Down, November 12th, 1862.
I thank you most sincerely for your kindness in writing to me, and for {your} very interesting letter. Your fact has surprised me greatly, and has alarmed me not a little, for if I am in error about Acropera I may be in error about Catasetum. Yet when I call to mind the state of the placentae in A. luteola, I am astonished that they should produce ovules. You will see in my book that I state that I did not look at the ovarium of A. Loddigesii. Would you have the kindness to send me word which end of the ovarium is meant by apex (that nearest the flower?), for I must try and get this species from Kew and look at its ovarium. I shall be extremely curious to hear whether the fruit, which is now maturing, produces a large number of good and plump seed; perhaps you may have seen the ripe capsules of other Vandeae, and may be able to form some conjecture what it ought to produce. In the young, unfertilised ovaria of many Vandeae there seemed an infinitude of ovules. In desperation it occurs to me as just possible, as almost everything in nature goes by gradation, that a properly male flower might occasionally produce a few seeds, in the same manner as female plants sometimes produce a little pollen. All your remarks seem to me excellent and very interesting, and I again thank you for your kindness in writing to me. I am pleased to observe that my description of the structure of Acropera seems to agree pretty well with what you have observed. Does it not strike you as very difficult to understand how insects remove the pollinia and carry them to the stigmas? Your suggestion that the mouth of the stigmatic cavity may become charged with viscid matter and thus secure the pollinia, and that the pollen-tubes may then protrude, seems very ingenious and new to me; but it would be very anomalous in orchids, i.e. as far as I have seen. No doubt, however, though I tried my best, I shall be proved wrong in many points. Botany is a new subject to me. With respect to the protrusion of pollen-tubes, you might like to hear (if you do not already know the fact) that, as I saw this summer, in the little imperfect flowers of Viola and Oxalis, which never open, the pollen-tubes always come out of the pollen-grain, whilst still in the anthers, and direct themselves in a beautiful manner to the stigma seated at some little distance. I hope that you will continue your very interesting observations.
LETTER 634. TO J. SCOTT. Down, November 19th {1862}.
I am much obliged for your letter, which is full of interesting matter. I shall be very glad to look at the capsule of the Acropera when ripe, and pray present my thanks to Mr. MacNab. (634/1. See Letter 608 (Lindley, December 15th, 1861). Also "Fertilisation of Orchids," Edition II., page 172, for an account of the observations on Acropera which were corrected by Scott.) I should like to keep it till I could get a capsule of some other member of the Vandeae for comparison, but ultimately all the seeds shall be returned, in case you would like to write any notice on the subject. It was, as I said (634/2. Letter 633.), only "in desperation" that I suggested that the flower might be a male and occasionally capable of producing a few seeds. I had forgotten Gartner's remark; in fact, I know only odds and ends of Botany, and you know far more. One point makes the above view more probable in Acropera than in other cases, viz. the presence of rudimentary placentae or testae, for I cannot hear that these have been observed in the male plants. They do not occur in male Lychnis dioica, but next spring I will look to male holly flowers. I fully admit the difficulty of similarity of stigmatic chamber in the two Acroperas. As far as I remember, the blunt end of pollen-mass would not easily even stick in the orifice of the chamber. Your view may be correct about abundance of viscid matter, but seems rather improbable. Your facts about female flowers occurring where males alone ought to occur is new to me; if I do not hear that you object, I will quote the Zea case on your authority in what I am now writing on the varieties of the maize. (634/3. See "Animals and Plants," Edition II., Volume I., page 339: "Mr. Scott has lately observed the rarer case of female flowers on a true male panicle, and likewise hermaphrodite flowers." Scott's paper on the subject is in "Trans. Bot. Soc. Edinburgh," Volume VIII. See Letter 151, Volume I.) I am glad to hear that you are now working on the most curious subject of parthenogenesis. I formerly fancied that I observed female Lychnis dioica seeded without pollen. I send by this post a paper on Primula, which may interest you. (634/4. "Linn. Soc. Journal," 1862.) I am working on the subject, and if you should ever observe any analogous case I should be glad to hear. I have added another very clever pamphlet by Prof. Asa Gray. Have you a copy of my Orchis book? If you have not, and would like one, I should be pleased to send one. I plainly see that you have the true spirit of an experimentalist and good observer. Therefore, I ask whether you have ever made any trials on relative fertility of varieties of plants (like those I quote from Gartner on the varieties of Verbascum). I much want information on this head, and on those marvellous cases (as some Lobelias and Crinum passiflora) in which a plant can be more easily fertilised by the pollen of another species than by its own good pollen. I am compelled to write in haste. With many thanks for your kindness.
LETTER 635. TO J. SCOTT. Down, 20th {1862?}.
What a magnificent capsule, and good Heavens, what a number of seeds! I never before opened pods of larger orchids. It did not signify a few seed being lost, as it would be hopeless to estimate number in comparison with other species. If you sow any, had you not better sow a good many? so I enclose small packet. I have looked at the seeds; I never saw in the British orchids nearly so many empty testae; but this goes for nothing, as unnatural conditions would account for it. I suspect, however, from the variable size and transparency, that a good many of the seeds when dry (and I have put the capsule on my chimney-piece) will shrivel up. So I will wait a month or two till I get the capsule of some large Vandeae for comparison. It is more likely that I have made some dreadful blunder about Acropera than that it should be male yet not a perfect male. May there be some sexual relation between A. Loddigesii and luteola; they seem very close? I should very much like to examine the capsule of the unimpregnated flower of A. Loddigesii. I have got both species from Kew, but whether we shall have skill to flower them I know not. One conjectures that it is imperfect male; I still should incline to think it would produce by seed both sexes. But you are right about Primula (and a very acute thought it was): the long-styled P. sinensis, homomorphically fertilised with own-form pollen, has produced during two successive homomorphic generations only long-styled plants. (635/1. In "Forms of Flowers," Edition II., page 216, a summary of the transmission of forms in the "homomorphic" unions of P. sinensis is given. Darwin afterwards used "illegitimate" for homomorphic, and "legitimate" for "heteromorphic" ("Forms of Flowers," Edition i., page 24).) The short-styled the same, i.e. produced short-styled for two generations with the exception of a single plant. I cannot say about cowslips yet. I should like to hear your case of the Primula: is it certainly propagated by seed?
LETTER 636. TO J. SCOTT. Down, December 3rd, {1862?}.
What a capital observer you are! and how well you have worked the primulas. All your facts are new to me. It is likely that I overrate the interest of the subject; but it seems to me that you ought to publish a paper on the subject. It would, however, greatly add to the value if you were to cover up any of the forms having pistil and anther of the same height, and prove that they were fully self-fertile. The occurrence of dimorphic and non-dimorphic species in the same genus is quite the same as I find in Linum. (636/1. Darwin finished his paper on Linum in December 1862, and it was published in the "Linn. Soc. Journal" in 1863.) Have any of the forms of Primula, which are non-dimorphic, been propagated for some little time by seed in garden? I suppose not. I ask because I find in P. sinensis a third rather fluctuating form, apparently due to culture, with stigma and anthers of same height. I have been working successive generations homomorphically of this Primula, and think I am getting curious results; I shall probably publish next autumn; and if you do not (but I hope you will) publish yourself previously, I should be glad to quote in abstract some of your facts. But I repeat that I hope you will yourself publish. Hottonia is dimorphic, with pollen of very different sizes in the two forms. I think you are mistaken about Siphocampylus, but I feel rather doubtful in saying this to so good an observer. In Lobelia the closed pistil grows rapidly, and pushes out the pollen and then the stigma expands, and the flower in function is monoecious; from appearance I believe this is the case with your plant. I hope it is so, for this plant can hardly require a cross, being in function monoecious; so that dimorphism in such a case would be a heavy blow to understanding its nature or good in all other cases. I see few periodicals: when have you published on Clivia? I suppose that you did not actually count the seeds in the hybrids in comparison with those of the parent-forms; but this is almost necessary after Gartner's observations. I very much hope you will make a good series of comparative trials on the same plant of Tacsonia. (636/2. See Scott in "Linn. Soc. Journal," VIII.) I have raised 700-800 seedlings from cowslips, artificially fertilised with care; and they presented not a hair's-breadth approach to oxlips. I have now seed in pots of cowslip fertilised by pollen of primrose, and I hope they will grow; I have also got fine seedlings from seed of wild oxlips; so I hope to make out the case. You speak of difficulties on Natural Selection: there are indeed plenty; if ever you have spare time (which is not likely, as I am sure you must be a hard worker) I should be very glad to hear difficulties from one who has observed so much as you have. The majority of criticisms on the "Origin" are, in my opinion, not worth the paper they are printed on. Sir C. Lyell is coming out with what, I expect, will prove really good remarks. (636/3. Lyell's "Antiquity of Man" was published in the spring of 1863. In the "Life and Letters," Volume III., pages 8, 11, Darwin's correspondence shows his deep disappointment at what he thought Lyell's half-heartedness in regard to evolution. See Letter 164, Volume I.) Pray do not think me intrusive; but if you would like to have any book I have published, such as my "Journal of Researches" or the "Origin," I should esteem it a compliment to be allowed to send it. Will you permit me to suggest one experiment, which I should much like to see tried, and which I now wish the more from an extraordinary observation by Asa Gray on Gymnadenia tridentata (in number just out of Silliman's N. American Journal) (636/4. In Gymnadenia tridentata, according to Asa Gray, the anther opens in the bud, and the pollen being somewhat coherent falls on the stigma and on the rostellum which latter is penetrated by the pollen-tubes. "Fertilisation of Orchids," Edition II., page 68. Asa Gray's papers are in "American Journal of Science," Volume XXXIV., 1862, and XXXVI., 1863.); namely, to split the labellum of a Cattleya, or of some allied orchis, remove caudicle from pollen-mass (so that no loose grains are about) and put it carefully into the large tongue-like rostellum, and see if pollen-tubes will penetrate, or better, see if capsule will swell. Similar pollen-masses ought to be put on true stigmas of two or three other flowers of same plants for comparison. It is to discover whether rostellum yet retains some of its primordial function of being penetrated by pollen-tubes. You will be sorry that you ever entered into correspondence with me. But do not answer till at leisure, and as briefly as you like. My handwriting, I know, is dreadfully bad. Excuse this scribbling paper, as I can write faster on it, and I have a rather large correspondence to keep up.
LETTER 637. TO J. SCOTT. Down, January 21st, 1863.
I thank you for your very interesting letter; I must answer as briefly as I can, for I have a heap of other letters to answer. I strongly advise you to follow up and publish your observations on the pollen-tubes of orchids; they promise to be very interesting. If you could prove what I only conjectured (from state of utriculi in rostellum and in stigma of Catasetum and Acropera) that the utriculi somehow induce, or are correlated with, penetration of pollen-tubes you will make an important physiological discovery. I will mention, as worth your attention (and what I have anxiously wished to observe, if time had permitted, and still hope to do)—viz., the state of tissues or cells of stigma in an utterly sterile hybrid, in comparison with the same in fertile parent species; to test these cells, immerse stigmas for 48 hours in spirits of wine. I should expect in hybrids that the cells would not show coagulated contents. It would be an interesting discovery to show difference in female organs of hybrids and pure species. Anyhow, it is worth trial, and I recommend you to make it, and publish if you do. The pollen-tubes directing themselves to stigma is also very curious, though not quite so new, but well worth investigation when you get Cattleya, etc., in flower. I say not so new, for remember small flowers of Viola and Oxalis; or better, see Bibliography in "Natural History Review," No. VIII., page 419 (October, 1862) for quotation from M. Baillon on pollen-tubes finding way from anthers to stigma in Helianthemum. I should doubt gum getting solid from {i.e. because of} continued secretion. Why not sprinkle fresh plaster of Paris and make impenetrable crust? (637/1. The suggestion that the stigma should be covered with a crust of plaster of Paris, pierced by a hole to allow the pollen-tubes to enter, bears a resemblance to Miyoshi's experiments with germinating pollen and fungal spores. See "Pringsheim's Jahrbucher," 1895; "Flora," 1894.) You might modify experiment by making little hole in one lower corner, and see if tubes find it out. See in my future paper on Linum pollen and stigma recognising each other. If you will tell me that pollen smells the stigma I will try and believe you; but I will not believe the Frenchman (I forget who) who says that stigma of Vanilla actually attracts mechanically, by some unknown force, the solid pollen-masses to it! Read Asa Gray in 2nd Review of my Orchis book on pollen of Gymnadenia penetrating rostellum. I can, if you like, lend you these Reviews; but they must be returned. R. Brown, I remember, says pollen-tubes separate from grains before the lower ends of tubes reach ovules. I saw, and was interested by, abstract of your Drosera paper (637/2. A short note on the irritability of Drosera in the "Trans. Bot. Soc. Edin." Volume VII.); we have been at very much the same work.