A letter to Dr. Asa Gray (September 5th, 1857) gives the substance of the paper in the "Gardeners" Chronicle":--
"Lately I was led to examine buds of kidney bean with the pollen shed; but I was led to believe that the pollen could HARDLY get on the stigma by wind or otherwise, except by bees visiting [the flower] and moving the wing petals: hence I included a small bunch of flowers in two bottles in every way treated the same: the flowers in one I daily just momentarily moved, as if by a bee; these set three fine pods, the other NOT ONE. Of course this little experiment must be tried again, and this year in England it is too late, as the flowers seem now seldom to set.
If bees are necessary to this flower"s self-fertilisation, bees must almost cross them, as their dusted right-side of head and right legs constantly touch the stigma.
"I have, also, lately been re-observing daily Lobelia fulgens--this in my garden is never visited by insects, and never sets seeds, without pollen be put on the stigma (whereas the small blue Lobelia is visited by bees and does set seed); I mention this because there are such beautiful contrivances to prevent the stigma ever getting its own pollen; which seems only explicable on the doctrine of the advantage of crosses."
The paper was supplemented by a second in 1858. ("Gardeners" Chronicle", 1858, page 828. In 1861 another paper on Fertilisation appeared in the "Gardeners" Chronicle", page 552, in which he explained the action of insects on Vinca major. He was attracted to the periwinkle by the fact that it is not visited by insects and never set seeds.) The chief object of these publications seems to have been to obtain information as to the possibility of growing varieties of leguminous plants near each other, and yet keeping them true. It is curious that the Papilionaceae should not only have been the first flowers which attracted his attention by their obvious adaptation to the visits of insects, but should also have const.i.tuted one of his sorest puzzles. The common pea and the sweet pea gave him much difficulty, because, although they are as obviously fitted for insect-visits as the rest of the order, yet their varieties keep true. The fact is that neither of these plants being indigenous, they are not perfectly adapted for fertilisation by British insects. He could not, at this stage of his observations, know that the co-ordination between a flower and the particular insect which fertilises it may be as delicate as that between a lock and its key, so that this explanation was not likely to occur to him. (He was of course alive to variety in the habits of insects. He published a short note in the "Entomologists Weekly Intelligencer", 1860, asking whether the Tineina and other small moths suck flowers.)
Besides observing the Leguminosae, he had already begun, as shown in the foregoing extracts, to attend to the structure of other flowers in relation to insects. At the beginning of 1860 he worked at Leschenaultia (He published a short paper on the manner of fertilisation of this flower, in the "Gardeners" Chronicle", 1871, page 1166.), which at first puzzled him, but was ultimately made out. A pa.s.sage in a letter chiefly relating to Leschenaultia seems to show that it was only in the spring of 1860 that he began widely to apply his knowledge to the relation of insects to other flowers. This is somewhat surprising, when we remember that he had read Sprengel many years before. He wrote (May 14):--
"I should look at this curious contrivance as specially related to visits of insects; as I begin to think is almost universally the case."
Even in July 1862 he wrote to Dr. Asa Gray:--
"There is no end to the adaptations. Ought not these cases to make one very cautious when one doubts about the use of all parts? I fully believe that the structure of all irregular flowers is governed in relation to insects. Insects are the Lords of the floral (to quote the witty "Athenaeum") world."
He was probably attracted to the study of Orchids by the fact that several kinds are common near Down. The letters of 1860 show that these plants occupied a good deal of his attention; and in 1861 he gave part of the summer and all the autumn to the subject. He evidently considered himself idle for wasting time on Orchids, which ought to have been given to "Variation under Domestication." Thus he wrote:--
"There is to me incomparably more interest in observing than in writing; but I feel quite guilty in trespa.s.sing on these subjects, and not sticking to varieties of the confounded c.o.c.ks, hens and ducks. I hear that Lyell is savage at me. I shall never resist Linum next summer."
It was in the summer of 1860 that he made out one of the most striking and familiar facts in the book, namely, the manner in which the pollen ma.s.ses in Orchis are adapted for removal by insects. He wrote to Sir J.D. Hooker July 12:--
"I have been examining Orchis pyramidalis, and it almost equals, perhaps even beats, your Listera case; the sticky glands are congenitally united into a saddle-shaped organ, which has great power of movement, and seizes hold of a bristle (or proboscis) in an admirable manner, and then another movement takes place in the pollen ma.s.ses, by which they are beautifully adapted to leave pollen on the two LATERAL stigmatic surfaces. I never saw anything so beautiful."
In June of the same year he wrote:--
"You speak of adaptation being rarely VISIBLE, though present in plants.
I have just recently been looking at the common Orchis, and I declare I think its adaptations in every part of the flower quite as beautiful and plain, or even more beautiful than in the Woodp.e.c.k.e.r. I have written and sent a notice for the "Gardeners" Chronicle" (June 9, 1860. This seems to have attracted some attention, especially among entomologists, as it was reprinted in the "Entomologists Weekly Intelligencer", 1860.), on a curious difficulty in the Bee Orchis, and should much like to hear what you think of the case. In this article I have incidentally touched on adaptation to visits of insects; but the contrivance to keep the sticky glands fresh and sticky beats almost everything in nature. I never remember having seen it described, but it must have been, and, as I ought not in my book to give the observation as my own, I should be very glad to know where this beautiful contrivance is described."
He wrote also to Dr. Gray, June 8, 1860:--
"Talking of adaptation, I have lately been looking at our common orchids, and I dare say the facts are as old and well-known as the hills, but I have been so struck with admiration at the contrivances, that I have sent a notice to the "Gardeners" Chronicle". The Ophrys apifera, offers, as you will see, a curious contradiction in structure."
Besides attending to the fertilisation of the flowers he was already, in 1860, busy with the h.o.m.ologies of the parts, a subject of which he made good use in the Orchid book. He wrote to Sir Joseph Hooker (July):--
"It is a real good joke my discussing h.o.m.ologies of Orchids with you, after examining only three or four genera; and this very fact makes me feel positive I am right! I do not quite understand some of your terms; but sometime I must get you to explain the h.o.m.ologies; for I am intensely interested on the subject, just as at a game of chess."
This work was valuable from a systematic point of view. In 1880 he wrote to Mr. Bentham:--
"It was very kind in you to write to me about the Orchideae, for it has pleased me to an extreme degree that I could have been of the LEAST use to you about the nature of the parts."
The pleasure which his early observations on Orchids gave him is shown in such extracts as the following from a letter to Sir J.D. Hooker (July 27, 1861):--
"You cannot conceive how the Orchids have delighted me. They came safe, but box rather smashed; cylindrical old cocoa- or snuff-canister much safer. I enclose postage. As an account of the movement, I shall allude to what I suppose is Oncidium, to make CERTAIN,--is the enclosed flower with crumpled petals this genus? Also I most specially want to know what the enclosed little globular brown Orchid is. I have only seen pollen of a Cattleya on a bee, but surely have you not unintentionally sent me what I wanted most (after Catasetum or Mormodes), viz. one of the Epidendreae?! I PARTICULARLY want (and will presently tell you why) another spike of this little Orchid, with older flowers, some even almost withered."
His delight in observation is again shown in a letter to Dr. Gray (1863). referring to Cruger"s letters from Trinidad, he wrote:--"Happy man, he has actually seen crowds of bees flying round Catasetum, with the pollinia sticking to their backs!"
The following extracts of letters to Sir J.D. Hooker ill.u.s.trate further the interest which his work excited in him:--
"Veitch sent me a grand lot this morning. What wonderful structures!
"I have now seen enough, and you must not send me more, for though I enjoy looking at them MUCH, and it has been very useful to me, seeing so many different forms, it is idleness. For my object each species requires studying for days. I wish you had time to take up the group.
I would give a good deal to know what the rostellum is, of which I have traced so many curious modifications. I suppose it cannot be one of the stigmas (It is a modification of the upper stigma.), there seems a great tendency for two lateral stigmas to appear. My paper, though touching on only subordinate points will run, I fear, to 100 MS. folio pages!
The beauty of the adaptation of parts seems to me unparalleled. I should think or guess waxy pollen was most differentiated. In Cypripedium which seems least modified, and a much exterminated group, the grains are single. In ALL OTHERS, as far as I have seen, they are in packets of four; and these packets cohere into many wedge-formed ma.s.ses in Orchis; into eight, four, and finally two. It seems curious that a flower should exist, which could AT MOST fertilise only two other flowers, seeing how abundant pollen generally is; this fact I look at as explaining the perfection of the contrivance by which the pollen, so important from its fewness, is carried from flower to flower" (1861).
"I was thinking of writing to you to-day, when your note with the Orchids came. What frightful trouble you have taken about Vanilla; you really must not take an atom more; for the Orchids are more play than real work. I have been much interested by Epidendrum, and have worked all morning at them; for heaven"s sake, do not corrupt me by any more"
(August 30, 1861).
He originally intended to publish his notes on Orchids as a paper in the Linnean Society"s Journal, but it soon became evident that a separate volume would be a more suitable form of publication. In a letter to Sir J.D. Hooker, September 24, 1861, he writes:--
"I have been acting, I fear that you will think, like a goose; and perhaps in truth I have. When I finished a few days ago my Orchis paper, which turns out 140 folio pages!! and thought of the expense of woodcuts, I said to myself, I will offer the Linnean Society to withdraw it, and publish it in a pamphlet. It then flashed on me that perhaps Murray would publish it, so I gave him a cautious description, and offered to share risks and profits. This morning he writes that he will publish and take all risks, and share profits and pay for all ill.u.s.trations. It is a risk, and heaven knows whether it will not be a dead failure, but I have not deceived Murray, and [have] told him that it would interest those alone who cared much for natural history. I hope I do not exaggerate the curiosity of the many special contrivances."
He wrote the two following letters to Mr. Murray about the publication of the book:]
Down, September 21 [1861].
My dear Sir,
Will you have the kindness to give me your opinion, which I shall implicitly follow. I have just finished a very long paper intended for Linnean Society (the t.i.tle is enclosed), and yesterday for the first time it occurred to me that POSSIBLY it might be worth publishing separately which would save me trouble and delay. The facts are new, and have been collected during twenty years and strike me as curious. Like a Bridgewater treatise, the chief object is to show the perfection of the many contrivances in Orchids. The subject of propagation is interesting to most people, and is treated in my paper so that any woman could read it. Parts are dry and purely scientific; but I think my paper would interest a good many of such persons who care for Natural History, but no others.
... It would be a very little book, and I believe you think very little books objectionable. I have myself GREAT doubts on the subject. I am very apt to think that my geese are swans; but the subject seems to me curious and interesting.
I beg you not to be guided in the least in order to oblige me, but as far as you can judge, please give me your opinion. If I were to publish separately, I would agree to any terms, such as half risk and half profit, or what you liked; but I would not publish on my sole risk, for to be frank, I have been told that no publisher whatever, under such circ.u.mstances, cares for the success of a book.
CHARLES DARWIN TO J. MURRAY. Down, September 24 [1861].
My dear Sir,
I am very much obliged for your note and very liberal offer. I have had some qualms and fears. All that I can feel sure of is that the MS.
contains many new and curious facts, and I am sure the Essay would have interested me, and will interest those who feel lively interest in the wonders of nature; but how far the public will care for such minute details, I cannot at all tell. It is a bold experiment; and at worst, cannot entail much loss; as a certain amount of sale will, I think, be pretty certain. A large sale is out of the question. As far as I can judge, generally the points which interest me I find interest others; but I make the experiment with fear and trembling,--not for my own sake, but for yours...
[On September 28th he wrote to Sir J.D. Hooker:--
"What a good soul you are not to sneer at me, but to pat me on the back.
I have the greatest doubt whether I am not going to do, in publishing my paper, a most ridiculous thing. It would annoy me much, but only for Murray"s sake, if the publication were a dead failure."
There was still much work to be done, and in October he was still receiving Orchids from Kew, and wrote to Hooker:--
"It is impossible to thank you enough. I was almost mad at the wealth of Orchids." And again--
"Mr. Veitch most generously has sent me two splendid buds of Mormodes, which will be capital for dissection, but I fear will never be irritable; so for the sake of charity and love of heaven do, I beseech you, observe what movement takes place in Cychnoches, and what part must be touched. Mr. V. has also sent me one splendid flower of Catasetum, the most wonderful Orchid I have seen."
On October 13th he wrote to Sir Joseph Hooker:--
"It seems that I cannot exhaust your good nature. I have had the hardest day"s work at Catasetum and buds of Mormodes, and believe I understand at last the mechanism of movements and the functions. Catasetum is a beautiful case of slight modification of structure leading to new functions. I never was more interested in any subject in my life than in this of Orchids. I owe very much to you."