Chapter 13

Obviously, this elimination conduces only to a partial purification. The conspicuous plants will be destroyed, but a greater number of hybrids will remain, still concealed by their resemblance to the general type and will be spared to repeat the same process next year. So while the variety may be freed every year from the impurities brought into it in the preceeding summer, the admixtures of the species [212] will continue during a number of years, and it may not be possible to get rid of them at all.

It is an often recurring assertion that white varieties of colored species are the most stable of all horticultural races. They are often said to be at least as constant as the species itself, and even to surpass it in this quality. With our present state of knowledge, the explanation of this general experience is easily given. For selection removes the effect of spontaneous crosses from the variety in each year, and renders it practically pure, while it is wholly inadequate to produce the same effects on the species, because of the concealed hybrids.

The explanation given in this simple instance may be applied to the case of different varieties of the same species, when growing together and crossed naturally by insects.

It would take too long to go into all the details that present themselves here to the student of nature and of gardens. I will only state, that since varieties differ principally from their species by the lack of some sharp character, one variety may be characterized by the lack of color of the flowers, another by the lack of pubescence, a third by being dwarfed, and so on. Every character must be studied separately in its effects on the offspring [213] of the crosses. And it is therefore easily seen, that the hybrids of two varieties may resemble neither of them, but revert to the species itself. This is necessarily and commonly the case, since it is always the older or positive characters that prevail in the hybrids and the younger or negative that lie hidden. So for instance, a blue dwarf larkspur, crossed with a tall white variety, must give a tall blue hybrid, reassuming in both characters the essentials of the species.

Keeping this rule in view, it will be easy to calculate what may be expected from spontaneous crosses for a wide range of occurrences, and thus to find an explanation of innumerable cases of apparent variability and reversion in the principle of vicinism. Students have only to recollect that specific characters prevail over varietal ones, and that every character competes only with its own antagonist. Or to give a sharper distinction: whiteness of flowers cannot be expected to be interchanged with pubescence of leaves.

In concluding I will point out another danger which in the principle of vicinism may be avoided. If you see a plant in a garden with all the characteristics of its species, how can you be sure that it is truly a representative of the species, and not a hybrid? The prevailing [214] characters are in either case the same. Perhaps on close inspection you may find in some cases a slight difference, some character being not as fully developed in the hybrid as in the species. But when such is not the case, or where the opportunity for such a closer examination is wanting, a hybrid may easily be taken for a specimen of the pure race. Now take the seeds of your plant and sow them. If you had not supposed it to be hybrid you will be astonished at finding among its progeny some of a wholly different type. You will be led to conclude that you are observing a sudden change in structure such as is usually called a sport.

Or in other words you may think that you are assisting at the origination of a new variety. If you are familiar with the principle of vicinism, you will refrain from such an inference and consider the supposition of a hybrid origin. But in former times, when this principle was still unknown and not even guessed at, it is evident that many mistakes must have been made, and that many an instance, which until now has been considered reliable proof of a so-called single variation, is in fact only a case of vicinism. In reading the sparse literature on sports, numerous cases will be found, which cannot stand this test. In many instances crossing must be looked to as an explanation, [215] and in other cases the evidence relied upon does not suffice to exclude this assumption. Many an old argument has of late lost its force by this test.

Returning to our starting point we may now state that regular reversions to a specific type characterize a form as a variety of that species. These reversions, however, are not due to an innate tendency, but to unobserved spontaneous crosses.

[217]

LECTURE VIII

LATENT CHARACTERS

No organism exhibits all of its qualities at any one time. Many of them are generally dormant and await a period of activity. For some of them this period comes regularly, while in others the awakening depends upon external influences, and consequently occurs very irregularly. Those of the first group correspond to the differences in age; the second constitute the responses of the plant to stimuli including wound-injuries.

Some illustrative examples may be quoted in order to give a precise idea of this general conception of dormant or latent characters. Seed leaves are only developed in the seed and the seedling; afterwards, during the entire lifetime of the plant, the faculty of producing them is not made use of. Every new generation of seeds however, bears the same kind of seed leaves, and hence it is manifest that it is the same quality, which shows itself from time to time.

The primary leaves, following the seed-leaves, are different in many species, from the later ones, and the difference is extremely pronounced in some cases of reduction. Often, when leaves are lacking in the adult plant, being replaced by flattened stalks as in the case of the acacias, or by thorns, or green stems and twigs as in the prickly broom or _Ulex europaeus_, the first leaves of the young plant may be more highly differentiated, being pinnate in the first case and bearing three leaflets in the second instance. This curious behavior which is very common, brings the plants, when young, nearer to their allies than in the adult state, and manifestly implies that the more perfect state of the leaves is latent throughout the life of the plant, with the exception of the early juvenile period.

_Eucalyptus Globulus_, the Australian gum tree, has opposite and broadly sessile leaves during the first years of its life. Later these disappear and are replaced by long sickle-shaped foliage organs, which seem to be scattered irregularly along the branches. The juvenile characters manifestly lie dormant during the adult period, and that this is so, may be shown artificially by cutting off the whole crown of the tree, when the stem responds by producing numerous new branches, which assume the [218] shape proper to the young trees, bearing sessile and opposite leaves.

It seems quite unnecessary to give further instances. They are familiar to every student. It is almost safe to say that every character has its periods of activity and of inactivity, and numbers of flowers and fruits can be mentioned as illustrations. One fact may be added to show that nearly every part of the plant must have the power of producing all or nearly all the characters of the individual to which it belongs. This proof is given by the formation of adventitious buds. These, when once formed, may grow out into twigs, with leaves and flowers and roots. They may even be separated from the plants and used as cuttings to reproduce the whole. Hence we may conclude that all tissues, which possess the power of producing adventitious buds, must conceal in a latent state, all the numerous characters required for the full development of the whole individual.

Adventitious buds may proceed from specialized cells, as on the margin of the leaves of _Bryophyllum calycinum_; or from the cells of special tissues, as in the epidermis of the begonias; or they may be provoked by wounds in nearly every part of the plant, provided it be able to heal the wound by swelling tissues or [219] callus. The best instance is afforded by elms and by the horse-chestnut. If the whole tree is hewn down the trunk tries to repair the injury by producing small granulations of tissue between the wood and the bark, which gradually coalesce while becoming larger. From this new ring of living matter innumerable buds arise, that expand into leafy branches, showing clearly that the old trunk possesses, in a latent state, all the qualities of the whole crown. Indeed, such injured stumps may be used for the production of copses and hedges.

All the hitherto recorded cases of latency have this in common, that they may become active during the life-time of any given individual once, or oftener. This may be called the ordinary type of latency.

Besides this there is another form of latent characters, in which this awakening power is extremely limited, or wholly absent. It is the systematic latency, which may be said to belong to species and varieties in the same way as the ordinary latency belongs to individuals. As this individual latency may show itself from time to time during the life of a given plant, the first may only become active from time to time during the whole existence of the variety or the species. It has no regular period of activity, nor may it be incited by artificial stimulation.

[220] It emerges from concealment only very rarely and only on its own initiative. Such instances of atavism have been described in previous lectures, and their existence has been proved beyond doubt.

Systematic latency explains the innumerable instances in which species are seen to lack definite characteristics which ordinarily do not fail, either in plants at large, or in the group or family to which the plant belongs. If we take for instance the broom-rape or _Orobanche_, or some other pale parasite, we explain their occurrence in families of plants with green leaves, by the loss of the leaves and of the green color. But evidently this loss is not a true one, but only the latency of those characters. And even this latency is not a complete one, as little scales remind us of the leaves, and traces of chlorophyll still exist in the tissues. Numerous other cases will present themselves to every practical botanist.

Taking for granted that characters, having once been acquired, may become latent, and that this process is of universal occurrence throughout the whole vegetable and animal kingdom, we may now come to a more precise and clear conception of the existing differences between species and varieties.

For this purpose we must take a somewhat [221] broader view of the whole evolution of the vegetable kingdom. It is manifest that highly developed plants have a larger number of characters than the lower groups. These must have been acquired in some way, during preceding times. Such evolution must evidently be called a process of improvement, or a progressive evolution. Contrasted to this is the loss, or the latency of characters, and this may be designated retrogressive or retrograde evolution. But there is still a third possibility. For a latent character may reassume its activity, return to the active state, and become once more an important part of the whole organization. This process may be designated as degressive evolution; it obviously completes the series of the general types of evolution.

Advancement in general in living nature depends on progressive evolution. In different parts of the vegetable kingdom, and even in different families this progression takes place on different lines. By this means it results in an ever increasing divergency between the several groups. Every step is an advance, and many a step must have been taken to produce flowering plants from the simplest unicellular algae.

But related to, and very intimately connected with this advancement is the retrogressive [222] evolution. It is equally universal, perhaps never failing. No great changes have been attained, without acquiring new qualities on one side, and reducing others to latency. Everywhere such retrogressions may be seen. The polypetalous genera _Pyrola_, _Ledum_, and _Monotropa_ among the sympetalous heaths, are a remarkable instance of this. The whole evolution of the monocotyledons from the lowest orders of dicotyledons implies the seeming loss of cambial growth and many other qualities. In the order of aroids, from the calamus-root or sweet flag, with its small but complete flowers, up to the reduced duckweeds (_Lemna_), almost an unbroken line of intermediate steps may be traced showing everywhere the concurrence of progressive and retrogressive evolution.

Degressive evolution is not so common by far, and is not so easy to recognize, but no doubt it occurs very frequently. It is generally called atavism, or better, systematic atavism, and the clearest cases are those in which a quality which is latent in the greater part of a family or group, becomes manifest in one of its members. Bracts in the inflorescence of crucifers are ordinarily wanting, but may be seen in some genera, _Erucastrum pollichii_ being perhaps the [223] most widely known instance, although other cases might easily be cited.

For our special purpose we may take up only the more simple cases that may be available for experimental work. The great lines of evolution of whole families and even of genera and of many larger species obviously lie outside the limits of experimental observation. They are the outcome of the history of the ancestors of the present types, and a repetition of their history is far beyond human powers. We must limit ourselves to the most recent steps, to the consideration of the smallest differences. But it is obvious that these may be included under the same heads as the larger and older ones. For the larger movements are manifestly to be considered only as groups of smaller steps, going in the same direction.

Hence we conclude, that even the smallest steps in the evolution of plants which we are able to observe, may be divided into progressive, retrogressive and degressive ones. The acquisition of a single new quality is the most simple step in the progressive line, the becoming latent and the reactivating of this same quality are the prototypes of the two other classes.

Having taken this theoretical point of view, it remains to inquire, how it concurs with the [224] various facts, given in former lectures and how it may be of use in our further discussions.

It is obvious that the differences between elementary species and varieties on the one hand, and between the positive and negative varieties as distinguished above, are quite comparable with our theoretical views. For we have seen that varieties can always be considered as having originated by an apparent loss of some quality of the species, or by the resumption of a quality which in allied species is present and visible. In our exposition of the facts we have of course limited ourselves to the observable features of the phenomena without searching for a further explanation. For a more competent inquiry however, and for an understanding of wider ranges of facts, it is necessary to penetrate deeper into the true nature of the implied causes.

Therefore we must try to show that elementary species are distinguished from each other by the acquisition of new qualities, and that varieties are derived from their species either by the reduction of one or more characteristics to the latent state, or by the energizing of dormant characters.

Here we meet with a great difficulty. Hitherto varieties and subspecies have never been clearly defined, or when they have been, it was [225] not by physiological, but only by morphological research. And the claims of these two great lines of inquiry are obviously very diverging. Morphological or comparative studies need a material standard, by which it may be readily decided whether certain groups of animals and plants are to be described or de-nominated as species, as subspecies or as varieties. To get at the inner nature of the differences is in most cases impossible, but a decision must be made. The physiological line of inquiry has more time at its disposal; it calls for no haste. Its experiments ordinarily cover years, and a conclusion is only to be reached after long and often weary trials. There is no making a decision on any matter until all doubtful points have been cleared up. Of course, large groups of facts remain uncertain, awaiting a closer inquiry, and the teacher is constrained to rely on the few known instances of thoroughly investigated cases. These alone are safe guides, and it seems far better to trust to them and to make use of them for the construction of sharp conceptions, which may help us to point out the lines of inquiry which are still open.

Leaving aside all such divisions and definitions, as were stamped with the name of provisional species and varieties by the great systematist, [226] Alphonse De Candolle, we may now try to give the proofs of our assertion, by using only those instances that have been thoroughly tested in every way.

We may at once proceed to the retrogressive or negative varieties. The arguments for the assumption that elementary species owe their origin to the acquisition of new qualities may well be left for later lectures when we shall deal with the experimental proofs in this matter.

There are three larger groups of facts, on which the assumption of latent characters in ordinary varieties rests. These are true atavism, incomplete loss of characters, and systematic affinity. Before dealing with each of these separately, it may be as well to recall once more that in former lectures we have treated the apparent losses only as modifications in a negative way, without contemplating the underlying causes.

Let us recall the cases of bud-atavism given by the whitish variety of the scarlet _Ribes_, by peaches and nectarines, and by conifers, including _Cephalotaxus_ and _Cryptomeria_. These and many other analogous facts go to prove the relation of the variety to the species. Two assumptions are allowable. In one the variety differs from the species by the total loss of the [227] distinctive character. In the other this character is simply reduced to an inactive or dormant state. The fact of its recurrence from time to time, accompanied by secondary characters previously exhibited, is a manifest proof of the existence of some relation between the lost and the resumed peculiarity. Evidently this relation cannot be accounted for on the assumption of an absolute disappearance; something must remain from which the old features may be restored.

This lengthy discussion may be closed by the citation of the cases, in which plants not only show developmental features of a former state, but also reproduce the special features they formerly had, but seemingly have lost. Two good illustrative examples may be given. One is afforded by the wheat-ear carnation, the other by the green dahlias, and both have occurred of late in my own cultures.

A very curious anomaly may from time to time be observed in large beds of carnations. It bears no flowers, but instead of them small green ears, which recall the ears of wheat. Thence the name of "Wheat-ear" carnation. On closer inspection it is easily seen how they originate. The normal flowers of the carnations are preceded by a small group of bracts, [228] which are arranged in opposite pairs and therefore constitute four rows.

In this variety the flower is suppressed and this loss is attended by a corresponding increase of the number of the pairs of bracts. This malformation results in square spikes or somewhat elongated heads consisting only of the greenish bracts. As there are no flowers, the variety is quite sterile, and as it is not regarded by horticulturists as an improvement on the ordinary bright carnations, it is seldom multiplied by layering. Notwithstanding this, it appears from time to time and has been seen in different countries and at different periods, and, what is of great importance for us, in different strains of carnations. Though sterile, and obviously dying out as often as it springs into existence, it is nearly two centuries old. It was described in the beginning of the 18th century by Volckamer, and afterwards by Jaeger, De Candolle, Weber, Masters, Magnus and many other botanists. I have had it twice, at different times and from different growers.

So far as I have been able to ascertain reversions of this curious carnation to normal flowers have not yet been recorded. Such a modification occurred last summer in my garden on a plant which had not been divided or layered, but on which the slender branches had [229] been left on the stem. Some of them remained true to the varietal type and bore only green spikes. Others reverted wholly or partially to the production of normal flowers. Some branches bore these only, others had spikes and flowers on neighboring twigs, and in still other instances little spikes had been modified in such manner that a more or less well developed flower was preceded by some part of an ear.

The proof that this retrograde modification was due to the existence of a character in the latent state was given by the color of the flowers. If the reverted bud had only lost the power of producing spikes, they would evidently simply have returned to the characteristics of the ordinary species, and their color would have been a pale pink. Instead of this, all flowers displayed corollas of a deep brown. They obviously reverted to their special progenitor, the chance variety from which they had sprung, and not to the common prototype of the species. Of course it was not possible to ascertain from which variety the plant had really originated, but the reproduction of any one clearly defined varietal mark is in itself proof enough of their origin, and of the latency of the dark brown flower-color in this special case.

A still better proof is afforded by a new type of green dahlia. The ordinary green dahlia [230] has large tufts of green bracts instead of flowering heads, the scales of the receptacle having assumed the texture and venation of leaves, and being in some measure as fleshy. But the green heads retain the form of the ordinary flower-heads, and as they have no real florets that may fade away, they remain unchanged on the plants, and increase in number through the whole summer. The new types of green dahlia however, with which I have now to deal, are distinguished by the elongation of the axis of the head, which is thereby changed into a long leafy stalk, attaining a length of several inches. These stalks continue growing for a very long time, and for the most part die without producing anything else than green fleshy scales.

This long-headed green dahlia originated at Haarlem some years ago, in the nursery of Messrs. Zocher & Co. It was seen to arise twice, from different varieties. Both of these were double-flowered, one a deep carmine with white tips on the rays, the other of a pale orange tint, known by the name of "Surprise." As they did not bear any florets or seeds, they were quite sterile. The strain arising from the carmine variety was kindly given to me by Messrs. Zocher & Co., and was propagated in my garden, while the other was kept in the nursery. In the earlier cultures both remained true to [231] their types, never producing true florets. No mark of the original difference was to be seen between them. But last summer (1903) both reverted to their prototypes, bearing relatively large numbers of ordinary double flowerheads among the great mass of green stalks. Some intermediate forms also occurred consisting of green-scaled stalks ending in small heads with colored florets.