v.3 Clejt Graft This is one oj several common methods oj grafting. The engrafted twigs are called cions, the branch on which they are placed is called the stock. The essential principle is that the inner bark, called the Cambium layer, oj the cion should be brought into intimate contact with the corresponding layer of bark oj the stock. Further details of grafting are shown in other pictures in this volume. LUTHER BURBANK HIS METHODS AND DISCOVERIES AND THEIR PRACTICAL APPLICATION PREPARED FROM HIS ORIGINAL FIELD NOTES COVERING MORE THAN 100,000 EXPERIMENTS MADE DURING FORTY YEARS DEVOTED TO PLANT IMPROVEMENT WITH THE ASSISTANCE OF The Luther Burbank Society AND ITS ENTIRE MEMBERSHIP UNDER THE EDITORIAL DIRECTION OF John Whitson and Robert John AND Henry Smith Williams, M. D., LL. D. VOLUME III ILLUSTRATED WITH 105 DIRECT COLOR PHOTOGRAPH PRINTS PRODUCED BY A NEW PROCESS DEVISED AND PERFECTED FOR USE IN THESE VOLUMES NEW YORK AND LONDON LUTHER BURBANK PRESS MCMXIV Copyright, 1914, by The Luther Burbank Society Entered at Stationers' Hall, London All rights reserved • -: ; : - •* •/ j •' Volume III — By Chapters Foreword ............................. Pae 3 I Planning a New Plant The First Step Practical Work —The First Step in « i II Plant Affinities — Choosing the Lines of Least Resistance III Practical Pollenation -A Survey of Working Method . IV Quantity Production -On Seedlin Their Care — A^ Survey of r/y — On Seedlings and (\Q V Grafting and Budding — Short-cuts to Quick Tests VI Letting the Bees Do Their Work —Nature Will Help Us All She Can VII Fixing Good Traits — How to Hold a Result Once Achieved VIII Recording the Experiments —Easy Wayi Keep Trai IX Final Selection — The Most Work of A List of Direct Color Photograph Prints 305 — Easy Ways to Keep Track of Progress . . — The Most Important Work of All .... 340358 FOREWORD TO VOLUME III With many examples of the actual work before us, Mr. Burbank now takes us into the interesting detail of method itself, treating the subjects of pollenation, grafting, plant affinities, fixing traits, selection, and spreading before us all of the processes which he has employed in his more than 100,000 separate experiments. The purpose has been to lead the reader, by easy and interesting stages, up to a point where a delineation of the actual processes may be most readily grasped. This, the third book, completes the consideration of general method, and with the two preceding volumes gives us an intelligent survey of Mr. Burbank's work. THE EDITORS. A Burbank Strawberry This picture represents a Jruit that was developed in accordance with Mr. Burbank's preconception as to what a perfect strawberry would be like. It is remarkably symmetrical in shape, of just the right size, and its qualities oj Jirmness oj texture and of flavor are ideal. It is the product of a long series of experiments in selective Breeding. John Burroughs pronounced it the best strawberry he had ever eaten. PLANNING A NEW PLANT THE FIRST STEPS IN PRACTICAL WORK SOMEONE has said that a painter is a man who can see the picture in the landscape. In similar fashion it may be said that a successful plant experimenter is one who can see new varieties of future plants when he looks at old existing varieties. But of course the painter, whatever his con- structive imagination, does not always see at first glance every detail of form and color that will ultimately appeal to him. Nor can the plant experimenter claim, by any manner of means, to know always from the outset just what his new plant creations will be like. There are numberless instances, indeed, in which a plant experimenter who operates on a large scale may make hybrid- izing experiments, merely to test the possibilities of crossing certain species, without having any precise and definite goal in view. [VOLUME III— CHAPTER I] LUTHER BURBANK But, on the other hand, it is necessary in the pursuit of practical plant developments to have a tolerably precise idea in mind as to the particular direction in which progress is desirable. Lacking such an ideal, the breeder of plants would be about as likely to produce new creations of value as an architect would be likely to construct a fine building by putting materials together at random without a carefully precon- ceived plan. "In what percentage of cases have you achieved the ideal at which you aimed in the production of new varieties of flowers or fruits?" a visitor asked me. The question is almost impossible of definite answer. When I first commenced, doubtless a very small proportion of my experiments came out as I expected. But now, with years of experi- ence to guide me, I may say that I practically always get something not far different from what I desire. In many cases, the result comes just about as I expected. But this is because I am working with plants that I have previously tested. With a new plant I am still sometimes in doubt. But if it is a case of poppies or walnuts or any one of a score or so of other plants that I have fully tested, I know just about what to expect. [8] PLANNING A NEW PLANT At best, however, I am very often reminded that each species has its own individuality and that even the most familiar plant may hold surprises in store for us. THE ROUGH SKETCH "But just how do you start out when you are seeking to create a new form of plant life?" I am constantly asked. And here again the answer is difficult. Every- thing depends upon the ultimate object. If I am seeking merely to test the possibilities of making certain crosses, or as it were feeling my way along new channels, I am more or less like a person groping in the dark. This form of vague experimentation is often full of interest. I have already given some instances of what may come to pass when we hybridize plants of widely separated species or of different genera. The reader will recall the case of the petunia with the tobacco habit and of the dewberry crossed with such remote cousins as apple and pear and mountain-ash. These experiments were made without a clearly defined object — except to ascertain whether it was possible to hybridize plants of such diverse character. And the results of the experiment, while of very great scientific interest, were not practically successful in a commercial sense. 19] LUTHER BURBANK I recall reading an address by the late Professor Newton, a distinguished American astronomer, on the subject of "dead work," in which he emphasized the fact that the main bulk of the experiments which any scientific worker must make will lead to no definite goal. A large part of the time of every experimenter must be given up to following trails that lead nowhere in particular or that end in blind cul de sacs. The work of the plant experimenter is no exception, but there is always an incentive to further effort in the knowledge that a path that seems to lead only into impenetrable mazes may presently bring one out into the light. To make the application to one illustrative case among many, I recall that for twenty-four successive seasons I attempted to hybridize certain species of Solanum before I finally succeeded in effecting a cross that gave me a single seed from which sprang the new race of Sunberries. But it must not be understood that the main bulk of my experiments are made in any hap- hazard manner. On the contrary my most important results have been attained by continuing the experimen- tation along rigidly predetermined lines and by methods of hybridizing and selection that my earlier work had fully established. Having served [10] Strawberries Showing Variation This picture illustrates the variation that may be shown by fruit growing on vines from the same lot of seed. By selection, each type of berry here shown might have its peculiarities accentuated, and as many new varieties developed. Mr. Burbank is always on the lookout Jor variation among his plants, and such variation Jorms the basis of his experiments in selective breeding. LUTHER BURBANK a long apprenticeship and tested the usual limits of making new plant combinations, I was presently able, like any other trained technician, to apply the knowledge thus acquired toward far more definite results than were at first possible. In the case of the Shasta daisy, the plans were all laid out beforehand as to just what type of flower I wished to produce. The ideal of a white blackberry was also, of course, a perfectly precise and definite one. Obviously the scented calla, the stoneless plum, the early bearing cherry, the sugar prune, and the spineless cactus are other instances in which the ideal pursued was as clearly conceived and as definitely outlined in advance of my earliest experiments as a cathedral is outlined in the mind of the architect before he commences his preliminary drawings. In one case as in the other the details may be modified as the work progresses, but the general idea of the structure aimed at — be it new fruit or new building — must be conceived with a good deal of definiteness from the outset. My original conception of a new plant creation, in the cases outlined and in a large number of others, certainly bore as close a resemblance to the final product achieved as the first rough drawing of the architect bears to his finished plans. [12] PLANNING A NEW PLANT "But how do you begin? What is the very first thing?" a visitor insists. The "very first thing" I have already described — it is the conception of an ideal, a mental picture of the new plant form desired. CLUES TO BE FOLLOWED It has occurred to me, for instance, that the cherry crop is not what it might be. I have learned that there is a steady market for early cherries and that a difference of a few days in the time of marketing may make a difference of more than one hundred per cent, in the price. And so I ask myself, why not create a new cherry that shall be ready for shipping at least two or three weeks earlier than any cherry now in the market? Of course, I reflect that my early cherry must have a number of other desirable qualities — large size, rich color, lusciousness of flavor. I know at the outset, or I presently learn, that it is desirable also, from the standpoint of the shipper, that my cherries shall grow on short stems. I know that the tree producing them must be hardy, capable of withstanding both cold winters and dry sum- mers, and that it must have an inherent vitality that will make it resistant to the attacks of insects and fungoid pests. Next I ask myself what warrant there is for [13] LUTHER BURBANK supposing that I can build such a fruit-structure as I have conceived. And here the answer is supplied solely by the use of imagination in connection with the inspection of existing races of cherries. I examine the best fruits already in the orchard and find that there is a large measure of variation between the cherries grown on different trees, as well as between the individual specimens on the same tree. In imagination I look back far into the past and inquire as to the racial history of this fruit. I am led to believe that certain among the ancestors of the cherry have grown in semi- tropical climates, and I know that even in the present day there are species, doubtless sprung from the same original stock, that grow far up into Canada. I ask myself why it is that the cherry shows such a propensity to vary, and I find an answer in the assumption that the existing cultivated races carry in their veins, so to speak — that is to say in their germ plasm — hereditary tendencies drawn from varied strains of a mixed ancestry. And I feel well assured that it should be possible, by accentuating the tendency to varia- tion through further hybridizing, and by careful selection, to combine and bring out in a more or [14] LUTHER BURBANK less remote generation of progeny of the existing cherries, a race that will furnish extreme examples, through reversion, of the limits of variation in each direction — and as regards each particular quality of fruit — that any ancestor reached. It will be obvious, then, that I am not preparing to make bricks without straw. I am counting well the materials with which I must work, just as the architect from the first stroke of his pencil bears in mind the materials of the future cathedral. I do not imagine that I can produce an apple from my cherry stalks, any more than the architect assumes that he can build a marble cathedral out of bricks. I know that there are sharply defined hereditary limitations beyond which the cherry cannot be made to go within any such period of time as that limiting my experiment. In other words, I do not ask the impossible, although it has often seemed to my critics that I have asked the highly improbable. But the results I have attained are in them- selves sufficient answer to the critic. If my vision has in some cases been the clearer, it is merely that my knowledge of plant life, drawn from the school of experience, has been wider. To the uninitiated observer it may have seemed [16] PLANNING A NEW PLANT that I set no limits to the transformations I attempted. In reality, my plan has always from the outset recognized most definite limits— although often enough the limits as I conceived them were quite different from those that had been set by theoretical botanists. AID FROM GALTON'S LAW In attempting to estimate the possibility of improvement in a given form of plant life, it is of value to recall the formula put forward by the late Sir Francis Gal ton; a formula often spoken of as Gallon's law. According to this estimate, the hereditary traits of any given organism are so intermingled that we may assume as a general rule that offspring of a given generation will inherit about half their tangible traits from their parents, one-quarter from their grandparents, one-eighth from their greatgrandparents, and so on in decreasing scale from each earlier generation. Stated otherwise, according to this rule, we should be able by observation of the parents of any given organism, to see presented half of the traits of the offspring; but we may expect that the offspring will manifest, as the other half of their inheritance, traits that have come to them, through the process of reversion or atavism, from remoter generations of the ancestral strain. [17] LUTHER BURBANK And this obviously gives opportunity for the appearance of an enormous variety of traits in any given generation that were not manifested in the preceding generation. Thus any given individual has normally, as a moment's reflection will show, four grand- parents, eight greatgrandparents, sixteen ancestors in the generation before that, then thirty-two, sixty-four, one hundred and twenty-eight, and so on in a geometrical ratio with each remoter generation. So the normal ancestral clan of any one of us numbers more than a thousand different individuals within the relatively limited period of time compassed by ten generations. And, according to the estimate of Galton, to which numberless cases of atavism give force, certain traits and tendencies of each and every one of these ancestors may make themselves manifest in the personality of any given descendant. Galton's studies, upon which his formula was based, were chiefly made with reference to human beings, but we now know that the laws of heredity apply with equal force to all kinds of living organisms, including plants; and whatever the limitations of Galton's law as a precise formula, there can be little question as to the general truth of the principle that he invoked. Hence the value of that search in imagination [18] 1^1 ?=L *w §?*- 51.1.1? ***•*§ Li j IlW'iliTl ^sa s.-.-^1-02- "S ^2 5 5f ilt|;ft(i?^, ^s^ri LUTHER BURBANK for the ancestors of our cherry in their widely separated habitats and with their widely diversi- fied traits and habits. But of course in making practical studies for the development of the mental blue print with its forecast of qualities of our new cherry, we must perforce be guided largely by the observed qualities of the parent stock with which we deal. Precisely what were the qualities of the remote ancestors, we can only infer. But we can see for ourselves what are the qualities of the fruit before us. We know, then, pretty definitely what we may expect as to one-half the traits of a hybrid that will result when we cross two varieties of cherries in our orchard. The other half must be somewhat matter of conjecture, to be revealed by the actual product or, as is practically the case, by succeeding generations. What we actually do, then, in practice, is to take flowers from a cherry tree that has been observed to bear fruit somewhat earlier than neighboring trees, and with this pollenize flowers of another tree that has been observed to produce fruit of exceptionally good quality. Pollenation accomplished, by the method elsewhere described, we can only mark the branch for future identifi- cation, and await results. [20] PLANNING A NEW PLANT The seed thus secured will be planted next season, and in due course we shall have a seedling which, when grafted on another tree to speed its maturing, will come to blossoming time — after another period of waiting — and finally show us the first fruits of our experiment. From this fruit we shall raise a new generation of seedlings which will reveal to us beyond peradventure a varied assortment of ancestral traits that the parental forms of our first hybridi- zation did not show. And from among these diversified forms, we shall be able, by a long series of selections and new hybridizations, to make our way toward the attainment of our original idea. The precise steps and the varying details through which this may be attained, will be dis- cussed in other chapters. Here we are concerned only with the general outline, and, this having been presented, we may leave our cherry in this interesting stage of partial construction. To be sure we have not seemingly advanced very far toward our ideal in these two generations; but in this our case is only comparable, after all, to that of the architect, who, when he has planned a building that shall ultimately tower toward the skies, must be content to see the workmen first begin digging in the opposite direction, to lay foundations far beneath the earth's surface. [21] LUTHER BURBANK This matter of the very doubtful result of the first stages of a hybridizing experiment should be emphasized, because otherwise the amateur is pretty sure to become discouraged at the outset and to proceed no farther. Many an experimenter has given up a quest because when the two varieties of plant were crossed the offspring seemed inferior as to the desired quality to either of the parents. But the experienced plant breeder knows that this is very often to be expected and that he should not be in the least discouraged by this result. It is necessary to go on to the next generation before we can hope to discover the real possibilities of the experiment. The simple fact is that, where varieties or species of plants that differ markedly as to certain qualities are hybridized, the offspring very fre- quently seems to present what has been spoken of as a mosaic of characters rather than a blending. It may and very commonly does manifest, as regards any given quality, the influence of one parent seemingly to the exclusion of the other. A familiar illustration of the same rule may be observed when a person having black eyes marries one having blue eyes. It is obvious that no individual child of this union can have both black eyes and blue eyes. In point of fact, it is a [22] The Chinese Pear The Oriental pears have not the characteristic shape oj the European pear. The latter was presumably modified in shape through selection at a comparatively recent period. It is supposed that the original home of the pear was Central Asia, and that the fruit was carried into Europe in prehistoric times. The pear is characterized by having a fibrous or woody deposit in the skin, indi- cated by the dotted appearance oj this Oriental specimen. LUTHER BURBANK matter of common observation that the offspring in such a case will have dark eyes. But it has also been observed that the blue eyes of one of the parents may reappear in the second generation. The tendency to blue eyes was entirely subordi- nated or submerged in one generation, yet it was by no means eliminated, as its reappearance in the next generation clearly proves. Similar instances without number may be studied from our plant experiments; for example, the case of the white blackberry. If flowers of this kind are fructified with pollen from flowers of a blackberry of the usual color, the hybrid progeny of the first generation will all bear black fruit. The quality of blackness has proved prepotent or dominant, and the opposed quality of whiteness has been totally subordinated so far as this generation is concerned. But if these black hybrid blackberries are cross-fertilized, from the seed thus produced there will spring a generation of brambles, some mem- bers of which will in due season produce white fruit precisely like that of the maternal ancestor. Such, it will be recalled, was indeed the experience in the development of my new race of white blackberries. [24] PLANNING A NEW PLANT To instill good qualities of fruit into the inferior original berry, it was necessary to cross with the large and well flavored Lawton black- berry. The immediate result was seemingly to oblit- erate the white-fruiting tendency altogether. But a wide experience of similar instances led me to continue the experiment, which for the moment seemed to be carrying me away from my ideal of a white blackberry; and the principle of reversion came to my aid in the next generation and gave me, as will be recalled, a berry that combined the light color of one of its grand- parents with the size and flavor of the other. I have already suggested that it aids the memory, and helps to give tangibility to the facts, to recall the Mendelian phrase which speaks of blackness versus whiteness in such a case as constituting a pair of unit characters; naming blackness as the dominant and whiteness as the recessive feature; and which gives us assurance that a fruit which shows the recessive character of whiteness in the second generation will there- after breed true, thus affording us evidence of definite progress toward the ideal of our experi- ment. AID FROM UNIT CHARACTERS As the principles that govern these cases are [25] LUTHER BURBANK of very wide application, it follows that there is very great advantage from the standpoint of the plant developer, in the discovery of pairs of unit characters and the demonstration of their relation toward each other as regards dominance and recessiveness. An interesting illustration of this is afforded by the experiments made by Professor R. F. Biffin, of Cambridge University, in the successful attempt to develop a new race of wheat. Professor Biffin through a series of experi- ments showed that when beardless ears of wheat are crossed with bearded ones, the beardless condition proves dominant, so that all the off- spring are smooth-eared; but that the recessive quality of bearded grain reappears in the second generation. The same thing held true for various other pairs of unit characters, such as red chaff versus white chaff, red grain versus white grain, hollow stem versus solid stem, and the like. Professor Biffin was able to make an imme- diate practical application of his experiments through which he developed a new race of wheat that is proving of great economic importance. It appears that the best races of British wheat have been peculiarly susceptible to the fungous pest known as rust. There are, somehow, certain races [26] Pi! Ms <> 3 "^ T""" ii e ~~ >!L ""* ro "^ M^|Ki»i?i| ' ftHUi^O&f ^ ?^?.fis-l « i r • i* 9 ft i il * I: LUTHER BURBANK of wheat that are immune to the pest; but unfor- tunately these produce a very poor quality of grain. Professor Biffin found that susceptibility and immunity to rust constitute a pair of unit char- acters, in which susceptibility is prepotent or dominant. When he crossed the susceptible grain with the immune one, he therefore produced an entire generation of susceptible grain. His experiment had seemingly gone backward, quite as in the case of my first generation of white blackberries. But in the ensuing generation the recessive character of immunity reasserted itself; and, combined with this desired character, in a certain proportion of the progeny, there appeared the other desired quality of a good head of grain of fine quality. So by the application of this principle of the segregation and recombination of unit characters Professor Biffin produced a new race of wheat in two or three generations, and this new race of wheat breeds true. We shall see this principle illustrated over and over in connection with the long series of my plant experiments. In case of the wheat, as in that of my white [28] PLANNING A NEW PLANT blackberry, the process was relatively simple because we were dealing only with two pairs of unit characters. Moreover, the case of wheat is further simplified by the fact that this plant is self-fertilized and under conditions of cultivation has become a very fixed race, little subject to variation. When we deal with races of fruits that tend to vary almost indefinitely, and when further we are concerned with ten or a dozen unit characters, the matter becomes vastly more involved, as we have previously seen illustrated. But the amateur will do well to begin his experiments with simple cases, dealing with only a single quality, say a particular color of flower, that he may thus learn to distinguish the prin- ciples here enunciated. In due course he may go on to apply these principles to more complicated experiments in plant hybridization. But unless he learns at the outset that certain characters that are submerged in the first hybrid generation will inevitably reappear in the second, he will constantly blunder in his interpretation of tenta- tive results. On the other hand, when he has learned to gauge his second-generation hybrids correctly, he is on the highway to success as a plant experi- menter. [29] Common, French, and Burbank Mangolds The large double Burbank marigolds have been developed from the small single flowers through selective breeding. By cross-fer- tilizing different varieties, a tendency to variation is induced, and by preserving /or seed purposes only those flowers which vary in the desired direction, any given quality is accentuated, until finally such a metamorphosis as this is brought about. The rapidity of development varies with different plants, but many generations are required to produce such a transformation as that shown in this picture. PLANT AFFINITIES CHOOSING THE LINES OF LEAST RESISTANCE WHY do not plants hybridize in a state of nature?" a visitor asked me. "You seem to get most of your new varieties by hybridizing old ones. Why does not Nature take a leaf from your note-book and produce new species in the same way?" And I was able to inform my facetious ques- tioner, much to his surprise, that the method he suggested was one that Nature had practiced from the beginning, and is constantly practicing all about us. We were standing near the gateway of my Sebastopol place. I pointed out across the road. "Why, just over by the roadside," I said, "you may see for yourself precisely such an experiment in hybridizing as I have made in the case of thousands of plants. Do you see those tarweeds there? Doubtless you are familiar with them. [VOLUME III — CHAPTER II] LUTHER BURBANK "There are two common species growing there together. One of them has large showy flowers, the other small and inconspicuous ones. The botanist calls the large flowered species Madia elegans, and the other M. saliva. The two species do not look much alike, and some botanists even classify them in different genera. "If you look at all closely you will see that there is a third form of plant, bearing some resemblance to each of them, growing among the others, and I can assure you that this is a natural hybrid between the two. "If you examine this hybrid, you will find that its branches are less spreading than those of its large-flowered parent, although not upright like those of the other parent; and that the stem is stouter than that of either parent. As to foliage, the hybrid plants have larger and thicker leaves than those of the large flowered tarweed, more closely resembling the other species in this respect, but the ray-flowers are intermediate in size and shape as well as color, the reddish- brown that characterizes the flower of the more conspicuous parent being reduced in the hybrid to a spot just in the top of the tube. "So here you are probably witnessing the creation of a new species in nature. You, of course, are an evolutionist and therefore are [32] LUTHER BURBANK with wolves, and foxes, and jackals of every species than with any member whatever of the cat family. Similarly, all cats — tigers, lions, leopards, along with the domestic tabby — give proof, in the chemical constitution of their blood, of a common origin. And, bringing the com- parison still nearer home, the blood of man is more like that of the chimpanzee, the gorilla, and the orang, than it is like that of any other creatures; and the monkey tribes of the Old World are more manlike in the constitution of their blood than are the monkeys of the New World. Dr. Nuttall's experiments comprised sixteen thousand individual tests, with a total of at least 586 species of mammals, birds, reptiles, batrachians, fishes, and crustaceans, coming from all parts of the globe. The biological implications of his experiments have been commented upon as follows: "Doubtless some hundreds of thousands of years have elapsed since the direct ancestors of men branched from a common stem with the direct ancestors of the gorilla. There has been no blending of blood in the intervening centuries. Cats have been cats and dogs dogs from geological epochs so remote that we hesitate to guess their span in terms of years. So the intimate chemical [50] PLANT AFFINITIES qualities that denote man or ape or cat or dog, each in contradistinction to all the others, must have been transmitted unmodified through count- less thousands of generations. "It taxes credulity to believe that such intan- gible properties could be transmitted unmodified through the blood streams of such myriads of individuals; but the evidence of the test-tubes proves that this has been done. "What makes the marvel greater is the fact that the bodies of the animals have meantime been so modified as to develop utterly divergent species — for example, the lion, the tiger, the puma, the leopard, and the house cat; different types of dogs, wolves, foxes and their allies. But in each case some intangible quality of the blood remains unchanged to prove the common origin. Blood is indeed thicker than water." The bearing of these extraordinary experi- ments upon the case in hand will be obvious. If animals carry in their veins generation after generation, through untold thousands of years, these intimate chemical conditions, then the same thing may well be supposed to be true of plants. And so the affinity shown between species that can be hybridized, and the antagonism between species that refuse to hybridize, can be explained on the basis of a fundamental intrinsic quality of [51] LUTHER BURBANK the protoplasm that is the foundation substance of life. This gives us a more profound and compre- hensive appreciation of the word "affinity" as applied to various species of plants than we could otherwise have. It also makes it in a measure comprehensible that the traits of remote ancestors should be carried latent in the tissues of the germ-plasm, as we have seen that they are carried, for untold generations. THE CONTINUITY OF THE GERM-PLASM This germ-plasm, which is the connecting link between one generation and another, is passed on, according to the prevalent idea, from parent to offspring, generation after generation, subject only to such modifications as may from time to time be imposed through environing influences. The physical mechanism that underlies this transfer we shall have occasion to examine in another connection when we discuss at some length the theories of heredity. For the moment it is enough to reflect that as the offspring in each successive generation spring from the substance of the parent, the germ-plasm may be thought of as a continuous stream uniting the remotest ancestor of any given strain with the most recent descendant. [52] A Mosaic Leaf When a purple-leaved plum is hybridized with a green-leaved plum, the hybrids of the first generation have green leaves, but purple leaves reappear among the varying hybrids oj the second generation. Occasionally, however, a cross between purple-leaved and green-leaved varieties, where the varieties lie near the limits oj affinity, results in a condition that might be described as a mosaic, the leaves par- taking oj the characteristics oj both ancestral strains. This picture illustrates such a case. LUTHER BURBANK Every tree in the orchard, for example, car- ries within its tissues a portion of protoplasmic matter that has come down to it through an almost infinite series of growths and divisions in unbroken succession from the first tree that ever developed on the earth — or, for that matter, from a vast series of more primitive organisms that were the progenitors of the first tree. And while this stream of primordial proto- plasm has been changed by an infinitesimal quantity in each successive era, it has retained even to the present the fundamental character- istics that it had from the outset. That such is the case seems little less than a miracle; that an almost microscopical speck of protoplasm which we term a pollen grain should contain the potentialities of thousands of genera- tions of ancestors, and should be able to transmit them with such force that the seed growing from the ovule fertilized by that pollen grain will inevitably produce, let us say, an apple tree, not a pear tree or a plum, is beyond comprehension. Yet we know it to be true. And so the plant hybridizer who consciously merges two different protoplasmic streams when he brings the pollen of one flower to the pistil of another, participates in what must be considered the most wonderful of all experiments. [54] PLANT AFFINITIES He brings tokens out of an almost infinite past to blend with the divergent tokens of another ancestral stream no less ancient. And it is not strange if he feels a certain impulse of elation when he reflects that his con- scious efforts have thus brought together ancestral tendencies that have long been separated and that now will appear in new combinations- stimulating such interplay of life-forces as may bring into being plant forms that may be described, without violence to the use of words, as new creations. —That the intimate record of cousinship, in all its grades, should be permanently graven in the protoplasm of every living thing, is a thought-com- pelling biological revelation. A Shirley Poppy — Showing Reproductive Organs The petals oj a flower are designed to attract insects. The essential organs are the pollen-bearing stamens and the pistil joining the ovule at the center oj the flower. This picture shows the large number of stamens of the poppy, each with a terminal anther, bearing pollen, growing in a circle about the pistil with its curiously rounded end, called a stigma, designed to receive the pollen. The office oj the insect is to transfer pollen from the stamens of one flower to the pistil of another. PRACTICAL POLLENATION A SURVEY OF WORKING METHODS ONCE upon a time — it may have been about the year five million B. C. — a plant imbued with nascent wisdom made a tacit compact with a fellow creature of the world at large that was fraught with strange and fateful meanings for races of beings yet unborn. The fellow creature in question was at that time probably the most highly developed citizen of the world, although in modern terminology he would be termed "merely an insect." The com- pact the plant made with him was to the effect that one should manufacture sweet nectar and freely supply it as food; and that the other in return should carry the fructifying pollen grains from flower to flower. Doubtless no more important compact was ever entered into in the history of animate crea- tion before or since. [VOLUME III — CHAPTER III] LUTHER BURBANK For out of this compact grew the rivalry that stimulated development and made possible the evolution of the whole race of plants that bear beautiful flowers and exhale sweet perfumes. But for this eventful alliance, there would never have developed in the world a conspicuously colored or a scented flower of any kind. And it requires no argument to show that a world without beautiful and sweet-scented flowers would be a world robbed of a large share of its attractions as an abiding place. But that is not all. The alliance between insect and flower did not merely suffice to give us things of beauty. It bespoke utility as well. It made possible the bringing together of germ-plasms from plants growing far apart, thus insuring virile and variant strains; and this determined in large measure the amount and direction of the evolu- tion of the highest orders of plants. For it must be observed that, with rare excep- tions, the higher plants are precisely those that long ago entered into this cooperative scheme whereby they trusted their fate absolutely to the insects. They hazarded much — for if anything should lead to the destruction of a few insect races, entire orders of plants would suffer race suicide. But if they risked much, they also profited much; for the cross-poll enizing effected [58] The Stigma of a Poppy, Greatly Enlarged The stigma of the flower may be variously modified to facilitate reception of the pollen. This picture shows the curious arrange- ment in the case of a poppy. The pollen grains deposited on this stigmatic surface send out little tubes that penetrate the stigma and ultimately make their way to the ovule or seed case, carrying the nucleus that unites with the nuclues of the egg cell, thus effecting fertilization. Each egg cell is fertilized by a single pollen nucleus. LUTHER BURBANK by the insects afforded the constant stimulus to variation that underlies all evolution, and enabled the plants that entered into the coalition presently to outstrip their fellows. Wherever you find a tribe of plants that shows great diversity of form, large numbers of species, and ready adaptability to improvement, you will as a rule find a tribe of so-called "entomophilous," or insect-loving flowers, dependent upon the winged messengers for the consummation of their matings. Vast responsibilities then were implied in this coalition of the plants and the insects; but the results have justified the hazard. PLANTS THAT DID NOT JOIN THE UNION We shall presently see illustrated in detail the curious adaptations of form and color and structure to which the plants of various species were led in their rivalry to secure the good graces of the insects and thus to make sure of perpet- uating their species. Every blossom of the entire orchard, every flower of the garden, and with a few exceptions all of the vegetables under cultivation furnish illustrations in point. But it should be recalled that there are large numbers of plants of a lower order that from the outset refused to enter into the coalition, and that even to this day have [60] PRACTICAL POLLENATION declared themselves independent of the plant- insect union. Plants of this non-union clan are the entire races of lowly mosses and lichens; a goodly num- ber of aquatic forms that maintain the appearance and manner of their remote ancestors; and the familiar tribe of ferns; and some of the trees which depend mainly upon the wind. All of these, and a large company of forms less familiar to the amateur, have obstinately retained throughout the ages the primeval habit of propagating their kind not with immobile pollen grains but with the aid of self-moving germ-cells. These motile germ-cells, of microscopic size, find their way through the water — supplied in case of land plants by a film of rain or of dew — from one plant to another, and effect cross-fertilization without calling in the aid of any allies. They do not need to attract insects, and so they have not adopted the system of advertising through the development of large and showy blossoms and nectar cups to which the members of the plant- insect alliance are obliged to resort. But if the lowly plants thus maintained their independence, they have done so at a very great sacrifice. They are not more independent than they are unprogressive ; and indeed they are unprogressive [61] Pollen-Bearing Pumpkin Blossom Many plants bear pollen and ovule on separate blossoms, so that self-fertilization is impossible. They are said to be dioecious. This picture shows a pumpkin blossom with the modified and coalesced stamens at its center bearing a quantity of pollen. The insects in visiting the blossom carry the pollen on their bodies or feet and transfer it to other flowers. Seed-Bearing Pumpkin Blossom This blossom is the mate oj the one shown on the opposite page. The stigma to receive the pollen occupies the same position that is occupied by the pollen-bearer in the other flower. The bulbous growth at the base of the flower is the ovary, or seed case, which, if the flower is fertilized, will develop into a pumpkin. LUTHER BURBANK precisely because of their independence. The method of cross-fertilization that they have adopted does indeed enable some of them to blend the strains of different individual plants; but in every instance the parents must be growing in the immediate vicinity of each other. Except by the accidental and most unusual transfer of a plant through the agency of a passing animal, there is hardly the remotest chance of effecting cross-fertilization between individual mosses or lichens or ferns growing in widely separated regions. But we have already seen that it is precisely this blending of traits brought from parents growing under different environing conditions that is chiefly responsible for making plants vary and furnishing the materials for evolutionary progress. So it goes without saying that the plants that are restricted, in the choice of possible mates, to individuals growing under the same conditions to which they themselves are subjected, cannot expect to change rapidly and therefore do not evolve in any such ratio as plants having the other habit. And in point of fact we find that the plants that retain this primitive custom of fertilization with the aid of motile germ cells, acting quite inde- pendently of insect or wind, are plants of a [64] Cross-Section oj a Cactus Blossom This is what is called a perfect flower — that is to say one that has both stamens and pistil. The stamens are grouped in a circle about the pistil, as in case of the poppy, this being the typical arrange- ment. The picture shows the seed case or ovary at the base of the pistil. Each ovule must receive the nucleus of a pollen grain or it will not develop. Where the stamens are thus clustered about the pistil, cross-fertilization is usually prevented by the maturing of the two sets of organs at different periods. LUTHER BURBANK low order of development, showing relatively little diversity of form and small capacity for adaptation. The most conspicuous of them with which the ordinary observer is familiar, namely the ferns, bear a striking resemblance in contour to plants of the remote Carboniferous Era, traces of which have been preserved in the coal beds. And there can be no doubt that this persistence of the primitive form has been largely due to the special method of fertilization which the ferns have retained. If it be permitted to carry personification one stage farther, we might say that the ancestors of the ferns belonged to a conservative family, jealous of its independence, and unwilling to enter into outside alliances. And the penalty of conservatism here, as so often in the range of human experience, has been racial stasis. PLANTS THAT HAVE LEFT THE UNION It would appear, however, that there are certain races of plants that were once members of the plant-insect alliance but which are now no longer in the union. These apostates include two quite different tribes of plants. On one hand there are numerous gigantic [66] PRACTICAL POLLENATION trees that no longer depend upon insects for the fertilization of their flowers. On the other hand there are little cowering plant-waifs that nestle close to the earth and which, in quite a different manner, also assert their independence. The trees that have thus revoked the treaty of alliance include such familiar forms as the pine, the oak and the walnut. These trees, and a goodly number of their fellows, long ago declared against further coop- eration with the insect, and adopted the method of producing large quantities of pollen and scattering it in the air to be carried by the wind to the pistillate flowers, which in some cases grow on neighboring branches and in other cases on quite different trees. The method is in one sense wasteful, inasmuch as it involves the production of vast quantities of pollen, only an infinitesimal portion of which will ever come in contact with a receptive pistil. And of course the production of this pollen must draw heavily on the energies of the living substance of the tree. But on the other hand the tree that thus depends upon the wind rather than upon the insects is under no necessity to develop large and conspicuously painted flowers. Nor need it [67] Two Gourd Blossoms to be Pollenated The Gourd belongs to the same family as the pump- kin, and, like that plant, bears the staminate and pistillate flowers separately. This arrangement makes it very easy to effect hybridization. Gourds and other squashes are cross-fertilized so readily that different species must be grown far apart if the strains are to be kept pure. Pollenating the Gourd Blossoms To effect fertilization, nothing more is necessary than to pluck a staminate flower, pulling away its corolla, to expose the pollen-bearing surface, and to insert this in the pistillate flower, gently dusting the stigma with pollen. The facility with which this may be effected makes the members of this family very attractive to the amateur experimenter. LUTHER BURBANK produce nectar to feed the insect allies, since these have been renounced. And it may very well chance that the saving of energy thus effected more than counterbalances the waste through excessive pollen production. At all events the plants that have adopted this system of pollenizing give evidence that their plan is not a bad one in the very fact of their extreme abundance. Moreover the "wind-loving" or "anemophilous" plants, as the botanist terms them, have not only produced a great variety of species and vast numbers of individuals, making up the bulk of our forests, but the individuals themselves are of such virility of constitution as to attain gigantic size. Indeed a moment's consideration makes it clear that the plants that had depended on the wind rather than on insects for fertilization are quite in a class by themselves in the matter of size, inasmuch as they constitute the bulk of our forest trees. This relation between size and habit of spreading the pollen broadcast on the winds cannot be altogether accidental. But whether the trees grew large because they had given up the alliance with the insects, or whether they gave up the alliance because they were growing large, it would be hard to say. [70] PRACTICAL POLLENATION We know that, in the main, insects tend to keep near the surface of the earth, and it may be that the plants that tended to grow very tall were relatively neglected by the insect messengers. But on the other hand, there are insects that haunt the highest trees, and we can hardly doubt that had even the tallest of plants desired to retain the services of insect messengers, races of these would have been developed that would have proved equal to the most exacting demands. What seems on the whole most probable, then, is that the trees have changed their allegiance from insect messengers to wind because of the very nature of the conditions under which they grew. By raising their heads high and higher into the air they obviously put themselves more in contact with the wind and thus make it increasingly possible to spread their pollen broadcast across wide stretches of territory. As a matter of fact we know that the pollen of pine trees in particular may be carried almost in clouds for scores and even hundreds and hundreds of miles. So there is every opportunity for the cross- fertilization of individual trees growing in widely separated territories; and there is therefore no restriction put upon the possibilities of progress [71] LUTHER BURBANK and evolution for these large-growing plants in penalty for their renunciation of the services of insect messengers. The case of the trees, then, simply illustrates the fact that there may be more than one way to effect a given purpose, and that a change of method may be no barrier to progress, even though the abandoned method still remains an admirable one for a vast coterie of organisms of slightly different habit. SELF-FERTILIZED PLANTS But the case of the other company of plants that have back-slidden from the insect alliance is altogether different. The plants in question do not make up any great conspicuous tribe, comparable to the forest trees, but are a miscellaneous company of lowly vegetables of unrelated families. Familiar examples are the wheat of the fields, peas and beans in our garden, and a certain number of the more obscure species of violets. The jewel weed, the fennel, the rue, and the nettle, are other somewhat less familiar yet not uncommon tribes of plants whose flowers are habitually self-fertilized. There can be no question that these plants are the descendants of tribes that were at one time members of the plant-insect union. The fact that [72] Strawberry Blossom Ready /or Pollenating Some varieties of strawberries bear perfect flowers as illustrated in this specimen, and others have blossoms that bear the pistils separately. It is necessary, in cultivating the straw- berry, to bear this in mind, and ij a pistillate variety is planted there must be pollen-bearers in the neighboring rows, otherwise there would be no crop oj berries. The bees are depended on to effect the transfer of pollen. In the perfect flowers, the stamens and pistils mature at different periods, to guard against self -fertilization. LUTHER BURBANK most of them retain more or less conspicuous flowers proves this beyond question. In the case of the wheat, which might be thought a possible exception, there is the evidence of certain species of wild wheat, growing to this day in Palestine, which have only partially renounced allegiance to the insects, still putting forth flowers that on occasion may be cross-fertilized with their aid or with that of the wind. Just why these various plants of different families have departed from the custom that has served their fellows so well, would be interesting matter for conjecture. Perhaps the most plausible suggestion is that the ancestors of the plants that now have closed flowers and thus depend exclusively upon cross-fertilization had fallen on evil days in which there was a dearth of insect messengers in the regions they inhabited. The story of the starved martins, told in an earlier chapter, furnishes a striking illustration of the fact that insects that ordinarily are abundant may in any given season fail to put in their appearance. And even if the insects themselves are abun- dant, the weather conditions, in a given season, may be such as to make it almost impossible for them to carry out their bargain by transferring [74] .s p| a n*& 2-. '£ a** 2 5 ;« to § o _ A • »•*• % 8 ^^§'3 5 a. LUTHER BURBANK This selected seedling he may nurture and use as part of his equipment for further experiment just as you retained the letter "E" as marking the beginning of your success in spelling the word "evolution." And as the plant developer continues his experiment with successive hybridizings and successive selections, he will be able in later generations to find individual seedlings that combine successively more and more of the qualities he is seeking. When, finally, he reaches the stage where the parent forms have between them all the desired qualities in superlative degree, he is somewhat in the position that you were in when only two of your lettered blocks remained in the bag. There is at least an even chance that he will find among his seedlings of the next generation one that will approximate his ideal, even though the number from which he selects is far smaller than the earlier groups. Thus by advancing step by step and using the ground gained as a new starting point the experimenter attains his end with comparative celerity, even though there would have been scarcely more chance of attaining that end with a single experiment than you would have had of spelling out the word "evolution" at a single series. [104] QUANTITY PRODUCTION But it must be fairly remembered that the probability of success is enhanced if at any of the earlier stages of the work you have opportunity to select the best plant among a large group instead of being restricted in choice to a few individuals; just as the chance of securing the block you seek in each successive drawing increases with the number of tests you are permitted. And in point of fact, this, or something like this, is the actual method in which the experiments of the plant developer are carried out, whenever he is attempting to construct a new fruit or flower or vegetable having a number of specified or clearly imagined qualities. In such a case, the wise experimenter does not hope to secure ideal results with a single hybridization; he seeks to group desired qualities of his flower or fruit together through successive crossings and selections. Keeping one supreme quality in mind and perhaps two or three others in the immediate background, he makes sure of first one and then another of these qualities, adding to them by successive crossings and selections and thus although advancing, as it were by indirection, and at first seeming to advance but slowly he may ultimately work with increasing certainty and approach his goal somewhat rapidly. [105] Germinating Seeds in Damp Cloth Seeds of various kinds, arranged in loose rows as here shown, may be rolled in a damp cloth and thus germinated preparatory to planting. This is one of the numerous shortcuts that Mr. Burbank practices. The method is some- times used merely to test the viability of seeds from different lots. ^^^^^mf^mummm *. ,. ' Germinated in Damp Cloth This shows the method illustrated on the opposite page, at a later stage. The four different groups of seeds have been germinated in the same roll of cloth, without mixing, and they are now ready to be transferred to the soil, either in the greenhouse or in the field. LUTHER BURBANK For example, our first cross, say in the case of a prune, may be made between two varieties that both show a fair quality of fruit. Careful attention to the result will guide us in the matter of the next experimental crossing. We soon discover which qualities are prepotent, and which tend to remain latent, and by selecting only individuals that show a tendency to vary in the desired direction, we introduce an element of direction into the experiment. I am accustomed to speak of this as "momen- tum of variation." We do not always know why a certain plant tends to vary in a given direction, but we may observe the fact, and the wise experi- menter is always on the lookout for this tendency, and ready to avail himself of the advantages it offers. Technical workers sometimes give the name "orthogenesis" to this tendency to vary in a certain direction, which I speak of as the plant's "momentum." Whatever aid we may gain in this way, how- ever, the manner of our advance is often devious. In fact, it is very likely to be somewhat comparable to the progress of a sailing ship which tacks this way and that, and which at times may seem to be progressing in the wrong direction, yet which in the end forges ahead. Take by way of illustration the case of our [108] ' ' 'lK5t * Pf jr* ^^ •C ;i * * *~* * Pumpkin Seeds Germinated in Wet Newspaper Sometimes Mr. Burbank employs a piece oj newspaper instead of a cloth to germinate seeds, merely wrapping the seeds loosely in the moistened paper. The efficacy oj the method is demonstrated in this picture. Seeds thus germinated will make rapid growth when transferred to the soil. LUTHER BURBANK stoneless plum. We discover soon that the stone seed is prepotent or dominant, and stonelessness latent or recessive. So we must be prepared to see the progeny of our first generation of hybrids all produce stony fruit. But a knowledge of the tendency of latent or recessive characters to reappear in successive generations comes to our aid, and we go on with the experiment with full confidence, even though for the moment we seem to be going backward rather than forward. In due course the second generation of plums appears with a number of stoneless specimens, the latent character having come to the surface. But these lack many of the good qualities that our perfected fruit must have, and in order to breed these qualities into the stock we must make a new cross; and this will involve the breeding in again of the tendency to bear stone fruit. So in three generations we shall find ourselves, as regards the essential quality of the stony seed, somewhat further back than we were in the beginning. But, on the other hand, our third generation fruit, even though it has a stony seed, has qualities of flesh that its stoneless ancestor altogether lacked; and in the fourth generation we shall be prepared to find individual seedlings that bear stoneless fruit of greatly improved quality. [110] QUANTITY PRODUCTION In each successive generation, then, we are dealing with better material — getting the chances grouped, if you will. WINNING AGAINST ODDS But, in a sense, we are running counter to the trend of heredity, because vastly the great proportion of the ancestors of our plum were bearers of stoned fruits. And so we must continue re-shuffling and dealing over, as it were, and watching results. We may lose in one generation what we gained in the generation before as regards the matter of stonelessness; even while on the whole advancing toward the production of a fruit of desired quality. But just in proportion as our ideal calls for the combination of numerous good qualities, does the attainment of that ideal become difficult. Even when, at let us say the fifth or sixth generation, we interbreed individuals that have the desired quality of stonelessness, we shall not at once secure what is desired; because our seedlings combine so many ancestral traits that they will not breed true. Even though they are all stoneless, there will be a great variation as to other qualities, and it is only by dealing with large numbers of seedlings that we can hope to find one or two that will show the desired com- bination of traits in high degree. [in] LUTHER BURBANK Perhaps the comparison may be thought somewhat whimsical; but I am led to make it because I thought it might serve to suggest the complexities and difficulties that attend a plant- breeding experiment that involves the blending of numerous desired characters. And the lesson that I wish pre-eminently to inculcate is this: You must make many experi- ments at plant-breeding before you can hope to secure the combination — -the sequence of qualities —that you desire. THE LOGIC OF QUANTITY PRODUCTION Now note the application: Each individual seedling of a hybrid strain represents a unique combination of ancestral traits, and constitutes in itself a new and unique experiment — equivalent to an independent deal of the cards. So the prob- ability of securing what we seek will be somewhat proportionate to the number of seedlings. This is particularly true in the case of such variable plants as the fruit trees of our orchards. The case is far simpler when we are dealing with plants that vary little in their qualities, or where we are breeding with only a single pair or two pairs of unit qualities in mind — say "hardness" of kernel and immunity to rust, as in Professor Biffin's experiments with wheat; or good flavor and whiteness as in my white blackberry. [112] HS-at •^ Ta m all 3 to rr§rsr & $ ~ ft--* «9 3 **> O- LUTHER BURBANK But where the varied traits sought to be com- bined in a Shasta daisy are in question; or the many qualities of a commercial cherry or prune, the case assumes new complexities. Hence it is that my records tell of tests applied to about half a million seedlings of the daisy; seven and one-half million seedlings of various plums, and the like. Hence also the constant necessity of what my neighbors speak of as ten-thousand-dollar bonfires in my orchard, when we burn seedlings that have been inspected and found wanting. To burn 65,000 hybrid blackberries in one pile, as I once did after saving perhaps half a dozen individual vines, seems like willful extravagance to the casual observer, but it is an unavoidable incident in the search for perfect fruits. Such prodigal use of material implies a large measure of experience in the handling of seeds and the growing of seedlings. In point of fact, it might be said that this is the most important part of a plant-breeder's task, so far as the practicalities of experiment are concerned. It is part and parcel of his daily routine. It is highly desirable, then, that the would-be experimenter should gain a clear understanding of the essentials of method of caring for seeds and cultivating seedlings. So it is my purpose in the [114] "Flat" With Layer of Gravel A layer of gravel is put at the bottom oj the germin- ating box to facilitate drainage and aeration, both oj which are very essential to the proper growth of the plant roots. Mr. Burbank regards this detail as very important. LUTHER BURBANK succeeding pages of this chapter to give a few practical hints as to various aspects of the subject. Thus summarized, the lessons I have learned in the hard school of experience may enable the reader to avoid some pitfalls and to make certain experimental shortcuts. KEEPING SEEDS OVER WINTER To begin at the beginning, let us note that the preservation of .seeds over winter calls for careful attention. All fruit seeds except those of apricots and almonds, when removed from the fruit, are at once placed in slightly moist, coarse sand or fine gravel or in sterilized sawdust In warm climates the boxes containing the seeds are then buried on the shady side of a building or tree where they will become neither too dry nor too wet. The object is to keep the kernels as nearly as possible in their original condition. If tree seeds, especially those of the cherry, the pear, and the plum once became thoroughly dry, it is difficult, and in some cases impossible, to induce them to germinate. An important function of the pulp of these fruits, in the original wild state, was, presumably, to keep the seeds moist until the season for germ motion. I have elsewhere called attention to the [116] ft-a S. ^ •r? - -g*'-0" Ills 2 a ^ LUTHER BURBANK exceptional difficulty of keeping stoneless plums and prune seeds in condition for growing. Not having the natural protection of the shell, they tend to germinate too early, and of course they are peculiarly subject to the attacks of insects and of fungous diseases. Such seeds may best be placed in cold storage as soon as collected and cleaned, and kept at freezing temperature. Seeds thus cared for will sometimes germinate almost as quickly and readily as beans or corn. They must not be planted too early in the spring, lest their too prompt germination subject them to injury from late frost. Incidentally, I may note that grafts sent to me from a cold climate have often been observed to start with greater promptness, and grow better than those from our own immediate vicinity where the winters are mild. Cold seems to rest the tissues and prepare them for rapid growth, just as treatment with narcotic drugs has been observed to do in certain interesting experiments that will elsewhere be referred to more at length. OUT OF DOOR PLANTING In California, plum seeds are usually planted in January or February, in a little furrow about an inch deep. A furrow may be made accurately and expeditiously with the aid of a triangular bit of board an inch or so wide nailed across another [118] *» ft *. ^ r s o £><£ *l - rfi'll'a." LUTHER BURBANK longer piece, so that when drawn along a garden line it makes a narrow furrow of exact width and uniform depth throughout. Plant the seeds about one-half inch to an inch apart, and cover with a thin layer of soil; then fill the furrow with sawdust. This is an important matter with cherry and plum seeds, especially with the stoneless ones which must be given every inducement to push through the soil. A heavy, compact soil placed over cherry and plum pits prevents a large number from pushing up to the light. For this reason a sawdust covering is preferred, and it also regulates the moisture with exactness, allows for sufficient aeration, and equalizes the temperature. Moreover, the saw- dust is distasteful to slugs, thrips, cut-worms, and other insect pests. Peach, nectarine, and apricot seeds are planted farther apart and a little deeper; quince, pear and apple seeds are planted about the same as plum seeds, both as to distance and depth, or in large lots may be rather thickly sown in drills or furrows six or eight inches wide and eighteen to thirty inches apart. For growing seedlings of conifers — pines and their allies — cold frames or shallow boxes are used filled with mellow sandy loam; or the seed may be sown broadcast or in rows in cold frames [120] "Flats" With Sprouting Seedlings All manner of seeds are sown in the "fiats" and Mr. Burbank has so perfected the method that he usually sprouts ninety-nine seeds in a hundred. The same method is used for the commonest flower and the rarest exotic. "Flats" grow- ing hybrid cactuses are side by side in his green- house with those containing orchard fruits, and flowers of many species. LUTHER BURBANK without boxes. The object of the cold frames is to shelter from hot sun and drying winds and in cold climates to prevent freezing. If the season is short or if warm weather comes on suddenly, it is sometimes desirable to soak seeds in water before planting. After being in the water several hours they should be drained and set in a warm place where germination can start quickly. In this way growth may sometimes be advanced by a week or more. But such forced germination is not usually necessary or desirable. If carried too far before planting, it endangers the growth. On the other hand, the very early plants often escape cut-worms and other insects by attaining a fair growth before these pests put in an appearance. BOXES FOR SEEDLINGS Valuable plants to be grown in large quantities from rare seeds, may best be started in small boxes or "flats" indoors, under glass or in sheds made of laths or slats so spaced as to allow free entrance to air and sunshine. Boxes of the right design and construction are far better for this purpose than pots or earthen pans. The boxes or "flats" that I have used for twenty years are made of redwood lumber. Where this cannot be obtained, cypress is nearly as good, but soft pine is not durable and should [122] The Seedlings in the Greenhouse 'Flats" are here shown at a later stage, when them c * i.uto uic uc re snuwn at a later stage, wnen some of the seedlings have been transplanted to give t room, and have attained considerable proportions. Much time is saved by thus developing the seedlings in the greenhouse over winter. LUTHER BURBANK be avoided. Eighteen inches square, outside measure, four and one-half inches deep, inside measure, is a good size. Two opposite sides are of common board lumber three-quarters or seven-eighths of an inch thick; the other sides, are a little less than half an inch thick. The bottoms are made of redwood "shakes" which are about one-fourth of an inch thick; two or more spaces of an eighth of an inch being left for drainage. Across the bottoms are nailed three strips which add rigidity and strength as well as affording better ventilation and drainage. After all the parts are carefully fitted, the joints are sometimes dipped in linseed oil, before being strongly nailed together. This gives durability and tends to prevent the nails from rusting out. These redwood boxes may be used for many years if sterilized once a year by being placed for about three or four minutes in boiling water. A suitable soil is the first requisite in raising seedlings in boxes. The mixture which I have generally found best for use in the early winter for raising seedlings in boxes in the greenhouse, is compounded about as follows : One-half clean, rather coarse, sharp sand; with about forty per cent, of some good pasture or forest soil which generally contains more or less leaf mould. To [124] 3 * LUTHER BURBANK this is often added ten per cent, finely powdered moss or peat. These mixtures, with the addition of about one or two per cent, of fine ground bone meal or superphosphate, make soils in which seeds of almost any kind of plants from any part of the earth will germinate. Seedlings thrive in this soil until they are ready for transplanting. If seeds of choice plants are to be grown, the soil is sterilized by a thorough scalding to destroy any fungus or insect pests. Usually we find it suits the plants better if a part of the soil last prepared is left over for use with the new mixture, like yeast for a loaf of bread, and I always prefer to have a little of the old on hand for this purpose. Common sharp sand, if the right texture can be obtained, is far better for cuttings than the soil just described. The sand found along creek or river banks is generally free from injurious insects or fungous diseases. But for rare cuttings and very choice seeds, this should be rinsed by pouring large quantities of water through it, at the same time stirring or jarring the material. In filling the boxes, coarse gravel, such as will just pass through a half -inch mesh, or a little smaller, is placed one-quarter to one-half inch deep over the bottom of the box. This ensures perfect drainage and sufficient aeration, both of [126] iftlffiftfr I iiii?Mit8 - & g-O «. •*"JS 4> ~ C i'fs S.S|| H-: I l1li;^t-:i4lrP i*HHj2hft ^ *1 ixl 11^1^1 1 ^^ -5s «. -e^stjScooK LETTING THE BEES DO THEIR WORK structure of the flower is such that an insect as it passes down the petals on its way to the nectary, brushes against the anthers, and dusts off the pollen. As the insect passes out, the stigma-shield protects the stigmatic surface completely. But as the insect visits another flower, its pollen-covered back comes in contact with the edge of the stigmatic shield and the pollen is scraped off against the receptive surface. These, then, are familiar illustrations of the really wonderful adaptations through .which it comes to pass that the bees carry out their part of the ancestral compact that ensures the plant such interchange of pollen as is essential to racial progress. Perhaps the most alluring feature of the entire coalition is that the bee performs its all-important function unwittingly in the course of the quest of sweets that appeal to its appetite. There is no compulsion in the matter; the plant depends upon the more powerful influence of persuasion. And to add to the satisfactoriness of the entire arrangement, from a human standpoint, it must be recalled that the efforts of the industrious insect, which thus make possible the work of the plant experimenter, result at the same time in storing the nectar gathered from the flowers to form one of the most delectable of foods. [219] Patagonian Squash The members of the squash tribe are readily fertilized, as illustrated in earlier pictures of this volume, but the hybrids tend to show extreme variation, and they are very difficult to fix. Mr. Burbank has shown, however, that it is not impossible to fix The new varieties by carejul selective breeding. here shown was developed from seed sent Jrom squash South America, and weight It has almost the solidity oj a cannon ball. FIXING GOOD TRAITS How TO HOLD A RESULT ONCE ACHIEVED IT IS traditional that you cannot teach an old dog new tricks. The maxim applies with full force to old plants. You may bend the twig and make a permanent twist in the future tree; but the hardened stock of the matured branch will return persistently if bent, and will break rather than change its form. Now there is something like the same differ- ence in flexibility between yoilng and old races of plants. Here is a variety of plant that has been developed in the orchard or garden, under man's influence, in the course of the past few generations. It tends to vary, and its progeny may be made to adapt themselves to different conditions; by selection, they may be developed into divers and sundry new races. But yonder palm tree has no such propensity to vary. Its ancestors have remained substantially [VOLUME III— CHAPTER VII] LUTHER BURBANK unchanged, true to their racial type, generation after generation, for untold centuries. It repre- sents an old, fixed, conservative stock. No one knows how to make it change, except within the narrowest of limits. There is a very essential time element, then, that is instrumental in determining the fixity or variability of a race of plants. A plant that has been bred true to a given type for long periods of time, as is the case with the generality of wild plants, will breed substantially true from seed, and as a rule will maintain its racial type even if transplanted to new surroundings. But, on the other hand, the generality of cultivated plants are of mixed ancestry. Man has attempted within recent generations, to change them and adapt them to his needs. He has constantly been hybridizing them, or placing them under conditions that resulted in their hybridization through the visits of bees; and he has selected and cultivated the individual speci- mens that tended to vary, and thus has fostered the habit of variability rather than that of fixity of character. In the case of most orchard fruits, as we have had occasion to observe more than once, so many strains are blended that propagation from seeds is quite out of the question; unless, indeed, it be [222] FIXING GOOD TRAITS desired to secure seedlings of varying qualities in the interests of experimentation, or in the attempt to develop still other varieties. One might plant a thousand acres with seeds of the Baldwin apple, without perhaps producing a single plant that would precisely duplicate the qualities of the fruit from which the seeds were taken. And the same thing is true in greater or less measure of the majority not merely of orchard fruits but of cultivated plants in general. The notable exceptions are annual plants that are habitually grown from seed, such as melons and peas in the garden, and the great tribe of cereals represented by wheat, oats, rye, and barley. The reason why all of these breed true from seed is that they are necessarily propagated in this way alone, and it has been essential that fixed races should be developed. Mankind depends largely upon the cereals for food, and his existence would be altogether pre- carious could he not have reasonable assurance that when he sows grain of a certain quality, he will secure a crop of grain of similar quality. The fixity of character of the cereals and various other plants, including peas, and beans, is enhanced and assured by the fact that the flowers of these plants are habitually self- fertilized. If you examine a head of wheat at [223] LUTHER BURBANK the right stage, you will find that you must pull open the little bracts in which the flowers are encased, in order to make the stamens and pistils visible. Under ordinary circumstances, insects cannot find access to them. The wind has no influence over them. Their normal habit is to fertilize the pistil of each individual flower with pollen from the stamens that grow within the same closed receptacle. This is inbreeding of the closest and most intimate character, and there is obviously no ordinary opportunity to introduce the element of variability which, as we have seen illustrated over and over, cross-fertilization brings. So the essential qualities that make wheat valuable have been aggregated in a few fixed combinations, and the resulting varieties of wheat, differing not very widely from one another, are never crossed, unless by artificial means to meet the special needs of the plant developer. They remain fixed because they are of pure lineage. MIXED ANCESTRY AND INBREEDING The case of the wheat is typical. Its develop- ment furnishes an illustration of the method through which many specialized races of animals and plants under domestication have been devel- oped. Indeed, it might almost be said that the [224] FIXING GOOD TRAITS one rule that has actuated the developer of special races has been to apply the principle of inbreeding. When an individual appeared in a herd or flock that showed certain peculiarities that the owner thought desirable, the natural and obvious way of perpetuating these was to breed from that individual; and then persistently, for a time, to inbreed the progeny in order to accentuate the desired trait. The result has often been all that could be expected. Take, for instance, the case of the trotting-horse. It is, I believe, a matter of record that practically the entire stock of trotters, as developed in America in the past hundred years, descended from a single ancestor, the celebrated "Messenger." This individual horse chanced through some accidental mixture of ancestral strains to combine in its organization the particular qualities of nerve and muscle that adapted it for rapid progress by trotting instead of by the more natural method of running: And as regards this quality or combination of qualities, the horse proved amazingly prepotent. Its descendants soon constituted a race of trotters. Pedigrees were kept; the best individuals of the new race were selected as breeders; closely related animals were mated; and the character- [225] LUTHER BURBANK istics that make for speed at the trotting gait were in a few generations so fixed that a new race of horse was produced. The principle thus illustrated applies with equal force to the breeding of plants. Indeed, it is possible here to hold even more rigidly to the idea of inbreeding, inasmuch as the individual flowers may be self-fertilized. We have just seen this illustrated in the case of the wheat and allied cereals. There is no question whatever that any given characteristic of a plant, once it appears, can be accentuated and fixed, first in individuals, and finally indelibly in the heredity of the descendants of the plant by systematic inbreeding. But, unfortunately, there are complications in the case of most experiments that the originator of new plants is called upon to undertake, that robs the method of its simplicity. The com- plications arise from the fact that the would-be originator of new races of fruit or flowers is usually seeking to develop not merely a single quality, but a number of qualities. And this alters the case fundamentally. In the case of the trotting-horse, the one all- essential quality desired is speed. The capacity to trot a full mile at high speed does, indeed, imply the possession of stamina and [226] FIXING GOOD TRAITS courage, as well as capacity for rapid action of the legs. These are qualities that are necessarily linked with the capacity for the right kind of muscular action. But beyond this there are very few qualities upon which the breeder must insist. It does not greatly matter whether the speedy animal is small or large; its color is mostly a matter of entire indifference; and it is taken as a matter of course that the record-breaking animal will be nervous in temperament, tender as a hothouse plant, and requiring such care and attention as would be only wasted upon a more plebeian animal. In a word, the breeder of trotting-horses fixes attention principally on the single quality of speed. But it is rare indeed that the would-be developer of a new plant can thus fix attention upon any single quality to the disregard of other qualities. On the contrary, as a rule, the plant experimenter, while he may have in mind one most important quality, must consider at the same time six or eight or ten or a dozen other qualities that are only a degree less essential. We have seen this illustrated again and again, and we shall have occasion to recall some of the specific characters involved in the course of the present discussion. [227] LUTHER BURBANK In reality, the task of the experimenter who would develop a new and really valuable variety of plum or cherry or apple or spineless cactus, is to be compared not with the task of breeding trotting-horses as they are, but rather to the task that would confront the breeder were he to attempt to develop a race of trotting-horses which should retain the capacity to trot a mile in less than two minutes, yet at the same time should be big and powerful enough to serve on occasion as draught horses; should be always of some pre- determined color, say bright bay; and should be as hardy and require as little attention as the toughest broncho. It requires no great amount of imagination to see that the task of breeding race horses would be quite different from what it is, were the specifications such as these. Yet I repeat that the qualities that the plant experimenter usually seeks to combine in his new variety of flower or fruit are at least as varied and as difficult to fix in combination as the qualities just suggested for the supposititious new breed of race horses. THE SHASTA ON THE WITNESS STAND Let us by way of illustration recall the case of the Shasta daisy which, the reader will remember, was developed by the union of three different Rose Cuttings — Developed by Selective Breeding A new variety of rose — developed by selective breed- ing— may be propagated by merely cutting the branches into sections, and planting them in the way illustrated on page 135. It is oj course necessary to have at least one bud, and preferably two or three, on each cutting. Mr. Burbank has devel- oped some very remarkable new varieties of roses, including one that received a gold medal at the St. Louis International Exposition in 1904. LUTHER BURBANK species of flowers, coming respectively from Europe, America, and Japan. It will be further recalled, that the ideal daisy that I had in mind for years before it became an actuality, showed in superlative degree a consid- erable variety of qualities that were not found in combination in any one of its ancestors. Indeed, the Shasta daisy, as ultimately developed, reveals a number of very conspicuous and important qualities that are not shown at all in any one of its known progenitors. To make the illustration specific, we may cite, among the qualities that are assembled in the finished product, the following: (1) extreme size, (2) dazzling whiteness, (3) broad rays, (4) double rays, (5) gracefully drooping rays, (6) keeping quality of flower, (7) smooth stem, (8) early and persistent blooming, (9) hardiness, (10) constant bearing. The perfected Shasta daisy manifests these qualities in supreme degree. As regards each and every one of them, it surpasses any of the parental forms from which it sprang; indeed, as to some of them, such as double and drooping rays, it shows an entire departure from all of its observed ancestors, harking back to the remoter forms of past ages. But to assemble these qualities in a single [230] FIXING GOOD TRAITS flower required about fifteen years of persistent effort, and the handling of probably not less than half a million individual seedlings. Generation after generation the plants were cross-pollenized and selected over and over, always with an eye not merely to a single quality, but to the ensemble of qualities. And always we were confronted with the difficulty that in reaching out to bring in some new quality, we were disturbing the balance of qualities already attained, and endangering the entire structure. When, for example, the final cross was made with the Japanese daisy, to secure if possible the element of whiteness shown pre-eminently by that flower, and add it to our mosaic, we, of necessity, brought in also from the Japanese parent, along with the quality of whiteness, such undesired qualities as crude, ungraceful stems and flowers. It was necessary to select and interbreed, and select again, for successive generations from among a multitude of the progeny of this cross, before a plant was finally secured that presented the desirable combination of qualities, retaining the whiteness of the Japanese parent, but rejecting its undesired characteristics of leaf and stem, and departing utterly from that particular parent in point of size. [231] LUTHER BURBANK But, although an individual was at last found that did combine all the desired qualities, the very fact that this individual had been built up by putting together this quality brought from one parent and that quality from another, with the rejection of antagonistic qualities in each case, makes it inevitable that the perfected Shasta should contain latent in its system a whole coterie of tendencies which are fighting for recog- nition, and which will make themselves felt in subsequent generations. Hence it is that when seeds are gathered from the perfected Shasta they will not give us a crop of flowers like their parent. On the contrary, they will show the utmost diversity of form and size and color, making tangible thus the persistent force of the hereditary tendencies that had been transmitted from divers ancestors, but which were submerged or made latent, simply because they were momentarily subordinated to opposing qualities, in the case of the perfect Shasta. The Shasta daisy, then, while individually almost a perfect embodiment of the ideal at which I aimed, is when reproduced, from seed, anything but a fixed type. Had it not been possible to propagate the plant by division and then by an unending series of successive divisions to produce an indefinite number of individuals, each precisely [232] FIXING GOOD TRAITS like the original because they were in a sense a part of it, my entire series of experiments in developing the new daisy would have been unavailing except for still further selection. But as the case stands, it was possible rapidly to develop an entire race of Shasta daisies by root-division, and thanks to this method the descendants, or, to speak somewhat more accu- rately, the sisters of the original Shasta daisy have become an enormously populous race, scattered to the remotest parts of the earth. Several other types of Shastas have been developed by new breeding experiments from the original stock, but to this day the race of Shasta daisies must be propagated from the root, and not grown from seed, unless one desires a conglom- erate progeny, departing in many ways from the form and quality of the immediate ancestor. FIXING A TYPE In all this, it must be recalled, the Shasta daisy does not differ from a large number of long- established cultivated plants that are everywhere recognized as being "fixed" races. One does not produce apples or pears or cherries or plums or blackberries or potatoes or sugar cane or horse radish, to say nothing of roses, ornamental shrubs and a great number of flower- ing plants, from seed. [233] Wild Grapes There are a good many species of wild grapes, and Mr. Burbank has utilized several of these in his experiments. His chiej work with this species has been done, however, with the common European cultivated grape, which has developed a large number oj varieties through being selected Jor various purposes during past centuries. In particular some oj his most important new varieties are descended Jrom a "bud sport," — that is, a variation that appeared "spontaneously" in a branch growing on an ordinary grape vine. FIXING GOOD TRAITS They are propagated by grafting or budding, or by rooting the stem or dividing the roots or planting the tuber. And the reason in each case is the same. The perfected variety originated from a single individual that combined a large number of desirable qualities, and the entire company of individual representatives of that variety, though they be numbered in millions, are not really descendants, but offshoots, of the original individual. Each cion or bud from a given tree will produce fruit precisely like that from the tree from which it is taken, because it is itself a part of the tree. And however widely new cions and buds from the first cion may be disseminated, they carry the same traits, because, rightly con- sidered, they are a part of the same individual organism. The Seckel pear tree that grows in your dooryard is, from the standpoint of heredity, a tree of the same generation with untold thou- sands of other Seckel pear trees that have grown here and there across the hemispheres for more than a hundred years — or since the first one appeared in the orchard of the Pennsylvanian whose name they bear. Were it not for the contradiction of terms, one might say that all Seckel pear trees constitute a single tree. [235] LUTHER BURBANK All these Seckel pear trees are essentially alike; they bear fruit that may vary in size and lusciousness with varying conditions, but that is everywhere essentially identical in flavor and in the characteristic qualities of texture and color. But if you plant the seeds of one of these pears you do not secure Seckel pears, unless by the merest chance, among the progeny. You secure instead, representatives of a galaxy of ancestors, no one of them individually just like the Seckel, although collectively they represent all the qualities of that fruit, plus almost numberless undesirable qualities. PAIRS OF QUALITIES The fact that our most familiar and best prized fruits and flowers show this lack of fixity, suggests that the inherent difficulties in the way of fixing the type of these plants so that they will breed true from seed are very great. Otherwise some one would long ago have remedied the defect, for the advantages of being able to grow these useful plants from seed are obvious. Nevertheless it should not be assumed that the task of fixing the type of a newly developed race of fruit or flowers is of necessity a hopeless one. The truth is that it would be possible to fix the type of almost any variety of plant, provided time enough and patience enough were devoted to the [236] FIXING GOOD TRAITS task, and the experiment were conducted on a wide enough scale. Indeed, nothing more would be necessary than to continue for an additional number of generations the same line of experi- mentation through which the new varieties were produced; attending carefully at all stages to the analysis of the different qualities that prove to be mutually antagonistic. To this end, the new terminology which endeavors to analyze the qualities of a given plant into complementary pairs of unit characters may prove very helpful, particularly to the inex- perienced investigator. Such an analysis has always been made, tacitly at any rate, by the successful plant experimenter. No one can think of the development of an early- fruiting cherry or prune without having in mind the quality of /afe-fruiting. To speak of a prune with high sugar content implies one with low sugar content. In a word, the desired quality of fruit or flower at which one aims is always balanced against the opposing quality — sweet fruit against sour, hardiness against tenderness, resistance to disease against susceptibility to disease, profuse bearing against scant bearing, thorny brier against smooth brier, black fruit against white fruit, and so on down the list. [237] LUTHER BURBANK It is only by constantly bearing these divergent pairs of qualities in mind that any experimenter can hope to advance toward the production of an ideal fruit or flower or vine. And it has always been so. OLD WINE IN NEW BOTTLES But there can be no question that the new terminology, as used by present day biologists, serves to give precision to the ideas of the plant experimenter, and enables him to analyze the results of his experiments in more precise terms than have hitherto been available. It will be convenient, therefore, and probably helpful to the reader, in making precise reference to some of the experiments in plant breeding already detailed, with special reference to the possibility of fixing the type of new races, if we discuss the matter in the new terminology. It will at once appear that when a plant developer attempts to fix a certain type, he is fundamentally changing his point of view. Hith- erto he has been concerned to make plants vary, in order that he might seize on new forms, and use them as material for developing the type at which he aims. And his success in developing a new race is largely dependent upon the extent to which he has been able to induce the plants with which he experiments to vary. [238] Seedless Grapes This shows one oj Mr. Burbank's choicest varieties oj new seedless grapes. They have been developed by selection, and are oj delicious quality, with an added attractiveness due to the entire absence oj seeds. Of course a fruit thus developed must be propagated by cuttings, and is not susceptible of Jurther improvement, unless there should be a reversion to the seed-bearing condition. LUTHER BURBANK So now, when he attempts to restore fixity to something that he has purposely made unstable, he is at once confronted with the danger of undoing much that he has accomplished. The measure of success that he can hope to attain will depend very largely upon the particular kind of unit characters that he has combined in the product that he now wishes to make stable. We have seen that, as between the opposing members of any pair of unit characters, it is usually discovered that one has prepotency or dominancy over the other. When blackberries of normal color, for example, are crossed with the white blackberry, the progeny are all black, because this color is the dominant member, and white the recessive or negative member of the pair of unit characters. But we saw also that the recessive trait will reappear in the succeeding generation, and that when it does reappear, it will, within certain limits, thereafter breed true. So, when in the second generation we again produce a white blackberry, we have a type which is fixed as regards the particular character of whiteness. In other words, our white blackberry, even though both its parents, and one grandparent, were black, may be considered a berry of pure white strain. From the moment of its appearance it is a fixed type as to color. [240] FIXING GOOD TRAITS But, unfortunately, it is not sufficient that the white blackberry should breed true as regards the quality of whiteness alone. There are other quali- ties of size and flavor that are equally essential. And these, it would appear, include sundry other pairs of unit characters — sweetness versus sour- ness, large size versus small size, profuse bearing versus scant bearing, and the like — that are repre- sented in our berry by mixed factors. In the Mendelian view, it will be recalled, there are always two factors representing any pair of unit characters. In the case of our white blackberry, in the Mendelian view, both factors for the unit char- acter blackness-versus-whiteness are of the white order; or in the technical phrase, the berry is "homozygous" for that pair of factors. But as regards, let us say, the factor for the unit character bigness-versus-smallness the case is different; for this character may chance to be represented by one factor of each type. In other words, resorting again to the technical language, the berry may be "heterozygous" as regards that character. In this particular generation, the quality of bigness prevails, because bigness is dominant to smallness. But the factor for smallness must have a hearing in the next generation. [241] LUTHER BURBANK Until we can produce a white blackberry that is "homozygous" for size-factors as well as for color-factors, we shall not obtain a fruit that will breed true to size as well as color. A similar analysis might be applied to the various other pairs of unit characters that are represented in any given fruit or flower. And the essential principle, stated in Mendelian terms, to be aimed at by the experimenter who would fix a newly developed type of plant so that it will breed true from seed, must be to render the plant "homozygous" for the factors of each pair of unit characters involved. If that can be done, the plant will breed true; if that cannot be done, the plant will not breed true. In the olden phrasing, this would be spoken of as "line" breeding — a method long familiar to every breeder of plants or animals. FIXING A TYPE IN THE SECOND GENERATION In actual practice, where only two or three unit characters are involved, it may be possible to produce a new type that breeds true, or is fixed, in the second generation. In such a case the time element may be ignored. , Take, by way of illustration, Professor Castle's guinea pigs, to which reference has more than once been made. Suppose we have as parent stock a black guinea pig with a smooth coat, and [242] S|! P *I:»«.KI S'S-l-s °° s ° c s1?-: ,-|§ Is *. . w « „ . " ^*> ^r. a sg ^-^ Ultir J^iS.'M S -< ft) LUTHER BURBANK a white guinea pig with a rough coat. Now we have already seen that blackness is dominant to whiteness as regards the coat of the guinea pig, and we must further understand that roughness of coat is known to be dominant to smoothness. We must expect, then (according to Professor Castle), that when a cross is made, the guinea pigs of the first filial generation will, unlike either parent, be black in color and rough as to coat. But, in the succeeding generation, the black, rough-coated guinea pigs being interbred, there will be a certain number of offspring that combine the dominant characters of blackness and rough- ness of coat, and will breed true to these; a certain number will be black and rough-coated, but will bear the latent characters of smoothness and whiteness of coat which will reappear in their progeny; and, finally, there will appear individ- uals combining the two recessive traits of white- ness and smoothness of coat. These white, smooth-coated individuals are obviously different from their parents, and different also from either of their grandparents. They constitute a new race, sprung into being in a single generation, and a race that will necessarily breed true as to the character of smooth coat and white coat, because they are "homozygous" as to the factors for both these recessive characters. [244] FIXING GOOD TRAITS Their progeny cannot be black because their germ-plasm contains no hereditary factor for blackness; nor can their progeny be rough-coated, because their germ-plasm contains no hereditary factors for rough-coatedness. Yet side by side with this new fixed race of smooth-coated white guinea pigs, there are, as we have seen, twins of the same fraternity that instead of being white and smooth in color, are black and rough. And these also constitute a new race that will breed true because they contain, as regards the unit character for color and for condition of hair, only the dominant factors of blackness and roughness. They also are "homozygotes," but they are of the opposite type — dominants instead of recessives. Meantime, we must not overlook the other members of the fraternity, twin brethren of these new races, which are individually black and rough, but which are "heterozygotes" as regards the unit characters under consideration, and hence will show progeny of variously mixed character- istics as to roughness or smoothness of coat, and as to black or white color. This illustration, perhaps, gives as tangible an impression as can well be gained, of the com- plexities that confront the experimenter when he attempts to fix a new type of animal or plant. [245] LUTHER BURBANK Even where only two unit characters are involved, the progeny of the second generation, as we have just seen, may break up into numerous races, some fixed and others variable. And, as we have previously pointed out, the complications thus introduced increase at a startling ratio when more characters are under consideration. Moreover, the matter is rendered increasingly difficult for the plant experimenter by the fact that he must often wait, particularly in the case of orchard fruits, for a term of years before he can know the result of any single breeding experiment. To sort out the pure types from the mixed ones of any given generation under these circumstances becomes a matter of enormous complexity. It could be done, by inbreeding representatives of the new type and carefully selecting the progeny for a series of generations. But in the end, all that would have been accom- plished, in the case say of a Shasta daisy or of a stoneless plum or a sugar prune, would be the production of seed that could be used to dissemi- nate the new variety. And in most cases we are justified in feeling that this would represent an undue expenditure of time and energy for a com- paratively insignificant result. For, as the case stands, even though the new form will not breed true from seed, it may be [246] FIXING GOOD TRAITS propagated indefinitely from roots or from the grafting of cions; so that in practice the failure to breed true from seed has little significance. Probably it is the fact of the relative unimpor- tance that our cultivated plants should breed true from seed that chiefly explains the failure of plant breeders in the past to fix the type of the best known fruits and vegetables and flowers. The same reasoning obviously applies to the newly developed varieties. While so much work remains to be done in the way of developing new types of fruit and flower, the most practiced experimenters will probably feel that they have not time and energy to spare for the fixing of the new races already developed. We shall have occasion to call attention to various exceptions to this rule in the course of our subsequent studies; particularly with refer- ence to certain annual plants. Here by "line" breeding for a few generations we may fix the new traits almost as firmly as the old traits are fixed in wild species. Again we shall learn in due course of new hybrid fruits like the sunberry, the primus berry, and the phenomenal berry that are fixed as to their chief properties from the very first hybrid generation. But as regards most of the new forms of fruit and flower that we have hitherto described, the rule holds with full force. [247] Label on Tree Graft This illustrates Mr. Burbank's method oj labeling a graft, in order to keep track of his experiments. The label is of wood, and the inscription is made with paint or pencil. The wire attaching the label should of course be loosely applied to avoid restricting the plant. RECORDING THE EXPERIMENTS EASY WAYS TO KEEP TRACK OF PROGRESS EVERY ONE has heard the story of the distinguished professor who devoted his entire life to the study of a particular species of mite, and who, on his deathbed, regretted that he had not confined his attention to the study of the respiratory organs of this insect, instead of trying to comprehend its entire structure. This specialist, like many another, felt that he had wasted his energy by attempting to cover too wide a field. He felt that his ten volumes or so on the anatomy of the mite could give but super- ficial treatment of a great subject. Whoever sympathizes with the attitude of mind revealed by this doubtless apocryphal yet truly symbolic tale, will have scant patience with my method of plant experimentation. For, far from confining attention to a single species, or even to [VOLUME III— CHAPTER VIII] LUTHER BURBANK a single genus or order, I have extended my observations to almost every form of plant, testing species by thousands, and individual specimens by hundreds of thousands or millions, in my experimental gardens; and only by exception has complete record been kept of all details of any given series of experiments, beyond the more or less fallacious records of memory. Yet I have had the good fortune to produce, I suppose, more forms of plant life that could justi- fiably be called new, than have been produced by any other single experimenter in our time. Had I stopped to make meticulous record of each experiment, I doubt if I should now know more than I do about even my less important products, and I surely should have been able to produce only a fraction of those that I have produced. METHODS AND RESULTS Yet it must not be supposed that I have alto- gether refrained from graphic recording of the progress of my tests. The fact is quite otherwise. I have kept in the aggregate a vast body of records, and have had them always at hand, under my eye from day to day, telling of the essentials of my hybridizing and other experiments. My "plan books" have been a constant aid to memory, and guide to further effort. My record books have set down in black and [250] 4. !««• t - "sg^ " s- 7 3 Kf. 1- s° ? LUTHER BURBANK white the unequivocal evidence of progress — or of failure to progress. Few salient facts as to the precise parentage of important hybrids and the exact methods by which variation has been brought about have failed to find explicit record, notwithstanding the omission of multitudes of details that to some observers might have seemed worthy of transcription. And if I have adopted in the field shortcut methods of recording selection, these have not lacked precision and accuracy, notwithstanding their time-saving character. In point of fact, all along the line I have endeavored to strike a happy medium between the waste of time that would result from the keeping of unduly elaborate records, and the waste of effort that would necessarily result if no records at all were kept. The reader who would clearly comprehend the nature of the compromise must bear in mind that I have, as a rule, had a practical object in view in conducting my experiments. It is true, as Professor Bailey has courteously said, that I constantly make experiments with plants for the mere love of the work. It is true also that my tests include hundreds of species from which I expect no very definite return. Yet it is further true that the main body of my experiments have [252] RECORDING THE EXPERIMENTS been concerned with flowers or fruits that seem to offer opportunities for practical improvement. I have usually been seeking, in the experiments to which most time has been given, to modify the plant in such a way as to make it a more beautiful and desirable garden ornament, or to modify a vegetable or fruit in such a way as to make it a more valuable food product. Such being the case, it will be understood that, with regard to large series of experiments, I have been concerned with results rather than with methods. As to the latter, it often happens that numberless experiments might be described in substantially the same terms. Once the prin- ciples of hybridization and selection have been clearly mastered, they may be applied to almost every variety of plant life. There are differences in the detail, but the broad outline is the same for each. ESSENTIALS VERSUS NON-ESSENTIALS It would then be but a waste of time to record over and over details as to these broader outlines of plant experimentation. Where anything of interest has appeared, any point as to which a plant shows differences from its fellows, this has become a matter for recording. Moreover, it has been my universal custom to make record of the first hybridizing or crossing [253] N^^^_".^. • V ^fe— " ^ Diagram o/ Tree Grq/is TAis is a leajjrom Mr. Burbank's plan book. It is a sort o/ shorthand or diagramatical record that condenses a large amount o/ information into small space, leaving room /or fiona/ records should they be called /or. Suc/r a page seems cryptic to the casual observert but to Mr. Burbank himself it is full oj meaning. RECORDING THE EXPERIMENTS through which any particular series of experi- ments is inaugurated. The parentage of the Shasta daisy, the white blackberry, the stoneless plum, the sugar prune, the plumcot, the thornless blackberry, the spineless cactus — these are matters of clearest and most unequivocal record. The results of the first crossing, through which matters of prepotency and of latency are determined, and through which the plant is given the impulse to variation, are also explicitly shown. But when, particularly in case of a fruit having complex characters, the experiment passes to stages of the third and fourth generations, involving tens of thousands or hundreds of thou- sands of seedlings, it is no longer possible to make detailed and explicit record, with exact count of the different combinations and variations devel- oped, for two very explicit and sufficient reasons. One reason is that the numbers of seedlings involved are so great that it would be physically impossible for any one carrying on hundreds of different series of tests at the same time to make numerical count in accordance with the statistical method adopted by workers who are experi- menting on a limited scale. The second reason is that even if such a count, showing the exact number or percentage of seed- lings with different combinations of traits, were [255] LUTHER BURBANK attempted, it would be unavailing unless vast companies of seedlings were preserved for the term of years necessary to bring them to fruitage. When one is concerned solely with numbers, or with such tangible qualities as color of hair in the case of Professor Castle's guinea pigs, or color of feather with Professor Davenport's fowls, it is an easy matter to check results, because the creatures under investigation manifest the quali- ties that are being tested from the moment of birth, or develop them at a very early age. But with plants the case is obviously different. Whereas we may judge something as to the character of fruit that a seedling will ultimately bear from observation of the seedling itself, yet for purposes of scientific record such predictions would be considered as worse than worthless. To know what percentage of seedlings of a given generation have really progressed toward the ideal of a sugar prune that will ripen August 1st instead of September 1st, let us say, it would be necessary to let all the seedlings grow for several years, or at the very least to wait two or three years for the grafted cions from each seedling to come to fruitage. The practical experimenter, seeking results, cannot possibly work in this way when he works on a large scale. [256] Record o/ Blooming Facsimile of another page from Mr. Burbank's record books. This one has to do with the time o/ blooming of various Jruit bearers, — a matter of great importance, because blossom- time is correlative with the time oj Jruiting, which often has important bearing on the value of a new fruit. It is a little surprising to learn that varieties that bloom early ripen their fruit relatively late, and vice versa. LUTHER BURBANK He must be content to select from among thousands of seedlings the one or five or ten or fifty that appear to him most promising. To these he must pin his faith, and all the rest must be destroyed to make room for other plants. Otherwise he would require not twenty odd acres, which make up the total area of my experi- mental farm, but hundreds or even thousands of acres. And to keep track of the multitudinous seedlings would require the aid not of the half dozen or so assistants whose cooperation makes my experiments possible, but of a small army of equally industrious workers. SYSTEMATIC WORK IMPERATIVE But, having thus outlined the limitations that necessarily attend work conducted on a large and comprehensive scale, let me now proceed to elab- orate somewhat the other side of the story. Let me outline the various practical methods of recording experiments that have been devel- oped in the course of my years of experience. Let me in particular point out some of the short- cuts that have made it possible for me to record the essentials, and even in important cases the details, of progress, with a minimum expenditure of time and labor. Among the essentials that cannot be overlooked [258] RECORDING THE EXPERIMENTS by any systematic and successful experimenter are the following: A general plan of the ground occupied by all the experiments must be made, and there must be clear record of each plant, shrub, or tree planted. It is important also to record the time when each one was grafted or budded; the date of all experi- ments in crossing any particular tree or plant; observations as to any anomalies of development; and finally, as a matter of course, the results obtained. As to these things, the memory, no matter how tenacious, must prove more or less untrustworthy. It is only the black and white record that can be depended upon. But plans may be outlined so simply that all these essentials may be recorded at the expense of very little time or labor. It is not much trouble, for example, to keep a plan book at hand, each page of which is devoted to a certain planting, its location on the grounds, and all other matters that are worthy of record. It will facilitate matters in such a book to have the records of planting arranged somewhat in the order in which the plantings have occurred during the season. If these records are made on large sheets of paper so plotted as to show the location of the various beds of plants, this will be an added [259] Grafting Record Yet another page of Mr. Burbank's record books, this one dealing with grafting. It will be seen that the records oj different years are associated, and it will be obvious that the records were intended for Mr. Burbank's own eye, to supplement the records oj memory. When thousands of experiments are being performed simultaneously, it is im- perative that the recording should be done with the least possible loss of time. RECORDING THE EXPERIMENTS convenience, as it will enable any particular lot of plants to be located, even if through some inad- vertence the label stakes, which are an absolute necessity, have been removed or lost by careless workmen. Often when planting in the field, letters or numbers are used on the stakes, corresponding with similar letters or numbers in the record book. LABEL STAKES AND LABELS As to the label stakes themselves, the ones that I habitually use for general field culture are about 20 inches long, 2 inches wide, and % of an inch in thickness. They are smoothly planed and painted about half way down on both sides with common white lead paint. One coat of paint is far better than two, for if a pencil is used the lightly painted surface takes the lead to advantage, and by bearing down heavily with the pencil, indentations are made in the wood that will resist the weather more effec- tually and thus give greater permanence to the record. It is desirable to make the label stakes of soft, smooth redwood or other durable wood. In the East the locust is an excellent substitute. It is advantageous to dip the end of the stake in car- bolic acid or in a solution of sulphate of copper to prevent decay. These stakes may be used over [261] LUTHER BURBANK and over again for many years, being planed off as the occasion requires and repainted. Many thousands of these label stakes are used each season on my experimental grounds. For smaller beds I use a stake usually 1 inch wide, from % to % of an inch thick, and from 10 to 14 inches in length. These smaller plant stakes may be purchased of dealers, and are prepared for use in the same way as the larger ones. For use on trees a special label is employed, to make records of budding, grafting, and varia- tion. This label is usually 5 or 6 inches long, and from 3-16 to % of an inch thick. It is notched at one end and attached to the branch of a tree with a piece of pliable galvanized iron wire. The wire should be loose enough to avoid any danger of strangling the branch. The labels are painted with white lead. They sometimes remain upon the trees for five, ten or even fifteen years. To inscribe these permanent labels, I use a thick black paint, composed of a mixture of lamp black, linseed oil, and a little turpentine, applying the mixture with a small camel's hair brush. The names of varieties, the parentage, and other important matters are thus recorded. Then, while the paint is still wet, some fine dry sand is sifted over the label so as to protect the paint from the weather. [262] Ripening Record This may 6e regarded as a companion record to the one shown on page 257. 7f records the highly important matter of the time of ripening oj various fruits. This particular page shows among other things that a seedling Japan raspberry ripened April 2$th, and a new hybrid blackberry May ist. Some of Mr. Burbank's finest new fruits, including the Burbank cherry and the Sugar prune are made doubly valuable by the fact that they ripen several weeks earlier than similar fruits of other varieties. Mr. Burbank is constantly encouraging fruits, by selective breeding, to modify or lengthen their season of fruiting. LUTHER BURBANK In addition to the labels and stakes I have just described, a small cardboard label of light weight is needed for making record of the hybridizing experiments. The common cardboard shipping tag about 1% inches by 3 inches in size with a reinforced eyelet hole, is generally used on plants with tender stems; and where the wind is likely to disturb larger labels, half or two-thirds of the cardboard may be cut off, leaving barely space to inscribe the record. Where these tags are used in extensive pollena- tions of many varieties on a single tree, it is not always necessary to write the record, for the same object may be accomplished by cutting off one corner of the card to indicate a certain variety of pollen, and a second corner to indicate another variety; additional varieties being represented by series of notches. Or the same end may be attained by punching holes in the card with a pocket steel punch. This plan saves much time, and the record is more permanent than if it were made with pencil. A large number of tags may be prepared at once with punch or scissors. Tags of this character are less likely to have their records erased by wasps and hornets, which often partially destroy labels when securing mate- rial for their paper homes. Conspicuous tags such as these are $lso of aid [264] RECORDING THE EXPERIMENTS later in the season when the seed is to be gathered; because they make known at a glance the facts as to parentage, and make it possible to keep sepa- rate the seeds of different varieties. The labels are tied to the plants with common twine, as wire or other hard substances would be likely to injure the tender stems when the wind moves the tags about. When numerous varieties of plants are grown in a single bed, we often nail a common tree label opposite each row on the board that borders the bed, instead of using a stake, as there is less danger of the label being displaced. It will be advan- tageous to place this label at the side of the bed away from that from which the prevailing winds and storms come. In this section of California the summer winds and winter storms come from the South, East, and Southwest, and in conjunction with the hot sunshine, they are very destructive to paint. So it is advantageous to face the labels toward the North. All of these are matters of minor detail, yet not without their importance. RECORDING RESULTS In making selection of individual plants that are to be preserved, or from which seed is to be gathered, the most convenient, and at the same time the most accurate method is the simple one [265] Budding Records It has sometimes been supposed that Mr. Burbank depends entirely on his memory ; and makes very Jew written records oj his experiments. This facsimile page, taken in connection with the others that have just been shown, will serve to dispel that erroneous notion. It will be seen that records are kept of the essentials of each different type of experiment, including not only methods, but results. In point oj Jact, although Mr. Burbank adopts many shortcuts, his records oj essentials are surpris- ingly comprehensive and complete, considering the enormous range oj his experiments. RECORDING THE EXPERIMENTS of tying a small strip of cloth about the stem of the plant. Visitors to my gardens are sure to notice that each bed of flowers has a half dozen or so plants that are thus decorated. In some cases two or three strings may be attached to a single plant, indicating degrees of excellence. Selection having been made in this way, the plants may be allowed to ripen their seeds, and in due course the work- men may gather them without further direction, placing them in labeled boxes to be stored for the winter. As regards new fruits, there is particular need of great accuracy, and here it is impossible to avoid a good deal of detail. It will not do at all, in dealing with a valuable addition to the list of fruits, to leave anything to memory as to its season of ripening, size and form, color, flavor, aroma, size of core or stone, length of stem, or any other essential quality. An exact record must be kept of these items, and for this purpose a book with removable unruled leaves is the most satisfactory. The fruit should be cut in half with a sharp knife. The incised surface may then be placed directly on the paper, and the outline of the fruit traced with pencil. The specimen may similarly be outlined in cross-section. This preserves a [267] "S»«~i RECORDING THE EXPERIMENTS graphic record of the exact size and form of the fruit. The main character of the inside of each fruit may be indicated, and by adding the date of ripening, the time of its earliest and medium ripening, the number of days it will remain in good condition upon the tree, its keeping quality when packed for shipment, and its susceptibility to the ravages of insect pests and fungoid disease, we have on a single sheet a fairly complete and very valuable record, together with a graphic rep- resentative of the size and form of the fruit itself. Record will be made in the same way in suc- cessive seasons of fruit from the same tree, with additional record of the appearance of any new characters or qualities. Comparison of the rec- ords will show whether the fruit on the young trees has increased in size, improved in quality, or varied in time of ripening from year to year. Not unfrequently the record of the third year shows a very considerable increase of good qualities over the first. After a record has been kept for four or five seasons, a fair estimate may be made of the gen- eral value of this particular fruit. If in addition we know the characteristics of the parent forms — whether the ancestors were hardy or tender, and the like — we are now in position to form a clear judgment as to the probable value of the fruit. [269] Punched Labels These labels are important time-savers. Each set of holes represents a specific plant or a different type oj experi- ment, the key to the labels being found in the record books. The upper left hand label here shown might indicate, for example, that the branch on which it is tied bears blossoms of the Sugar prune (one hole at base of label), fertilized by pollen of the Conquest stoneless prune (three holes at center of label). Such labels as this are loosely attached to the stem of the plant that has been pollenized or grafted. They are per- manent, and they minimize mistakes. RECORDING THE EXPERIMENTS Such a record as this is essential to actual progress. It is important, if for no other reason, to prevent the experimenter from deceiving him- self. It is very easy to imagine that a certain product that has caused one much trouble is better than some other; or that a fruit of a given tree is larger than some rival variety. But the record book enables one to put the matter to a precise and definite test; it makes self-deception impos- sible; and it affords an invaluable guide to further experimentation. There are thousands of graphic records such as these on the shelves of my library. I would not think of attempting to conduct an intricate series of experiments looking to the development of a new fruit without the aid of these plan books. When the experiment is finally completed, a series of these loose leaves, properly collated, furnishes a complete record of the various hybrid- izings and selections — resulting sometimes in better and sometimes in worse fruits — through which success has finally been achieved. These records are in themselves sufficient answer to any one who imagines that the plant experimenter works haphazard, merely because he does not always adopt a biometric method. After all, from the standpoint of the consumer [271] Notched Labels This is another type oj short-hand label similar in its use and interpretation to the ones shown on page 270. Each series oj notches tells a different story, and the key to their inter- pretation is found in the record books. Two notchest /or example, might represent the Alaska daisy and three notches the variety named Westralia. The top label above would then record the hybridizing oj these two varieties. (The small notches for the string are not counted.) Mr. Burbank finds this method oj keeping track oj experi- ments highly satisfactory. RECORDING THE EXPERIMENTS who makes up the main bulk of the population, and whose tastes and needs are the criterion by which the plant experimenter's results will be judged, it is the final product rather than the precise method by which it is attained that is important. But the ideal at which the plant experimenter aims would probably never have been realized had he not given himself the aid of some such system of quick and accurate records as my plan books present. —Once the principles of hybridiza- tion and selection have been clearly mastered, they may be applied to almost every variety of plant life. There are differences in detail, but the broad outline is the same for each. Variation in Corn SeeJ Af r. Burfcanfe Jinds material for most of his experi- ments in variations as to one quality or another that appear among plants oj the same species. It may or may not be necessary to accentuate variation by hybridizing experiments. The range oj ^ariation that may be shown in the seed oj a single species is illustrated in this lot of kernels oj corn, which show surprising diversity in shape, size, and color. Numberless new varieties could be devel- oped through selective breeding from such a lot oj seed as this. FINAL SELECTION THE MOST IMPORTANT TASK OF ALL IN FARMING districts of the Mississippi Valley they have a curious custom in selling cattle at auction. They drive a herd of cows together and the auctioneer asks his audience to bid for first choice, no individual animal being specified. The highest bidder makes his choice, and the cow he selects is taken from the herd. Then the auctioneer starts over, receiving bids for "first choice" among the remaining animals. This process is repeated again and again until all the exceptional animals have been selected. A curious result of the method is that it very commonly happens that different bidders have their eyes on different animals. Farmer A, who bid highest at the outset, did not have in mind the animal for which farmer B was bidding. And so it often happens that after six or eight selections have been made a cow still remains that was [VOLUME III — CHAPTER IX] LUTHER BURBANK regarded by some of the bidders as the very best one of the entire herd. A man who bid unsuccessfully again and again may thus, in some cases, finally have his choice precisely as if he had made the highest bid at the outset. The obvious explanation both of the method and of its somewhat anomalous results is found in the fact that individuals differ in their judgment as to what constitute the superior qualities of a cow. Each bidder has noted an animal that particularly appeals to him, and each is backing his own judgment in making selection. The result is a process of elimination that may or may not select from the herd the best animals at the very outset. "But what have cows and their selection to do with the development of new varieties of plants?" you ask. Nothing direct and obvious to be sure. But it has often occurred to me that the process of selec- tion at the Iowa auctions is closely comparable to that which is employed by the plant experimenter in the course of his every-day work. In lieu of a herd of cattle, he deals with a group of seedlings. But his task is precisely like that of the auction bidder in that he must select from among scores of plants of the same kind, and often of closely [276] Chilian Beans This lot oj beans affords another striking instance of the wide range of variation among the seeds of individual plants of the same species. It may be assumed 'that these widely diver- gent seeds represent various heredities. Each plant is, in point of fact, a mosaic of characteristics inherited from many lines of ancestry. In Mr. Burbank's phrase, heredity is the sum of past environments. A curious and important feature of the matter is that the different racial strains may be segregated, as illustrated in this lot of beans. LUTHER BURBANK similar appearance, the one that seems to him the choice of the entire lot; and then in succession the second and third and fourth best, until he has chosen possibly six or eight individuals out of a group of hundreds or thousands. These six or eight individuals will be preserved for use in further experiments. They are the ones with which the attempt to improve the variety to which they belong will be carried out. And the ultimate success of the entire experi- ment in plant breeding will very largely be deter- mined by the perspicacity with which the selection of these few individuals was made. Nor can we doubt that it must often happen, in the case of the seedlings as in that of the cattle, that after the final selection has been made there remain, unknown to the experimenter and in contravention of his judgment, better plants among those rejected than any one that he has chosen. It could not be otherwise when we consider the large numbers involved, the variety of plant characteristics, and the great diversity of traits represented in a single generation of hybridized seedlings. Yet, on the other hand, experience should enable the experimenter to choose with a relative degree of certainty, and it is possible to acquire a degree of skill, based on careful obser- vation of the minute details of plant structure, [278] FINAL SELECTION that will give full assurance of a capacity to select with at least a large measure of success. A HALF HOUR IN THE ORCHARD It is usually a surprise to any visitor who comes to my orchard at a time when I am making selec- tions among seedlings of many kinds to observe my method. Many people have expressed astonishment when they have seen me wralk rapidly along a row of plum trees saying: "Kill this one, and that one, and that; save this one, and that one yonder"; indicating the choice between plants to be saved and those to be destroyed so rapidly that the men following me can scarcely tie strings to the selected ones as fast as they are chosen. In this way I may test from five to ten thousand young trees as I walk along the row, scarcely pausing for more than what seems the most casual glance. But my eye takes in the important thing. I know just what I am looking for. And if my judgment in the matter had not proved in the main good, the output from my orchard would have been quite different from what it has been. I may recall by way of illustration an experi- ence in which my selective judgment was put to a practical test — no different a test, to be sure, from thousands that I myself have made, but having added interest because it was made by another. [279] LUTHER BURBANK It chanced that a well known judge, who is also a horticultural enthusiast, who had been very much interested in my work, was visiting me at a time when I was sorting out plum trees from among a lot of several thousand seedlings about a foot high. I had a man carrying them away as fast as selected. They were thrown in three piles, the first containing those I had declared to be the best ones for continuing the test; the second pile containing those I thought possibly worth trying; and the third pile those that seemed to me no good at all. The judge watched me for a few minutes and then said: "You are picking them altogether too fast. You cannot possibly tell like that which are good and which are not." I replied : "Wait and see, or test the matter for yourself if you wish." "Very well," said my visitor, "I will do so." And therewith he selected a few seedlings from each of the piles and took them home with him to graft on trees of his own. Of course it was necessary to wait two or three years for results. But when the time came, the judge very cheerfully admitted that I had been quite right all along the line. The cions from my discarded pile bore fruit that was almost worthless; those from the intermediate pile gave [280] 1-gif*! LUTHER BURBANK fairly good fruit; and from the pile of my first choice seedlings he secured a fruit of such quality that he named it the Klondike, declaring that it gave him more good plums than he had ever had before from a similar tree. I cite the incident as showing the possibility of gauging fruiting qualities of a seedling at a time when the plant itself is a mere sapling a few inches in height. The capacity to make such selection has sometimes been spoken of as intuition; but it is really a matter of observation and practice. One learns through long experience to judge what characteristics of the seedlings are suggestive of possibilities of fruit-bearing. And after all this is no more than judging the man of the future by observation of the child of to-day. THE CORRELATION OF PARTS If we were to state the matter a little more technically we might say that such selective judg- ment as I have just illustrated is based on a knowledge of the correlation between the different parts or members of a plant's organization. It was first prominently brought out, I believe, by the French naturalist Cuvier something over a hundred years ago that there is always a correlation between the different structures of a given animal, to accord with its habits of life. [282] FINAL SELECTION For instance the teeth and claws of a cat are associated with its carnivorous habits and are linked with a certain structure of legs and muscles adapting the creature to spring forward with great celerity upon its prey. A somewhat different structure of body and limb is associated with the talonless feet of the dog tribe which are adapted to rapid running for prolonged periods rather than to sudden leaping and clutching. It was by careful study of the correlation of parts, of which these are only crude and familiar examples, that Cuvier was enabled to gain an insight into the characteristics of fossil animals of which only small fragments of skeletons were preserved in the rocks. The science of comparative anatomy was the outgrowth of his observations. Now it is at once obvious to anyone who studies plants attentively that their structure also shows a corresponding and no less invariable correlation of parts. The more conspicuous illustrations of this are obvious to the most casual observer — various adaptations of form of tree and shrub and vine to their natural surroundings are so patent that they cannot escape attention. But of course the plant experimenter must deal with correlations of a very delicate order. He [283] too C too fj _. ^J K v. g .1 «1?2 ^ ?-* ^^vS 3 ton^ ^5 tor.^3 ^ 0:1 111 ~« «* ^ 3 §•* 5 : ^flail •s s s I •5 -Si FINAL SELECTION is called upon to make nice distinctions between individual seedlings of the same variety. All will have the same general formation of stem and leaf. He must look, then, for details of variation that would altogether escape the notice of the untrained observer. But that such differences exist, and that they are signs that to the practiced eye are of the utmost importance, any successful plant experimenter can testify. It would obviously be futile to attempt a detailed description of the nice shades of distinction between various seedlings of the same race upon which the plant experimenter depends in forming his selective judgments. That, clearly, must be matter for practical observation. It can be learned nowhere but in the field. But perhaps two or three illustrations may be given that will at least serve in a general way to suggest what manner of traits are taken into consideration when the plant experimenter is choosing the individuals with which he is to continue his experiment. A FEW PRACTICAL HINTS In selecting raspberry or blackberry plants for color of fruit, for example, there is almost always a correlation of the plant and fruit that will foretell the future crop. I have observed in thousands of instances that [285] LUTHER BURBANK vines that have purple spines and canes will in future produce berries that are dark purple or dark red in color. Pinkish leaves, on the other hand, foretell fruit of light pink or red color; plants with yellowish vines and foliage may be expected to produce berries of a yellowish color. Very pale foliage and canes usually indicate that the crop will be of a whitish or amber color. A knowledge of this correlation between vine and fruit was of great service to me in my later experiments for the development of the race of white blackberries. It enabled me to select for transplantation and particular care vines that would produce the type of berry I was seeking. It was not necessary to await the time of fruiting in order to gauge progress. The correlation of characters between the vine and the fruit of the grape is not always quite so clearly established, yet it is often observable. Grape tentacles may give clear indication of the size and flavor of the future bunches of fruit. Long before a grape vine has come to the age of fruiting, the taste of the tendrils may give a fair idea of the flavor of the grapes it will ultimately bear. Moreover the seedling vines that produce bushy stems that are small and much branched, and have small leaves, will almost invariably produce [286] FINAL SELECTION meager clusters of small fruit of poor quality. So the wise experimenter will root out such vines without letting them come to maturity. Among plums and peaches the correlation of characters is exceedingly valuable. The case of the plum seedlings already cited suggests the possibility of p re-judgment of fruit from observation of small seedlings. There are a good many characters of leaf and twig that are almost too intangible for description, like the changing expressions of the human face, or like delicately graded colors, yet which to the practiced eye are full of meaning. COLOR OF FOLIAGE A GUIDE A broad general distinction that is fairly obvious to any observer is found in the color of the foliage. It may be expected that a plum or peach seedling having foliage of a reddish purple color will produce fruit dark-colored not only in skin but in flesh. And of course the selection made from any given lot of seedlings will depend largely upon the particular qualities that one desires to develop. But, as repeatedly pointed out, in practical work one is usually looking for a combination of qualities; and, by the same token, one usually inspects his seedlings for the combination of characteristics of stem and leaf and color. He [287] •8 I * few Mil : « FINAL SELECTION seldom has his choice determined by a single characteristic, obvious or otherwise. SELECTING FOR A SINGLE CHARACTER Yet there are cases where an experimenter is working with a single plant-characteristic in view, as, for example, when I successfully attempted to develop scented callas and dahlias and verbenas. Here, obviously, the task of selection is com- paratively simple. We are dealing in each case with a flower that has certain desired qualities of color that are firmly fixed in its heredity. The one conspicuous point of variation among thousands of specimens is the presence or absence of a pleasing aroma. It is necessary, then, merely to select the individual plants that have the most pleasing perfume and to use these only for carrying on the experiment. By making such selection generation after generation, choosing always the sweet- scented and rejecting the others, it proves possible to accentuate and fix the quality of perfume-pro- duction without altering the other characteristics of the respective flowers in question. Again the quality sought may be a particular color of blossom, and it may be desirable to pay attention to this only, practically disregarding all other qualities. Such, for example, was the case with my experiments with the crimson [289] LUTHER BURBANK Eschscholtzia, commonly known as the California poppy. The blossoms of the plant from which my new type of poppy was developed, had a narrow strip of crimson on the inner side of one petal. This was an anomaly that appeared "spon- taneously." Doubtless it was due to some crossing of ancestral strains that brought out a latent character that had long been suppressed. But as to this we can only surmise. The simple fact of the matter was that a blossom did appear that had this narrow strip of crimson on one petal. I seized on this individual blossom as offering material for an experiment in color variation. Seeds from this plant produced the next year several plants that had a trifle more crimson on their blossoms. The following year there was still further improvement, as plants appeared that showed a much larger invasion of the flower petals by the crimson coloring. And by selecting year after year blossoms that showed this increasing tendency to adopt the new color, I produced presently a plant that bore blossoms of a beautiful uniform clear crimson. No trace of the original color remained. This furnishes a very good illustration of selection for color where the material consisted [290] II »1*JI s-§.*|S Q I- §- LUTHER BURBANK of a small strip of an unusual color appearing on blossoms otherwise of a fixed hue. But the same method of selection may some- times be applied to the improvement of the shade of color, or even to the development of a new color, from a flower that shows only a faint departure in shade from the normal. And the same principle of selection, followed out in precisely the same manner, applies to the development of flowers or fruits of varying size, of larger or smaller stem, abundance of blossom, profusion of leaf or flower or fruit, and the like. It is equally possible to alter the proportions of the chemical constituents of a plant in certain instances. The case of my sugar prunes, which were developed to have a sugar content of more than 23 per cent, as against the 15 per cent, of their ancestral type, will be recalled. In a similar way the sugar beet has by mere selection been developed until the races now cultivated contain several times the proportion of sugar of the ancestral beets even of twenty years ago. An interesting experiment in causing the progeny of a certain plant to vary in opposite directions through selection, has been made at the Illinois Agricultural Station. Here the quality under consideration was the protein content — that [292] FINAL SELECTION is to say the amount of nitrogeneous matter— in the kernels of a given variety of corn. The specimen with which the experiment started showed on analysis 10.92 per cent, of protein. Selection was made, among the ears of corn grown from this seed, of the individual specimens having the highest protein content on one hand, and those having the lowest protein content on the other. By continuing this double selection for ten generations, two races of corn were developed, one of which produced seed having an average protein content of 14.26 per cent., while the other, grown in the same field, showed a decrease to 8.64 per cent. This experiment illustrates the possibility of selecting out and fixing new races varying widely as to a single important quality of grain among the descendants of a parent plant of relatively fixed strain. In point of fact no plant is so fixed that its individual members do not show variation ; none so fixed that it does not supply material with which the experimenter may work in producing new varieties. Another illustration of the same thing was given by an allied series of experiments at the Illinois Station at which selection was made with reference, to the height of the ear on the corn [293] 1.1 ^ g «o"°°° 2 S £ & • *«j ^j ~o &o •! |r§ g-o-l $> *:•*-.;: o iniiy.1 :ii:|!ls|s FINAL SELECTION stalk. Seed from the same cob was planted in two fields and grown always under closely similar conditions. But in one field selection was made for breeding purposes from stalks having the ears higher from the ground than the average; and in the other field from ears that were lower than the average. At the end of five years the two fields were so widely diversified that the average height of the ear from the ground in one of them was less than three feet (33.2 inches), whereas in the other field the average height of the ears was fully six feet (72.4 inches). One could not well ask a more striking illustration than this of the possibility of devel- oping new races, differing as to some conspicuous character, by simple selection from a given stock. The case of my winter rhubarb, which came to have a relatively gigantic stalk, will be recalled as of similar import; although in that case the experiment was complicated by having to bear in mind various other qualities in addition to mere size of stalk. My giant corn and the corn with the rainbow-striped leaves are other examples. SOME ALARMING FIGURES But, as repeatedly pointed out, the experiment usually is complicated by the necessity for considering more qualities than one whenever [295] LUTHER BURBANK selection is made with an eye to the production of a commercially valuable variety of flower or fruit or vegetable, and not merely for the purpose of scientific record. We have seen this illustrated again and again; and we have seen also how great are the complications which result when we are called upon to make a selection that will give us not merely one quality — merely size or a given color or sugar content — but a combination of six or eight or ten qualities, all presented in superlative measure. We have seen that the chance of securing any given combination of qualities decreases at a startling geometrical ratio in proportion as the number of qualities increase. The precise formula, as calculated by the biometricians, runs something like this. In case a single pair of qualities is in question — say high protein content versus low protein content in corn — the chances are, if the two strains are crossed, that there will appear in the second generation of their progeny one offspring in four that closely resembles each parent. But when we are considering two qualities- say protein content and height of ear on the stalk— in combination, the chance that there will be an individual of the offspring like each parent [296] FINAL SELECTION in the progeny of the second generation, is only one in 16. And when three qualities are in question the ratio jumps to one in 64; with four qualities it advances to one in 256; with five qualities, to one in 1026. When eight qualities are in question, the chance of producing one offspring showing precisely the combination of qualities of each parent is only one in 98,496. And when we deal with ten qualities we encounter the altogether disconcerting ratio of one to 1,575,936! All of which makes it very clear that the wise plant experimenter does not depend upon mere chance to give him the combination of desired qualities in the production of a new form of flower or fruit. He must make his selection, in any given generation, with reference to one or two pre-eminently desirable qualities, and must be content to accept for the moment such other qualities, however undesirable, as are associated with the desired ones. MULTIPLE SELECTION For example, in developing a stoneless plum, my earliest selections were made with an eye to stonelessness alone. Then as I gradually developed a race of plums in which I was certain of finding a fairly large [297] Inferior Plum Seedling This seedling should be compared with the ones shown on pages 300 and 302. It will be seen that the one here shown is only about ten inches high, and that it branches irregularly, and has a curved stalk, forecasting a tree oj small size and poor formation. The leaves also are small in size and oj inferior quality. FINAL SELECTION proportion of individuals growing stoneless fruit, I could select among these the ones that combined with stonelessness the largest proportion of other good qualities, such as size and color and flavor and abundant bearing. When presently I had, through selection, developed a somewhat fixed strain that combined the qualities of stonelessness with fair size and good flavor, I could then select among the many individuals showing these qualities the particular ones that showed them in fullest measure; and at the same time I could now have in mind one or two other qualities — say color of fruit and keeping quality — and be guided in my selection by a consideration of these traits in addition to the others that had already been fairly fixed. Thus the matter of selection, even when many qualities are to be combined in the ultimate product, is not quite so hopelessly complex as the calculations of the biometricians might lead one to suppose. Yet it is assuredly complex enough to test the patience and the ingenuity of the experimenter to the last degree. So the amateur who enters this fascinating field will do well to begin with simple cases, paying heed to a single quality of any flower or fruit with which he experiments; endeavoring to advance along one line till he acquires skill [299] Fairly Good Plum Seedlings These seedlings are intermediate in quality between the one shown on page 298 and that on page 302. They are about eighteen inches tall, and the better one is of upright growth with large well-Jormed leaves. But it has a rather scant equipment of branches. FINAL SELECTION enough through practice to attempt more complex experiments. Let him, for example, increase the perfume of some familiar garden plant, or develop a race having large blossoms, or one having peculiar brilliancy of color. Any flower bed will show him, among different specimens of the same species, enough of variation to furnish material for his first selection. And he is almost sure to find encouragement through discovery, among the plants grown from this seed, of some that will show the particular quality he has in mind in a more pronounced degree than did the parent plant. So here he will have material for further selection, and step by step he can progress in successive seasons, often more rapidly than he had dared to hope, toward the production of the new variety at which he aims. Of course the time will presently come when the amateur who thus begins with what may be called the alphabet of plant experimentation, will wish to advance to more complicated projects. He will wish to urge his plants along a little more rapidly on the path of variation by means of hybridization. But even here, as will be obvious on a moment's reflection, the experimenter is still dealing with [301] Perfect Plum Seedling This seedling contrasts markedly with the one on page 298 and considerably with those on page 300. It is nearly two feet tall; its stalk is relatively thick, and absolutely perpcn- dicular; it has a fine spread of symmetrically arranged branches; and its leaves (shown in smaller scale than the others because of the larger size of the tree) are oj perfect shape and of fine color and texture. This seedling may be depended on to make a tree far superior to those shown in the other illustrations. Mr. Burbank confidently selects this seedling as the future bearer of valuable fruit. FINAL SELECTION selection. For of course he will not make his hybridizing tests at random, but will select for his parent stock individuals that manifest in pronounced degree the qualities that he wishes to combine in his projected new race. So when we pass from the stage of simple selection of "spontaneous" variations, to the stage of inducing variations along given lines by cross-breeding, we are not abandoning selection but are only dealing with selection in its more complicated aspects. Rightly understood, then, it is not too much to say that the entire task of the plant developer is a matter of selection. First, he may select varieties as nature presents them to him. Second, he may through selective breeding improve these varieties. Next he selects among these and makes combinations for further variations; and then he is ready for a new series of selections. So from first to last it is only the same story presented in different aspects. How important a part does selection play in the life about us! Whether it be in animal or human life, whether it be the selection of materials for a nest, an appropriate club or stone to lay low an enemy, the selection of materials for a dynamo or a pyramid, or of words to convey certain thoughts, aspirations or emotions; but selection [303] LUTHER BURBANK alone from among all the materials supplied by nature no matter how skillfully carried out can never produce the artist's ideal in pigments or marble, the architect's vision of a great struc- ture for the shelter of thousands — universal standards of excellence — unless their production is accomplished by means other than metrical and statistical! The beginning is selection and the end is selection. [END OF VOLUME III] LIST OF DIRECT COLOR PHOTOGRAPH PRINTS IN VOLUME III Apple p.,* Two Burbank Apples 196 Artichoke Giant Artichoke in Blossom , 89 Balloon Berry Balloon Berry Blossom 79 Balloon Berry Bush After Pollenation 81 Bean New Chilian Beans 243 (See also reierence under "Variation.") Budding Cutting a Bud 181 Cutting the Bark to Receive a Bud 182 The Bud Graft Completed 183 Bud After a Year's Growth 185 Bud With Stock Cut Away 187 Cactus Cross-Section of a Cactus Blossom. 65 Mr. Burbank Selecting Cactus Seedlings 284 Chestnut Pollen-Bearing Chestnut Blossom 92 Stigmatic Chestnut Blossoms 93 Corn Hybrid Corn 33 Ten Corn Variations 37 Corn Tassels Bearing Kernels 41 Corn Tassel Growing From the Ear 45 Corn Seventeen Feet High 49 Corn Self-PoIIenated and Crossed With Teosinte 95 Corn Insufficiently PoIIenated 97 Selected Ears of Corn Drying 101 Selected Corn Seed 105 Protecting Test Corn from the Breezes 142 (See also reference under "Variation.") LIST OF ILLUSTRATIONS (Continued) Dandelion p.ge Wild and Improved Dandelions 203 Germination Germinating Seeds in Damp Cloth 106 Seeds Germinated in Damp Cloth 107 Pumpkin Seeds Germinated in Wet Newspaper 109 The Burbank "Flats" 113 " Flat" With Layer of Gravel . . ,. 115 "Flat" Partly Filled with Dirt 117 " Flat" After Seeds Are Sown 119 Gourd Two Gourd Blossoms to be PoIIenated 68 PoIIenating the Gourd Blossoms 69 Grafting Cleft Graft Frontispiece Complete Grafting Outfit 144 Cutting Cion for Whip Graft 149 Cutting Stock for Whip Graft 151 Whip Graft in Place 153 Whip Graft Waxed 155 Whip Graft Bandaged 157 Grafted Wood 161 Cutting Stock for Cleft Graft 163 Cleft Grafts Waxed and Unwaxed 165 Side Graft in Position 167 Side Graft on a Root 169 The Root Graft Completed 171 Crown or Bark Graft 173 Bridge Graft 175 Cutting Stock for Top Graft 177 Top Grafts in Place 179 Quince Cion Ready for Grafting 189 Cions Showing One, Two, and Three Buds 193 (See also "Budding.") Grape Pollen-Bearing Grape 86 Wild Grapes 234 Seedless Grapes 239 Hybridization A Mosaic Leaf 53 LIST OF ILLUSTRATIONS (Continued) Labels and Records Page Labels on Tree Graft 248 Label on Greenhouse "Flat" : 251 Diagram of Tree Grafts 254 Record of Blooming 257 Grafting Record 260 Ripening Record 263 Budding Records . . . 266 Stakes to Mark Divisions of a Verbena Bed 268 Punched Labels 270 Notched Labels 272 Marigold Common, French, and Burbank Marigolds 30 Peach Wild Flowering Peach, and Improved Variety 21 Two Burbank Peaches. . 2ii Pear A Japanese Pear 19 The Chinese Pear 23 Pears with Blended Heredities 27 Pear Seedlings 281 Plum Inferior Plum Seedling 298 Fairly Good Plum Seedling 300 Perfect Plum Seedling 302 PoIIenizing Complete Kit of PoIIenizing Tools 83 (See also pictures listed under "Balloon Berry," "Cactus," "Chestnut," "Gourd," "Pumpkin," etc.) Poppy A Shirley Poppy — Showing Reproductive Organs 56 The Stigma of a Poppy, Greatly Enlarged 59 Pumpkin Pollen-Bearing Pumpkin Blossom 62 Seed-Bearing Pumpkin Blossom 63 Rose Rose Cuttings 229 LIST OF ILLUSTRATIONS (Continued) Seedlings P^ "Flats" with Sprouting Seedlings i 121 Seedlings in the Greenhouse 123 Protecting Seedlings from the Birds 125 Mr. Burbank's Only Cold Frame 127 Mr. Burbank's Cloth Screens 129 (See also references under "Pear" and "Plum.") Squash Patagonian Squash 220 Strawberry A Burbank Strawberry. 6 Strawberries Showing Variation. 1 1 A Giant Strawberry 15 Strawberry Blossom Ready for PoIIenating 73 Strawberry Plant After PoIIenation 75 Strawberry Bed Before Selection 288 A Standard Strawberry 291 Strawberry Bed After Selection 294 Transplanting Setting Out Plants from the "Flat" 131 Making a Trench for Cuttings 133 Setting Out Cuttings in the Trench. 135 Tamping the Dirt Around the Cuttings 137 Seedling Trees Awaiting Transplantation 139 Variation Variation in Corn Seed 274 Chilian Beans 277 Verbena Verbena Bed 209 UNIVERSITY OF CALIFORNIA LIBRARY BERKELEY THIS BOOK IS DUE ON THE LAST DATE STAMPED BELOW Books not returned on time are subject to a fine of 50c t>er volume after the third day overdue, increasing to $1.00 per volume after the sixth day. Books not in demand may be renewed if application is made before of loan period. an is1. MAR 28 f9 MAft 3^92S NO APR 30 192'3 00V FEB 9 19 '-' RECEIVED JUl 0 o CIRCULATION DEPT 50m-7.'16 YD 10834 MMBM U.C. BERKELEY LIBRARIE 3^358 UNIVERSITY OF CALIFORNIA LIBRARY