The Variation of Animals and Plants under Domestication — Volume 1

Chapter 40

Chapter 404,220 wordsPublic domain

1 page 444) declares that he has found the types of all our cultivated varieties in wild seedlings, but then he looks on these seedlings as so many aboriginal stocks.)

[CEREALIA.

I will now enter on details. The cereals cultivated in Europe consist of four genera--wheat, rye, barley, and oats. Of wheat the best modern authorities (9/18. See A. De Candolle 'Geograph. Bot.' 1855 page 928 et seq. Godron 'De l'Espece' 1859 tome 2 page 70; and Metzger 'Die Getreidearten' etc. 1841.) make four or five, or even seven distinct species; of rye, one; of barley, three; and of oats, two, three, or four species. So that altogether our cereals are ranked by different authors under from ten to fifteen distinct species. These have given rise to a multitude of varieties. It is a remarkable fact that botanists are not universally agreed on the aboriginal parent-form of any one cereal plant. For instance, a high authority writes in 1855 (9/19. Mr. Bentham in his review entitled 'Hist. Notes on cultivated Plants' by Dr. A. Targioni- Tozzetti in 'Journal of Hort. Soc.' volume 9 1855 page 133. He informs me that he still retains the same opinion.), "We ourselves have no hesitation in stating our conviction, as the result of all the most reliable evidence, that none of these Cerealia exist, or have existed, truly wild in their present state, but that all are cultivated varieties of species now growing in great abundance in S. Europe or W. Asia." On the other hand, Alph. De Candolle (9/20. 'Geograph. Bot.' page 928. The whole subject is discussed with admirable fulness and knowledge.) has adduced abundant evidence that common wheat (Triticum vulgare) has been found wild in various parts of Asia, where it is not likely to have escaped from cultivation: and there is some force in M. Godron's remark, that, supposing these plants to be escaped seedlings (9/21. Godron 'De l'Espece' tome 2 page 72. A few years ago the excellent, though misinterpreted, observations of M. Fabre led many persons to believe that wheat was a modified descendant of Aegilops; but M. Godron (tome 1 page 165) has shown by careful experiments that the first step in the series, viz. Aegilops triticoides, is a hybrid between wheat and Ae. ovata. The frequency with which these hybrids spontaneously arise, and the gradual manner in which the Ae. triticoides becomes converted into true wheat, alone leave any doubt with respect to M. Godron's conclusions.), as they have propagated themselves in a wild state for several generations, their continued resemblance to cultivated wheat renders it probable that the latter has retained its aboriginal character. But the strong tendency to inheritance, which most of the varieties of wheat evince, as we shall presently see, is here greatly undervalued. Much weight must also be attributed to a remark by Professor Hildebrand (9/22. 'Die Verbreitungsmittel der Pflanzen' 1873 page 129.) that when the seeds or fruit of cultivated plants possess qualities disadvantageous to them as a means of distribution, we may feel almost sure that they no longer retain their aboriginal condition. On the other hand, M. De Candolle insists strongly on the frequent occurrence in the Austrian dominions of rye and of one kind of oats in an apparently wild condition. With the exception of these two cases, which however are rather doubtful, and with the exception of two forms of wheat and one of barley, which he believes to have been found truly wild, M. De Candolle does not seem fully satisfied with the other reported discoveries of the parent-forms of our other cereals. With respect to oats, according to Mr. Buckmann (9/23. Report to British Association for 1857 page 207.), the wild English Avena fatua can be converted by a few years of careful cultivation and selection into forms almost identical with two very distinct cultivated races. The whole subject of the origin and specific distinctness of the various cereal plants is a most difficult one; but we shall perhaps be able to judge a little better after considering the amount of variation which wheat has undergone.

Metzger describes seven species of wheat, Godron refers to five, and De Candolle to only four. It is not improbable that, besides the kinds known in Europe, other strongly characterised forms exist in the more distant parts of the world; for Loiseleur-Deslongchamps (9/24. Considerations sur les Cereales' 1842-43 page 29.) speaks of three new species or varieties, sent to Europe in 1822 from Chinese Mongolia, which he considers as being there indigenous. Moorcroft (9/25. 'Travels in the Himalayan Provinces' etc. 1841 volume 1 page 224.) also speaks of Hasora wheat in Ladakh as very peculiar. If those botanists are right who believe that at least seven species of wheat originally existed, then the amount of variation in any important character which wheat has undergone under cultivation has been slight; but if only four or a lesser number of species originally existed, then it is evident that varieties have arisen so strongly marked, that they have been considered by capable judges as specifically distinct. But the impossibility of deciding which forms ought to be ranked as species and which as varieties, makes it useless to specify in detail the differences between the various kinds of wheat. Speaking generally, the organs of vegetation differ little (9/26. Col. J. Le Couteur on the 'Varieties of Wheat' pages 23, 79.); but some kinds grow close and upright, whilst others spread and trail along the ground. The straw differs in being more or less hollow, and in quality. The ears (9/27. Loiseleur-Deslongchamps 'Consid. sur les Cereales' page 11.) differ in colour and in shape, being quadrangular, compressed, or nearly cylindrical; and the florets differ in their approximation to each other, in their pubescence, and in being more or less elongated. The presence or absence of barbs is a conspicuous difference, and in certain Gramineae serves even as a generic character (9/28. See an excellent review in Hooker 'Journ. of Botany' volume 8 page 82 note.); although, as remarked by Godron (9/29. 'De l'Espece' tome 2 page 73.) the presence of barbs is variable in certain wild grasses, and especially in those such as Bromus secalinus and Lolium temulentum, which habitually grow mingled with our cereal crops, and which have thus unintentionally been exposed to culture. The grains differ in size, weight, and colour; in being more or less downy at one end, in being smooth or wrinkled, in being either nearly globular, oval, or elongated; and finally in internal texture, being tender or hard, or even almost horny, and in the proportion of gluten which they contain.

Nearly all the races or species of wheat vary, as Godron (9/30. Ibid tome 2 page 75.) has remarked, in an exactly parallel manner,--in the seed being downy or glabrous, and in colour,--and in the florets being barbed or not barbed, etc. Those who believe that all the kinds are descended from a single wild species may account for this parallel variation by the inheritance of a similar constitution, and a consequent tendency to vary in the same manner; and those who believe in the general theory of descent with modification may extend this view to the several species of wheat, if such ever existed in a state of nature.

Although few of the varieties of wheat present any conspicuous difference, their number is great. Dalbret cultivated during thirty years from 150 to 160 kinds, and excepting in the quality of the grain they all kept true; Colonel Le Couteur possessed upwards of 150, and Philippar 322 varieties. (9/31. For Dalbret and Philippar see Loiseleur-Deslongchamps 'Consid. sur les Cereales' pages 45, 70. Le Couteur on Wheat pages 6, 14-17.) As wheat is an annual, we thus see how strictly many trifling differences in character are inherited through many generations. Colonel Le Couteur insists strongly on this same fact. In his persevering and successful attempts to raise new varieties, he found that there was only one "secure mode to ensure the growth of pure sorts, namely, to grow them from single grains or from single ears, and to follow up the plan by afterwards sowing only the produce of the most productive so as to form a stock." But Major Hallett (9/32. See his Essay on 'Pedigree in Wheat' 1862; also paper read before the British Association 1869 and other publications.) has gone much farther, and by the continued selection of plants from the grains of the same ear, during successive generations, has made his 'Pedigree in Wheat' (and other cereals) now famous in many quarters of the world. The great amount of variability in the plants of the same variety is another interesting point, which would never have been detected except by an eye long practised to the work; thus Colonel Le Couteur relates (9/33. 'Varieties of Wheat' Introduction page 6. Marshall in his 'Rural Economy of Yorkshire' volume 2 page 9 remarks that "in every field of corn there is as much variety as in a herd of cattle.") that in a field of his own wheat, which he considered at least as pure as that of any of his neighbours, Professor La Gasca found twenty-three sorts; and Professor Henslow has observed similar facts. Besides such individual variations, forms sufficiently well marked to be valued and to become widely cultivated sometimes suddenly appear: thus Mr. Shirreff has had the good fortune to raise in his lifetime seven new varieties, which are now extensively grown in many parts of Britain. (9/34. 'Gardener's Chronicle' and 'Agricult. Gazette' 1862 page 963.)

As in the case of many other plants, some varieties, both old and new, are far more constant in character than others. Colonel Le Couteur was forced to reject some of his new sub-varieties, which he suspected had been produced from a cross, as incorrigibly sportive. On the other hand Major Hallett (9/35. 'Gardener's Chronicle' November 1868 page 1199.) has shown how wonderfully constant some varieties are, although not ancient ones, and although cultivated in various countries. With respect to the tendency to vary, Metzger (9/36. 'Getreidearten' 1841 s. 66, 91, 92, 116, 117.) gives from his own experience some interesting facts: he describes three Spanish sub-varieties, more especially one known to be constant in Spain, which in Germany assumed their proper character only during hot summers; another variety kept true only in good land, but after having been cultivated for twenty-five years became more constant. He mentions two other sub-varieties which were at first inconstant, but subsequently became, apparently without any selection, accustomed to their new homes, and retained their proper character. These facts show what small changes in the conditions of life cause variability, and they further show that a variety may become habituated to new conditions. One is at first inclined to conclude with Loiseleur-Deslongchamps, that wheat cultivated in the same country is exposed to remarkably uniform conditions; but manures differ, seed is taken from one soil to another, and, what is far more important, the plants are exposed as little as possible to struggle with other plants, and are thus enabled to exist under diversified conditions. In a state of nature each plant is confined to that particular station and kind of nutriment which it can seize from the other plants by which it is surrounded.

Wheat quickly assumes new habits of life. The summer and winter kinds were classed by Linnaeus as distinct species; but M. Monnier (9/37. Quoted by Godron 'De l'Espece' volume 2 page 74. So it is, according to Metzger 'Getreidearten' s. 18, with summer and winter barley.) has proved that the difference between them is only temporary. He sowed winter-wheat in spring, and out of one hundred plants four alone produced ripe seeds; these were sown and resown, and in three years plants were reared which ripened all their seed. Conversely, nearly all the plants raised from summer-wheat, which was sown in autumn, perished from frost; but a few were saved and produced seed, and in three years this summer-variety was converted into a winter-variety. Hence it is not surprising that wheat soon becomes to a certain extent acclimatised, and that seed brought from distant countries and sown in Europe vegetates at first, or even for a considerable period (9/38. Loiseleur-Deslongchamps 'Cereales' part 2 page 224. Le Couteur page 70. Many other accounts could be added.) differently from our European varieties. In Canada the first settlers, according to Kalm (9/39. 'Travels in North America' 1753-1761 English translation volume 3 page 165.), found their winters too severe for winter-wheat brought from France, and their summers often too short for summer-wheat; and they thought that their country was useless for corn crops until they procured summer-wheat from the northern parts of Europe, which succeeded well. It is notorious that the proportion of gluten differs much under different climates. The weight of the grain is also quickly affected by climate: Loiseleur-Deslongchamps (9/40. 'Cereales' part 2 pages 179-183.) sowed near Paris 54 varieties, obtained from the South of France and from the Black Sea, and 52 of these yielded seed from 10 to 40 per cent heavier than the parent-seed. He then sent these heavier grains back to the South of France, but there they immediately yielded lighter seed.

All those who have closely attended to the subject insist on the close adaptation of numerous varieties of wheat to various soils and climates even within the same country; thus Colonel Le Couteur (9/41. 'On the Varieties of Wheat' Introduction page 7. See Marshall 'Rural Econ. of Yorkshire' volume 2 page 9. With respect to similar cases of adaptation in the varieties of oats see some interesting papers in the 'Gardener's Chronicle and Agricult. Gazette' 1850 pages 204, 219.) says, "It is the suitableness of each sort to each soil that will enable the farmer to pay his rent by sowing one variety, where he would be unable to do so by attempting to grow another of a seemingly better sort." This may be in part due to each kind becoming habituated to its conditions of life, as Metzger has shown certainly occurs, but it is probably in main part due to innate differences between the several varieties.

Much has been written on the deterioration of wheat; that the quality of the flour, size of grain, time of flowering, and hardness, may be modified by climate and soil, seems nearly certain; but that the whole body of any one sub-variety ever becomes changed into another and distinct sub-variety, there is no reason to believe. What apparently does take place, according to Le Couteur (9/42. 'On the Varieties of Wheat' page 59. Mr. Shirreff and a higher authority cannot be given ('Gardener's Chronicle and Agricult. Gazette' 1862 page 963), says "I have never seen grain which has either been improved or degenerated by cultivation, so as to convey the change to the succeeding crop.), is, that some one sub-variety out of the many which may always be detected in the same field is more prolific than the others, and gradually supplants the variety which was first sown.

With respect to the natural crossing of distinct varieties the evidence is conflicting, but preponderates against its frequent occurrence. Many authors maintain that impregnation takes place in the closed flower, but I am sure from my own observation that this is not the case, at least with those varieties to which I have attended. But as I shall have to discuss this subject in another work, it may be here passed over.]

In conclusion, all authors admit that numerous varieties of wheat have arisen; but their differences are unimportant, unless, indeed, some of the so-called species are ranked as varieties. Those who believe that from four to seven wild species of Triticum originally existed in nearly the same condition as at present, rest their belief chiefly on the great antiquity of the several forms. (9/43. Alph. De Candolle 'Geograph. Bot.' page 930. ) It is an important fact, which we have recently learnt from the admirable researches of Heer (9/44. 'Pflanzen der Pfahlbauten' 1866.), that the inhabitants of Switzerland, even so early as the Neolithic period, cultivated no less than ten cereal plants, namely, five kinds of wheat, of which at least four are commonly looked at as distinct species, three kinds of barley, a panicum, and a setaria. If it could be shown that at the earliest dawn of agriculture five kinds of wheat and three of barley had been cultivated, we should of course be compelled to look at these forms as distinct species. But, as Heer has remarked, agriculture even at the Neolithic period, had already made considerable progress; for, besides the cereals, peas, poppies, flax, and apparently apples, were cultivated. It may also be inferred, from one variety of wheat being the so called Egyptian, and from what is known of the native country of the panicum and setaria, as well as from the nature of the weeds which then grew mingled with the crops, that the lake-inhabitants either still kept up commercial intercourse with some southern people or had originally proceeded as colonists from the South.

Loiseleur-Deslongchamps (9/45. 'Les Cereales' page 94.) has argued that, if our cereal plants have been greatly modified by cultivation, the weeds which habitually grow mingled with them would have been equally modified. But this argument shows how completely the principle of selection has been overlooked. That such weeds have not varied, or at least do not vary now in any extreme degree, is the opinion of Mr. H.C. Watson and Professor Asa Gray, as they inform me; but who will pretend to say that they do not vary as much as the individual plants of the same sub-variety of wheat? We have already seen that pure varieties of wheat, cultivated in the same field, offer many slight variations, which can be selected and separately propagated; and that occasionally more strongly pronounced variations appear, which, as Mr. Shirreff has proved, are well worthy of extensive cultivation. Not until equal attention be paid to the variability and selection of weeds, can the argument from their constancy under unintentional culture be of any value. In accordance with the principles of selection we can understand how it is that in the several cultivated varieties of wheat the organs of vegetation differ so little; for if a plant with peculiar leaves appeared, it would be neglected unless the grains of corn were at the same time superior in quality or size. the selection of seed-corn was strongly recommended (9/46. Quoted by Le Couteur page 16.) in ancient times by Columella and Celsus; and as Virgil says,--

I've seen the largest seeds, tho' view'd with care, Degenerate, unless th' industrious hand Did yearly cull the largest."

But whether in ancient times selection was methodically pursued we may well doubt, when we hear how laborious the work has been found by Le Coutour and Hallett. Although the principle of selection is so important, yet the little which man has effected, by incessant efforts (9/47. A. De Candolle 'Geograph. Bot.' page 932.) during thousands of years, in rendering the plants more productive or the grains more nutritious than they were in the time of the old Egyptians, would seem to speak strongly against its efficacy. But we must not forget that at each successive period the state of agriculture and the quantity of manure supplied to the land will have determined the maximum degree of productiveness; for it would be impossible to cultivate a highly productive variety, unless the land contained a sufficient supply of the necessary chemical elements.

We now know that man was sufficiently civilised to cultivate the ground at an immensely remote period; so that wheat might have been improved long ago up to that standard of excellence which was possible under the then existing state of agriculture. One small class of facts supports this view of the slow and gradual improvement of our cereals. In the most ancient lake-habitations of Switzerland, when men employed only flint-tools, the most extensively cultivated wheat was a peculiar kind, with remarkably small ears and grains. (9/48. O. Heer 'Die Pflanzen der Pfahlbauten' 1866. The following passage is quoted from Dr. Christ in 'Die Fauna der Pfahlbauten, von Dr. Rutimeyer' 1861 s. 225.) "Whilst the grains of the modern forms are in section from seven to eight millimetres in length, the larger grains from the lake-habitations are six, seldom seven, and the smaller ones only four. The ear is thus much narrower, and the spikelets stand out more horizontally, than in our present forms." So again with barley, the most ancient and most extensively cultivated kind had small ears, and the grains were "smaller, shorter, and nearer to each other, than in that now grown; without the husk they were 2 1/2 lines long, and scarcely 1 1/2 broad, whilst those now grown have a length of three lines, and almost the same in breadth." (9/49. Heer as quoted by Carl Vogt 'Lectures on Man' English translation page 355.) These small-grained varieties of wheat and barley are believed by Heer to be the parent-forms of certain existing allied varieties, which have supplanted their early progenitors.

Heer gives an interesting account of the first appearance and final disappearance of the several plants which were cultivated in greater or less abundance in Switzerland during former successive periods, and which generally differed more or less from our existing varieties. The peculiar small-eared and small-grained wheat, already alluded to, was the commonest kind during the Stone period; it lasted down to the Helvetico-Roman age, and then became extinct. A second kind was rare at first, but afterwards became more frequent. A third, the Egyptian wheat (T. turgidum), does not agree exactly with any existing variety, and was rare during the Stone period. A fourth kind (T. dicoccum) differs from all known varieties of this form. A fifth kind (T. monococcum) is known to have existed during the Stone period only by the presence of a single ear. A sixth kind, the common T. spelta, was not introduced into Switzerland until the Bronze age. Of barley, besides the short-eared and small-grained kind, two others were cultivated, one of which was very scarce, and resembled our present common H. distichum. During the Bronze age rye and oats were introduced; the oat- grains being somewhat smaller than those produced by our existing varieties. The poppy was largely cultivated during the Stone period, probably for its oil; but the variety which then existed is not now known. A peculiar pea with small seeds lasted from the Stone to the Bronze age, and then became extinct; whilst a peculiar bean, likewise having small seeds, came in at the Bronze period and lasted to the time of the Romans. These details sound like the descriptions given by palaeontologists of the first appearance, the increasing rarity, and final extinction or modification of fossil species, embedded in the successive stages of a geological formation.

Finally, every one must judge for himself whether it is more probable that the several forms of wheat, barley, rye, and oats are descended from between ten and fifteen species, most of which are now either unknown or extinct, or whether they are descended from between four and eight species, which may have either closely resembled our present cultivated forms, or have been so widely different as to escape identification. In this latter case we must conclude that man cultivated the cereals at an enormously remote period, and that he formerly practised some degree of selection, which in itself is not improbable. We may, perhaps, further believe that, when wheat was first cultivated the ears and grains increased quickly in size, in the same manner as the roots of the wild carrot and parsnip are known to increase quickly in bulk under cultivation.

[MAIZE OR INDIAN CORN: Zea mays.

Botanists are nearly unanimous that all the cultivated kinds belong to the same species. It is undoubtedly (9/50. See Alph. De Candolle's long discussion in his 'Geograph. Bot.' page 942. With respect to New England see Silliman's 'American Journal' volume 44 page 99.) of American origin, and was grown by the aborigines throughout the continent from New England to Chili. Its cultivation must have been extremely ancient, for Tschudi (9/51. 'Travels in Peru' English translation page 177.) describes two kinds, now extinct or not known in Peru, which were taken from tombs apparently prior to the dynasty of the Incas. 'But there is even stronger evidence of antiquity, for I found on the coast of Peru (9/52. 'Geolog. Observ. on S. America' 1846 page 49.) heads of maize, together with eighteen species of recent sea-shell, embedded in a beach which had been upraised at least 85 feet above the level of the sea. In accordance with this ancient cultivation, numerous American varieties have arisen. The aboriginal form has not as yet been discovered in the wild state. A peculiar kind (9/53. This maize is figured in Bonafous' magnificent work, 'Hist. Nat. du Mais' 1836 P1. v. bis, and in the 'Journal of Hort. Soc.'