Chapter D: Eversporting Varieties (14)

Secondly the instability seems to be a constant quality, although the words themselves are at first sight, contradictory. I mean to convey the conception that the degree of instability remains unchanged during successive generations. This is a very curious fact, and strongly reminds us of the hereditary conditions of striped-flower varieties. But, on the contrary, the atavists, which are here the individuals with the stature and the characteristics of the _lamarckiana_, have become _lamarckiana_s in their hereditary qualities, too. If their seed is saved and sown, their progeny does not contain any _scintillans_, or at least no more than might arise by ordinary mutations.

One other inconstant new species is to be noted, but as it was very rare both in the field and in my cultures, and as it was difficult of cultivation, little can as yet be said about it. It is the _Oenothera elliptica_, with narrow elliptical leaves and also with elliptical petals. It repeats [546] its type only in a very small proportion of its seed.

All in all we thus have a group of a dozen new types, springing from an original form in one restricted locality, and seen to grow there, or arising in the garden from seeds collected from the original locality. Without any doubt the germs of the new types are fully developed within the seed, ready to be evolved at the time of germination. More favorable conditions in the field would no doubt allow all of the described new species to unfold their attributes there, and to come into competition with each other and with the common parents. But obviously this is only of secondary importance, and has no influence on the fact that a number of new types, analogous to the older swarms of _Draba_, _Viola_ and of many other polymorphous species, have been seen to arise directly in the wild state.

[547]

LECTURE XIX

EXPERIMENTAL PEDIGREE-CULTURES

The observation of the production of mutants in the field at Hilversum, and the subsequent cultivation of the new types in the garden at Amsterdam, gives ample proof of the mutability of plants. Furthermore it furnishes an analogy with the hypothetical origin of the swarms of species of _Draba_ and _Viola_. Last but not least important it affords material for a complete systematic and morphologic study of the newly arisen group of forms.

The physiologic laws, however, which govern this process are only very imperfectly revealed by such a study. The instances are too few. Moreover the seeds from which the mutants spring, escape observation. It is simply impossible to tell from which individual plants they have been derived. The laevifolia and the brevistylis have been found almost every year, the first always recurring on the same spot, the second on various parts of the original field. It is therefore allowable to assume a common [548] origin for all the observed individuals of either strain. But whether, besides this, similar strains are produced anew by the old _lamarckiana_ group, it is impossible to decide on the sole ground of these field-observations.

The same holds good with the other novelties. Even if one of them should germinate repeatedly, without ever opening its flowers, the possibility could not be excluded that the seeds might have come originally from the same capsule but lain dormant in the earth during periods of unequal length.

Other objections might be cited that can only be met by direct and fully controlled experiments. Next to the native locality comes the experimental garden. Here the rule prevails that every plant must be fertilized with pollen of its own, or with pollen of other individuals of known and recorded origin. The visits of insects must be guarded against, and no seeds should be saved from flowers which have been allowed to open without this precaution. Then the seeds of each individual must be saved and sown separately, so as to admit of an appreciation, and if necessary, a numerical determination of the nature of its progeny. And last but not least the experiments should be conducted in a similar manner during a series of successive years.

[549] I have made four such experiments, each comprising the handling of many thousands of individual plants, and lasting through five to nine generations. At the beginning the plants were biennial, as in the native locality, but later I learned to cultivate them in annual generations. They have been started from different plants and seeds, introduced from the original field into my garden at Amsterdam.

It seems sufficient to describe here one of these pedigree-cultures, as the results of all four were similar. In the fall of 1886 I took nine large rosettes from the field, planted them together on an isolated spot in the garden, and harvested their seeds the next year. These nine original plants are therefore to be considered as constituting the first generation of my race. The second generation was sown in 1888 and flowered in 1889. It at once yielded the expected result. 15,000 seedlings were tested and examined, and among them 10 showed diverging characters. They were properly protected, and proved to belong to two new types. 5 of them were _lata_ and 5 _nanella_. They flowered next year and displayed all the characters as described in our preceding lecture. Intermediates between them and the general type were not found, and no indication of their appearance was noted in their parents. [550] They came into existence at once, fully equipped, without preparation or intermediate steps. No series of generations, no selection, no struggle for existence was needed. It was a sudden leap into another type, a sport in the best acceptation of the word. It fulfilled my hopes, and at once gave proof of the possibility of the direct observation of the origin of species, and of the experimental control thereof.

The third generation was in the main a repetition of the second. I tried some 10,000 seedlings and found three _lata_ and three _nanella_, or nearly the same proportion as in the first instance. But besides these a _rubrinervis_ made its appearance and flowered the following year. This fact at once revealed the possibility that the instability of _lamarckiana_ might not be restricted to the three new types now under observation. Hence the question arose how it would be possible to obtain other types or to find them if they were present. It was necessary to have better methods of cultivation and examination of the young plants. Accordingly I devoted the three succeeding years to working on this problem.

I found that it was not at all necessary to sow any larger quantities of seed, but that the young plants must have room enough to develop into full and free rosettes. Moreover I observed [551] that the attributes of _lata_ and _nanella_, which I now studied in the offspring of my first mutants, were clearly discernible in extreme youth, while those of _rubrinervis_ remained concealed some weeks longer. Hence I concluded that the young plants should be examined from time to time until they proved clearly to be only normal _lamarckiana_. Individuals exhibiting any deviation from the type, or even giving only a slight indication of it, were forthwith taken out of the beds and planted separately, under circumstances as favorable as possible. They were established in pots with well-manured soil and kept under glass, but fully exposed to sunshine. As a rule they grew very fast, and could be planted out early in June. Some of them, of course, proved to have been erroneously taken for mutants, but many exhibited new characters.

All in all I had 334 young plants which did not agree with the parental type. As I examined some 14,000 seedlings altogether, the result was estimated at about 2.5%. This proportion is much larger than in the yields of the two first generations and illustrates the value of improved methods. No doubt many good mutations had been overlooked in the earlier observations.

As was to be expected, _lata_ and _nanella_ [552] were repeated in this third generation (1895). I was sure to get nearly all of them, without any important exceptions, as I now knew how to detect them at almost any age. In fact, I found many of them; as many as 60 _nanella_ and 73 _lata_, or nearly 5% of each. _Rubrinervis_ also recurred, and was seen in 8 specimens. It was much more rare than the two first-named types.

But the most curious fact in that year was the appearance of _oblonga_. No doubt I had often seen it in former years, but had not attached any value to the very slight differences from the type, as they then seemed to me. I knew now that any divergence was to be esteemed as important, and should be isolated for further observation. This showed that among the selected specimens not less than 176, or more than 1% belonged to the _oblonga_ type. This type was at that time quite new to me, and it had to be kept through the winter, to obtain stems and flowers. It proved to be as uniform as its three predecessors, and especially as sharply contrasted with _lamarckiana_. The opportunity for the discovery of any intermediates was as favorable as could be, because the distinguishing marks were hardly beyond doubt at the time of the selection and removal of the young plants. But no connecting links were found.

[553] The same holds good for _albida_, which appeared in 15 specimens, or in 0.1%, of the whole culture. By careful cultivation these plants proved not to be sickly, but to belong to a new, though weak type. It was evident that I had already seen them in former years, but having failed to recognize them had allowed them to be destroyed at an early age, not knowing how to protect them against adverse circumstances. Even this time I did not succeed in getting them strong enough to keep through the winter.

Besides these, two new types were observed, completing the range of all that have since been recorded to regularly occur in this family. They were _scintillans_ and _gigas_. The first was obtained in the way just described. The other hardly escaped being destroyed, not having showed itself early enough, and being left in the bed after the end of the selection. But as it was necessary to keep some rosettes through the winter in order to have biennial flowering plants to furnish seeds, I selected in August about 30 of the most vigorous plants, planted them on another bed and gave them sufficient room for their stems and branches in the following summer. Most of them sent up robust shoots, but no difference was noted till the first flowers opened. One plant had a much larger crown of bright blossoms than any of the others. [554] As soon as these flowers faded away, and the young fruits grew out, it became clear that a new type was showing itself. On that indication I removed all the already fertilized flowers and young fruits, and protected the buds from the visits of insects. Thus the isolated flowers were fertilized with their own pollen only, and I could rely upon the purity of the seed saved. This lot of seeds was sown in the spring of 1897 and yielded a uniform crop of nearly 300 young _gigas_ plants.

Having found how much depends upon the treatment, I could gradually decrease the size of my cultures. Evidently the chance of discovering new types would be lessened thereby, but the question as to the repeated production of the same new forms could more easily and more clearly be answered in this way. In the following year (1896) I sowed half as many seeds as formerly, and the result proved quite the same. With the exception of _gigas_ all the described forms sprang anew from the purely fertilized ancestry of normal _lamarckiana_s. It was now the fifth generation of my pedigree, and thus I was absolutely sure that the descendants of the mutants of this year had been pure and without deviation for at least four successive generations.

Owing partly to improved methods of selection, [555] partly no doubt to chance, even more mutants were found this year than in the former. Out of some 8,000 seedlings I counted 377 deviating ones, or nearly 5%, which is a high proportion. Most of them were _oblonga_ and _lata_, the same types that had constituted the majority in the former year.

_Albida_, _nanella_ and _rubrinervis_ appeared in large numbers, and even _scintillans_, of which I had but a single plant in the previous generation, was repeated sixfold.

New forms did not arise, and the capacity of my strain seemed exhausted. This conclusion was strengthened by the results of the next three generations, which were made on a much smaller scale and yielded the same, or at least the mutants most commonly seen in previous years.

Instead of giving the figures for these last two years separately, I will now summarize my whole experiment in the form of a pedigree. In this the normal _lamarckiana_ was the main line, and seeds were only sown from plants after sufficient isolation either of the plants themselves, or in the latter years by means of paper bags enclosing the inflorescences. I have given the number of seedlings of _lamarckiana_ which were examined each year in the table below. Of course by far the largest number of them were [556] thrown away as soon as they showed their differentiating characters in order to make room for the remaining ones. At last only a few plants were left to blossom in order to perpetuate the race. I have indicated for each generation the number of mutants of each of the observed forms, placing them in vertical columns underneath their respective heads. The three first generations were biennial, but the five last annual.

PEDIGREE OF A MUTATING FAMILY
OF _OENOTHERA LAMARCKIANA_ IN THE
EXPERIMENTAL GARDEN AT AMSTERDAM

Gener: O.gig. albida obl. rubrin. Lam. nanella lata. scint.
VIII. 5 1 0 1700 21 1
VII. 9 0 3000 11
VI. 11 29 3 1800 9 5 1
V. 25 135 20 8000 49 142 6
IV. 1 15 176 8 14000 60 73 1
III. 1 10000 3 3
II. 15000 5 5
I. 9

It is most striking that the various mutations of the evening-primrose display a great degree of regularity. There is no chaos of forms, no indefinite varying in all degrees and in all directions. Quite on the contrary, it is at once evident that very simple rules govern the whole phenomenon.

I shall now attempt to deduce these laws from [557] my experiment. Obviously they apply not only to our evening-primroses, but may be expected to be of general validity. This is at once manifest, if we compare the group of new mutants with the swarms of elementary forms which compose some of the youngest systematic species, and which, as we have seen before, are to be considered as the results of previous mutations. The difference lies in the fact that the evening-primroses have been seen to spring from their ancestors and that the _drabas_ have not. Hence the conclusion that in comparing the two we must leave out the pedigree of the evening-primroses and consider only the group of forms as they finally show themselves. If in doing so we find sufficient similarity, we are justified in the conclusion that the _drabas_ and others have probably originated in the same way as the evening-primroses. Minor points of course will differ, but the main lines cannot have complied with wholly different laws. All so-called swarms of elementary species obviously pertain to a single type, and this type includes our evening-primroses as the only controlled case.

Formulating the laws of mutability for the evening-primroses we therefore assume that they hold good for numerous other corresponding cases.

[558] I. The first law is, that new elementary species appear suddenly, without intermediate steps.

This is a striking point, and the one that is in the most immediate contradiction to current scientific belief. The ordinary conception assumes very slow changes, in fact so slow that centuries are supposed to be required to make the differences appreciable. If this were true, all chance of ever seeing a new species arise would be hopelessly small. Fortunately the evening-primroses exhibit contrary tendencies. One of the great points of pedigree-culture is the fact that the ancestors of every mutant have been controlled and recorded. Those of the last year have seven generations of known _lamarckiana_ parents preceding them. If there had been any visible preparation towards the coming mutation, it could not have escaped observation. Moreover, if visible preparation were the rule, it could hardly go on at the same time and in the same individuals in five or six diverging directions, producing from one parent, _gigas_ and _nanella_, _lata_ and _rubrinervis_, _oblonga_ and _albida_ and even _scintillans_.

On the other hand the mutants, that constitute the first representatives of their race, exhibit all the attributes of the new type in full display at once. No series of generations, no selection, [559] no struggle for existence are needed to reach this end. In previous lectures I have mentioned that I have saved the seeds of the mutants whenever possible, and have always obtained repetitions of the prototype only. Reversions are as absolutely lacking as is also a further development of the new type. Even in the case of the inconstant forms, where part of the progeny yearly return to the stature of _lamarckiana_, intermediates are not found. So it is also with _lata_, which is pistillate and can only be propagated by cross-fertilization. But though the current belief would expect intermediates at least in this case, they do not occur. I made a pedigree-culture of lata during eight successive generations, pollinating them in different ways, and always obtained cultures which were partly constituted of _lata_ and partly of _lamarckiana_ specimens. But the _lata_s remained _lata_ in all the various and most noticeable characters, never showing any tendency to gradually revert into the original form.

Intermediate forms, if not occurring in the direct line from one species to another, might be expected to appear perhaps on lateral branches. In this case the mutants of one type, appearing in the same year, would not be a pure type, but would exhibit different degrees of deviation from the parent. The best would then have to [560] be chosen in order to get the new type in its pure condition. Nothing of the kind, however, was observed. All the _oblonga_-mutants were pure _oblongas_. The pedigree shows hundreds of them in the succeeding years, but no difference was seen and no material for selection was afforded. All were as nearly equal as the individuals of old elementary species.

II. New forms spring laterally from the main stem.

The current conception concerning the origin of species assumes that species are slowly converted into others. The conversion is assumed to affect all the individuals in the same direction and in the same degree. The whole group changes its character, acquiring new attributes. By inter-crossing they maintain a common line of progress, one individual never being able to proceed much ahead of the others.

The birth of the new species necessarily seemed to involve the death of the old one. This last conclusion, however, is hard to understand. It may be justifiable to assume that all the individuals of one locality are ordinarily intercrossed, and are moreover subjected to the same external conditions. They might be supposed to vary in the same direction if these conditions were changed slowly. But this could of course have no possible influence on the plants of the [561] same species growing in distant localities, and it would be improbable they should be affected in the same way. Hence we should conclude that when a species is converted into a new type in one locality this is only to be considered as one of numerous possible ones, and its alteration would not in the least change the aspect of the remainder of the species.

But even with this restriction the general belief is not supported by the evidence of the evening-primroses. There is neither a slow nor a sudden change of all the individuals. On the contrary, the vast majority remain unchanged; thousands are seen exactly repeating the original prototype yearly, both in the native field and in my garden. There is no danger that _lamarckiana_ might die out from the act of mutating, nor that the mutating strain itself would be exposed to ultimate destruction from this cause.

In older swarms, such as _Draba_ or _Helianthemum_, no such center, around which the various forms are grouped, is known. Are we to conclude therefore that the main strain has died out? Or is it perhaps concealed among the throng, being distinguished by no peculiar character? If our _gigas_ and _rubrinervis_ were growing in equal numbers with the _lamarckiana_ in the native field, would it be possible to decide [562] which of them was the progenitor of the others? Of course this could be done by long and tedious crossing experiments, showing atavism in the progeny, and thereby indicating the common ancestor. But even this capacity seems to be doubtful and connected only with the state of mutability and to be lost afterwards. Therefore if this period of mutation were ended, probably there would be no way to decide concerning the mutual relationship of the single species.

Hence the lack of a recognizable main stem in swarms of elementary species makes it impossible to answer the question concerning their common origin.

Another phase of the opposition between the prevailing view and my own results seems far more important. According to the current belief the conversion of a group of plants growing in any locality and flowering simultaneously would be restricted to one type. In my own experiments several new species arose from the parental form at once, giving a wide range of new forms at the same time and under the same conditions.

III. New elementary species attain their full constancy at once.

Constancy is not the result of selection or of improvement. It is a quality of its own. It can neither be constrained by selection if it is absent [563] from the beginning, nor does it need any natural or artificial aid if it is present. Most of my new species have proved constant from the first. Whenever possible, the original mutants have been isolated during the flowering period and artificially self-fertilized. Such plants have always given a uniform progeny, all children exhibiting the type of the parent. No atavism was observed and therefore no selection was needed or even practicable.

Briefly considering the different forms, we may state that the full experimental proof has been given for the origin of _gigas_ and _rubrinervis_, for _albida_ and _oblonga_, and even for _nanella_, which is to be considered as of a varietal nature; with _lata_ the decisive experiment is excluded by its unisexuality. _laevifolia_ and _brevistylis_ were found originally in the field, and never appeared in my cultures. No observations were made as to their origin, and seeds have only been sown from later generations. But these have yielded uniform crops, thereby showing that there is no ground for the assumption that these two older varieties might behave otherwise than the more recent derivatives.

_Scintillans_ and _elliptica_ constitute exceptions to the rule given. They repeat their character, from pure seed, only in part of the offspring. I have tried to deliver the _scintillans_ from this [564] incompleteness of heredity, but in vain. The succeeding generations, if produced from true representatives of the new type, and with pure fertilization, have repeated the splitting in the same numerical proportions. The instability seems to be here as permanent a quality as the stability in other instances. Even here no selection has been adequate to change the original form.

IV. Some of the new strains are evidently elementary species, while others are to be considered as retrograde varieties.

It is often difficult to decide whether a given form belongs to one or another of these two groups. I have tried to show that the best and strictest conception of varieties limits them to those forms that have probably originated by retrograde or degressive steps. Elementary species are assumed to have been produced in a progressive way, adding one new element to the store. Varieties differ from their species clearly in one point, and this is either a distinct loss, or the assumption of a character, which may be met with in other species and genera. _laevifolia_ is distinguished by the loss of the crinkling of the leaves, _brevistylis_ by the partial loss of the epigynous qualities of the flowers, and _nanella_ is a dwarf. These three new forms are therefore [565] considered to constitute only retrograde steps, and no advance. This conclusion has been fully justified by some crossing experiments with _brevistylis_, which wholly complies with Mendel's law, and in one instance with _nanella_, which behaves in the same manner, if crossed with _rubrinervis_.

On the other hand, _gigas_ and _rubrinervis_, _oblonga_ and _albida_ obviously bear the characters of progressive elementary species. They are not differentiated from _lamarckiana_ by one or two main features. They diverge from it in nearly all organs, and in all in a definite though small degree. They may be recognized as soon as they have developed their first leaves and remain discernible throughout life. Their characters refer chiefly to the foliage, but no less to the stature, and even the seeds have peculiarities. There can be little doubt but that all the attributes of every new species are derived from one principal change. But why this should affect the foliage in one manner, the flowers in another and the fruits in a third direction, remains obscure. To gain ever so little an insight into the nature of these changes, we may best compare the differences of our evening-primroses with those between the two hundred elementary species of _Draba_ and other similar instances. In doing so we find the same main [566] feature, the minute differences in nearly all points.

V. The same new species are produced in a large number of individuals.

This is a very curious fact. It embraces two minor points, viz: the multitude of similar mutants in the same year, and the repetition thereof in succeeding generations. Obviously there must be some common cause. This cause must be assumed to lie dormant in the _Lamarckiana_s of my strain, and probably in all of them, as no single parent-plant proved ever to be wholly destitute of mutability. Furthermore the different causes for the sundry mutations must lie latent together in the same parent-plant. They obey the same general laws, become active under similar conditions, some of them being more easily awakened than others. The germs of the _oblonga_, _lata_ and _nanella_ are especially irritable, and are ready to spring into activity at the least summons, while those of _gigas_, _rubrinervis_ and _scintillans_ are far more difficult to arouse.

These germs must be assumed to lie dormant during many successive generations. This is especially evident in the case of _lata_ and _nanella__, which appeared in the first year of the pedigree culture and which since have been repeated yearly, and have been seen to arise by mutation [567] also during the last season (1903). Only _gigas_ appeared but once, but then there is every reason to assume that in larger sowings or by a prolongation of the experiments it might have made a second appearance.

Is the number of such germs to be supposed to be limited or unlimited? My experiment has produced about a dozen new forms. Without doubt I could easily have succeeded in getting more, if I had had any definite reason to search for them. But such figures are far from favoring the assumption of indefinite mutability. The group of possible new forms is no doubt sharply circumscribed. Partly so by the morphologic peculiarities of _lamarckiana_, which seem to exclude red flowers, composite leaves, etc. No doubt there are more direct reasons for these limits, some changes having taken place initially and others later, while the present mutations are only repetitions of previous ones, and do not contribute new lines of development to those already existing. This leads us to the supposition of some common original cause, which produced a number of changes, but which itself is no longer at work, but has left the affected qualities, and only these, in the state of mutability.

In nature, repeated mutations must be of far greater significance than isolated ones. How [568] great is the chance for a single individual to be destroyed in the struggle for life? Hundreds of thousands of seeds are produced by _lamarckiana_ annually in the field, and only some slow increase of the number of specimens can be observed. Many seeds do not find the proper circumstances for germination, or the young seedlings are destroyed by lack of water, of air, or of space. Thousands of them are so crowded when becoming rosettes that only a few succeed in producing stems. Any weakness would have destroyed them. As a matter of fact they are much oftener produced in the seed than seen in the field with the usual unfavorable conditions; the careful sowing of collected seeds has given proof of this fact many times.

The experimental proof of this frequency in the origin of new types, seems to overcome many difficulties offered by the current theories on the probable origin of species at large.

VI. The relation between mutability and fluctuating variability has always been one of the chief difficulties of the followers of Darwin. The majority assumed that species arise by the slow accumulation of slight fluctuating deviations, and the mutations were only to be considered as extreme fluctuations, obtained, in the main, by a continuous selection of small differences in a constant direction.

[569] My cultures show that quite the opposite is to be regarded as fact. All organs and all qualities of _lamarckiana_ fluctuate and vary in a more or less evident manner, and those which I had the opportunity of examining more closely were found to comply with the general laws of fluctuation. But such oscillating changes have nothing in common with the mutations. Their essential character is the heaping up of slight deviations around a mean, and the occurrence of continuous lines of increasing deviations, linking the extremes with this group. Nothing of the kind is observed in the case of mutations. There is no mean for them to be grouped around and the extreme only is to be seen, and it is wholly unconnected with the original type. It might be supposed that on closer inspection each mutation might be brought into connection with some feature of the fluctuating variability. But this is not the case. The dwarfs are not at all the extreme variants of structure, as the fluctuation of the height of the _lamarckiana_ never decreases or even approaches that of the dwarfs. There is always a gap. The smallest specimens of the tall type are commonly the weakest, according to the general rule of the relationship between nourishment and variation, but the tallest dwarfs are of course the most robust specimens of their group. [570] Fluctuating variability, as a rule, is subject to reversion. The seeds of the extremes do not produce an offspring which fluctuates around their parents as a center, but around some point on the line which combines their attributes with the corresponding characteristic of their ancestors, as Vilmorin has put it. No reversion accompanies mutation, and this fact is perhaps the completest contrast in which these two great types of variability are opposed to each other.

The offspring of my mutants are, of course, subject to the general laws of fluctuating variability. They vary, however, around their own mean, and this mean is simply the type of the new elementary species.

VII. The mutations take place in nearly all directions.

Many authors assume that the origin of species is directed by unknown causes. These causes are assumed to work in each single case for the improvement of the animals and plants, changing them in a manner corresponding in a useful way to the changes that take place in their environment. It is not easy to imagine the nature of these influences nor how they would bring about the desired effect.

This difficulty was strongly felt by Darwin, and one of the chief purposes of his selection theory may be said to have been the attempt [571] to surmount it. Darwin tried to replace the unknown cause by natural agencies, which lie under our immediate observation. On this point Darwin was superior to his predecessors, and it is chiefly due to the clear conception of this point that his theory has gained its deserved general acceptance. According to Darwin, changes occur in all directions, quite independently of the prevailing circumstances. Some may be favorable, others detrimental, many of them without significance, neither useful nor injurious. Some of them will sooner or later be destroyed, while others will survive, but which of them will survive, is obviously dependent upon whether their particular changes agree with the existing environic conditions or not. This is what Darwin has called the struggle for life. It is a large sieve, and it only acts as such. Some fall through and are annihilated, others remain above and are selected, as the phrase goes. Many are selected, but more are destroyed; daily observation does not leave any doubt upon this point.

How the differences originate is quite another question. It has nothing to do with the theory of natural selection nor with the struggle for life. These have an active part only in the accumulation of useful qualities, and only in so [572] far as they protect the bearers of such characters against being crowded out by their more poorly constituted competitors.

However, the differentiating characteristics of elementary species are only very small. How widely distant they are from the beautiful adaptative organizations of orchids, of insectivorous plants and of so many others! Here the difference lies in the accumulation of numerous elementary characters, which all contribute to the same end. Chance must have produced them, and this would seem absolutely improbable, even impossible, were it not for Darwin's ingenious theory. Chance there is, but no more than anywhere else. It is not by mere chance that the variations move in the required direction. They do go, according to Darwin's view, in all directions, or at least in many. If these include the useful ones, and if this is repeated a number of times, cumulation is possible; if not, there is simply no progression, and the type remains stable through the ages. Natural selection is continually acting as a sieve, throwing out the useless changes and retaining the real improvements. Hence the accumulation in apparently predisposed directions, and hence the increasing adaptations to the more specialized conditions of life. It must be obvious to any one who can free himself from the current ideas, [573] that this theory of natural selection leaves the question as to how the changes themselves are brought about, quite undecided. There are two possibilities, and both have been propounded by Darwin. One is the accumulation of the slight deviations of fluctuating variability, the other consists of successive sports or leaps taking place in the same direction.

In further lectures a critical comparison of the two views will be given. Today I have only to show that the mutations of the evening-primroses, though sudden, comply with the demands made by Darwin as to the form of variability which is to be accepted as the cause of evolution and as the origin of species.

Some of my new types are stouter and others weaker than their parents, as shown by _gigas_ and _albida_. Some have broader leaves and some narrower, _lata_ and _oblonga_. Some have larger flowers (_gigas_) or deeper yellow ones (_rubrinervis_), or smaller blossoms (_scintillans_), or of a paler hue (_albida_). In some the capsules are longer (_rubrinervis_), or thicker (_gigas_), or more rounded (_lata_), or small (_oblonga_), and nearly destitute of seeds (_brevistylis_). The unevenness of the surface of the leaves may increase as in _lata_, or decrease as in _laevifolia_. The tendency to become annual prevails in _rubrinervis_, but _gigas_ tends to become [574] biennial. Some are rich in pollen, while _scintillans_ is poor. Some have large seeds, others small. _Lata_ has become pistillate, while _brevistylis_ has nearly lost the faculty to produce seeds. Some undescribed forms were quite sterile, and some I observed which produced no flowers at all. From this statement it may be seen that nearly all qualities vary in opposite directions and that our group of mutants affords wide material for the sifting process of natural selection. On the original field the _laevifolia_ and _brevistylis_ have held their own during sixteen years and probably more, without, however, being able to increase their numbers to any noticeable extent. Others perish as soon as they make their appearance, or a few individuals are allowed to bloom, but probably leave no progeny.

But perhaps the circumstances may change, or the whole strain may be dispersed and spread to new localities with different conditions. Some of the latter might be found to be favorable to the robust _gigas_, or to _rubrinervis_, which requires a drier air, with rainfall in the springtime and sunshine during the summer. It would be worth while to see whether the climate of California, where neither _O. lamarckiana_ nor _O. biennis_ are found wild, would not exactly [575] suit the requirements of the new species _rubrinervis_ and _gigas_.

NOTE. _Oenothera_s are native to America and all of the species growing in Europe have escaped from gardens directly, or may have arisen by mutation, or by hybridization of introduced species. A fixed hybrid between _O. cruciata_ and _O. biennis_ constituting a species has been in cultivation for many years. The form known as _O. biennis_ in Europe, and used by de Vries in all of the experiments described in these lectures, has not yet been found growing wild in America and is not identical with the species bearing that name among American botanists. Concerning this matter Professor de Vries writes under date of Sept. 12, 1905: "The '_biennis_' which I collected in America has proved to be a motley collection of forms, which at that time I had no means of distinguishing. No one of them, so far as they are now growing in my garden is identical with our _biennis_ of the sand dunes." The same appears to be the case with _O. muricata_. Plants from the Northeastern American seaboard, identifiable with the species do not entirely agree with those raised from seed received from Holland.

_O. lamarckiana_ has not been found growing wild in America in recent years although the evidence at hand seems to favor the conclusion that it was seen and collected in the southern states in the last century. (See MacDougal, Vail, Shull, and Small: Mutants and Hybrids of the _Oenotheras_. Publication 24. Carnegie Institution. Washington, D.C., 1905.) EDITOR.

[576] LECTURE XX

THE ORIGIN OF WILD SPECIES AND VARIETIES

New species and varieties occur from time to time in the wild state. Setting aside all theoretical conceptions as to the common origin of species at large, the undoubted fact remains that new forms are sometimes met with. In the case of the peloric toad-flax the mutations are so numerous that they seem to be quite regular. The production of new species of evening-primroses was observed on the field and afterwards duplicated in the garden. There is no reason to think that these cases are isolated instances. Quite on the contrary they seem to be the prototypes of repeated occurrences in nature.

If this conception is granted, the question at once arises, how are we to deal with analogous cases, when fortune offers them, and what can we expect to learn from them?

A critical study of the existing evidence seems to be of great importance in order to ascertain the best way of dealing with new facts, and of estimating the value of the factors concerned. [577] It is manifest that we must be very careful and conservative in dealing with new facts that are brought to our attention, and every effort should be made to bring additional evidence to light. Many vegetable anomalies are so rare that they are met with only by the purest chance, and are then believed to be wholly new. When a white variety of some common plant is met with for the first time we generally assume that it originated on that very spot and only a short time previously. The discovery of a second locality for the same variety at once raises the question as to a common origin in the two instances. Could not the plants of the second locality have arisen from seeds transported from the first?

White varieties of many species of blue-bells and gentians are found not rarely, white-flowering plants of heather, both of _Erica Tetralix_ and _Calluna vulgaris_ occur on European heaths; white flowers of _Brunella vulgaris_, _Ononis repens_, _Thymus vulgaris_ and others may be seen in many localities in the habitats of the colored species. Pelories of labiates seem to occur often in Austria, but are rare in Holland; white bilberries (_Vaccinium Myrtillus_) have many known localities throughout Europe, and nearly all the berry-bearing species in the large heath family are recorded as having white varieties.

[578] Are we to assume a single origin for all the representatives of such a variety, as we have done customarily for all the representatives of a wild species? Or can the same mutation have been repeated at different times and in distant localities? If a distinct mutation from a given species is once possible, why should it not occur twice or thrice?

A variety which seems to be new to us may only appear so, because the spot where it grows had hitherto escaped observation. _Lychnis preslii_ is a smooth variety of _Lychnis diurna_ and was observed for the first time in the year 1842 by Sekera. It grew abundantly in a grove near Munchengratz in southern Hungary. It was accompanied by the ordinary hairy type of the species. Since then it has been observed to be quite constant in the same locality, and some specimens have been collected for me there lately by Dr. Nemec, of Prague. No other native localities of this variety have been discovered, and there can be no doubt that it must have arisen from the ordinary campion near the spot where it still grows. But this change may have taken place some years before the first discovery, or perhaps one or more centuries ago. This could only be known if it could be proved that the locality had been satisfactorily investigated previously, and that the variety had not [579] been met with. Even in this case only something would be discovered about the time of the change, but nothing about its real nature.

So it is in many cases. If a variety is observed in a number of specimens at the time of its first discovery, and at a locality not studied previously, it takes the aspect of an old form of limited distribution, and little can be learned as to the circumstances under which it arose. If on the contrary it occurs in very small numbers or perhaps even in a single individual, and if the spot where it is found is located so that it could hardly have escaped previous observation, then the presumption of a recent origin seems justified.

What has to be ascertained on such occasions to give them scientific value? Three points strike me as being of the highest importance. First, the constancy of the new type; secondly, the occurrence or lack of intermediates, and last, but not least, the direct observation of a repeated production.

The first two points are easily ascertained. Whether the new type is linked with its more common supposed ancestor by intermediate steps is a query which at once strikes the botanist. It is usually recorded in such cases, and we may state at once that the general result is, that such intermediates do not occur. This is [580] of the highest importance and admits of only two explanations. One is that intermediates may be assumed to have preceded the existent developed form, and to have died out afterwards. But why should they have done so, especially in cases of recent changes? On the other hand the intermediates may be lacking because they have never existed, the change having taken place by a sudden leap, such as the mutations described in our former lectures. It is manifest that the assumption of hypothetical intermediates could only gain some probability if they had been found in some instance. Since they do not occur, the hypothesis seems wholly unsupported.

The second point is the constancy of the new type. Seeds should be saved and sown. If the plant fertilizes itself without the aid of insects, as do some evening-primroses, the seed saved from the native locality may prove wholly pure, and if it does give rise to a uniform progeny the constancy of the race may be assumed to be proved, provided that repeated trials do not bring to light any exceptions. If the offspring shows more than one type, cross-fertilization is always to be looked to as the most probable cause, and should be excluded, in order to sow pure seeds. Garden-experiments of this kind, and repeated trials, should always be combined [581] with the discovery of a presumed mutation. In many instances the authors have realized the importance of this point, and new types have been found constant from the very beginning. Many cases are known which show no reversions and even no partial reversions. This fact throws a distinct light on our first point, as it makes the hypothesis of a slow and gradual development still more improbable.

My third point is of quite another nature and has not as yet been dealt with. But as it appeals to me as the very soul of the problem, it seems necessary to describe it in some detail. It does not refer to the new type itself, nor to any of its morphologic or hereditary attributes, but directly concerns the presumed ancestors themselves.

The peloric toad-flax in my experiment was seen to arise thrice from the same strain. Three different individuals of my original race showed a tendency to produce peloric mutations, and they did so in a number of their seeds, exactly as the mutations of the evening-primroses were repeated nearly every year. Hence the inference, that whenever we find a novelty which is really of very recent date, the parent-strain which has produced it might still be in existence on the same spot. In the case of shrubs or perennials the very parents might yet be found. [582] But it seems probable, and is especially proved in the case of the evening-primroses, that all or the majority of the representatives of the whole strain have the same tendency to mutate. If this were a general rule, it would suffice to take some pure seeds from specimens of the presumed parents and to sow and multiply the individuals to such an extent that the mutation might have a chance to be repeated.

Unfortunately, this has not as yet been done, but in my opinion it should be the first effort of any one who has the good luck to discover a new wild mutation. Specimens of the parents should be transplanted into a garden and fertilized under isolated conditions. Seeds saved from the wild plant would have little worth, as they might have been partly fertilized by the new type itself.

After this somewhat lengthy discussion of the value of observations surrounding the discovery of new wild mutations, we now come to the description of some of the more interesting cases. As a first example, I will take the globular fruited shepherd's purse, described by Solms Laubach as _Capsella heegeri_. Professor Heeger discovered one plant with deviating fruits, in a group of common shepherd's purses in the market-place near Landau in Germany, in the fall of 1897. They were nearly spherical, [583] instead of flat and purse-shaped. Their valves were thick and fleshy, while those of the ordinary form are membranaceous and dry. The capsules hardly opened and therefore differed in this point from the shepherd's purse, which readily loosens both its valves as soon as it is ripe.

Only one plant was observed; whence it came could not be determined, nor whether it had arisen from the neighboring stock of C_apsella_ or not. The discoverer took some seed to his garden and sent some to the botanical garden at Strassburg, of which Solms-Laubach is the director. The majority of the seeds of course were sowed naturally on the original spot. The following year some of the seeds germinated and repeated the novelty. The leaves, stems and flowers were those of the common shepherd's purse, but no decision could be reached concerning the type of this generation before the first flowers had faded and the rounded capsules had developed. Then it was seen that the _heegeri_ came true from seed. It did so both in the gardens and on the market-place, where it was observed to have multiplied and spread in some small measure. The same was noted the following year, but then the place was covered with gravel and all the plants destroyed. It is not recorded to have been seen wild since.

[584] Intermediate forms have not been met with. Some slight reversions may occur in the autumn on the smallest and weakest lateral branches. Such reversions, however, seem to be very rare, as I have tried in vain to produce them on large and richly branched individuals, by applying all possible inducements in the form of manure and of cutting, to stimulate the production of successive generations of weaker side branches.

This constancy was proved by the experiments of Solms-Laubach, which I have repeated in my own garden during several years with seed received from him. No atavists or deviating specimens have been found among many hundreds of flowering plants.

It is important to note that within the family of the crucifers the form of the capsule and the attributes of the valves and seeds are usually considered to furnish the characteristics of genera, and this point has been elucidated at some length by Solms-Laubach. There is, however, no sufficient reason to construe a new genus on the ground of Heeger's globular fruited shepherd's purse; but as a true elementary species, and even as a good systematic species it has proved itself, and as such it is described by Solms-Laubach, who named it in honor of its discoverer.

Exactly analogous discoveries have been [586] instead of displaying a bright yellow cup. _O. cruciata_ grows in the Adirondack Mountains, in the states of New York and Vermont, and seems to be abundant there. It has been introduced into botanical gardens and yielded a number of hybrids, especially with _O. biennis and _O. lamarckiana_, and the narrow petals of the parent-species may be met with in combination with the stature and vegetative characteristics of these last named species. _O. cruciata_ has a purple foliage, while _biennis_ and _lamarckiana_ are green, and many of the hybrids may instantly be recognized by their purple color.

The curious attribute of the petals is not to be considered simply as a reduction in size. On anatomical inquiry it has been found that these narrow petals bear some characteristics which, on the normal plants, are limited to the calyx. Stomata and hairs, and the whole structure of the surface and inner tissues on some parts of these petals are exactly similar to those of the calyx, while on others they have retained the characteristics of petals. Sometimes there may even be seen by the naked eye green longitudinal stripes of calyx-like structure alternating with bright yellow petaloid parts. For these reasons the cruciata character may be considered as a case of sepalody of the petals, or of the petals being partly converted into sepals.

[587] It is worth while to note that as a monstrosity this occurrence is extremely rare throughout the whole vegetable kingdom, and only very few instances have been recorded.

Two cases of sudden mutations have come to my knowledge, producing this same anomaly in allied species. One has been already alluded to; it pertains to the common evening-primrose or _Oenothera biennis_, and one is a species belonging to another genus of the same family, the great hairy willow-herb or _Epilobium _hirsutum_. I propose to designate both new forms by the varietal name of _cruciata_, or _cruciatum_.

_Oenothera biennis cruciata_ was found in a native locality of the _O. biennis itself. It consisted of only one plant, showing in all its flowers the _cruciata_ marks. In all other respects it resembled wholly the _biennis_, especially in the pure green color of its foliage, which at once excluded all suspicion of hybrid origin with the purple _O. cruciata_. Moreover in our country this last occurs only in the cultivated state in botanical gardens.