Lecture XI Striped Flowers
Terminology is an awkward thing. It is as disagreeable to be compelled to make new names, as to be constrained to use the old faulty ones. Different readers may associate different ideas with the same terms, and unfortunately this is the case with much of the terminology of the science of heredity and variability. What are species and what are varieties? How many different conceptions are conveyed by the terms constancy and variability? We are compelled to use them, but we are not at all sure that we are rightly understood when we do so.
Gradually new terms arise and make their way. They have a more limited applicability than the old ones, and are more narrowly circumscribed. They are not to supplant the older terms, but permit their use in a more general way.
[310] One of these doubtful terms is the word sport. It often means bud-variation, while in other cases it conveys the same idea as the old botanical term of mutation. But then all sorts of seemingly sudden variations are occasionally designated by the same term by one writer or another, and even accidental anomalies, such as teratological ascidia, are often said to arise by sports.
If we compare all these different conceptions, we will find that their most general feature is the suddenness and the rarity of the phenomenon. They convey the idea of something unexpected, something not always or not regularly occurring. But even this demarcation is not universal, and there are processes that are regularly repeated and nevertheless are called sports. These at least should be designated by another name.
In order to avoid confusion as far as possible, with the least change in existing terminology, I shall use the term “ever-sporting varieties” for such forms as are regularly propagated by seed, and of pure and not hybrid origin, but which sport in nearly every generation. The term is a new one, but the facts are for the most part new, and require to be considered in a new light. Its meaning will become clearer at once when the illustrations afforded by [311] striped flowers are introduced. In the following discussion it will be found most convenient to give a summary of what is known concerning them, and follow this by a consideration of the detailed evidence obtained experimentally, which supports the usage cited.
The striped variety of the larkspur of our gardens is known to produce monochromatic flowers, in addition to striped ones. They may be borne by the same racemes, or on different branches, or some seedlings from the same parent-plant may bear monochromatic flowers while others may be striped. Such deviations are usually called sports. But they occur yearly and regularly and may be observed invariably when the cultures are large enough. Such a variety I shall call “ever-sporting.” The striped larkspur is one of the oldest garden varieties. It has kept its capacity of sporting through centuries, and therefore may in some sense be said to be quite stable. Its changes are limited to a rather narrow circle, and this circle is as constant as the peculiarities of any other constant species or variety. But within this circle it is always changing from small stripes to broad streaks, and from them to pure colors. Here the variability is a thing of absolute constancy, while the constancy consists in eternal changes. Such apparent [312] contradictions are unavoidable, when we apply the old term to such unusual though not at all new cases. Combining the stability and the qualities of sports in one word, we may evidently best express it by the new term of eversporting variety.
We will now discuss the exact nature of such varieties, and of the laws of heredity which govern them. But before doing so, I might point out, that this new type is a very common one. It embraces most of the so-called variable types in horticulture, and besides these a wide range of anomalies.
Every ever-sporting variety has at least two different types, around and between which it varies in numerous grades, but to which it is absolutely limited. Variegated leaves fluctuate between green and white, or green and yellow, and display these colors in nearly all possible patterns. But there variability ends, and even the patterns are ordinarily narrowly prescribed in the single varieties. Double flowers afford a similar instance. On one side the single type, on the other the nearly wholly double model are the extreme limits, between which the variability is confined. So it is also with monstrosities. The race consists of anomalous and normal individuals, and displays between them all possible combinations of normal and monstrous [313] parts. But its variability is restricted to this group. And large as the group may seem on first inspection, it is in reality very narrow. Many monstrosities, such as fasciated branches, pitchers, split leaves, peloric flowers, and others constitute such ever-sporting varieties, repeating their anomalies year by year and generation after generation, changing as much as possible, but remaining absolutely true within their limits as long as the variety exists.
It must be a very curious combination of the unit-characters which causes such a state of continuous variability. The pure quality of the species must be combined with the peculiarity of the variety in such a way, that the one excludes the other, or modifies it to some extent, although both never fully display themselves in the same part of the same plant. A corolla cannot be at once monochromatic and striped, nor can the same part of a stem be twisted and straight. But neighboring organs may show the opposite attributes side by side.
In order to look closer into the real mechanism of this form of variability, and of this constant tendency to occasional reversions, it will be best to limit ourselves first to a single case, and to try to gather all the evidence, which can be obtained by an examination of the hereditary relations of its sundry constituents.
[314] This may best be done by determining the degree of inheritance for the various constituents of the race during a series of years. It is only necessary to apply the two precautions of excluding all cross-fertilization, and of gathering the seeds of each individual separately. We do not need to ascertain whether the variety as such is permanent; this is already clear from the simple fact of its antiquity in so many cases. We wish to learn what part each individual, or each group of individuals with similar characters, play in the common line of inheritance. In other words, we must build up a genealogical tree, embracing several generations and a complete set of the single cases occurring within the variety, in order to allow of its being considered as a part of the genealogy of the whole. It should convey to us an idea of the hereditary relations during the life-time of the variety.
It is manifest that the construction of such a genealogical tree requires a number of separate experiments. These should be extended over a series of years. Each should include a number of individuals large enough to allow the determination of the proportion of the different types among the offspring of a single plant. A species which is easily fertilized by its own pollen, and which bears capsules with [315] large quantities of seeds, obviously affords the best opportunities. As such, I have chosen the common snapdragon of the gardens, Antirrhinum majus. It has many striped varieties, some tall, others of middle height, or of dwarfed stature. In some the ground-color of the flowers is yellow, in others it is white, the yellow disappearing, with the exception of a large mark in the throat. On these ground-colors the red pigment is seen lying in streaks of pure carmine, with white intervals where the yellow fails, but combined with yellow to make a fiery red, and with yellow intervals when that color is present. This yellow color is quite constant and does not vary in any marked degree, notwithstanding the fact that it seems to make narrower and broader stripes, according to the parts of the corolla left free by the red pigment. But it is easily seen that this appearance is only a fallacious one.
The variety of snapdragon chosen was of medium height and with the yellow ground-color, and is known by horticulturists as A. majus luteum rubro-striatum. As the yellow tinge showed itself to be invariable; I may limit my description to the red stripes.
Some flowers of this race are striped, others are not. On a hasty survey there seem to be three types, pure yellow, pure red, and stripes [316] with all their intermediate links of narrower and broader, fewer and more numerous streaks. But on a close inspection one does not succeed in finding pure yellow racemes. Little lines of red may be found on nearly every flower. They are the extreme type on this side of the range of variability. From them an almost endless range of patterns passes over to the broadest stripes and even to whole sections of a pure red. But then, between these and the wholly red flowers we observe a gap, which may be narrower by the choice of numerous broad striped individuals, but which is never wholly filled up. Hence we see that the red flowers are a separate type within the striped variety.
This red type springs yearly from the striped form, and yearly reverts to it. This is what in the usual descriptions of this snapdragon, is called its sporting. The breadth of the streaks is considered to be an ordinary case of variability, but the red flowers appear suddenly, without the expected links. Therefore they are to be considered as sports. Similarly the red forms may suddenly produce striped ones, and this too is to be taken as a sport, according to the usual conception of the word.
Such sports may occur in different ways. Either by seeds, or by buds, or even within the single spikes. Both opposite reversions, [317] from striped to red and from red to stripes, occur by seed, even by the strictest exclusion of cross-fertilization. As far as my experiments go, they are the rule, and parent-plants that do not give such reversions, at least in some of their offspring, are very rare, if not wholly wanting. Bud-variations and variations within the spike I have as yet only observed on the striped individuals, and never on the red ones, though I am confident that they might appear in larger series of experiments. Both cases are more common on individuals with broad stripes than on plants bearing only the narrower red lines, as might be expected, but even on the almost purely yellow individuals they may be seen from time to time. Bud-variations produce branches with spikes of uniform red flowers. Every bud of the plant seems to have equal chances to be transformed in this way. Some striped racemes bear a few red flowers, which ordinarily are inserted on one side of the spike only. As they often cover a sharply defined section of the raceme, this circumstance has given rise to the term of sectional variability to cover such cases. Sometimes the section is demarcated on the axis of the flower-spike by a brownish or reddish color, sharply contrasting with the green hue of the remaining parts. Sectional variation may be looked at as a [318] special type of bud-variation, and from this point of view we may simplify our inquiry and limit ourselves to the inheritance of three types, the striped plants, the red plants and the red asexual variants of the striped individuals. In each case the heredity should be observed not only for one, but at least for two successive generations.
Leaving these introductory remarks I now come at once to the genealogical tree, as it may be deduced from my experiments:
Year
1896 95% Striped 84% Red
1895 Striped Individual Red Indiv.
\ /
1895 98% Striped 71% Red
1894 Striped branches. Red branches.
\ /
1894 98% Striped 76% Red
1893 90% Striped Indiv. 10% Red Indiv.
\ /
1892 Striped Individual
This experiment was begun in the year 1892 with one individual out of a large lot of striped plants grown from seeds which I had purchased from a firm in Erfurt. The capsules were gathered separately from this individual and about 40 flowering plants were obtained from the seeds in the following year. Most of them had neatly striped flowers, some displayed broader stripes and spare flowers were seen with one [319] half wholly red. Four individuals were found with only uniform red flowers. These were isolated and artificially pollinated, and the same was done with some of the best striped individuals. The seeds from every parent were sown separately, so as to allow the determination of the proportion of uniform red individuals in the progeny.
Neither group was constant in its offspring. But as might be expected, the type of the parent plant prevailed in both groups, and more strongly so in the instances with the striped, than with the red ones. Or, in other words seed-reversions were more numerous among the already reverted reds than among the striped type itself. I counted 2% reversion in the latter case, but 24% from the red parents.
Among the striped plants from the striped parents, I found some that produced bud variations. I succeeded in isolating these red flowering branches in paper bags and in pollinating them with their own pollen, and subjected the striped spikes of the same individuals to a similar treatment. Three individuals gave a sufficient harvest from both types, and these six lots of seeds were sown separately. The striped flowers repeated their character in 98% of their offspring, the red twigs in only 71%, the [320] remaining individuals sporting into the opposite group.
In the following year I continued the experiment with the seeds of the offspring of the red bud-variations. The striped individuals gave 95%, but in the red ones only 84% of the progeny remained true to the parent type.
From these figures it is manifest that the red and striped types differ from one another not only in their visible attributes, but also in the degree of their heredity. The striped individuals repeat their peculiarity in 90-98% of their progeny, 2-10% sporting into the uniform red color. On the other hand the red individuals are constant in 71-84% of their offspring, while 16-29% go over to the striped type. Or, briefly, both types are inherited to a high degree, but the striped type is more strictly inherited than the red one.
Moreover the figures show that the degree of inheritance is not contingent upon the question as to how the sport may have arisen. Bud-sports show the same degree of inheritance as seed-sports. Sexual and asexual variability therefore seem to be one and the same process in this instance. But the deeper meaning of this and other special features of our genealogical tree are still awaiting further investigation. It seems that much important evidence might [321] come from an extension of this line of work. Perhaps it might even throw some light on the intimate nature of the bud-variations of ever-sporting varieties in general. Sectional variations remain to be tested as to the degree of inheritance exhibited, and the different occurrences as to the breadth of the streaks require similar treatment.
In ordinary horticultural practice it is desirable to give some guarantee as to what may be expected to come from the seeds of brightly striped flowers. Neither the pure red type, nor the nearly yellow racemes are the object of the culture, as both of them may be had pure from their, own separate varieties. In order to insure proper striping, both extremes are usually rejected and should be rooted out as soon as the flowering period begins. Similarly the broad-striped ones should be rejected, as they give a too large amount of uniform red flowers. Clearly, but not broadly striped individuals always yield the most reliable seed.
Summing up once more the results of our pedigree-experiment, we may assert that the striped variety of the snapdragon is wholly permanent, including the two opposite types of uniform color and of stripes. It must have been so since it first originated from the invariable uniform [322] varieties, about the middle of the last century, in the nursery of Messrs. Vilmorin, and probably it will remain so as long as popular taste supports its cultivation. It has never been observed to transgress its limits or to sport into varieties without reversions or sports. It fluctuates from one extreme to the other yearly, always recurring in the following year, or even in the same summer by single buds. Highly variable within its limits, it is absolutely constant or permanent, when considered as a definite group.
Similar cases occur not rarely among cultivated plants. In the wild state they seem to be wholly wanting. Neither are they met with as occasional anomalies nor as distinct varieties. On the contrary, many garden-flowers that are colored in the species, and besides this have a white or yellow variety, have also striped sorts. The oldest instance is probably the marvel of Peru, Mirabilis Jalappa, which already had more than one striped variety at the time of its introduction from Peru into the European gardens, about the beginning of the seventeenth century. Stocks, liver-leaf (Hepatica), dame’s violet (Hesperis), Sweet William (Dianthus barbatus), and periwinkles (Vinca minor) seem to be in the same condition, as their striped varieties were already quoted [323] by the writers of the same century. Tulips, hyacinths, Cyclamen, Azalea, Camellia, and even such types of garden-plants as the meadow crane’s-bill (Geranium pratense have striped varieties. It is always the red or blue color which occurs in stripes, the underlying ground being white or yellow, according to the presence or absence of the yellow in the original color mixture.
All these varieties are known to be permanent, coming true during long series of successive generations. But very little is known concerning the more minute details of their hereditary qualities. They come from seed, when this is taken from striped individuals, and thence revert from time to time to the corresponding monochromatic type. But whether they would do so when self-fertilized, and whether the reversionary individuals are always bound to return towards the center of the group or towards the opposite limit, remains to be investigated. Presumably there is nowhere a real transgression of the limits, and never or only very rarely and at long intervals of time a true production of another race with other hereditary qualities.
In order to satisfy myself on these points, I made some pedigree-cultures with the striped forms of dame’s violet (Hesperis matronalis) [324] and of Clarkia pulchella. Both of them are ever-sporting varieties. The experiments were conducted during five generations with the violet, and during four with the striped Clarkia, including the progeny of the striped and of the monochromatic red offspring of a primitive striped plant. I need not give the figures here for the numerical relations between the different types of each group, and shall limit myself to the statement that they behaved in exactly the same manner as the snapdragon.
It is worth while to dwell a moment on the capacity of the individuals with red flowers to reproduce the striped type among their offspring. For it is manifest that this latter quality must have lain dormant in them during their whole life. Darwin has already pointed out that when a character of a grandparent, which is wanting in the progeny, reappears in the second generation, this quality must always be assumed to have been present though latent in the intermediate generation. To the many instances given by him of such alternative inheritance, the monochromatic reversionists of the striped varieties are to be added as a new type. It is moreover, a very suggestive type, since the latency is manifestly of quite another character than for instance in the case of Mendelian hybrids, and probably more allied to those instances, [325] where secondary sexual marks, which are as a rule only evolved by one sex, are transferred to the offspring through the other.
Stripes are by no means limited to flowers. They may affect the whole foliage, or the fruits and the seeds, and even the roots. But all such cases occur much more rarely than the striped flowers. An interesting instance of striped roots is afforded by radishes. White and red varieties of different shapes are cultivated. Besides them sometimes a curious motley sort may be seen in the markets, which is white with red spots, which are few and narrow in some samples, and more numerous and broader in others. But what is very peculiar and striking is the circumstance, that these stripes do not extend in a longitudinal, but in a transverse direction. Obviously this must be the effect of the very notable growth in thickness. Assuming that the colored regions were small in the beginning, they must have been drawn out during the process of thickening of the root, and changed into transverse lines. Rarely a streak may have had its greatest extension in a transverse direction from the beginning, in which case it would only be broadened and not definitely changed in its direction.
This variety being a very fine one, and more agreeable to the eye than the uniform colors, is [326] being more largely cultivated in some countries. It has one great drawback: it never comes wholly true from seed. It may be grown in full isolation, and carefully selected, all red or nearly monochromatic samples being rooted out long before blooming, but nevertheless the seed will always produce some red roots. The most careful selection, pursued through a number of years, has not been sufficient to get rid of this regular occurrence of reversionary individuals. Seed-growers receive many complaints from their clients on this account, but they are not able to remove the difficulty. This experience is in full agreement with the experimental evidence given by the snapdragon, and it would certainly be very interesting to make a complete pedigree-culture with the radishes to test definitely their compliance with the rules observed for striped flowers.
Horticulturists in such cases are in the habit of limiting themselves to the sale of so-called mixed seeds. From these no client expects purity, and the normal and hereditary diversity of types is here in some sense concealed under the impurities included in the mixture from lack of selection. Such cases invite scrutiny, and would, no doubt, with the methods of isolation, artificial pollination, and the sowing of the seeds separately from each parent, yield [327] results of great scientific value. Any one who has a garden, and sufficient perseverance to make pure cultures during a series of years might make important contributions to scientific knowledge in this way.
Choice might be made from among a wide range of different types. A variety of corn called “Harlequin” shows stripes on its kernels, and one ear may offer nearly white and nearly red seeds and all the possible intermediate steps between them. From these seeds the next generation will repeat the motley ears, but some specimens will produce ears of uniform kernels of a dark purple, showing thus the ordinary way of reversion. Some varieties of beans have spotted seeds, and among a lot of them one may be sure to find some purely red ones. It remains to be investigated what will be their offspring, and whether they are due to partial or to individual variation.
The cockscomb (Celosia cristata) has varieties of nearly all colors from white and yellow to red and orange, and besides them some striped varieties occur in our gardens, with the stripes going from the lower parts of the stem up to the very crest of the comb. They are on sale as constant varieties, but nothing has as yet been recorded concerning their peculiar behavior in the inheritance of the stripes. [328] Striped grapes, apples and other fruits might be mentioned in this connection.
Before leaving the striped varieties, attention is called to an interesting deduction, which probably gives an explanation of one of the most widely known instances of ever-sporting garden plants. Striped races always include two types. Both of them are fertile, and each of them reproduces in its offspring both its own and the alternate type. It is like a game of ball, in which the opposing parties always return the ball. But now suppose that only one of the types were fertile and the other for some reason wholly sterile, and assume the reversionary, or primitive monochromatic individuals to be fertile, and the derivative striped specimens to bloom without seed. If this were the case, knowledge concerning the hereditary qualities would be greatly limited. In fact the whole pedigree would be reduced to a monochromatic strain, which would in each generation sport in some individuals into the striped variety. But, being sterile, they would not be able to propagate themselves.
Such seems to be the case with the double flowered stocks. Their double flowers produce neither stamens nor pistils, and as each individual is either double or single in all its flowers, the doubles are wholly destitute of seed. [329] Nevertheless, they are only reproduced by seed from single flowers, being an annual or biennial species.
Stocks are a large family, and include a wonderful variety of colors, ranging from white and yellow to purple and red, and with some variations toward blue. They exhibit also diversity in the habit of growth. Some are annuals, including the ten-week and pyramidal forms; others are intermediates and are suitable for pot-culture; and the biennial sorts include the well-known “Brompton” and “Queen” varieties. Some are large and others are small or dwarf. For their brightness, durability and fragrance, they are deservedly popular. There are even some striped varieties. Horticulturists and amateurs generally know that seed can be obtained from single stocks only, and that the double flowers never produce any. It is not difficult to choose single plants that will produce a large percentage of double blossoms in the following generation. But only a percentage, for the experiments of the most skilled growers have never enabled them to save seed, which would result entirely in double flowering plants. Each generation in its turn is a motley assembly of singles and doubles.
Before looking closer into the hereditary peculiarities of this old and interesting ever-sporting [330] variety, it may be as well to give a short description of the plants with double flowers. Generally speaking there are two principal types of doubles. One is by the conversion of stamens into petals, and the other is an anomaly, known under the name of petalomany.
The change of stamens into petals is a gradual modification. All intermediate steps are easily to be found. In some flowers all stamens may be enlarged, in others only part of them. Often the broadened filaments bear one or two fertile anthers. The fertility is no doubt diminished, but not wholly destroyed. Individual specimens may occur, which cannot produce any seed, but then others of the same lot may be as fertile as can be desired. As a whole, such double varieties are regularly propagated by seed.
Petalomany is the tendency of the axis of some flowers never to make any stamens or pistils, not even in altered or rudimentary form. Instead of these, they simply continue producing petals, going on with this production without any other limit than the supply of available food. Numerous petals fill the entire space within the outer rays, and in the heart of the flower innumerable young ones are developed half-way, not obtaining food enough to attain [331] full size. Absolute sterility is the natural consequence of this state of things.
Hence it is impossible to have races of petalomanous types. If the abnormality happens to show itself in a species, which normally propagates itself in an asexual way, the type may become a vegetative variety, and be multiplied by bulbs, buds or cuttings, etc. Some cultivated anemones and crowfoots (Ranunculus) are of this character, and even the marsh-marigold (Caltha palustris) has a petalomanous variety. I once found in a meadow such a form of the meadow-buttercup (Ranunculus acris), and succeeded in keeping it in my garden for several years, but it did not make seeds and finally died. Camellias are known to have both types of double flowers. The petalomanous type is highly regular in structure, so much so as to be too uniform in all its parts to be pleasing, while the conversion of stamens into petals in the alternative varieties gives to these flowers a more lively diversity of structure. Lilies have a variety called Lilium candidum flore pleno, in which the flowers seem to be converted into a long spike of bright, white narrow bracts, crowded on an axis which never seems to cease their production.
It is manifestly impossible to decide how all such sterile double flowers have originated. [332] Perhaps each of them originally had a congruent single-flowered form, from which it was produced by seed in the same way as the double stocks now are yearly. If this assumption is right, the corresponding fertile line is now lost; it has perhaps died out, or been masked. But it is not absolutely impossible that such strains might one day be discovered for one or another of these now sterile varieties.
Returning to the stocks we are led to the conception that some varieties are absolutely single, while others consist of both single-flowered and double-flowered individuals. The single varieties are in respect to this character true to the original wild type. They never give seed which results in doubles, providing all intercrossing is excluded. The other varieties are ever-sporting, in the sense of this term previously assumed, but with the restriction that the sports are exclusively one-sided, and never return, owing to their absolute sterility.
The oldest double varieties of stocks have attained an age of a century and more. During all this time they have had a continuous pedigree of fertile and single-flowered individuals, throwing off in each generation a definite number of doubles. This ratio is not at all dependent on chance or accident, nor is it even variable to a remarkable degree. Quite on the contrary [333] it is always the same, or nearly the same, and it is to be considered as an inherent quality of the race. If left to themselves, the single individuals always produce singles and doubles in the same quantity; if cultivated after some special method, the proportion may be slightly changed, bringing the proportion of doubles up to 60% or even more.
Ordinarily the single and double members of such a race are quite equal in the remainder of their attributes, especially in the color of their flowers. But this is not always the case. The colors of such a race may repeat for themselves the peculiarities of the ever-sporting characters. It often happens that one color is more or less strictly allied to the doubles, and another to the singles. This sometimes makes it difficult to keep the various colors true. There are certain sorts, which invariably exhibit a difference in color between the single and the double flowers. The sulphur-yellow varieties may be adduced as illustrative examples, because in them the single flowers always come white. Hence in saving seed, it is impossible so to select the plant, that an occasional white does not also appear among the double flowers, agreeing in this deviation with the general rule of the eversporting varieties.
I commend all the above instances to those [334] who wish to make pedigree-cultures. The cooperation of many is needed to bring about any notable advancement, since the best way to secure isolation is to restrict one’s self to the culture of one strain, so as to avoid the intermixture of others. So many facts remain doubtful and open to investigation, that almost any lot of purchased seed may become the starting point for interesting researches. Among these the sulphur-yellow varieties should be considered in the first place.
In respect to the great questions of heredity, the stocks offer many points of interest. Some of these features I will now try to describe, in order to show what still remains to be done, and in what manner the stocks may clear the way for the study of the ever-sporting varieties.
The first point, is the question, which seeds become double-flowered and which single-flowered plants? Beyond all doubt, the determination has taken place before the ripening of the seed. But though the color of the seed is often indicative of the color of the flowers, as in some red or purple varieties, and though in balsams and some other instances the most “highly doubled” flowers are to be obtained from the biggest and plumpest seeds, no such rule seems to exist respecting the double stocks. Now if one half of the seeds gives doubles, and [335] the other half singles, the question arises, where are the singles and the doubles to be found on the parent-plant? The answer is partly given by the following experiment. Starting from the general rule of the great influence of nutrition on variability, it may be assumed that those seeds will give most doubles, that are best fed. Now it is manifest that the stem and larger branches are, in a better condition than the smaller twigs, and that likewise the first fruits have better chances than the ones formed later. Even in the same pod the uppermost seeds will be in a comparatively disadvantageous position. This conception leads to an experiment which is the basis of a practical method much used in France in order to get a higher percentage of seeds of double-flowering plants.
This method consists in cutting off, in the first place the upper parts of all the larger spikes, in the second place, the upper third part of each pod, and lastly all the small and weak twigs. In doing so the percentage is claimed to go up to 67-70%, and in some instances even higher. This operation is to be performed as soon as the required number of flowers have ceased blossoming. All the nutrient materials, destined for the seeds, are now forced to flow into these relatively few embryos, and it is clear that [336] they will be far better nourished than if no operation were made.
In order to control this experiment some breeders have made the operation on the fruits when ripe, instead of on the young pods, and have saved the seeds from the upper parts separately. This seed, produced in abundance, was found to be very poor in double flowers, containing only some 20-30%. On the contrary the percentage of doubles in the seed of the lower parts was somewhat augmented, and the average of both would have given the normal proportion of 50%.
Opposed to the French method is the German practice of cultivating stocks, as I have seen it used on a very large scale at Erfurt and at other places. The stocks are grown in pots on small scaffolds, and not put on or into the earth. The obvious aim of this practice is to keep the earth in the pots dry, and accordingly they are only scantily watered. In consequence they cannot develop as fully as they would have done when planted directly in the beds, and they produce only small racemes and no weak twigs, eliminating thereby without further operation the weaker seeds as by the French method. The effect is increased by planting from 6-10 separate plants in each pot.
It would be very interesting to make comparative [337] trials of both methods, in order to discover the true relation between the practice and the results reached. Bath should also be compared with cultures on open plots, which are said to give only 50% of doubles. This last method of culture is practiced wherever it is desired to produce great quantities of seeds at a low cost. Such trials would no doubt give an insight into the relations of hereditary characters to the distribution of the food within the plant.
A second point is the proportional increase of the double-flowering seeds with age. If seed is kept for two or three years, the greater part of the grains will gradually die, and among the remainder there is found on sowing, a higher percentage of double ones. Hence we may infer that the single-flowered seeds are shorter lived than the doubles, and this obviously points to a greater weakness of the first. It is quite evident that there is some common cause for these facts and for the above cited experience, that the first and best pods give more doubles. Much, however, remains to be investigated before a satisfactory answer can be made to these questions.
A third point is the curious practice, called by the French “esimpler,” and which consists in pulling out the singles when very young. It seems to be done at an age when the flower-buds [338] are not yet visible, or at least are not far enough developed to show the real distinctive marks. Children may be employed to choose and destroy the singles. There are some slight differences in the fullness and roundness of the buds and the pubescence of the young leaves. Moreover the buds of the doubles are said to be sweeter to the taste than those of the singles. But as yet I have not been able to ascertain, whether any scientific investigation of this process has ever been made, though according to some communications made to me by the late Mr. Cornu, the practice seems to be very general in the environs of Paris. In summer large fields may be seen, bearing exclusively double flowers, owing to the weeding out of the singles long before flowering.
Bud-variation is the last point to be taken up. It seems to be very rare with stocks, but some instances have been recorded in literature. Darwin mentions a double stock with a branch bearing single flowers, and other cases are known to have occurred. But in no instance does the seed of such a bud-variant seem to have been saved. Occasionally other reversions also occur. From time to time specimens appear with more luxurious growth and with divergent instead of erect pods. They are called, in Erfurt, “generals” on account [339] of their stiff and erect appearance, and they are marked by more divergent horns crowning the pods. They are said to produce only a relatively small number of doubles from their seeds, and even this small number might be due to fertilization with pollen of their neighbors. I saw some of these reversionary types; when inspecting the nurseries of Erfurt, but as they are, as a rule, thrown out before ripening their seed, nothing is exactly known about their real hereditary qualities.
Much remains to be cleared up, but it seems that one of the best means to find a way through the bewildering maze of the phenomena of inheritance, is to make groups of related forms and to draw conclusions from a comparison of the members of such groups. Such comparisons must obviously give rise to questions, which in their turn will directly lead to experimental investigation.
[340]
LECTURE XII
FIVE-LEAVED CLOVER
Every one knows the “four-leaved” clover. It is occasionally found on lawns, in pastures and by the roadsides. Specimens with five leaflets may be found now and then in the same place, or on the same plant, but these are rarer. I have often seen isolated plants with quaternate leaves, but only rarely have I observed individuals with more than one such leaf.
The two cases are essentially dissimilar. They may appear to differ but little morphologically, but from the point of view of heredity they are quite different. Isolated quaternate leaves are of but little interest, while the occurrence of many on the same individual indicates a distinct variety. In making experiments upon this point it is necessary to transplant the divergent individuals to a garden in order to furnish them proper cultural conditions and to keep them under constant observation. When a plant bearing a quaternate leaf is thus transplanted however, it rarely repeats the [341] anomaly. But when plants with two or more quaternate leaves on the same individual are chosen it indicates that it belongs to a definite race, which under suitable conditions may prove to become very rich in the anomalies in question.
Obviously it is not always easy to decide definitely whether a given individual belongs to such a race or not. Many trials may be necessary to secure the special race. I had the good fortune to find two plants of clover, bearing one quinate and several quaternate leaves, on an excursion in the neighborhood of Loosdrecht in Holland. After transplanting them into my garden, I cultivated them during three years and observed a slowly increasing number of anomalous leaves. This number in one summer amounted to 46 quaternate and 16 quinate leaves, and it was evident that I had secured an instance of the rare “five-leaved” race which I am about to describe.
Before doing so it seems desirable to look somewhat closer into the morphological features of the problem. Pinnate and palmate leaves often vary in the number of their parts. This variability is generally of the nature of a common fluctuation, the deviations grouping themselves around an average type in the ordinary way. Ash leaves bear five pairs, and [342] the mountain-ash (Sorbus Aucuparia) has six pairs of leaflets in addition to the terminal one. But this number varies slightly, the weaker leaves having less, the stronger more pairs than the average. Such however, is not the case, with ternate leaves, which seem to be quite constant. Four leaflets occur so very rarely that one seems justified in regarding them rather as an anomaly than, as a fluctuation. And this is confirmed by the almost universal absence of two-bladed clover-leaves.
Considering the deviation as an anomaly, we may look into its nature. Such an inquiry shows that the supernumerary leaflets owe their origin to a splitting of one or more of the normal ones. This splitting is not terminal, as is often the case with other species, and as it may be seen sometimes in the clover. It is for the most part lateral. One of the lateral nerves grows out becoming a median nerve of the new leaflet. Intermediate steps are not wanting, though rare, and they show a gradual separation of some lateral part of a leaflet, until this division reaches the base and divides the leaflet into two almost equal parts. If this splitting occurs in one leaflet we get the “four-leaved” Clover, if it occurs in two there will be five leaflets. And if, besides this, the terminal leaflet produces a derivative on one or both of its sides, [343] we obtain a crown of six or seven leaflets on one stalk. Such were often met with in the race I had under cultivation, but as a rule it did not exceed this limit.
The same phenomenon of a lateral doubling of leaflets may of course be met with in other instances. The common laburnum has a variety which often produces quaternate and quinate leaves, and in strawberries I have also seen instances of this abnormality. It occurs also in pinnate leaves, and complete sets of all the intermediate links may often be found on the false or bastard-acacia (Robinia Pseud-Acacia).
Opposed to this increase of the number of leaflets, and still more rare and more curious is the occurrence of “single-leaved” varieties among trees and herbs with pinnate or ternate leaves. Only very few instances have been described, and are cultivated in gardens. The ashes and the bastard-acacia may be quoted among trees, and the “one-leaved” strawberry among herbs. Here it seems that several leaflets have been combined into one, since this one is, as a rule, much larger than the terminal leaflet of an ordinary leaf of the same species. These monophyllous varieties are interesting also on account of their continuous but often incomplete reversion to the normal type.
[344] Pinnate and palmate leaves are no doubt derivative types. They must have originated from the ordinary simple leaf. The monophylly may therefore be considered as a reversion to a more primitive state and the monophyllous varieties may be called atavistic.
On the other hand we have seen that these atavistic varieties may revert to their nearest progenitors, and this leads to the curious conception of positive and negative atavism. For if the change of compound leaves into single ones is a retrograde or negative step, the conversion of single or ternate leaves into pinnate and palmate ones must evidently be considered in this case as positive atavism.
This discussion seems to throw some light on the increase of leaflets in the clover. The pea family, or the group of papilionaceous plants, has pinnate leaves ordinarily, which, according to our premises, must be considered as a derivative type. In the clovers and their allies this type reverts halfway to the single form, producing only three leaflets on each stalk. If now the clover increases its number of leaflets, this may be considered as a reversion to its nearest progenitors, the papilionaceous plants with pinnate leaves. Hence a halfway returning and therefore positive atavism. And as I have already mentioned in a former lecture, pinnate [345] leaves are also sometimes produced by my new race of clover.
Returning to the original plants of this race, it is evidently impossible to decide whether they were really the beginning of a new strain, and had originated themselves by some sudden change from the common type, or whether they belonged to an old variety, which had propagated itself perhaps during centuries, unobserved by man. But the same difficulty generally arises when new varieties are discovered. Even the behavior of the plants themselves or of their progeny does not afford any means of deciding the question. The simplest way of stating the matter therefore, is to say that I accidentally found two individuals of the “five-leaved” race. By transplanting them into my garden, I have isolated them and kept them free from cross-fertilization with the ordinary type. Moreover, I have brought them under such conditions as are necessary for the full development of their characters. And last but not least, I have tried to improve this character as far as possible by a very rigid and careful selection.
The result of all this effort has been a rapid improvement of my strain. I saved the seed of the original plants in 1889 and cultivated the second generation in the following year. It [346] showed some increase of the anomaly, but not to a very remarkable degree. In the flowering period I selected four plants with the largest number of quaternate and quinate leaves and destroyed all the others. I counted in the average 25 anomalous organs on each of them. From their seed I raised the third generation of my culture in the year 1891.
This generation included some 300 plants, on which above 8,000 leaves were counted. More than 1,000 were quaternate or quinate, the ternate leaves being still in the majority. But the experiment clearly showed that “four-leaved” clovers may be produced in any desired quantity, provided that the seed of the variety is available. In the summer only three, four and five leaflets on one stalk were seen, but towards the fall, and after the selection of the best individuals, this number increased and came up to six and seven in some rare instances.
The selection in this year was by no means easy. Nearly all the individuals produced at least some quaternate leaves, and thereby showed the variety to be quite pure. I counted the abnormal organs on a large group of the best plants, and selected 20 excellent specimens from them, with more than one-third of all their leaves changed in the desired manner. Having brought my race up to this point, I [347] was able to introduce a new and far more easy mark, afforded by the seedlings, for my selections. This mark has since remained constant, and has brought about a rapid continuance of the improvement, without necessitating such large cultures.
This seedling in the various species of clover usually begins with a first leaf above the cotyledons of a different structure from those that follow. It has only one blade instead of three. But in my variety the increase of the number of the leaflets may extend to these primary organs, and make them binate or even ternate. Now it is obvious that an individual, which begins with a divided primary leaf, will have a greater tendency to produce a large number of supernumerary leaflets than a plant which commences in the ordinary way. Or in other words, the primary leaves afford a sure criterion for the selection, and this selection may be made in the seed-pans. In consequence, no young individual with an undivided primary leaf was planted out. Choosing the 20 or 30 best specimens in the seed-pan, no further selection was required, and the whole lot could be left to cross-fertilization by insects.
The observation of this distinguishing mark in the young seedlings has led to the discovery of another quality as a starting-paint for further [348] selection. According to the general rule of pedigree-culture, the seeds of each individual plant are always saved and sowed separately. This is done even with such species as the clover, which are infertile when self-pollinated, and which are incapable of artificial pollination on the required scale, since each flower produces only one seed. My clover was always left free to be pollinated by insects. Obviously this must have led to a diminution of the differentiating characters of the individual plants. But this does not go far enough to obliterate the differences, and the selection made among the seedlings will always throw out at least a large part of those that have suffered from the cross.
Leaving this discussion, we may inquire closer into the nature of the new criterion afforded by the seedlings. Two methods present themselves. First, the choice of the best seedlings. In the second place it becomes possible to compare the parent-plants by counting the number of deviating seedlings. This leads to the establishment of a percentage for every single parent, and gives data for comparisons. Two or three hundreds of seeds from a parent may easily be grown in one pan, and in this way a sufficiently high degree of accuracy may be reached. Only those parents that give [349] the highest percentage are chosen, and among their progeny only the seedlings with trifoliolate primary leaves are planted out. The whole procedure of the selection is by this means confined to the glasshouse during the spring, and the beds need not be large, nor do they require any special care during the summer.
By this method I brought my strain within two years up to an average of nearly 90% of the seedlings with a divided primary leaf. Around this average the real numbers fluctuated between the maximum of 99% and the minimum of 70% or thereabouts. This condition was reached by the sixth generation in the year 1894, and has since proved to be the limit, the group of figures remaining practically the same during all the succeeding generations.
Such selected plants are very rich in leaves with four, five and six blades. Excluding the small leaves at the tops of the branches, and those on the numerous weaker side-branches, these three groups include the large majority of all the stronger leaves. In summer the range is wider, and besides many trifoliolate leaves the curiously shaped seven-bladed ones are not at all rare. In the fall and in the winter the range of variability is narrowed, and at first sight the plants often seem to bear only quinquefoliolate leaves.
[350] I have cultivated a new generation of this race nearly every year since 1894, using always the strictest selection. This has led to a uniform type, but has not been adequate to produce any further improvement. Obviously the extreme limit, under the conditions of climate and soil, has been reached. This extreme type is always dependent upon repeated selection. No constant variety, in the older sense, has been obtained, nor was any indication afforded that such a type might ever be produced. On the contrary, it is manifest that the new form belongs to the group of ever-sporting varieties. It is never quite free from the old atavistic type of the trifoliolate leaves, and invariably, when external conditions become less favorable, this atavistic form is apt to gain dominion over the more refined varietal character. Reversions always occur, both partial and individual.
Some instances of these reversions may now be given. They are not of such a striking character as those of the snapdragon. Intermediate steps are always occurring, both in the leaves themselves, and in the percentages of deviating seedlings of the several parent plants.
On normal plants of my variety the quinquefoliolate leaves usually compose the majority, when there are no weak lateral branches, or when they are left out of consideration. Next [351] to these come the fours and the sixes, while the trifoliolate and seven-bladed types are nearly equal in number. But out of a lot of plants, grown from seed of the same parent, it is often possible to choose some in which one extreme prevails, and others with a preponderating number of leaves with the other extreme number of leaflets. If seed from these extremes are saved separately, one strain, that with numerous seven-bladed leaves will remain true to the type, but the other will diverge more or less, producing leaves with a varying number of subdivisions.
Very few generations of such opposite selection are required to reduce the race to an utterly poor one. In three years I was able to nearly obliterate the type of my variety. I chose the seedlings with an undivided primary leaf, cultivated them and counted their offspring separately after the sowing. I found some parents with only 2-3% of seedlings with divided primary leaves. And by a repeated selection in this retrograde direction I succeeded in getting a great number of plants, which during the whole summer made only very few leaves with more than three blades. But an absolute reversion could no more be reached in this direction than in the normal one. Any sowing without selection would be [352] liable to reduce the strain to an average condition.
The production of varietal and of atavistic leaves is dependent to a high degree on external conditions. It agrees with the general rule, that favorable circumstances strengthen the varietal peculiarities, while unfavorable conditions increase the number of the parts with the atavistic attribute. These influences may be seen to have their effect on the single individuals, as well as on the generations growing from their seed. I cannot cite here all the experimental material, but a single illustrative example may be given. I divided a strong individual into two parts, planted one in rich soil and the other in poor sand, and had both pollinated by bees with the pollen of some normal individuals of my variety growing between them. The seeds of both were saved and sown separately, and the two lots of offspring cultivated close to each other under the same external conditions. In the beginning no difference was seen, but as soon as the young plants had unfolded three or four leaves, the progeny of the better nourished half of the parent plant showed a manifest advance. This difference increased rapidly and was easily seen in the beds, even before the flowering period.
This experience probably gives an explanation [353] why the quinquefoliolate variety is so seldom met with in the wild state. For even if it did occur more often, the plants would hardly find circumstances favorable enough for the full development of their varietal character. They must often be so poor in anomalous leaves as to be overlooked, or to be taken for instances of the commonly occurring quadrifoliolate leaves and therefore as not indicating the true variety.
In the beginning of my discussion I have asserted the existence of two different races of “four-leaved” clovers, a poor one and a rich one, and have insisted on a sharp distinction between them. This distinction partly depends on experiments with clover, but in great part on tests with other plants. The previously mentioned circumstance, that clover cannot be pollinated on a sufficiently large scale otherwise than by insects, prevents trials in more than one direction at the same time and in the same garden. For this reason I have chosen another species of clover to be able to give proof or disproof of the assertion quoted.
This species is the Italian, or crimson clover, which is sometimes also called scarlet clover (Trifolium incarnatum). It is commonly used in Europe as a crop on less fertile soils than are required by the red clover. It is annual [354] and erect and more or less hairy, and has stouter leaves than other kinds of clover. It has oblong or cylindrical heads with bright crimson flowers, and may be considered as one of the most showy types. As an annual it has some manifest advantages over the perennial species, especially in giving its harvest of hay at other seasons of the year.
I found some stray quaternate leaves of this plant some years ago, and tried to win from them, through culture and selection, a race that would be as rich in these anomalies as the red clover. But the utmost care and the most rigid selection, and all the attention I could afford, failed to produce any result. It is now ten years since I commenced this experiment, and more than once I have been willing to give it up. Last year (1903) I cultivated some hundreds of selected plants, but though they yielded a few more instances of the desired anomaly than in the beginning, no trace of a truly rich race could be discovered. The experimental evidence of this failure shows at least that stray “four-leaves” may occur, which do not indicate the existence of a true “four-” or “five-leaved” variety.
This conception seems destined to become of great value in the appreciation of anomalies, as they are usually found, either in the wild state [355] or in gardens. And before describing the details of my unsuccessful pedigree-culture, it may be as well to give some more instances of what occurs in nature.
Stray anomalies are of course rare, but not so rare that they might not be found in large numbers when perseveringly sought for. Pitcher-like leaves may be found on many trees and shrubs and herbs, but ordinarily one or only two of them are seen in the course of many years on the same plant, or in the same strain. In some few instances they occur annually or nearly so, as in some individuals of the European lime-tree (Tilia parvifolia) and of the common magnolia (Magnolia obovata). Many of our older cultivated plants are very rich in anomalies of all kinds, and Cyclamen, Fuchsia, Pelargonium and some others are notorious sources of teratologic phenomena. Deviations in flowers may often be seen, consisting of changes in the normal number of the several organs, or alterations in their shape and color. Leaves may have two tips, instead of one, the mid-vein being split near the apex, and the fissure extending more or less towards the base. Rays of the umbels of umbelliferous plants may grow together and become united in groups of two or more, and in the same way the fruits of [356] the composites may be united into groups. Many other instances could easily be given.
If we select some of these anomalies for breeding-experiments, our results will not agree throughout, but will tend to group themselves under two heads. In some cases the isolation of the deviating individuals will at once show the existence of a distinct variety, which is capable of producing the anomaly in any desired number of instances; only dependent on a favorable treatment and a judicious selection. In other cases no treatment and no selection are adequate to give a similar result, and the anomaly remains refractory despite all our endeavors to breed it. The cockscomb and the peloric fox-glove are widely known instances of permanent anomalies, and others will be dealt with in future lectures. On the other hand I have often tried in vain to win an anomalous race from an accidental deviation, or to isolate a teratologic variety out of more common aberrations. Two illustrative examples may be quoted. In our next lecture we shall deal with a curious phenomenon in poppies, consisting in the change of the stamens into pistils and giving rise to a bright crown of secondary capsules around the central one. Similar anomalies may be occasionally met with in other species of the same genus. But they are rare, and may show [357] the conversion of only a single stamen in the described manner. I observed this anomaly in a poppy called Papaver commutatum, and subjected it during several years to a rigid selection of the richest individuals. No amelioration was to be gained and the culture had to be given up. In the same way I found on the bulbous buttercup (Ranunculus bulbosus) a strain varying largely in the number of the petals, amounting often to 6-8, and in some flowers even yet to higher figures. During five succeeding years I cultivated five generations, often in large numbers, selecting always those which had the highest number of petals, throwing out the remainder and saving the seed only from the very best plants. I got a strain of selected plants with an average number of nine petals in every flower, and found among 4,000 flowers four having 20 petals or more, coming up even to 31 in one instance. But such rare instances had no influence whatever on the selection, since they were not indicative of individual qualities, but occurred quite accidentally on flowers of plants having only the average number of petals. Now double flowers are widely known to occur in other species of the buttercups, both in the cultivated varieties and in some wild forms. For this reason it might be expected that through a continuous selection of [358] the individuals with the largest numbers a tendency to become double would be evolved. Such, however, was not the case. No propensity to vary in any definite direction could be observed. Quite on the contrary, an average condition was quickly reached, and then remained constant, strongly counteracting all selection.
Such experiences clearly show that the same anomaly may occur in different species, and no doubt in strains of the same species from different localities, according to at least two different standards. The one is to be called the poor, and the other the rich variety. The first always produces relatively few instances of the deviation, the last is apt to give as many of them as desired. The first is only half-way a variety, and therefore would deserve the name of a half-race; the second is not yet a full constant variety, but always fluctuates to and fro between the varietal and the specific mark, ever-sporting in both directions. It holds a middle position between a half-race and a variety, and therefore might be called a “middle-race.” But the term ever-sporting variety seems more adequate to convey a right idea of the nature of this curious type of inheritance.
From this discussion it will be seen that the behavior of the crimson clover is not to be considered [359] as an exception, but as a widely occurring type of phenomenon, occurring perhaps in all sorts of teratologic deviations, and in wide ranges of species and genera. Hence it may be considered worth while to give some more details of this extended experiment.
Ten years ago (1894-5) I bought and sowed about a pound of seed of the crimson clover. Among many thousands of normal seedlings I found two with three and one with four cotyledons. Trusting to the empirical rules of correlation, I transplanted these three individuals in order to isolate them in the flowering period.
One of them produced during the ensuing summer one four-bladed and one five-bladed leaf. The seeds were saved separately and sown the following spring and the expected result could soon be seen. Among some 250 individual plants I counted 22 with one or two deviations, and 10 with from three to nine four- or five-bladed leaves. Proportions nearly similar have been observed repeatedly. Better nourished individuals have produced more deviating leaves on one plant, partly owing to the larger number of stems and branches, and poor or average specimens have mostly been without any aberration or with only one or two abnormal leaves. No further improvement could be attained. Quadrifoliolate leaves were always rare, never [360] attaining a number that would put its stamp on a whole bed. I have endeavored to get some six- and seven-bladed crimson clover leaves, but in vain; selection, culture of many hundreds of individuals, manure, and the best possible treatment has not been adequate to produce them. Of course I am quite convinced that a repetition of my experiment on a far larger scale would yield the desired types, but then only in such rare instances that they would have no influence whatever on the average, or on the improvement of the race. The eighth generation in the year 1903 has not been noticeably better than the second and third generations after the first selection.
In comparing this statement with the results gained in the experiment with the red clover, the difference is at once striking. In one case a rich variety was isolated, and, by better treatment and sharp methods of selection, was brought up in a few years to its highest pitch of development. In the other case a very weak race was shown to exist, and no amount of work and perseverance was adequate to improve it to any noticeable degree.
I wish to point out that the decision of what is to be expected from deviating specimens may become manifest within one or two generations. Even the generation grown from the seeds of [361] the first observed aberrant-individuals, if gathered after sufficient isolation during the period of blossoming, may show which type of inheritance is present, whether it is an unpromising half-race, or a richly endowed sporting variety. I have kept such strains repeatedly after the first isolation, and a special case, that of cotyledoneous aberrations, will be dealt with later. The first generation always gave a final decision, provided that a suitable method of cultivation for the species under observation was found at the beginning. This however, is a condition, which it is not at all easy to comply with, when new sorts are introduced into a garden. Especially so when they had been collected in the wild state. Often one or two years, sometimes more, are necessary to find the proper method of sowing, manuring, transplanting and, other cultural methods satisfactory to the plants. Many wild species require more care and more manure in gardens than the finest garden flowers. And a large number are known to be dependent on very particular conditions of soil.
One of the most curious features of anomalies, which has been learned from accumulated instances, is the fact that they obey definite laws as to their occurrence on the different parts of the plant. Obviously such laws are [362] not apparent as long as each plant produces only one or two, or, at most, a few instances of the same deviation. On the contrary, any existing regularity must betray itself, as soon as a larger number of instances is produced. A rule of periodicity becomes most clearly manifest in such cases.
This rule is shown by no other race in a more undoubted and evident manner than by the “five-leaved” clover. Evidently the several degrees of deviation, going from three to seven leaflets, may be regarded as responses to different degrees of variation, and their distribution over the stems and branches, or over the whole plant, may be considered as the manifestation of the ever-changing internal tendency to vary.
Considered from this point of view, my plants always showed a definite periodicity in this distribution, which is the same for the whole plant. Each of them, and each of the larger branches, begin with atavistic leaves or with slight deviations. These are succeeded by greater deviations, but only the strongest axes show as many as seven leaflets on a stalk. This ordinarily does not occur before the height of development is reached, and often only towards its close. Then the deviation diminishes rapidly, returning often to atavistic leaves at the summit of the stem or branch. I give the numbers of the [363] leaves of a branch, in their order from the base to the top. They were as follows:
3. 4. 5. 6. 7. 5. 5. 4.
But this is a selected case, and such regular examples of the expected periodicity are rarely found. Often one or more of the various steps are lacking, or even leaves with smaller numbers may be interspersed among those with larger numbers of leaflets. But while the regularity of the periodicity is in some degree diminished by such occurrences, yet the rule always holds good, when taken broadly. It may be expressed by stating that the bases and apices have on the average fewer leaflets on each leaf than the middle parts of the stem and branches, and that the number of leaflets gradually increases from the base toward a maximum, which is reached in organs on the middle or upper part of the axis, and then diminishes from this toward the apex.
This periodicity is not limited to the stems and branches, considered singly, but also holds good in a comparison made between the branches of a single stem, in regard to their relative places on that stem. So it is also for the whole plant. The first stems, produced by the subterranean axis, ordinarily show only a low maximum deviation: the next succeeding being [364] more divergent and the last ones returning to less differentiated forms.
It is evident that on a given stem the group of deviating leaves will be extended upward and downward, with the increase of the number of these organs. This shows that a stem, or even a plant, promises a higher degree of differentiation if it commences with its aberration earlier. Hence it becomes possible to discern the most promising individuals in early youth, and this conclusion leads to a very easy and reliable method of selection, which may be expressed simply as follows: the seedlings which commence earliest with the production of four- and five-foliolate leaves are the best and should be selected for the continuance of the race. And it is easily seen that this rule agrees with that given above, and which was followed in my pedigree-culture.
Furthermore it is seen that there is a complete agreement between the law of periodicity and the responses of the deviations to nourishment and other conditions of life. Weak plants only produce low degrees of deviation, the stronger the individual becomes, the higher it reaches in the scale of differentiation, and the more often it develops leaves with five or more blades. Whether weakness or strength are derived from outer causes, or from the internal [365] succession of the periods of life, is evidently of no consequence, and in this way the law of periodicity may be regarded as a special instance of the more general law of response to external conditions.
The validity of this law of periodicity is of course not limited to our “five-leaved” clover. Quite on the contrary it is universal in eversporting varieties. Moreover it may be ascertained and studied in connection with the most widely different morphologic abnormalities, and therefore affords easily accessible material for statistical inquiry. I will now give some further instances, but wish to insist first upon the necessity of an inquiry on a far larger scale, as the evidence as yet is very scanty.
The great celandine (Chelidonium majus) has a very curious double variety. Its flowers are simpler and much more variable than in ordinary garden-varieties. The process of doubling consists mainly in a change of stamens into petals. This change is dependent on the season. On each stem the earliest flowers are single. These are succeeded by blossoms with one or two converted stamens, and towards the summer this number increases gradually, attaining 10-11 and in some instances even more altered filaments. Each year the same succession may be seen repeating itself on the stems of [366] the old roots. Double tuberous begonias are ordinarily absolutely sterile throughout the summer, but towards autumn the new flowers become less and less altered, producing some normal stamens and pistils among the majority of metamorphosed organs. From these flowers the seeds are saved. Sometimes similar flowers occur at the beginning of the flowering-period. Double garden-camomiles (Chrysanthemum inodorum plenissimum) and many other double varieties of garden-plants among the great family of the composites are very sensitive to external agencies, and their flower-heads are fuller the more favorable the external conditions. Towards the autumn many of them produce fewer and fewer converted heads and often only these are fertile and yield seeds.
Ascidia afford another instance of this periodicity, though ordinarily they are by far too rare to show any regularity in their distribution. However, it is easy to observe that on lime-trees they prefer the lower parts of each twig, while on magnolias the terminal leaves of the branches are often pitcher-bearing. Ascidia of the white clover have been found in numbers, in my own experiment-garden, but always in the springtime. The thickleaved saxifrage (Saxifraga crassifolia) is often very productive of ascidia, especially in [367] the latter part of the season, and as these organs may be developed to very different degrees, they afford fine material for the study of the law of periodicity. On a garden-cytisus (Cytisus candicans attleyanus) I once had the good fortune to observe a branch with ascidia, which ordinarily are very rare in this species. It had produced seven ascidia in all, each formed by the conversion of one leaflet on the trifoliolate leaves. The first six leaves were destitute of this malformation and were quite normal. Then followed a group of five leaves, constituting the maximum of the period. The first bore one small pitcher-like blade, the second and third, each one highly modified organ, the fourth, two ascidia, and the last, one leaflet with slightly connate margins. The whole upper part of the branch was normal, with the exception of the seventeenth leaf, which showed a slight change in the same direction. All in all, the tendency to produce ascidia increased from the beginning to the tenth leaf, and decreased from this upward.
The European Venus’ looking-glass was observed in my garden to produce some quaternate and some quinate flowers on the same specimens. The quinate were placed at the end of the branches, those with four petals and sepals lower down. The peloric fox-glove shows the [368] highest degree of metamorphy in the terminal flowers of the stem itself, the weaker branches having but little tendency towards the formation of the anomaly. The European pine or Pinus sylvestris ordinarily has two needles in each sheath, but trifoliolate sheaths occur on the stems and stronger branches, where they prefer, as a rule, the upper parts of the single annual shoots. Camellia japonica is often striped in the fall and during the winter, but when flowering in the spring it returns to the monochromatic type.
Peloric flowers are terminal in some cases, but occur in the lower parts of the flower-spikes in others. Some varieties of gladiolus commence on each spike with more or less double flowers, which, higher up, are replaced by single ones. A wide range of bulbs and perennial garden-plants develop their varietal characters only partly when grown from seed and flowering for the first time. The annual garden-forget-me-not of the Azores (Myosotis azorica) has a variety with curiously enlarged flowers, often producing 20 or more corolla-segments in one flower. But this number gradually diminishes as the season advances. It would be quite superfluous to give further proof of the general validity of the law of periodicity in ever-sporting varieties.
[369]
LECTURE XIII
PISTILLODY IN POPPIES
One of the most curious anomalies that may be met with in ornamental garden-plants is the conversion of stamens into pistils. It is neither common nor rare, but in most cases the change is so slight comparatively that it is ordinarily overlooked. In the opium-poppy, on the contrary, it is very showy, and heightens the ornamental effect of the young fruits after the fading of the flowers. Here the central capsule is surrounded by a large crown of metamorphosed stamens.
This peculiarity has attracted the attention both of horticulturists and of botanists. As a rule not all the stamens are changed in this way but only those of the innermost rows. The outer stamens remain normal and fertile, and the flowers, when pollinated with their own pollen, bear as rich a harvest of seeds as other opium-poppies. The change affects both the filament and the anther, the former of which is dilated into a sheath. Within this sheath perfect [370] and more or less numerous ovules may be produced. The anthers become rudimentary and in their place broad leafy flaps are developed, which protrude laterally from the tip and constitute the stigmas. Ordinarily these altered organs are sterile, but in some instances a very small quantity of seed is produced, and when testing their viability I succeeded in raising a few plants from them.
The same anomaly occurs in other plants. The common wall-flower (Cheiranthus Cheiri) and the houseleek (Sempervivum tectorum) are the best known instances. Both have repeatedly been described by various investigators. In compiling the literature of this subject it is very interesting to observe the two contrasting views respecting the nature of this anomaly. Some writers, and among them Masters in his “Vegetable Teratology” consider the deviations to be merely accidental. According to them some species are more subject to this anomaly than others, and the houseleek is said to be very prone to this change. Goeppert, Hofmeister and others occasionally found the pistilloid poppies in fields or gardens, and sowed their seeds in order to ascertain whether the accidental peculiarity was inheritable or not. On the other hand De Candolle in his “Prodromus” mentions the pistilloid wall-flowers as a distinct [371] variety, under the name of Cheiranthus Cheiri gynantherus, and the analogous form of the opium-poppy is not at all an accidental anomaly, but an old true horticultural variety, which can be bought everywhere under the names of Papaver somniferum monstruosum or polycephalum. Since it is an annual plant, only the seeds are for sale, and this at once gives a sufficient proof of its heredity. In all cases, where it was met with accidentally by botanists, it is to be assumed that stray seeds had been casually mixed with those of other varieties, or that the habit had been transmitted by a spontaneous cross.
Wherever opportunity led to experiments on heredity, distinct races were found to be in possession of this quality, while others were not. It is of no use to cultivate large numbers of wall-flowers in the hope of one day seeing the anomaly arise; the only means is to secure the strain from those who have got it. With poppies the various varieties are so often intercrossed by bees, that the appearance of an accidental change may sometimes be produced, and in the houseleek the pistilloid warily seems to be the ordinary one, the normal strain being very rare or perhaps wholly wanting.
Our three illustrative examples are good and permanent races, producing their peculiar qualities [372] regularly and abundantly. In this respect they are however very variable and dependent on external circumstances. Such a regularity is not met with in other instances. Often pedigree-experiments lead to poor races, betraying their tendency to deviate only from time to time and in rare cases. Such instances constitute what we have called in a former lecture, “half races,” and their occurrence indicates that the casual observation of an anomaly is not in itself adequate to give an opinion as to the chance of repetition in sowing experiments. A large number of species seem to belong to this case, and their names may be found in the above mentioned work by Masters and elsewhere. But no effort has yet been made to separate thoroughly the pistilloid half-races from the corresponding ever-sporting varieties. Some plants are recorded as being more liable to this peculiarity than others.
Stamens are sometimes replaced by open carpels with naked ovules arising from their edges and even from their whole inner surfaces. This may be seen in distinct strains of the cultivated bulbous Begonia, and more rarely in primroses. Here the apex of the carpellary leaf is sometimes drawn out into a long style, terminated by a flattened spatulate stigma.
The pistillody of the stamens is frequently [373] combined with another deviation in the poppies. This is the growing together of some of the altered stamens so as to constitute smaller or larger connate groups. Often two are united, sometimes three, four or more. Flowers with numerous altered stamens are seldom wholly free from this most undesirable secondary anomaly. I call it undesirable with respect to experiments on the variability of the character. For it may easily be seen that while it is feasible to count the stamens even when converted into pistils, it is not possible when groups of them are more or less intimately united into single bodies. This combination makes all enumeration difficult and inaccurate and often wholly unreliable. In such cases the observation is limited to a computation of the degree of the change, rather than to a strict numerical inquiry. Happily the responses to the experimental influences are so marked and distinct that even this method of describing them has proved to be wholly sufficient.
In extreme instances I have seen all the changed stamens of a flower of the opium-poppy united into a single body, so as to form a close sheath all around the central ovary. Lesser sheaths, surrounding one-half or one-third of the capsule are of course less rarely met with. Leaving this description of the outer appearance [374] of our anomaly, we may now consider it from the double point of view of inheritance and variability.
The fact of inheritance is shown by the experience of many authors, and by the circumstance already quoted, that the variety has been propagated from seed for more than half a century, and may be obtained from various seed merchants. In respect to the variability, the variety belongs to the ever-sporting group, constituting a type which is more closely related to the “five-leaved” clover than to the striped flowers or even the double stocks.
It fluctuates around an average type with half filled crowns, going as far as possible in both directions, but never transgressing either limit. It is even doubtful whether the presumable limits are, under ordinary circumstances, ever reached. Obviously one extreme would be the conversion of all the stamens, and the other the absolute deficiency of any marked tendency to such a change. Both may occur, and will probably be met with from time to time. But they must be extremely rare, since in my own extensive experiments, which were strictly controlled, I never was able to find a single instance of either of them. Some of the outer stamens have always remained unchanged, yielding enough pollen for the artificial pollination of [375] the central ovary, and on the other hand some rudiments of hardened filaments were always left, even if they were reduced to small protuberances on the thalamus of the flower. Between these extremes all grades occur. From single, partially or wholly changed stamens upwards to 150 and over, all steps may be seen. It is a true fluctuating variability. There is an average of between 50 and 100, constituting a nearly filled crown around the central capsule. Around this average the smaller deviations are most numerous and the larger ones more rare. The inspection of any bed of the variety suffices to show that, taken broadly, the ordinary laws of fluctuating variability are applicable. No counting of the single individuals is required to dispel all doubts on this point.
Moreover all intermediate steps respecting the conversion of the single stamens may nearly always be seen. Rarely all are changed into normal secondary ovaries with a stigma and with a cavity filled with ovules. Often the stigma is incomplete or even almost wanting, in other instances the ovules are lacking or the cavity itself is only partially developed. Not rarely some stamens are reduced and converted into thin hard stalks, without any appearance of an ovary at their tip. But then the demarcation [376] between them and the thalamus fails, so that they cannot be thrown off when the flower fades away, but remain as small stumps around the base of the more fully converted filaments. This fact would frequently render the enumeration of the altered organs quite unreliable.
For these reasons I have chosen a group of arbitrary stages in order to express the degree of deviation for a given lot of plants. The limits were chosen so as to be sufficiently trustworthy and easy to ascertain. In each group the members could be counted, and a series of figures was reached by this means which allowed of a further comparison of the competing sets of plants.
It should be stated that in such experiments and especially in the case of such a showy criterion as the pistilloid heads afford after the time of flowering is over, the inspection of the controlling beds at once indicates the result of the experiment. Even a hasty survey is in most cases sufficient to get a definite conclusion. Where this is not the case, the counting of the individuals of the various groups often does not add to the evidence, and the result remains uncertain. On the other hand, the impression made by the groups of plants on the experimenter and on his casual visitors, cannot well be conveyed to the readers of his account by [377] other means than by figures. For this reason the result of the experiments is expressed in this way.
I made six groups. The first includes the cases where the whole circle is reduced to small rudiments. The second shows 1-10 secondary capsules. The two following constitute half a crown around the central fruit, the third going up to this limit, the fourth going from this limit to a nearly filled circle. Wholly filled circles of secondary capsules without gaps give the two last degrees, the fifth requiring only continuity of the circle, the sixth displaying a large and bright crown all around the central head. The fifth group ordinarily includes from 90-100 altered stamens, while the sixth has from 100-150 of these deviating parts.
In ordinary cultures the third and fourth group, with their interrupted crowns, predominate. Large crowns are rare and flowers which at first sight seem to be wholly normal, occur only under circumstances definitely known to be unfavorable to growth, and to the development of the anomaly.
Having reached by this means a very simple and easy method of stating the facts shown by equal lots under contrasting influences, we will now make use of it to inquire into the relation [378] of this exceptionally high degree of variability to the inner and outer conditions of life.
As a rule, all experiments show the existence of such a relation. Unfavorable conditions reduce the numbers of altered stamens, favorable circumstances raise it to its highest point. This holds true for lots including hundreds of specimens, but also for the sundry heads of one bed, and often for one single plant.
We may compare the terminal flower with those of the lateral branches on a plant, and when no special influences disturb the experiment, the terminal head ordinarily bears the richest crown. If the first has more than 100 metamorphosed parts, the latter have often less than 50 on the same plant. In poor soil, terminal heads are often reduced to 10-20 monstrous organs, and in such cases I found the lateral flowers of the same plants ordinarily with less than 10 altered stamens. In some cases I allowed the branches of the third and fourth degree, in other words, the side twigs of the first branches of my selected plants to grow out and produce flowers in the fall. They were ordinarily weak, sometimes very small, having only 5-9 stigmas on their central fruit. Secondary capsules were not seen on such flowers, even when the experiment was repeated on a [379] somewhat larger scale and during a series of years.
Among the same lot of plants individual differences almost always occur. They are partly due to inequalities already existing in the seeds, and partly to the diversity of the various parts of the same bed. Some of the plants become stout and have large terminal heads. Others remain very weak, with a slender stem, small leaves and undersized flowers. The height and thickness of the stem, the growth of the foliage and of the axillary buds are the most obvious measures of the individual strength of the plant. The development of the terminal flower and the size of its ovary manifestly depends largely on this individual strength, as may be seen at once by the inspection of any bed of opium-poppies. Now this size of the head can easily be measured, either by its height or circumference, or by its weight. Moreover we can arrange them into a series according to their size. If we do this with the polycephalous variety, the relation between individual strength and degree of metamorphosis at once becomes manifest. The largest heads have the brightest crowns, and the number of supernumerary carpels diminishes in nearly exact proportion to the size of the fruits. Fruits with less than 50 altered stamens weighed on an average 5 grams, [380] those with 50-100 such organs 7 grams and those with a bright crown 10 grams, the appendices being removed before the weighing. Corresponding results have been reached by the comparison of the height of the capsules with their abnormal surroundings. The degree of development of the monstrosity is shown by this observation to be directly dependent on, and in a sense proportionate to the individual strength of the plant.
The differences between the specimens grown from a single lot of seeds, for instance from the seeds of one self-fertilized capsule are, as I have said, partly due to the divergences which are always present in a bed, even if the utmost care has been taken to make it as uniform as possible. These local differences are ordinarily underrated and overlooked, and it is often considered to be sufficient to cultivate small lots of plants under apparently similar conditions on neighboring beds, to be justified in imputing all the observed deviations of the plants to hereditary inequalities. This of course is true for large lots, whenever the averages only are compared. In smaller experiments the external conditions of the single individuals should always be considered carefully. Lots of one or two square meters suffice for such comparisons, but smaller lots are always subject to chances and [381] possibilities, which should never be left out of consideration.
Therefore I will now point out some circumstances, which are ordinarily different on various parts of one and the same bed.
In the first place comes the inequality of the seeds themselves. Some of them will germinate earlier and others later. Those that display their cotyledons on a sunny day will be able to begin at once with the production of organic food. Others appear in bad weather, and will thus be retarded in their development. These effects are of a cumulative nature as the young plants must profit by every hour of sunshine, according to the size of the cotyledons. Any inequality between two young seedlings is apt to be increased by this cumulative effect.
The same holds good for the soil of the bed. It is simply impossible to mix the manure so equally that all individuals receive the same amount of it from the very beginning. I am in the habit of using manures in a dry and pulverized condition, of giving definite quantities to each square meter, and of taking the utmost care to get equal distribution and mixture with the soil, always being present myself during this most important operation. Nevertheless it is impossible to make the nourishment exactly equal for all the plants of even a small bed.
[382] Any inequality from this cause will increase the difference in the size of the young leaves, augment the inequality of their production of organic matter and for this reason go on in an ever increasing rate.
Rain and spraying, or on the other hand dryness of the soil, have still greater consequences. The slightest unevenness of the surface will cause some spots to dry rapidly and others to retain moisture during hours and even sometimes during days.
Seeds, germinating in such little moist depressions grow regularly and rapidly, while those on the dryer elevations may be retarded for hours and days, before fully unfurling their seed-leaves. After heavy rains these differences may be observed to increase continually, and in some instances I found that plants were produced only on the wet spots, while the dry places remained perfectly bare. From this the wet spots seem to be the most favorable, but on the other hand, seeds may come to germinate there too numerously and so closely that the young plants will be crowded together and find neither space nor light enough, for a free and perfect development. The advantage may change to disadvantage in this way unless the superfluous individuals are weeded out in due time.
[383] From all these and other reasons some plants will be favored by the external conditions from the beginning, while others will be retarded, and the effects will gradually increase until at last they become sufficient to account for a considerable amount of individual variability. There is no doubt that the difference in the strength of the plant and in the size of the capsules, going from 5-10 grams for a single fruit, are for the most part due to these unavoidable circumstances. I have tried all conceivable means to find remedies for these difficulties, but only by sowing my seeds in pans in a glass-house have I been able to reach more constant and equal conditions. But unfortunately such a method requires the planting out of the young seedlings in the beginning of the summer, and this operation is not without danger for opium-poppies, and especially not without important influence on the monstrosity of the pistilloid variety. Consequently my sowings of this plant have nearly always been made in the beds.
In order to show how great the influence of all these little things may become, we only have to make two sowings on neighboring beds and under conditions which have carefully been made as equal as possible. If we use for these controlling experiments seeds from one and the same capsule, it will soon become evident that [384] no exact similarity between the two lots may be expected. Such differences as may be seen in these cases are therefore never to be considered of value when comparing two lots of seeds of different origin, or under varying conditions. No amount of accuracy in the estimation of the results of a trial, or in the counting out of the several degrees of the anomaly, is adequate to overcome the inaccuracy resulting from these differences.
It is certainly of great importance to have a correct conception in regard to the influence of the surrounding conditions on the growth of a plant and on the development of the attribute we are to deal with. No less important is the question of the sensibility of the plants to these factors. Obviously this sensibility must not be expected to remain the same during the entire life-period, and periods of stronger and of weaker responses may be discerned.
In the first place it is evident that external or inner influences are able to change the direction of the development of an organ only so long as this development is not yet fully finished. In the young flower-bud of the pistilloid poppy there must evidently be some moment in which it is definitely decided whether the young stamens will grow out normally or become metamorphosed into secondary pistils. From this [385] moment no further change of external conditions is able to produce a corresponding change in the degree of the anomaly. The individual strength of the whole plant may still be affected in a more or less manifest degree, but the number of converted stamens of the flower has been definitely fixed. The sensitive period has terminated.
In order to determine the exact moment of this termination of the period of sensibility, I have followed the development of the flower buds during the first weeks of the life of the young plants. The terminal flower may already be seen in young plants only seven weeks old, with a stem not exceeding 5-6 cm. in height and a flower-bud with a diameter of nearly 1 mm., in which the stamens and secondary pistils are already discernible, but still in the condition of small rounded protuberances on the thalamus. Though it is not possible at that time to observe any difference between the future normal and converted stamens, it does not seem doubtful that the development is so far advanced, that in the inner tissues the decision has already definitely been taken. In the next few days this decision rapidly becomes visible, and the different parts of the normal stamens and the metamorphosed carpels soon become apparent. From this observation it [386] can be inferred that the sensitive period of the anomaly is limited for the terminal flower-head, to the first few weeks of the life of the young plants. The secondary heads manifestly leave this period at a somewhat later stage.
In order to prove the accuracy of this conclusion I have tried to injure the anomalies after the expiration of the first six or seven weeks. I deprived them of their leaves, and damaged them in different ways. I succeeded in making them very weak and slender, without being able to diminish the number of the supernumerary carpels. The proportionality of the size of the central fruit and the development of the surrounding crown can often be modified or even destroyed by this means, and the apparent exceptions from this rule, which are often observed, may find their explanation in this way.
In the second place I have tried to change the development of the anomaly during the period of sensibility, and even in the last part of it. This experiment succeeded fully when carried out within the fifth or sixth week after the beginning of the germination. As means of injury I transplanted the young plants. To this end I sowed my seeds in pans in unmanured soil, planted them out in little pots with richly prepared earth, grew them in these during a few weeks and afterwards transferred them to the [387] beds, taking care that the pats were removed, but the balls of earth not broken.
In consequence of this treatment the plants became very large and strong, with luxuriant foliage and relatively numerous large flowers and fruits. But almost without exception they were poor in anomalous stamens, at least so on the terminal heads. On a lot of some 70 plants more than 50 had less than half a crown of secondary capsules, while from the same packet of seed the control-plants gave in an equal number more than half of filled crowns on all plants with the exception of five weak specimens.
It is curious to compare such artificially injured plants with the ordinary cultures. Strong stems and heavy fruits, which otherwise are always indicative of showy crowns, now bear fruits wholly or nearly destitute of any anomalous change. The commonly prevailing rule seems to be reversed, showing thereby the possibility of abolishing the correlation between individual strength and anomaly by an artificial encroachment upon the normal conditions.
Aside from these considerations the experiments clearly give proof of the existence of a period of sensibility limited to the first weeks of the life of the plant for the terminal flower. This knowledge enables us to explain many apparent [388] parent abnormalities, which may occur in the experiments.
We now may take a broader view of the period of sensibility. Evidently the response to external influences will be greater the younger the organ. Sensibility will gradually diminish, and the phenomena observed in the last part of this period may be considered as the last remainder of a reaction which previously must have been much stronger and much readier, providing that it would be possible to isolate them from, and contrast them with, the other responses of the same plant.
With the light thus cast upon the question, we may conclude that the sensitive period commences not only at the beginning of the germination, but must also be considered to include the life of the seed itself. From the moment of fertilization and the formation of the young embryo the development must be subjected to the influence of external agencies which determine the direction it will take and the degree of development it will finally be able to acquire. Probably the time of growth of the embryo and of the ripening of the seed correspond exactly to the period of highest sensibility. This period is only interrupted during the resting stage of the seed, to be repeated in germination. Afterwards the sensibility [389] slowly and gradually decreases, to end with the definite decision of all further growth sometime before the outer form of the organ becomes visible under the microscope. The last period of life includes only an expansion of the tissues, which may still have some influence on their final size, but not on their form. This has been definitely arrested before the end of the sensitive period, and ordinarily before the commencement of that rapid development, which is usually designated by the name of growth, as contrasted with evolution.
Within the seed the evolution of the young plant manifestly depends upon the qualities and life-conditions of the parent-plant. The stronger this is, and the more favorable circumstances it is placed under, the more food will be available for the seed, and the healthier will be the development of the embryo. Only well-nourished plants give well-nourished seeds, and the qualities of each plant are for this reason at least, partly dependent on the properties of its parents and even of its grandparents.
From these considerations the inference is forced upon us that the apparently hereditary differences, which are observed to exist among the seeds of a species or a variety and even of a single strain or a single parent-plant, may for a large part, and perhaps wholly, be the result [390] of the life-conditions of their parents and grandparents. Within the race all ssvariability would in this way be reduced to the effects of external circumstances. Among these nourishment is no doubt the most momentous, and this to such a degree that older writers designated the external conditions by the term nourishment. According to Knight nutrition reigns supreme in the whole realm of variability, the kind of food and the method of nourishment coming into consideration only in a secondary way. The amount of useful nutrition is the all-important factor.
If this is so, and if nutrition decides the degree of deviation of any given character, the widest deviating individuals are the best nourished ones. The best nourished not only during the period of sensibility of the attribute under consideration, but also in the broadest sense of the word.
This discussion casts a curious light upon the whole question of selection. Not of course upon the choice of elementary species or varieties out of the original motley assembly which nature and old cultures offer us, but upon the selection of the best individuals for isolation and for the improvement of the race. These are, according to my views, only the best nourished ones. Their external conditions have been the [391] most favorable, not only from the beginning of their own life in the field, but also during their embryonic stages, and even during the preparation of these latter in the life of their parents and perhaps even their grandparents. Selection then, would only be the choice of the best nourished individuals.
In connection with the foregoing arguments I have tried to separate the choicest of the poppies with the largest crown of pistilloid stamens, from the most vigorous individuals. As we have already seen, these two attributes are as a rule proportional to one another. Exceptions occur, but they may be explained by some later changes in the external circumstances, as I have also pointed out. As a rule, these exceptions are large fruits with comparatively too few converted stamens; they are exactly the contrary from what is required for a selection. Or plants, which from the beginning were robust, may have become crowded together by further growth, and for these reasons become weaker than their congeners, though retaining the full development of the staminodal crown, which was fixed during the sensitive period and before the crowding. I have searched my beds yearly for several years in vain to find individuals which might recommend themselves for selection without having the stamp of permanent, [392] or at least temporarily better, nourishment. No starting-point for such an independent selection has ever been met with.
Summing up the consequences of this somewhat extended discussion, we may state it as a rule that a general proportion between the individual strength and the degree of development of the anomaly exists. And from this point of view it is easy to see that all external causes which are known to affect the one, must be expected to influence the other also.
It will therefore hardly be necessary to give a full description of all my experiments on the relations of the monstrosity to external conditions. A hasty survey will suffice.
This survey is not only intended to convey an idea of the relations of pistilloid poppies to their environment, but may serve as an example of the principle involved. According to my experience with a large range of other anomalies, the same rule prevails everywhere. And this rule is so simple that exact knowledge of one instance may be considered as sufficient to enable us to calculate from analogy what is to be expected from a given treatment of any other anomaly. Our appreciation of observed facts and the conditions to be chosen for intended cultures are largely dependent on such calculations. What I am now going to describe [393] is to be considered therefore as an experimental basis for such expectations.
First of all comes the question how many individuals are to be grown in a given place. When sowing plants for experimental purposes it is always best to sow in rows, and to give as few seeds to each row as possible, so as to insure all necessary space to the young plants. On the other hand the seeds do not all germinate, and after sowing too thinly, gaps may appear in the rows. This would cause not only a loss of space, but an inequality between the plants in later life, as those nearest the gaps would have more space and more light, and a larger area for their roots than those growing in the unbroken rows. Hence the necessity of using large quantities of seed and of weeding out a majority of young plants on the spots where the greatest numbers germinate.
Crowded cultures as a rule, will give weak plants with thin stems, mostly unbranched and bearing only small capsules. According to the rule, these will produce imperfect crowns of secondary pistils. The result of any culture will thus be dependent to a high degree on the number of individuals per square meter. I have sown two similar and neighboring beds with the thoroughly mixed seeds of parent-plants of the same strain and culture, using as much [394] as 2.5 cu. cm. per square meter. On one of the beds I left all the germinating plants untouched and nearly 500 of them flowered, but among them 360 were almost without pistillody, and only 10 had full crowns. In the other bed I weeded away more than half of the young plants, leaving only some 150 individuals and got 32 with a full crown, nearly 100 with half crowns and only 25 apparently without monstrosity.
These figures are very striking. From the same quantity of seed, in equal spaces, by similar exposure and treatment I got 10 fully developed instances in one and 32 in the other case. The weeding out of supernumerary individuals had not only increased the percentage of bright crowns, but also their absolute number per square meter. So the greatest number of anomalies upon a given space may be obtained by taking care that not too many plants are grown upon it: any increase of the number beyond a certain limit will diminish the probability of obtaining these structures. The most successful cultures may be made after the maximum number of individuals per unit of area has been determined. A control-experiment was made under the same conditions and with the same seed, but allowing much less for the same space. I sowed only 1 cu. cm. on my bed of 2 square meters, and thereby avoided [395] nearly all weeding out. I got 120 plants, and among them 30 with full crowns of converted stamens, practically the same number as after the weeding out in the first experiment. This shows that smaller quantities of seed give an equal chance for a greater number of large crowns, and should therefore always be preferred, as it saves both seed and labor.
Weeding out is a somewhat dangerous operation in a comparative trial. Any one who has done it often, knows that there is a strong propensity to root out the weaker plants and to spare the stronger ones. Obviously this is the best way for ordinary purposes, but for comparisons evidently one should not discriminate. This rule is very difficult in practice, and for this reason one should never sow more than is absolutely required to meet all requirements.
Our second point is the manuring of the soil. This is always of the highest importance, both for normal and for anomalous attributes. The conversion of the stamens into pistils is in a large measure dependent upon the conditions of the soil. I made a trial with some 800 flowering plants, using one sample of seed, but sowing one-third on richly manured soil, one-third on an unprepared bed of my garden, and one-third on nearly pure sand. In all other respects the three groups were treated in the same way. Of [396] the manured plants one-half gave full crowns, of the non-manured only one-fifth, and on the sandy soil a still smaller proportion. Other trials led to the same results. I have often made use of steamed and ground horn, which is a manure very rich in nitrogenous substances. One-eighth of a kilo per square meter is an ample amount. And its effect was to increase the number of full crowns to an exceptional degree.
In the controlling trial and under ordinary circumstances this figure reached some 50%, but with ground horn it came up as high as 90%. We may state this result by the very striking assertion that the number of large crowns in a given culture may be nearly doubled by rich manure.
All other external conditions act in a similar manner. The best treatment is required to attain the best result. A sunny exposure is one of the most essential requisites, and in some attempts to cultivate my poppies in the shade, I found the pistillody strongly reduced, not a single full crown being found in the whole lot. Often the weather may be hurtful, especially during the earlier stages of the plants. I protected my beds during several trials by covering them with glass for a few weeks, until the young plants reached the glass covering. I got a normal number of full crowns, some 55%, at a time [397] when the weather was so bad as to reduce the number in the control experiments to 10%.
It would be quite superfluous to give more details or to describe additional experiments. Suffice to say, that the results all point in the same direction, and that pistillody of the poppies always clearly responds to the treatment, especially to external conditions during the first few weeks, that is, during the period of sensitiveness. The healthier and the stronger the plants the more fully they will develop their anomaly.
In conclusion something is to be said about the choice of the seed. Obviously it is possible to compare seeds of different origin by sowing and treating them in the same way, giving attention to all the points above mentioned. In doing so the first question will be, whether there is a difference between the seeds of strong plants with a bright crown around the head and those of weaker individuals with lesser development of the anomaly. It is evident that such a difference must be expected, since the nutrition of the seed takes place during the period of the greatest sensitiveness.
But the experiments will show whether this effect holds good against the influences which tend to change the direction of the development of the anomaly during the time of germination. [398] The result of my attempt has shown that the choice of the seeds has a manifest influence upon the ultimate development of the monstrosity, but that this influence is not strong enough to overwhelm all other factors.
The choice of the fullest or smallest crowns may be repeated during succeeding generations, and each time compared with a culture under average conditions. By this means we come to true selection-experiments, and these result in a notable and rapid change of the whole strain. By selecting the brightest crowns I have come up in three years from 40 to 90 and ultimately to 120 converted stamens in the best flower of my culture, and in selecting the smallest crowns I was able in three years to exclude nearly all good crowns, and to make cultures in which heads with less than half-filled crowns constituted the majority. But such selected strains always remain very sensitive to treatment, and by changing the conditions the effect may be wholly lost in a single year, or even turned in the contrary direction. In other words, the anomaly is more dependent on external conditions during the germinating period than on the choice of the seeds, providing these belong to the pistilloid variety and have not deteriorated by some crossing with other sorts.
At the beginning of this lecture I stated that [399] no selection is adequate to produce either a pure strain of brightly crowned flower-heads without atavism, or to conduce to an absolute and permanent loss of the anomaly. During a series of years I have tested my plants in both directions, but without the least effect. Limits are soon reached on both sides, and to transgress these seems quite impossible.
Taking these limits as the marks of the variety, and considering all fluctuations between them as responses to external influences working during the life of the individual or governing the ripening of the seeds, we get a clear picture of a permanent ever-sporting type. The limits are absolutely permanent during the whole existence of this already old variety. They never change. But they include so wide a range of variability, that the extremes may be said to sport into one another, so much the more so as one of the extremes is to be considered morphologically as the type of the variation, while the other extreme can hardly be distinguished from the normal form of the species.
[400]
LECTURE XIV
MONSTROSITIES
I have previously dealt with the question of the hereditary tendencies that cause monstrosities. These tendencies are not always identical for the same anomaly. Two different types may generally, be distinguished. One of them constitutes a poor variety, the other a rich one. But this latter is abundant and the first one is poor in instances of exactly the same conformation. Therefore the difference only lies in the frequency of the anomaly, and not in its visible features. In discovering an instance of any anomaly it is therefore impossible to tell whether it belongs to a poor or to a rich race. This important question can only be answered by direct sowing-experiments to determine the degree of heredity.
Monstrosities are often considered as accidents, and rightfully so, at least as long as they are considered from a morphological point of view. Physiology of course excludes all accidentality. And in our present ease it shows [401] that some internal hereditary quality is present, though often latent, and that the observed anomalies are to be regarded as responses of this innate tendency to external conditions. Our two types differ in the frequency of these responses. Rare in the poor race, they are numerous in the rich variety. The external conditions being the same for both, the hereditary factor must be different. The tendency is weak in the one and strong in the other. In both cases, according to my experience, it may be weakened or strengthened by selection and by treatment. Often to a very remarkable degree, but not so far as to transgress the limits between the two races. Such transgression may apparently be met with from time to time, but then the next generation generally shows the fallacy of the conclusion, as it returns more or less directly to the type from which the strain had been derived. Monstrosities should always be studied by physiologists from this point of view. Poor and rich strains of the same anomaly seem at first sight to be so nearly allied that it might be thought to be very easy to change the one into the other. Nevertheless such changes are not on record, and although I have made several attempts in this line, I never succeeded in passing the limit. I am quite convinced that sometime [402] a method will be discovered of arbitrarily producing such conversions, and perhaps the easiest way to attain artificial mutations may lie concealed here. But as yet not the slightest indication of this possibility is to be found, save the fallacious conclusions drawn from too superficial observations.
Unfortunately the poor strains are not very interesting. Their chance of producing beautiful instances of the anomaly for which they are cultivated is too small. Exceptions to this rule are only afforded by those curious and rare anomalies, which command general attention, and of which, therefore, instances are always welcome. In such cases they are searched for with perseverance, and the fact of their rarity impresses itself strongly on our mind.
Twisted stems are selected as a first example. This monstrosity, called biastrepsis, consists of strongly marked torsions as are seen in many species with decussate leaves, though as a rule it is very rare. Two instances are the most generally known, those of the wild valerian (Valeriana officinalis) and those of cultivated and wild sorts of teasels (Dipsacus fullonum, D. sylvestris, and others). Both of these I have cultivated during upwards of fifteen years, but with contradictory results. The valerian is a perennial herb, multiplying itself yearly by [403] slender rootstocks or runners producing at their tips new rosettes of leaves and in the center of these the flowering stem. My original plant has since been propagated in this manner, and during several years I preserved large beds with hundreds of stems, in others I was compelled to keep my culture within more restricted limits. This plant has produced twisted stems of the curious shape, with a nearly straight flag of leaves on one side, described by De Candolle and other observers, nearly every year. But only one or two instances of abnormal stems occurred in each year, and no treatment has been found that proved adequate to increase this number in any appreciable manner. I have sown the seeds of this plant repeatedly, either from normal or from twisted stems, but without better results. It was highly desirable to be able to offer instances of this rare and interesting peculiarity to other universities and museums, but no improvement of the race could be reached and I have been constrained to give it up. My twisted valerian is a poor race, and hardly anything can be done with it. Perhaps, in other countries the corresponding rich race may be hidden somewhere, but I have never had the good fortune of finding it.
This good fortune however, I did have with the wild teasel or Dipsacus sylvestris. [404] Stems of this and of allied species are often met with and have been described by several writers, but they were always considered as accidents and nobody had ever tried to cultivate them. In the summer of 1885 I saw among a lot of normal wild teasels, two nicely twisted stems in the botanical garden of Amsterdam. I at once proposed to ascertain whether they would yield a hereditary race and had all the normal individuals thrown away before the flowering time. My two plants flowered in this isolated condition and were richly pollinated by insects. Of course, at that time, I knew nothing of the dependency of monstrosities on external conditions, and made the mistake of sowing the seeds and cultivating the next generation in too great numbers on a small space. But nevertheless the anomaly was repeated, and the aberrant individuals were once more isolated before flowering. The third generation repeated the second, but produced sixty twisted stems on some 1,600 individuals. The result was very striking and quite sufficient for all further researches, but the normal condition of the race was not reached. This was the case only after I had discovered the bad effects of growing too many plants in a limited space. In the fourth generation I restricted my whole culture to about 100 individuals, and by this simple [405] means at once got up to 34% of twisted stems. This proportion has since remained practically the same. I have selected and isolated my plants during five succeeding generations, but without any further result, the percentage of twisted stems fluctuating between 30 and about 45 according to the size of the cultures and the favorableness or unfavorableness of the weather.
It is very interesting to note that all depends on the question whether one has the good fortune of finding a rich race or not, as this pedigree-culture shows. Afterwards everything depends on treatment and very little on selection. As soon as the treatment becomes adequate, the full strength of the race at once displays itself, but afterwards no selection is able to improve it to any appreciable amount. Of course, in the long run, the responses will be the same as those of the pistilloid poppies on the average, and some influence of selection will show itself on closer scrutiny.
Compared with the polycephalous poppies my race of twisted teasels is much richer in atavists. They are never absent, and always constitute a large part of each generation and each bed, comprising somewhat more than half of the individuals. Intermediate stages between them and the wholly twisted stems are not wanting, [406] and a whole series of steps may easily be observed from sufficiently large cultures. But they are always relatively rare, and any lot of plants conveys the idea of a dimorphous race, the small twisted stems contrasting strongly with the tall straight ones.
A sharper contrast between good representatives of a race and their atavists is perhaps to be seen in no other instance. All the details contribute to the differentiation in appearance. The whole stature of the plants is affected by the varietal mark. The atavists are not, as in the case of the poppies, obviously allied with the type by a full range of intermediate steps, but quite distant from it by their rarity. There seems to be a gap in the same way as between the striped flowers of the snapdragon and their uniform red atavists, while with the poppies the atavists may be viewed as being only the extremes of a series of variations fluctuating around some average type.
From this reason it is as interesting to appreciate the hereditary position of the atavists of twisted varieties as it was for the red-flowered descendants of the striped flowers. In order to ascertain this relation it is only necessary to isolate some of them during the blooming-period. I made this experiment in the summer of 1900 with the eighth generation of my race, and contrived [407] to isolate three groups of plants by the use of parchment bags, covering them alternately, so the flowers of only one group were accessible to insects, at a time. I made three groups, because the atavists show two different types. Some specimens have decussate stems, others bear all their leaves in whorls of three, but in respect to the hereditary tendency of the twisting character this difference does not seem to be of any importance.
In this way I got three lots of seeds and sowed enough of them to have three groups of plants each containing about 150-200 well developed stems. Among these I counted the twisted individuals, and found nearly the same numbers for all three. The twisted parents gave as many as 41% twisted children, but the decussate atavists gave even somewhat more, viz., 44%, while the ternate specimens gave 37%. Obviously the divergences between these figures are too slight to be dwelt upon, but the fact that the atavists are as true or nearly as true inheritors of the twisted race as the best selected individuals is clearly proved by this experience.
It is evident that here we have a double race, including two types, which may be combined in different degrees. These combinations determine a wide range of changes in the stature of the plants, and it seems hardly right to use the [408] same term for such changes as for common variations. It is more a contention of opposite characters than a true phenomenon of simple variability. Or perhaps we might say that it is the effect of the cooperation of a very variable mark, the twisting, with a scarcely varying attribute of the normal structure of the stem. Between the two types an endless diversity prevails, but outwardly there are limits which are never transgressed. The double race is as permanent, and in this sense as constant, as any ordinary simple variety, both in external form, and in its intimate hereditary qualities.
I have succeeded in discovering some other rich races of twisted plants. One of them is the Sweet William (Dianthus barbatus), which yielded, after isolation, in the second generation, 25% of individuals with twisted stems, and as each individual produces often 10 and more stems, I had a harvest of more than half a thousand of instances of this curious, and ordinarily very rare anomaly. My other race is a twisted variety of Viscaria oculata, which is still in cultivation, as it has the very consistent quality of being an annual. It yielded last summer (1903) as high a percentage as 65 of twisted individuals, many of them repeating the monstrosity on several branches. After some occasional observations Gypsophila paniculata [409] seems to promise similar results. On the other hand I have sowed in vain the seeds of twisted specimens of the soapwort and the cleavewort (Saponaria officinalis and Galium Aparine). These and some others seem to belong to the same group as the valerian and to constitute only poor or so-called half-races.
Next to the torsions come the fasciated stems. This is one of the most common of all malformations, and consists, in its ordinary form, of a flat ribbon-like expansion of the stems or branches. Below they are cylindrical, but they gradually lose this form and assume a flattened condition. Sometimes the rate of growth is unequal on different portions or on the opposite sides of the ribbon, and curvatures are produced and these often give to the fasciation a form that might be compared with a shepherd’s crook. It is a common thing for fasciated branches and stems to divide at the summit into a number of subdivisions, and ordinarily this splitting occurs in the lower part, sometimes dividing the entire fasciated portion. In biennial species the rosette of the root-leaves of the first year may become changed by the monstrosity, the heart stretching in a transverse direction so as to become linear. In the next year this line becomes the base from which the stem grows. In such cases the fasciated stems [410] are broadened and flattened from the very beginning, and often retain the incipient breadth throughout their further development. Species of primroses (Primula japonica and others), of buttercups (Ranunculus bulbosus), the rough hawksbeard (Crepis biennis), the Aster Tripolium, and many others could be given as instances.
Some of these are so rare as to be considered as poor races, and in cultural trials do not produce the anomaly except in a very few instances. Heads of rye are found in a cleft condition from time to time, single at their base and double at the top, but this anomaly is only exceptionally repeated from seed. Flattened stems of Rubia tinctorum are not unfrequently met with on the fields, but they seem to have as little hereditary tendency as the split rye (Secale Cereale). Many other instances could be given. Both in the native localities and in pedigree-cultures such ribboned stems are only seen from time to time, in successive years, in annual and biennial as well as in perennial species. The purple pedicularis (Pedicularis palustris) in the wild state, and the sunflower among cultivated plants, may be cited instead of giving a long list of analogous instances.
On the other hand rich races of flattened stems are not entirely lacking. They easily betray [411] themselves by the frequency of the anomaly, and therefore may be found, and tried in the garden. Under adequate cultivation they are here as rich in aberrant individuals as the twisted races quoted above, producing in good years from 30-40% and often more instances. I have cultivated such rich races of the dandelion (Taraxacum officinale), of Thrincia hirta, of the dame’s violet (Hesperis matronalis), of the hawkweed (Picris hieracioides), of the rough hawksbeard (Crepis biennis), and others.
Respecting the hereditary tendencies these rich varieties with flattened stems may be put in the same category with the twisted races. Two points however, seem to be of especial interest and to deserve a separate treatment.
The common cockscomb or Celosia cristata, one of the oldest and most widely cultivated fasciated varieties may be used to illustrate the first point. In beds it is often to be seen in quite uniform lots of large and beautiful crests, but this uniformity is only secured by careful culture and selection of the best individuals. In experimental trials such selection must be avoided, and in doing so a wide range of variability at once shows itself. Tall, branched stems with fan-shaped tops arise, constituting a series of steps towards complete atavism. This last [412] however, is not to be reached easily. It often requires several successive generations grown from seed collected from the most atavistic specimens. And even such selected strains are always reverting to the crested type. There is no transgression, no springing over into a purely atavistic form, such as may be supposed to have once been the ancestor of the present cockscomb. The variety includes crests and atavists, and may be perpetuated from both. Obviously every gardener would select the seeds of the brightest crests, but with care the full crests may be recovered, even from the worst reversionists in two or three generations. It is a double race of quite the same constitution as the twisted teasels.
My second point is a direct proof of this assertion, but made with a fasciated variety of a wild species. I took for my experiment the rough hawksbeard. In the summer of 1895 I isolated some atavists of the fifth generation of my race, which, by ordinary selection, gave in the average from 20-40% of fasciated stems. My isolated atavists bore abundant fruit, and from these I had the next year a set of some 350 plants, out of which about 20% had broadened and linear rosettes. This proportion corresponds with the degree of inheritance which is shown in many years by the largest and strongest [413] fasciated stems. It strengthens our conclusion as to the innermost constitution of the double races or ever-sporting varieties.
Twisted stems and fasciations are very striking monstrosities. But they are not very good for further investigation. They require too much space and too much care. The calculation of a single percentage requires the counting of some hundreds of individuals, taking many square meters for their cultivation, and this, as my best races are biennial, during two years. For this reason the countings must always be very limited, and selection is restrained to the most perfect specimens.
Now the question arises, whether this mark is the best upon which to found selection. This seems to be quite doubtful. In the experiments on the heredity of the atavists, we have seen that they are, at least often, in no manner inferior to even the best inheritors of the race. This suggests the idea that it is not at all certain that the visible characters of a given individual are a trustworthy measure of its value as to the transmission of the same character to the offspring. In other words, we are confronted with the existence of two widely different groups of characters in estimating the hereditary tendency. One is the visible quality of the individuals and the other is the direct observation [414] of the degree in which the attribute is transmitted. These are by no means parallel, and seem in some sense to be nearly independent of each other. The fact that the worst atavists may have the highest percentage of varietal units seems to leave no room for another explanation.
Developing this line of thought, we gradually arrive at the conclusion that the visible attribute of a varying individual is perhaps the most untrustworthy and the most unreliable character for selection, even if it seems in many cases practically to be the only available one. The direct determination of the degree of heredity itself is obviously preferable by far. This degree is expressed by the proportion of its inheritors among the offspring, and this figure therefore should be elevated to the highest rank, as a measure of the hereditary qualities. Henceforward we will designate it by the name of hereditary percentage.
In scientific experiments this figure must be determined for every plant of a pedigree-culture singly, and the selection should be founded exclusively or at least mainly on it. It is easily seen that this method requires large numbers of individuals to be grown and counted. Some two or three hundred progeny of one plant are needed to give the decisive figure for this one [415] individual, and selection requires the comparison of at least fifty or more individuals. This brings the total amount of specimens to be counted up to some tens of thousands. In practice, where important interests depend upon the experiments, such numbers are usually employed and often exceeded, but for the culture of monstrosities, other methods are to be sought in order to avoid these difficulties.
The idea suggests itself here that the younger the plants are, when showing their distinguishing marks, the more of them may be grown on a small space. Hence the best way is to choose such attributes, as may already be seen in the young seedlings, in the very first few weeks of their lives. Fortunately the seed-leaves themselves afford such distinctive marks, and by this means the plants may be counted in the pans, requiring no culture at all in the garden. Only the selected individuals need be grown to ripen their seeds, and the whole selection may be made in the spring, in the glasshouse. Instead of being very troublesome, the determination of the hereditary percentages becomes a definite reduction of the size of the experiments. Moreover it may easily be effected by any one who cares for experimental studies, but has not the means required for cultures on a larger scale. And lastly, there are [416] a number of questions about heredity, periodicity, dependency on nourishment and other life conditions, and even about hybridizing, which may be answered by this new method.
Seed-leaves show many deviations from the ordinary shape, especially in dicotyledonous plants. A very common aberration is the multiplication of their number, and three seed-leaves in a whorl are not rarely met with. The whorl may even consist of four, and in rare cases of five or more cotyledons. Cleft cotyledons are also to be met with, and the fissure may extend varying distances from the tips. Often all these deviations may be seen among the seedlings of one lot, and then it is obvious that together they constitute a scale of cleavages, the ternate and quaternate whorls being only cases where the cleaving has reached its greatest development. All in all it is manifest that here we are met by one type of monstrosity, but that this type allows of a wide range of fluctuating variability. For brevity’s sake all these cleft and ternate, double cleft and quaternate cotyledons and even the higher grades are combined under one common name and indicated as tricotyls.
A second aberration of young seed-plants is exactly opposite to this. It consists of the union of the two seed-leaves into a single organ. This ordinarily betrays its origin by [417] having two separate apices, but not always. Such seedlings are called syncotyledonous or syncotyls. Other monstrosities have been observed from time to time, but need not be mentioned here.
It is evident that the determination of the hereditary percentage is very easy in tricotylous or syncotylous cultures. The parent plants must be carefully isolated while blooming. Many species pollinate themselves in the absence of bees; from these the insects are to be excluded. Others have the stamens and stigmas widely separated and have to be pollinated artificially. Still others do not lend themselves to such operations, but have to be left free to the visits of bees and of humble-bees, this being the only means of securing seed from every plant. At the time of the harvest the seeds should be gathered separately from each plant, and this precaution should also be observed in studies of the hereditary percentage at large, and in all scientific pedigree-cultures. Every lot of seeds is to be sown in a separate pan, and care must be taken to sow such quantities the three to four hundred seedlings will arise from each. As soon as they display their cotyledons, they are counted, and the number is the criterion of the parent-plant. Only parent-plants with the highest percentages are selected, and out of [418] their seedlings some fifty or a hundred of the best ones are chosen to furnish the seeds for the next generation.
This description of the method shows that the selection is a double one. The first feature is the hereditary percentage. But then not all the seedlings of the selected parents can be planted out, and a choice has to be made. This second selection may favor the finest tricotyls, or the strongest individuals, or rely on some other character, but is unavoidable.
We now come to the description of the cultures. Starting points are the stray tricotyls which are occasionally found in ordinary sowings. In order to increase the chance of finding them, thousands of seeds of the same species must be inspected, and the range of species must be widened as much as possible.
Material for beginning such experiments is easily obtained, and almost any large sample of seeds will be found suitable. Some tricotyls will be found among every thousand seedlings in many species, while in others ten or a hundred times, as many plants must be examined to secure them, but species with absolutely pure dicotylous seeds are very rare.
The second phase of the experiment, however, is not so promising. Some species are rich, and others are poor in this anomaly. This difference [419] often indicates what can be expected from further culture. Stray tricotyls point to poor species or half-races, while more frequent deviations suggest rich or double-races. In both cases however, the trial must be made, and this requires the isolation of the aberrant individuals and the determination of their hereditary percentage.
In some instances the degree of their inheritance is only a very small one. The isolated tricotyls yield 1 or 2% of inheritors, in some cases even less, or upwards up to 3 or 4%. If the experiment is repeated, no amelioration is observed, and this result remains the same during a series of successive generations. In the case of Polygonum convolvulus, the Black bindweed, I have tried as many as six generations without ever obtaining more than 3%. With other species I have limited myself to four successive years with the same negative result, as with spinage, the Moldavian dragon-head, (Dracocephalum moldavicum), and two species of corn catch-fly (Silene conica and S. conoidea).
Such poor races hardly afford a desirable material for further inquiries. Happily the rich races, though rare, may be discovered also from time to time. They seem to be more common among cultivated plants and horticultural as well as agricultural species may be used. Hemp [420] and mercury (Mercurialis annua) among the first, snapdragon, poppies, Phacelia, Helichrysum, and Clarkia among garden-flowers may be given as instances of species containing the rich tricotylous double races.
It is very interesting to note how strong the difference is between such cases and those which only yield poor races. The rich type at once betrays itself. No repeated selection is required. The stray tricotyls themselves, that are sought out from among the original samples, give hereditary percentages of a much higher type after isolation than those quoted above. They come up to 10-20% and in some cases even to 40%. As may be expected, individual differences occur, and it must even be supposed that some of the original tricotyls may not be pure, but hybrids between tricotylous and dicotylous parents. These are at once eliminated by selection, and if only the tricotyls which have the highest percentages are chosen for the continuance of the new race, the second generation comes up with equal numbers of dicotyls and tricotyls among the seedlings. The figures have been observed to range from 51-58% in the majority of the cases, and average 55%, rarely diverging somewhat more from this average.
Here we have the true type of an ever-sporting variety. Every year it produces in the [421] same way heirs and atavists. Every plant, if fertilized with its own pollen, gives rise to both types. The parent itself may be tricotylous or dicotylous, or show any amount of multiplication and cleavage in its seed-leaves, but it always gives the entire range among its progeny of the variation. One may even select the atavists, pollinate them purely and repeat this in a succeeding generation without any chance of changing the result. On an average the atavists may give lower hereditary figures, but the difference will be only slight.
Such tricotylous double races offer highly interesting material for inquiries into questions of heredity, as they have such a wide range of variability. There is little danger in asserting that they go upwards to nearly 100%, and downwards to 0%, diverging symmetrically on both sides of their average (50-55%). These limits they obviously cannot transgress, and are not even able to reach them. Samples of seed consisting only of tricotyls are very rare, and when they are met with the presumption is that they are too few to betray the rare aberrants they might otherwise contain. Experimental evidence can only be reached by the culture of a succeeding generation, and this always discloses the hidden qualities, showing that the double [422] type was only temporarily lost, but bound to return as soon as new trials are made.
This wide range of variability between definite limits is coupled with a high degree of sensibility and adequateness to the most diverging experiments. Our tricotylous double races are perhaps more sensitive to selection than any other variety, and equally dependent on outer circumstances. Here, however, I will limit myself to a discussion of the former point.
In the second generation after the isolation of stray tricotylous seedlings the average condition of the race is usually reached, but only by some of the strongest individuals, and if we continue the race, sowing or planting only from their offspring, the next generation will show the ordinary type of variability, going upwards in some and downwards in other instances. With the Phacelia and the mercury and some others I had the good luck in this one generation to reach as high as nearly 90% of tricotylous seedlings, a figure indicating that the normal dicotylous type had already become rare in the race. In other cases 80% or nearly 80% was easily attained. Any further divergence from the average would have required very much larger sowings, the effect of selection between a limited number of parents being only to retain the high degree once [423] reached; so for instance with the mercury, I had three succeeding generations of selection after reaching the average of 55%, but their extremes gave no increasing advance, remaining at 86, 92 and 91%.
If we compare these results with the effects of selection in twisted and fasciated races, we observe a marked contrast. Here they reached their height at 30-40%, and no number of generations had the power of making any further improvement. The tricotyls come up in two generations to a proportion of about 54%, which shows itself to correspond to the average type. And as soon as this is reached, only one generation is required to obtain a very considerable improvement, going up to 80 or even 90%.
It is evident that the cause of this difference does not lie in the nature of the monstrosity, but is due to the criterion upon which the selection is made. Selection of the apparently best individuals is one method, and it gives admirable results. Selection on the ground of the hereditary percentages is another method and gives results which are far more advantageous than the former.
In the lecture on the pistillody of the poppies we limited ourselves to the selection of the finest individuals and showed that there is always a manifest correlation between the individual [424] strength of the plant and the degree of development of its anomaly. The same holds good with other monstrosities, and badly nourished specimens of rich races with twisted or fasciated stems always tend to reversion. This reversion, however, is not necessarily correlated with the hereditary percentage and therefore does not always indicate a lessening of the degree of inheritance. This shows that even in those cases an improvement may be expected, if only the means can be found to subject the twisted and the fasciated races to the same sharp test as the tricotylous varieties.
Much remains to be done, and the principle of the selection of parents according to the average constitution of their progeny seems to be one of the most promising in the whole realm of variability.
Besides tricotylous, the syncotylous seedlings may be used in the same way. They are more rarely met with, and in most instances seem to belong only to the unpromising half-races. The black bindweed (Polygonum Convolvulus), the jointed charlock (Raphanus Raphanistrum), the glaucous evening-primrose (Oenothera glauca) and many other plants seem to contain such half-races. On the other hand I found a plant of Centranthus macrosiphon yielding as much as 55% of syncotylous children [425] and thereby evidently betraying the nature of a rich or double race. Likewise the mercury was rich in such deviations. But the best of all was the Russian sunflower, and this was chosen for closer experiments.
In the year of 1888 I had the good luck to isolate some syncotylous seedlings and of finding among them one with 19% of inheritors among its seeds. The following generation at once surpassed the ordinary average and came up in three individuals to 76, 81 and even 89%. My race was at once isolated and ameliorated by selection. I have tried to improve it further and selected the parents with the highest percentages during seven more generations; but without any remarkable result. I got figures of 90% and above, coming even in one instance up to the apparent purity of 100%. These, however, always remained extremes, the averages fluctuating yearly between 80-90% or thereabouts, and the other extremes going nearly every year downwards to 50%, the value which would be attained, if no selection were made.
Contra-selection is as easily made as normal selection. According to our present principle it means the choice of the parents with the smallest hereditary percentage. One might easily imagine that by this means the dicotylous seedlings could be rendered pure. This, however, [426] is not at all the case. It is easy to return from so highly selected figures as for instance 95% to the average about of 50%, as regression to mediocrity is always an easy matter. But to transgress this average on the lower side seems to be as difficult as it is on the upper side. I continued the experiment during four succeeding generations, but was not able to go lower than about 10%, and could not even exclude the high figures from my strain. Parents with 65-75% of syncotylous seedlings returned in each generation, notwithstanding the most careful contra-selection. The attribute is inherent in the race, and is not to be eliminated by so simple a means as selection, nor even by a selection on the ground of hereditary percentages.
We have dealt with torsions and fasciations and with seedling variations at some length, in order to point out the phases needing investigation according to recent views. It would be quite superfluous to consider other anomalies in a similar manner, as they all obey the same laws. A hasty survey may suffice to show what prospects they offer to the student of nature.
First of all come the variegated leaves. They are perhaps the most variable of all variations. They are evidently dependent on external circumstances, and by adequate nutrition the leaves may even become absolutely white or [427] yellowish, with only scarcely perceptible traces of green along the veins. Some are very old cultivated varieties, as the wintercress, or Barbarea vulgaris. They continuously sport into green, or return from this normal color, both by seeds and by buds. Sports of this kind are very often seen on shrubs or low trees, and they may remain there and develop during a long series of years. Bud-sports of variegated holly, elms, chestnuts, beeches and others might be cited. One-sided variegation on leaves or twigs with the opposite side wholly green are by no means rare. It is very curious to note that variegation is perhaps the most universally known anomaly, while its hereditary tendencies are least known.
Cristate and plumose ferns are another instance. Half races or rare accidental cleavages seem to be as common with ferns as cultivated double races, which are very rich in beautiful crests. But much depends on cultivation. It seems that the spores of crested leaves are more apt to reproduce the variety than those of normal leaves, or even of normal parts of the same leaves. But the experiments on which this assertion is made are old and should be repeated. Other cases of cleft leaves should also be tested. Ascidia are far more common than is usually believed. Rare instances point [428] to poor races, but the magnolias and lime-trees are often so productive of ascidia as to suggest the idea of ever-sporting varieties. I have seen many hundred ascidia on one lime-tree, and far above a hundred on the magnolia. They differ widely in size and shape, including in some cases two leaves instead of one, or are composed of only half a leaf or of even still a smaller part of the summit. Rich ascidia-bearing varieties seem to offer notable opportunities for scientific pedigree-cultures.
Union of the neighboring fruits and flowers on flower-heads, of the rays of the umbellifers or of the successive flowers of the racemes of cabbages and allied genera, seem to be rare. The same holds good for the adhesion of foliar to axial organs, of branches to stems and other cases of union. Many of these cases return regularly in each generation, or may at least be seen from time to time in the same strains. Proliferation of the inflorescence is very common and changes in the position of staminate and pistillate flowers are not rare. We find starting points for new investigations in almost any teratological structure. Half-races and double-races are to be distinguished and isolated in all cases, and their hereditary qualities, the periodicity of the recurrence of the anomaly, the dependency on external circumstances [429] and many other questions have to be answered.
Here is a wide field for garden experiments easily made, which might ultimately yield much valuable information on many questions of heredity of universal interest.
[430]
LECTURE XV
DOUBLE ADAPTATIONS
The chief object of all experimentation is to obtain explanations of natural phenomena. Experiments are a repetition of things occurring in nature with the conditions so guarded and so closely followed that it is possible to make a clear analysis of facts and their causes, it being rightfully assumed that the laws are the same in both cases.
Experiments on heredity and the experience of the breeder find their analogy in the succession of generations in the wild state. The stability of elementary species and of retrograde varieties is quite the same under both conditions. Progression and retrogression are narrowly linked everywhere, and the same laws govern the abundance of forms in cultivated and in wild plants.
Elementary species and retrograde varieties are easily recognizable. Ever-sporting varieties on the contrary are far less obvious, and in many cases their hereditary relations have [431] had to be studied anew. A clear analogy between them and corresponding types of wild plants has yet to be pointed out. There can be no doubt that such analogy exists; the conception that they should be limited to cultivated plants is not probable. Striped flowers and variegated leaves, changes of stamens into carpels or into petals may be extremely rare in the wild state, but the “five-leaved” clover and a large number of monstrosities cannot be said to be typical of the cultivated condition. These, however, are of rare occurrence, and do not play any important part in the economy of nature.
In order to attain a better solution of the problem we must take a broader view of the facts. The wide range of variability of ever-sporting varieties is due to the presence of two antagonistic characters which cannot be evolved at the same time and in the same organ, because they exclude one another. Whenever one is active, the other must be latent. But latency is not absolute inactivity and may often only operate to encumber the evolution of the antagonistic character, and to produce large numbers of lesser grades of its development. The antagonism however, is not such in the exact meaning of the word; it is rather a mutual exclusion, because one of the opponents simply takes the place of the other when absent, or supplements [432] it to the extent that it may be only imperfectly developed. This completion ordinarily occurs in all possible degrees and thus causes the wide range of the variability. Nevertheless it may be wanting, and in the case of the double stocks only the two extremes are present.
It is rather difficult to get a clear conception of the substitution, and it seems necessary to designate the peculiar relationship between the two characters forming such a pair by a simple name. They might be termed alternating, if only it were clearly understood that the alternation may be complete, or incomplete in all degrees. Complete alternation would result in the extremes, the incomplete condition in the intermediate states. In some cases as with the stocks, the first prevails, while in other cases, as with the poppies, the very extremes are only rarely met with.
Taking such an alternation as a real character of the ever-sporting varieties, a wide range of analogous cases is at once revealed among the normal qualities of wild plants. Alternation is here almost universal. It is the capacity of young organs to develop in two diverging directions. The definitive choice must be made in extreme youth, or often at a relatively late period of development. Once made, this [433] choice is final, and a further change does not occur in the normal course of things.
The most curious and most suggestive instance of such an alternation is the case of the water-persicaria or Polygonum amphibium. It is known to occur in two forms, one aquatic and the other terrestrial. These are recorded in systematic works as varieties, and are described under the names of P. amphibium var. natans Moench, and P. amphibium var. terrestre Leers or P. amphibium var. terrestris Moench. Such authorities as Koch in his German flora, and Grenier and Godron in their French flora agree in the conception of the two forms as varieties.
Notwithstanding this, the two varieties may often be observed to sport into one another. They are only branches of the same plant, grown under different conditions. The aquatic form has floating or submerged stems with oblong or elliptic leaves, which are glabrous and have long petioles. The terrestrial plants are erect, nearly simple, more or less hispid throughout, with lanceolate leaves and short petioles, often nearly sessile. The aquatic form flowers regularly, producing its peduncle at right angles from the floating stems, but the terrestrial specimens are ordinarily seen without flower-spikes, which are but rarely met with, at least as far as my own experience goes. Intermediate [434] forms are very rare, perhaps wholly wanting, though in swamps the terrestrial plants may often vary widely in the direction of the floating type.
That both types sport into each other has long been recognized in field-observations, and has been the ground for the specific name of amphibium, though in this respect herbarium material seems usually to be scant. The matter has recently been subjected to critical and experimental studies by the Belgian botanist Massart, who has shown that by transplanting the forms into the alternate conditions, the change may always be brought about artificially. If floating plants are established on the shore they make ascending hairy stems, and if the terrestrial shoots are submerged, their buds grow into long and slack, aquatic stems. Even in such experiments, intermediates are rare, both types agreeing completely with the corresponding models in the wild state.
Among all the previously described cases of horticultural plants and monstrosities there is no clearer case of an ever-sporting variety than this one of the water-persicaria. The var. terrestris sports into the var. natans, and as often as the changing life conditions may require it. It is-true that ordinary sports occur without our discerning the cause and without [435] any relation to adaptation. This however is partly due to our lack of knowledge, and partly to the general rule that in nature only such sports as are useful are spared by natural selection, and what is useful we ordinarily term adaptive.
Another side of the question remains to be considered. The word variety, as is now becoming generally recognized, has no special meaning whatever. But here it is assumed in the clearly defined sense of a systematic variety, which includes all subdivisions of species. Such subdivisions may be, from a biological point of view, elementary species and also be eversporting varieties. They may be retrograde varieties, and the two alternating types may be described as separate varieties.
It is readily granted that many writers would not willingly accept this conclusion. But it is simply impossible to avoid it. The two forms of the water-persicaria must remain varieties, though they are only types of the different branches of a single plant.
If not, hundreds and perhaps thousands of analogous cases are at once exposed to doubt, and the whole conception of systematic varieties would have to be thrown over. Biologists of course would have no objection to this, but the student of the flora of any given country [436] or region requires the systematic subdivisions and should always use his utmost efforts to keep them as they are. There is no intrinsic difficulty in the statement that different parts of the same plant should constitute different varieties.
In some cases different branches of the same plant have been described as species. So for instance with the climbing forms of figs. Under the name of Ficus repens a fine little plant is quite commonly cultivated as a climber in flower baskets. It is never seen bearing figs. On the other hand a shrub of our hothouses called Ficus stipulata, is cultivated in pots and makes a small tree which produces quite large, though non-edible figs. Now these two species are simply branches of the same plant. If the repens is allowed to climb up high along the walls of the hothouses, it will at last produce stipulate branches with the corresponding fruits. Ficus radicans is another climbing form, corresponding to the shrub Ficus ulmifolia of our glasshouses. And quite the same thing occurs with ivy, the climbing stems of which never flower, but always first produce erect and free branches with rhombic leaves. These branches have often been used as cuttings and yield little erect and richly flowering shrubs, which are known in [437] horticulture under the varietal name of Hedera Helix arborea.
Manifestly this classification is as nearly right as that of the two varieties of the water-persicaria. Going one step further, we meet with the very interesting case of alpine plants. The vegetation of the higher regions of mountains is commonly called alpine, and the plants show a large number of common features, differentiating them from the flora of lower stations. The mountain plants have small and dense foliage, with large and brightly-colored flowers. The corresponding forms of the lowlands have longer and weaker stems, bearing their leaves at greater distances, the leaves themselves being more numerous. The alpine forms, if perennial, have thick, strongly developed and densely branched rootstocks with heavy roots, in which a large amount of food material is stored up during the short summer, and is available during the long winter months of the year.
Some species are peculiar to such high altitudes, while many forms from the lowlands have no corresponding type on the mountains. But a large number of species are common to both regions, and here the difference of course is most striking. Lotus corniculatus and Calamintha Acinos, Calluna vulgaris and Campanula [438] rotundifolia may be quoted as instances, and every botanist who has visited alpine regions may add other examples. Even the edelweiss of the Swiss Alps, Gnaphalium Leontopodium, loses its alpine characters, if cultivated in lowland gardens. Between such lowland and alpine forms intermediates regularly occur. They may be met with whenever the range of the species extends from the plains upward to the limit of eternal snow.
In this case the systematists formerly enumerated the alpine plants as forma alpestris, but whenever the intermediate is lacking the term Varietas alpestris was often made use of.
It is simply impossible to decide concerning the real relation between the alpine and lowland types without experiments. About the middle of the last century it was quite a common thing to collect plants not only for herbarium-material, but also for the purpose of planting them in gardens and thus to observe their behavior under new conditions. This was done with the acknowledged purpose of investigating the systematic significance of observed divergencies. Whenever these held good in the garden they were considered to be reliable, but if they disappeared they were regarded as the results of climatic conditions, or of the influence of soil or nourishment. Between [439] these two alternatives, many writers have tried to decide, by transplanting their specimens after some time in the garden, into arid or sandy soil, in order to see whether they would resume their alpine character.
Among the systematists who tested plants in this way, Nageli especially, directed his attention to the hawkweeds or Hieracium. On the Swiss Alps they are very small and exhibit all the characters of the pure alpine type. Thousands of single plants were cultivated by him in the botanical garden of Munich, partly from seed and partly from introduced rootstocks. Here they at once assumed the tall stature of lowland forms. The identical individual, which formerly bore small rosettes of basal leaves, with short and unbranched flower-stalks, became richly leaved and often produced quite a profusion of flower-heads on branched stems. If then they were transplanted to arid sand, though remaining in the same garden and also under the same climatic conditions they resumed their alpine characters. This proved nutrition to be the cause of the change and not the climate.
The latest and most exact researches on this subject are due to Bonnier, who has gone into all the details of the morphologic as well as of the physiologic side of the problem. [440] His purpose was the study of partial variability under the influence of climate and soil. In every experiment he started from a single individual, divided it into two parts and planted one half on a mountain and the other half on the plain. The garden cultures were made chiefly at Paris and Fontainebleau, the alpine cultures partly in the Alps, partly in the Pyrenees. From time to time the halved plants were compared with each other, and the cultures lasted, as a rule, during the lifetime of the individual, often covering many years.
The common European frostweed or Helianthemum vulgare will serve to illustrate his results. A large plant growing in the Pyrenees in an altitude of 2,400 meters was divided. One half was replanted on the same spot, and the other near Cadeac, at the base of the mountain range (740 M.). In order to exclude the effect of a change of soil, a quantity of the earth from the original locality was brought into the garden and the plant put therein. Further control experiments were made at Paris. As soon as the two halved individuals commenced to grow and produced new shoots, the influence of the different climates made itself felt. On the mountain, the underground portions remained strong and dense, the leaves and internodes small and hairy, the flowering stems nearly [441] procumbent, the flowers being large and of a deep yellow. At Cadeac and at Paris the whole plant changed at once, the shoots becoming elongated and loose, with broad and flattened, rather smooth leaves and numerous pale-hued flowers. The anatomical structure exhibited corresponding differences, the intercellular spaces being small in the alpine plant and large in the one grown in the lowlands, the wood-tissues strong in the first and weak in the second case.
The milfoil (Achillea Millefolium) served as a second example, and the experiments were carried on in the same localities. The long and thick rootstocks of the alpine plant bearing short stems only with a few dense corymbs contrasted markedly with the slender stems, loose foliage and rich groups of flowerheads of the lowland plant. The same differences, in inner and outer structures were observed in numerous instances, showing that the alpine type in these cases is dependent on the climate, and that the capacity for assuming the antagonistic characters is present in every individual of the species. The external conditions decide which of them becomes active and which remains inactive, and the case seems to be exactly parallel to that of the water-persicaria.
In the experiments of Bonnier the influence of the soil was, as a rule, excluded by transplanting [442] part of the original earth with the transplanted half of the plant. From this he concluded that the observed changes were due to the inequality of the climate. This involved three main factors, light, moisture and temperature. On the mountains the light is more intense, the air drier and cooler. Control-experiments were made on the mountains, depriving the plants of part of the light. In various ways they were more or less shaded, and as a rule responded to this treatment in the same way as to transplantation to the plain below. Bonnier concluded that, though more than one factor takes part in inciting the morphologic changes, light is to be considered as the chief agency. The response is to be considered as a useful one, as the whole structure of the alpine varieties is fitted to produce a large amount of organic material in a short time, which enables the plants to thrive during the short summers and long winters of their elevated stations.
In connection with these studies on the influences of alpine climates, Bonnier has investigated the internal structure of arctic plants, and made a series of experiments on growth in continuous electric light. The arctic climate is cold, but wet, and the structure of the leaves is correspondingly loose, though the plants become [443] as small as on the Alps. Continuous electric light had very curious effects; the plants became etiolated, as if growing in darkness, with the exception that they assumed a deep green tinge. They showed more analogy with the arctic than with the alpine type.
The influence of the soil often produces changes similar to that of climate. This was shown by the above cited experiments of Nageli with the hawkweeds, and may easily be controlled in other cases. The ground-honeysuckle or Lotus corniculatus grows in Holland partly on the dry and sandy soil of the dunes, and occasionally in meadows. It is small and dense in the first case, with orange and often very darkly colored petals, while it is loose and green in the meadows, with yellower flowers. Numerous analogous cases might be given. On mountain slopes in South Africa, and especially in Natal, a species of composite is found, which has been introduced into culture and is used as a hanging plant. It is called Othonna crassifolia and has fleshy, nearly cylindrical leaves, and exactly mimics some of the crassulaceous species. On dry soil the leaves become shorter and thicker and assume a reddish tinge, the stems remain short and woody and bear their leaves in dense rosettes. On moist and rich garden-soil this aspect becomes [444] changed at once, the stems grow longer and of a deeper green. Intermediates occur, but notwithstanding this the two extremes constitute clearly antagonistic types.
The flora of the deserts is known to exhibit a similar divergent type. Or rather two types, one adapted to paucity of water, and the other to a storage of fluid at one season in order to make use of it at other times, as is the case with the cactuses. Limiting ourselves to the alternate group, we observe a rich and dense branching, small and compact leaves and extraordinarily long roots. Here the analogy with the alpine varieties is manifest, and the dryness of the soil evidently affects the plants in a similar way, as do the conditions of life in alpine regions. The question at once comes up as to whether here too we have only instances of partial variability, and whether many of the typical desert-species would lose their peculiar character by cultivation under ordinary conditions. The varieties of Monardella macrantha, described by Hall, from the San Jacinto Mountain, Cal., are suggestive of such an intimate analogy with the cases studied by Bonnier, that it seems probable that they might yield similar results, if tested by the same method.
Leaving now the description of these special [445] cases, we may resume our theoretical discussion of the subject, and try to get a clearer insight into the analogy of ever-sporting varieties and the wild species quoted. All of them may be characterized by the general term of dimorphism. Two types are always present, though not in the same individual or in the same organ. They exclude one another, and during their juvenile stage a decision is taken in one direction or in the other. Now, according to the theory of natural selection, wild species can only retain useful or at least innocuous qualities, since all mutations in a wrong direction must perish sooner or later. Cultivated species on the other hand are known to be largely endowed with qualities, which would be detrimental in the wild condition. Monstrosities are equally injurious and could not hold their own if left to themselves.
These same principles may be applied to ever-sporting or antagonistic pairs of characters. According to the theory of mutations such pairs may be either useful or useless. But only the useful will stand further test, and if they find suitable conditions will become specific or varietal characters. On this conclusion it becomes at once clear, why natural dimorphism is, as a rule, a very useful quality, while the cultivated dimorphous varieties [446] strike us as something unnatural. The relation between cause and effect, is in truth other than it might seem to be at first view, but nevertheless it exists, and is of the highest importance.
From this same conclusion we may further deduce some explanation of the hereditary races characterized by monstrosities. It is quite evident that the twisted teasels are inadequate for the struggle with their tall congeners, or with the surrounding plants. Hence the conclusion that a pure and exclusively twisted race would soon die out. The fact that such races are not in existence finds its explanation in this circumstance, and therefore it does not prove the impossibility or even the improbability that some time a pure twisted race might arise. If chance should put such an accidental race in the hands of an experimenter, it could be protected and preserved, and having no straight atavistic branches, but being twisted in all its organs, might yield the most curious conceivable monstrosity, surpassing even the celebrated dwarf twisted shrubs of Japanese horticulturists.
Such varieties however, do not exist at present. The ordinary twisted races on the other hand, are found in the wild state and have only to be isolated and cultivated to yield large numbers [447] of twisted individuals. In nature they are able to maintain themselves during long centuries, quite as well as normal species and varieties. But they owe this quality entirely to their dimorphous character. A twisted race of teasels might consist of successive generations of tall atavistic individuals, and produce yearly some twisted specimens, which might be destroyed every time before ripening their seeds. Reasoning from the evidence available, and from analogous cases, the variety would, even under such extreme circumstances, be able to last as long as any other good variety or elementary species. And it seems to me that this explanation makes clear how it is possible that varieties, which are potentially rich in their peculiar monstrosity, are discovered from time to time among plants when tested by experimental methods.
Granting these conclusions, monstrosities on the one side, and dimorphous wild species on the other, constitute the most striking examples of the inheritance of latent characters.
The bearing of the phenomena of dimorphism upon the principles of evolution formulated by Lamarck, and modified by his followers to constitute Neo-Lamarckianism, remains to be considered. Lamarck assumed that the external conditions directly affected the organisms in [448] such a way as to make them better adapted to life, under prevailing circumstances. Nageli gave to this conception the name “Theory of direct causation” (Theorie der directen Bewirkung), and it has received the approval of Von Wettstein, Strasburger and other German investigators. According to this conception a plant, when migrating from lowlands into the mountains would slowly be changed and gradually assume alpine habits. Once acquired this habit would become fixed and attain the rank of specific characters. In testing this theory by field-observations and culture-experiments, the defenders of the Nagelian principle could easily produce evidence upon the first point. The change of lowland-plants into alpine varieties can be brought about in numerous cases, and corresponding changes under the influence of soil, or climate, or life-conditions are on record for the most various characters and qualities.
The second point, however, is as difficult to prove as the first is of easy treatment. If after hundreds and thousands of years of exposure to alpine or other extreme conditions a fixed change is proved to have taken place, the question remains unanswered, whether the change has been a gradual or a sudden one. Darwin pointed out that long periods of life afford a [449] chance for a sudden change in the desired direction, as well as for the slow accumulation of slight deviations. Any mutations in a wrong direction would at once be destroyed, but an accidental change in a useful way would be preserved, and multiply itself. If in the course of centuries this occurred, they would be nearly sure to become established, however rare at the outset. Hence the positive assertion is scarcely capable of direct proof.
On the other hand the negative assertion must be granted full significance. If the alpine climate has done no more than produce a transitory change, it is clear that thousands of years do not, necessarily, cause constant and specific alterations. This requirement is one of the indispensable supports of the Lamarckian theory. The matter is capable of disproof however, and such disproof seems to be afforded by the direct evidence of the present condition of the alpine varieties at large, and by many other similar cases.
Among these the observations of Holtermann on some desert-plants of Ceylon are of the highest value. Moreover they touch questions which are of wide importance for the study of the biology of American deserts. For this reason I may be allowed to introduce them here at some length.
[450] The desert of Kaits, in Northern Ceylon, nourishes on its dry and torrid sands some species, represented by a large number of individuals, together with some rarer plants. The commonest forms are Erigeron Asteroides, Vernonia cinerea, Laurea pinnatifida, Vicoa auriculata, Heylandia latebrosa and Chrysopogon montanus. In direct contrast with the ordinary desert-types they have a thin epidermis, with exposed stomata, features that ordinarily were characteristic of species of moister regions. They are annuals, growing rapidly, blooming and ripening their seeds before the height of the dry season. Evidently they are to be considered as the remainder of the flora of a previous period, when the soil had not yet become arid. They might be called relics. Of course they are small and dwarf-like, when compared with allied forms.
These curious little desert-plants disprove the Nagelian views in two important points. First, they show that extreme conditions do not necessarily change the organisms subjected to them, in a desirable direction. During the many centuries that these plants must have existed in the desert in annual generations, no single feature in the anatomical structure has become changed. Hence the conclusion that small leaves, abundant rootstocks and short [451] stems, a dense foliage, a strongly cuticularized epidermis, few and narrow air-cavities in the tissues and all the long range of characteristics of typical desert-plants are not a simple result of the influence of climate and soil. There is no direct influence in this sense.
The second point, in which Nageli’s idea is broken down by Holtermann’s observations, results from the behavior of the plants of the Kaits desert when grown or sown on garden soil. When treated in this way they at once lose the only peculiarity which might be considered as a consequence of the desert-life of their ancestors, their dwarf stature. They behave exactly like the alpine plants in Bonnier’s experiments, and with even more striking differences. In the desert they attain a height of a few centimeters, but in the garden they attain half a meter and more in height. Nothing in the way of stability has resulted from the action of the dry soil, not even in such a minor point as the height of the stems.
From the facts and discussions we may conclude that double adaptation is not induced by external influences, at least not in any way in which it might be of use to the plant. It may arise by some unknown cause, or may not be incited at all. In the first case the plant becomes capable of living under the alternating [452] circumstances, and if growing near the limits of such regions it will overlap and get into the new area. All other species, which did not acquire the double habit, are of course excluded, with such curious exceptions as those of Kaits. The typical vegetation under such extreme conditions however, finds explanation quite as well by the one as by the other view.
Leaving these obvious cases of double adaptation, there still remains one point to be considered. It is the dwarf stature of so many desert and alpine plants. Are these dwarfs only the extremes of the normal fluctuating variability, or is their stature to be regarded as the expression of some peculiar adaptive but latent quality? It is as yet difficult to decide this question, because statistical studies of this form of variability are still wanting. The capacity of ripening the seed on individuals of dwarf stature however, is not at all a universal accompaniment of a variable height. Hence it cannot be considered as a necessary consequence of it. On the other hand the dwarf varieties of numerous garden-plants, as for instance: of larkspurs, snapdragon, opium-poppies and others are quite stable and thence are obviously due to peculiar characteristics. Such characteristics, if combined with tall stature into a pair of antagonists, would yield a double [453] adaptation, and on such a base a hypothetical explanation could no doubt be rested. Instead of discussing this problem from the theoretical side, I prefer to compare those species which are capable of assuming a dwarf stature under less uncommon conditions than those of alpine and desert-plants. Many weeds of our gardens and many wild species have this capacity. They become very tall, with large leaves, richly branched stems and numerous flowers in moist and rich soil. On bad soil, or if germinating too late, when the season is drier, they remain very small, producing only a few leaves and often limiting themselves to one flower-head. This is often seen with thorn-apples and amaranths, and even with oats and rye, and is notoriously the case with buckwheat. Gauchery has observed that the extremes differ often as much from one another as 1:10. In the case of the Canadian horseweed or Erigeron canadensis, which is widely naturalized in Europe, the tallest specimens are often twenty-five times as tall as the smallest, the difference increasing to greater extremes, if besides the main stem, the length of the numerous branches of the tall plants are taken into consideration. Other instances studied by the French investigator are Erythraea pulchella and Calamintha Acinos.
[454] Dimorphism is of universal occurrence in the whole vegetable kingdom. In some cases it is typical, and may easily be discerned from extreme fluctuating variability. In others the contrast is not at all obvious, and a closer investigation is needed to decide between the two possibilities. Sometimes the adaptive quality is evident, in other cases it is not. A large number of plants bear two kinds of leaves linked with one another by intermediate forms. Often the first leaves of a shoot, or those of accidentally strong shoots, exhibit deviating shapes, and the usefulness of such occurrences seems to be quite doubtful. The elongation of stems and linear leaves, and the reduction of lateral organs in darkness, is manifestly an adaptation. Many plants have stolons with double adaptations which enable them to retain their character of underground stems with bracts or to exchange it for the characteristics of erect stems with green leaves according to the outer circumstances. In some shrubs and trees the capacity of a number of buds to produce either flowers or shoots with leaves seems to be in the same condition. The capacity of producing spines is also a double adaptation, active on dry and arid soil and latent in a moist climate or under cultivation, as with the wild and cultivated apple, and in the experiments of Lothelier [455] with Berberis, Lycium and other species, which lose their spines in damp air.
In some conifers the evolution of horizontal branches may be modified by simply turning the buds upside down. Or the lateral branches can be induced to become erect stems by cutting off the normal summit of a tree. Numerous organs and functions lie dormant until aroused by external agencies, and many other cases could be cited, showing the wide occurrence of double adaptation.
There are, however, two points, which should not be passed over without some mention. One of them is the influence of sun and shade on leaves, and the other the atavistic forms, often exhibited during the juvenile period.
The leaves of many plants, and especially those of some shrubs and trees, have the capacity of adapting themselves either to intense or to diffuse light. On the circumference of the crown of a tree the light is stronger and the leaves a small and thick, with a dense tissue. In the inner parts of the crown the light is weak and the leaves are broader in order to get as much of it as possible. They become larger but thinner, consisting often of a small number of cell layers. The definitive formation is made in extreme youth, often even during the previous summer, at the time of the [456] very first evolution of the young organs within the buds. Iris, and Lactuca Scariola or the prickly lettuce, and many other plants afford similar instances. As the definitive decision must be made in these cases long before the direct influence of the conditions which would make the change useful is felt, it is hardly conceivable how they could be ascribed to this cause.
It is universally known that many plants show deviating features when very young, and that these often remind us of the characters of their probable ancestors. Many plants that must have been derived from their nearest systematic relatives, chiefly by reductions, are constantly betraying this relation by a repetition of the ancestral marks during their youth.
There can be hardly a doubt that the general law of natural selection prevails in such cases as it does in others. Or stated otherwise, it is very probable, that in most cases the atavistic characters have been retained during youth because of their temporary usefulness. Unfortunately, our knowledge of utility of qualities is as yet, very incomplete. Here we must assume that what is ordinarily spared by natural selection is to be considered as useful, [457] until direct experimental investigations have been made.
So it is for instance with the submerged leaves of water-plants. As a rule they are linear, or if compound, are reduced to densely branching filiform threads. Hence we may conclude that this structure is of some use to them. Now two European and some corresponding American species of water-parsnip, the Sium latifolium and Berula angustifolia with their allies, are umbellifers, which bear pinnate instead of bi- or tri-pinnate leaves. But the young plants and even the young shoots when developing from the rootstocks under water comply with the above rule, producing very compound, finely and pectinately dissected leaves. From a systematic point of view these leaves indicate the origin of the water-parsnips from ordinary umbellifers, which generally have bi- and tripinnate leaves.
Similar cases of double adaptation, dependent on external conditions at different periods of the evolution of the plant are very numerous. They are most marked among leguminous plants, as shown by the trifoliolate leaves of the thorn-broom and allies, which in the adult state have green twigs destitute of leaves.
As an additional instance of dimorphism and probable double adaptation to unrecognized external [458] conditions I might point to the genus Acacia. As we have seen in a previous lecture some of the numerous species of this genus bear bi-pinnate leaves, while others have only flattened leaf-stalks. According to the prevailing systematic conceptions, the last must have been derived from the first by the loss of the blades and the corresponding increase of size and superficial extension of the stalk. In proof of this view they exhibit, as we have described, the ancestral characters in the young plantlets, and this production of bi-pinnate leaves has probably been retained at the period of the corresponding negative mutations, because of some distinct, though still unknown use.
Summarizing the results of this discussion, we may state that useful dimorphism, or double adaptation, is a substitution of characters quite analogous to the useless dimorphism of cultivated ever-sporting varieties and the stray occurrence of hereditary monstrosities. The same laws and conditions prevail in both cases.
[459]
E. MUTATIONS