Contrasted with these groups of constant forms are three inconstant types which we now take up. They belong to two different groups, according to the cause of their inconstancy. In one species which I call _O. lata_, the question of stability or instability must remain wholly unsolved, as only pistillate flowers are produced, and no seed can be fertilized save by the use of the pollen of another form, and therefore by hybridization. The other head comprises two fertile forms, _O.

scintillans_ and _O. elliptica_, which may easily be fertilized with their own pollen, but which gave a progeny only partly similar to the parents.

The _Oenothera lata_ is a very distinct form [541] which was found more than once in the field, and recently (1902) in a luxuriant flowering specimen. It has likewise been raised from seeds collected in different years at the original station. It is also wholly pistillate. Apparently the anthers are robust, but they are dry, wrinkled and nearly devoid of contents. The inner wall of cells around the groups of pollen grow out instead of being resorbed, partly filling the cavity which is left free by the miscarriage of the pollen-grains. This miscarriage does not affect all the grains in the same degree, and under the microscope a few of them with an apparently normal structure may be seen. But the contents are not normally developed, and I have tried in vain to obtain fertilization with a large number of flowers. Only by cross-fertilization does _O. lata_ produce seeds, and then as freely as the other species when self-fertilized. Of course its chance of ever founding a wild type is precluded by this defect.

_O. lata_ is a low plant, with a limp stem, bent tips and branches, all very brittle, but with dense foliage and luxuriant growth. It has bright yellow flowers and thick flower-buds. But for an unknown reason the petals are apt to unfold only partially and to remain wrinkled throughout the flowering time. The stigmas are slightly divergent from the normal type, [542] also being partly united with one another, and laterally with the summit of the style, but without detriment to their function.

Young seedlings of _lata_ may be recognized by the very first leaves.

They have a nearly orbicular shape and are very sharply set off against their stalk. The surface is very uneven, with convexities and concavities on both sides. This difference is lessened in the later leaves, but remains visible throughout the whole life of the plant, even during the flowering season. Broad, sinuate leaves with rounded tips are a sure mark of _O. lata_. On the summits of the stems and branches they are crowded so as to form rosettes.

Concerning inheritance of these characteristics nothing can be directly a.s.serted because of the lack of pollen. The new type can only be perpetuated by crosses, either with the parent form or some other mutant. I have fertilized it, as a rule, with _lamarckiana_ pollen, but have often also used that from _nanella_ and others. In doing so, the _lata_ repeats its character in part of its offspring. This part seems to be independent of the nature of the pollen used, but is very variable according to external circ.u.mstances. On the average one-fourth of the offspring become _lata_, the others a.s.suming the type of the pollen-parent, if this was a _lamarckiana_ or [543] partly this type and partly that of any other of the new species derived from _lamarckiana_, that might have been used as the pollen-parent. This average seems to be a general rule, recurring in all experiments, and remaining unchanged through a long series of successive generations. The fluctuations around this mean go up to nearly 50% and down nearly to 1%, but, as in other cases, such extreme deviations from the average are met with only exceptionally.

The second category includes the inconstant but perfectly fertile species. I have already given the names of the only two forms, which deserve to be mentioned here.

One of them is called _scintillans_ or the shiny evening-primrose, because its leaves are of a deep green color with smooth surfaces, glistening in the sunshine. On the young rosettes these leaves are somewhat broader, and afterwards somewhat narrower than those of _O.

lamarckiana_ at the corresponding ages. The plants themselves always remain small, never reaching the stature of the ancestral type. They are likewise much less branched. They can easily be cultivated in annual generations, but then do not become as fully developed and as fertile, as when flowering in the second year. The flowers have the same structure as those of the _lamarckiana_, but are of a smaller size.

[544] Fertilizing the flowers artificially with their own pollen, excluding the visiting insects by means of paper bags, and saving and sowing the seed of each individual separately, furnishes all the requisites for the estimation of the degree of stability of this species. In the first few weeks the seed-pans do not show any unequality, and often the young plants must be replanted at wider intervals, before anything can be made out with certainty. But as soon as the rosettes begin to fill it becomes manifest that some of them are more backward than others in size. Soon the smaller ones show their deeper green and broader leaves, and thereby display the attributes of the _scintillans_. The other grow faster and stronger and exhibit all the characteristics of ordinary _lamarckiana_s.

The numerical proportion of these two groups has been found different on different occasions. Some plants give about one-third _scintillans_ and two-thirds _lamarckiana_, while the progeny of individuals of another strain show exactly the reverse proportion.

Two points deserve to be noticed. First the progeny of the _scintillans_ appears to be mutable in a large degree, exceeding even the _lamarckiana_. The same forms that are produced most often by the parent-family are also most ordinarily [545] met with among the offspring of the shiny evening-primrose. They are _oblonga_, _lata_ and _nanella_. _Oblonga_ was observed at times to const.i.tute as much as 1% or more of the sowings of _scintillans_, while _lata_ and _nanella_ were commonly seen only in a few scattering individuals, although seldom lacking in experiments of a sufficient size.

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 compet.i.tion 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, a.n.a.logous 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 a.n.a.logy 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 a.s.sume 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 const.i.tuting 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 repet.i.tion 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 quant.i.ties 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 circ.u.mstances as favorable as possible. They were established in pots with well-manured soil and kept under gla.s.s, 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 ill.u.s.trates 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 circ.u.mstances. 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 const.i.tuted 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 a.s.sume 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 a.s.sumes 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_.