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

LECTURE XVI

THE ORIGIN OF THE PELORIC TOAD-FLAX

I have tried to show previously that species, in the ordinary sense of the word, consist of distinct groups of units. In systematic works these groups are all designated by the name of varieties, but it is usually granted that the units of the system are not always of the same value. Hence we have distinguished between elementary species and varieties proper. The first are combined into species whose common original type is now lost or unknown, and from their characters is derived an hypothetical image of what the common ancestor is supposed to have been. The varieties proper are derived in most cases from still existing types, and therefore are subjoined to them. A closer investigation has shown that this derivation is ordinarily produced by the loss of some definite attribute, or by the re-acquisition of an apparently [460] lost character. The elementary species, on the other hand, must have arisen by the production of new qualities, each new acquisition constituting the origin of a new elementary form.

Moreover we have seen, that such improvements and such losses constitute sharp limits between the single unit-forms. Every type, of course, varies around an average, and the extremes of one form may sometimes reach or even overlap those of the nearest allies, but the offspring of the extremes always return to the type. The transgression is only temporary and a real transition of one form to another does not come within ordinary features of fluctuating variability. Even in the cases of eversporting varieties, where two opposite types are united within one race, and where the succeeding individuals are continually swinging from one extreme to the other, passing through a wide range of intermediate steps, the limits of the variety are as sharply defined and as free from real transgression as in any other form.

In a complete systematic enumeration of the real units of nature, the elementary species and varieties are thus observed to be discontinuous and separated by definite gaps. Every unit may have its youth, may lead a long life in the adult state and may finally die. But through [461] the whole period of its existence it remains the same, at the end as sharply defined from its nearest allies as in the beginning. Should some of the units die out, the gaps between the neighboring ones will become wider, as must often have been the case. Such segregations, however important and useful for systematic distinctions, are evidently only of secondary value, when considering the real nature of the units themselves.

We may now take up the other side of the problem. The question arises as to how species and varieties have originated. According to the Darwinian theory they have been produced from one another, the more highly differentiated ones from the simpler, in a graduated series from the most simple forms to the most complicated and most highly organized existing types. This evolution of course must have been regular and continuous, diverging from time to time into new directions, and linking all organisms together into one common pedigree. All lacunae in our present system are explained by Darwin as due to the extinction of the forms, which previously filled them.

Since Lamarck first propounded the conception of a common origin for all living beings, much has been done to clear up our ideas as to the real nature of this process. The broader [462] aspect of the subject, including the general pedigree of the animal and vegetable kingdom, may be said to have been outlined by Darwin and his followers, but this phase of the subject lies beyond the limits of our present discussion.

The other phase of the problem is concerned with the manner in which the single elementary species and varieties have sprung from one another. There is no reason to suppose that the world is reaching the end of its development, and so we are to infer that the production of new species and varieties is still going on. In reality, new forms are observed to originate from time to time, both wild and in cultivation, and such facts do not leave any doubt as to their origin from other allied types, and according to natural and general laws.

In the wild state however, and even with cultivated plants of the field and garden, the conditions, though allowing of the immediate observation of the origination of new forms, are by no means favorable for a closer inquiry into the real nature of the process. Therefore I shall postpone the discussion of the facts till another lecture, as their bearing will be more easily understood after having dealt with more complete cases.

These can only be obtained by direct experimentation. Comparative studies, of course, [463] are valuable for the elucidation of general problems and broad features of the whole pedigree, but the narrower and more practical question as to the genetic relation of the single forms to one another must be studied in another way, by direct experiment. The exact methods of the laboratory must be used, and in this case the garden is the laboratory. The cultures must be guarded with the strictest care and every precaution taken to exclude opportunities for error. The parents and grandparents and their offspring must be kept pure and under control, and all facts bearing upon the birth or origin of the new types should be carefully recorded.

Two great difficulties have of late stood in the way of such experimental investigation. One of them is of a theoretical, the, other of a practical nature. One is the general belief in the supposed slowness of the process, the other is the choice of adequate material for experimental purposes. Darwin's hypothesis of natural selection as the means by which new types arise, is now being generally interpreted as stating the slow transformation of ordinary fluctuating divergencies from the average type into specific differences. But in doing so it is overlooked that Quetelet's law of fluctuating variability was not yet discovered at the time, when Darwin propounded his theory. So there [464] is no real and intimate connection between these two great conceptions. Darwin frequently pointed out that a long period of time might be needed for slow improvements, and was also a condition for the occurrence of rare sports. In any case those writers have been in error, according to my opinion, who have refrained from experimental work on the origin of species, on account of this narrow interpretation of Darwin's views. The choice of the material is quite another question, and obviously all depends upon this choice. Promising instances must be sought for, but as a rule the best way is to test as many plants as possible. Many of them may show nothing of interest, but some might lead to the desired end.

For to-day's lecture I have chosen an instance, in which the grounds upon which the choice was based are very evident. It is the origin of the peloric toad-flax (Linaria vulgaris peloria).

The ground for this choice lies simply in the fact that the peloric toad-flax is known to have originated from the ordinary type at different times and in different countries, under more or less divergent conditions. It had arisen from time to time, and hence I presumed that there was a chance to see it arise again. If this should happen under experimental circumstances [465] the desired evidence might easily be gathered. Or, to put it in other words, we must try to arrange things so as to be present at the time when nature produces another of these rare changes.

There was still another reason for choosing this plant for observational work. The step from the ordinary toad-flax to the peloric form is short, and it appears as if it might be produced by slow conversion. The ordinary species produces from time to time stray peloric flowers. These occur at the base of the raceme, or rarely in the midst of it. In other species they are often seen at the summit. Terminal pelories are usually regular, having five equal spurs. Lateral pelories are generally of zygomorphic structure, though of course in a less degree than the normal bilabiate flowers, but they have unequal spurs, the middle one being of the ordinary length, the two neighboring being shorter, and those standing next to the opposite side of the flower being the shortest of all. This curious remainder of the original, symmetrical structure of the flower seems to have been overlooked hitherto by the investigators of peloric toad-flaxes.

The peloric variety of this plant is characterized by its producing only peloric flowers. No single bilabiate or one-spurred flower remains.

[466] I once had a lot of nearly a hundred specimens of this fine variety, and it was a most curious and beautiful sight to observe the many thousands of nearly regular flowers blooming at the same time. Some degree of variability was of course present, even in a large measure. The number of the spurs varied between four and six, transgressing these limits in some instances, but never so far as to produce really one-spurred flowers. Comparing this variety with the ordinary type, two ways of passing over from the one to the other might be imagined. One would entail a slow increase of the number of the peloric flowers on each plant, combined with a decrease of the number of the normal ones, the other a sudden leap from one extreme to the other without any intermediate steps. The latter might easily be overlooked in field observations and their failure may not have the value of direct proof. They could never be overlooked, on the other hand, in experimental culture.

The first record of the peloric toad-flax is that of Zioberg, a student of Linnaeus, who found it in the neighborhood of Upsala. This curious discovery was described by Rudberg in his dissertation in the year 1744. Soon afterwards other localities were discovered by Link near Gottingen in Germany about 1791 and afterwards [467] in the vicinity of Berlin, as stated by Ratzeburg, 1825. Many other localities have since been indicated for it in Europe, and in my own country some have been noted of late, as for instance near Zandvoort in 1874 and near Oldenzaal in 1896. In both these last named cases the peloric form arose spontaneously in places which had often been visited by botanists before the recorded appearance, and therefore, without any doubt, they must have been produced directly and independently by the ordinary species which grows in the locality. The same holds good for other occurrences of it. In many instances the variety has been recorded to disappear after a certain lapse of time, the original specimens dying out and no new ones being produced. Linaria is a perennial herb, multiplying itself easily by buds growing on the roots, but even with this means of propagation its duration seems to have definite limits.

There is one other important point arguing strongly for the independent appearance of the peloric form in its several localities. It is the difficulty of fertilization and the high degree of sterility, even if artificially pollinated. Bees and bumble-bees are unable to crawl into the narrow tubular flowers, and to bring the fertilizing pollen to the stigma. Ripe capsules with seeds [468] have never been seen in the wild state. The only writer who succeeded in sowing seeds of the peloric variety was Wildenow and he got only very few seedlings. But even in artificial pollination the result is the same, the anthers seeming to be seriously affected by the change. I tried both self-fertilization and cross-pollination, and only with utmost care did I succeed in saving barely a hundred seeds. In order to obtain them I was compelled to operate on more than a thousand flowers on about a dozen peloric plants.

The variety being wholly barren in nature, the assumption that the plants in the different recorded localities might have a common origin is at once excluded. There must have been at least nearly as many mutations as localities. This strengthens the hope of seeing such a mutation happen in one's own garden. It should also be remembered that peloric flowers are known to have originated in quite a number of different species of Linaria, and also with many of the allied species within the range of the Labiatiflorae.

I will now give the description of my own experiment. Of course this did not give the expected result in the first year. On the contrary, it was only after eight years' work that I had the good fortune of observing the mutation. [469] But as the whole life-history of the preceding generations had been carefully observed and recorded, the exact interpretation of the fact was readily made.

My culture commenced in the year 1886. I chose some plants of the normal type with one or two peloric flowers besides the bilabiate majority which I found on a locality in the neighborhood of Hilversum in Holland. I planted the roots in my garden and from them had the first flowering generation in the following summer. From their seeds I grew the second generation in three following years. They flowered profusely and produced in 1889 only one, and in 1890 only two peloric structures. I saved the seeds in 1889 and had in 1890-1891 the third generation. These plants likewise flowered only in the second year, and gave among some thousands of symmetrical blossoms, only one five-spurred flower. I pollinated this flower myself, and it produced abundant fruit with enough seeds for the entire culture in 1892, and they only were sown.

Until this year my generations required two years each, owing to the perennial habit of the plants. In this way the prospects of the culture began to decrease, and I proposed to try to heighten my chances by having a new generation yearly. With this intention I sowed the [470] selected seeds in a pan in the glasshouse of my laboratory and planted them out as soon as the young stems had reached a length of some few centimeters. Each seedling was put in a separate pot, in heavily manured soil. The pots were kept under glass until the beginning of June, and the young plants produced during this period a number of secondary stems from the curious hypocotylous buds which are so characteristic of the species. These stems grew rapidly and as soon as they were strong enough, the plants were put into the beds. They all, at least nearly all, some twenty specimens, flowered in the following month.

I observed only one peloric flower among the large number present. I took the plant bearing this flower and one more for the culture of the following year, and destroyed all others. These two plants grew on the same spot, and were allowed to fertilize each other by the agency of the bees, but were kept isolated from any other congener. They flowered abundantly, but produced only one-spurred bilabiate flowers during the whole summer. They matured more than 10 cu. cm. of seeds.

It is from this pair of plants that my peloric race has sprung. And as they are the ancestors of the first closely observed case of peloric mutation, [471] it seems worth while to give some details regarding their fertilization.

Isolated plants of Linaria vulgaris do not produce seed, even if freely pollinated by bees. Pollen from other plants is required. This requirement is not at all restricted to the genus Linaria, as many instances are known to occur in different families. It is generally assumed that the pollen of any other individual of the same species is capable of producing fertilization, although it is to be said that a critical examination has been made in but few instances.

This, however, is not the case, at least not in the present instance. I have pollinated a number of plants, grown from seed of the same strain and combined them in pairs, and excluded the visits of insects, and pollen other than that of the plant itself and that of the specimen with which it was paired. The result was that some pairs were fertile and others barren. Counting these two groups of pairs, I found them nearly equal in number, indicating thereby that for any given individual the pollen of half of the others is potent, but that of the other half impotent. From these facts we may conclude the presence of a curious case of dimorphy, analogous to that proposed for the primroses, but without visible differentiating marks in the flowers. At least such opposite characters [472] have as yet not been ascertained in the case of our toad-flax.

In order to save seed from isolated plants it is necessary, for this reason, to have at least two individuals, and these must belong to the two physiologically different types. Now in the year 1892, as in other years, my plants, though separated at the outset by distances of about 20 cm. from each other, threw out roots of far greater length, growing in such a way as to abolish the strict isolation of the individuals. Any plot may produce several stems from such roots, and it is manifestly impossible to decide whether they all belong to one original plant or to the mixed roots of several individuals. No other strains were grown on the same bed with my plants however, and so I considered all the stems of the little group as belonging to one plant. But their perfect fertility showed, according to the experience described, that there must have been at least two specimens mingled together.

Returning now to the seeds of this pair of plants, I had, of course, not the least occasion to ascribe to it any higher value than the harvest of former years. The consequence was that I had no reason to make large sowings, and grew only enough young plants to have about 50 in bloom in the summer of 1894. Among [473] these, stray peloric flowers were observed in somewhat larger number than in the previous generations, 11 plants bearing one or two, or even three such abnormalities. This however, could not be considered as a real advance, since such plants may occur in varying, though ordinarily small numbers in every generation.

Besides them a single plant was seen to bear only peloric flowers; it produced racemes on several stems and their branches. All were peloric without exception. I kept it through the winter, taking care to preserve a complete isolation of its roots. The other plants were wholly destroyed. Such annihilation must include both the stems and roots and the latter of course requires considerable labor. The following year, however, gave proof of the success of the operation, since my plant bloomed luxuriously for the second time and remained true to the type of the first year, producing peloric flowers exclusively.

Here we have the first experimental mutation of a normal into a peloric race. Two facts were clear and simple. The ancestry was known for over a period of four generations, living under the ordinary care and conditions of an experimental garden, isolated from other toad-flaxes, but freely fertilized by bees or at times by myself. This ancestry was quite constant as to [474] the peloric peculiarity, remaining true to the wild type as it occurs everywhere in my country, and showing in no respect any tendency to the production of a new variety.

The mutation took place at once. It was a sudden leap from the normal plants with very rare peloric flowers to a type exclusively peloric. No intermediate steps were observed. The parents themselves had borne thousands of flowers during two summers, and these were inspected nearly every day, in the hope of finding some pelories and of saving their seed separately. Only one such flower was seen. If there had been more, say a few in every hundred flowers, it might be allowable to consider them as previous stages, showing a preparation of the impending change. But nothing of this kind was observed. There was simply no visible preparation for the sudden leap.

This leap, on the other hand, was full and complete. No reminiscence of the former condition remained. Not a single flower on the mutated plant reverted to the previous type. All were thoroughly affected by the new attribute, and showed the abnormally augmented number of spurs, the tubular structure of the corolla and the round and narrow entrance of its throat. The whole plant departed absolutely from the old type of its progenitors.

[475] Three ways were open to continue my experiment. The first was indicated by the abundant harvest from the parent-plants of the mutation. It seemed possible to compare the numerical proportion of the mutated seeds with those of normal plants. In order to ascertain this proportion I sowed the greatest part of my 10 cu. cm. of seed and planted some 2,000 young plants in little pots with well-manured soil. I got some 1,750 flowering plants and observed among them 16 wholly peloric individuals. The numerical proportion of the mutation was therefore in this instance to be calculated equal to about 1% of the whole crop.

This figure is of some importance. For it shows that the chance of finding mutations requires the cultivation of large groups of individuals. One plant in each hundred may mutate, and cultures of less than a hundred specimens must therefore be entirely dependent on chance for the appearance of new forms, even if such should accidentally have been produced and lay dormant in the seed. In other cases mutations may be more numerous, or on the contrary, more rare. But the chance of mutative changes in larger numbers is manifestly much reduced by this experiment, and they may be expected to form a very small proportion of the culture.

[476] The second question which arose from the above result was this. Could the mutation be repeated? Was it to be ascribed to some latent cause which might be operative more than once? Was there some hidden tendency to mutation, which, ordinarily weak, was strengthened in my cultures by some unknown influence? Was the observed mutation to be explained by a common cause with the other cases recorded by field-observations? To answer this question I had only to continue my experiment, excluding the mutated individuals from any intercrossing with their brethren. To this end I saved the seeds from duly isolated groups in different years and sowed them at different times. For various causes I was not prepared to have large cultures from these seeds, but notwithstanding this, the mutation repeated itself. In one instance I obtained two, in another, one peloric plant with exclusively many-spurred flowers. As is easily understood, these were related as "nieces" to the first observed mutants. They originated in quite the same way, by a sudden leap, without any preparation and without any intermediate steps.

Mutation is proved by this experience to be of an iterative nature. It is the expression of some concealed condition, or as it is generally [477] called, of some hidden tendency. The real nature of this state of the hereditary qualities is as yet wholly unknown. It would not be safe to formulate further conclusions before the evidence offered by the evening-primroses is considered.

Thirdly, the question arises, whether the mutation is complete, not only as to the morphologic character, but also as to the hereditary constitution of the mutated individuals. But here unfortunately the high degree of sterility of the peloric plants, as previously noted, makes the experimental evidence a thing of great difficulty. During the course of several years I isolated and planted together the peloric individuals already mentioned, all in all some twenty plants. Each individual was nearly absolutely sterile when treated with its own pollen, and the aid of insects was of no avail. I intercrossed my plants artificially, and pollinated more than a thousand flowers. Not a single one gave a normal fruit, but some small and nearly rudimentary capsules were produced, bearing a few seeds. From these I had 119 flowering plants, out of which 106 were peloric and 13 one-spurred. The great majority, some 90%, were thus shown to be true to their new type. Whether the 10% reverting ones were truly atavists, or whether they were [478] only vicinists, caused by stray pollen grains from another culture, cannot of course be decided with sufficient certitude.

Here I might refer to the observations concerning the invisible dimorphous state of the flowers of the normal toad-flax. Individuals of the same type, when fertilized with each other, are nearly, but not absolutely, sterile. The yield of seeds of my peloric plants agrees fairly well with the harvest which I have obtained from some of the nearly sterile pairs of individuals in my former trial. Hence the suggestion is forced upon us that perhaps, owing to some unknown cause, all the peloric individuals of my experiment belonged to one and the same type, and were sterile for this reason only. If this is true, then it is to be presumed that all previous investigators have met the same condition, each having at hand only one of the two required types. And this discussion has the further advantage of showing the way, in which perhaps a full and constant race of peloric toad-flaxes may be obtained. Two individuals of different type are required to start from. They seem as yet never to have arisen from one group of mutations. But if it were possible to combine the products of two mutations obtained in different countries and under different conditions, there would be a chance [479] that they might belong to the supposed opposite types, and thus be fertile with one another. My peloric plants are still available, and the occurrence of this form elsewhere would give material for a successful experiment. The probability thereof is enhanced by the experience that my peloric plants bear large capsules and a rich harvest of seeds when fertilized from plants of the normal one-spurred race, while they remain nearly wholly barren by artificial fertilization with others. I suppose that they are infertile with the normal toad-flaxes of their own sexual disposition, but fertile with those of the opposite constitution. At all events the fact that they may bear abundant seed when properly pollinated is an indication of successful experiments on the possibility of gaining a hereditary race with exclusively peloric flowers. And such a race would be a distinct gain for sundry physiologic inquiries, and perhaps not wholly destitute of value from an horticultural point of view.

Returning now to the often recorded occurrence of peloric toad-flaxes in the wild state and recalling our discussion about the improbability of a dispersion from one locality to another by seed, and the probability of independent origin for most of these cases, we are confronted with the conception that a latent [480] tendency to mutation must be universally present in the whole species. Another observation, although it is of a negative character, gains in importance from this point of view. I refer to the total lack of intermediate steps between normal and peloric individuals. If such links had ordinarily been produced previous to the purely peloric state they would no doubt have been observed from time to time. This is so much the more probable as Linaria is a perennial herb, and the ancestors of a mutation might still be in a flowering condition together with their divergent offspring. But no such intermediates are on record. The peloric toad-flaxes are, as a rule, found surrounded by the normal type, but without intergrading forms. This discontinuity has already been insisted upon by Hofmeister and others, even at the time when the theory of descent was most under discussion, and any link would surely have been produced as a proof of a slow and continuous change. But no such proof has been found, and the conclusion seems admissible that the mutation of toad-flaxes ordinarily, if not universally, takes place by a sudden step. Our experiment may simply be considered as a thoroughly controlled instance of an often recurring phenomenon. It teaches us how, in the [481] main, the peloric mutations must be assumed to proceed.

This conception may still be broadened. We may include in it all similar occurrences, in allied and other species. There is hardly a limit to the possibilities which are opened up by this experience. But it will be well to refrain from hazardous theorizing, and consider only those cases which may be regarded as exact repetitions of the same phenomenon and of which our culture is one of the most recent instances on record. We will limit ourselves to the probable origin of peloric variations at large, of which little is known, but some evidence may be derived from the recorded facts. Only one case can be said to be directly analogous to our observations.

This refers to the peloric race of the common snapdragon, or Antirrhinum majus of our gardens. It is known to produce peloric races from time to time in the same way as does the toadflax. But the snapdragon is self-fertile and so is its peloric variety. Some cases are relatively old, and some of them have been recorded and in part observed by Darwin. Whence they have sprung and in what manner they were produced, seems never to have been noted. Others are of later origin, and among these one or two varieties have been accidentally produced [482] in the nursery of Mr. Chr. Lorenz in Erfurt, and are now for sale, the seeds being guaranteed to yield a large proportion of peloric individuals. The peloric form in this case appeared at once, but was not isolated, and was left free to visiting insects, which of course crossed it with the surrounding varieties. Without doubt the existence of two color-varieties of the peloric type, one of a very dark red, indicating the "Black prince" variety as the pollen-parent, and the other with a white tube of the corolla, recalling the form known as "Delila," is due to these crossings. I had last year (1903) a large lot of plants, partly normal and partly peloric, but evidently of hybrid origin, from seeds from this nursery, showing moreover all intermediate steps between nearly wholly peloric individuals and apparently normal ones. I have saved the seeds of the isolated types and before seeing the flowers of their offspring, nothing can be said about the purity and constancy of the type, when freed from hybrid admixtures. The peloric snapdragon has five small unequal spurs at the base of its long tube, and in this respect agrees with the peloric toad-flax.

Other pelories are terminal and quite regular, and occur in some species of Linaria, where I observed them in Linaria dalmatica. The [483] terminal flowers of many branches were large and beautifully peloric, bearing five long and equal spurs. About their origin and inheritance nothing is known.

A most curious terminal pelory is that of the common foxglove or Digitalis purpurea. As we have seen in a previous lecture, it is an old variety. It was described and figured for the first time by Vrolik of Amsterdam, and the original specimens of his plates are still to be seen in the collections of the botanic garden of that university. Since his time it has been propagated by seed as a commercial variety, and may be easily obtained. The terminal flower of the central stem and those of the branches only are affected, all other flowers being wholly normal. Almost always it is accompanied by other deviations, among which a marked increase of the number of the parts of the corolla and other whorls is the most striking. Likewise supernumerary petals on the outer side of the corolla, and a production of a bud in the center of the capsule may be often met with. This bud as a rule grows out after the fading away of the flower, bursting through the green carpels of the unripe fruit, and producing ordinarily a secondary raceme of flowers. This raceme is a weak but exact repetition of the first, bearing symmetrical foxgloves all [484] along and terminating in a peloric structure. On the branches these anomalies are more or less reduced, according to the strength of the branch, and conforming to the rule of periodicity, given in our lecture on the "five-leaved" clover. Through all this diminution the peloric type remains unchanged and therefore becomes so much the purer, the weaker the branches on which it stands.

I am not sure whether such peloric flowers have ever been purely pollinated and their seed saved separately, but I have often observed that the race comes pure from the seed of the zygomorphic flowers. It is as yet doubtful whether it is a half race or a double race, and whether it might be purified and strengthened by artificial selection. Perhaps the determination of the hereditary percentage described when dealing with the tricotyls might give the clue to the acquisition of a higher specialized race. The variety is old and widely disseminated, but must be subjected to quite a number of additional experiments before it can be said to be sufficiently understood.

The most widespread peloric variety is that of Gloxinia. It has erect instead of drooping flowers; and with the changed position the structure is also changed. Like other pelories it has five equal stamens instead of four unequal [485] ones, and a corolla with five equal segments instead of an upper and a lower lip. It shows the peloric condition in all of its flowers and is often combined with a small increase of the number of the parts of the whorls. It is for sale under the name of erecta, and may be had in a wide range of color-types. It seems to be quite constant from seed.

Many other instances of peloric flowers are on record. Indian cress or Tropaeolum majus loses the spur in some double varieties and with it most of its symmetrical structure; it seems to be considered justly as a peloric malformation. Other species produce such anomalies only from time to time and nothing is known about their hereditary tendency. One of the most curious instances is the terminal flower of the raceme of the common laburnum, which loses its whole papilionaceous character and becomes as regularly quinate as a common buttercup.

Some families are more liable to pelorism than others. Obviously all the groups, the flowers of which are not symmetrical, are to be excluded. But then we find that labiates and their allies among the dicotyledonous plants, and orchids among the monocotyledonous ones are especially subjected to this alteration. In both groups many genera and a long list of species [486] could be quoted as proof. The family of the labiates seems to be essentially rich in terminal pelories, as for instance in the wild sage or Salvia and the dead-nettle or Lamium. Here the pelories have long and straight corolla-tubes, which are terminated by a whorl of four or five segments. Such forms often occur in the wild state and seem to have a geographic distribution as narrowly circumscribed as in the case of many small species. Those of the labiates chiefly belong to southern Europe and are unknown at least in some parts of the other countries. On the contrary terminal pelories of Scrophularia nodosa are met with from time to time in Holland. Such facts clearly point to a common origin, and as only the terminal flowers are affected by the malformation, the fertility of the whole plant is evidently not seriously infringed upon.

Before leaving the labiates, we may cite a curious instance of pelorism in the toad-flax, which is quite different from the ordinary peloric variety. This latter may be considered from a morphologic standpoint to be owing to a five-fold repetition of the middle part of the underlip. This conception would at once explain the occurrence of five spurs and of the orange border all around the corolla-tube. We might readily imagine that any other of the five [487] parts of the corolla could be repeated five-fold, in which case there would be no spur, and no orange hue on the upper corolla-ring. Such forms really occur, though they seem to be more rare than the five-spurred pelories. Very little is known about their frequency and hereditary qualities.

Orchids include a large number of peloric monstrosities and moreover a wild pelory which is systematically described not only as a separate species but even as a new genus. It bears the name of Uropedium lindenii, and is so closely related to Cypripedium caudatum that many authors take it for the peloric variety of this plant. It occurs in the wild state in some parts of Mexico, where the Cypripedium also grows. Its claims to be a separate genus are lessened by the somewhat monstrous condition of the sexual organs, which are described as quite abnormal. But here also, intermediates are lacking, and this fact points to a sudden origin.

Many cases of pelorism afford promising material for further studies of experimental mutations. The peloric toad-flax is only the prototype of what may be expected in other cases. No opportunity should be lost to increase the as yet too scanty, evidence on this point.

[488]

LECTURE XVII

THE PRODUCTION OF DOUBLE FLOWERS

Mutations occur as often among cultivated plants as among those in the wild state. Garden flowers are known to vary markedly. Much of their variability, however, is due to hybridism, and the combination of characters previously separate has a value for the breeder nearly equal the production of really new qualities. Nevertheless there is no doubt that some new characters appear from time to time.

In a previous lecture we have seen that varietal characters have many features in common. One of them is their frequent recurrence both in the same and in other, often very distantly related, species. This recurrence is an important factor in the choice of the material for an experimental investigation of the nature of mutations.

Some varieties are reputed to occur more often and more readily than others. White-colored varieties, though so very common, seem for the most part to be of ancient date, but only few [489] have a known origin, however. Without any doubt many of them have been found in a wild state and were introduced into culture. On the other hand double flowers are exceedingly rare in the wild state, and even a slight indication of a tendency towards doubling, the stray petaloid stamens, are only rarely observed growing wild. In cultivation, however, double flowers are of frequent occurrence; hence the conclusion that they have been produced in gardens and nurseries more frequently than perhaps any other type of variety.

In the beginning of my experimental work I cherished the hope of being able to produce a white variety. My experiments, however, have not been successful, and so I have given them up temporarily. Much better chances for a new double variety seemed to exist, and my endeavors in this direction have finally been crowned with success.

For this reason I propose to deal now with the production of double flowers, to inquire what is on record about them in horticultural literature, and to give a full description of the origin thereof in an instance which it was my good fortune to observe in my garden.

Of course the historical part is only a hasty survey of the question and will only give such evidence as may enable us to get an idea of the [490] chances of success for the experimental worker. In the second half of the seventeenth century (1671), my countryman, Abraham Munting, published a large book on garden plants with many beautiful figures. It is called "Waare Oeffeninge der Planters," or "True Exercises With Plants." The descriptions pertain to ordinary typical species in greater part, but garden varieties receive special attention. Among these a long list of double flowers are to be seen. Double varieties of poppies, liverleaf (Hepatica), wallflowers (Cheiranthus), violets, Caltha, Althaea, Colchicum, and periwinkles (Vinca), and a great many other common flowers were already in cultivation at that time.

Other double forms have been since added. Many have been introduced from Japan, especially the Japanese marigold, Chrysanthemum indicum. Others have been derived from Mexico, as for instance the double zinnias. The single dahlias only seem to have been originally known to the inhabitants of Mexico. They were introduced into Spain at about 1789, and the first double ones were produced in Louvain, Belgium, in 1814. The method of their origin has not been described, and probably escaped the originators themselves. But in historical records we find the curious statement that it took place after three years' work. This indicates [491] a distinct plan, and the possibility of carrying it to a practical conclusion within a few years' time.

Something more is known about other cases. Garden anemones, Anemone coronaria, are said to have become double in the first half of the last century in an English nursery. The owner, Williamson, observing in his beds a flower with a single broadened stamen, saved its seeds separately, and in the next generations procured beautifully filled flowers. These he afterwards had crossed by bees with a number of colored varieties, and in this way succeeded in producing many new double types of anemone.

The first double petunia is known to have suddenly and accidentally arisen from ordinary seed in a private garden at Lyons about 1855. From this one plant all double races and-varieties have been derived by natural and partly by artificial crosses. Carriere, who reported this fact, added that likewise other species were known at that time to produce new double varieties rapidly. The double fuchsias originated about the same time (1854) and ten years later the range of double varieties of this plant had become so large that Carriere found it impossible to enumerate all of them.

Double carnations seem to be relatively old, double corn-flowers and double blue-bells being [492] of a later period. A long list could easily be made, to show that during the whole history of horticulture double varieties have arisen from time to time. As far as we can judge, such appearances have been isolated and sudden. Sometimes they sprang into existence in the full display of their beauty, but most commonly they showed themselves for the first time, exhibiting only spare supernumerary petals. Whenever such sports were worked up, a few years sufficed to reach the entire development of the new varietal attribute.

From this superficial survey of historical facts, the inference is forced upon us that the chance of producing a new double variety is good enough to justify the attempt. It has frequently succeeded for practical purposes, why should it not succeed as well for purely scientific investigation? At all events the type recommends itself to the student of nature, both on account of its frequency, and of the apparent insignificance of the first step, combined with the possibility of rapidly working up from this small beginning of one superfluous petal towards the highest degree of duplication.

Compared with the tedious experimental production of the peloric toad-flax, the attempt to produce a double flower has a distinct attraction. The peloric toad-flax is nothing new; the [493] experiment was only a repetition of what presumably takes place often within the same species. To attempt to produce a double variety we may choose any species, and of course should select one which as yet has not been known to produce double flowers. By doing so we will, if we succeed, produce something new. Of course, it does not matter whether the new variety has an horticultural interest or not, and it seems preferable to choose a wild or little cultivated species, to be quite sure that the variety in question is not already in existence. Finally the prospect of success seems to be enhanced if a species is chosen, the nearest allies of which are known to have produced double flowers.

For these reasons and others I chose for my experiment the corn-marigold, or Chrysanthemum segetum. It is also called the golden cornflower. In the wheat and rye fields of central Europe it associates with the blue-bottle or blue corn-flower. It is sometimes cultivated and the seeds are offered for sale by many nurserymen. It has a cultivated variety, called grandiflorum, which is esteemed for its brilliancy and long succession of golden bloom. This variety has larger flower-heads, surrounded with a fuller border of ray-florets. The species belongs to a genus many species of which have produced [494] double varieties. One of them is the Japanese marigold, others are the carinatum and the imbricatum species. Nearly allied are quite a number of garden-plants with double flower-heads, among which are the double camomiles.

My attention was first drawn to the structure of the heads and especially to the number of the ray-florets of the corn-marigold. The species appertains to that group of composites which have a head of small tubular florets surrounded by a broad border of rays. These rays, when counted, are observed to occur in definite numbers, which are connected with each other by a formula, known as "the series" of Braun and Schimper. In this formula, which commences with 1 and 2, each number is equal to the sum of the two foregoing figures. Thus 5, 8 and 13 are very frequent occurrences, and the following number, 21, is a most general one for apparently full rays, such as in daisies, camomiles, Arnica and many other wild and cultivated species.

These numbers are not at all constant. They are only the averages, around which the real numbers fluctuate. There may even be an overlapping of the extremes, since the fluctuation around 13 may even go beyond 8 and 21, and so on. But such extremes are only found in stray flowers, occurring on the same [495] individuals with the lesser degrees of deviation.

Now the marigold averages 13, and the grandiflorum 21 rays. The wild species is pure in this respect, but the garden-variety is not. The seeds which are offered for sale usually contain a mixture of both forms and their hybrids. So I had to isolate the pure types from this mixture and to ascertain their constancy and mutual independency. To this end I isolated from the mixture first the 13-rayed, and afterwards the 21-rayed types. As the marigolds are not sufficiently self-fertile, and are not easily pollinated artificially, it seemed impossible to carry on these two experiments at the same time and in the same garden. I devoted the first three years to the lower form, isolated some individuals with 12-13 rays out of the mixture of 1892 and counted the ray-florets on the terminal head of every plant of the ensuing generation next year. I cultivated and counted in this way above 150 individuals and found an average of exactly 13 with comparatively few individuals displaying 14 or only 12 rays, and with the remainder of the plants grouped symmetrically around this average. I continued the experiment for still another year and found the same group of figures. I was then satisfied as to the purity of the isolated strain. Next year I sowed a new mixture in [496] order to isolate the reputed pure grandiflorum type. During the beginning of the flowering period I ruthlessly threw away all plants displaying less than 21 rays in the first or terminal head. But this selection was not to be considered as complete, because the 13-rayed race may eventually transgress its boundary and come over to the 21 and more. This made a second selection necessary. On the selected plants all the secondary heads were inspected and their ray-florets counted. Some individuals showed an average of about 13 and were destroyed. Others gave doubtful figures and were likewise eliminated, and only 6 out of a lot of nearly 300 flowering plants reached an average of 21 for all of the flowers.