Chapter 246

(590/1. The genera Scaevola and Leschenaultia, to which the following letter refers, belong to the Goodeniaceae (Goodenovieae, Bentham & Hooker), an order allied to the Lobeliaceae, although the mechanism of fertilisation resembles rather more nearly that of Campanula. The characteristic feature of the flower in this order is the indusium, or, as Delpino (590/2. Delpino's observations on Dichogamy, summarised by Hildebrand in "Bot. Zeitung," 1870, page 634.) calls it, the "collecting cup": this cuplike organ is a development of the style, and serves the same function as the hairs on the style of Campanula, namely, that of taking the pollen from the anthers and presenting it to the visiting insect. During this stage the immature stigma is at the bottom of the cup, and though surrounded by pollen is incapable of being pollinated. In most genera of the order the pollen is pushed out of the indusium by the growth of the style or stigma, very much as occurs in Lobelia or the Compositae. Finally the style emerges from the indusium (590/3. According to Hamilton ("Proc. Linn. Soc. N. S. Wales," X., 1895, page 361) the stigma rarely grows beyond the indusium in Dampiera. In the same journal (1885-6, page 157, and IX., 1894, page 201) Hamilton has given a number of interesting observations on Goodenia, Scaevola, Selliera, Brunonia. There seem to be mechanisms for cross- and also for self-fertilisation.), the stigmas open out and are pollinated from younger flowers. The mechanism of fertilisation has been described by F. Muller (590/4. In a letter to Hildebrand published in the "Bot. Zeitung," 1868, page 113.), and more completely by Delpino (loc. cit.).

Mr. Bentham wrote a paper (590/5. "Linn. Soc. Journal," 1869, page 203.) on the style and stigma in the Goodenovieae, where he speaks of Mr. Darwin's belief that fertilisation takes place outside the indusium. This statement, which we imagine Mr. Bentham must have had from an unpublished source, was incomprehensible to him as long as he confined his work to such genera as Goodenia, Scaevola, Velleia, Coelogyne, in which the mechanism is much as above described; but on examining Leschenaultia the meaning became clear. Bentham writes of this genus:--"The indusium is usually described as broadly two-lipped, without any distinct stigma. The fact appears to be that the upper less prominent lip is stigmatic all over, inside and out, with a transverse band of short glandular hairs at its base outside, while the lower more prominent lip is smooth and glabrous, or with a tuft of rigid hairs. Perhaps this lower lip and the upper band of hairs are all that correspond to the indusium of other genera; and the so-called upper lip, outside of which impregnation may well take place, as observed by Mr. Darwin, must be regarded as the true stigma."

Darwin's interest in the Goodeniaceae was due to the mechanism being apparently fitted for self-fertilisation. In 1871 a writer signing himself F.W.B. made a communication to the "Gardeners' Chronicle" (590/6. 1871, page 1103.), in which he expresses himself as "agreeably surprised" to find Leschenaultia adapted for self-fertilisation, or at least for self-pollinisation. This led Darwin to publish a short note in the same journal, in which he describes the penetration of pollen-tubes into the viscid surface on the outside of the indusium. (590/7. 1871, page 1166. He had previously written in the "Journal of Horticulture and Cottage Gardener," May 28th, 1861, page 151:--"Leschenaultia formosa has apparently the most effective contrivance to prevent the stigma of one flower ever receiving a grain of pollen from another flower; for the pollen is shed in the early bud, and is there shut up round the stigma within a cup or indusium. But some observations led me to suspect that nevertheless insect agency here comes into play; for I found by holding a camel-hair pencil parallel to the pistil, and moving it as if it were a bee going to suck the nectar, the straggling hairs of the brush opened the lip of the indusium, entered it, stirred up the pollen, and brought out some grains. I did this to five flowers, and marked them. These five flowers all set pods; whereas only two other pods set on the whole plant, though covered with innumerable flowers...I wrote to Mr. James Drummond, at Swan River in Australia,...and he soon wrote to me that he had seen a bee cleverly opening the indusium and extracting pollen.") He also describes how a brush, pushed into the flower in imitation of an insect, presses "against the slightly projecting lower lip of the indusium, opens it, and some of the hairs enter and become smeared with pollen." The yield of pollen is therefore differently arranged in Leschenaultia; for in the more typical genera it depends on the growth of the style inside the indusium. Delpino, however (see Hildebrand's version, loc. cit.), describes a similar opening of the cup produced by pressure on the hairs in some genera of the order.)

Down, June 7th [1860].

Best and most beloved of men, I supplicate and entreat you to observe one point for me. Remember that the Goodeniaceae have weighed like an incubus for years on my soul. It relates to Scaevola microcarpa. I find that in bud the indusium collects all the pollen splendidly, but, differently from Leschenaultia, cannot be afterwards easily opened. Further, I find that at an early stage, when the flower first opens, a boat-shaped stigma lies at the bottom of the indusium, and further that this stigma, after the flower has some time expanded, grows very rapidly, when the plant is kept hot, and pushes out of the indusium a mass of pollen; and at same time two horns project at the corners of the indusium. Now the appearance of these horns makes me suppose that these are the stigmatic surfaces. Will you look to this? for if they be by the relative position of the parts (with indusium and stigma bent at right angles to style) [I am led to think] that an insect entering a flower could not fail to have [its] whole back (at the period when, as I have seen, a whole mass of pollen is pushed out) covered with pollen, which would almost certainly get rubbed on the two horns. Indeed, I doubt whether, without this aid, pollen would get on to the horns. What interests me in the case is the analogy in result with the Lobelia, but by very different means. In Lobelia the stigma, before it is mature, pushes by its circular brush of hairs the pollen out of the conjoined anthers; here the indusium collects pollen, and then the growth of the stigma pushes it out. In the course of about 1 1/2 hour, I found an indusium with hairs on the outer edge perfectly clogged with pollen, and horns protruded, which before the 1 1/2 hour had not one grain of pollen outside the indusium, and no trace of protruding horns. So you will see how I wish to know whether the horns are the true stigmatic surfaces. I would try the case experimentally by putting pollen on the horns, but my greenhouse is so cold, and my plant so small, and in such a little pot, that I suppose it would not seed...

The little length of stigmatic horns at the moment when pollen is forced out of the indusium, compared to what they ultimately attain, makes me fancy that they are not then mature or ready, and if so, as in Lobelia, each flower must be fertilised by pollen from another and earlier flower.

How curious that the indusium should first so cleverly collect pollen and then afterwards push it out! Yet how closely analogous to Campanula brushing pollen out of the anther and retaining it on hairs till the stigma is ready. I am going to try whether Campanula sets seed without insect agency.