Recently some remarkable experiments bearing closely on these questions have been published by F. Rosen.[10] They concern the forms of Erophila (Draba) verna, celebrated in the history of evolutionary theory as the plants especially chosen by Alexis Jordan for the exposition of his views on these subjects.
The "species" contains a profusion of forms dissimilar in many structural characters, such as the size and shape of leaves, flowers, fruits, etc. Of these forms many grow in association. Jordan found, on experiment, that each, to the number of some two hundred, bred true, and that therefore, the conventional assumption that polymorphism of this kind must mean great contemporary variability had no foundation in fact. So far indeed is the evidence from favouring the belief that such forms are in any way transitional or indeterminate, that, as is well known, Jordan used it with every plausibility to support the doctrine of the fixity of species. To certain aspects of Jordan's work we will return later in this chapter, but the matter is in the present connection of especial interest for the reason that Rosen has lately found by experiment that some of these presumably very closely allied forms, crossed together, gave hybrids more or less sterile. In the case of the offspring of one pair of forms only (E. cochleata and stricta) was the fertility undiminished, and the various degrees of sterility found in the other crosses ranged up to the extreme infertility of the hybrids between E. stricta × elata. From this cross ten plants were bred. Of these the four strongest were chosen to breed from, but two of the four proved totally sterile; one had only bad seeds; and from the fourth a single seedling was raised which in its turn proved to be sterile. From the less sterile hybrids F2 families were raised, with the usual experience that in this and subsequent generations the sterility diminished among extracted forms, new and true-breeding types with complete fertility being thus derived from the original cross.[11]
The production of sterility as a consequence of crossing plants so nearly approaching each other as these Erophila "species" do is not a little interesting, and the fact well exemplifies the futility of the various attempts to frame general expressions as to specific properties or behaviour. Commenting on his results Rosen argues that the polymorphic group commonly called by systematists Erophila (Draba) verna may now be regarded as having arisen by crossing, as did his own types mentioned above. The question, however, what species were the original progenitors of the group cannot be answered. Rosen considers that no form which he knows satisfies the requirements, and that it or they must be supposed to be lost. This conclusion will recall the similar problem raised by the Oenothera mutants (Chap. V); and unsatisfactory as it may be to have recourse to such hypotheses we must remember the possibility that as a consequence of hybridisation, subsequent segregation and recombination of factors, species may have thus actually, as we may say, exploded, and left nothing but a polymorphic group of miscellaneous types to represent them in posterity. If this way of regarding the phenomena be a true one, the sterility now seen when some of the group are re-crossed, becomes analogous to that "reversion or crossing" which we now so well understand to be a consequence of the recombination of characters separated at some previous point in the history of descent. In the partial sterility of the contemporary hybrid we see this character reappearing, formed now as it was on the occasion of the original cross, by the meeting of complementary factors.
Another case that may be mentioned in this connection is that of the crosses between various culinary peas (Pisum sativum) and a peculiar form found by Mr. Arthur Sutton growing ostensibly in a wild state in Palestine. This Palestine Pea is low growing, rarely reaching 18 inches. It is in general appearance like a small and poorly grown field pea. The stems are thin and rather hard. The most obvious differences which distinguish this from other field peas are the marked serration of the stipules, and the development of pith in the pods. Such pith is often present in the pods of peas more or less, but in the Palestines it is so strongly developed as almost to form a lomentum. Curiously enough, though the flowers are purple much as those of ordinary field peas, there is no coloured spot in the axils. On the other hand, the stems have coloured stripes running up from the axils. Though this plant differs so little from domesticated peas, all crosses with them either failed, or produced hybrids quite or almost quite sterile. This was Mr. Sutton's experience, and on repeating the experiments with material kindly given by him I found the same result.[12]
In a large series of crosses some seeds died or gave rise to feeble plants. Of the plants which lived, few gave any seed. The seed, however, that was obtained from F1 plants grew well enough, and the F2 plants proved, as often in such cases, fertile. In these, indeed, no sign of sterility was noticeable. The experiment is being repeated in various ways, for, as the genetic behaviour of peas is comparatively well known, the subject is an exceptionally favourable one for these investigations.
Such an example shows the confusion produced the moment we attempt to harmonize conceptions of specific difference with results attained by experimental methods. It has been usual to regard the field pea (P. arvense) as a species distinct from the edible pea (P. sativum). De Candolle and others regard the field pea as derived from a form wild in Italy, but the origin of the edible pea is considered to be unknown. From breeding experiments we find no sterility whatever in the crosses between the various arvense and sativum types, nor in the crosses made between them and several other peculiar types from various countries; whereas this Palestine Pea, which only differs from a small arvense in what might have been thought trivial characters,[13] either fails to cross altogether or gives a sterile product, whatever type be chosen as the other parent.
Examples of this kind have at least the merit that they lead to more precise delimitations of the problem. We are confronted with two distinct alternatives.
1. We may apply the term Species promiscuously to all distinct forms. If we do so it must be clearly understood that we cannot even rule out the several combinations of "presences and absences" represented by the various types whether wild or domesticated. For we may feel perfectly assured that at least all the arvense and all the sativum types yet subjected to experimental tests are on precisely the same level in this respect. There is no distinction, logical or physiological, to be drawn between them. Some contain more factors, and others contain fewer. In some the re-combinations have been brought about by natural variation or crossing, while the same consequences in the others have resulted from man's interference.
2. We may follow the conventions of systematists and distinguish the outstanding or conspicuous forms such as arvense, quadratum, sativum and perhaps a few more as species, and leave the rest unheeded. If this course is followed it must be clearly understood and permitted as a piece of pure pragmatism, deliberately adopted for the convenience of cataloguers and collectors, without regard to any natural fact or system whatsoever.
But while following either the one plan or the other we shall be still awaiting the answer, which only genetic experiment can provide, to the question whether among the various types there are some which differ from the rest in a peculiar way: whether by having groups of characters linked together in especially durable combinations, or by possessing ingredients which cause greater or less disturbance in the processes of cell-division, and especially in the processes of gametic maturation, when they are united by fertilisation with complementary ingredients.