SigPhi · Charles Darwin

The Variation of Animals and Plants under Domestication

Page 38 of 72

produced offspring before they had acquired with advancing age their new characters, nothing is more probable than that they would have transmitted them to some of their offspring, who in this case would in appearance have received such characters from their grand-parents or more distant progenitors. We should then have had a case of reversion, that is, of the reappearance in the child of an ancestral character, actually present, though during youth completely latent, in the parent; and this we may safely conclude is what occurs in all reversions to progenitors, however remote.

This view of the latency in each generation of all the characters which appear through reversion, is also supported by their actual presence in some cases during early youth alone, or by their more frequent appearance and greater distinctness at this age than during maturity.

We have seen that this is often the case with the stripes on the legs and faces of the several species of the horse genus. The Himalayan rabbit, when crossed, sometimes produces offspring which revert to the parent silver-grey breed, and we have seen that in purely bred animals pale-grey fur occasionally reappears during early youth. Black cats, we may feel assured, would occasionally produce by reversion tabbies; and on young black kittens, with a pedigree[66] known to have been long pure, faint traces of stripes may almost always be seen which afterwards disappear. Hornless Suffolk cattle occasionally produce by reversion horned animals; and Youatt[67] asserts that even in hornless individuals “the rudiment of a horn may be often felt at an early age.”

No doubt it appears at first sight in the highest degree improbable that in every horse of every generation there should be a latent capacity and tendency to produce stripes, though these may not appear once in a thousand generations; that in every white, black, or other coloured pigeon, which may have transmitted its proper colour during centuries, there should be a latent capacity in the plumage to become blue and to be marked with certain characteristic bars; that in every child in a six-fingered family there should be the capacity for the production of an additional digit; and so in other cases. Nevertheless, there is no more inherent improbability in this being the case than in a useless and rudimentary organ, or even in only a tendency to the production of a rudimentary organ, being inherited during millions of generations, as is well known to occur with a multitude of organic beings. There is no more inherent improbability in each domestic pig, during a thousand generations, retaining the capacity and tendency to develop great tusks under fitting conditions, than in the young calf having retained, for an indefinite number of generations rudimentary incisor teeth, which never protrude through the gums.

I shall give at the end of the next chapter a summary of the three preceding chapters; but as isolated and striking cases of reversion have here been chiefly insisted on, I wish to guard the reader against supposing that reversion is due to some rare or accidental combination of circumstances. When a character, lost during hundreds of generations, suddenly reappears, no doubt some such combination must occur; but reversions, to the immediately preceding generations may be constantly observed, at least, in the offspring of most unions. This has been universally recognised in the case of hybrids and mongrels, but it has been recognised simply from the difference between the united forms rendering the resemblance of the offspring to their grandparents or more remote progenitors of easy detection. Reversion is likewise almost invariably the rule, as Mr. Sedgwick has shown, with certain diseases. Hence we must conclude that a tendency to this peculiar form of transmission is an integral part of the general law of inheritance.

_Monstrosities._—A large number of monstrous growths and of lesser anomalies are admitted by every one to be due to an arrest of development, that is, to the persistence of an embryonic condition. But many monstrosities cannot be thus explained; for parts of which no trace can be detected in the embryo, but which occur in other members of the same class of animals occasionally appear, and these may probably with truth be attributed to reversion. As, however, I have treated this subject as fully as I could in my ‘Descent of Man’ (ch. 1 2nd edition), I will not here recur to it.

When flowers which have normally an irregular structure become regular or peloric, the change is generally looked at by botanists as a return to the primitive state. But Dr. Maxwell Masters,[68] who has ably discussed this subject, remarks that when, for instance, all the sepals of a Tropæolum become green and of the same shape, instead of being coloured with one prolonged into a spur, or when all the petals of a Linaria become simple and regular, such cases may be due merely to an arrest of development; for in these flowers all the organs during their earliest condition are symmetrical, and, if arrested at this stage of growth, they would not become irregular. If, moreover, the arrest were to take place at a still earlier period of development, the result would be a simple tuft of green leaves; and no one probably would call this a case of reversion. Dr. Masters designates the cases first alluded to as regular peloria; and others, in which all the corresponding parts assume a similar form of irregularity, as when all the petals in a Linaria become spurred, as irregular peloria. We have no right to attribute these latter cases to reversion, until it can be shown that the parent-form, for instance, of the genus Linaria had had all its petals spurred; for a chance of this nature might result from the spreading of an anomalous structure, in accordance with the law, to be discussed in a future chapter, of homologous parts tending to vary in the same manner. But as both forms of peloria frequently occur on the same individual plant of the Linaria,[69] they probably stand in some close relation to one another. On the doctrine that peloria is simply the result of an arrest of development, it is difficult to understand how an organ arrested at a very early period of growth should acquire its full functional perfection;—how a petal, supposed to be thus arrested, should acquire its brilliant colours, and serve as an envelope to the flower, or a stamen produce efficient pollen; yet this occurs with many peloric flowers. That pelorism is not due to mere chance variability, but either to an arrest of development or to reversion, we may infer from an observation made by Ch. Morren[70] namely, that families which have irregular flowers often “return by these monstrous growths to their regular form; whilst we never see a regular flower realise the structure of an irregular one.”

Some flowers have almost certainly become more or less completely peloric through reversion, as the following interesting case shows.

_Corydalis tuberosa_ properly has one of its two nectaries colourless, destitute of nectar, only half the size of the other, and therefore, to a certain extent, in a rudimentary state; the pistil is curved towards the perfect nectary, and the hood, formed of the inner petals, slips off the pistil and stamen in one direction alone, so that, when a bee sucks the perfect nectary, the stigma and stamens are exposed and rubbed against the insect’s body. In several closely allied genera, as in Dielytra, etc., there are two perfect nectaries, the pistil is straight, and the hood slips off on either side, according as the bee sucks either nectary. Now, I have examined several flowers of _Corydalis tuberosa,_ in which both nectaries were equally developed and contained nectar; in this we see only the redevelopment of a partially aborted organ; but with this redevelopment the pistil becomes straight, and the hood slips off in either direction, so that these flowers have acquired the perfect structure, so well adapted for insect agency, of Dielytra and its allies. We cannot attribute these coadapted modifications to chance, or to correlated variability; we must attribute them to reversion to a primordial condition of the species.

The peloric flowers of Pelargonium have their five petals in all respects alike, and there is no nectary so that they resemble the symmetrical flowers of the closely allied genus Geranium; but the alternate stamens are also sometimes destitute of anthers, the shortened filaments being left as rudiments, and in this respect they resemble the symmetrical flowers of the closely allied genus Erodium.

Hence we may look at the peloric flowers of Pelargonium as having reverted to the state of some primordial form, the progenitor of the three closely related genera of Pelargonium, Geranium, and Erodium.

In the peloric form of _Antirrhinum majus,_ appropriately called the “_Wonder,_” the tubular and elongated flowers differ wonderfully from those of the common snapdragon; the calyx and the mouth of the corolla consist of six equal lobes, and include six equal instead of four unequal stamens. One of the two additional stamens is manifestly formed by the development of a microscopically minute papilla, which may be found at the base of the upper lip of the flower of the common snapdragons in the nineteen plants examined by me. That this papilla is a rudiment of a stamen was well shown by its various degrees of development in crossed plants between the common and the peloric Antirrhinum. Again, a peloric _Galeobdolon luteum,_ growing in my garden, had five equal petals, all striped like the ordinary lower lip, and included five equal instead of four unequal stamens; but Mr. R.

Keeley, who sent me this plant, informs me that the flowers vary greatly, having from four to six lobes to the corolla, and from three to six stamens.[71] Now, as the members of the two great families to which the Antirrhinum and Galeobdolon belong are properly pentamerous, with some of the parts confluent and others suppressed, we ought not to look at the sixth stamen and the sixth lobe to the corolla in either case as due to reversion, any more than the additional petals in double flowers in these same two families. But the case is different with the fifth stamen in the peloric Antirrhinum, which is produced by the redevelopment of a rudiment always present, and which probably reveals to us the state of the flower, as far as the stamens are concerned, at some ancient epoch. It is also difficult to believe that the other four stamens and the petals, after an arrest of development at a very early embryonic age, would have come to full perfection in colour, structure, and function, unless these organs had at some former period normally passed through a similar course of growth. Hence it appears to me probable that the progenitor of the genus Antirrhinum must at some remote epoch have included five stamens and borne flowers in some degree resembling those now produced by the peloric form. The conclusion that peloria is not a mere monstrosity, irrespective of any former state of the species, is supported by the fact that this structure is often strongly inherited, as in the case of the peloric Antirrhinum and Gloxinia and sometimes in that of the peloric Lastly I may add that many instances have been recorded of flowers, not generally considered as peloric, in which certain organs are abnormally augmented in number. As an increase of parts cannot be looked at as an arrest of development, nor as due to the redevelopment of rudiments, for no rudiments are present, and as these additional parts bring the plant into closer relationship with its natural allies, they ought probably to be viewed as reversions to a primordial condition.

These several facts show us in an interesting manner how intimately certain abnormal states are connected together; namely, arrests of development causing parts to become rudimentary or to be wholly suppressed,—the redevelopment of parts now in a more or less rudimentary condition,—the reappearance of organs of which not a vestige can be detected,—and to these may be added, in the case of animals, the presence during youth, and subsequent disappearance, of certain characters which occasionally are retained throughout life.

Some naturalists look at all such abnormal structures as a return to the ideal state of the group to which the affected being belongs; but it is difficult to conceive what is meant to be conveyed by this expression. Other naturalists maintain, with greater probability and distinctness of view, that the common bond of connection between the several foregoing cases is an actual, though partial, return to the structure of the ancient progenitor of the group. If this view be correct, we must believe that a vast number of characters, capable of evolution, lie hidden in every organic being. But it would be a mistake to suppose that the number is equally great in all beings. We know, for instance, that plants of many orders occasionally become peloric; but many more cases have been observed in the Labiatæ and Scrophulariaceæ than in any other order; and in one genus of the Scrophulariaceæ, namely Linaria, no less than thirteen species have been described in this condition.[73] On this view of the nature of peloric flowers, and bearing in mind certain monstrosities in the animal kingdom, we must conclude that the progenitors of most plants and animals have left an impression, capable of redevelopment, on the germs of their descendants, although these have since been profoundly modified.

The fertilised germ of one of the higher animals, subjected as it is to so vast a series of changes from the germinal cell to old age,—incessantly agitated by what Quatrefages well calls the _tourbillon vital,_—is perhaps the most wonderful object in nature. It is probable that hardly a change of any kind affects either parent, without some mark being left on the germ. But on the doctrine of reversion, as given in this chapter, the germ becomes a far more marvellous object, for, besides the visible changes which it undergoes, we must believe that it is crowded with invisible characters, proper to both sexes, to both the right and left side of the body, and to a long line of male and female ancestors separated by hundreds or even thousands of generations from the present time: and these characters, like those written on paper with invisible ink, lie ready to be evolved whenever the organisation is disturbed by certain known or unknown conditions.

REFERENCES [1] Youatt on Sheep, pp. 20, 234. The same fact of loose horns occasionally appearing in hornless breeds has been observed in [2] Youatt on Cattle, pp. 155, 174.

[3] Youatt on Sheep, 1838, pp. 17, 145.

[4] I have been informed of this fact through the Rev. W. D. Fox on the excellent authority of Mr. Wilmot: _see also_ remarks on this subject in an article in the ‘Quarterly Review,’ 1849, p. 395.

with much experience on this subject, has likewise assured me that this sometimes occurs.

sheep often running wild in the Bavarian Alps; but, on making further inquiries at my request, he found that they are not able to establish themselves; they generally perish from the frozen snow clinging to their wool, and they have lost the skill necessary to pass over steep icy slopes. On one occasion two ewes survived the winter, but their lambs perished.

[11] _See_ some excellent remarks on this subject by Mr. Wallace [12] Dureau de la Malle ‘Comptes Rendus,’ tom. xli., 1855, p. 807.

From the statements above given, the author concludes that the wild pigs of Louisiana are not descended from the European _Sus scrofa._ [13] Capt. W. Allen, in his ‘Expedition to the Niger,’ states that fowls have run wild on the island of Annobon, and have become modified in form and voice. The account is so meagre and vague that it did not appear to me worth copying; but I now find that Dureau de la Malle (‘Comptes Rendus,’ tom. xli., 1855, p. 690) advances this as a good instance of reversion to the primitive stock, and as confirmatory of a still more vague statement in classical times by Varro.

[16] Mr. Sedgwick gives many instances in the ‘British and Foreign [17] In his edit. of ‘Youatt on the Pig,’ 1860, p. 27.

practical article on the subject in ‘Gard. Chronicle,’ 1856, p. 620. I could add a vast number of references, but they would be superfluous.

[19] Kölreuter gives curious cases in his ‘Dritte Fortsetzung,’ 1766, ss. 53, 59; and in his well-known ‘Memoirs on Lavatera and Jalapa.’

Gärtner, ‘Bastarderzeugung,’ ss. 437, 441, etc. Naudin in his p. 25.

[20] Quoted by Mr. Sedgwick in ‘Med.-Chirurg. Review,’ April, 1861, p.

485. Dr. H. Dobell in ‘Med.-Chirurg. Transactions,’ vol. xlvi., gives an analogous case in which, in a large family, fingers with thickened joints were transmitted to several members during five generations; but when the blemish once disappeared it never reappeared.

[22] ‘Nouvelles Archives du Muséum,’ tom. i. p. 25. Alex. Braun (in his ‘Rejuvenescence,’ Ray Soc., 1853, p. 315) apparently holds a similar opinion.

[23] Mr. Teebay in ‘The Poultry Book,’ by Mr. Tegetmeier, 1866, p. 72.

[24] Quoted by Hofacker ‘Ueber die Eigenschaften,’ etc., s. 98.

[25] Azara, ‘Essais Hist. Nat. de Paraguay,’ tom. ii. 1801, p. 372.

[26] These facts are given on the high authority of Mr. Hewitt, in [29] Ibid., 1860, p. 343. I am glad to find that so experienced a breeder of cattle as Mr. Willoughby Wood, (‘Gard. Chron.’ 1869, p.

1216), admits my principle of a cross giving a tendency to reversion.

[32] ‘Mém. présentés par divers Savans à l’Acad. Royale,’ tom. vi.

1835, p. 338.

[36] Sclater, in ‘Proc. Zoolog. Soc.,’ 1862, p. 163: this species is the Ghor-Khur of N.W. India, and has often been called the Hemionus of Pallas. _See also_ Mr. Blyth’s excellent paper in ‘Journal of Asiatic Soc. of Bengal,’ vol. xviii., 1860, p. 229.

[37] Another species of wild ass, the true _E. hemionus_ or _Kiang,_ which ordinarily has no shoulder-stripes, is said occasionally to have them; and these, as with the horse and ass, are sometimes double: _see_ Mr. Blyth in the paper just quoted and in ‘Indian Sporting Review,’ 1856, p. 320: and Col. Hamilton Smith in ‘Nat. Library, [38] Figured in the ‘Gleanings from the Knowsley Menageries,’ by Dr.

J. E. Gray.

[39] ‘Darwin’sche Theorie und ihre Stellung zu Moral und Religion,’ p.

85.

[40] Cases of both Spanish and Polish hens sitting are given in the [41] ‘The Poultry Book,’ by Mr. Tegetmeier, 1866, pp. 119, 163. The author, who remarks on the two negatives (‘Journ. of Hort.,’ 1862, p.

325), states that two broods were raised from a Spanish cock and Silver-pencilled Hamburgh hen, neither of which are incubators, and no less than seven out of eight hens in these two broods “showed a perfect obstinacy in sitting.” The Rev. E. S. Dixon (‘Ornamental Poultry,’ 1848, p. 200) says that chickens reared from a cross between Golden and Black Polish fowls, are “good and steady birds to sit.” Mr.

B. P. Brent informs me that he raised some good sitting hens by crossing Pencilled Hamburgh and Polish breeds. A cross-bred bird from a Spanish non-incubating cock and Cochin incubating hen is mentioned in the ‘Poultry Chronicle,’ vol. iii. p. 13, as an “exemplary mother.”

On the other hand, an exceptional case is given in the ‘Cottage Gardener,’ 1860, p. 388, of a hen raised from a Spanish cock and black Polish hen which did not incubate.

[45] As stated by Mr. Orton, in his ‘Physiology of Breeding,’ p. 12.

[46] M. E. de Selys-Longchamps refers (‘Bulletin Acad. Roy. de Bruxelles,’ tom. xii. No. 10) to more than seven of these hybrids shot in Switzerland and France. M. Deby asserts (‘Zoologist,’ vol. v., 1845-46, p. 1254) that several have been shot in various parts of Belgium and Northern France. Audubon (‘Ornitholog. Biography,’ vol.

iii. p. 168), speaking of these hybrids, says that, in North America, they “now and then wander off and become quite wild.”

[49] Dr. P. Broca, on ‘Hybridity in the Genus Homo,’ Eng. translat., 1864, p. 39.

[53] Prof. Weismann, in his very curious essay on the different forms (‘Saison-Dimorphismus der Schmetterlinge,’ pp. 27, 28), has come to a similar conclusion, namely, that any cause which disturbs the organisation, such as the exposure of the cocoons to heat or even to much shaking, gives a tendency to reversion.

[56] In his ‘Essays on Nat. Hist.,’ 1838, Mr. Hewitt gives analogous cases with hen-pheasants in ‘Journal of Horticulture,’ July 12, 1864, p. 37. Isidore Geoffroy Saint-Hilaire, in his ‘Essais de Zoolog. Gen.’

(‘suites a Buffon,’ 1842, pp. 496-513), has collected such cases in ten different kinds of birds. It appears that Aristotle was well aware of the change in mental disposition in old hens. The case of the female deer acquiring horns is given at p. 513.

p. 31.

[60] _See_ Steenstrup on the ‘Obliquity of Flounders’: in ‘Annals and Mag. of Nat. Hist.’ May, 1865, p. 361. I have given an abstract of Malm’s explanation of this wonderful phenomenon in the ‘Origin of [61] Dr. E. von Martens, in ‘Annals and Mag. of Nat. Hist.’ March, 1866, p. 209.

appended remarks on the apparently capricious development of the thoracic limbs on the right and left sides in the higher crustaceans.

opportunity of examining these remarkable feathers through the kindness of Mr. Tegetmeier.

Lecture, Royal Institution, March 16, 1860. On same subject see Peyritsch has collected a large number of very interesting cases, Sitzb. d. k. Akad. d. Wissensch.: Wien. b. LX. and especially b.

[70] In his discussion on some curious peloric Calceolarias, quoted in [71] For other cases of six divisions in peloric flowers of the 192.

[72] Godron, reprinted from the ‘Mémoires de l’Acad. de Stanislas,’ 1868.

CHAPTER XIV. INHERITANCE _continued_—FIXEDNESS OF CHARACTER—PREPOTENCY—SEXUAL LIMITATION—CORRESPONDENCE OF AGE.

FIXEDNESS OF CHARACTER APPARENTLY NOT DUE TO ANTIQUITY OF INITANCE—PREPOTENCY OF TRANSMISSION IN INDIVIDUALS OF THE SAME FAMILY, IN CROSSED BREEDS AND SPECIES; OFTEN STRONGER IN ONE SEX THAN THE OTHER; SOMETIMES DUE TO THE SAME CHARACTER BEING PRESENT AND VISIBLE IN ONE BREED AND LATENT IN THE OTHER—INHERITANCE AS LIMITED BY SEX—NEWLY-ACQUIRED CHARACTERS IN OUR DOMESTICATED ANIMALS OFTEN TRANSMITTED BY ONE SEX ALONE, SOMETIMES LOST BY ONE SEX ALONE—INHERITANCE AT CORRESPONDING PERIODS OF LIFE—THE IMPORTANCE OF THE PRINCIPLE WITH RESPECT TO EMBRYOLOGY; AS EXHIBITED IN DOMESTICATED ANIMALS: AS EXHIBITED IN THE APPEARANCE AND DISAPPEARANCE OF INHERITED DISEASES; SOMETIMES SUPERVENING EARLIER IN THE CHILD THAN IN THE PARENT—SUMMARY OF THE THREE PRECEDING CHAPTERS.

In the last two chapters the nature and force of Inheritance, the circumstances which interfere with its power, and the tendency to Reversion, with its many remarkable contingencies, were discussed. In the present chapter some other related phenomena will be treated of, as fully as my materials permit.

It is a general belief amongst breeders that the longer any character has been transmitted by a breed, the more fully it will continue to be transmitted. I do not wish to dispute the truth of the proposition that inheritance gains strength simply through long continuance, but I doubt whether it can be proved. In one sense the proposition is little better than a truism; if any character has remained constant during many generations, it will be likely to continue so, if the conditions of life remain the same. So, again, in improving a breed, if care be taken for a length of time to exclude all inferior individuals, the breed will obviously tend to become truer, as it will not have been crossed during many generations by an inferior animal. We have previously seen, but without being able to assign any cause, that, when a new character appears, it is occasionally from the first constant, or fluctuates much, or wholly fails to be transmitted. So it is with the aggregate of slight differences which characterise a new variety, for some propagate their kind from the first much truer than others. Even with plants multiplied by bulbs, layers, etc., which may in one sense be said to form parts of the same individual, it is well known that certain varieties retain and transmit through successive bud-generations their newly-acquired characters more truly than others. In none of these, nor in the following cases, does there appear to be any relation between the force with which a character is transmitted and the length of time during which it has been transmitted. Some varieties, such as white and yellow hyacinths and white sweet-peas, transmit their colours more faithfully than do the varieties which have retained their natural colour. In the Irish family, mentioned in the twelfth chapter, the peculiar tortoiseshell-like colouring of the eyes was transmitted far more faithfully than any ordinary colour. Ancon and Mauchamp sheep and niata cattle, which are all comparatively modern breeds, exhibit remarkably strong powers of inheritance. Many similar cases could be adduced.

As all domesticated animals and cultivated plants have varied, and yet are descended from aboriginally wild forms, which no doubt had retained the same character from an immensely remote epoch, we see that scarcely any degree of antiquity ensures a character being transmitted perfectly true. In this case, however, it may be said that changed conditions of life induce certain modifications, and not that the power of inheritance fails; but in every case of failure, some cause, either internal or external, must interfere. It will generally be found that the organs or parts which in our domesticated productions have varied, or which still continue to vary,—that is, which fail to retain their former state,—are the same with the parts which differ in the natural species of the same genus. As, on the theory of descent with modification, the species of the same genus have been modified since they branched off from a common progenitor, it follows that the characters by which they differ from one another have varied, whilst other parts of the organisation have remained unchanged; and it might be argued that these same characters now vary under domestication, or fail to be inherited, from their lesser antiquity. But variation in a state of nature seems to stand in some close relation with changed conditions of life, and characters which have already varied under such conditions would be apt to vary under the still greater changes consequent on domestication, independently of their greater or less antiquity.

Fixedness of character, or the strength of inheritance, has often been judged of by the preponderance of certain characters in the crossed offspring between distinct races; but prepotency of transmission here comes into play, and this, as we shall immediately see, is a very different consideration from the strength or weakness of inheritance.[1] It has often been observed that breeds of animals inhabiting wild and mountainous countries cannot be permanently modified by our improved breeds; and as these latter are of modern origin, it has been thought that the greater antiquity of the wilder breeds has been the cause of their resistance to improvement by crossing; but it is more probably due to their structure and constitution being better adapted to the surrounding conditions. When plants are first subjected to culture, it has been found that, during several generations, they transmit their characters truly, that is, do not vary, and this has been attributed to ancient characters being strongly inherited: but it may with equal or greater probability be consequent on changed conditions of life requiring a long time for their cumulative action. Notwithstanding these considerations, it would perhaps be rash to deny that characters become more strongly fixed the longer they are transmitted; but I believe that the proposition resolves itself into this,—that characters of all kinds, whether new or old, tend to be inherited, and that those which have already withstood all counteracting influences and been truly transmitted, will, as a general rule, continue to withstand them, and consequently be faithfully inherited.

_Prepotency in the Transmission of Character._ When individuals, belonging to the same family, but distinct enough to be recognised, or when two well-marked races, or two species, are crossed, the usual result, as stated in the previous chapter, is, that the offspring in the first generation are intermediate between their parents, or resemble one parent in one part and the other parent in another part. But this is by no means the invariable rule; for in many cases it is found that certain individuals, races, and species, are prepotent in transmitting their likeness. This subject has been ably discussed by Prosper Lucas,[2] but is rendered extremely complex by the prepotency sometimes running equally in both sexes, and sometimes more strongly in one sex than in the other; it is likewise complicated by the presence of secondary sexual characters, which render the comparison of crossed breeds with their parents difficult.

It would appear that in certain families some one ancestor, and after him others in the same family, have had great power in transmitting their likeness through the male line; for we cannot otherwise understand how the same features should so often be transmitted after marriages with many females, as in the case of the Austrian Emperors; and so it was, according to Niebuhr, with the mental qualities of certain Roman families.[3] The famous bull Favourite is believed[4] to have had a prepotent influence on the shorthorn race. It has also been observed[5] with English racehorses that certain mares have generally transmitted their own character, whilst other mares of equally pure blood have allowed the character of the sire to prevail. A famous black greyhound, Bedlamite, as I hear from Mr. C. M. Brown “invariably got all his puppies black, no matter what was the colour of the bitch;” but then Bedlamite “had a preponderance of black in his blood, both on the sire and dam side.”

The truth of the principle of prepotency comes out more clearly when distinct races are crossed. The improved Shorthorns, notwithstanding that the breed is comparatively modern, are generally acknowledged to possess great power in impressing their likeness on all other breeds; and it is chiefly in consequence of this power that they are so highly valued for exportation.[6] Godine has given a curious case of a ram of a goat-like breed of sheep from the Cape of Good Hope, which produced offspring hardly to be distinguished from himself, when crossed with ewes of twelve other breeds. But two of these half-bred ewes, when put to a merino ram, produced lambs closely resembling the merino breed.

Girou de Buzareingues[7] found that of two races of French sheep the ewes of one, when crossed during successive generations with merino rams, yielded up their character far sooner than the ewes of the other race. Sturm and Girou have given analogous cases with other breeds of sheep and with cattle, the prepotency running in these cases through the male side; but I was assured on good authority in South America, that when niata cattle are crossed with common cattle, though the niata breed is prepotent whether males or females are used, yet that the prepotency is strongest through the female line. The Manx cat is tailless and has long hind legs; Dr. Wilson crossed a male Manx with common cats, and, out of twenty-three kittens, seventeen were destitute of tails; but when the female Manx was crossed by common male cats all the kittens had tails, though they were generally short and imperfect.[8] In making reciprocal crosses between pouter and fantail pigeons, the pouter-race seemed to be prepotent through both sexes over the fantail.

But this is probably due to weak power in the fantail rather than to