Darwin and Modern Science [291]
vigorous growth by floral biology.
While the influence of the work on the biology of flowers was extraordinarily great, it could not fail to elicit opinions at variance with Darwin's conclusions. The opposition was based partly on reasons valueless as counterarguments, partly on problems which have still to be solved; to some extent also on that tendency against teleological conceptions which has recently become current. This opposing trend of thought is due to the fact that many biologists are content with teleological explanations, unsupported by proof; it is also closely connected with the fact that many authors estimate the importance of natural selection less highly than Darwin did. We may describe the objections which are based on the widespread occurrence of self- fertilisation and geitonogamy as of little importance. Darwin did not deny the occurrence of self-fertilisation, even for a long series of generations; his law states only that "Nature abhors PERPETUAL self- fertilisation." (It is impossible (as has been attempted) to express Darwin's point of view in a single sentence, such as H. Muller's statement of the "Knight-Darwin law." The conditions of life in organisms are so various and complex that laws, such as are formulated in physics and chemistry, can hardly be conceived.) An exception to this rule would therefore occur only in the case of plants in which the possibility of cross-pollination is excluded. Some of the plants with cleistogamous flowers might afford examples of such cases. We have already seen, however, that such a case has not as yet been shown to occur. Burck believed that he had found an instance in certain tropical plants (Anonaceae, Myrmecodia) of the complete exclusion of cross-fertilisation. The flowers of these plants, in which, however,--in contrast to the cleistogamous flowers--the corolla is well developed, remain closed and fruit is produced.
Loew (E. Loew, "Bemerkungen zu Burck...", "Biolog. Centralbl." XXVI. (1906).) has shown that cases occur in which cross-fertilisation may be effected even in these "cleistopetalous" flowers: humming birds visit the permanently closed flowers of certain species of Nidularium and transport the pollen. The fact that the formation of hybrids may occur as the result of this shows that pollination may be accomplished.
The existence of plants for which self-pollination is of greater importance than it is for others is by no means contradictory to Darwin's view. Self- fertilisation is, for example, of greater importance for annuals than for perennials as without it seeds might fail to be produced. Even in the case of annual plants with small inconspicuous flowers in which self- fertilisation usually occurs, such as Senecio vulgaris, Capsella bursa- pastoris and Stellaria media, A. Bateson (Anna Bateson, "The effects of cross-fertilisation on inconspicuous flowers", "Annals of Botany", Vol. I. 1888, page 255.) found that cross-fertilisation gave a beneficial result, although only in a slight degree. If the favourable effects of sexual reproduction, according to Darwin's view, are correlated with change of environment, it is quite possible that this is of less importance in plants which die after ripening their seeds ("hapaxanthic") and which in any case constantly change their situation. Objections which are based on the proof of the prevalence of self-fertilisation are not, therefore, pertinent. At first sight another point of view, which has been more recently urged, appears to have more weight.
W. Burck (Burck, "Darwin's Kreuzeungsgesetz...", "Biol. Centralbl". XXVIII. 1908, page 177.) has expressed the opinion that the beneficial results of cross-fertilisation demonstrated by Darwin concern only hybrid plants. These alone become weaker by self-pollination; while pure species derive no advantage from crossing and no disadvantage from self-fertilisation. It is certain that some of the plants used by Darwin were of hybrid origin. (It is questionable if this was always the case.) This is evident from his statements,
While the influence of the work on the biology of flowers was extraordinarily great, it could not fail to elicit opinions at variance with Darwin's conclusions. The opposition was based partly on reasons valueless as counterarguments, partly on problems which have still to be solved; to some extent also on that tendency against teleological conceptions which has recently become current. This opposing trend of thought is due to the fact that many biologists are content with teleological explanations, unsupported by proof; it is also closely connected with the fact that many authors estimate the importance of natural selection less highly than Darwin did. We may describe the objections which are based on the widespread occurrence of self- fertilisation and geitonogamy as of little importance. Darwin did not deny the occurrence of self-fertilisation, even for a long series of generations; his law states only that "Nature abhors PERPETUAL self- fertilisation." (It is impossible (as has been attempted) to express Darwin's point of view in a single sentence, such as H. Muller's statement of the "Knight-Darwin law." The conditions of life in organisms are so various and complex that laws, such as are formulated in physics and chemistry, can hardly be conceived.) An exception to this rule would therefore occur only in the case of plants in which the possibility of cross-pollination is excluded. Some of the plants with cleistogamous flowers might afford examples of such cases. We have already seen, however, that such a case has not as yet been shown to occur. Burck believed that he had found an instance in certain tropical plants (Anonaceae, Myrmecodia) of the complete exclusion of cross-fertilisation. The flowers of these plants, in which, however,--in contrast to the cleistogamous flowers--the corolla is well developed, remain closed and fruit is produced.
Loew (E. Loew, "Bemerkungen zu Burck...", "Biolog. Centralbl." XXVI. (1906).) has shown that cases occur in which cross-fertilisation may be effected even in these "cleistopetalous" flowers: humming birds visit the permanently closed flowers of certain species of Nidularium and transport the pollen. The fact that the formation of hybrids may occur as the result of this shows that pollination may be accomplished.
The existence of plants for which self-pollination is of greater importance than it is for others is by no means contradictory to Darwin's view. Self- fertilisation is, for example, of greater importance for annuals than for perennials as without it seeds might fail to be produced. Even in the case of annual plants with small inconspicuous flowers in which self- fertilisation usually occurs, such as Senecio vulgaris, Capsella bursa- pastoris and Stellaria media, A. Bateson (Anna Bateson, "The effects of cross-fertilisation on inconspicuous flowers", "Annals of Botany", Vol. I. 1888, page 255.) found that cross-fertilisation gave a beneficial result, although only in a slight degree. If the favourable effects of sexual reproduction, according to Darwin's view, are correlated with change of environment, it is quite possible that this is of less importance in plants which die after ripening their seeds ("hapaxanthic") and which in any case constantly change their situation. Objections which are based on the proof of the prevalence of self-fertilisation are not, therefore, pertinent. At first sight another point of view, which has been more recently urged, appears to have more weight.
W. Burck (Burck, "Darwin's Kreuzeungsgesetz...", "Biol. Centralbl". XXVIII. 1908, page 177.) has expressed the opinion that the beneficial results of cross-fertilisation demonstrated by Darwin concern only hybrid plants. These alone become weaker by self-pollination; while pure species derive no advantage from crossing and no disadvantage from self-fertilisation. It is certain that some of the plants used by Darwin were of hybrid origin. (It is questionable if this was always the case.) This is evident from his statements,