Чарльз Дарвин
More Letters of Charles Darwin — Volume 2
With respect to your case of Aristolochia, I think further observation would convince you that it is not fertilised only by larvae, for in a nearly parallel case of an Arum and a Aristolochia, I found that insects flew from flower to flower. I would suggest to you to observe any cases of flowers which catch insects by their probosces, as occurs with some of the Apocyneae (593/1. Probably Asclepiadeae. See H. Muller, "Fertilisation of Flowers," page 396.); I have never been able to conceive for what purpose (if any) this is effected; at the same time, if I tempt you to neglect your zoological work for these miscellaneous observations I shall be guilty of a great crime.
To return for a moment to the indusium: how curious it is that the pollen should be thus collected in a special receptacle, afterwards to be swept out by insects' agency!
I am surprised at what you tell me about the fewness of the flowers of your native orchids which produce seed-capsules. What a contrast with our temperate European species, with the exception of some species of Ophrys! — I now know of three or four cases of self-fertilising orchids, but all these are provided with means for an occasional cross.
I am sorry to say Dr. Cruger is dead from a fever.
I received yesterday your paper in the "Botanische Zeitung" on the wood of climbing plants. (593/2. Fritz Muller, "Ueber das Holz einiger um Desterro wachsenden Kletterpflanzen." "Botanische Zeitung," 1866, pages 57, 65.) I have read as yet only your very interesting and curious remarks on the subject as bearing on the change of species; you have pleased me by the very high compliments which you pay to my paper. I have been at work since March 1st on a new English edition (593/3. The 4th Edition.) of my "Origin," of which when published I will send you a copy. I have much regretted the time it has cost me, as it has stopped my other work. On the other hand, it will be useful for a new third German edition, which is now wanted. I have corrected it largely, and added some discussions, but not nearly so much as I wished to do, for, being able to work only two hours daily, I feared I should never get it finished. I have taken some facts and views from your work "Fur Darwin"; but not one quarter of what I should like to have quoted.
LETTER 594. TO A.G. MORE. Down, June 24th, 1860.
I hope that you will forgive the liberty which I take in writing to you and requesting a favour. Mr. H.C. Watson has given me your address, and has told me that he thought that you would be willing to oblige me. Will you please to read the enclosed, and then you will understand what I wish observed with respect to the bee-orchis. (594/1. Ophrys apifera.) What I especially wish, from information which I have received since publishing the enclosed, is that the state of the pollen-masses should be noted in flowers just beginning to wither, in a district where the bee-orchis is extremely common. I have been assured that in parts of Isle of Wight, viz., Freshwater Gate, numbers occur almost crowded together: whether anything of this kind occurs in your vicinity I know not; but, if in your power, I should be infinitely obliged for any information. As I am writing, I will venture to mention another wish which I have: namely, to examine fresh flowers and buds of the Aceras, Spiranthes, marsh Epipactis, and any other rare orchis. The point which I wish to examine is really very curious, but it would take too long space to explain. Could you oblige me by taking the great trouble to send me in an old tin canister any of these orchids, permitting me, of course, to repay postage? It would be a great kindness, but perhaps I am unreasonable to make such a request. If you will inform me whether you have leisure so far to oblige me, I would tell you my movements, for on account of my own health and that of my daughter, I shall be on the move for the next two or three weeks.
I am sure I have much cause to apologise for the liberty which I have taken...
LETTER 595. TO A.G. MORE. Down, August 3rd, 1860.
I thank you most sincerely for sending me the Epipactis {palustris}. You can hardly imagine what an interesting morning's work you have given me, as the rostellum exhibited a quite new modification of structure. It has been extremely kind of you to take so very much trouble for me. Have you looked at the pollen-masses of the bee-Ophrys? I do not know whether the Epipactis grows near to your house: if it does, and any object takes you to the place (pray do not for a moment think me so very unreasonable as to ask you to go on purpose), would you be so kind {as} to watch the flowers for a quarter of an hour, and mark whether any insects (and what?) visit these flowers.
I should suppose they would crawl in by depressing the terminal portion of the labellum; and that when within the flower this terminal portion would resume its former position; and lastly, that the insect in crawling out would not depress the labellum, but would crawl out at back of flower. (595/1. The observations of Mr. William Darwin on Epipactis palustris given in the "Fertilisation of Orchids," Edition II., 1877, page 99, bear on this point. The chief fertilisers are hive-bees, which are too big to crawl into the flower. They cling to the labellum, and by depressing it open up the entrance to the flower. Owing to the elasticity of the labellum and its consequent tendency to spring up when released, the bees, "as they left the flower, seemed to fly rather upwards." This agrees with Darwin's conception of the mechanism of the flower as given in the first edition of the Orchid book, 1862, page 100, although at that time he imagined that the fertilising insect crawled into the flower. The extreme flexibility and elasticity of the labellum was first observed by Mr. More (see first edition, page 99). The description of the flower given in the above letter to Mr. More is not quite clear; the reader is referred to the "Fertilisation of Orchids," loc. cit.) An insect crawling out of a recently opened flower would, I believe, have parts of the pollen-masses adhering to the back or shoulder. I have seen this in Listera. How I should like to watch the Epipactis.
If you can it any time send me Spiranthes or Aceras or O. ustulata, you would complete your work of kindness.
P.S. — If you should visit the Epipactis again, would you gather a few of the lower flowers which have been opened for some time and have begun to wither a little, and observe whether pollen is well cleared out of anther-case. I have been struck with surprise that in nearly all the lower flowers sent by you, though much of the pollen has been removed, yet a good deal of pollen is left wasted within the anthers. I observed something of this kind in Cephalanthera grandiflora. But I fear that you will think me an intolerable bore.
LETTER 596. TO A.G. MORE. Down, August 5th, 1860.
I am infinitely obliged for your most clearly stated observations on the bee-orchis. It is now perfectly clear that something removes the pollen-masses far more with you than in this neighbourhood. But I am utterly puzzled about the foot-stalk being so often cut through. I should suspect snails. I yesterday found thirty-nine flowers, and of them only one pollen-mass in three flowers had been removed, and as these were extremely much-withered flowers I am not quite sure of the truth of this. The wind again is a new element of doubt. Your observations will aid me extremely in coming to some conclusion. (596/1. Mr. More's observations on the percentage of flowers in which the pollinia were absent are quoted in "Fertilisation of Orchids," Edition I., page 68.) I hope in a day or two to receive some day-moths, on the probosces of which I am assured the pollen-masses of the bee-orchis still adhere (596/2. He was doomed to disappointment. On July 17th, 1861, he wrote to Mr. More: — "I found the other day a lot of bee-Ophrys with the glands of the pollinia all in their pouches. All facts point clearly to eternal self-fertilisation in this species; yet I cannot swallow the bitter pill. Have you looked at any this year?")...
I wrote yesterday to thank you for the Epipactis. For the chance of your liking to look at what I have found: take a recently opened flower, drag gently up the stigmatic surface almost any object (the side of a hooked needle), and you will find the cap of the hemispherical rostellum comes off with a touch, and being viscid on under-surface, clings to needle, and as pollen-masses are already attached to the back of rostellum, the needle drags out much pollen. But to do this, the curiously projecting and fleshy summits of anther-cases must at some time be pushed back slightly. Now when an insect's head gets into the flower, when the flap of the labellum has closed by its elasticity, the insect would naturally creep out by the back-side of the flower. And mark when the insect flies to another flower with the pollen-masses adhering to it, if the flap of labellum did not easily open and allow free ingress to the insect, it would surely rub off the pollen on the upper petals, and so not leave it on stigma. It is to know whether I have rightly interpreted the structure of this whole flower that I am so curious to see how insects act. Small insects, I daresay, would crawl in and out and do nothing. I hope that I shall not have wearied you with these details.
If you would like to see a pretty and curious little sight, look to Orchis pyramidalis, and you will see that the sticky glands are congenitally united into a saddle-shaped organ. Remove this under microscope by pincers applied to foot-stalk of pollen-mass, and look quickly at the spontaneous movement of the saddle-shaped organs and see how beautifully adapted to seize proboscis of moth.
LETTER 597. TO J.D. HOOKER December 4th {1860}.
Many thanks about Apocynum and Meyen.
The latter I want about some strange movements in cells of Drosera, which Meyen alone seems to have observed. (597/1. No observations of Meyen are mentioned in "Insectivorous Plants.") It is very curious, but Trecul disbelieves that Drosera really clasps flies! I should very much wish to talk over Drosera with you. I did chloroform it, and the leaves which were already expanded did not recover thirty seconds of exposure for three days. I used the expression weight for the bit of hair which caused movement and weighed 1/78000 of a grain; but I do not believe it is weight, and what it is, I cannot after many experiments conjecture. (597/2. The doubt here expressed as to whether the result is due to actual weight is interesting in connection with Pfeffer's remarkable discovery that a smooth object in contact with the gland produces no effect if the plant is protected from all vibration; on an ordinary table the slight shaking which reaches the plant is sufficient to make the body resting on the gland tremble, and thus produce a series of varying pressures — under these circumstances the gland is irritated, and the tentacle moves. See Pfeffer, "Untersuchungen aus d. bot. Institut zu Tubingen," Volume I., 1885, page 483; also "Insectivorous Plants," Edition II., page 22.) The movement in this case does not depend on the chemical nature of substance. Latterly I have tried experiments on single glands, and a microscopical atom of raw meat causes such rapid movement that I could see it move like hand of clock. In this case it is the nature of the object. It is wonderful the rapidity of the absorption: in ten seconds weak solution of carbonate of ammonia changes not the colour, but the state of contents within the glands. In two minutes thirty seconds juice of meat has been absorbed by gland and passed from cell to cell all down the pedicel (or hair) of the gland, and caused the sap to pass from the cells on the upper side of the pedicel to the lower side, and this causes the curvature of the pedicel. I shall work away next summer when Drosera opens again, for I am much interested in subject. After the glandular hairs have curved, the oddest changes take place — viz., a segregation of the homogeneous pink fluid and necessary slow movements in the thicker matter. By Jove, I sometimes think Drosera is a disguised animal! You know that I always so like telling you what I do, that you must forgive me scribbling on my beloved Drosera. Farewell. I am so very glad that you are going to reform your ways; I am sure that you would have injured your health seriously. There is poor Dana has done actually nothing — cannot even write a letter — for a year, and it is hoped that in another YEAR he may quite recover.
After this homily, good night, my dear friend. Good heavens, I ought not to scold you, but thank you, for writing so long and interesting a letter.
LETTER 598. TO E. CRESY. Down, December 12th {1860?}.
After writing out the greater part of my paper on Drosera, I thought of so many points to try, and I wished to re-test the basis of one large set of experiments, namely, to feel still more sure than I am, that a drop of plain water never produces any effect, that I have resolved to publish nothing this year. For I found in the record of my daily experiments one suspicious case. I must wait till next summer. It will be difficult to try any solid substances containing nitrogen, such as ivory; for two quite distinct causes excite the movement, namely, mechanical irritation and presence of nitrogen. When a solid substance is placed on leaf it becomes clasped, but is released sooner than when a nitrogenous solid is clasped; yet it is difficult (except with raw meat and flies) to be sure of the result, owing to differences in vigour of different plants. The last experiments which I tried before my plants became too languid are very curious, and were tried by putting microscopical atoms on the gland itself of single hairs; and it is perfectly evident that an atom of human hair, 1/76000 of a grain (as ascertained by weighing a length of hair) in weight, causes conspicuous movement. I do not believe (for atoms of cotton thread acted) it is the chemical nature; and some reasons make me doubt whether it is actual weight; it is not the shadow; and I am at present, after many experiments, confounded to know what the cause is. That these atoms did really act and alter the state of the contents of all the cells in the glandular hair, which moved, was perfectly clear. But I hope next summer to make out a good deal more...
LETTER 599. TO J.D. HOOKER. Down, May 14th {1861}.
I have been putting off writing from day to day, as I did not wish to trouble you, till my wish for a little news will not let me rest...
I have no news to tell you, for I have had no interesting letters for some time, and have not seen a soul. I have been going through the "Cottage Gardener" of last year, on account chiefly of Beaton's articles (599/1. Beaton was a regular contributor to the "Cottage Gardener," and wrote various articles on cross breeding, etc., in 1861. One of these was in reply to a letter published in the "Cottage Gardener," May 14th, 1861, page 112, in which Darwin asked for information as to the Compositae and the hollyhock being crossed by insect visitors. In the number for June 8th, 1861, page 211, Darwin wrote on the variability of the central flower of the carrot and the peloria of the central flower in Pelargonium. An extract from a letter by Darwin on Leschenaultia, "Cottage Gardener," May 28th, 1861, page 151, is given in Letter 590, note.); he strikes me as a clever but d — d cock-sure man (as Lord Melbourne said), and I have some doubts whether to be much trusted. I suspect he has never recorded his experiment at the time with care. He has made me indignant by the way he speaks of Gartner, evidently knowing nothing of his work. I mean to try and pump him in the "Cottage Gardener," and shall perhaps defend Gartner. He alludes to me occasionally, and I cannot tell with what spirit. He speaks of "this Mr. Darwin" in one place as if I were a very noxious animal.
Let me have a line about poor Henslow pretty soon.
(599/2. In a letter of May 18th, 1861, Darwin wrote again: — )
By the way, thanks about Beaton. I have now read more of his writings, and one answer to me in "Cottage Gardener." I can plainly see that he is not to be trusted. He does not well know his own subject of crossing.
LETTER 600. TO J.D. HOOKER.
(600/1. Part of this letter has been published in "Life and Letters," III., page 265.)
2, Hesketh Crescent, Torquay {1861}.
...The beauty of the adaptation of parts seems to me unparalleled. I should think or guess {that} 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 masses 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. By the way, Cephalanthera has single pollen-grains, but this seems to be a case of degradation, for the rostellum is utterly aborted. Oddly, the columns of pollen are here kept in place by very early penetration of pollen-tubes into the edge of the stigma; nevertheless, it receives more pollen by insect agency. Epithecia {Dichaea} has done me one good little turn. I often speculated how the caudicle of Orchis had been formed. (600/2. The gradation here suggested is thoroughly worked out in the "Fertilisation of Orchids," Edition I., page 323, Edition II., page 257.) I had noticed slight clouds in the substance half way down; I have now dissected them out, and I find they are pollen-grains fairly embedded and useless. If you suppose the pollen-grains to abort in the lower half of the pollinia of Epipactis, but the parallel elastic threads to remain and cohere, you have the caudicle of Orchis, and can understand the few embedded and functionless pollen-grains. I must not look at any more exotic orchids: hearty thanks for your offer. But if you would make one single observation for me on Cypripedium, I should be glad. Asa Gray writes to me that the outside of the pollen-masses is sticky in this genus; I find that the whole mass consists of pollen-grains immersed in a sticky brownish thick fluid. You could tell by a mere lens and penknife. If it is, as I find it, pollen could not get on the stigma without insect aid. Cypripedium confounds me much. I conjecture that drops of nectar are secreted by the surface of the labellum beneath the anthers and in front of the stigma, and that the shield over the anthers and the form of labellum is to compel insects to insert their proboscis all round both organs. (600/3. This view was afterwards given up.) It would be troublesome for you to look at this, as it is always bothersome to catch the nectar secreting, and the cup of the labellum gets filled with water by gardener's watering.
I have examined Listera ovata, cordata, and Neottia nidus avis: the pollen is uniform; I suspect you must have seen some observation founded on a mistake from the penetration and hardening of sticky fluid from the rostellum, which does penetrate the pollen a little.
It is mere virtue which makes me not wish to examine more orchids; for I like it far better than writing about varieties of cocks and hens and ducks. Nevertheless, I have just been looking at Lindley's list in the "Vegetable Kingdom," and I cannot resist one or two of his great division of Arethuseae, which includes Vanilla. And as I know so well the Ophreae, I should like (God forgive me) any one of the Satyriadae, Disidae and Corycidae.
I fear my long lucubrations will have wearied you, but it has amused me to write, so forgive me.
LETTER 601. TO J.D. HOOKER.
(601/1. Part of the following letter is published in the "Life and Letters," the remainder, with the omission of part bearing on the Glen Roy problem, is now given as an example of the varied botanical assistance Darwin received from Sir Joseph Hooker. For the part relating to Verbascum see the "Variation of Animals and Plants," Edition II., 1875, Volume II., page 83. The point is that the white and yellow flowered plants which occur in two species of Verbascum are undoubted varieties, yet "the sterility which results from the crossing of the differently coloured varieties of the same species is fully as great as that which occurs in many cases when distinct species are crossed."
The sterility of the long-styled form (B) of Linum grandiflorum, with its own pollen is described in "Forms of Flowers," Edition II., page 87: his conclusions on the short-styled form (A) differ from those in the present letter.)
September 28th {1861}.
I am going to beg for help, and I will explain why I want it.
You offer Cypripedium; I should be very glad of a specimen, and of any good-sized Vandeae, or indeed any orchids, for this reason: I never thought of publishing separately, and therefore did not keep specimens in spirits, and now I should be very glad of a few woodcuts to illustrate my few remarks on exotic orchids. If you can send me any, send them by post in a tin canister on middle of day of Saturday, October 5th, for Sowerby will be here.
Secondly: Have you any white and yellow varieties of Verbascum which you could give me, or propagate for me, or LEND me for a year? I have resolved to try Gartner's wonderful and repeated statement, that pollen of white and yellow varieties, whether used on the varieties or on DISTINCT species, has different potency. I do not think any experiment can be more important on the origin of species; for if he is correct we certainly have what Huxley calls new physiological species arising. I should require several species of Verbascum besides the white and yellow varieties of the same species. It will be tiresome work, but if I can anyhow get the plants, it shall be tried.
Thirdly: Can you give me seeds of any Rubiaceae of the sub-order Cinchoneae, as Spermacoce, Diodia, Mitchella, Oldenlandia? Asa Gray says they present two forms like Primula. I am sure that this subject is well worth working out. I have just almost proved a very curious case in Linum grandiflorum which presents two forms, A and B. Pollen of A is perfectly fertile on stigma of A. But pollen of B is absolutely barren on its own stigma; you might as well put so much flour on it. It astounded me to see the stigmas of B purple with its own pollen; and then put a few grains of similar-looking pollen of A on them, and the germen immediately and always swelled; those not thus treated never swelling.
Fourthly: Can you give me any very hairy Saxifraga (for their functions) {i.e. the functions of the hairs}?
I send you a resume of my requests, to save you trouble. Nor would I ask for so much aid if I did not think all these points well worth trying to investigate.
My dear old friend, a letter from you always does me a world of good. And, the Lord have mercy on me, what a return I make.
LETTER 602. TO J.D. HOOKER. Down, October 4th {1861}.
Will you have the kindness to read the enclosed, and look at the diagram. Six words will answer my question. It is not an important point, but there is to me an irresistible charm in trying to make out homologies. (602/1. In 1880 he wrote to Mr. Bentham: "It was very kind of 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." — "Life and Letters," III., page 264.) You know the membranous cup or clinandrum, in many orchids, behind the stigma and rostellum: it is formed of a membrane which unites the filament of the normal dorsal anther with the edges of the pistil. The clinandrum is largely developed in Malaxis, and is of considerable importance in retaining the pollinia, which as soon as the flower opens are quite loose.
The appearance and similarity of the tissues, etc., at once gives suspicion that the lateral membranes of the clinandrum are the two other and rudimentary anthers, which in Orchis and Cephalanthera, etc., exist as mere papillae, here developed and utilised.
Now for my question. Exactly in the middle of the filament of the normal anther, and exactly in the middle of the lateral membrane of the clinandrum, and running up to the same height, are quite similar bundles of spiral vessels; ending upwards almost suddenly. Now is not this structure a good argument that I interpret the homologies of the sides of clinandrum rightly? (602/2. Though Robert Brown made use of the spiral vessels of orchids, yet according to Eichler, "Bluthendiagramme," 1875, Volume I., page 184, Darwin was the first to make substantial additions to the conclusions deducible from the course of the vessels in relation to the problem of the morphology of these plants. Eichler gives Darwin's diagram side by side with that of Van Tieghem without attempting to decide between the differences in detail by which they are characterised.)
I find that the great Bauer does not draw very correctly! (602/3. F. Bauer, whom Pritzel calls "der grosste Pflanzenmaler." The reference is to his "Illustrations of Orchidaceous Plants, with Notes and Prefatory Remarks by John Lindley," London, 1830-38, Folio. See "Fertilisation of Orchids," Edition II., page 82.) And, good Heavens, what a jumble he makes on functions.
LETTER 603. TO J.D. HOOKER. Down, October 22nd. {1861}.
Acropera is a beast, — stigma does not open, everything seems contrived that it shall NOT be anyhow fertilised. There is something very odd about it, which could only be made out by incessant watching on several individual plants.
I never saw the very curious flower of Canna; I should say the pollen was deposited where it is to prevent inevitable self-fertilisation. You have no time to try the smallest experiment, else it would be worth while to put pollen on some stigmas (supposing that it does not seed freely with you). Anyhow, insects would probably carry pollen from flower to flower, for Kurr states the tube formed by pistil, stamen and "nectarblatt" secretes (I presume internally) much nectar. Thanks for sending me the curious flower.
Now I want much some wisdom; though I must write at considerable length, your answer may be very brief.
(FIGURE 8. — FLORAL DIAGRAM OF AN ORCHID. The "missing bundle" could not be found in some species.)
In R. Brown's admirable paper in the "Linnean Transacts." (603/4. Volume XVI., page 685.) he suggests (and Lindley cautiously agrees) that the flower of orchids consists of five whorls, the inner whorl of the two whorls of anthers being all rudimentary, and when the labellum presents ridges, two or three of the anthers of both whorls {are} combined with it. In the ovarium there are six bundles of vessels: R. Brown judged by transverse sections. It occurred to me, after what you said, to trace the vessels longitudinally, and I have succeeded well. Look at my diagram {Figure 8} (which please return, for I am transported with admiration at it), which shows the vessels which I have traced, one bundle to each of fifteen theoretical organs, and no more. You will see the result is nothing new, but it seems to confirm strongly R. Brown, for I have succeeded (perhaps he did, but he does not say so) in tracing the vessels belonging to each organ in front of each other to the same bundle in the ovarium: thus the vessels going to the lower sepal, to the side of the labellum, and to one stigma (when there are two) all distinctly branch from one ovarian bundle. So in other cases, but I have not completely traced (only seen) that going to the rostellum. But here comes my only point of novelty: in all orchids as yet looked at (even one with so simple a labellum as Gymnadenia and Malaxis) the vessels on the two sides of the labellum are derived from the bundle which goes to the lower sepal, as in the diagram. This leads me to conclude that the labellum is always a compound organ. Now I want to know whether it is conceivable that the vessels coming from one main bundle should penetrate an organ (the labellum) which receives its vessels from another main bundle? Does it not imply that all that part of the labellum which is supplied by vessels coming from a lateral bundle must be part of a primordially distinct organ, however closely the two may have become united? It is curious in Gymnadenia to trace the middle anterior bundle in the ovarium: when it comes to the orifice of the nectary it turns and runs right down it, then comes up the opposite side and runs to the apex of the labellum, whence each side of the nectary is supplied by vessels from the bundles, coming from the lower sepals. Hence even the thin nectary is essentially, I infer, tripartite; hence its tendency to bifurcation at its top. This view of the labellum always consisting of three organs (I believe four when thick, as in Mormodes, at base) seems to me to explain its great size and tripartite form, compared with the other petals. Certainly, if I may trust the vessels, the simple labellum of Gymnadenia consists of three organs soldered together. Forgive me for writing at such length; a very brief answer will suffice. I am desperately interested in the subject: the destiny of the whole human race is as nothing to the course of vessels of orchids...
