Fluctuating variability may be regarded from two different points of view. The multiformity of a bed of flowers is often a desirable feature, and all means which widen the range of fluctuation are therefore used to enhance this feature, and variability affords specimens, which surpass the average, by yielding a better or larger product.
In the case of fruits and other cultivated forms, it is of course profitable to propagate from the better specimens only, and if possible only from the very best. Obviously the best are the extremes of the whole range of diverging forms, and moreover the extremes on one side of the group. Almost always the best for practical purposes is that in which some quality is strengthened. Cases occur however, in which it is desirable to diminish an injurious peculiarity as far as possible, and in these instances the opposite extreme is the most profitable one.
These considerations lead us to a discussion [743] of the results of the choice of extremes, which it may be easily seen is a matter of the greatest practical importance. This choice is generally designated as selection, but as with most of the terms in the domain of variability, the word selection has come to have more than one meaning. Facts have accumulated enormously since the time of Darwin, a more thorough knowledge has brought about distinctions, and divisions at a rapidly increasing rate, with which terminology has not kept pace. Selection includes all kinds of choice. Darwin distinguished between natural and artificial selection, but proper subdivisions of these conceptions are needed.
In the fourth lecture we dealt with this same question, and saw that selection must, in the first place, make a choice between the elementary species of the same systematic form. This selection of species or species-selection was the work of Le Couteur and Patrick Shirreff, and is now in general use in practice where it has received the name of variety-testing. This clear and unequivocal term however, can hardly be included under the head of natural selection. The poetic terminology of selection by nature has already brought about many difficulties that should be avoided in the future. On the other hand, the designation of the process as a natural [744] selection of species complies as closely as possible with existing terminology, and does not seem liable to any misunderstanding.
It is a selection between species. Opposed to it is the selection within the species. Manifestly the first should precede the second, and if this sequence is not conscientiously followed it will result in confusion. This is evident when it is considered that fluctuations can only appear with their pure and normal type in pure strains, and that each admixture of other units is liable to be shown by the form of the curves. More over, selection chooses single individuals, and a single plant, if it is not a hybrid, can scarcely pertain to two different species. The first choice therefore is apt to make the strain pure.
In contrasting selection between species with that within the species, of course elementary species are meant, including varieties. The terms would be of no consequence if only rightly understood. For the sake of clearness we might designate the last named process with the term of intra-specific selection, and it is obvious that this term is applicable both to natural and to artificial selection.
Having previously dealt with species-selection at sufficient length, we may now confine ourselves to the consideration of the intra-specific [745] selection process. In practice it is of secondary importance, and in nature it takes a very subordinate position. For this reason it will be best to confine further discussions to the experience of the breeders.
Two different ways are open to make fluctuating variability profitable. Both consist in the multiplication of the chosen extremes, and this increase may be attained in a vegetative manner, or by the use of seeds. Asexual and sexual propagation are different in many respects, and so they are also in the domain of variability.
In order to obtain a clear comprehension of this difference, it is necessary to start from the distinction between individual and partial fluctuations, as given in the last lecture. This distinction may be discussed more understandingly if the causes of the variability are taken into consideration. We have dealt with them at some length, and are now aware that inner conditions only, determine averages, while some fluctuation around them is allowable, as influenced by external conditions. These outward influences act throughout life. At the very first they impress their stamp on the whole organism, and incite a lasting change in distinct directions. This is the period of the development of the germ within the seed; it begins with the fusion of the sexual cells, and each of them may be influenced [746] to a noticeable degree before this union. This is the period of the determination of individual variability. As soon as ramifications begin, the external conditions act separately on every part, influencing some to a greater and others to a lesser degree. Here we have the beginning of partial variability. At the outset all parts may be affected in the same way and in the same measure, but the chances of such an agreement, of course, rapidly diminish. This is partly due to differences in exposure, but mainly to alterations of the sensibility of the organs themselves.
It is difficult to gain a clear conception of the contrast between individual and partial variability, and neither is it easy to appreciate their cooperation rightly. Perhaps the best way is to consider their activity as a gradual narrowing of possibilities. At the outset the plant may develop its qualities in any measure, nothing being as yet fixed. Gradually however, the development takes a definite direction, for better or for worse. Is a direction once taken, then it becomes the average, around which the remaining possibilities are grouped. The plant or the organ goes on in this way, until finally it reaches maturity with one of the thousands of degrees of development, between which at the beginning it had a free choice.
[747] Putting this discussion in other terms, we find every individual and every organ in the adult state corresponding with a single ordinate of the curve. The curve indicates the range of possibilities, the ordinate shows the choice that has been made. Now it is clear at once that this choice has not been made suddenly but gradually. Halfway of the development, the choice is halfway determined, but the other half is still undefined. The first half is the same for all the organs of the plant, and is therefore termed individual; the second differs in the separate members, and consequently is known as partial. Which of the two halves is the greater and which the lesser, of course depends on the cases considered.
Finally we may describe a single example, the length of the capsules of the evening-primrose. This is highly variable, the longest reaching more than twice the length of the smallest. Many capsules are borne on the same spike, and they are easily seen to be of unequal size. They vary according to their position, the size diminishing in the main from the base upwards, especially on the higher parts. Likewise the fruits of weaker lateral branches are smaller. Curves are easily made by measuring a few hundred capsules from corresponding parts of different plants, or even by limiting the [748] inquiry to a single individual. These curves give the partial variability, and are found to comply with Quetelet's law.
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.
Vegetative propagation has the great advantage of exempting the varieties from regression to mediocrity, which always follows multiplication by seeds. It affords the possibility of keeping the extremes constant, and this is not its only advantage. Another, likewise highly interesting, side of the question is the uniformity of the whole strain. This is especially important in the case of fruits, though ordinarily it is regarded as a matter of course, but there are some exceptions which give proof of the real importance of the usual condition. For example, the walnut-tree. Thousands of acres of walnut-orchards consist of seedling trees grown from nuts of unknown parentage. The result is a great diversity in the types of trees and in the size and shape of the nuts, and this diversity is an obvious disadvantage to the industry. The cause lies in the enormous difficulties attached to grafting or budding of these trees, which make this method very expensive and to a high degree uncertain and unsatisfactory.
[767] After this hasty survey of the more reliable facts of the practice of an asexual multiplication of the extremes of fluctuating variability, we may now return to the previously mentioned theoretical considerations. These are concerned with an estimation of the chances of the occurrence of deviations, large enough to exhibit commercial value. This chance may be calculated on the basis of Quetelet's law, whenever the agreement of the fluctuation of the quality under consideration has been empirically determined. In the discussion of the methods of comparing two curves, we have pointed to the quartiles as the decisive points, and to the necessity of drawing the curves so that these points are made to overlie one another, on each side of the average. If now we calculate the binomium of Newton for different values of the exponent, the sum of the coefficients is doubled for each higher unit of the exponent, and at the same time the extreme limit of the curve is extended one step farther. Hence it is possible to calculate a relation between the value of the extreme and the number of cases required. It would take us too long to give this calculation in detail, but it is easily seen that for each succeeding step the number of individuals must be doubled, though the length of the steps, or the amount of increase of the quality [768] remains the same. The result is that many thousands of seedlings are required to go beyond the ordinary range of variations, and that every further improvement requires the doubling of the whole culture. If ten thousand do not give a profitable deviation, the next step requires twenty thousand, the following forty thousand, and so on. And all this work would be necessary for the improvement of a single quality, while practice requires the examination and amelioration of nearly all the variable characters of the strain.
Hence the rule that great results can only be obtained by the use of large numbers, but it is of no avail to state this conclusion from a scientific point of view. Scientific experimenters will rarely be able to sacrifice fifty thousand plants to a single selection. The problem is to introduce the principle into practice and to prove its direct usefulness and reliability. It is to Luther Burbank that we owe this great achievement. His principles are in full harmony with the teachings of science. His methods are hybridization and selection in the broadest sense and on the largest scale. One very illustrative example of his methods must suffice to convey an idea of the work necessary to produce a new race of superlative excellency. Forty thousand blackberry and raspberry [769] hybrids were produced and grown until the fruit matured. Then from the whole lot a single variety was chosen as the best. It is now known under the name of "Paradox." All others were uprooted with their crop of ripening berries, heaped up into a pile twelve feet wide, fourteen feet high and twenty-two feet long, and burned. Nothing remained of that expensive and lengthy experiment, except the one parent-plant of the new variety. Similar selections and similar amount of work have produced the famous plums, the brambles and the blackberries, the Shasta daisy, the peach almond, the improved blueberries, the hybrid lilies, and the many other valuable fruits and garden-flowers that have made the fame of Burbank and the glory of horticultural California.

[770]

LECTURE XXVII

INCONSTANCY OF IMPROVED RACES

The greater advantages of the asexual multiplication of extremes are of course restricted to perennial and woody plants. Annual and biennial species cannot as a rule, be propagated in this way, and even with some perennials horticulturists prefer the sale of seeds to that of roots and bulbs. In all these cases it is clear that the exclusion of the individual variability, which was shown to be an important point in the last lecture, must be sacrificed.
Seed-propagation is subject to individual as well as to fluctuating variability. The first could perhaps be designated by another term, embryonic variability, since it indicates the fluctuations occurring during the period of development of the germ. This period begins with the fusion of the male and female elements and is largely dependent upon the vigor of these cells at the moment, and on the varying qualities they may have acquired. It comprises in the main the time of the ripening of the seed, and [771] might perhaps best be considered to end with the beginning of the resting stage of the ripe seed. Hence it is clear that the variability of seed-propagated annual races has a wider range than that of perennials, shrubs and trees. At present it is difficult to discern exactly the part each of these two main factors plays in the process. Many indications are found however, that make it probable that embryonic variability is wider, and perhaps of far greater importance than the subsequent partial fluctuations. The high degree of similarity between the single specimens of a vegetative variety, and the large amount of variability in seed-races strongly supports this view. The propagation and multiplication of the extremes of fluctuating variability by means of seeds requires a close consideration of the relation between seedling and parent. The easiest way to get a clear conception of this relation is to make use of the ideas concerning the dependency of variability upon nourishment. Assuming these to be correct in the main, and leaving aside all minor questions, we may conclude that the chosen extreme individual is one of the best nourished and intrinsically most vigorous of the whole culture. On account of these very qualities it is capable of nourishing all of its organs better and also its seeds. In other words, the seeds [772] of the extreme individuals have exceptional chances of becoming better nourished than the average of the seeds of the race. Applying the same rule to them, it is easily understood that they will vary, by reason of this better nourishment, in a direction corresponding to that of their parent.
This discussion gives a very simple explanation of the acknowledged fact that the seeds of the extremes are in the main the best for the propagation of the race. It does not include however, all the causes for this preferment. Some are of older date and due to previous influences.
A second point in our discussion is the appreciation of the fact that a single individual may be chosen to gather the seed from, and that these seeds, and the young plants they yield, are as a rule, numerous. Hence it follows that we are to compare their average and their extremes with the qualities of the parents. Both are of practical as well as of theoretical interest. The average of the progeny is to be considered as the chief result of the selection in the previous generation, while the extremes, at least those which depart in the same direction, are obviously the means of further improvement of the race.
Thus our discussion should be divided into [773] two heads. One of these comprises the relation of the average of the progeny to the exceptional qualities of the chosen parent, and the other the relation of exceptional offspring to the exceptional parents.
Let us consider the averages first. Are they to be expected to be equal to the unique quality of the parent, or perhaps to be the same as the average of the whole unselected race? Neither of these cases occur. Experience is clear and definite on this important point. Vilmorin, when making the first selections to improve the amount of sugar in beets, was struck with the fact that the average of the progeny lies between that of the original strain and the quality of the chosen parent. He expressed his observation by stating that the progeny are grouped around and diverge in all directions from some point, placed on the line which unites their parent with the type from which it sprang. All breeders agree on this point, and in scientific experiments it has often been confirmed. We shall take up some illustrative examples presently, but in order to make them clear, it is necessary to give a closer consideration to the results of Vilmorin.
From his experience it follows that the average of the progeny is higher than that of the race at large, but lower than the chosen parent. [774] In other words, there is a progression and a regression. A progression in relation to the whole race, and a regression in comparison with the parent. The significance of this becomes clear at once, if we recall the constancy of the variety which could be obtained from the selected extreme in the case of vegetative multiplication. The progression is what the breeder wants, the regression what he detests. Regression is the permanency of part of the mediocrity which the selection was invoked to overcome. Manifestly it is of the highest interest that the progression should be as large, and the regression as small as possible. In order to attain this goal the first question is to know the exact measure of progression and regression as they are exhibiting themselves in the given cases, and the second is to inquire into the influences, on which this proportion may be incumbent.
At present our notions concerning the first point are still very limited and those concerning the second extremely vague. Statistical inquiries have led to some definite ideas about the importance of regression, and these furnish a basis for experimental researches concerning the causes of the phenomenon. Very advantageous material for the study of progression and regression in the realm of fluctuating variability is afforded by the [775] ears of corn or maize. The kernels are arranged in longitudinal rows, and these rows are observed to occur in varying, but always even, numbers. This latter circumstance is due to the fact that each two neighboring rows contain the lateral branches of a single row of spikelets, the ages of which however, are included in the fleshy body of the ear. The variation of the number of the rows is easily seen to comply with Quetelet's law, and often 30 or 40 ears suffice to give a trustworthy curve. Fritz Muller made some experiments upon the inheritance of the number of the rows, in Brazil. He chose a race which averaged 12 rows, selected ears with 14, 16 and 18 rows, etc., and sowed their kernels separately. In each of-these cultures he counted the rows of the seeds on the ears of all the plants when ripe, and calculated their average. This average, of course, does not necessarily correspond to a whole number, and fractions should not be neglected.
According to Vilmorin's rule he always found some progression of the average and some regression. Both were the larger, the more the parent-ear differed from the general average, but the proportion between both remained the same, and seems independent of the amount of the deviation. Putting the deviation at 5, the progression calculated from his figures is [776] 2 and the regression 3. In other words the average of the progeny has gained over the average of the original variety slightly more than one-third, and slightly less than one-half of the parental deviation. I have repeated this experiment of Fritz Miller's and obtained nearly the same regression of three-fifths, though working with another variety, and under widely different climatic conditions.
The figures of Fritz Muller were, as given below, in one experiment. In the last column I put the improvement calculated for a proportion of two-fifths above the initial average of 12.

Rows on
parent ears		Average of rows
of progeny		12 + 2/5ths of
Difference
1412.612.8
1614.113.6
1815.214.4
2015.815.2
2216.116

Galton, in his work on natural inheritance, describes an experiment with the seeds of the sweet pea or Lathyrus odoratus. He determined the average size in a lot of purchased seeds, and selected groups of seeds of different, but for each group constant, sizes. These were sown, and the average of the seeds was determined anew in the subsequent harvest they yielded. These figures agreed with the rule of Vilmorin and were calculated in the manner [777] given for the test of the corn. The progression and regression were found to be proportionate to the amount of the deviation. The progression of the average was one-third, and the regression in consequence two-thirds of the total deviation. The amelioration is thus seen to be nearly, though not exactly, the same as in the previous case.
From the evidence of the other corresponding experiments, and from various statistical inquiries it seems that the value of the progression is nearly the same in most cases, irrespective of the species used and the quality considered. It may be said to be from one-third to one-half of the parental deviation, and in this form the statement is obviously of wide and easy applicability.
Our figures also demonstrate the great preeminence of vegetative varieties above the improved strains multiplied by seeds. They have a definite relation. Asexually multiplied strains may be said to be generally two times or even three times superior to the common offspring. This is a difference of great practical importance, and should never be lost sight of in theoretical considerations of the productive capacity of selection. Multiplication by seed however, has one great advantage over the asexual method; it may be repeated. The [778] selection is not limited to a single choice, but may be applied in two or more succeeding generations. Obviously such a repetition affords a better chance of increasing the progression of the average and of ameliorating the race to a greater degree than would be possible by a single choice. This principle of repeated selection is at present the prominent feature of race improvement. Next to variety-testing and hybridizing it is the great source of the steady progression of agricultural crops. From a practical standpoint the method is clear and as perfect as might be expected, but this is not the side of the problem with which we are concerned here. The theoretical analysis and explanation of the results obtained, however, is subject to much doubt, and to a great divergence of conceptions. So it is also with the application of the practical processes to those occurring in nature. Some assume that here repeated selection is only of subordinate importance, while others declare that the whole process of evolution is due to this agency. This very important point however, will be reserved for the next lecture, and only the facts available at present will be considered here.
As a first example we may take the ray-florets of the composites. On a former occasion we have dealt with their fluctuation in number and [779] found that it is highly variable and complies in the main with Quetelet's law. Madia elegans, a garden species, has on the average 21 rays on each head, fluctuating between 16 and 25 or more. I saved the seeds of a plant with only 17 rays on the terminal head, and got from them a culture which averaged 19 rays, which is the mean between 21 and 17. In this second generation I observed the extremes to be 22 and 12, and selected a plant with 13 rays as the parent for a continuation of the experiment. The plants, which I got from its seeds, averaged 18 and showed 22 and 13 as extremes. The total progression of the average was thus, in two generations, from 21 to 18, and the total regression from 13 to 18, and the proportion is thus seen to diminish by the repetition rather than to increase.
This experiment, however, is of course too imperfect upon which to found general conclusions. It only proves the important fact that the improved average of the second generation is not the starting-point for the further improvement. But the second generation allows a choice of an extreme, which diverges noticeably more from the mean than any individual of the first culture, and thereby gives a larger amount of absolute progression, even if the proportion between progression and regression remains [780] the same. The repetition is only an easy method of getting more widely deviating extremes; whether it has, besides this, another effect, remains doubtful. In order to be able to decide this question, it is necessary to repeat the selection during a series of generations. In this way the individual faults may be removed as far as possible. I chose an experiment of Fritz Muller, relating to the number of rows of grains on the ears exactly as in the case above referred to, and which I have repeated in my experimental garden at Amsterdam.
I started from a variety known to fructify fairly regularly in our climate, and exhibiting in the mean 12-14 rows, but varying between 8 and 20 as exceptional cases. I chose an ear with 16 rows and sowed its seeds in 1887. A number of plants were obtained, from each of which, one ear was chosen in order to count its rows. An average of 15 rows was found with variations complying with Quetelet's law. One ear reached 22 rows, but had not been fertilized, some others had 20 rows, and the best of these was chosen for the continuation of the experiment. I repeated the sowing during 6 subsequent generations in the same way, choosing each time the most beautiful ear from among those with the greatest number of rows. Unfortunately with the increase of the number the [781] size of the grains decreases, the total amount of nourishment available for all of them remaining about the same. Thus the kernels and consequently the new plants became smaller and weaker, and the chance of fertilization was diminished in the ears with the highest number of rows. Consequently the choice was limited, and after having twice chosen a spike with 20 and once one with 24 rows, I finally preferred those with the intermediate number of 22.
This repeated choice has brought the average of my race up from 13 to 20, and thus to the extreme limit of the original variety. Seven years were required to attain this result, or on an average the progression was one row in a year. This augmentation was accompanied by an accompanying movement of the whole group in the same direction. The extreme on the side of the small numbers came up from 8 to 12 rows, and cobs with 8 or 10 rows did not appear in my race later than the third generation. On the other side the extreme reached 28, a figure never reached by the original variety as cultivated with us, and ears with 24 and 26 rows have been seen during the four last generations in increasing numbers.
This slow and gradual amelioration was partly due to the mode of pollination of the corn. [782] The pollen falls from the male spikes on the ears of the same plant, but also is easily blown on surrounding spikes. In order to get the required amount of seed it is necessary in our climate to encroach as little as possible upon free pollination, aiding the self-pollination, but taking no precautions against intercrossing. It is assumed that the choice of the best ears indicates the plants which have had the best pollen-parents as well as the best pistil parents, and that selection here, as in other cases, corrects the faults of free intercrossing. But it is granted that this correction is only a slow one, and accounts in a great degree for the slowness of the progression. Under better climatic conditions and with a more entire isolation of the individuals, it seems very probable that the same result could have been reached in fewer generations.
However this may be, the fact is that by repeated selection the strain can be ameliorated to a greater extent than by a single choice. This result completely agrees with the general experience of breeders and the example given is only an instance of a universal rule. It has the advantage of being capable of being recorded in a numerical way, and of allowing a detailed and definite description of all the succeeding generations. The entire harvest of all [783] of them has been counted and the figures combined into curves, which at once show the whole course of the pedigree-experiment. These curves have in the main taken the same shape, and have only gradually been moved in the chosen direction.
Three points are now to be considered in connection with this experiment. The first is the size of the cultures required for the resulting amelioration. In other words, would it have been possible to attain an average of 20 rows in a single experiment? This is a matter of calculation, and the calculation must be based upon the experience related above, that the progression in the case of maize is equal to two-fifths of the parental deviation. A cob with 20 rows means a deviation of 7 from the average of 13, the incipient value of my race. To reach such an average at once, an ear would be required with 7 x 5/2 = 17-1/2 rows above the average, or an ear with 30-32 rows. These never occur, but the rule given in a previous lecture gives a method of calculating the probability of their occurrence, or in other words, the number of ears required to give a chance of finding such an ear. It would take too long to give this calculation here, but I find that approximately 12,000 ears would be required to give one with 28 rows, which was the highest number attained in [784] my experiment, while 100,000 ears would afford a chance of one with 32 rows*. Had I been able to secure and inspect this number of ears, perhaps I would have needed only a year to get an average of 20 rows. This however, not being the case, I have worked for seven years, but on the other hand have cultivated all in all only about one thousand individuals for the entire experiment.
Obviously this reduction of the size of the experiment is of importance. One hundred thousand ears of corn could of course, be secured directly from trade or from some industrial culture, but corn is cultivated only to a small extent in Holland, and in most cases the requisite number of individuals would be larger than that afforded by any single plantation.
Repeated selection is thereby seen to be the means of reducing the size of the required cultures to possible measures, not only in the experimental-garden, but also for industrial purposes. A selection from among 60,000-100,000 individuals may be within reach of Burbank, but of few others. As a rule they prefer a longer time with a smaller lot of plants. This

* On about 200 ears the variability ranges from 8-22 rows, and
this leads approximately to one row more by each doubling of
the numbers of instances. One ear with 22 rows in 200 would
thus lead to the expectation of one ear with 32 rows in
100,000 ears.

[785] is exactly what is gained by repeated selections. To my mind this reduction of the size of the cultures is probably the sole effect of the repetition. But experience is lacking on this point, and exact comparisons should be made whenever possible, between the descendants of a unique but extreme choice, and a repeated but smaller selection. The effect of the repetition on the nourishment of the chosen representatives should be studied, for it is clear that a plant with 22 rows, the parents and grandparents of which had the same number, indicates a better condition of internal qualities than one with the same number of rows, produced accidentally from the common race. In this way it may perhaps be possible to explain, why in my experiment an ear with 22 rows gave an average offspring with 20, while the calculation, founded on the regression alone would require a parental ear with 32 rows.
However, as already stated, this discussion is only intended to convey some general idea as to the reduction of the cultures by means of repeated selections, as the material at hand is wholly inadequate for any closer calculation. This important point of the reduction may be illustrated in still another manner.
The sowing of very large numbers is only required because it is impossible to tell from the [786] inspection of the seeds which of them will yield the desired individual. But what is impossible in the inspection of the seeds may be feasible, at least in important measure, in the inspection of the plants which bear the seeds. Whenever such an inspection demonstrates differences, in manifest connection with the quality under consideration, any one will readily grant that it would be useless to sow the seeds of the worst plants, and that even the whole average might be thrown over, if it were only possible to point out a number of the best. But it is clear that by this inspection of the parent plants the principle of repeated selection is introduced for two succeeding generations, and that its application to a larger series of generations is only a question of secondary importance.
Summing up our discussion of this first point we may assert that repeated selection is only selection on a small and practical scale, while a single choice would require numbers of individuals higher than are ordinarily available.
A second discussion in connection with our pedigree-culture of corn is the question whether the amelioration obtained was of a durable nature, or only temporary. In other words, whether the progeny of the race would remain constant, if cultivated after cessation of the selection. In order to ascertain this, [787] I continued the culture during several generations, choosing ears with less than the average number of rows. The excellence of the race at once disappeared, and the ordinary average of the variety from which I had started seven years before, returned within two or three seasons. This shows that the attained improvement is neither fixed nor assured and is dependent on continued selection. This result only confirms the universal experience of breeders, which teaches the general dependency of improved races on continued selection. Here a striking contrast with elementary species or true varieties is obvious. The strains which nature affords are true to their type; their average condition remains the same during all the succeeding generations, and even if it should be slightly altered by changes in the external conditions, it returns to the type, as soon as these changes come to an end. It is a real average, being the sum of the contribution of all the members of the strain. Improved races have only an apparent average, which is in fact biased by the exclusion of whole groups of individuals. If left to themselves, their appearance changes, and the real average soon returns. This is the common experience of breeders.
A third point is to be discussed in connection [788] with the detailed pedigree-cultures. It is the question as to what might be expected from a continuation of improvement selection. Would it be possible to obtain any imaginable deviation from the original type, and to reach independency from further selection? This point has not until now attracted any practical interest, and from a practical point of view and within the limits of ordinary cultures, it seems impossible to obtain a positive answer. But in the theoretical discussion of the problems of descent it has become of the highest importance, and therefore requires a separate treatment, which will be reserved for the next lecture.
Here we come upon another equally difficult problem. It relates to the proportion of embryonic or individual fluctuation, to partial variation as involved in the process of selection. Probably all qualities which may be subjected to selection vary according to both principles, the embryonic decision giving only a more definite average, around which the parts of the individual are still allowed to oscillate. It is so with the corn, and whenever two or more ears are ripening or even only flowering on the same plant, differences of a partial nature may be seen in the number of their rows. These fluctuations are only small however, ordinarily not exceeding two and rarely four [789] rows. Choosing always the principal ear, the figures may be taken to indicate the degree of personal deviation from the average of the race. But whenever we make a mistake, and perchance sow from an ear, the deviation of which was largely due to partial variation, the regression should be expected to become considerably larger. Hence it must be conceded that exact calculations of the phenomena of inheritance are subject to much uncertainty, resulting from our very imperfect knowledge concerning the real proportion of the contributing factors, and the difficulty of ascertaining their influence in any given case. Here also we encounter more doubts than real facts, and much remains to be done before exact calculations may become of real scientific value.
Returning to the question of the effects of selection in the long run, two essentially different cases are to be considered. Extremes may be selected from among the variants of ordinary fluctuating variability, or from ever-sporting varieties. These last we have shown to be double races. Their peculiar and wide range of variability is due to the substitution of two characters, which exclude one another, or if combined, are diminished in various degrees. Striped flowers and stocks, "five-leaved" clover, pistilloid opium-poppies and numerous other [790] monstrosities have been dealt with as instances of such ever-sporting varieties.
Now the question may be put, what would be the effect of selection if in long series of years one of the two characters of such a double race were preferred continuously, to the complete exclusion of the other. Would the race become changed thereby? Could it be affected to such a degree as to gradually lose the inactive quality, and cease to be a double race?
Here manifestly we have a means by which to determine what selection is able to accomplish. Physiologic experiments may be said to be too short to give any definite evidence. But cases may be cited where nature has selected during long centuries and with absolute constancy in her choice. Moreover unconscious selections by man have often worked in an analogous manner, and many cultivated plants may be put to the test concerning the evidence they might give on this point. Stating beforehand the result of this inquiry, we may assert that long-continued selection has absolutely no appreciable effect. Of course I do not deny the splendid results of selection during the first few years, nor the necessity of continued selection to keep the improved races to the height of their ameliorated qualities. I only wish to state that the work [791] of selection here finds its limit and that centuries and perhaps geologic periods of continued effort in the same direction are not capable of adding anything more to the initial effect. Some illustrative examples may suffice to prove the validity of this assertion. Every botanist who has studied the agricultural practice of plant-breeding, or the causes of the geographic distribution of plants, will easily recall to his mind numerous similar cases. Perhaps the most striking instance is afforded by cultivated biennial plants. The most important of them are forage-beets and sugar-beets. They are, of course, cultivated only as biennials, but some annual specimens may be seen each year and in nearly every field. They arise from the same seed as the normal individuals, and their number is obviously dependent on external conditions, and especially on the time of sowing. Ordinary cultures often show as much as 1% of these useless plants, but the exigencies of time and available labor often compel the cultivator to have a large part of his fields sown before spring. In central Europe, where the climate is unfavorable at this season, the beets respond by the production of far larger proportions of annual specimens, their number coming often up to 20% or more, thus constituting noticeable losses in the product [792] of the whole field. Rimpau, who has made a thorough study of this evil and has shown its dependency on various external conditions, has also tried to find methods of selection with the aim of overcoming it, or at least of reducing it to uninjurious proportions. But in these efforts he has reached no practical result. The annuals are simply inexterminable.
Coming to the alternative side of the problem it is clear that annuals have always been excluded in the selection. Their seeds cannot be mixed with the good harvest, not even accidentally, since they have ripened in a previous year. In order to bear seeds in the second year beets must be taken from the field, and kept free from frost through the winter. The following spring they are planted out, and it is obvious that even the most careless farmer is not liable to mix them with annual specimens. Hence we may conclude that a strict and unexcelled process of selection has been applied to the destruction of this tendency, not only for sugar-beets, since Vilmorin's time, when selection had become a well understood process, but also for forage-beets since the beginning of beet culture. Although unconscious, the selection of biennials must have been uninterrupted and strict throughout many centuries.
It has had no effect at all. Annuals are seen [793] to return every year. They are ineradicable. Every individual is in the possession of this latent quality and liable to convert it into activity as soon as the circumstances provoke its appearance, as proved by the increase of annuals in the early sowings. Hence the conclusion that selection in the long run is not adequate to deliver plants from injurious qualities. Other proofs could be given by other biennials, and among them the stray annual plants of common carrots are perhaps the most notorious. In my own cultures of evening-primroses I have preferred the annuals and excluded the biennials, but without being able to produce a pure annual race. As soon as circumstances are favorable, the biennials return in large numbers. Cereals give analogous proofs. Summer and winter varieties have been cultivated separately for centuries, but in trials it is often easy to convert the one into the other. No real and definite isolation has resulted from the effect of the long continued unconscious selection.
Striped flowers, striped fruits, and especially striped radishes afford further examples. It would be quite superfluous to dwell upon them. Selection always tends to exclude the monochromatic specimens, but does not prevent their return in every generation. Numerous [794] rare monstrosities are in the same category, especially when they are of so rare occurrence as not to give any noticeable contribution to the seed-production, or even if they render their bearers incapable of reproduction. In such cases the selection of normal plants is very severe or even absolute, but the anomalies are by no means exterminated. Any favorable circumstances, or experimental selection in their behalf shows them to be still capable of full development. Numerous cases of such subordinate hereditary characters constitute the greater part of the science of vegetable teratology.
If it should be objected that all these cases cover too short a time to be decisive, or at least fail in giving evidence relative to former times, alpine plants afford a proof which one can hardly expect to be surpassed. During the whole present geologic epoch they have been subjected to the never failing selection of their climate and other external conditions. They exhibit a full and striking adaptation to these conditions, but also possess the latent capacity for assuming lowland characters as soon as they are transported into such environment. Obviously this capacity never becomes active on the mountains, and is always counteracted by selection. This agency is evidently without any effect, for as we have seen when dealing [795] with the experiments of Nageli, Bonnier and others, each single individual may change its habits and its aspect in response to transplantation. The climate has an exceedingly great influence on each individual, but the continuance of this influence is without permanent result.
So much concerning ever-sporting varieties and double adaptations. We now come to the effects of a continuous selection of simple characters.
Here the sugar-beets stand preeminent. Since Vilmorin's time they have been selected according to the amount of sugar in their roots, and the result has been the most striking that has ever been attained, if considered from the standpoint of practice. But if critically examined, with no other aim than a scientific appreciation of the improvement in comparison with other processes of selection, the support of the evidence for the theory of accumulative influence proves to be very small.
The amount of sugar is expressed by percentage-figures. These however, are dependent on various causes, besides the real quantity of sugar produced. One of these causes is the quantity of watery fluid in the tissues, and this in its turn is dependent on the culture in dryer or moister soil, and on the amount of moisture in the air, and the same variety of sugar-beets [796] yields higher percentage-figures in a dry region than in a wet one. This is seen when comparing, for instance, the results of the analyses from the sandy provinces of Holland with those from the clay-meadows, and it is very well known that Californian beets average as high as 26% or more, while the best European beets remain at about 20%. As far as I have been able to ascertain, these figures however, are not indicative of any difference of race, but simply direct responses to the conditions of climate and of soil.
Apart from these considerations the improvement reached in half a century or in about twenty to thirty generations is not suggestive of anything absolute. Everything is fluctuating now, even as it was at the outset, and equally dependent on continual care. Vilmorin has given some figures for the beets of the first generations from which he started his race. He quotes 14% as a recommendable amount, and 7 and 21 as the extreme instances of his analyses. However incorrect these figures may be, they coincide to a striking degree with the present condition of the best European races. Of course minor values are excluded each year by the selection, and in consequence the average value has increased. For the year 1874 we find a standard of 10-14% considered as normal, [797] bad years giving 10%, good years from 12% to 14% in the average. Extreme instances exceeded 17%. From that time the practice of the polarization of the juice for the estimate of the sugar has rapidly spread throughout Europe, and a definite increase of the average value soon resulted. This however, often does not exceed 14%, and beets selected in the field for the purpose of polarization come up to an average of 15 to 16%, varying downward to less than 10% and upward to 20 and 21%. In the main the figures are the same as those of Vilmorin, the range of variability has not been reduced, and higher extremes are not reached. An average increase of 1% is of great practical importance, and nothing can excel the industry and care displayed in the improvement of the beet-races. Notwithstanding this a lasting influence has not been exercised; the methods of selection have been improved, and the number of polarized beets has been brought up to some hundreds of thousands in single factories, but the improvement is still as dependent upon continuous selection as it was half a century ago.
The process is practically very successful, but the support afforded by it to the selection theory vanishes on critical examination.

[798]