Species and Varieties, Their Origin by Mutation Part 5

[92]

LECTURE IV

SELECTION OF ELEMENTARY SPECIES

The improvement of cultivated plants must obviously begin with already existing forms. This is true of old cultivated sorts as well as for recent introductions. In either case the starting-point is as important as the improvement, or rather the results depend in a far higher degree on the adequate choice of the initial material than on the methodical and careful treatment of the chosen varieties. This however, has not always been appreciated as it deserves, nor is its importance at present universally recognized. The method of selecting plants for the improvement of the race was discovered by Louis Vilmorin about the middle of the last century. Before his time selection was applied to domestic animals, but Vilmorin was the first to apply this principle to plants. As is well known, he used this method to increase the amount of sugar in beets and thus to raise their value as forage-crops, with such success, that his plants have since been used for the production [93] of sugar. He must have made some choice among the numerous available sorts of beets, or chance must have placed in his hands one of the most appropriate forms. On this point however, no evidence is at hand.

Since the work of Vilmorin the selection-principle has increased enormously in importance, for practical purposes as well as for the theoretical aspect of the subject. It is now being applied on a large scale to nearly all ornamental plants. It is the one great principle now in universal practice as well as one of preeminent scientific value. Of course, the main arguments of the evolution theory rest upon morphologic, systematic, geographic and paleontologic evidence. But the question as to how we can coordinate the relation between existing species and their supposed ancestors is of course one of a physiologic nature. Direct observation or experiments were not available for Darwin and so he found himself constrained to make use of the experience of breeders. This he did on a broad scale, and with such success that it was precisely this side of his arguments that played the major part in convincing his contemporaries.

The work of the breeders previous to Darwin's time had not been very critically performed. Recent a.n.a.lyses of the evidence obtained [94] from them show that numerous types of variability were usually thrown together. What type in each case afforded the material, which the breeder in reality made use of, has only been inquired into in the last few decades. Among those who have opened the way for thorough and more scientific treatment are to be mentioned Rimpau and Von Rumker of Germany and W.M. Hays of America.

Von Rumker is to be considered as the first writer, who sharply distinguished between two phases of methodical breeding-selection. One side he calls the production of new forms, the other the improvement of the breed. He dealt with both methods extensively. New forms are considered as spontaneous variations occurring or originating without human aid. They have only to be selected and isolated, and their progeny at once yields a constant and pure race. This race retains its character as long as it is protected against the admixture of other minor varieties, either by cross-pollination, or by accidental seeds.

Improvement, on the other hand, is the work of man. New varieties of course can only be isolated if chance offers them; the improvement is not inc.u.mbent on chance. It does not create really anything new, but develops characters, which were already existing. It brings [95] the race above its average, and must guard constantly against the regression towards this average which usually takes place.

Hays has repeatedly insisted upon the principle of the choice of the most favorable varieties as the foundation for all experiments in improving races. He a.s.serts that half the battle is won by choosing the variety which is to serve as a foundation stock, while the other half depends upon the selection of parent-plants within the chosen variety.

Thus the choice of the variety is the first principle to be applied in every single case; the so-called artificial selection takes only a secondary place. Calling all minor units within the botanic species by the common name of varieties, without regard to the distinction between elementary species and retrograde varieties, the principle is designated by the term of "variety-testing." This testing of varieties is now, as is universally known, one of the most important lines of work of the agricultural experiment stations. Every state and every region, in some instances even the larger farms, require a separate variety of corn, or wheat, or other crops. They must be segregated from among the hundreds of generally cultivated forms, within each single botanic species. Once found, the type may be ameliorated according to the local conditions [96] and needs, and this is a question of improvement.

The fact that our cultivated plants are commonly mixtures of different sorts, has not always been known. The first to recognize it seems to have been the Spanish professor of botany, Mariano Lagasca, who published a number of Spanish papers dealing with useful plants and botanical subjects between 1810 and 1830, among them a catalogue of plants cultivated in the Madrid Botanical Garden. Once when he was on a visit to Colonel Le Couteur on his farm in Jersey, one of the Channel Islands off the coast of France, in discussing the value of the fields of wheat, he pointed out to his host, that they were not really pure and uniform, as was thought at that time, and suggested the idea that some of the const.i.tuents might form a larger part in the harvest than others.

In a single field he succeeded in distinguis.h.i.+ng no less than 23 varieties, all growing together. Colonel Le Couteur took the hint, and saved the seeds of a single plant of each supposed variety separately.

These he cultivated and multiplied till he got large lots of each and could compare their value. From among them he then chose the variety producing the greatest amount of the finest, whitest and most nutritious flour. This he eventually placed in the [97] market under the name of "Talavera de Bellevue." It is a tall, white variety, with long and slender white heads, almost without awns, and with fine white pointed kernels. It was introduced into commerce about 1830, and is still one of the most generally cultivated French wheats. It was highly prized in the magnificent collection of drawings and descriptions of wheats, published by Vilmorin under the t.i.tle "Les meilleurs bles" and is said to have quite a number of valuable qualities, branching freely and producing an abundance of good grain and straw. It is however, sensitive to cold winters in some degree and thereby limited in its distribution. Hallett, the celebrated English wheat-breeder, tried in vain to improve the peculiar qualities of this valuable production of Le Couteur's.

Le Couteur worked during many years along this line, long before the time when Vilmorin conceived the idea of improvement by race selections, and he used only the simple principle of distinguis.h.i.+ng and isolating the members of his different fields. Later he published his results in a work on the varieties, peculiarities and cla.s.sification of wheat (1843), which though now very rare, has been the basis and origin of the principle of variety-testing.

The discovery of Lagasca and Le Couteur was [98] of course not applicable to the wheat of Jersey alone. The common cultivated sorts of wheat and other grains were mixtures then as they are even now. Improved varieties are, or at least should be, in most cases pure and uniform, but ordinary sorts, as a rule, are mixtures. Wheat, barley and oats are self-fertile and do not mix in the field through cross-pollination.

Every member of the a.s.semblage propagates itself, and is only checked by its own greater or less adaptation to the given conditions of life.

Rimpau has dealt at large with the phenomenon as it occurs in the northern and middle parts of Germany. Even Rivett's "Bearded wheat,"

which was introduced from England as a fine improved variety, and has become widely distributed throughout Germany, cannot keep itself pure.

It is found mingled almost anywhere with the old local varieties, which it was destined to supplant. Any lot of seed exhibits such impurities, as I have had the opportunity of observing myself in sowings in the experimental-garden. But the impurities are only mixtures, and all the plants of Rivett's "Bearded wheat," which of course const.i.tute the large majority, are of pure blood. This may be confirmed when the seeds are collected and sown separately in cultures that can be carefully guarded.

[99] In order to get a closer insight into the causes of this confused condition of ordinary races, Rimpau made some observations on Rivett's wheat. He found that it suffers from frost during winter more than the local German varieties, and that from various causes, alien seeds may accidentally, and not rarely, become mixed with it. The thres.h.i.+ng-machines are not always as clean as they should be and may be the cause of an accidental mixture. The manure comes from stables, where straw and the dust from many varieties are thrown together, and consequently living kernels may become mixed with the dung. Such stray grains will easily germinate in the fields, where they find more congenial conditions than does the improved variety. If winter arrives and kills quant.i.ties of this latter, the accidental local races will find ample s.p.a.ce to develop. Once started, they will be able to multiply so rapidly, that in one or two following generations they will const.i.tute a very considerable portion of the whole harvest. In this way the awnless German wheat often prevails over the introduced English variety, if the latter is not kept pure by continuous selection.

The Swiss wheat-breeder Risler made an experiment which goes to prove the certainty of the explanation given by Rimpau. He observed on his farm at Saleves near the lake of Geneva that after a lapse of time the "Galland wheat" deteriorated and a.s.sumed, as was generally believed, the characters of the local sorts. In order to ascertain the real cause of this apparent change, he sowed in alternate rows in a field, the "Galland" and one of the local varieties. The "Galland" is a race with obvious characters and was easily distinguished from the other at the time when the heads were ripe. They are bearded when flowering, but afterwards throw off the awns. The kernels are very large and yield an extraordinarily good, white flour.

During the first summer all the heads of the "Galland" rows had the deciduous awns but the following year these were only seen on half of the plants, the remainder having smooth heads, and the third year the "Galland" had nearly disappeared, being supplanted by the competing local race. The cause of this rapid change was found to be twofold.

First the "Galland," as an improved variety, suffers from the winter in a far higher degree than the native Swiss sorts, and secondly it ripens its kernels one or two weeks later. At the time of harvest it may not have become fully ripe, while the varieties mixed with it had reached maturity. The wild oat, _Avena fatua_, is very common in [101] Europe from whence it has been introduced in the United States. In summers which are unfavorable to the development of the cultivated oats it may be observed to multiply with an almost incredible rapidity. It does not contribute to the harvest, and is quite useless. If no selection were made, or if selection were discontinued, it would readily supplant the cultivated varieties.

From these several observations and experiments it may be seen, that it is not at all easy to keep the common varieties of cereals pure and that even the best are subject to the encroachment of impurities. Hence it is only natural that races of cereals, when cultivated without the utmost care, or even when selected without an exact knowledge of their single const.i.tuents, are always observed to be more or less in a mixed condition. Here, as everywhere with cultivated and wild plants, the systematic species consist of a number of minor types, which pertain to different countries and climates, and are growing together in the same climate and under the same external conditions. They do not mingle, nor are their differentiating characters destroyed by intercrossing. They each remain pure, and may be isolated whenever and wherever the desirability for such a proceeding should arise. The purity of [102] the races is a condition implanted in them by man, and nature always strives against this arbitrary and one-sided improvement. Numerous slight differences in characters and numerous external influences benefit the minor types and bring them into compet.i.tion with the better ones.

Sometimes they tend to supplant the latter wholly, but ordinarily sooner or later a state of equilibrium is reached, in which henceforth the different sorts may live together. Some are favored by warm and others by cool summers, some are injured by hard winters while others thrive then and are therefore relatively at an advantage. The mixed condition is the rule, purity is the exception.

Different sorts of cereals are not always easily distinguishable by the layman and therefore I will draw your attention to conditions in meadows, where a corresponding phenomenon can be observed in a much simpler way.

Only artificial pasture-grounds are seen to consist of a single species of gra.s.s or clover. The natural condition in meadows is the occurrence of clumps of gra.s.ses and some clovers, mixed up with perhaps twenty or more species of other genera and families. The numerical proportion of these const.i.tuents is of great interest, and has been studied at Rothamstead in England and on a number of other farms. It is [103]

always changing. No two successive years show exactly the same proportions. At one time one species prevails, at another time one or two or more other species. The weather during the spring and summer benefits some and hurts others, the winter may be too cold for some, but again harmless for others, the rainfall may partly drown some species, while others remain uninjured. Some weeds may be seen flowering profusely during some years, while in other summers they are scarcely to be found in the same meadow. The whole population is in a fluctuating state, some thriving and others deteriorating. It is a continuous response to the ever changing conditions of the weather. Rarely a species is wholly annihilated, though it may apparently be so for years; but either from seeds or from rootstocks, or even from neighboring lands, it may sooner or later regain its foothold in the general struggle for life.

This phenomenon is a very curious and interesting one. The struggle for life, which plays so considerable a part in the modern theories of evolution, may be seen directly at work. It does not alter the species themselves, as is commonly supposed, but it is always changing their numerical proportion. Any lasting change in the external conditions will of course alter the average oscillation and the influence [104] of such alterations will manifest itself in most cases simply in new numerical proportions. Only extremes have extreme effects, and the chance for the weaker sorts to be completely overthrown is therefore very small.

Any one, who has the opportunity of observing a waste field during a series of years, should make notes concerning the numerical proportions of its inhabitants. Exact figures are not at all required; approximate estimates will ordinarily prove to be sufficient, if only the standard remains the same during the succeeding years.

The entire ma.s.s of historic evidence goes to prove that the same conditions have always prevailed, from the very beginning of cultivation up to the present time. The origin of the cultivation of cereals is to be sought in central Asia. The recent researches of Solms Laubach show it to be highly probable that the historic origin of the wheat cultivated in China, is the same as that of the wheat of Egypt and Europe. Remains of cereals are found in the graves of Egyptian mummies, in the mounds of waste material of the lake-dwellings of Central Europe, and figures of cereals are to be seen on old Roman coins. In the sepulchre of King Ra-n-Woser of the Fifth Dynasty of Egypt, who lived about 2000 years B.C., two [105] tombs have recently been opened by the German Oriental Society. In them were found quant.i.ties of the tares of the _Tritic.u.m dicocc.u.m_, one of the more primitive forms of wheat. In other temples and pyramids and among the stones of the walls of Dashur and El Kab studied by Unger, different species and varieties of cereals were discovered in large quant.i.ties, that showed their ident.i.ty with the present prevailing cultivated races of Egypt.

The inhabitants of the lake-dwellings in Switzerland possessed some varieties of cereals, which have entirely disappeared. They are distinguished by Heer under special names. The small barley and the small wheat of the lake-dwellers are among them. All in all there were ten well distinguished varieties of cereals, the Panic.u.m and the Setaria or millet being of the number. Oats were evidently introduced only toward the very last of the lake-dwelling period, and rye is of far later introduction into western Europe. Similar results are attained by the examination of the cereals figured by the Romans of the same period.

All these are archaeologic facts, and give but slight indications concerning the methods of cultivation or the real condition of the cultivated races of that time. Virgil has left us some knowledge of the requirements of methodical [106] culture of cereals of his time. In his poem _Georgics_ (I. 197) the following lines are found:

_Vidi lecta din, et multo spectata labore Degenerare tamen, ni vis humana quotannis Maxima quaeque manu legeret_.

(The chosen seed, through years and labor improved, Was seen to run back, unless yearly Man selected by hand the largest and fullest of ears.)

Elsewhere Virgil and also some lines of Columella and Varro go to prove in the same way that selection was applied by the Romans to their cereals, and that it was absolutely necessary to keep their races pure.

There is little doubt, but that it was the same principle as that which has led, after many centuries, to the complete isolation and improvement of the very best races of the mixed varieties. It further proves that the mixed conditions of the cereals was known to man at that time, although distinct ideas of specific marks and differences were of course still wholly lacking. It is proof also that cultivated cereals from the earliest times must have been built up of numerous elementary forms.

Moreover it is very probable, that in the lapse of centuries a goodly number of such types must have disappeared. [107] Among the vanished forms are the special barley and wheat of the lake-dwellings, the remains of which have been accidentally preserved, but most of the forms must have disappeared without leaving any trace.

This inference is supported by the researches of Solms-Laubach, who found that in Abyssinia numerous primitive types of cereals are still in culture. They are not adequate to compete with our present varieties, and would no doubt also have disappeared, had they not been preserved by such quite accidental and almost primitive isolation.

Closing this somewhat long digression into history we will now resume our discussion concerning the origin of the method of selecting cereals for isolation and segregate-cultivation. Some decades after Le Couteur, this method was taken up by the celebrated breeder Patrick Sheriff of Haddington in Scotland. His belief, which was general at that time, was "That cultivation has not been found to change well defined kinds, and that improvement can be best attained by selecting new and superior varieties, which nature occasionally produces, as if inviting the husbandman to stretch forth his hand and cultivate them."

Before going into the details of Sheriff's work it is as well to say something concerning [108] the use of the word "selection." This word was used by Sheriff as seen in the quotation given, and it was obviously designed to convey the same idea as the word "lecta" in the quotation from Virgil. It was a choice of the best plants from among known mixed fields, but the chosen individuals were considered to be representatives of pure and constant races, which could only be isolated, but not ameliorated. Selection therefore, in the primitive sense of the word, is the choice of elementary species and varieties, with no other purpose than that of keeping them as pure as possible from the admixture of minor sorts. The Romans attained this end only imperfectly, simply because the laws governing the struggle for life and the compet.i.tion of numerous sorts in the fields were unsuspected by them.

Le Couteur and Sheriff succeeded in the solution of the problem, because they had discovered the importance of isolation. The combination of a careful choice with subsequent isolation was all they knew about it, and it was one of the great achievements to which modern agriculture owes its success.

The other great principle was that of Vilmorin. It was the improvement within the race, or the "amelioration of the race" as it was termed by him. It was introduced into [109] England by F.F. Hallett of Brighton in Suss.e.x, who at once called it "pedigree-culture," and produced his first new variety under the very name of "Pedigree-wheat." This principle, which yields improved strains, that are not constant but dependent on the continued and careful choice of the best plants in each succeeding generation, is now generally called "selection." But it should always be remembered that according to the historic evolution of the idea, the word has the double significance of the distinction and isolation of constant races from mixtures, and that of the choice of the best representatives of a race during all the years of its existence. Even sugar-beets, the oldest "selected" agricultural plants, are far from having freed themselves from the necessity of continuous improvement.

Without this they would not remain constant, but would retrograde with great rapidity.

The double meaning of the word selection still prevailed when Darwin published his "Origin of Species." This was in the year 1859, and at that time s.h.i.+rreff was the highest authority and the most successful breeder of cereals. Vilmorin's method had been applied only to beets, and Hallett had commenced his pedigree-cultures only a few years before and his first publication of the "Pedigree-wheat" [110] appeared some years later at the International Exhibition of London in 1862. Hence, whenever Darwin speaks of selection, s.h.i.+rreff's use of the word may as well be meant as that of Vilmorin.

However, before going deeper into such theoretical questions, we will first consider the facts, as given by s.h.i.+rreff himself.

During the best part of his life, in fact during the largest part of the first half of the nineteenth century, s.h.i.+rreff worked according to a very simple principle. When quite young he had noticed that sometimes single plants having better qualities than the average were seen in the fields. He saved the grains, or sometimes the whole heads of such plants separately, and tried to multiply them in such manner as to avoid intermixtures.

His first result was the "Mungoswell's wheat." In the spring of 1819 he observed quite accidentally in a field of the farm of that name, a single plant which attracted his attention by a deeper green and by being more heavily headed out. Without going into further details, he at once chose this specimen as the starting point of a new race. He destroyed the surrounding plants so as to give it more s.p.a.ce, applied manure to its roots, and tended it with special care. It yielded 63 heads and nearly [111] 2500 grains. All of these were sown the following fall, and likewise in the succeeding years the whole harvest was sown in separate lots. After two years of rapid multiplication it proved to be a good new variety and was brought into commerce. It has become one of the prominent varieties of wheat in East Lothian, that county of Scotland of which Haddington is the princ.i.p.al borough.

The grains of "Mungoswell's wheat" are whiter than those of the allied "Hunter's wheat," more rounded but otherwise of the same size acid weight. The straw is taller and stronger, and each plant produces more culms and more heads.

s.h.i.+rreff a.s.sumed, that the original plant of this variety was a sport from the race in which he had found it, and that it was the only instance of this sport. He gives no details about this most interesting side of the question, omitting even to tell the name of the parent variety. He only a.s.serts that it was seen to be better, and afterwards proved so by the appreciation of other breeders and its success in trade. He observed it to be quite constant from the beginning, no subsequent selection being needed. This important feature was simply a.s.sumed by him to be true as a matter of course.

[112] Some years afterwards, in the summer of 1824, he observed a large specimen of oats in one of the fields of the same farm. Being at that time occupied in making a standard collection of oats for a closer comparison of the varieties, he saved the seeds of that plant and sowed them in a row in his experiment-field. It yielded the largest culms of the whole collection and bore long and heavy kernels with a red streak on the concave side and it excelled all other sorts by the fine qualities of its very white meal. In the unequal length of its stalks it has however a drawback, as the field appears thinner and more meager than it is in reality. "Hopetown oats," as it is called, has found its way into culture extensively in Scotland and has even been introduced with success into England, Denmark and the United States. It has been one of the best Scottish oats for more than half a century.

The next eight years no single plant judged worthy of selection on his own farm attracted s.h.i.+rreff's attention. But in the fall of 1832 he saw a beautiful plant of wheat on a neighboring farm and he secured a head of it with about 100 grains. From this he produced the "Hopetown wheat."

After careful separation from the kernels this original ear was preserved, and was afterwards exhibited at the Stirling Agricultural [113] Museum. The "Hopetown wheat" has proved to be a constant variety, excelling the ordinary "Hunter's wheat" by larger grains and longer heads; it yields likewise a straw of superior quality and has become quite popular in large districts of England and Scotland, where it is known by the name of "White Hunter's" from its origin and the brilliant whiteness of its heads.

In the same way s.h.i.+rreff's oats were discovered in a single plant in a field where it was isolated in order to be brought into commerce after multiplication. It has won the surname of "Make-him-rich." Nothing is on record about the details of its origin.

Four valuable new varieties of wheat and oats were obtained in this way in less than forty years. Then s.h.i.+rreff changed his ideas and his method of working. Striking specimens appeared to be too rare, and the expectation of a profitable result too small. Therefore he began work on a larger scale. He sought and selected during the summer of 1857 seventy heads of wheat, each from a single plant showing some marked and presumably favorable peculiarity. These were not gathered on one field, but were brought together from all the fields to which he had access in his vicinity. The grains of each of these selected heads were [114] sown separately, and the lots compared during their whole life-period and chiefly at harvest time. Three of the lots were judged of high excellence, and they alone were propagated, and proving to be constant new varieties from the outset were given to the trade under the names of "s.h.i.+rreff's bearded white," "s.h.i.+rreff's bearded red," and "Pringle's wheat." They have found wide acceptance, and the first two of them are still considered by Vilmorin as belonging to the best wheats of France.

This second method of s.h.i.+rreff evidently is quite a.n.a.logous to the principle of Lagasca and Le Couteur. The previous a.s.sumption that new varieties with striking features were being produced by nature from time to time, was abandoned, and a systematic inquiry into the worth of all the divergent const.i.tuents of the fields was begun. Every single ear at once proved to belong to a constant and pure race, but most of these were only of average value. Some few however, excelled to a degree, which made them worth multiplying, and to be introduced into trade as separate varieties.