the number of possible forms is 2, raised to the power of the number of factors concerned. Thus where one factor is concerned there are only 2^1 = 2 possible forms, where ten factors are concerned there are 2^{10} = 1024 possible forms differing from one another in at most ten and at least one character. Where the factors interact upon one another this number will, of course, be considerably increased. If the heterozygous form is different in appearance from the homozygous form, there are three possible forms connected with each factor; for ten such factors the possible number of individuals would be 3^{10} = 59,049; for twenty such factors the possible number of different individuals would be 3^{20} = 3,486,784,401. The presence or absence of a comparatively small number of factors in a species carries with it the possibility of an enormous range of individual variation. But every one of these individuals has a perfectly definite constitution which can {137} be determined in each case by the ordinary methods of Mendelian analysis. For in every instance the variation depends upon the presence or absence of definite factors carried in by the gametes from whose union the individual results. And as these factors separate out cleanly in the gametes which the individual forms, such variations as depend upon them are transmitted strictly according to the Mendelian scheme. Provided that the constitution of the gametes is unchanged, the heredity of such variation is independent of any change in the conditions of nutrition or environment which may operate upon the individual producing the gametes.

But, as everybody knows, an individual organism, whether plant or animal, reacts, and often reacts markedly, to the environmental conditions under which its life is passed. More especially is this to be seen where such characters as size or weight are concerned. More sunlight or a richer soil may mean stronger growth in a plant, better nutrition may result in a finer animal, superior education may lead to a more intelligent man. But although the changed conditions produce a direct effect upon the individual, we have no indisputable evidence that such alterations are connected with alterations in the nature of the gametes which the individual produces. And without this such variations cannot be perpetuated through heredity, but the conditions which produce the effect must always be renewed in each {138} successive generation. We are led, therefore, to the conclusion that two sorts of variations exist, those which are due to the presence of specific factors in the organism and those which are due to the direct effect of the environment during its lifetime. The former are known as mutations, and are inherited according to the Mendelian scheme; the latter have been termed fluctuations, and at present we have no valid reason for supposing that they are ever inherited. For though instances may be found in which effects produced during the lifetime of the individual would appear to affect the offspring, this is not necessarily due to heredity. Thus plants which are poorly nourished and grown under adverse conditions may set seed from which come plants that are smaller than the normal although grown under most favorable conditions. It is natural to attribute the smaller size of the offspring to the conditions under which the parents were grown, and there is no doubt that we should be quite right in doing so. Nevertheless, it need have nothing to do with heredity. As we have already pointed out, the seed is a larval plant which draws its nourishment from the mother. The size of the offspring is affected because the poorly nourished parent offered a bad environment to the young plant, and not because the gametes of the parent were changed through the adverse conditions under which it grew. The parent in this case is not only the producer of gametes, but also a part of the environment of the young {139} plant, and it is in this latter capacity that it affects its offspring. Wherever, as in plants and mammals, the organism is parasitic upon the mother during its earlier stages, the state of nutrition of the latter will almost certainly react upon it, and in this way a semblance of transmitted weakness or vigour is brought about. Such a connection between mother and offspring is purely one of environment, and it cannot be too strongly emphasised that it has nothing to do with the ordinary process of heredity.

The distinction between these two kinds of variation, so entirely different in their causation, renders it possible to obtain a clearer view of the process of evolution than that recently prevalent. As Darwin long ago realised, any theory of evolution must be based upon the facts of heredity and variation. Evolution only comes about through the survival of certain variations and the elimination of others. But to be of any moment in evolutionary change a variation must be inherited. And to be inherited it must be represented in the gametes. This, as we have seen, is the case for those variations which we have termed mutations. For the inheritance of fluctuations, on the other hand, of the variations which result from the direct action of the environment upon the individual, there is no indisputable evidence. Consequently we have no reason for regarding them as playing any part in the production of that succession of temporarily stable forms which we term evolution. In {140} the light of our present knowledge we must regard the mutation as the basis of evolution—as the material upon which natural selection works. For it is the only form of variation of whose heredity we have any certain knowledge.

It is evident that this view of the process of evolution is in some respects at variance with that generally held during the past half century. There we were given the conception of an abstract type representing the species, and from it most of the individuals diverged in various directions, though, generally speaking, only to a very small extent. It was assumed that any variation, however small, might have a selection value, that is to say, could be transmitted to the offspring. Some of these would possess it in a less and some in a greater degree than the parent. If the variation were a useful one, those possessing to a rather greater extent would be favoured through the action of natural selection at the expense of their less fortunate brethren, and would leave a greater number of offspring, of whom some possessed it in an even more marked degree than themselves. And so it would go on. The process was a cumulative one. The slightest variation in a favourable direction gave natural selection a starting-point to work on. Through the continued action of natural selection on each successive generation the useful variation was gradually worked up, until at last it reached the magnitude of a specific {141} distinction. Were it possible in such a case to have all the forms before us, they would present the appearance of a long series imperceptibly grading from one extreme to the other.

Upon this view are made two assumptions not unnatural in the absence of any exact knowledge of the nature of heredity and variation. It was assumed, in the first place that variation was a continuous process, and, second, that any variation could be transmitted to the offspring. Both of these assumptions have since been shown to be unjustified. Even before Mendel's work became known Bateson had begun to call attention to the prevalence of discontinuity in variation, and a few years later this was emphasised by the Dutch botanist Hugo de Vries in his great work on The Mutation Theory. The ferment of new ideas was already working in the solution, and under the stimulus of Mendel's work they have rapidly crystallised out. With the advent of heredity as a definite science we have been led to revise our views as to the nature of variation, and consequently in some respects as to the trend of evolution. Heritable variation has a definite basis in the gamete, and it is to the gamete, therefore, not to the individual, that we must look for the initiation of this process. Somewhere or other in the course of their production is added or removed the factor upon whose removal or addition the new variation owes its existence. The new variation springs into being by a {142} sudden step, not by a process of gradual and almost imperceptible augmentation. It is not continuous but discontinuous, because it is based upon the presence or absence of some definite factor or factors—upon discontinuity in the gametes from which it sprang. Once formed, its continued existence is subject to the arbitrament of natural selection. If of value in the struggle for existence natural selection will decide that those who possess it shall have a better chance of survival and of leaving offspring than those who do not possess it. If it is harmful to the individual natural selection will soon bring about its elimination. But if the new variation is neither harmful nor useful there seems no reason why it should not persist.

In this way we avoid a difficulty that beset the older view. For on that view no new character could be developed except by the piling up of minute variations through the action of natural selection. Consequently any character found in animals and plants must be supposed to be of some definite use to the individual. Otherwise it could not have developed through the action of natural selection. But there are plenty of characters to which it is exceedingly difficult to ascribe any utility, and the ingenuity of the supporters of this view has often been severely taxed to account for their existence. On the more modern view this difficulty is avoided. The origin of a new variation is independent of natural {143} selection, and provided that it is not directly harmful there is no reason why it should not persist. In this way we are released from the burden of discovering a utilitarian motive behind all the multitudinous characters of living organisms. For we now recognise that the function of natural selection is selection and not creation. It has nothing to do with the formation of the new variation. It merely decides whether it is to survive or to be eliminated.

One of the arguments made use of by supporters of the older view is that drawn from the study of adaptation. Animals and plants are as a rule remarkably well adapted to living the life which their surroundings impose upon them, and in some cases this adaptation is exceedingly striking. Especially is this so in the many instances of what is called protective coloration, where the animal comes to resemble its surroundings so closely that it may reasonably be supposed to cheat even the keenest sighted enemy. Surely, we are told, such perfect adaptation could hardly have arisen through the mere survival of chance sports. Surely there must be some guiding hand moulding the species into the required shape. The argument is an old one. For John Ray that guiding hand was the superior wisdom of the Creator: for the modern Darwinian it is Natural Selection controlling the direction of variation. Mendelism certainly offers no suggestion of any such controlling force. It interprets the {144} variations of living forms in terms of definite physiological factors, and the diversity of animal and plant life is due to the gain or loss of these factors, to the origination of new ones, or to fresh combinations among those already in existence. Nor is there any valid reason against the supposition that even the most remarkable cases of resemblance, such as that of the leaf insect, may have arisen through a process of mutation. Experience with domestic plants and animals shows that the most bizarre forms may arise as sports and perpetuate themselves. Were such forms, arising under natural conditions, to be favoured by natural selection owing to a resemblance to something in their environment we should obtain a striking case of protective adaptation. And here it must not be forgotten that those striking cases to which our attention is generally called are but a very small minority of the existing forms of life.

For that special group of adaptation phenomena classed under the head of Mimicry, Mendelism seems to offer an interpretation simpler than that at present in vogue. This perhaps may be more clearly expressed by taking a specific case. There is in Africa a genus of Danaine butterflies known as Amauris, and there are reasons for considering that the group to which it belongs possesses properties which render it unpalatable to vertebrate enemies such as birds or monkeys. In the same region is also found the genus Euralia belonging to the entirely {145} different family of the Nymphalidae, to which there is no evidence for assigning the disagreeable properties of the Danaines. Now the different species of Euralia show remarkably close resemblances to the species of Amauris, which are found flying in the same region, and it is supposed that by "mimicking" the unpalatable forms they impose upon their enemies and thereby acquire immunity from attack. The point at issue is the way in which this seemingly purposeful resemblance has been brought about.

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