Darwiniana : Essays — Volume 02

by Thomas Henry Huxley

Of such works published subsequently to 1859, several are monographic discussions of topics briefly dealt with in the "Origin," which, it must always be recollected, was considered by the author to be merely an abstract of an opus majus.

The earliest of the books which may be placed in this category, "On the Various Contrivances by which Orchids are Fertilised by Insects," was published in 1862, and whether we regard its theoretical significance, the excellence of the observations and the ingenuity of the reasonings which it records, or the prodigious mass of subsequent investigation of which it has been the parent, it has no superior in point of importance. The conviction that no theory of the origin of species could be satisfactory which failed to offer an explanation of the way in which mechanisms involving adaptations of structure and function to the performance of certain operations are brought about, was, from the first, dominant in Darwin's mind. As has been seen, he rejected Lamarck's views because of their obvious incapacity to furnish such an explanation in the case of the great majority of animal mechanisms, and in that of all those presented by the vegetable world.

So far back as 1793, the wonderful work of Sprengel had established, beyond any reasonable doubt, the fact that, in a large number of cases, a flower is a piece of mechanism the object of which is to convert insect visitors into agents of fertilisation. Sprengel's observations had been most undeservedly neglected and well-nigh forgotten; but Robert Brown having directed Darwin's attention to them in 1841, he was attracted towards the subject, and verified many of Sprengel's statements. (III, p. 258.) It may be doubted whether there was a living botanical specialist, except perhaps Brown, who had done as much. If, however, adaptations of this kind were to be explained by natural selection, it was necessary to show that the plants which were provided with mechanisms for ensuring the aid of insects as fertilisers, were by so much the better fitted to compete with their rivals. This Sprengel had not done. Darwin had been attending to cross fertilisation in plants so far back as 1839, from having arrived, in the course of his speculations on the origin of species, at the conviction "that crossing played an important part in keeping specific forms constant" (I, p. 90). The further development of his views on the importance of cross fertilisation appears to have taken place between this time and 1857, when he published his first papers on the fertilisation of flowers in the "Gardener's Chronicle." If the conclusion at which he ultimately arrived, that cross fertilisation is favourable to the fertility of the parent and to the vigour of the offspring, is correct, then it follows that all those mechanisms which hinder self-fertilisation and favour crossing must be advantageous in the struggle for existence; and, the more perfect the action of the mechanism, the greater the advantage. Thus the way lay open for the operation of natural selection in gradually perfecting the flower as a fertilisation-trap. Analogous reasoning applies to the fertilising insect. The better its structure is adapted to that of the trap, the more will it be able to profit by the bait, whether of honey or of pollen, to the exclusion of its competitors. Thus, by a sort of action and reaction, a two-fold series of adaptive modifications will be brought about.

In 1865, the important bearing of this subject on his theory led Darwin to commence a great series of laborious and difficult experiments on the fertilisation of plants, which occupied him for eleven years, and furnished him with the unexpectedly strong evidence in favour of the influence of crossing which he published in 1876, under the title of "The Effects of Cross and Self Fertilisation in the Vegetable Kingdom." Incidentally, as it were, to this heavy piece of work, he made the remarkable series of observations on the different arrangements by which crossing is favoured and, in many cases, necessitated, which appeared in the work on "The Different Forms of Flowers in Plants of the same Species" in 1877.

In the course of the twenty years during which Darwin was thus occupied in opening up new regions of investigation to the botanist and showing the profound physiological significance of the apparently meaningless diversities of floral structure, his attention was keenly alive to any other interesting phenomena of plant life which came in his way. In his correspondence, he not unfrequently laughs at himself for his ignorance of systematic botany; and his acquaintance with vegetable anatomy and physiology was of the slenderest. Nevertheless, if any of the less common features of plant life came under his notice, that imperious necessity of seeking for causes which nature had laid upon him, impelled, and indeed compelled, him to inquire the how and the why of the fact, and its bearing on his general views. And as, happily, the atavic tendency to frame hypotheses was accompanied by an equally strong need to test them by well-devised experiments, and to acquire all possible information before publishing his results, the effect was that he touched no topic without elucidating it.

Thus the investigation of the operations of insectivorous plants, embodied in the work on that topic published in 1875, was started fifteen years before, by a passing observation made during one of Darwin's rare holidays.

"In the summer of 1860, I was idling and resting near Hartfield, where two species of Drosera abound; and I noticed that numerous insects had been entrapped by the leaves. I carried home some plants, and on giving them some insects saw the movements of the tentacles, and this made me think it possible that the insects were caught for some special purpose. Fortunately, a crucial test occurred to me, that of placing a large number of leaves in various nitrogenous and non-nitrogenous fluids of equal density; and as soon as I found that the former alone excited energetic movements, it was obvious that here was a fine new field for investigation." (I, p. 95.)

The researches thus initiated led to the proof that plants are capable of secreting a digestive fluid like that of animals, and of profiting by the result of digestion; whereby the peculiar apparatuses of the insectivorous plants were brought within the scope of natural selection. Moreover, these inquiries widely enlarged our knowledge of the manner in which stimuli are transmitted in plants, and opened up a prospect of drawing closer the analogies between the motor processes of plants and those of animals.

So with respect to the books on "Climbing Plants" (1875), and on the "Power of Movement in Plants" (1880), Darwin says;--

"I was led to take up this subject by reading a short paper by Asa Gray, published in 1858. He sent me some seeds, and on raising some plants I was so much fascinated and perplexed by the revolving movements of the tendrils and stems, which movements are really very simple, though appearing at first sight very complex, that I procured various other kinds of climbing plants and studied the whole subject.... Some of the adaptations displayed by climbing plants are as beautiful as those of orchids for ensuring cross-fertilisation." (I, p. 93.)