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Чарльз Дарвин
The Variation of Animals and Plants under Domestication — Volume 2

With plants the period of vegetation is easily changed and is inherited, as in the case of summer and winter wheat, barley, and vetches; but to this subject we shall immediately return under acclimatisation. Annual plants sometimes become perennial under a new climate, as I hear from Dr. Hooker is the case with the stock and mignonette in Tasmania. On the other hand, perennials sometimes become annuals, as with the Ricinus in England, and as, according to Captain Mangles, with many varieties of the heartsease. Von Berg (24/40. 'Flora' 1835 b. 2 page 504.) raised from seed of Verbascum phoeniceum, which is usually a biennial, both annual and perennial varieties. Some deciduous bushes become evergreen in hot countries. (24/41. Alph. de Candolle 'Geograph. Bot.' tome 2 page 1078.) Rice requires much water, but there is one variety in India which can be grown without irrigation. (24/42. Royle 'Illustrations of the Botany of the Himalaya' page 19.) Certain varieties of the oat and of our other cereals are best fitted for certain soils. (24/43. 'Gardener's Chronicle' 1850 pages 204, 219.) Endless similar facts could be given in the animal and vegetable kingdoms. They are noticed here because they illustrate analogous differences in closely allied natural species, and because such changed habits of life, whether due to habit, or to the direct action of external conditions, or to so-called spontaneous variability, would be apt to lead to modifications of structure.

ACCLIMATISATION.

From the previous remarks we are naturally led to the much disputed subject of acclimatisation. There are two distinct questions: Do varieties descended from the same species differ in their power of living under different climates? And secondly, if they so differ, how have they become thus adapted? We have seen that European dogs do not succeed well in India, and it is asserted (24/44. Rev. R. Everest 'Journal As. Soc. of Bengal' volume 3 page 19.), that no one has there succeeded in keeping the Newfoundland long alive; but then it may be argued, and probably with truth, that these northern breeds are specifically distinct from the native dogs which flourish in India. The same remark may be made with respect to different breeds of sheep, of which, according to Youatt (24/45. Youatt on 'Sheep' 1838 page 491.), not one brought "from a torrid climate lasts out the second year," in the Zoological Gardens. But sheep are capable of some degree of acclimatisation, for Merino sheep bred at the Cape of Good Hope have been found far better adapted for India than those imported from England. (24/46. Royle 'Prod. Resources of India' page 153.) It is almost certain that all the breeds of the fowl are descended from one species; but the Spanish breed, which there is good reason to believe originated near the Mediterranean (24/47. Tegetmeier 'Poultry Book' 1866 page 102.), though so fine and vigorous in England, suffers more from frost than any other breed. The Arrindy silk moth introduced from Bengal, and the Ailanthus moth from the temperate province of Shan Tung, in China, belong to the same species, as we may infer from their identity in the caterpillar, cocoon, and mature states (24/48. Dr. R. Paterson in a paper communicated to Bot. Soc. of Canada quoted in the 'Reader' 1863 November 13.); yet they differ much in constitution: the Indian form "will flourish only in warm latitudes," the other is quite hardy and withstands cold and rain.

[Plants are more strictly adapted to climate than are animals. The latter when domesticated withstand such great diversities of climate, that we find nearly the same species in tropical and temperate countries; whilst the cultivated plants are widely dissimilar. Hence a larger field is open for inquiry in regard to the acclimatisation of plants than of animals. It is no exaggeration to say that with almost every plant which has long been cultivated, varieties exist which are endowed with constitutions fitted for very different climates; I will select only a few of the more striking cases, as it would be tedious to give all. In North America numerous fruit-trees have been raised, and in horticultural publications, — for instance, in that by Downing, — lists are given of the varieties which are best able to withstand the severe climate of the northern States and Canada. Many American varieties of the pear, plum, and peach are excellent in their own country, but until recently, hardly one was known that succeeded in England; and with apples (24/49. See remarks by Editor in 'Gardener's Chronicle' 1848 page 5.), not one succeeds. Though the American varieties can withstand a severer winter than ours, the summer here is not hot enough. Fruit-trees have also originated in Europe with different constitutions, but they are not much noticed, because nurserymen here do not supply wide areas. The Forelle pear flowers early, and when the flowers have just set, and this is the critical period, they have been observed, both in France and England, to withstand with complete impunity a frost of 18 deg and even 14 deg Fahr., which killed the flowers, whether fully expanded or in bud, of all other kinds of pears. (24/50. 'Gardener's Chronicle' 1860 page 938. Remarks by Editor and quotation from Decaisne.) This power in the flower of resisting cold and afterwards producing fruit does not invariably depend, as we know on good authority (24/51. J. de Jonghe of Brussels in 'Gardener's Chronicle' 1857 page 612.), on general constitutional vigour. In proceeding northward, the number of varieties which are found capable of resisting the climate rapidly decreases, as may be seen in the list of the varieties of the cherry, apple, and pear, which can be cultivated in the neighbourhood of Stockholm. (24/52. Ch. Martius 'Voyage Bot. Cotes Sept. de la Norvege' page 26.) Near Moscow, Prince Troubetzkoy planted for experiment in the open ground several varieties of the pear, but one alone, the Poire sans Pepins, withstood the cold of winter. (24/53. 'Journal de l'Acad. Hort. de Gand' quoted in 'Gardener's Chronicle' 1859 page 7.) We thus see that our fruit-trees, like distinct species of the same genus, certainly differ from each other in their constitutional adaptation to different climates.

With the varieties of many plants, the adaptation to climate is often very close. Thus it has been proved by repeated trials "that few if any of the English varieties of wheat are adapted for cultivation in Scotland" (24/54. 'Gardener's Chronicle' 1851 page 396.); but the failure in this case is at first only in the quantity, though ultimately in the quality, of the grain produced. The Rev. M.J. Berkeley sowed wheat-seed from India, and got "the most meagre ears," on land which would certainly have yielded a good crop from English wheat. (24/55. Ibid 1862 page 235.) In these cases varieties have been carried from a warmer to a cooler climate; in the reverse case, as "when wheat was imported directly from France into the West Indian Islands, it produced either wholly barren spikes or furnished with only two or three miserable seeds, while West Indian seed by its side yielded an enormous harvest." (24/56. On the authority of Labat quoted in 'Gardener's Chronicle' 1862 page 235.) Here is another case of close adaptation to a slightly cooler climate; a kind of wheat which in England may be used indifferently either as a winter or summer variety, when sown under the warmer climate of Grignan, in France, behaved exactly as if it had been a true winter wheat. (24/57. MM. Edwards and Colin 'Annal. des Sc. Nat.' 2nd series Bot. tome 5 page 22.)

Botanists believe that all the varieties of maize belong to the same species; and we have seen that in North America, in proceeding northward, the varieties cultivated in each zone produce their flowers and ripen their seed within shorter and shorter periods. So that the tall, slowly maturing southern varieties do not succeed in New England, and the New English varieties do not succeed in Canada. I have not met with any statement that the southern varieties are actually injured or killed by a degree of cold which the northern varieties can withstand with impunity, though this is probable; but the production of early flowering and early seeding varieties deserves to be considered as one form of acclimatisation. Hence it has been found possible, according to Kalm, to cultivate maize further and further northwards in America. In Europe, also, as we learn from the evidence given by Alph. De Candolle, the culture of maize has extended since the end of the last century thirty leagues north of its former boundary. (24/58. 'Geograph. Bot.' page 337.) On the authority of Linnaeus (24/59. 'Swedish Acts' English translation 1739-40 volume 1. Kalm in his 'Travels' volume 2 page 166 gives an analogous case with cotton-plants raised in New Jersey from Carolina seed.), I may quote an analogous case, namely, that in Sweden tobacco raised from home-grown seed ripens its seed a month sooner and is less liable to miscarry than plants raised from foreign seed.

With the Vine, differently from the maize, the line of practical culture has retreated a little southward since the middle ages (24/60. De Candolle 'Geograph. Bot.' page 339.); but this seems due to commerce being now easier, so that it is better to import wine from the south than to make it in northern districts. Nevertheless the fact of the vine not having spread northward shows that acclimatisation has made no progress during several centuries. There is, however, a marked difference in the constitution of the several varieties, — some being hardy, whilst others, like the muscat of Alexandria, require a very high temperature to come to perfection. According to Labat (24/61. 'Gardener's Chronicle' 1862 page 235.), vines taken from France to the West Indies succeed with extreme difficulty, whilst those imported from Madeira or the Canary Islands thrive admirably.

Gallesio gives a curious account of the naturalisation of the Orange in Italy. During many centuries the sweet orange was propagated exclusively by grafts, and so often suffered from frosts, that it required protection. After the severe frost of 1709, and more especially after that of 1763, so many trees were destroyed, that seedlings from the sweet orange were raised, and, to the surprise of the inhabitants, their fruit was found to be sweet. The trees thus raised were larger, more productive, and hardier than the old kinds; and seedlings are now continually raised. Hence Gallesio concludes that much more was effected for the naturalisation of the orange in Italy by the accidental production of new kinds during a period of about sixty years, than had been effected by grafting old varieties during many ages. (24/62. Gallesio 'Teoria della Riproduzione Veg.' 1816 page 125; and 'Traite du Citrus' 1811 page 359.) I may add that Risso (24/63. 'Essai sur l'Hist. des Orangers' 1813 page 20 etc.) describes some Portuguese varieties of the orange as extremely sensitive to cold, and as much tenderer than certain other varieties.

The peach was known to Theophrastus, 322 B.C. (24/64. Alph. de Candolle 'Geograph. Bot.' page 882.) According to the authorities quoted by Dr. F. Rolle (24/65. 'Ch. Darwin's Lehre von der Entstehung' etc. 1862 s. 87.), it was tender when first introduced into Greece, and even in the island of Rhodes only occasionally bore fruit. If this be correct, the peach, in spreading during the last two thousand years over the middle parts of Europe, must have become much hardier. At the present day different varieties differ much in hardiness: some French varieties will not succeed in England; and near Paris, the Pavie de Bonneuil does not ripen its fruit till very late in the season, even when grown on a wall; "it is, therefore, only fit for a very hot southern climate." (24/66. Decaisne quoted in 'Gardener's Chronicle' 1865 page 271.)

I will briefly give a few other cases. A variety of Magnolia grandiflora, raised by M. Roy, withstands a temperature several degrees lower than that which any other variety can resist. With camellias there is much difference in hardiness. One particular variety of the Noisette rose withstood the severe frost of 1860 "untouched and hale amidst a universal destruction of other Noisettes." In New York the "Irish yew is quite hardy, but the common yew is liable to be cut down." I may add that there are varieties of the sweet potato (Convolvulus batatas) which are suited for warmer, as well as for colder, climates. (24/67. For the magnolia see Loudon's 'Gardener's Mag.' volume 13 1837 page 21. For camellias and roses see 'Gardener's Chronicle' 1860 page 384. For the yew 'Journal of Hort.' March 3, 1863 p 174. For sweet potatoes see Col. von Siebold in 'Gardener's Chronicle' 1855 page 822.)]

The plants as yet mentioned have been found capable of resisting an unusual degree of cold or heat, when fully grown. The following cases refer to plants whilst young. In a large bed of young Araucarias of the same age, growing close together and equally exposed, it was observed (24/68. The Editor 'Gardener's Chronicle' 1861 page 239.), after the unusually severe winter of 1860-61, that, "in the midst of the dying, numerous individuals remained on which the frost had absolutely made no kind of impression." Dr. Lindley, after alluding to this and other similar cases, remarks, "Among the lessons which the late formidable winter has taught us, is that, even in their power of resisting cold, individuals of the same species of plants are remarkably different." Near Salisbury, there was a sharp frost on the night of May 24, 1836, and all the French beans (Phaseolus vulgaris) in a bed were killed except about one in thirty, which completely escaped. (24/69. Loudon's 'Gardener's Mag.' volume 12 1836 page 378.) On the same day of the month, but in the year 1864, there was a severe frost in Kent, and two rows of scarlet- runners (P. multiflorus) in my garden, containing 390 plants of the same age and equally exposed, were all blackened and killed except about a dozen plants. In an adjoining row of "Fulmer's dwarf bean" (P. vulgaris), one single plant escaped. A still more severe frost occurred four days afterwards, and of the dozen plants which had previously escaped only three survived; these were not taller or more vigorous than the other young plants, but they escaped completely, with not even the tips of their leaves browned. It was impossible to behold these three plants, with their blackened, withered, and dead brethren all around, and not see at a glance that they differed widely in constitutional power of resisting frost.

This work is not the proper place to show that wild plants of the same species, naturally growing at different altitudes or under different latitudes, become to a certain extent acclimatised, as is proved by the different behaviour of their seedlings when raised in another country. In my 'Origin of Species' I have alluded to some cases, and I could add many others. One instance must suffice: Mr. Grigor, of Forres (24/70. 'Gardener's Chronicle' 1865 page 699. Mr. G. Maw gives ('Gardener's Chronicle' 1870 page 895) a number of striking cases; he brought home from southern Spain and northern Africa several plants, which he cultivated in England alongside specimens from northern districts; and he found a great difference not only in their hardiness during the winter, but in the behaviour of some of them during the summer.), states that seedlings of the Scotch fir (Pinus sylvestris), raised from seed from the Continent and from the forests of Scotland, differ much. "The difference is perceptible in one-year-old, and more so in two-year- old seedlings; but the effects of the winter on the second year's growth almost uniformly make those from the Continent quite brown, and so damaged, that by the month of March they are quite unsaleable, while the plants from the native Scotch pine, under the same treatment, and standing alongside, although considerably shorter, are rather stouter and quite green, so that the beds of the one can be known from the other when seen from the distance of a mile." Closely similar facts have been observed with seedling larches.

[Hardy varieties would alone be valued or noticed in Europe; whilst tender varieties, requiring more warmth, would generally be neglected; but such occasionally arise. Thus Loudon (24/71. 'Arboretum et Fruticetum' volume 3 page 1376.) describes a Cornish variety of the elm which is almost an evergreen, and of which the shoots are often killed by the autumnal frosts, so that its timber is of little value. Horticulturists know that some varieties are much more tender than others: thus all the varieties of the broccoli are more tender than cabbages; but there is much difference in this respect in the sub-varieties of the broccoli; the pink and purple kinds are a little hardier than the white Cape broccoli, "but they are not to be depended on after the thermometer falls below 24 deg Fahr.;" the Walcheren broccoli is less tender than the Cape, and there are several varieties which will stand much severer cold than the Walcheren. (24/72. Mr. Robson in 'Journal of Horticulture' 1861 page 23.) Cauliflowers seed more freely in India than cabbages. (24/73. Dr. Bonavia 'Report of the Agri. — Hort. Soc. of Oudh' 1866.) To give one instance with flowers: eleven plants raised from a hollyhock, called the Queen of the Whites (24/74. 'Cottage Gardener' 1860 April 24 page 57.) were found to be much more tender than various other seedlings. It may be presumed that all tender varieties would succeed better under a climate warmer than ours. With fruit-trees, it is well known that certain varieties, for instance of the peach, stand forcing in a hot-house better than others; and this shows either pliability of organisation or some constitutional difference. The same individual cherry-tree, when forced, has been observed during successive years gradually to change its period of vegetation. (24/75. 'Gardener's Chronicle' 1841 page 291.) Few pelargoniums can resist the heat of a stove, but Alba Multiflora will, as a most skilful gardener asserts, "stand pine-apple top and bottom heat the whole winter; without looking any more drawn than if it had stood in a common greenhouse; and Blanche Fleur seems as if it had been made on purpose for growing in winter, like many bulbs, and to rest all summer." (24/76. Mr. Beaton in 'Cottage Gardener' March 20, 1860 page 377. Queen Mab will also stand stove heat. See 'Gardener's Chronicle' 1845 page 226.) There can hardly be a doubt that the Alba Multiflora pelargonium must have a widely different constitution from that of most other varieties of this plant; it would probably withstand even an equatorial climate.

We have seen that according to Labat the vine and wheat require acclimatisation in order to succeed in the West Indies. Similar facts have been observed at Madras: "two parcels of mignonette-seed, one direct from Europe, the other saved at Bangalore (of which the mean temperature is much below that of Madras), were sown at the same time: they both vegetated equally favourably, but the former all died off a few days after they appeared above ground; the latter still survive, and are vigorous, healthy plants." "So again, turnip and carrot seed saved at Hyderabad are found to answer better at Madras than seed from Europe or from the Cape of Good Hope." (24/77. 'Gardener's Chronicle' 1841 page 439.) Mr. J. Scott of the Calcutta Botanic Gardens, informs me that seeds of the sweet-pea (Lathyrus odoratus) imported from England produce plants, with thick, rigid stems and small leaves, which rarely blossom and never yield seed; plants raised from French seed blossom sparingly, but all the flowers are sterile; on the other hand, plants raised from sweet-peas grown near Darjeeling in Upper India, but originally derived from England, can be successfully cultivated on the plains of India; for they flower and seed profusely, and their stems are lax and scandent. In some of the foregoing cases, as Dr. Hooker has remarked to me, the greater success may perhaps be attributed to the seeds having been more fully ripened under a more favourable climate; but this view can hardly be extended to so many cases, including plants, which, from being cultivated under a climate hotter than their native one, become fitted for a still hotter climate. We may therefore safely conclude that plants can to a certain extent become accustomed to a climate either hotter or colder than their own; although the latter cases have been more frequently observed.]

We will now consider the means by which acclimatisation may be effected, namely, through the appearance of varieties having a different constitution, and through the effects of habit. In regard to new varieties, there is no evidence that a change in the constitution of the offspring necessarily stands in any direct relation with the nature of the climate inhabited by the parents. On the contrary, it is certain that hardy and tender varieties of the same species appear in the same country. New varieties thus spontaneously arising become fitted to slightly different climates in two different ways; firstly, they may have the power, either as seedlings or when full-grown, of resisting intense cold, as with the Moscow pear, or of resisting intense heat, as with some kinds of Pelargonium, or the flowers may withstand severe frost, as with the Forelle pear. Secondly, plants may become adapted to climates widely different from their own, from flowering and fruiting either earlier or later in the season. In both these cases the power of acclimatisation by man consists simply in the selection and preservation of new varieties. But without any direct intention on his part of securing a hardier variety, acclimatisation may be unconsciously effected by merely raising tender plants from seed, and by occasionally attempting their cultivation further and further northwards, as in the case of maize, the orange and the peach.

How much influence ought to be attributed to inherited habit or custom in the acclimatisation of animals and plants is a much more difficult question. In many cases natural selection can hardly have failed to have come into play and complicated the result. It is notorious that mountain sheep resist severe weather and storms of snow which would destroy lowland breeds; but then mountain sheep have been thus exposed from time immemorial, and all delicate individuals will have been destroyed, and the hardiest preserved. So with the Arrindy silk-moths of China and India; who can tell how far natural selection may have taken a share in the formation of the two races, which are now fitted for such widely different climates? It seems at first probable that the many fruit-trees which are so well fitted for the hot summers and cold winters of North America, in contrast with their poor success under our climate, have become adapted through habit; but when we reflect on the multitude of seedlings annually raised in that country, and that none would succeed unless born with a fitting constitution, it is possible that mere habit may have done nothing towards their acclimatisation. On the other hand, when we hear that Merino sheep, bred during no great number of generations at the Cape of Good Hope — that some European plants raised during only a few generations in the cooler parts of India, withstand the hotter parts of that country much better than the sheep or seeds imported directly from England, we must attribute some influence to habit. We are led to the same conclusion when we hear from Naudin (24/78. Quoted by Asa Gray in 'Am. Journ. of Sc.' 2nd series January 1865 page 106.) that the races of melons, squashes, and gourds, which have long been cultivated in Northern Europe, are comparatively more precocious, and need much less heat for maturing their fruit, than the varieties of the same species recently brought from tropical regions. In the reciprocal conversion of summer and winter wheat, barley, and vetches into each other, habit produces a marked effect in the course of a very few generations. The same thing apparently occurs with the varieties of maize, which, when carried from the Southern States of America, or into Germany, soon became accustomed to their new homes. With vine-plants taken to the West Indies from Madeira, which are said to succeed better than plants brought directly from France, we have some degree of acclimatisation in the individual, independently of the production of new varieties by seed.

The common experience of agriculturists is of some value, and they often advise persons to be cautious in trying the productions of one country in another. The ancient agricultural writers of China recommend the preservation and cultivation of the varieties peculiar to each country. During the classical period, Columella wrote, "Vernaculum pecus peregrino longe praestantius est." (24/79. For China see 'Memoire sur les Chinois' tome 11 1786 page 60. Columella is quoted by Carlier in 'Journal de Physique' tome 24 1784.)

I am aware that the attempt to acclimatise either animals or plants has been called a vain chimera. No doubt the attempt in most cases deserves to be thus called, if made independently of the production of new varieties endowed with a different constitution. With plants propagated by buds, habit rarely produces any effect; it apparently acts only through successive seminal generations. The laurel, bay, laurestinus, etc., and the Jerusalem artichoke, which are propagated by cuttings or tubers, are probably now as tender in England as when first introduced; and this appears to be the case with the potato, which until recently was seldom multiplied by seed. With plants propagated by seed, and with animals, there will be little or no acclimatisation unless the hardier individuals are either intentionally or unconsciously preserved. The kidney-bean has often been advanced as an instance of a plant which has not become hardier since its first introduction into Britain. We hear, however, on excellent authority (24/80. Messrs. Hardy and Son in 'Gardener's Chronicle' 1856 page 589.) that some very fine seed, imported from abroad, produced plants "which blossomed most profusely, but were nearly all but abortive, whilst plants grown alongside from English seed podded abundantly;" and this apparently shows some degree of acclimatisation in our English plants. We have also seen that seedlings of the kidney-bean occasionally appear with a marked power of resisting frost; but no one, as far as I can hear, has ever separated such hardy seedlings, so as to prevent accidental crossing, and then gathered their seed, and repeated the process year after year. It may, however, be objected with truth that natural selection ought to have had a decided effect on the hardiness of our kidney- beans; for the tenderest individuals must have been killed during every severe spring, and the hardier preserved. But it should be borne in mind that the result of increased hardiness would simply be that gardeners, who are always anxious for as early a crop as possible, would sow their seed a few days earlier than formerly. Now, as the period of sowing depends much on the soil and elevation of each district, and varies with the season; and as new varieties have often been imported from abroad, can we feel sure that our kidney-beans are not somewhat hardier? I have not been able, by searching old horticultural works, to answer this question satisfactorily.

On the whole the facts now given show that, though habit does something towards acclimatisation, yet that the appearance of constitutionally different individuals is a far more effective agent. As no single instance has been recorded either with animals or plants of hardier individuals having been long and steadily selected, though such selection is admitted to be indispensable for the improvement of any other character, it is not surprising that man has done little in the acclimatisation of domesticated animals and cultivated plants. We need not, however, doubt that under nature new races and new species would become adapted to widely different climates, by variation, aided by habit, and regulated by natural selection.

[ARRESTS OF DEVELOPMENT: RUDIMENTARY AND ABORTED ORGANS.

Modifications of structure from arrested development, so great or so serious as to deserve to be called monstrosities, are not infrequent with domesticated animals, but, as they differ much from any normal structure, they require only a passing notice. Thus the whole head may be represented by a soft nipple-like projection, and the limbs by mere papillae. These rudiments of limbs are sometimes inherited, as has been observed in a dog. (24/81. Isid. Geoffroy Saint-Hilaire 'Hist. Nat. des Anomalies' 1836 tome 2 pages 210, 223, 224, 395; 'Philosoph. Transact.' 1775 page 313.)

Many lesser anomalies appear to be due to arrested development. What the cause of the arrest may be, we seldom know, except in the case of direct injury to the embryo. That the cause does not generally act at an extremely early embryonic period we may infer from the affected organ seldom being wholly aborted, — a rudiment being generally preserved. The external ears are represented by mere vestiges in a Chinese breed of sheep; and in another breed, the tail is reduced "to a little button, suffocated in a manner, by fat." (24/82. Pallas quoted by Youatt on 'Sheep' page 25.) In tailless dogs and cats a stump is left. In certain breeds of fowls the comb and wattles are reduced to rudiments; in the Cochin-China breed scarcely more than rudiments of spurs exist. With polled Suffolk cattle, "rudiments of horns can often be felt at an early age" (24/83. Youatt on 'Cattle' 1834 page 174.); and with species in a state of nature, the relatively great development of rudimentary organs at an early period of life is highly characteristic of such organs. With hornless breeds of cattle and sheep, another and singular kind of rudiment has been observed, namely, minute dangling horns attached to the skin alone, and which are often shed and grow again. With hornless goats, according to Desmarest (24/84. 'Encyclop. Method.' 1820 page 483: see page 500, on the Indian zebu casting its horns. Similar cases in European cattle were given in the third chapter.), the bony protuberance which properly supports the horn exists as a mere rudiment.