The attribution attempted here of the essential difference in the development of hybrids to a permanent or temporary union of the differing cell elements can, of course, only claim the value of an hypothesis for which the lack of definite data offers a wide field. Some justification of the opinion expressed lies in the evidence afforded by Pisum that the behaviour of each pair of differentiating characters in hybrid union is independent of the other differences between the two original plants, and, further, that the hybrid produces just so many kinds of egg and pollen cells as there are possible constant combination forms. The differentiating characters of two plants can finally, however, only depend upon differences in the composition and grouping of the elements which exist in the foundation-cells [fertilised ova] of the same in vital interaction[47].
Even the validity of the law formulated for Pisum requires still to be confirmed, and a repetition of the more important experiments is consequently much to be desired, that, for instance, relating to the composition of the hybrid fertilising cells. A differential [element] may easily escape the single observer[48], which although at the outset may appear to be unimportant, may yet accumulate to such an extent that it must not be ignored in the total result. Whether the variable hybrids of other plant species observe an entire agreement must also be first decided experimentally. In the meantime we may assume that in material points a difference in principle can scarcely occur, since the unity in the developmental plan of organic life is beyond question.
In conclusion, the experiments carried out by Kölreuter, Gärtner, and others with respect to the transformation of one species into another by artificial fertilisation merit special mention. A special importance has been attached to these experiments, and Gärtner reckons them among “the most difficult of all in hybridisation.”
If a species A is to be transformed into a species B, both must be united by fertilisation and the resulting hybrids then be fertilised with the pollen of B; then, out of the various offspring resulting, that form would be selected which stood in nearest relation to B and once more be fertilised with B pollen, and so continuously until finally a form is arrived at which is like B and constant in its progeny. By this process the species A would change into the species B. Gärtner alone has effected thirty such experiments with plants of genera Aquilegia, Dianthus, Geum, Lavatera, Lychnis, Malva, Nicotiana, and Œnothera. The period of transformation was not alike for all species. While with some a triple fertilisation sufficed, with others this had to be repeated five or six times, and even in the same species fluctuations were observed in various experiments. Gärtner ascribes this difference to the circumstance that “the specific [typische] force by which a species, during reproduction, effects the change and transformation of the maternal type varies considerably in different plants, and that, consequently, the periods within which the one species is changed into the other must also vary, as also the number of generations, so that the transformation in some species is perfected in more, and in others in fewer generations.” Further, the same observer remarks “that in these transformation experiments a good deal depends upon which type and which individual be chosen for further transformation.”
If it may be assumed that in these experiments the constitution of the forms resulted in a similar way to that of Pisum, the entire process of transformation would find a fairly simple explanation. The hybrid forms as many kinds of egg cells as there are constant combinations possible of the characters conjoined therein, and one of these is always of the same kind as the fertilising pollen cells. Consequently there always exists the possibility with all such experiments that even from the second fertilisation there may result a constant form identical with that of the pollen parent. Whether this really be obtained depends in each separate case upon the number of the experimental plants, as well as upon the number of differentiating characters which are united by the fertilisation. Let us, for instance, assume that the plants selected for experiment differed in three characters, and the species ABC is to be transformed into the other species abc by repeated fertilisation with the pollen of the latter; the hybrids resulting from the first cross form eight different kinds of egg cells, viz.:
ABC, ABc, AbC, aBC, Abc, aBc, abC, abc.
These in the second year of experiment are united again with the pollen cells abc, and we obtain the series
AaBbCc + AaBbc + AabCc + aBbCc + Aabc + aBbc + abCc + abc.
Since the form abc occurs once in the series of eight components, it is consequently little likely that it would be missing among the experimental plants, even were these raised in a smaller number, and the transformation would be perfected already by a second fertilisation. If by chance it did not appear, then the fertilisation must be repeated with one of those forms nearest akin, Aabc, aBbc, abCc. It is perceived that such an experiment must extend the farther the smaller the number of experimental plants and the larger the number of differentiating characters in the two original species; and that, furthermore, in the same species there can easily occur a delay of one or even of two generations such as Gärtner observed. The transformation of widely divergent species could generally only be completed in five or six years of experiment, since the number of different egg cells which are formed in the hybrid increases in square ratio with the number of differentiating characters.
Gärtner found by repeated experiments that the respective period of transformation varies in many species, so that frequently a species A can be transformed into a species B a generation sooner than can species B into species A. He deduces therefrom that Kölreuter’s opinion can hardly be maintained that “the two natures in hybrids are perfectly in equilibrium.” It appears, however, that Kölreuter does not merit this criticism, but that Gärtner rather has overlooked a material point, to which he himself elsewhere draws attention, viz. that “it depends which individual is chosen for further transformation.” Experiments which in this connection were carried out with two species of Pisum demonstrated that as regards the choice of the fittest individuals for the purpose of further fertilisation it may make a great difference which of two species is transformed into the other. The two experimental plants differed in five characters, while at the same time those of species A were all dominant and those of species B all recessive. For mutual transformation A was fertilised with pollen of B, and B with pollen of A, and this was repeated with both hybrids the following year. With the first experiment B/A there were eighty-seven plants available in the third year of experiment for the selections of individuals for further crossing, and these were of the possible thirty-two forms; with the second experiment A/B seventy-three plants resulted, which agreed throughout perfectly in habit with the pollen parent; in their internal composition, however, they must have been just as varied as the forms of the other experiment. A definite selection was consequently only possible with the first experiment; with the second some plants selected at random had to be excluded. Of the latter only a portion of the flowers were crossed with the A pollen, the others were left to fertilise themselves. Among each five plants which were selected in both experiments for fertilisation there agreed, as the following year’s culture showed, with the pollen parent:—