Darwin and Modern Science [280]
The further development of floral biology by Darwin is in the first place closely connected with the book on the fertilisation of Orchids. It is noteworthy that the title includes the sentence,--"and on the good effects of intercrossing."
The purpose of the book is clearly stated in the introduction:--"The object of the following work is to show that the contrivances by which Orchids are fertilised, are as varied and almost as perfect as any of the most beautiful adaptations in the animal kingdom; and, secondly, to show that these contrivances have for their main object the fertilisation of each flower by the pollen of another flower." ("Fertilisation of Orchids", page 1.) Orchids constituted a particularly suitable family for such researches. Their flowers exhibit a striking wealth of forms; the question, therefore, whether the great variety in floral structure bears any relation to fertilisation (In the older botanical literature the word fertilisation is usually employed in cases where POLLINATION is really in question: as Darwin used it in this sense it is so used here.) must in this case possess special interest.
Darwin succeeded in showing that in most of the orchids examined self- fertilisation is either an impossibility, or, under natural conditions, occurs only exceptionally. On the other hand these plants present a series of extraordinarily beautiful and remarkable adaptations which ensure the transference of pollen by insects from one flower to another. It is impossible to describe adequately in a few words the wealth of facts contained in the Orchid book. A few examples may, however, be quoted in illustration of the delicacy of the observations and of the perspicuity employed in interpreting the facts.
The majority of orchids differ from other seed plants (with the exception of the Asclepiads) in having no dust-like pollen. The pollen, or more correctly, the pollen-tetrads, remain fastened together as club-shaped pollinia usually borne on a slender pedicel. At the base of the pedicel is a small viscid disc by which the pollinium is attached to the head or proboscis of one of the insects which visit the flower. Darwin demonstrated that in Orchis and other flowers the pedicel of the pollinium, after its removal from the anther, undergoes a curving movement. If the pollinium was originally vertical, after a time it assumed a horizontal position. In the latter position, if the insect visited another flower, the pollinium would exactly hit the sticky stigmatic surface and thus effect fertilisation. The relation between the behaviour of the viscid disc and the secretion of nectar by the flower is especially remarkable. The flowers possess a spur which in some species (e.g. Gymnadenia conopsea, Platanthera bifolia, etc.) contains honey (nectar), which serves as an attractive bait for insects, but in others (e.g. our native species of Orchis) the spur is empty. Darwin held the opinion, confirmed by later investigations, that in the case of flowers without honey the insects must penetrate the wall of the nectarless spurs in order to obtain a nectar-like substance. The glands behave differently in the nectar-bearing and in the nectarless flowers. In the former they are so sticky that they at once adhere to the body of the insect; in the nectarless flowers firm adherence only occurs after the viscid disc has hardened. It is, therefore, adaptively of value that the insects should be detained longer in the nectarless flowers (by having to bore into the spur),--than in flowers in which the nectar is freely exposed. "If this relation, on the one hand, between the viscid matter requiring some little time to set hard, and the nectar being so lodged that moths are delayed in getting it; and, on the other hand, between the viscid matter being at first as viscid as ever it will become, and the nectar lying all ready for rapid suction, be accidental, it is a fortunate accident for the plant. If not accidental, and I cannot believe it to be accidental, what a singular case of adaptation!" ("Fertilisation