Чарльз Дарвин
More Letters of Charles Darwin — Volume 2
In making longitudinal sections of the fertilised ovary before mentioned, I found the basal portion entirely destitute of ovules, their place being substituted by transparent cellular ramification of the placentae. As I traced the placentae upwards, the ovules appeared, becoming gradually more abundant towards its apex. A transverse section near the apex of the ovary, however, still exhibited a more than ordinary placental development — i.e. {congenitally?} considered — each end giving off two branches, which meet each other in the centre of the ovary, the ovules being irregularly and sparingly disposed upon their surfaces.
In regard to the mere question of fertilisation, then, I am perfectly satisfied, but there are other points which require further elucidation. Among these I may particularly refer to the contracted stigmatic chamber, and the slight viscidity of its disk. The latter, however, may be a consequence of uncongenial conditions — as you do not mention particularly its examination by any author in its natural habitat. If such be the case, the contracted stigmatic chamber will offer no real difficulty, should the viscous exudations be only sufficient to render the mouth adhesive. For, as I have already shown, the pollen-tubes may be emitted in this condition, and effect fecundation without being in actual contact with the stigmatic surface, as occurs pretty regularly in the fertilisation of the Stapelias, for example. But, indeed, your own discovery of the independent germinative capabilities of the pollen-grains of certain Orchidaceae is sufficiently illustrative of this.
I may also refer to the peculiar abnormal condition that many at least of the ovaries present in a comparative examination of the placentae, and of which I beg to suggest the following explanation, though it is as yet founded on limited observations. In examining certain young ovaries of A. Loddigesii, I found some of them filled with the transparent membranous fringes of more or less distinctly cellular matter, which, from your description of the ovaries of luteola, appears to differ simply in the greater development in the former species. Again, in others I found small mammillary bodies, which appeared to be true ovules, though I could not perfectly satisfy myself as to the existence of the micropyle or nucleus. I unfortunately neglected to apply any chemical test. The fact, however, that in certain of the examined ovaries few or none of the latter bodies occurred — the placenta alone being developed in an irregular membranous form, taken in conjunction with the results of my experiments — before alluded to — on their fertilisation, leads me to infer that two sexual conditions are presented by the flowers of this plant. In short, that many of the ovaries are now normally abortive, though Nature occasionally makes futile efforts for their perfect development, in the production of ovuloid bodies; these then I regard as the male flowers. The others that are still capable of fertilisation, and likewise possessing male organs, are hermaphrodite, and must, I think, from the results of your comparative examinations, present a somewhat different condition; as it can scarcely be supposed that ovules in the condition you describe could ever be fertilised.
This is at least the most plausible explanation I can offer for the different results in my experiments on the fertilisation of apparently similar morphologically constructed flowers; others may, however, occur to you. Here there is not, as in the Catasetum, any external change visible in the respective unisexual and bisexual flowers. And yet it would appear from your researches that the ovules of Acropera are in a more highly atrophied condition than occurs in Catasetum, though, as you likewise remark, M. Neumann has never succeeded in fertilising C. tridentatum. If there be not, then, an arrangement of the reproductive structures, such as I have indicated, how can the different results in M. Neumann's experiments and mine be accounted for? However, as you have examined many flowers of both A. luteola and Loddigesii, such a difference in the ovulary or placental structures could scarcely have escaped your observation. But, be this as it may, the — to me at least — demonstrated fact still remains, that certain flowers of A. Loddigesii are capable of fertilisation, and that, though there are good grounds for supposing that important physiological changes are going on in the sexual phenomena of this species, there is no evidence whatever for supposing that external morphological changes have so masked certain individuals as to prevent their recognition.
I would now, sir, in conclusion beg you to excuse me for this infringement upon your valuable time, as I have been induced to write you in the belief that you have had negative results from other experimenters, before you ventured to propose your theoretical explanation, and consequently that you have been unknowingly led into error. I will continue, as opportunities present themselves, to examine the many peculiarities you have pointed out in this as well as others of the Orchid family; and at present I am looking forward with anxiety for the maturation of the ovary of A. Loddigesii, which will bear testimony to the veracity of the remarks I have ventured to lay before you.
LETTER 632. TO J.D. HOOKER. Down, 18th {November 1862}.
Strange to say, I have only one little bother for you to-day, and that is to let me know about what month flowers appear in Acropera Loddigesii and luteola; for I want extremely to beg a few more flowers, and if I knew the time I would keep a memorandum to remind you. Why I want these flowers is (and I am much alarmed) that Mr. J. Scott, of Bot. Garden of Edinburgh (do you know anything of him?) has written me a very long and clever letter, in which he confirms most of my observations; but tells me that with much difficulty he managed to get pollen into orifice, or as far as mouth of orifice, of six flowers of A. Loddigesii (the ovarium of which I did not examine), and two pods set; one he gathered, and saw a very few ovules, as he thinks, on the large and mostly rudimentary placenta. I shall be most curious to hear whether the other pod produces a good lot of seed. He says he regrets that he did not test the ovules with chemical agents: does he mean tincture of iodine? He suggests that in a state of nature the viscid matter may come to the very surface of stigmatic chamber, and so pollen-masses need not be inserted. This is possible, but I should think improbable. Altogether the case is very odd, and I am very uneasy, for I cannot hope that A. Loddigesii is hermaphrodite and A. luteola the male of the same species. Whenever I can get Acropera would be a very good time for me to look at Vanda in spirits, which you so kindly preserved for me.
LETTER 633. TO J. SCOTT.
(633/1. The following is Darwin's reply to the above letter from Scott. In the first edition of "Fertilisation of Orchids" (page 209) he assumed that the sexes in Acropera, as in Catasetum, were separate. In the second edition (page 172) he writes: "I was, however, soon convinced of my error by Mr. Scott, who succeeded in artificially fertilising the flowers with their own pollen. A remarkable discovery by Hildebrand (633/2. "Bot. Zeitung," 1863 and 1865.), namely, that in many orchids the ovules are not developed unless the stigma is penetrated by the pollen-tubes...explains the state of the ovarium in Acropera, as observed by me." In regard to this subject see Letter 608.)
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.
LETTER 638. TO J. SCOTT. Down, February 16th {1863}.
Absence from home has prevented me from answering you sooner. I should think that the capsule of Acropera had better be left till it shows some signs of opening, as our object is to judge whether the seeds are good; but I should prefer trusting to your better judgment. I am interested about the Gongora, which I hope hereafter to try myself, as I have just built a small hot-house.
Asa Gray's observations on the rostellum of Gymnadenia are very imperfect, yet worth looking at. Your case of Imatophyllum is most interesting (638/1. A sucker of Imatophyllum minatum threw up a shoot in which the leaves were "two-ranked instead of four-ranked," and showed other differences from the normal. — "Animals and Plants," Edition II., Volume I., page 411.); even if the sport does not flower it will be worth my giving. I did not understand, or I had forgotten, that a single frond on a fern will vary; I now see that the case does come under bud-variation, and must be given by me. I had thought of it only as proof {of} inheritance in cryptogams; I am much obliged for your correction, and will consult again your paper and Mr. Bridgeman's. (638/2. The facts are given in "Animals and Plants," Edition II., Volume I., page 408.) I enclose varieties of maize from Asa Gray. Pray do not thank me for trusting you; the thanks ought to go the other way. I felt a conviction after your first letter that you were a real lover of Natural History.
If you can advance good evidence showing that bisexual plants are more variable than unisexual, it will be interesting. I shall be very glad to read the discussion which you are preparing. I admit as fully as any one can do that cross-impregnation is the great check to endless variability; but I am not sure that I understand your view. I do not believe that the structure of Primula has any necessary relation to a tendency to a dioecious structure, but seeing the difference in the fertility of the two forms, I felt bound unwillingly to admit that they might be a step towards dioeciousness; I allude to this subject in my Linum paper. (638/3. "Linn. Soc. Journal," 1863.) Thanks for your answers to my other queries. I forgot to say that I was at Kew the other day, and I find that they can give me capsules of several Vandeae.
LETTER 639. TO J. SCOTT. Down, March 24th {1863}.
Your letter, as every one you have written, has greatly interested me. If you can show that certain individual Passifloras, under certain known or unknown conditions of life, have stigmas capable of fertilisation by pollen from another species, or from another individual of its own species, yet not by its own individual pollen (its own individual pollen being proved to be good by its action on some other species), you will add a case of great interest to me; and which in my opinion would be quite worth your publication. (639/1. Cases nearly similar to those observed by Scott were recorded by Gartner and Kolreuter, but in these instances only certain individuals were self-impotent. In "Animals and Plants," Edition II., Volume II., page 114, where the phenomenon is fully discussed, Scott's observations ("Trans. Bot. Soc. Edin." 1863) are given as the earliest, except for one case recorded by Lecoq ("Fecondation," 1862). Interesting work was afterwards done by Hildebrand and Fritz Muller, as illustrated in many of the letters addressed to the latter.) I always imagined that such recorded cases must be due to unnatural conditions of life; and think I said so in the "Origin." (639/2. See "Origin of Species," Edition I., page 251, for Herbert's observations on self-impotence in Hippeastrum. In spite of the uniformness of the results obtained in many successive years, Darwin inferred that the plants must have been in an "unnatural state.") I am not sure that I understand your result, {nor} whether it means what I have above obscurely expressed. If you can prove the above, do publish; but if you will not publish I earnestly beg you to let me have the facts in detail; but you ought to publish, for I may not use the facts for years. I have been much interested by what you say on the rostellum exciting pollen to protrude tubes; but are you sure that the rostellum does excite them? Would not tubes protrude if placed on parts of column or base of petals, etc., near to the stigma? Please look at the "Cottage Gardener" (or "Journal of Horticulture") (639/3. "Journal of Horticulture" and "Cottage Gardener," March 31st, 1863. A short note describing Cruger's discovery of self-fertilisation in Cattleya, Epidendrum, etc., and referring to the work of "an excellent observer, Mr. J. Scott." Darwin adds that he is convinced that he has underrated the power of tropical orchids occasionally to produce seeds without the aid of insects.) to be published to-morrow week for letter of mine, in which I venture to quote you, and in which you will see a curious fact about unopened orchid flowers setting seed in West Indies. Dr. Cruger attributes protrusion of tubes to ants carrying stigmatic secretion to pollen (639/4. In Cruger's paper ("Linn. Soc. Journ." VIII., 1865; read March 3rd 1864) he speaks of the pollen-masses in situ being acted on by the stigmatic secretion, but no mention is made of the agency of ants. He describes the pollen-tubes descending "from the {pollen} masses still in situ down into the ovarian canal."); but this is mere hypothesis. Remember, pollen-tubes protrude within anther in Neottia nidus-avis. I did think it possible or probable that perfect fertilisation might have been effected through rostellum. What a curious case your Gongora must be: could you spare me one of the largest capsules? I want to estimate the number of seed, and try my hand if I can make them grow. This, however, is a foolish attempt, for Dr. Hooker, who was here a day or two ago, says they cannot at Calcutta, and yet imported species have seeded and have naturally spread on to the adjoining trees! Dr. Cruger thinks I am wrong about Catasetum: but I cannot understand his letter. He admits there are three forms in two species; and he speaks as if the sexes were separate in some and that others were hermaphrodites (639/5. Cruger ("Linn. Soc. Journal," VIII., page 127) says that the apparently hermaphrodite form is always sterile in Trinidad. Darwin modified his account in the second edition of the orchid book.); but I cannot understand what he means. He has seen lots of great humble-bees buzzing about the flowers with the pollinia sticking to their backs! Happy man!! I have the promise, but not yet surety, of some curious results with my homomorphic seedling cowslips: these have not followed the rule of Chinese Primula; homomorphic seedlings from short-styled parent have presented both forms, which disgusts me.
