Popular Science Monthly/Volume 69/October 1906/Scientific Aspects of Luther Burbank's Work

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SCIENTIFIC ASPECTS OF LUTHER BURBANK'S WORK
By VERNON L. KELLOGG

PROFESSOR OF ENTOMOLOGY, STANFORD UNIVERSITY

MR. BURBANK has so far not formulated any new or additional laws of species-change, nor do his observations and results justify any such formulation, and we may rest in the belief that he has no new fundamental laws to reveal. He has indeed the right to formulate, if he cares to, some valuable and significant special conclusions touching certain already recognized evolution factors, in particular, the influence on variability of the two long-known variation producing factors of hybridization and modification of environment. His reliance on the marked increase in variability to be got after a crossing in the second and third generations over that obvious in the first will come as a surprise to most men first getting acquainted with his work. He has got more starts for his new things from these generations than in any other way. He is wholly clear and convinced in his own mind as to the inheritance of acquired characters; 'acquired characters are inherited or I know nothing of plant life' he says; and also convinced that the only unit in organic nature is the individual, not the species; that the so-called species are wholly mutable and dependent for their apparent fixity solely on the length of time through which their so-called phyletic characters have been ontogenetically repeated. He does not agree at all with de Vries that mutations in plants occur only at certain periodic times in the history of the species, but rather that, if they occur at all, they do so whenever the special stimulus derived from unusual nutrition or general environment can be brought to bear on them. He finds in his breeding work no prepotency of either sex as such in inheritance, though any character or group of characters may be prepotent in either sex. He believes that no sharp line can be drawn between the fluctuating or so-called Darwinian variations and those less usual, large, discontinuous ones called sports. Ordinary fluctuating variation goes on under ordinary conditions of nutrition, but with extraordinary environmental conditions come about extraordinary variation results, namely, discontinuous, sport or mutational variation. These variations are the effects of past environment also, having remained latent until opportunity for their development occurs. Starvation causes reversions, but reversions can also be produced by unusually rich nutrition. New variations are developed most often, as far as environmental influences go, by rich soil and generally favorable conditions. So-called new qualities are usually, if not always (the fact may sometimes not be obvious), simply new combinations of old qualities, both latent and obvious. To get a new and pleasing odor it may often be sufficient simply to lose one bad element in an old odor. So one might go on for some pages with specific conclusions or deductions reached by Burbank on a basis of experience. But it is true that he has at his command the knowledge of no new fundamental scientific principles to give him advantage over us. And yet none of us has done what Burbank has been able to do, although many of us have tried. What then is it that Burbank brings to his work of modifying organisms swiftly and extremely and definitely that others do not?

To answer this it will be advisable to analyze, in general terms, at least, the various processes which either singly, or in combinations of two or three, or all together, are used by Mr. Burbank in his work. We may roughly classify these processes and means. First, there is the importation from foreign countries, through many correspondents, of a host of various kinds of plants, some of economic value in their native land and some not, any of which grown under different conditions here may prove specially vigorous or prolific or hardy, or show other desirable changes or new qualities. Among these importations are often special kinds particularly sought for by Burbank to use in his multiple hybridizations; kinds closely related to our native or to already cultivated races which, despite many worthless characteristics, may possess one or more particular, valuable ones needed to be added to a race already useful to make it more useful. Such an addition makes a new race.

Second, the production of variations, abundant and extreme, by various methods, as (a) the growing under new and, usually, more favorable environment (food supply, water, temperature, light, space, etc.) of various wild or cultivated forms, and (b) by hybridizations between forms closely related, less closely related and, finally, as dissimilar as may be (not producing sterility), this hybridizing being often immensely complicated by multiplying crosses, i. e., the offspring from one cross being immediately crossed with a third form, and the offspring of this with still another form, and so on. These hybridizations are made sometimes with very little reference to the actual useful or non-useful characteristics of the crossed parents, with the primary intention of producing an unsettling or instability in the heredity, of causing, as Burbank sometimes says, 'perturbations' in the plants, so as to get just as wide and as large variation as possible. Other crosses are made, of course, in the deliberate attempt to blend, to mix, to add together, two desirable characteristics, each possessed by only one of the crossed forms. Some crosses are made in the attempt to extinguish an undesirable characteristic.

Third, there is always immediately following the unusual production of variations, the recognition of desirable modifications and the intelligent and effective selection of them, i. e., the saving of those plants to produce seed or cuttings which show the desirable variations and the discarding of all the others. In Burbank's gardens the few tenderly cared for little potted plants or carefully grafted seedlings represent the surviving fittest, and the great bonfires of scores of thousands of uprooted others, the unfit, in this close mimicry of Darwin and Spencer's struggle and survival in nature.

It is precisely in this double process of the recognition and selection of desirable variations that Burbank's genius comes into particular play. Bight here he brings something to bear on his work that few other men have been able to do. It is the extraordinary keenness of perception, the delicacy of recognition of desirable variations in their (usually) small and to most men imperceptible beginnings. Is it a fragrance that is sought? To Burbank in a bed of hundreds of seedling walnuts scores of the odors of the plant kingdom are arising and mingling from the fresh green leaves, but each, mind you, from a certain single seedling or perhaps from a similar pair or trio. But to me or to you, until the master prover points out two or three of the more dominant single odors, the impression on the olfactories is simply (or confusedly) that of one soft elusive fragrance of fresh green leaves. Similarly Burbank is a master at seeing, and a master at feeling. And besides he has his own unique knowledge of correlations. Does this plum seedling with its score of leaves on its thin stem have those leaves infinitesimally plumper, smoother or stronger, or with more even margins and stronger petiole or what not else, than any other among a thousand similar childish trees? Then it is saved, for it will bear a larger, or a sweeter, or a firmer sort of plum, or more plums than the others. So to the bonfires with the others and to the company of the elect with this 'fittest' one. Now this recognition, this knowledge of correlations in plant structure, born of the exercise of a genius for perceiving through thirty years of opportunity for testing and perfecting it, is perhaps the most important single thing which Burbank brings to his work that other men do not (at least in such unusual degree of reliability). Enormous industry, utter concentration and single-mindedness, deftness in manipulation, fertility in practical resource, has Burbank—and so have numerous other breeders and experimenters. But in his perception of variability in its forming, his recognition of its possibilities of outcome, and in his scientific knowledge of correlations, a knowledge that is real, for it is one that is relied on and built on, and is at the very foundation of his success, Burbank has an advantage of true scientific character over his fellow workers, and in it he makes a genuine contribution to scientific knowledge of plant biology, albeit this knowledge is so far only proved to be attainable and to exist. It is not yet exposed in its details and may never be, however unselfish be the owner of it. For the going to oblivion of scientific data of an extent and value equivalent, I may estimate roughly, to those now issuing from any half dozen experimental laboratories of variation and heredity, is the crying regret of all evolution students acquainted with the situation. The recently assumed relations of Mr. Burbank to the Carnegie Institution are our present chief hope for at least a lessening of this loss.

But let us follow our saved plum seedling. Have we now to wait the six or seven years before a plum tree comes into bearing to know by actual seeing and testing what new sort of plum we have? No; and here again is one of Burbank' s contributions (not wholly original to be sure, but original in the extent and perfection of its development) to the scientific aspects of plant-breeding. This saved seedling and other similar saved ones (for from the examination of 20,000 seedlings, say, Burbank will find a few tens or even scores in which he has faith of reward) will be taken from their plots and grafted on to the sturdy branches of some full-grown vigorous plum tree, so that in the next season or second next our seedling stem will bear its flowers and fruits. Here are years saved. Twenty, forty, sixty, different seedlings grafted on to one strong tree (in a particular instance Burbank had 600 plum grafts on a single tree!); and each seedling-stem certain to bear its own kind of leaf and flower and fruit. For we have long known that the scion is not materially influenced by the stock nor the stock by the scion; that is not modified radically, although grafting sometimes increases or otherwise modifies the vigor of growth and the extent of the root system of the stock.

If now the fruit from our variant seedling is sufficiently desirable; if it produces earlier or later, sweeter or larger, firmer or more abundant, plums, we have a new race of plums, a 'new creation' to go into that thin catalogue of results. For by simply subdividing the wood of the new branch, i. e., making new grafts from it, the new plum can be perpetuated and increased at will. Simple, is it not? No, it is anything but that in the reality of doing it; but in the scientific aspects of it, easily understandable.

Perhaps it may not be amiss to call attention to what must be the familiar knowledge of most of us, and that is the fact that many (probably most) cultivated plants must be reproduced by division, that is by cuttings, buds or grafts, and not by seeds, in order to grow 'true.' For a piece of a cultivated plant will grow out to be very much like the individual it was cut from, but the seeds will not, in most cases, reproduce faithfully the parents, but will produce a very variable lot of individuals, most of them strongly reversionary in character. Grow peach trees from the stones of your favorite peach and see what manner of peaches you get; but if you want to be sure of more peaches like the ones you enjoy, graft scions from your tree on to other trees. Indeed one of the plant-breeder's favorite methods of making a start for new things, of getting the requisite beginning wealth and eccentricity of variation, is to grow seedlings, especially from cross-bred varieties. Burbank will give you a thousand dollars for a pinch of horse-radish seed. Sugar-cane seed is needed. The amelioration of many kinds of fruit and flowers and vegetables is checked, because in our carelessness we have allowed these kinds to get into that condition of seedlessness which almost all cultivated races tend toward when grown from cuttings. In our oranges and grape-fruit and in a score of other fruits, the elimination of seeds is exactly one of the modifications we have bred and selected for, in order to make the fruits less troublesome in their eating. But when we lose the seeds entirely of a whole group of related plant kinds we may find ourselves, as we have found ourselves actually in many cases, at the end of our powers of amelioration of these plant sorts. Burbank believes that the very fact that plants when grown asexually always sooner or later lose their power to produce seeds is almost sufficient proof (if such proof is needed) that acquired characters are transmitted.

Another of Burbank's open secrets of success is the great range of his experimentation—nothing is too bold for him to attempt, the chances of failure are never too great to frighten him. And another secret is the great extent, as regards material used, of each experiment. His beds of seedlings contain hundreds, often thousands, of individuals where other men are content with hundreds. Another element in his work is his prodigality of time. Experiments begun twenty years ago are actually still under way.

In all that I have so far written, I have purposely kept to general statements applicable to Burbank's work as a whole. My readers might be more interested, perhaps, to have some illustrations of the application of various processes of making new sorts of things, some analytical account of the history of various specific 'new creations,' but considerations of space practically forbid this. Just a few briefly described examples must suffice. More than is generally imagined, perhaps, Burbank uses pure selection to get new things. From the famous golden orange colored California poppy (Escholtzia) he has produced a fixed new crimson form by selection alone. That is, noticing, somewhere, sometime, an Escholtzia individual varying slightly redder, he promptly took possession of it, raised young poppies from its seeds, selected from among them those varying in a similar direction, raised new generations from them and so on until now he who wishes may have his California poppies of a strange glowing crimson for the price of a little package of seed, where formerly he was perforce content with the golden orange. For me the golden orange suffices, but that does not detract from my eager interest in the flower-painting methods of Mr. Burbank. Even more striking a result is his blue Shirley poppy, produced also solely by repeated selection from the crimson field poppy of Europe. "We have long had various shades of black and crimson and white poppies, but no shade of blue. Out of 200,000 seedlings I found one showing a faintest trace of sky blue and planted the seed from it, and got next year one pretty blue one out of many thousand, and now I have one almost pure blue."

But another brilliant new poppy was made in a different way. The pollen of Papaver pilosum, a butter-colored poppy, was put on the pistils of the Bride, a common pure white variety of Papaver somniferum (double), and in the progeny of this cross was got a fire-colored single form. The character of singleness was common to the ancestors of both parents, the character of fire color in the lineage of somniferum only, although the red of the new form is brighter than ever before known in the somnifera series. Both characteristics were absent (or rather latent) in both parents. And yet the perturbing influence of the hybridization brought to the fore again these ancestral characters. The foliage of this fire poppy is intermediate in type between that of the two parents.

The history of the stoneless and seedless plum, now being slowly developed by Burbank, shows an interesting combination of selection, hybridization and reselecting. Mr. Burbank found a plum in a small wild plum species with only a part of a stone. He crossed this wild species with the French prune; in the first generation he got most individuals with whole stones, some with parts of a stone, and even some with no stone. Through three generations he has now carried his line by steadily selecting, and the percentage of no-stone fruits is slowly increasing, while quality, beauty and productiveness are also increasing at the same time.

The plum-cot is the result of crossing the Japanese plum and the apricot. The plum-cot, however, has not yet become a fixed variety and may never be, as it tends to revert to the plum and apricot about equally, although with also a tendency to remain fixed, which tendency may be made permanent.

Most of Burbank's plums and prunes are the result of multiple crossings in which the Japanese plums have played an important part. Hundreds of thousands of seedlings have been grown and carefully worked over in the twenty years of experimenting with plums, and single trees

PSM V69 D373 Escholtzia california poppy by seedling selection.png

California Poppy (Escholtzia) rendered bright crimson as a result of selection only, without crossing.

have been made to carry as many as 600 varying seedling grafts. The Bartlett plum, cross of the bitter Chinese Simoni and the Delaware, itself a Simoni hybrid, has the exact fragrance and flavor of the Bartlett pear. The Climax, a successful shipping plum, is also a cross of the Simoni and the Japanese triflora. This Chinese Simoni produces almost no pollen, but few grains of it ever having been obtained. But these few grains have enabled Burbank to revolutionize the whole plum shipping industry. The sugar prune, which promises to supplant the French prune in California, is a selected product of a second or third generation variety of the Petite d'Agen, a somewhat variable French plum.

Next in extent probably to Burbank's work with plums and prunes

PSM V69 D373 Stoneless plums.png

One of the Stoneless Plums (Center) and Two of its Parents. On the right hand the common French prune.

is his long and successful experimentation with berries. This has extended through twenty-five years of constant attention, has involved the use, in hybridizations, of forty different species of Rubus, and has resulted in the origination of a score of new commercial varieties, mostly obtained through various hybridizations of dewberries, blackberries and raspberries. Among these may specially be mentioned the Primus, a hybrid of the western dewberry (R. ursinus) and the Siberian raspberry (R. cratægifolius), fixed in the first generation, which ripens its main crop before most of the standard varieties of raspberries and blackberries commence to bloom. (Mr. Burbank does not recommend this for general cultivation; the 'Phenomenal' and Himalaya are

PSM V69 D374 Early and abundant chestnut tree.png

Early and Abundant Bearing Chestnut Tree. One of hundreds of similar ones, due to crossing and selection. These bear fruit at the age of one and a half years and are never without fruit.

better.) In this Primus berry, we have the exceptional instance of a strong variation, due to hybridization, breeding true from the time of Its first appearance. It usually takes about six generations to fix a new variety, but like de Vries's evening primrose mutations, the Primus berry is a fixed new form from the time of its beginning. An interesting feature of Mr. Burbank's brief account, in his 'New Creations' catalogue of 1894, of the berry experimentation, is a reproduction of a photograph showing "a sample pile of brush 12 ft. wide, 11 ft. high, and 22 ft. long, containing 65,000 two-and three-year-old seedling berry bushes (40,000 blackberry X raspberry hybrids and 25,000 Shaffer X Gregg hybrids) all dug up with their crop of ripening berries." The photograph is introduced to give the reader some idea of the work necessary to produce a satisfactory new race of berries. "Of the 40,000 blackberry X raspberry hybrids of this kind 'Phenomenal' is the only one now in existence. From the other 25,000 hybrids, two dozen bushes were reserved for further trial."

An astonishing result of the hybridization between the black walnut. Juglans nigra, used as the pistillate parent, and the California walnut, J. californica, staminate parent, are walnut trees which grow with such an amazing vigor and rapidity that they increase in size at least twice as fast as the combined growth of both parents. Many tremendous growers are got in the first generation, but in the second there are included

PSM V69 D375 Rhodante naglesi improved by size and color by selection.png

Rhodanthe naglesi, improved in size and color by selection only.

some of the most rapidly growing trees, perhaps, in the world. This hybrid has clean-cut, glossy bright green leaves from two to three feet long with a sweet odor like that of apples, but it produces few nuts. Curiously enough the result of hybridization by using the pollen of nigra on pistils of californica produces in abundance large nuts of a quality superior to that possessed by either parent.

The famous Shasta daisy is the result of a multiple crossing between an American and a European species of field daisy and then between these hybrids and a Japanese form. The fragrant calla, known as 'Fragrance' is descended from a single individual found by Burbank while critically examining a block of Little Gem calla seedlings. He was surprised in this examination by a fragrance resembling that of violets or water-lilies; as be had long been seeking a fragrant calla,

PSM V69 D376 Heuchera crested by seedling selection.png

Heuchera Leaves made Crested by Selection of Varying Seedlings.

the individual giving this perfume was carefully hunted out. No farther selecting was done; this plant was the single ancestor of the fragrant new race.

And so one might go on for pages, but with slight variations in detail all these pages would tell only the same story: the stimulating or inducing of variability by environmental influences and by hybridizations; the search after, and keen recognition of, promising special variations; the selection of the plants showing these variations; rearing new generations from them, repeated selection, and new hybridizations to eliminate this characteristic or introduce that, and on until a desirable combination is found. Then the careful fixing of this form or type by repeated selection through several generations.

But an end must be made of this. Let us, in a paragraph, simply sum up the essential things in the scientific aspects of Burbank's work. No new revelations to science of an overturning character; but the revelation of the possibilities of accomplishment, based on general principles already known, by an unusual man. No new laws of evolution, but new facts, new data, new canons for special cases. No new principle or process to substitute for selection, but a new proof of the possibilities of the effectiveness of the old principle. No new categories of variations, but an illuminating demonstration of the possibilities of stimulating variability and of the reality of this general variability as the fundamental transforming factor. No new evidence either to help the Darwinian factors to their death-lied, or to strengthen their lease on life; for the 'man' factor in all the selecting phenomena in Burbank's gardens excludes all 'natural' factors. Here are some of Burbank's own words, touching these matters that scientific men are particularly interested in, in his work:

All scientists have found that preconceived notions, dogmas, and all personal prejudice must be set aside, listening patiently, quietly and reverently to the lessons one by one which mother nature has to teach, shedding light on that which before was a mystery, so that all who will may see and know.

Crossing gives the raiser of new plants the only means of uniting the best qualities of each, but just as often the worst qualities of each are combined and transmitted, so that to be of any value it must be followed by rigid and persistent selection, and in crossing, as in budding and grafting, the affinities can only be demonstrated by actual test.

All wild plants of any species are under almost identical environments, having their energies taxed to the utmost in the fierce struggle for existence. Any great variation under such circumstances is not likely to occur, and is much more likely to be stamped out at once by its struggling competitors, unless the variation should be of special use in competition, in which case it will survive, and all others may be supplanted by it. Thus we see how new species are often produced by nature, but this is not her only mode. Crosses and hybrids are very often found growing wild where two somewhat similar species grow contiguous, and if the combination happens to be a useful one,

PSM V69 D377 Hybrid opuntia without thorns or bristles.png

Hybrid Opuntia, without Thorns or Bristles, with a parent from which these appendages are to be removed by crossing and selection.

as it often does, the new creation is encouraged by nature; then time and environment fix it, and man comes on the scene, perhaps ages later, and discovers it, and, not knowing all the facts, wonders where the connecting links have gone. It is botanically classified as a new species, which it is most certainly.

In cultivated plants the life struggle is removed, and here we find variation almost the rule rather than the exception.

Varieties are the product of fixed laws, never of chance, and with a knowledge of these laws we can improve the products of nature, by employing nature's forces, in ameliorating old or producing new species and varieties better adapted to our necessities and tastes. Better food, more sunshine, less arduous competition, will of themselves induce variation in individual plants which will be more or less transmitted to their seedlings, which, selected consecutively through a certain number of generations, will become permanent. Environment here exerts an influence as in all chemical cosmical and celestial movements. These small increments from environmental forces may produce a gradual or sudden change according to circumstances. The combustion of food liberates the moving force, environment guides it as it does the planets.

When once the persistent type is broken up, old latent forces may be liberated and types buried in the dim past reappear. This, called atavism, is a concentration of ancestral forces—reverberating echoes—from varieties long since passed away, exhibiting themselves when from some cause, for instance crossing, present forces are in a state of antagonism, division, perturbation or weakness. These echoes, if collected by crossing and selection, produce combinations of superlative importance and value.

Finally, in any summation of the scientific aspects of Burbank's work must be mentioned the hosts of immensely valuable data regarding the inheritance of characteristics, the influence of epigenetic factors in development, the possibilities of plant variability, and what not else important to evolution students, mostly going unrecorded, except as they are added in mass to the already too heavy burden carried by the master of the laboratory, and as they are summed up in those actual results which the world gratefully knows as Burbank's 'new creations.'