Volume Ii Part 26 (2/2)
I thank you much for your remarks about my crossing notions, to which, I may add, I was led by exactly the same idea as yours, viz., that crossing must be one means of eliminating variation, and then I wished to make out how far in animals and vegetables this was possible.
Papilionaceous flowers are almost dead floorers to me, and I cannot experimentise, as castration alone often produces sterility. I am surprised at what you say about Compositae and Gramineae. From what I have seen of latter they seemed to me (and I have watched wheat, owing to what L. de Longchamps has said on their fertilisation in bud) favourable for crossing; and from Ca.s.sini's observations and Kolreuter's on the adhesive pollen, and C.C. Sprengel's, I had concluded that the Compositae were eminently likely (I am aware of the pistil brus.h.i.+ng out pollen) to be crossed. (586/1. This is an instance of the curious ignorance of the essential principles of floral mechanism which was to be found even among learned and accomplished botanists such as Gray, before the publication of the ”Fertilisation of Orchids.” Even in 1863 we find Darwin explaining the meaning of dichogamy in a letter to Gray.) If in some months' time you can find time to tell me whether you have made any observations on the early fertilisation of plants in these two orders, I should be very glad to hear, as it would save me from great blunder. In several published remarks on this subject in various genera it has seemed to me that the early fertilisation has been inferred from the early shedding of the pollen, which I think is clearly a false inference. Another cause, I should think, of the belief of fertilisation in the bud, is the not-rare, abnormal, early maturity of the pistil as described by Gartner. I have hitherto failed in meeting with detailed accounts of regular and normal impregnation in the bud. Podostemon and Subularia under water (and Leguminosae) seem and are strongest cases against me, as far as I as yet know. I am so sorry that you are so overwhelmed with work; it makes your VERY GREAT kindness to me the more striking.
It is really pretty to see how effectual insects are. A short time ago I found a female holly sixty measured yards from any other holly, and I cut off some twigs and took by chance twenty stigmas, cut off their tops, and put them under the microscope: there was pollen on every one, and in profusion on most! weather cloudy and stormy and unfavourable, wind in wrong direction to have brought any.
LETTER 587. TO J.D. HOOKER. Down, January 12th [1858].
I want to ask a question which will take you only few words to answer.
It bears on my former belief (and Asa Gray strongly expressed opinion) that Papilionaceous flowers were fatal to my notion of there being no eternal hermaphrodites. First let me say how evidence goes. You will remember my facts going to show that kidney-beans require visits of bees to be fertilised. This has been positively stated to be the case with Lathyrus grandiflorus, and has been very partially verified by me.
Sir W. Macarthur tells me that Erythrina will hardly seed in Australia without the petals are moved as if by bee. I have just met the statement that, with common bean, when the humble-bees bite holes at the base of the flower, and therefore cease visiting the mouth of the corolla, ”hardly a bean will set.” But now comes a much more curious statement, that [in] 1842-43, ”since bees were established at Wellington (New Zealand), clover seeds all over the settlement, WHICH IT DID NOT BEFORE.” (587/1. See Letter 362, Volume I.) The writer evidently has no idea what the connection can be. Now I cannot help at once connecting this statement (and all the foregoing statements in some degree support each other, as all have been advanced without any sort of theory) with the remarkable absence of Papilionaceous plants in N. Zealand. I see in your list Clianthus, Carmichaelia (four species), a new genus, a shrub, and Edwardsia (is latter Papilionaceous?). Now what I want to know is whether any of these have flowers as small as clover; for if they have large flowers they may be visited by humble-bees, which I think I remember do exist in New Zealand; and which humble-bees would not visit the smaller clover. Even the very minute little yellow clover in England has every flower visited and revisited by hive-bees, as I know by experience. Would it not be a curious case of correlation if it could be shown to be probable that herbaceous and small Leguminosae do not exist because when [their] seeds [are] washed ash.o.r.e (!!!) no small bees exist there. Though this latter fact must be ascertained. I may not prove anything, but does it not seem odd that so many quite independent facts, or rather statements, should point all in one direction, viz., that bees are necessary to the fertilisation of Papilionaceous flowers?
LETTER 588. TO JOHN LUBBOCK (Lord Avebury). Sunday [1859].
Do you remember calling my attention to certain flowers in the truss of Pelargoniums not being true, or not having the dark shade on the two upper petals? I believe it was Lady Lubbock's observation. I find, as I expected, it is always the central or sub-central flower; but what is far more curious, the nectary, which is blended with the peduncle of the flowers, gradually lessens and quite disappears (588/1. This fact is mentioned in Maxwell Masters' ”Vegetable Teratology” (Ray Society's Publications), 1869, page 221.), as the dark shade on the two upper petals disappears. Compare the stalk in the two enclosed parcels, in each of which there is a perfect flower.
Now, if your gardener will not be outrageous, do look over your geraniums and send me a few trusses, if you can find any, having the flowers without the marks, sending me some perfect flowers on same truss. The case seems to me rather a pretty one of correlation of growth; for the calyx also becomes slightly modified in the flowers without marks.
LETTER 589. TO MAXWELL MASTERS. Down, April 7th [1860].
I hope that you will excuse the liberty which I take in writing to you and begging a favour. I have been very much interested by the abstract (too brief) of your lecture at the Royal Inst.i.tution. Many of the facts alluded to are full of interest for me. But on one point I should be infinitely obliged if you could procure me any information: namely, with respect to sweet-peas. I am a great believer in the natural crossing of individuals of the same species. But I have been a.s.sured by Mr. Cattell (589/1. The nurseryman he generally dealt with.), of Westerham, that the several varieties of sweet-pea can be raised close together for a number of years without intercrossing. But on the other hand he stated that they go over the beds, and pull up any false plant, which they very naturally attribute to wrong seeds getting mixed in the lot. After many failures, I succeeded in artificially crossing two varieties, and the offspring out of the same pod, instead of being intermediate, was very nearly like the two pure parents; yet in one, there was a trace of the cross, and these crossed peas in the next generation showed still more plainly their mongrel origin. Now, what I want to know is, whether there is much variation in sweet-peas which might be owing to natural crosses.
What I should expect would be that they would keep true for many years, but that occasionally, perhaps at long intervals, there would be a considerable amount of crossing of the varieties grown close together.
Can you give, or obtain from your father, any information on this head, and allow me to quote your authority? It would really be a very great favour and kindness.
LETTER 590. TO J.D. HOOKER.
(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].
<script>