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.

Thus far we have an ordinary case of reversion. But the important side of the phenomenon was, that each plant exactly "recollected" from which parent it had sprung. All of those in my garden reverted to the carmine florets with white tips, and all of those in the nursery to the pale orange color and the other characteristics of the "Surprise" variety.

It seems absolutely evident, that no simple loss can account for this difference. Something of the character of the parent-varieties must have remained in the plant. And whatever conception we may formulate of these vestigial characters it is clear that the simplest and most obvious idea is their preservation in a dormant or latent state. Assuming that the distinguishing marks have only become inactive by virescence, it is manifest that on returning each will show its own peculiarities, as recorded above. Our second point was the incomplete loss of [232] the distinguishing quality in some varieties. It is of general occurrence, though often overlooked. Many white varieties of colored flowers give striking instances, among them many of the most stable and most prized garden-flowers. If you look at them separately or in little bouquets they seem to be of irreproachable purity. But if you examine large beds a pale hue will become visible. In many cases this tinge is so slight as to be only noticeable in a certain illumination, or by looking in an oblique direction across the bed; in others it is at once evident as soon as it has been pointed out. It always reminds the observer of the color of the species to which the variety belongs, being bluish in violets and harebells, reddish in godetias and phloxes, in Silene Armeria and many others. It proves that the original color quality of the species has not wholly, but only partly disappeared. It is dormant, but not entirely obliterated; latent, but not totally concealed; inactive, but only partially so. Our terminology is an awkward one; it practically assumes, as it so often does in other cases, a conventional understanding, not exactly corresponding to the simple meaning of the words. But it would be cumbrous to speak always of partial inactivity, incomplete latency or half awakening qualities. Even such words as sub-latent, [233] which would about express the real state of things, would have little chance of coming into general use.

Such sub-latent colors are often seen on special parts in white varieties of flowers. In many cases it is the outer side of the petals which recalls the specific color, as in some white roses. In violets it is often on the spur that the remains of the original pigment are to be seen. In many instances it is on the tips of the petals or of the segments of the corolla, and a large number of white or yellow flowers betray their affinity to colored species by becoming red or bluish at the edges or on the outer side.

The reality of such very slight hues, and their relation to the original pigment of the species may in some cases be proved by direct experiment. If it is granted that latency is not an absolute quality, then it will be readily accepted, that even latency must be subjected to the laws of gradual variation or fluctuating variability. We will deal with these laws in a later lecture but every one knows that greater deviations than the ordinary may be attained by sowing very large numbers and by selecting from among them the extreme individuals and sowing anew from their seed. In this way the slightest tinge of any latent color may be [234] strengthened, not indeed to the restoration of the tinge of the species, but at least so far as to leave no doubt as to the identity of the visible color of the species and the latent or sublatent one of the variety.

I made such an experiment with the peach leaved harebell or Campanula persicifolia. The white variety of this species, which is often met with in our gardens, shows a very pale bluish hue when cultivated in large quantities, which however is subject to individual variations. I selected some plants with a decided tinge, flowered them separately, sowed their seeds, and repeated this during two generations. The result was an increase of the color on the tips of the segments of the corolla in a few individuals, most of them remaining as purely white as the original strain. But in those few plants the color was very manifest, individually variable in degree, but always of the same blue as in the species itself.

Many other instances could be given. Smooth varieties are seldom absolutely so, and if scattering hairs are found on the leaves or only on some more or less concealed parts, they correspond in their character to those of the species. So it is with prickles, and even the thornless thorn-apple has fruits with surfaces far from smooth. The thornless horse-chestnut [235] has in some instances such evident protuberances on the valves of its fruits, that it may seem doubtful whether it is a pure and stable variety.

Systematic latency may betray itself in different ways, either by normal systematic marks, or by atavism. With the latter I shall deal at length on another occasion, and therefore I will give here only one very clear and beautiful example. It is afforded by the common red clover. Obviously the clovers, with their three leaflets in each leaf, stand in the midst of the great family of papilionaceous plants, the leaves of which are generally pinnate. Systematic affinity suggests that the "three leaved" forms must have been derived from pinnate ancestors, evidently by the reduction of the number of the leaflets. In some species of clover the middle of the three is more or less stalked, as is ordinarily the case in pinnate leaves; in others it is as sessile as are its neighbors. In a subsequent chapter I will describe a very fine variety, which sometimes occurs in the wild state and may easily be isolated and cultivated. It is an ordinary red clover with five leaflets instead of three, and with this number varying between three and seven, instead of being nearly wholly stable as in the common form. It produces from time to time pinnate leaves, [236] very few indeed, and only rarely, but then often two or three or even more on the same individual. Intermediate stages are not wanting, but are of no consequence here. The pinnate leaves obviously constitute a reversion to some prototype, to some ancestor with ordinary papilionaceous leaves. They give proof of the presence of the common character of the family, concealed here in a latent state. Any other explanation of this curious anomaly would evidently be artificial. On the other hand nothing is really known about the ancestors of clover, and the whole conception rests only on the prevailing views of the systematic relationships in this family. But, as I have already said, further proof must be left for a subsequent occasion.

Many instances, noted in our former lectures, could be quoted here. The systematic distribution of rayed and rayless species and varieties among the daisy-group of the composites affords a long series of examples. Accidental variations in both directions occur. The Canada fleabane or Erigeron canadensis, the tansy or Tanacetum vulgare and some others may at times be seen with ray-florets, and according to Murr, they may sometimes be wanting in Aster Tripolium, Bellis perennis, some species of Anthemis, Arnica montana and in a number [237] of other well-known rayed species. Another instance may be quoted; it has been pointed out by Grant Allen, and refers to the dead-nettle or Lamium album. Systematically placed in a genus with red-flowering species, we may regard its white color as due to the latency of the general red pigment.

But if the flower of this plant is carefully examined, it will be found in most cases not to be purely white, but to have some dusky lines and markings on its lower lip. Similar devices are observed on the lip of the allied Lamium maculatum, and in a less degree on the somewhat distant Lamium purpureum. With Lamium maculatum or spotted dead-nettle, the affinity is so close that even Bentham united the two in a single species, considering the ordinary dead-nettle only as a variety of the dappled purple type. For the support of this conception of a specific or varietal retrograde change many other facts are afforded by the distribution of the characteristic color and of the several patterns of the lips of other labiates, and our general understanding of the relationships of the species and genera in this family may in a broad sense be based on the comparison of these seemingly subordinate characteristics.

The same holds good in many other cases, and systematists have often become uncertain [238] as to the true value of some form, by its relationship to the allied types in the way of retrogressive modification. Color-differences are so showy, that they easily overshadow other characters. The white and the blue thorn-apple, the white and the red campion (Lychnis vespertina and diurna) and many other illustrative cases could be given, in which two forms are specifically separated by some authors, but combined by others on the ground of the retrograde nature of some differentiating mark.

Hitherto we have dealt with negative characters and tried to prove that the conception of latency of the opposite positive characteristics is a more natural explanation of the phenomenon than the idea of a complete loss. We have now to consider the positive varieties, and to show that it is quite improbable that here the species have struck out for themselves a wholly new character. In some instances such may have been the case, but then I should prefer to treat these rather as elementary species. But in the main we will have to assume the latency of the character in the species and its reassumption by the variety when originating, as the most probable explanation.

Great stress is laid upon this conception by the fact, that positive varieties are so excessively rare when compared with the common occurrence [239] of negative ones. Indeed, if we put aside the radiate and the color-varieties of flowers and foliage, hardly any cases can be cited. We have dealt with this question in a former lecture, and may now limit ourselves to the positive color-varieties.

The latency of the faculty of producing the red pigment in leaves must obviously be accepted for nearly the whole vegetable kingdom. Oaks and elms, the beautiful climbing species of Ampelopsis, many conifers, as for instance Cryptomeria japonica, some brambles, the Guelder-rose (Viburnum Opulus) and many other trees and shrubs assume a more or less bright red color in the fall. During summer this tendency must have been dormant, and that this is so, is shown by the young leaves of oaks and others, which, when unfolding in the spring show a similar but paler hue. Moreover, there is a way of awakening the concealed powers at any time. We have only to inflict small wounds on the leaves, or to cut through the nerves or to injure them by a slight bruising, and the leaves frequently respond with an intense reddening of the living tissues around and especially above the wounds. Azolla caroliniana, a minute mosslike floating plant allied to the ferns, responds to light and cold with a reddish tinge, and to shade or warmth with a pure green. The foliage [240] of many other plants behaves likewise, as also do apples and peaches on the insolated sides of the fruits. It is quite impossible to state these groups of facts in a more simple way than by the statement that the tendency to become red is almost generally present, though latent in leaves and stems, and that it comes into activity whenever a stimulus provokes it.

Now it must be granted that the energizing of such a propensity under ordinary circumstances is quite another thing from the origination of a positive variety by the evolution of the same character. In the variety the activity has become independent of outer influences or dependent upon them in a far lesser degree. The power of producing the red pigments is shown to be latent by the facts given above, and we see that in the variety it is no longer latent but is in perfect and lasting activity throughout the whole life of the plant.

Red varieties of white flowers are much more rare. Here the latency of the red pigment may be deduced partly from general arguments like those just given, partly from the special systematic relations in the given cases. Hildebrand has clearly worked out this mode of proof. He showed by the critical examination of a large number of instances that the occurrence of the red-flowered varieties is contingent upon the [241] existence of red species in the same genus, or in some rare cases, in nearly allied genera. Colors that are not systematically present in the group to which a white species belongs are only produced in its varieties in extremely rare cases.

We may quote some special rules, indicated by Hildebrand. Blue species are n the main very rare, and so are blue varieties of white species also. Carnations, Asiatic or cultivated buttercups (Ranunculus asiaticus), Mirabilis, poppies, Gladiolus, Dahlia, and some other highly cultivated or very old garden-plants have not been able to produce true blue flowers. But the garden-anemone (Anemone coronaria) has allies with very fine blue flowers. The common stock has bluish varieties and is allied to Aubretia and Hesperis, and gooseberries have a red form, recalling the ordinary currant. In nearly all other instances of blue or red varieties every botanist will be able to point out some allied red or blue species, as an indication of the probable source of the varietal character.

Dark spots on the lower parts of the petals of some plants afford another instance, as in poppies and in the allied Glaucium, where they sometimes occur as varietal and in other cases as specific marks.

The yellow fails in many highly developed [242] flowers, which are not liable to produce yellow variations, as in Salvia, Aster, Centaurea, Vinca, Polygala and many others. Even the rare pale yellowish species of some of these genera have no tendency in this direction. The hyacinths are the most remarkable, if not the sole known instance of a species having red and blue and white and yellow varieties, but here the yellow is not the bright golden color of the buttercups.

The existence of varietal colors in allied species obviously points to a common cause, and this cause can be no other than the latency of the pigment in the species that do not show it.

The conception of latency of characters as the common source of the origination of varieties, either in the positive or in the negative way, leads to some rules on variability, which are known under the names given to them by Darwin. They are the rules of repeated, homologous, parallel and analogous variability. Each of them is quite general, and may be recognized in instances from the most widely distant families. Each of them is quite evident and easily understood on the principle of latency.

By the term of repeated variability is meant the well-known phenomenon, that the same variety has sprung at different times and in different [243] countries from the same species. The repetition obviously indicates a common internal cause. The white varieties of blue- and red-flowered plants occur in the wild state so often, and in most of the instances in so few individuals that a common pedigree is absolutely improbable. In horticulture this tendency is widely and vexatiously known, since the repetition of an old variety does not bring any advantage to the breeder. The old name of "conquests," given by the breeders of hyacinths, tulips and other flower-bulbs to any novelty, in disregard of the common occurrence of repetitions, is an indication of the same experience in the repeated appearance of certain varieties.

The rule of parallel variations demands that the same character occasionally makes its appearance in the several varieties or races, descended from the same species, and even in widely distinct species. This is a rule, which is very important for the general conception of the meaning of the term variety as contrasted with elementary species. For the recurrence of the same deviation always impresses us as a varietal mark. Laciniated leaves are perhaps the most beautiful instance, since they occur in so many trees and shrubs, as the walnut tree, the beech, the birch, the hazelnut, and even in [244] brambles and some garden-varieties of the turnip (Brassica).

In such cases of parallel variations the single instances obviously follow the same rules and are therefore to be designated as analogous. Pitchers or ascidia, formed by the union of the margins of a leaf, are perhaps the best proof. They were classified by Morren under two heads, according to their formation from one or more leaves. Monophyllous pitchers obey the same law, viz.: that the upper side of the leaf has become the inner side of the pitcher. Only one exception to this rule is known to me. It is afforded by the pitchers of the banyan or holy fig-tree, Ficus religiosus, but it does not seem to belong to the same class as other pitchers, since as far as it has been possible to ascertain the facts, these pitchers are not formed by a few leaves as in all other cases, but by all the leaves of the tree.

In some cases pitchers are only built up of part of the leaf-blade. Such partial malformations obey a rule, that is common to them and to other foliar enations, viz.: that the side of the leaf from which they emerge, is always their outer side. The inner surface of these enations corresponds to the opposite side of the leaf, both in color and in anatomical structure. The last of the four rules above mentioned is [245] that of the homologous variability. It asserts that the same deviation may occur in different, but homologous parts of the same plant. We have already dealt with some instances, as the occurrence of the same pigment in the flowers and foliage, in the fruits and seeds of the same plant, as also illustrated by the loss of the red or blue tinge by flowers and berries. Other instances are afforded by the curious fact that the division of the leaves into numerous and small segments is repeated by the petals, as in the common celandine and some sorts of brambles.

It would take too long to make a closer examination of the numerous cases which afford proof of these statements. Suffice it to say that everywhere the results of close inspection point to the general rule, that the failure of definite qualities both in species and in varieties must, in a great number of cases, be considered as only apparent. Hidden from view, occasionally reappearing, or only imperfectly concealed, the same character must be assumed to be present though latent.

In the case of negative or retrogressive varieties it is the transition from the active into a dormant state to which is due the origin of the variety. Positive varieties on the contrary owe their origin to the presence of some character [246] in the species in the latent state, and to the occasional re-energizing thereof.

Specific or varietal latency is not the same thing as the ordinary latency of characters that only await their period of activity, or the external influence which will awake them. They are permanently latent, and could well be designated by the word perlatent. They spring into activity only by some sudden leap, and then at once become independent of ordinary external stimulation.



[247]

LECTURE IX

CROSSES OF SPECIES AND VARIETIES

In the foregoing lectures I have tried to show that there is a real difference between elementary species and varieties. The first are of equal rank, and together constitute the collective or systematic species. The latter are usually derived from real and still existing types. Elementary species are in a sense independent of each other, while varieties are of a derivative nature.

Furthermore I have tried to show that the ways in which elementary or minor species must have originated from their common ancestor must be quite different from the mode of origin of the varieties. We have assumed that the first come into existence by the production of something new, by the acquirement of a character hitherto unnoticed in the line of their ancestors. On the contrary, varieties, in most cases, evidently owe their origin to the loss of an already existing character, or in other less frequent cases, to the re-assumption of a quality [248] formerly lost. Some may originate in a negative, others in a positive manner, but in both cases nothing really new is acquired.

This distinction holds good for all cases in which the relationship between the forms in question is well known. It seems entirely justifiable therefore to apply it also to cases in which the systematic affinity is doubtful, as well as to instances in which it is impossible to arrive at any taxonomic conclusions. The extreme application of the principle would no doubt disturb the limits between many species and varieties as now recognized. It is not to be forgotten however that all taxonomic distinctions, which have not been confirmed by physiologic tests are only provisional, a view acknowledged by the best systematists. Of course the description of newly discovered forms can not await the results of physiologic inquiries; but it is absolutely impossible to reach definite conclusions on purely morphologic evidence. This is well illustrated by the numerous discords of opinion of different authors on the systematic worth of many forms.

Assuming the above mentioned principle as established, and disregarding doubtful cases as indicated, the term progressive evolution is used to designate the method in which elementary species must have originated. It is the [249] manner in which all advance in the animal and vegetable kingdoms must have taken place, continuously adding new characters to the already existing number. Contrasted with this method of growing differentiation, are the retrogressive modifications, which simply retrace a step, and the degressive changes in which a backward step is retraced and old characters revived. No doubt both of these methods have been operative on a large scale, but they are evidently not in the line of general advancement.

In all of these directions we see that the differentiating marks show more or less clearly that they are built up of units. Allied forms are separated from each other without intermediates. Transitions are wholly wanting, although fallaciously apparent in some instances owing to the wide range of fluctuating variability of the forms concerned, or to the occurrence of hybrids and subvarieties.

These physiologic units, which in the end must be the basis for the distinction of the systematic units, may best be designated by the term of "unit-characters." Their internal nature is as yet unknown to us, and we will not now look into the theories, which have been propounded as to the probable material basis underlying them. For our present purpose the empirical evidence of the general occurrence of [250] sharp limits between nearly related characters must suffice. As Bateson has put it, species are discontinuous, and we must assume that their characters are discontinuous also.

Moreover there is as yet no reason for trying to make a complete analysis of all the characters of a plant. No doubt, if attained, such an analysis would give us a deep insight into the real internal construction of the intricate properties of organisms in general. But taxonomic studies in this direction are only in their infancy and do not give us the material required for such an analysis. Quite on the contrary, they compel us to confine our study to the most recently acquired, or youngest characters, which constitute the differentiating marks between nearly allied forms.

Obviously this is especially the case in the realm of the hybrids, since only nearly related forms are able to give hybrid offspring. In dealing with this subject we must leave aside all questions concerning more remote relationships.

It is not my purpose to treat of the doctrine of hybridization at any length. Experience is so rapidly increasing both in a practical and in a purely scientific direction that it would take an entire volume to give only a brief survey of the facts and of all the proposed theories.

[251] For our present purposes we are to deal with hybrids only in so far as they afford the means of a still better distinction between elementary species and varieties. I will try to show that these two contrasting groups behave in quite a different manner, when subjected to crossing experiments, and that the hope is justified that some day crosses may become the means of deciding in any given instance, what is to be called a species, and what a variety, on physiologic grounds. It is readily granted that the labor required for such experiments, is perhaps too great for the results to be attained, but then it may be possible to deduce rules from a small series of experiments, which may lead us to a decision in wider ranges of cases.

To reach such a point of view it is necessary to compare the evidence given by hybrids, with the conclusions already attained by the comparison of the differentiating characteristics of allied forms.

On this ground we first have to inquire what may be expected respecting the internal nature and the outcome of the process of crossing in the various cases cited in our former discussion.

We must always distinguish the qualities, which are the same in both parents, from those that constitute the differentiating marks in every single cross. In respect to the first [252] group the cross is not at all distinguished from a normal fertilization, and ordinarily these characters are simply left out of consideration. But it should never be forgotten that they constitute the enormous majority, amounting to hundreds and thousands, whereas the differentiating marks in each case are only one or two or a few at most. The whole discussion is to be limited to these last-named exceptions. We must consider first what would be the nature of a cross when species are symmetrically combined, and what must be the case when varieties are subjected to the same treatment. In so doing, I intend to limit the discussion to the most typical cases. We may take the crosses between elementary species of the same or of very narrowly allied systematic species on the one side, and on the other, limit treatment to the crossing of varieties with the species, from which they are supposed to have sprung by a retrograde modification. Crosses of different varieties of the same species with one another obviously constitute a derivative case, and should only be discussed secondarily. And crosses of varieties with positive or depressive characters have as yet so rarely been made that we may well disregard them.

Elementary species differ from their nearest allies by progressive changes, that is by the acquirement [253] of some new character. The derivative species has one unit more than the parent. All other qualities are the same as in the parent. Whenever such a derivative is combined with its parent the result for these qualities will be exactly as in a normal fertilization. In such ordinary cases it is obvious that each character of the pollen-parent is combined with the same character of the pistil-parent. There may be slight individual differences, but each unit character will become opposed to, and united with, the same unit-character in the other parent. In the offspring the units will thus be paired, each pair consisting of two equivalent units. As to their character the units of each single pair are the same, only they may exhibit slight differences as to the degree of development of this character.

Now we may apply this conception to the sexual combination of two different elementary species, assuming one to be the derivative of the other. The differentiating mark is only present in one of the parents and wanting in the other. While all other units are paired in the hybrid, this one is not. It meets with no mate, and must therefore remain unpaired. The hybrid of two such elementary species is in some way incomplete and unnatural. In the ordinary course of things all individuals derive [254] their qualities from both parents; for each single mark they possess at least two units. Practically but not absolutely equal, these two opponents always work together and give to the offspring a likeness to both parents. No unpaired qualities occur in normal offspring; these constitute the essential features of the hybrids of species and are at the same time the cause of their wide deviations from the ordinary rules.

Turning now to the varieties, we likewise need discuss their differentiating marks only. In the negative types, these consist of the apparent loss of some quality which was active in the species. But it was pointed out in our last lecture that such a change is an apparent loss. On a closer inquiry we are led to the assumption of a latent or dormant state. The presumably lost characters have not absolutely, or at least not permanently disappeared. They show their presence by some slight indication of the quality they represent, or by occasional reversions. They are not wanting, but only latent.

Basing our discussion concerning the process of crossing on this conception, and still limiting the discussion to one differentiating mark, we come to the inference, that this mark is present and active in the species, and present but dormant in the variety. Thus it is present in both, and as all other characters not differentiating [255] find their mates in the cross, so these two will also meet one another. They will unite just as well as though they were both active or both dormant. For essentially they are the same, only differing in their degree of activity. From this we can infer, that in the crossing of varieties, no unpaired remainder is left, all units combining in pairs exactly as in ordinary fertilization.

Setting aside the contrast between activity and latency in this single pair, the procedure in the inter-crossing of varieties is the same as in ordinary normal fertilization.

Summarizing this discussion we may conclude that in normal fertilization and in the inter-crossing of varieties all characters are paired, while in crosses between elementary species the differentiating marks are not mated.

In order to distinguish these two great types of fertilization we will use the term bisexual for the one and unisexual for the other. The term balanced crosses then conveys the idea of complete bisexuality, all unit-characters combining in pairs. Unbalanced crosses are those in which one or more units do not find their mates and therefore remain unpaired. This distinction was proposed by Macfarlane when studying the minute structure of plant-hybrids in comparison with that of their parents (1892).

[256] In the first place it shows that a species hybrid may inherit the distinguishing marks of both parents. In this way it may become intermediate between them, having some characters in common with the pollen-parent and others with the pistil-parent. As far as these characters do not interfere with each other, they may be fully developed side by side, and in the main this is the way in which hybrid characters are evolved. But in most cases our existing knowledge of the units is far too slender to give a complete analysis, even of these distinguishing marks alone. We recognize the parental marks more or less clearly, but are not prepared for exact delimitations. Leaving these theoretical considerations, we will pass to the description of some illustrative examples.

In the first place I will describe a hybrid between two species of Oenothera, which I made some years ago. The parents were the common evening-primrose or Oenothera biennis and of its small-flowered congener, Oenothera muricata. These two forms were distinguished by Linnaeus as different species, but have been considered by subsequent writers as elementary species or so-called systematic varieties of one species designated with the name of the presumably older type, the O. biennis. Varietal differences in a physiologic sense they [257] do not possess, and for this reason afford a pure instance of unbalanced union, though differing in more than one point.

I have made reciprocal crosses, taking at one time the small-flowered and at the other the common species as pistillate parent. These crosses do not lead to the same hybrid as is ordinarily observed in analogous cases; quite on the contrary, the two types are different in most features, both resembling the pollen-parent far more than the pistil-parent. The same curious result was reached in sundry other reciprocal crosses between species of this genus. But I will limit myself here to one of the two hybrids.

In the summer of 1895 I castrated some flowers of O. muricata, and pollinated them with O. biennis, surrounding the flowers with paper bags so as to exclude the visits of insects. I sowed the seeds in 1896 and the hybrids were biennial and flowered abundantly the next year and were artificially fertilized with their own pollen, but gave only a very small harvest. Many capsules failed, and the remaining contained only some few ripe seeds.

From these I had in the following year the second hybrid generation, and in the same way I cultivated also the third and fourth. These were as imperfectly fertile as the first, and in [258] some years did not give any seed at all, so that the operation had to be repeated in order to continue the experiment. Last summer (1903) I had a nice lot of some 25 biennial specimens blooming abundantly. All in all I have grown some 500 hybrids, and of these about 150 specimens flowered.

These plants were all of the same type, resembling in most points the pollen-parent, and in some others the pistil-parent of the original cross. The most obvious characteristic marks are afforded by the flowers, which in O. muricata are not half so large as in biennis, though borne by a calyx-tube of the same length. In this respect the hybrid is like the biennis bearing the larger flowers. These may at times seem to deviate a little in the direction of the other parent, being somewhat smaller and of a slightly paler color. But it is very difficult to distinguish between them, and if biennis and hybrid flowers were separated from the plants and thrown together, it is very doubtful whether one would succeed in separating them.

The next point is offered by the foliage. The leaves of O. biennis are broad, those of O. muricata narrow. The hybrid has the broad leaves of O. biennis during most of its life and at the time of flowering. Yet small deviations in the [259] direction of the other parent are not wanting, and in winter the leaves of the hybrid rosettes are often much narrower than those of O. biennis, and easily distinguishable from both parents. A third distinction consists in the density of the spike. The distance between the insertion of the flowers of O. biennis is great when compared with that of O. muricata. Hence the flowers of the latter species are more crowded and those of O. biennis more dispersed, the spikes of the first being densely crowned with flowers and flower-buds while those of O. biennis are more elongated and slender. As a further consequence the O. biennis opens on the same evening only one, two or three flowers on the same spike, whereas O. muricata bears often eight or ten or more flowers at a time. In this respect the hybrid is similar to the pistil-parent, and the crowding of the broad flowers at the top of the spikes causes the hybrids to be much more showy than either of the parent types.

Other distinguishing marks are not recorded by the systematists, or are not so sharply separated as to allow of the corresponding qualities of the hybrids being compared with them.

This hybrid remains true to the description given. In some years I cultivated two generations [260] so as to be able to compare them with one another, but did not find any difference. The most interesting point however, is the likeness between the first generation, which obviously must combine in its internal structure the units of both parents, and the second and later generations which are only of a derivative nature. Next to this stands the fact that in each generation all individuals are alike. No reversion to the parental forms either in the whole type or in the single characteristics has ever been observed, though the leaves of some hundreds, and the spikes and flowers of some 150 individual plants have been carefully examined. No segregation or splitting up takes place.

Here we have a clear, undoubted and relatively simple, case of a true and pure species hybrid. No occurrence of possible varietal characteristics obscures the result, and in this respect this hybrid stands out much more clearly than all those between garden-plants, where varietal marks nearly always play a most important part.

From the breeder's point of view our hybrid Oenothera would be a distinct gain, were it not for the difficulty of its propagation. But to enlarge the range of the varieties this simple and stable form would need to be treated anew, by [261] crossing it with the parent-types. Such experiments however, have miscarried owing to the too stable nature of the unit-characters.

This stability and this absence of the splitting shown by varietal marks in the offspring of hybrids is one of the best proofs of unisexual unions. It is often obscured by the accompanying varietal marks, or overlooked for this reason. Only in rare cases it is to be met with in a pure state and some examples are given of this below.

Before doing so, I must call your attention to another feature of the unbalanced unions. This is the diminution of the fertility, a phenomenon universally known as occurring in hybridizations. It has two phases. First, the diminished chance of the crosses themselves of giving full crops of seed, as compared with the pure fertilization of either parent. And, secondly, the fertility of the hybrids themselves. Seemingly, all grades of diminished fertility occur and the oldest authors on hybrids have pointed out that a very definite relation exists between the differences of the parents and the degree of sterility, both of the cross and of the hybrid offspring. In a broad sense these two factors are proportionate to each other, the sterility being the greater, the lesser the affinity between the parents. Many writers have [262] tried to trace this rule in the single cases, but have met with nearly unsurmountable difficulties, owing chiefly to our ignorance of the units which form the differences between the parents in the observed cases.

In the case of Oenothera muricata x biennis the differentiating units reduce the fertility to a low degree, threatening the offspring with almost complete infertility and extinction. But then we do not know whether these characters are really units, or perhaps only seemingly so and are in reality composed of smaller entities which as yet we are not able to segregate. And as long as we are devoid of empirical means of deciding such questions, it seems useless to go farther into the details of the question of the sterility. It should be stated here however, that pure varietal crosses, when not accompanied by unbalanced characters, have never showed any tendency to diminished fertility. Hence there can be little doubt that the unpaired units are the cause of this decrease in reproductive power.

The genus Oenothera is to a large degree devoid of varietal characteristics, especially in the subgenus Onagra, to which biennis, muricata, lamarckiana and some others belong. On the other hand it seems to be rich in elementary species, but an adequate study of [263] them has as yet not been made. Unfortunately many of the better systematists are in the habit of throwing all these interesting forms together, and of omitting their descriptive study. I have made a large number of crosses between such undescribed types and as a rule got constant hybrid races. Only one or two exceptions could be quoted, as for instance the Oenothera brevistylis, which in its crosses always behaves as a pure retrogressive variety. Instead of giving an exhaustive survey of hybrids, I simply cite my crosses between lamarckiana and biennis, as having nearly the aspect of the last named species, and remaining true to this in the second generation without any sign of reversion or of splitting. I have crossed another elementary species, the Oenothera hirtella with some of my new and with some older Linnean species, and got several constant hybrid races. Among these the offspring of a cross between muricata and hirtella is still in cultivation. The cross was made in the summer of 1897 and last year (1903) I grew the fourth generation of the hybrids. These had the characters of the muricata in their narrow leaves, but the elongated spikes and relatively large flowers of the hirtella parent, and remained true to this type, showing only slight fluctuations and never reverting or segregating [264] the mixed characters. Both parents bear large capsules with an abundance of seed, but in the hybrids the capsules remain narrow and weak, ripening not more than one-tenth the usual quantity of seed. Both parents are easily cultivated in annual generations and the same holds good for the hybrid. But whereas the hybrid of muricata and biennis is a stout plant, this type is weak with badly developed foliage, and very long strict spikes. Perhaps it was not able to withstand the bad weather of the last few years.

A goodly number of constant hybrids are described in literature, or cultivated in fields and gardens. In such cases the essential question is not whether they are now constant, but whether they have been so from the beginning, or whether they prove to be constant whenever the original cross is repeated. For constant hybrids may also be the issue of incipient splittings, as we shall soon see.

Among other examples we may begin with the hybrid alfalfa or hybrid lucerne (Medicago media). It often originates spontaneously between the common purple lucerne or alfalfa and its wild ally with yellow flowers and procumbent stems, the Medicago falcata. This hybrid is cultivated in some parts of Germany on a large scale, as it is more productive than [265] the ordinary lucerne. It always comes true from seed and may be seen in a wild state in parks and on lawns. It is one of the oldest hybrids with a pure and known lineage. The original cross has been repeated by Urban, who found the hybrid race to be constant from the beginning.

Another very notorious constant hybrid race is the Aegilops speltaeformis. It has been cultivated in botanic gardens for more than half a century, mostly in annual or biennial generations. It is sufficiently fertile and always comes true. Numerous records have been made of it, since formerly it was believed by Fabre and others to be a spontaneous transition from some wild species of grass to the ordinary wheat, not a cross. Godron, however, showed that it can be produced artificially, and how it has probably sprung into existence wherever it is found wild. The hybrid between Aegilops ovata, a small weed, and the common wheat is of itself sterile, producing no good pollen. But it may be fertilized by the pollen of wheat and then gives rise to a secondary hybrid, which is no other than the Aegilops speltaeformis. This remained constant in Godron's experiments during a number of generations, and has been constant up to the present time.

[266] Constant hybrids have been raised by Millardet between several species of strawberries. He combined the old cultivated forms with newly discovered types from American localities. They ordinarily showed only the characteristics of one of their parents and did not exhibit any new combination of qualities, but they came true to this type in the second and later generations.

In the genus Anemone, Janczewski obtained the same results. Some characters of course may split, but others remain constant, and when only such are present, hybrid races result with new combinations of characters, which are as constant as the best species of the same genus. The hybrids of Janczewski were quite fertile, and he points out that there is no good reason why they should not be considered as good new species. If they had not been produced artificially, but found in the wild state, their origin would have been unknown, and there can be no doubt that they would have been described by the best systematists as species of the same value as their parents. Such is especially the case with a hybrid between Anemone magellanica and the common Anemone sylvestris.

Starting from similar considerations Kerner von Marilaun pointed out the fact long ago that many so-called species, of rare occurrence, [267] standing between two allied types, may be considered to have originated by a cross. Surely a wide field for abuse is opened by such an assertion, and it is quite a common habit to consider intermediate forms as hybrids, on the grounds afforded by their external characters alone, and without any exact knowledge of their real origin and often without knowing anything as to their constancy from seed. All such apparent explanations are now slowly becoming antiquated and obsolete, but the cases adduced by Kerner seem to stand this test.

Kerner designates a willow, Salix ehrhartiana as a constant hybrid between Salix alba and S. pentandra. Rhododendron intermedium is an intermediate form between the hairy and the rusty species from the Swiss Alps, R. hirsutum and R. ferrugineum, the former growing on chalky, and the other on silicious soils. Wherever both these types of soil occur in the same valley and these two species approach one another, the hybrid R. intermedium is produced, and is often seen to be propagating itself abundantly. As is indicated by the name, it combines the essential characters of both parents.

Linaria italica is a hybrid toad-flax between L. genistifolia and L. vulgaris, a cross which I have repeated in my garden. Drosera obovata [268] is a hybrid sundew between D. anglica and D. rotundifolia. Primula variabilis is a hybrid between the two common primroses, P. officinalis and P. grandiflora. The willow-herb (Epilobium), the self-heal (Brunella) and the yellow pond-lilies (Nuphar) afford other instances of constant wild hybrids.

Macfarlane has discovered a natural hybrid between two species of sundew in the swamps near Atco, N.J. The parents, D. intermedia and D. filiformis, were growing abundantly all around, but of the hybrid only a group of eleven plants was found. A detailed comparison of the hybrid with its parents demonstrated a minute blending of the anatomical peculiarities of the parental species.

Luther Burbank of Santa Rosa, California, has produced a great many hybrid brambles, the qualities of which in many respects surpass those of the wild species. Most of them are only propagated by cuttings and layers, not being stable from seed. But some crosses between the blackberry and the raspberry (R. fruticosus and R. idaeus) which bear good fruit and have become quite popular, are so fixed in their type as to reproduce their composite characters from seed with as much regularity as the species of Rubus found in nature. Among them are the "Phenomenal" and the [269] "Primus." The latter is a cross between the Californian dewberry and the Siberian raspberry and is certainly to be regarded as a good stable species, artificially produced. Bell Salter crossed the willow-herbs Epilobium tetragonum and E. montanum, and secured intermediate hybrids which remained true to their type during four successive generations.

Other instances might be given. Many of them are to be found in horticultural and botanical journals which describe their systematic and anatomical details. The question of stability is generally dealt with in an incidental manner, and in many cases it is difficult to reach conclusions from the facts given. Especially disturbing is the circumstance that from a horticultural point of view it is quite sufficient that a new type should repeat itself in some of its offspring to be called stable, and that for this reason absolute constancy is rarely proved.

The range of constant hybrids would be larger by far were it not for two facts. The first is the absolute sterility of so many beautiful hybrids, and the second is the common occurrence of retrogressive characters among cultivated plants. To describe the importance of both these groups of facts would take too much [270] time, and therefore it seems best to give some illustrative examples instead.

Among the species of Ribes or currant, which are cultivated in our gardens, the most beautiful are without doubt the Californian and the Missouri currant, or Ribes sanguineum and R. aureum. A third form, often met with, is "Gordon's currant," which is considered to be a hybrid between the two. It has some peculiarities of both parents. The leaves have the general form of the Californian parent, but are as smooth as the Missouri species. The racemes or flower-spikes are densely flowered as in the red species, but the flowers themselves are of a yellow tinge, with only a flesh-red hue on the outer side of the calyx. It grows vigorously and is easily multiplied by cuttings, but it never bears any fruit. Whether it would be constant, if fertile, is therefore impossible to decide. Berberis ilicifolia is considered as a hybrid between the European barberry (B. vulgaris) and the cultivated shrub Mahonia aquifolia. The latter has pinnate leaves, the former undivided ones. The hybrid has undivided leaves which are more spiny than those of the European parent, and which are not deciduous like them, but persist during the winter, a peculiarity inherited from the Mahonia. As far as I [271] have been able to ascertain, this hybrid never produces seed.

Another instance of an absolutely sterile hybrid is the often quoted Cytisus adami. It is a cross between the common laburnum (Cytisus Laburnum) and another species of the same genus, C. purpureus, and has some traits of both. But since the number of differentiating marks is very great in this case, most of the organs have become intermediate. It is absolutely sterile. But it has the curious peculiarity of splitting in a vegetative way. It has been multiplied on a large scale by grafting and was widely found in the parks and gardens of Europe during the last century. Nearly all these specimens reverted from time to time to the presumable parents. Not rarely a bud of Adam's laburnum assumed all the qualities of the common laburnum, its larger leaves, richer flowered racemes, large and brightly yellow flowers and its complete fertility. Other buds on the same tree reverted to the purple parent, with its solitary small flowers, its dense shrublike branches and very small leaves. These too are fertile, though not producing their seeds as abundantly as the C. Laburnum reversions. Many a botanist has sown the seeds of the latter and obtained only pure common C. Laburnum plants. I had a lot of nearly a hundred seedlings [272] myself, many of which have already flowered, bearing the leaves and flowers of the common species. Seeds of the purple reversions have also been sown, and also yielded the parental type only.

Why this most curious hybrid sports so regularly and why others always remain true to their type is as yet an open question.

But recalling our former consideration of this subject the supposition seems allowable that the tendency to revert is not connected with the type of the hybrid, but is apt to occur in some rare individuals of every type. But since most of the sterile hybrids are only known to us in a single individual and its vegetative offspring, this surmise offers an explanation of the rare occurrence of sports.

Finally, we must consider some of the so called hybrid races or strains of garden-plants. Dahlia, Gladiolus, Amaryllis, Fuchsia, Pelargonium and many other common flowers afford the best known instances. Immeasurable variability seems here to be the result of crossing. But on a closer inspection the range of characters is not so very much wider in these hybrid races than in the groups of parent species which have contributed to the origin of the hybrids. Our tuberous begonias owe their variability to at least seven original parent species, [273] and to the almost incredible number of combinations which are possible between their characters. The first of these crosses was made in the nursery of Veitch and Sons near London by Seden, and the first hybrid is accordingly known as Begonia sedeni and is still to be met with. It has been superseded by subsequent crosses between the sedeni itself and the Veitchi and rosiflora, the davisii, the clarkii and others. Each of them contributed its advantageous qualities, such as round flowers, rosy color, erect flower stalks, elevation of the flowers above the foliage and others. New crosses are being made continuously, partly between the already existing hybrids and partly with newly introduced wild species. Only rarely is it possible to get pure seeds, and I have not yet been able to ascertain whether the hybrids would come true from seed. Specific and varietal characters may occur together in many of the several forms, but nothing is as yet accurately known as to their behavior in pure fertilizations. Constancy and segregation are thrown together in such a manner that extreme variability results, and numerous beautiful types may be had, and others may be expected from further crosses. For a scientific analysis, however, the large range of recorded facts and the written history, which at first sight [274] seems to have no lacunae, are not sufficient. Most of the questions remain open and need investigation. It would be a capital idea to try to repeat the history of the begonias or any other hybrid race, making all the described crosses and then recording the results in a manner requisite for complete and careful scientific investigations.

Many large genera of hybrid garden-flowers owe their origin to species rich in varieties or in elementary subspecies. Such is the case with the gladiolus and the tulips. In other cases the original types have not been obtained from the wild state but from the cultures of other countries.

The dahlias were cultivated in Mexico when first discovered by Europeans, and the chrysanthemums have been introduced from the old gardens of Japan. Both of them consisted of various types, which afterwards have been increased chiefly by repeated intercrossing.

The history of many hybrid races is obscure, or recorded by different authorities in a different way. Some have derived their evidence from one nursery, some from another, and the crosses evidently may have been different in different places. The early history of the gladiolus is an instance. The first crosses are recorded to have been made between _Gladiolus_ [275] _psittacinus_ and _G. cardinalis_, and between their hybrid, which is still known under the name of gandavensis_ and the _purpureo-auratus_. But other authors give other lines of descent. So it is with _Amaryllis_, which is said by De Graaff to owe its stripes to _A. vittata_, its fine form to _A. brasiliensis_, the large petals to _A. psittacina_, the giant flowers to _A. leopoldi_, and the piebald patterns to _A. pardina_. But here, too, other authors give other derivations.

Summarizing the results of our inquiry we see in the first place how very much remains to be done. Many old crosses must be repeated and studied anew, taking care of the purity of the cross as well as of the harvesting of the seeds. Many supposed facts will be shown to be of doubtful validity. New facts have to be gathered, and in doing so the distinction between specific and varietal marks must be taken strictly into account. The first have originated as progressive mutations; they give unbalanced crosses with a constant offspring, as far as experience now goes. The second are chiefly due to retrograde modifications, and will be the subject of the next lecture.

[276]

LECTURE X

MENDEL'S LAW OF BALANCED CROSSES

In the scientific study of the result of crosses, the most essential point is the distinction of the several characters of the parents in their combination in the hybrids and their offspring. From a theoretical point of view it would be best to choose parents which would differ only in a single point. The behavior of the differentiating character might then easily be seen.

Unfortunately, such simple cases do not readily occur. Most species, and even many elementary species are distinguished by more than one quality. Varieties deviating only in one unit-character from the species, are more common. But a closer inspection often reveals some secondary characters which may be overlooked in comparative or descriptive studies, but which reassume their importance in experimental crossings.

In a former lecture we have dealt with the qualities which must be considered as being due to the acquisition of new characters. If we [277] compare the new form in this case with the type from which it has originated, it may be seen that the new character does not find its mate, or its opposite, and it will be unpaired in the hybrid.

In the case of retrogressive changes the visible modification is due, at least in the best known instances, to the reduction of an active quality to a state of inactivity or latency. Now if we make a cross between a species and its variety, the differentiating character will be due to the same internal unit, with no other difference than that it is active in the species and latent in the variety. In the hybrid these two corresponding units will make a pair. But while all other pairs in the same hybrid individuals consist of like antagonists, only this pair consists of slightly unlike opponents.

This conception of varietal crosses leads to three assertions, which seem justifiable by actual experience.

First, there is no reason for a diminution of the fertility, as all characters are paired in the hybrid, and no disturbance whatever ensues in its internal structure. Secondly, it is quite indifferent, how the two types are combined, or which of them is chosen as pistillate and which as staminate parent. The deviating pair will have the same constitution in both cases, being [278] built up of one active and one dormant unit. Thirdly this deviating pair will exhibit the active unit which it contains, and the hybrid will show the aspect of the parent in which the character was active and not that of the parent in which it was dormant. Now the active quality was that of the species, and its latent state was found in the variety. Hence the inference that hybrids between a species and its retrograde variety will bear the aspect of the species. This attribute may be fully developed, and then the hybrid will not be distinguishable from the pure species in its outer appearance. Or the character may be incompletely evolved, owing to the failure of cooperation of the dormant unit. In this case the hybrid will be in some sense intermediate between its parents, but these instances are more rare than the alternate ones, though presumably they may play an important part in the variability of many hybrid garden-flowers.

All of these three rules are supported by a large amount of evidence. The complete fertility of varietal hybrids is so universally acknowledged that it is not worth while to give special instances. With many prominent systematists it has become a test between species and varieties, and from our present point of view this assumption is correct. Only the test is of little use in practice, as fertility may be diminished [279] in unbalanced unions in all possible degrees, according to the amount of difference between the parents. If this amount is slight, if for instance, only one unit-character causes the difference, the injury to fertility may, be so small as to be practically nothing. Hence we see that this test would not enable us to judge of the doubtful cases, although it is quite sufficient as a proof in cases of wider differences.

Our second assertion related to the reciprocal crosses. This is the name given to two sexual combinations between the same parents, but with interchanged places as to which furnishes the pollen. In unbalanced crosses of the genus Oenothera the hybrids of such reciprocal unions are often different, as we have previously shown. Sometimes both resemble the pollen parent more, in other instances the pistil-parent. In varietal crosses no such divergence is as yet known. It would be quite superfluous to adduce single cases as proofs for this rule, which was formerly conceived to hold good for hybrids in general. The work of the older hybridists, such as Koelreuter and Gaertner affords numerous instances.

Our third rule is of a wholly different nature. Formerly the distinction between elementary species and varieties was not insisted upon, and the principle which stamps retrograde changes [280] as the true character of varieties is a new one. Therefore it is necessary to cite a considerable amount of evidence in order to prove the assertion that a hybrid bears the active character of its parent-species and not the inactive character of the variety chosen for the cross.

We may put this assertion in a briefer form, stating that the active character prevails in the hybrid over its dormant antagonist. Or as it is equally often put, the one dominates and the other is recessive. In this terminology the character of the species is dominant in the hybrid while that of the variety is recessive. Hence it follows that in the hybrid the latent or dormant unit is recessive, but it does not follow that these three terms have the same meaning, as we shall see presently. The term recessive only applies to the peculiar state into which the latent character has come in the hybrid by its pairing with the antagonistic active unit.

In the first place it is of the highest importance to consider crosses between varieties of recorded origin and the species from which they have sprung. When dealing with mutations of celandine we shall see that the laciniated form originated from the common celandine in a garden at Heidelberg about the year 1590. Among my Oenotheras one of the eldest of the recent productions is the O. brevistylis or short [281] styled species which was seen for the first time in the year 1889. The third example offered is a hairless variety of the evening campion, Lychnis vespertina, found the same year, which hitherto had not been observed.

For these three cases I have made the crosses of the variety with the parent-species, and in each case the hybrid was like the species, and not like the variety. Nor was it intermediate. Here it is proved that the older character dominates the younger one.

In most cases of wild, and of garden-varieties, the relation between them and the parent-species rests upon comparative evidence. Often the variety is known to be younger, in other cases it may be only of local occurrence, but ordinarily the historic facts about its origin have never been known or have long since been forgotten.

The easiest and most widely known varietal crosses are those between varieties with white flowers and the red- or blue-flowered species. Here the color prevails in the hybrid over the lack of pigment, and as a rule the hybrid is as deeply tinted as the species itself, and cannot be distinguished from it, without an investigation of its hereditary qualities. Instances may be cited of the white varieties of the snapdragon, of the red clover, the long-spurred violet (Viola [282] cornuta) the sea-shore aster (Aster Tripolium), corn-rose (Agrostemma Githago), the Sweet William (Silene Armeria), and many garden flowers, as for instance, the Clarkia pulchella, the Polemonium coeruleum, the Veronica longifolia, the gloxinias and others. If the red hue is combined with a yellow ground-color in the species, the variety will be yellow and the hybrid will have the red and yellow mixture of the species as for instance, in the genus Geum. The toad-flax has an orange-colored palate, and a variety occurs in which the palate is of the same yellow tinge as the remaining parts of the corolla. The hybrid between them is in all respects like the parent-species.

Other instances could be given. In berries the same rule prevails. The black nightshade has a variety with yellow berries, and the black color returns in the hybrid. Even the foliage of some garden-plants may afford instances, as for instance, the purplish amaranth (Amaranthus caudatus). It has a green variety, but the hybrid between the two has the red foliage of the species.

Special marks in leaves and in flowers follow the same rule. Some varieties of the opium poppy have large black patches at the basal end of the petals, while in others this pattern is entirely white. In crossing two such varieties, [283] for instance, the dark "Mephisto" with the white-hearted "Danebrog," the hybrid shows the active character of the dark pattern.

Hairy species crossed with their smooth varieties produce hairy hybrids, as in some wheats, in the campion (Lychnis), in Biscutella and others. The same holds good for the crosses between spiny species and their unarmed derivatives, as in the thorn-apple, the corn-crowfoot (Ranunculus arvensis) and others.

Lack of starch in seeds is observed in some varieties of corn and of peas. When such derivatives are crossed with ordinary starch-producing types, the starch prevails in the hybrid.

It would take too much time to give further examples. But there is still one point which should be insisted upon. It is not the systematic relation of the two parents of a cross, that is decisive, but only the occurrence of the same quality, in the one in an active, and in the other in an inactive condition. Hence, whenever this relation occurs between the parents of a cross, the active quality prevails in the hybrid, even when the parents differ from each other in other respects so as to be distinguished as systematic species. The white and red campions give a red hybrid, the black and pale henbane (Hyoscyamus niger and H. pallidus) give a hybrid [284] with the purple veins and center in the corolla of the former, the white and blue thornapple produce a blue hybrid, and so on. Instances of this sort are common in cultivated plants.

Having given this long list of examples of the rule of the dominancy of the active character over the opposite dormant unit, the question naturally arises as to how the antagonistic units are combined in the hybrid. This question is of paramount importance in the consideration of the offspring of the hybrids. But before taking it up it is as well to learn the real signification of recessiveness in the hybrids themselves.

Recessive characters are shown by those rare cases, in which hybrids revert to the varietal parent in the vegetative way. In other words by bud-variations or sports, analogous to the splitting of Adam's laburnum into its parents, by means of bud-variation already described. But here the wide range of differentiating characters of the parents of this most curious hybrid fail. The illustrative examples are extremely simple, and are limited to the active and inactive condition of only one quality.

An instance is given by the long-leaved veronica (Veronica longifolia), which has bluish flowers in long spikes. The hybrid between [285] this species and its white variety has a blue corolla. But occasionally it produces some purely white flowers, showing its power of separating the parental heritages, combined in its internal structures. This reversion is not common, but in thousands of flowering spikes one may expect to find at least one of them. Sometimes it is a whole stem springing from the underground system and bearing only white flowers on all its spikes. In other instances it is only a side branch which reverts and forms white flowers on a stem, the other spikes of which remain bluish. Sometimes a spike even differentiates longitudinally, bearing on one side blue and on the other white corollas, and the white stripe running over the spike may be seen to be long and large, or narrow and short in various degrees. In such cases it is evident that the heritages of the parents remain uninfluenced by each other during the whole life of the hybrid, working side by side, but the active element always prevails over its latent opponent which is ready to break free whenever an opportunity is offered.

It is now generally assumed that this incomplete mixture of the parental qualities in a hybrid, this uncertain and limited combination is the true cause of the many deviations, exhibited by varietal hybrids when compared with their [286] parents. Partial departures are rare in the hybrids themselves, but in their offspring the divergence becomes the rule.

Segregation seems to be a very difficult process in the vegetative way, but it must be very easy in sexual reproduction, indeed so easy as to show itself in nearly every single instance.

Leaving this first generation, the original hybrids, we now come to a discussion of their offspring. Hybrids should be fertilized either by their own pollen, or by that of other individuals born from the same cross. Only in this case can the offspring be considered as a means of arriving at a decision as to the internal nature of the hybrids themselves. Breeders generally prefer to fertilize hybrids with the pollen of their parents. But this operation is to be considered as a new cross, and consequently is wholly excluded from our present discussion. Hence it follows that a clear insight into the heredity of hybrids may be expected only from scientific experiments. Furthermore some of the diversity observed as a result of ordinary crosses, may be due to the instability of the parents themselves or at least of one of them, since breeders ordinarily choose for their crosses some already very variable strain. Combining such a strain with the desirable qualities of some newly imported species, a new strain may [287] result, having the new attribute in addition to all the variability of the old types. In scientific experiments made for the purpose of investigating the general laws of hybridity, such complex cases are therefore to be wholly excluded. The hereditary purity of the parents must be considered as one of the first conditions of success.

Moreover the progeny must be numerous, since neither constancy, nor the exact proportions in the case of instability, can be determined with a small lot of plants.

Finally, and in order to come to a definite choice of research material, we should keep in mind that the chief object is to ascertain the relation of the offspring to their parents. Now in nearly all cases the seeds are separated from the fruits and from one another, before it becomes possible to judge of their qualities. One may open a fruit and count the seeds, but ordinarily nothing is noted as to their characters. In this respect no other plant equals the corn or maize, as the kernels remain together on the spike, and as it has more than one variety characterized by the color, or constitution, or other qualities of the grains. A corn-grain, however, is not a seed, but a fruit containing a seed. Hence the outer parts pertain to the parent plant and only the innermost ones to the [288] seedling and therefore to the following generation. Fruit-characters thus do not offer the qualities we need, only the qualities resulting from fertilizations are characteristic of the new generation. Such attributes are afforded in some cases by the color, in others by the chemical constitution.

We will choose the latter, and take the sugarcorn in comparison with the ordinary or starch producing forms for our starting point. Both sugar- and starch-corns have smooth fruits when ripening. No difference is to be seen in the young ripe spikes. Only the taste, or a direct chemical analysis might reveal the dissimilarity. But as soon as the spikes are dried, a diversity is apparent. The starchy grains remain smooth, but the sugary kernels lose so much water that they become wrinkled. The former becomes opaque, the latter more or less transparent. Every single kernel may instantly be recognized as belonging to either of the types in question, even if but a single grain of the opposite quality might be met with on a spike. Kernels can be counted on the spike, and since ordinary spikes may bear from 300-500 grains and often more, the numerical relation of the different types may be deduced with great accuracy.

Coming now to our experiment, both starchy [289] and sugary varieties are in this respect wholly constant, when cultivated separately. No change is to be seen in the spikes. Furthermore it is very easy to make the crosses. The best way is to cultivate both types in alternate rows and to cut off the staminate panicles a few days before they open their first flowers. If this operation is done on all the individuals of one variety, sparing all the panicles of the other, it is manifest that all the plants will become fertilized by the latter, and hence that the castrated plants will only bear hybrid seeds.

The experiment may be made in two ways; by castrating the sugary or the starchy variety. In both cases the hybrid kernels are the same. As to their composition they repeat the active character of the starchy variety. The sugar is only accumulated as a result of an incapacity of changing it into starch, and the lack of this capacity is to be considered as a retrogressive varietal mark. The starch-producing unit character, which is active in the ordinary sorts of corns, is therefore latent in sugar-corn.

In order to obtain the second generation, the hybrid grains are sown under ordinary conditions, but sufficiently distant from any other variety of corn to insure pure fertilization. The several individuals may be left to pollinate [290] each other, or they may be artificially pollinated with their own pollen.

The outcome of the experiments is shown by the spikes, as soon as they dry. Each spike bears two sorts of kernels irregularly dispersed over its surface. In this point all the spikes are alike. On each of them one may see on the first inspection that the majority of the kernels are starch-containing seeds, while a minor part becomes wrinkled and transparent according to the rule for sugary seeds. This fact shows at once that the hybrid race is not stable, but has differentiated the parental characters, bringing those of the varietal parent to perfect purity and isolation. Whether the same holds good for the starchy parent, it is impossible to judge from the inspection of the spikes, since it has been seen in the first generation that the hybrid kernels are not visibly distinguished from those of the pure starch-producing grains.

It is very easy to count the number of both sorts of grains in the spike of such a hybrid. In doing so we find, that the proportion is nearly the same on all the spikes, and only slight variations would be found in hundreds of them. One-fourth of the seeds are wrinkled and three-fourths are always smooth. The number may vary in single instances and be a little more or a little less than 25%, ranging, for [291] instance, from 20 to 27%, but as a rule, the average is found nearly equal to 25%.