Chapter 42

Besides this limited study, we may compare the numerous individuals of one locality or of a large plot of cultivated plants with one another. In doing so, we are struck with the fact that some plants have large and others small fruits. We now limit ourselves to the main spike of each plant, and perhaps to its lower parts, so as to avoid as far as possible the impression made by the partial fluctuations. The differences remain, and are sufficient to furnish an easy comparison with the general law. In order to do this, we take from each plant a definite number of capsules and measure their average length. In some experiments I took the twenty lowermost capsules of the main spikes. In this way one average was obtained for each plant, and combining these into a curve, it was found that these fluctuations also came under Quetelet's law. Thus the individual averages, and the fluctuations around each of them, follow the same rule. The first are a measure for the whole plant, the second only for its parts. As a general resume we can assert that, as a rule, a quality is determined in some degree during the earlier stages of the organism, and that this determination is valid throughout its [749] life. Afterwards only the minor points remain to be regulated. This makes it at once clear that the range of individual and partial variability together must be wider than that of either of them, taken alone. Partial fluctuations cannot, of course, be excluded. Thus our comparison is limited to individual and partial variability on one side, and partial fluctuations alone on the other side.

Intra-specific selection is thus seen to fall under two heads: a selection between the individuals, and a choice within each of them. The first affords a wider and the latter a narrower field.

Individual variability, considered as the result of outward influences operative during extreme youth, can be excluded in a very simple manner. Obviously it suffices to exclude extreme youth, in other words, to exclude the use of seeds. Multiplication in a vegetative way, by grafting and budding, by runners or roots, or by simple division of rootstocks and bulbs is the way in which to limit variability to the partial half. This is all we may hope to attain, but experience shows that it is a very efficient means of limitation. Partial fluctuations are generally far smaller than individual and partial fluctuations together.

Individual variability in the vegetable kingdom [750] might be called seed-variation, as opposed to partial or bud-fluctuation. And perhaps these terms are more apt to convey a clear conception of the distinction than any other. The germ within the unripe seed is easily understood to be far more sensitive to external conditions than a bud.

Multiplication of extremes by seed is thus always counteracted by individual variability, which at once reopens all, or nearly all, the initial possibilities. Multiplication by buds is exempt from this danger and thus leads to a high degree of uniformity. And this uniformity is in many cases exactly what the breeder endeavors to obtain.

We will treat of this reopening of previous possibilities under the head of regression in the next lecture. It is not at all absolute, at least not in one generation. Part of the improvement remains, and favors the next generation. This part may be estimated approximately as being about one-third or one-half of the improvement attained. Hence the conclusion that vegetative multiplication gives rise to varieties which are as a rule twice or thrice as good as selected varieties of plants propagated by seeds. Hence, likewise the inference that breeders generally prefer vegetative multiplication of improved forms, and apply it in all possible cases. [751] Of course the application is limited, and forage crops and the greater number of vegetables will always necessarily be propagated by seed.

Nature ordinarily prefers the sexual way. Asexual multiplications, although very common with perennial plants, appear not to offer important material for selection. Hence it follows that in comparing the work of nature with that of man, the results of selection followed by vegetative propagation should always be carefully excluded. Our large bulb-flowers and delicious fruits have nothing in common with natural products, and do not yield a standard by which to judge nature's work.

It is very difficult for a botanist to give a survey of what practice has attained by the asexual multiplication of extremes. Nearly all of the large and more palatable fruits are due to such efforts. Some flowers and garden-plants afford further instances. By far the greatest majority of improved asexual varieties, however, are not the result of pure intra-specific selection. They are due largely to the choice of the best existing elementary species, and to some extent to crosses between them, or between distinct systematic species. In practice selection and hybridization go hand in hand and it is often difficult to ascertain what part of [752] the result is due to the one, and what to the other factor.

The scientist, on the contrary, has nothing to do with the industrial product. His task is the analysis of the methods, in order to reach a clear appreciation of the influence of all the competing factors. This study of the working causes leads to a better understanding of the practical processes, and may become the basis of improvement in methods.

Starting from these considerations, we will now give some illustrative examples, and for the first, choose one in which hybridization is almost completely excluded.

Sugar-canes have long been considered to be plants without seed. Their numerous varieties are propagated only in a vegetative way. The stems are cut into pieces, each bearing one or two or more nodes with their buds. An entire variety, though it may be cultivated in large districts and even in various countries, behaves with respect to variability as a single individual. Its individual fluctuability has been limited to the earliest period of its life, when it arose from an unknown seed. The personal characters that have been stamped on this one seed, partly by its descent, and partly in the development of its germ during the period of ripening, have become the indelible characters [753] of the variety, and only the partial fluctuability, due to the effect of later influences, can now be studied statistically.

This study has for its main object the production of sugar in the stems, and the curves, which indicate the percentage of this important substance in different stems of the same variety, comply with Quetelet's law. Each variety has its own average, and around this the data of the majority of the stems are densely crowded, while deviations on both sides are rare and become the rarer the wider they are. The "Cheribon" cane is the richest variety cultivated in Java, and has an average of 19% sugar, while it fluctuates between 11% and 28%. "Chunnic" averages 14%, "Black Manilla" 13% and "White Manilla" 10%; their highest and lowest extremes diverge in the same manner, being for the last named variety 1% and 15%.

This partial variability is of high practical interest, because on it a selection may be founded. According to the conceptions described in a previous lecture, fluctuating variability is the result of those outward factors that determine the strength of development of the plant or the organ. The inconstancy of the degree of sensibility, combined with the ever-varying weather conditions preclude any close proportionality, but apart from this difficulty there is, in the [754] main, a distinct relation between organic strength and the development of single qualities. This correlation has not escaped observation in the case of the sugar-cane, and it is known that the best grown stocks are generally the richest in sugar. Now it is evident that the best grown and richest stems will have the greater chance of transmitting these qualities to the lateral-buds. This at once gives, a basis for vegetative selection, upon which it is not necessary to choose a small number of very excellent stems, but simply to avoid the planting of all those that are below the average. By this means the yield of the cultures has often noticeably been enhanced.

As far as experience goes, this sort of selection, however profitable, does not conduce to the production of improved races. Only temporary ameliorations are obtained, and the selection must be made in the same manner every year. Moreover the improvement is very limited and does not give any promise of further increase. In order to reach this, one has to recur to the individual fluctuability, and therefore to seed.

Nearly half a century ago, Parris discovered, on the island of Barbados, that seeds might occasionally be gathered from the canes. These, however, yielded only grass-like plants of no real value. The same observation was made [755] shortly afterwards in Java and in other sugar producing countries. In the year 1885, Soltwedel, the director of one of the experiment stations for the culture of sugar-cane in Java, conceived the idea of making use of seedlings for the production of improved races. This idea is a very practical one, precisely because of the possibility of vegetative propagation. If individuals would show the same range as that of partial fluctuability, then the choice of the extremes would at once bring the average up to the richness of the best stocks. Once attained, this average would be fixed, without further efforts.

Unfortunately there is one great drawback. This is the infertility of the best variety, that of the "Cheribon" cane. It flowers abundantly in some years, but it has never been known to produce ripe seeds. For this reason Soltwedel had to start from the second best sort, and chose the "Hawaii" cane. This variety usually yields about 14% sugar, and Soltwedel found among his seedlings one that showed 15%. This fact was quite unexpected at that time, and excited widespread interest in the new method, and since then it has been applied to numerous varieties, and many thousands of seedlings have been raised and tested as to their sugar-production.

[756] From a scientific point of view the results are quite striking. From the practical standpoint, however, the question is, whether the "Hawaii" and other fertile varieties are adequate to yield seedlings, which will surpass the infertile "Cheribon" cane. Now "Hawaii" averages 14% and "Cheribon" 19%, and it is easily understood that a "Hawaii" seedling with more than 19% can be expected only from very large sowings. Hundreds of thousands of seedlings must be cultivated, and their juice tested, before this improvement can be reached. Even then, it may have no significance for practical purposes. Next to the amount of sugar comes the resistance to the disease called "Sereh," and the new race requires to be ameliorated in this important direction, too. Other qualities must also be considered, and any casual deterioration in other characters would make all progress illusory. For these reasons much time is required to attain distinct improvements.

These great difficulties in the way of selecting extremes for vegetative propagation are of course met with everywhere. They impede the work of the breeder to such a degree, that but few men are able to surmount them. Breeding new varieties necessitates the bending of every effort to this purpose, and a clear conception of [757] the manifold aspects of this intricate problem. These fall under two heads, the exigencies of practice, and the physiologic laws of variability. Of course, only the latter heading comes within the limits of our discussion which includes two main points. First comes the general law of fluctuation that, though slight deviations from the average may be found by thousands, or rather in nearly every individual, larger and therefore important deviations are very rare. Thousands of seedlings must be examined carefully in order to find one or two from which it might be profitable to start a new race. This point is the same for practical and for scientific investigation. In the second place however, a digression is met with. The practical man must take into consideration all the varying qualities of his improved strains. Some of them must be increased and others be decreased, and their common dependency on external conditions often makes it very difficult to discover the desired combinations. It is obvious, however, that the neglect of one quality may make all improvement of other characters wholly useless. No augmentation of sugar-percentage, of size and flavor of fruits can counterbalance an increase in sensitiveness to disease, and so it is with other qualities also.

[758] Improved races for scientific investigation can be kept free from infection, and protected against numerous other injuries. In the experimental garden they may find conditions which cannot be realized elsewhere. They may show a luxuriant growth, and prove to be excellent material for research, but have features which, having been overlooked at the period of selection, would at once condemn them if left to ordinary conditions, or to the competition of other species.

Considering all these obstacles, it is only natural that breeders should use every means to reach their goal. Only in very rare instances do they follow methods analogous to scientific processes, which tend to simplify the questions as much as possible. As a rule, the practical way is the combination of as many causes of variability as possible. Now the three great sources of variability are, as has been pointed out on several occasions, the original multiformity of the species, fluctuating variability, and hybridization. Hence, in practical experiments, all three are combined. Together they yield results of the highest value, and Burbank's improved fruits and flowers give testimony to the practical significance of this combination.

From a scientific point of view however, it is [759] ordinarily difficult, if not impossible, to discern the part which each of the three great branches of variability has taken in the origination of the product. A full analysis is rarely possible, and the treatment of one of the three factors must necessarily remain incomplete.

Notwithstanding these considerations, I will now give some examples in order to show that fluctuating variability plays a prominent part in these improvements. Of course it is the third in importance in the series. First comes the choice of the material from the assemblage of species, elementary species and varieties. Hybridization comes next in importance. But even the hybrids of the best parents may be improved, because they are no less subject to Quetelet's law than any other strain. Any large number of hybrids of the same ancestry will prove this, and often the excellency of a hybrid variety depends chiefly, or at least definitely, on the selection of the best individuals. Being propagated only in a vegetative way, they retain their original good qualities through all further culture and multiplication.

As an illustrative example I will take the genus _Canna_. Originally cultivated for its large and bright foliage only, it has since become a flowering plant of value. Our garden strains have originated by the crossing of [760] a number of introduced wild species, among which the _Canna indica_ is the oldest, now giving its name to the whole group. It has tall stems and spikes with rather inconspicuous flowers with narrow petals. It has been crossed with _C. nepalensis_ and _C. warczewiczii_, and the available historic evidence points to the year 1846 as that of the first cross. This was made by Annee between the _indica_ and the _nepalensis_; it took ten years to multiply them to the required degree for introduction into commerce. These first hybrids had bright foliage and were tall plants, but their flowers were by no means remarkable.

Once begun, hybridization was widely practiced. About the year 1889 Crozy exhibited at Paris the first beautifully flowering form, which he named for his wife, "Madame Crozy." Since that time he and many others, have improved the flowers in the shape and size, as well as in color and its patterns. In the main, these ameliorations have been due to the discovery and introduction of new wild species possessing the required characters. This is illustrated by the following incident. In the year 1892 I visited Mr. Crozy at Lyons. He showed me his nursery and numerous acquisitions, those of former years as well as those that were quite new, and which were in the process of rapid [761] multiplication, previous to being given to the trade. I wondered, and asked, why no pure white variety was present. His answer was "Because no white species had been found up to the present time, and there is no other means of producing white varieties than by crossing the existing forms with a new white type."

Comparing the varieties produced in successive periods, it is very easy to appreciate their gradual improvement. On most points this is not readily put into words, but the size of the petals can be measured, and the figures may convey at least some idea of the real state of things. Leaving aside the types with small flowers and cultivated exclusively for their foliage, the oldest flowers of _Canna_ had petals of 45 mm. length and 13 mm. breadth. The ordinary types at the time of my visit had reached 61 by 21 mm., and the "Madame Crozy" showed 66 by 30 mm. It had however, already been surpassed by a few commercial varieties, which had the same length but a breadth of 35 mm. And the latest production, which required some years of propagation before being put on the market, measured 83 by 43 mm. Thus in the lapse of some thirty years the length had been doubled and the breadth tripled, giving flowers with broad corollas and with petals [762] joined all around, resembling the best types of lilies and _Amaryllis_.

Striking as this result unquestionably is, it remains doubtful as to what part of it is due to the discovery and introduction of new large flowered species, and what to the selection of the extremes of fluctuating variability. As far as I have been able to ascertain however, and according to the evidence given to me by Mr. Crozy, selection has had the largest part in regard to the size, while the color-patterns are introduced qualities.

The scientific analysis of other intricate examples is still more difficult. To the practical breeder they often seem very simple, but the student of heredity, who wishes to discern the different factors, is often quite puzzled by this apparent simplicity. So it is in the case of the double lilacs, a large number of varieties of which have recently been originated and introduced into commerce by Lemoine of Nancy. In the main they owe their origin to the crossing and recrossing of a single plant of the old double variety with the numerous existing single-flowered sorts.

This double variety seems to be as old as the culture of the lilacs. It was already known to Munting, who described it in the year 1671. Two centuries afterwards, in 1870, a new description [763] was given by Morren, and though more than one varietal name is recorded in his paper, it appears from the facts given that even at that time only one variety existed. It was commonly called _Syringa vulgaris azurea plena_, and seems to have been very rare and without real ornamental value.

Lemoine, however, conceived the desirability of a combination of the doubling with the bright colors and large flower-racemes of other lilacs, and performed a series of crosses. The "_azurea plena_" has no stamens, and therefore must be used in all crosses as the pistil-parent; its ovary is narrowly inclosed in the tube of the flower, and difficult to fertilize. On the other hand, new crosses could be made every year, and the total number of hybrids with different pollen-parents was rapidly increased. After five years the hybrids began to flower and could be used for new crosses, yielding a series of compound hybrids, which however, were not kept separate from the products of the first crosses.

Gradually the number of the flowering specimens increased, and the character of doubling was observed to be variable to a high degree. Sometimes only one supernumerary petal was produced, sometimes a whole new typical corolla was extruded from within the first. In the same [764] way the color and the number of the flowers on each raceme were seen to vary. Thousands of hybrids were produced, and only those which exhibited real advantages were selected for trade. These were multiplied by grafting, and each variety at present consists only of the buds of one original individual and their products. No constancy from seed is assumed, many varieties are even quite sterile.

Of course, no description was given of the rejected forms. It is only stated that many of them bore either single or poorly filled flowers, or were objectionable in some other way. The range of variability, from which the choices were made, is obscure and only the fact of the selection is prominent. What part is due to the combination of the parental features and what to the individual fluctuation of the hybrid itself cannot be ascertained.

So it is in numerous other instances. The dahlias have been derived from three or more original species, and been subjected to cultivation and hybridization in an ever-increasing scale for a century. The best varieties are only propagated in the vegetative way, by the roots and buds, or by grafting and cutting. Each of them is, with regard to its hereditary qualities, only one individual, and the individual characters were selected at the same time with the [765] varietal and hybrid characters. Most of them are very inconstant from seed and as a rule, only mixtures are offered for sale in seed-lists. Which of their ornamental features are due to fluctuating deviation from an average is of course unknown. _Amaryllis_ and _Gladiolus_ are surrounded with the same scientific uncertainties. Eight or ten, or even more, species have been combined into one large and multiform strain, each bringing its peculiar qualities into the mixed mass. Every hybrid variety is one individual, being propagated by bulbs only. Colors and color-patterns, shape of petals and other marks, have been derived from the wild ancestors, but the large size of many of the best varieties is probably due to the selection of the extremes of fluctuating variability. So it is with the begonias of our gardens, which are also composite hybrids, but are usually sown on a very large scale. Flowers of 15 cm. diameter are very showy, but there can be no doubt about the manner in which they are produced, as the wild species fall far short of this size.

Among vegetables the potatoes afford another instance. Originally quite a number of good species were in culture, most of them having small tubers. Our present varieties are due to hybridization and selection, each of them being propagated only in the vegetative way.

[766] Selection is founded upon different qualities, according to the use to be made of the new sort. Potatoes for the factory have even been selected for their amount of starch, and in this case at least, fluctuating variability has played a very important part in the improvement of the race.