LETTER 605. TO J.D. HOOKER.
(605/2. The following letter is of interest because it relates to one of the two chief difficulties Darwin met with in working out the morphology of the orchid flower. In the orchid (605/3. Edition I., page 303.) he wrote, "This anomaly {in Habenaria} is so far of importance, as it throws some doubt on the view which I have taken of the labellum being always an organ compounded of one petal and two petaloid stamens." That is to say, it leaves it open for a critic to assert that the vessels which enter the sides of the labellum are lateral vessels of the petal and do not necessarily represent petaloid stamens. In the sequel he gives a satisfactory answer to the supposed objector.)
Down, November 10th, {1861}.
For the love of God help me. I believe all my work (about a fortnight) is useless. Look at this accursed diagram (Figure 9) of the butterfly-orchis {Habenaria}, which I examined after writing to you yesterday, when I thought all my work done. Some of the ducts of the upper sepal (605/4. These would be described by modern morphologists as lower, not upper, sepals, etc. Darwin was aware that he used these terms incorrectly.) and upper petal run to the wrong bundles on the column. I have seen no such case.
This case apparently shows that not the least reliance can be placed on the course of ducts. I am sure of my facts.
There is great adhesion and extreme displacement of parts where the organs spring from the top of the ovarium. Asa Gray says ducts are very early developed, and it seems to me wonderful that they should pursue this course. It may be said that the lateral ducts in the labellum running into the antero-lateral ovarian bundle is no argument that the labellum consists of three organs blended together.
In desperation (and from the curious way the base of upper petals are soldered at basal edges) I fancied the real form of upper sepal, upper petal and lower sepal might be as represented by red lines, and that there had been an incredible amount of splitting of sepals and petals and subsequent fusion.
This seems a monstrous notion, but I have just looked at Bauer's drawing of allied Bonatea, and there is a degree of lobing of petals and sepals which would account for anything.
Now could you spare me a dry flower out of your Herbarium of Bonatea speciosa (605/5. See "Fertilisation of Orchids," Edition I., page 304 (note), where the resemblances between the anomalous vessels of Bonatea and Habenaria are described. On November 14th, 1861, he wrote to Sir Joseph: "You are a true friend in need. I can hardly bear to let the Bonatea soak long enough."), that I might soak and look for ducts. If I cannot explain the case of Habenaria all my work is smashed. I was a fool ever to touch orchids.
LETTER 606. TO J.D. HOOKER. Down, November 17th {1861}.
What two very interesting and useful letters you have sent me. You rather astound me with respect to value of grounds of generalisation in the morphology of plants. It reminds me that years ago I sent you a grass to name, and your answer was, "It is certainly Festuca (so-and-so), but it agrees as badly with the description as most plants do." I have often laughed over this answer of a great botanist...Lindley, from whom I asked for an orchid with a simple labellum, has most kindly sent me a lot of what he marks "rare" and "rarissima" of peloric orchids, etc., but as they are dried I know not whether they will be of use. He has been most kind, and has suggested my writing to Lady D. Nevill, who has responded in a wonderfully kind manner, and has sent a lot of treasures. But I must stop; otherwise, by Jove, I shall be transformed into a botanist. I wish I had been one; this morphology is surprisingly interesting. Looking to your note, I may add that certainly the fifteen alternating bundles of spiral vessels (mingled with odd beadlike vessels in some cases) are present in many orchids. The inner whorl of anther ducts are oftenest aborted. I must keep clear of Apostasia, though I have cast many a longing look at it in Bauer. (606/1. Apostasia has two fertile anthers like Cypripedium. It is placed by Engler and Prantl in the Apostasieae or Apostasiinae, among the Orchideae, by others in a distinct but closely allied group.)
I hope I may be well enough to read my own paper on Thursday, but I have been very seedy lately. (606/2. "On the two Forms, or Dimorphic Condition, in the Species of the Genus Primula," "Linn. Soc. Journ." 1862. He did read the paper, but it cost him the next day in bed. "Life and Letters," III., page 299.) I see there is a paper at the Royal on the same night, which will more concern you, on fossil plants of Bovey (606/3. Oswald Heer, "The Fossil Flora of Bovey Tracey," "Phil. Trans. R. Soc." 1862, page 1039.), so that I suppose I shall not have you; but you must read my paper when published, as I shall very much like to hear what you think. It seems to me a large field for experiment. I shall make use of my Orchid little volume in illustrating modification of species doctrine, but I keep very, very doubtful whether I am not doing a foolish action in publishing. How I wish you would keep to your old intention and write a on plants. (606/4. Possibly a similar to that described in Letter 696.)
LETTER 607. TO G. BENTHAM. Down, November 26th {1861}.
Our notes have crossed on the r I know it is an honour to have a paper in the "Transactions," and I am much obliged to you for proposing it, but I should greatly prefer to publish in the "Journal." Nor does this apply exclusively to myself, for in old days at the Geological Society I always protested against an abstract appearing when the paper itself might appear. I abominate also the waste of time (and it would take me a day) in making an abstract. If the referee on my paper should recommend it to appear in the "Transactions," will you be so kind as to lay my earnest request before the Council that it may be permitted to appear in the "Journal?"
You must be very busy with your change of residence; but when you are settled and have some leisure, perhaps you will be so kind as to give me some cases of dimorphism, like that of Primula. Should you object to my adding them to those given me by A. Gray? By the way, I heard from A. Gray this morning, and he gives me two very curious cases in Boragineae.
LETTER 608. TO JOHN LINDLEY.
(608/1. In the following fragment occurs the earliest mention of Darwin's work on the three sexual forms of Catasetum tridentatum. Sir R. Schomburgk (608/2. "Trans. Linn. Soc." XVII., page 522.) described Catasetum tridentatum, Monacanthus viridis and Myanthus barbatus occurring on a single plant, but it remained for Darwin to make out that they are the male, female and hermaphrodite forms of a single species. (608/3. "Fertilisation of Orchids," Edition I., page 236; Edition II., page 196.)
With regard to the species of Acropera (Gongora) (608/4. Acropera Loddigesii = Gongora galeata: A. luteola = G. fusca ("Index Kewensis").) he was wrong in his surmise. The apparent sterility seems to be explicable by Hildebrand's discovery (608/5. "Bot. Zeitung," 1863 and 1865.) that in some orchids the ovules are not developed until pollinisation has occurred. (608/6. "Fertilisation of Orchids," Edition II., page 172. See Letter 633.))
Down, December 15th {1861}.
I am so nearly ready for press that I will not ask for anything more; unless, indeed, you stumbled on Mormodes in flower. As I am writing I will just mention that I am convinced from the rudimentary state of the ovules, and from the state of the stigma, that the whole plant of Acropera luteola (and I believe A. Loddigesii) is male. Have you ever seen any form from the same countries which could be the females? Of course no answer is expected unless you have ever observed anything to bear on this. I may add {judging from the} state of the ovules and of the pollen {that}:—
Catasetum tridentatum is male (and never seeds, according to Schomburgk, whom you have accidentally misquoted in the "Vegetable Kingdom"). Monacanthus viridis is female. Myanthus barbatus is the hermaphrodite form of same species.
LETTER 609. TO J.D. HOOKER. Down, December 18th {1861}.
Thanks for your note. I have not written for a long time, for I always fancy, busy as you are, that my letters must be a bore; though I like writing, and always enjoy your notes. I can sympathise with you about fear of scarlet fever: to the day of my death I shall never forget all the sickening fear about the other children, after our poor little baby died of it. The "Genera Plantarum" must be a tremendous work, and no doubt very valuable (such a odd as it may appear, would be very useful even to me), but I cannot help being rather sorry at the length of time it must take, because I cannot enter on and understand your work. Will you not be puzzled when you come to the orchids? It seems to me orchids alone would be work for a man's lifetime; I cannot somehow feel satisfied with Lindley's classification; the Malaxeae and Epidendreae seem to me very artificially separated. (609/1. Pfitzer (in the "Pflanzenfamilien") places Epidendrum in the Laeliinae-Cattleyeae, Malaxis in the Liparidinae. He states that Bentham united the Malaxideae and Epidendreae.) Not that I have seen enough to form an opinion worth anything.
Your African plant seems to be a vegetable Ornithorhynchus, and indeed much more than that. (609/2. See Sir J.D. Hooker, "On Welwitschia, a new genus of Gnetaceae." "Linn. Soc. Trans." XXIV., 1862-3.) The more I read about plants the more I get to feel that all phanerogams seem comparable with one class, as lepidoptera, rather than with one kingdom, as the whole insecta. (609/3. He wrote to Hooker (December 28th, 1861): "I wrote carelessly about the value of phanerogams; what I was thinking of was that the sub-groups seemed to blend so much more one into another than with most classes of animals. I suspect crustacea would show more difference in the extreme forms than phanerogams, but, as you say, it is wild speculation. Yet it is very strange what difficulty botanists seem to find in grouping the families together into masses.")
Thanks for your comforting sentence about the accursed ducts (accursed though they be, I should like nothing better than to work at them in the allied orders, if I had time). I shall be ready for press in three or four weeks, and have got all my woodcuts drawn. I fear much that publishing separately will prove a foolish job, but I do not care much, and the work has greatly amused me. The Catasetum has not flowered yet!
In writing to Lindley about an orchid which he sent me, I told him a little about Acropera, and in answer he suggests that Gongora may be its female. He seems dreadfully busy, and I feel that I have more right to kill you than to kill him; so can you send me one or at most two dried flowers of Gongora? if you know the habitat of Acropera luteola, a Gongora from the same country would be the best, but any true Gongora would do; if its pollen should prove as rudimentary as that of Monacanthus relatively to Catasetum, I think I could easily perceive it even in dried specimens when well soaked.
I have picked a little out of Lecoq, but it is awful tedious hunting.
Bates is getting on with his natural history travels in one volume. (609/4. H.W. Bates, the "Naturalist on the Amazons," 1863. See Volume I., Letters 123, 148, also "Life and Letters," Volume II., page 381.) I have read the first chapter in MS., and I think it will be an excellent and very well written; he argues, in a good and new way to me, that tropical climate has very little direct relation to the gorgeous colouring of insects (though of course he admits the tropics have a far greater number of beautiful insects) by taking all the few genera common to Britain and Amazonia, and he finds that the species proper to the latter are not at all more beautiful. I wonder how this is in species of the same restricted genera of plants.
If you can remember it, thank Bentham for getting my Primula paper printed so quickly. I do enjoy getting a subject off one's hands completely.
I have now got dimorphism in structure in eight natural orders just like Primula. Asa Gray sent me dried flowers of a capital case in Amsinkia spectabilis, one of the Boragineae. I suppose you do not chance to have the plant alive at Kew.
LETTER 610. TO A.G. MORE. Down, June 7th, 1862.
If you are well and have leisure, will you kindly give me one bit of information: Does Ophrys arachnites occur in the Isle of Wight? or do the intermediate forms, which are said to connect abrthis species and the bee-orchis, ever there occur?
Some facts have led me to suspect that it might just be possible, though improbable in the highest degree, that the bee {orchis} might be the self-fertilising form of O. arachnites, which requires insects' aid, something {in the same way} as we have self-fertilising flowers of the violet and others requiring insects. I know the case is widely different, as the bee is borne on a separate plant and is incomparably commoner. This would remove the great anomaly of the bee being a perpetual self-fertiliser. Certain Malpighiaceae for years produce only one of the two forms. What has set my head going on this is receiving to-day a bee having one alone of the best marked characters of O. arachnites. (610/1. Ophrys arachnites is probably more nearly allied to O. aranifera than to O. apifera. For a case somewhat analogous to that suggested see the description of O. scolopax in "Fertilisation of Orchids," Edition II., page 52.) Pray forgive me troubling you.
LETTER 611. TO G. BENTHAM. Down, June 22nd {1862?}.
Here is a piece of presumption! I must think that you are mistaken in ranking Hab{enaria} chlorantha (611/1. In Hooker's "Students' Flora," 1884, page 395, H. chlorantha is given as a subspecies of H. bifolia. Sir J.D. Hooker adds that they are "according to Darwin, distinct, and require different species of moths to fertilise them. They vary in the position and distances of their anther-cells, but intermediates occur." See "Fertilisation of Orchids," Edition II., page 73.) as a variety of H. bifolia; the pollen-masses and stigma differ more than in most of the best species of Orchis. When I first examined them I remember telling Hooker that moths would, I felt sure, fertilise them in a different manner; and I have just had proof of this in a moth sent me with the pollinia (which can be easily recognised) of H. chlorantha attached to its proboscis, instead of to the sides of its face, as an H. bifolia.
Forgive me scribbling this way; but when a man gets on his hobby-horse he always is run away with. Anyhow, nothing here requires any answer.
LETTER 612. TO J.D. HOOKER. Down, {September} 14th {1862}.
Your letter is a mine of wealth, but first I must scold you: I cannot abide to hear you abuse yourself, even in joke, and call yourself a stupid dog. You, in fact, thus abuse me, because for long years I have looked up to you as the man whose opinion I have valued more on any scientific subject than any one else in the world. I continually marvel at what you know, and at what you do. I have been looking at the "Genera" (612/1. "Genera Plantarum," by Bentham and Hooker, Volume I., Part I., 1862.), and of course cannot judge at all of its real value, but I can judge of the amount of condensed facts under each family and genus.
I am glad you know my feeling of not being able to judge about one's own work; but I suspect that you have been overworking. I should think you could not give too much time to Wellwitchia (I spell it different every time I write it) (612/2. "On Welwitschia," "Linn. Soc. Trans." {1862}, XXIV., 1863.); at least I am sure in the animal kingdom monographs cannot be too long on the osculant groups.
Hereafter I shall be excessively glad to read a paper about Aldrovanda (612/3. See "Insectivorous Plants," page 321.), and am very much obliged for reference. It is pretty to see how the caught flies support Drosera; nothing else can live.
Thanks about plants with two kinds of anthers. I presume (if an included flower was a Cassia) (612/4. Todd has described a species of Cassia with an arrangement of stamens like the Melastomads. See Chapter 2.X.II.) that Cassia is like lupines, but with some stamens still more rudimentary. If I hear I will return the three Melastomads; I do not want them, and, indeed, have cuttings. I am very low about them, and have wasted enormous labour over them, and cannot yet get a glimpse of the meaning of the parts. I wish I knew any botanical collector to whom I could apply for seeds in their native land of any Heterocentron or Monochoetum; I have raised plenty of seedlings from your plants, but I find in other cases that from a homomorphic union one generally gets solely the parent form. Do you chance to know of any botanical collector in Mexico or Peru? I must not now indulge myself with looking after vessels and homologies. Some future time I will indulge myself. By the way, some time I want to talk over the alternation of organs in flowers with you, for I think I must have quite misunderstood you that it was not explicable.
I found out the Verbascum case by pure accident, having transplanted one for experiment, and finding it to my astonishment utterly sterile. I formerly thought with you about rarity of natural hybrids, but I am beginning to change: viz., oxlips (not quite proven), Verbascum, Cistus (not quite proven), Aegilops triticoides (beautifully shown by Godron), Weddell's and your orchids (612/5. For Verbascum see "Animals and Plants," Edition II., Volume I., page 356; for Cistus, Ibid., Edition II., Volume I., page 356, Volume II., page 122; for Aegilops, Ibid., Edition II., Volume I., page 330, note.), and I daresay many others recorded. Your letters are one of my greatest pleasures in life, but I earnestly beg you never to write unless you feel somewhat inclined, for I know how hard you work, as I work only in the morning it is different with me, and is only a pleasant relaxation. You will never know how much I owe to you for your constant kindness and encouragement.
LETTER 613. TO JOHN LUBBOCK (Lord Avebury). Cliff Cottage, Bournemouth, Hants, September 2nd {1862}.
Hearty thanks for your note. I am so glad that your tour answered so splendidly. My poor patients (613/1. Mrs. Darwin and one of her sons, both recovering from scarlet fever.) got here yesterday, and are doing well, and we have a second house for the well ones. I write now in great haste to beg you to look (though I know how busy you are, but I cannot think of any other naturalist who would be careful) at any field of common red clover (if such a field is near you) and watch the hive-bees: probably (if not too late) you will see some sucking at the mouth of the little flowers and some few sucking at the base of the flowers, at holes bitten through the corollas. All that you will see is that the bees put their heads deep into the {flower} head and rout about. Now, if you see this, do for Heaven's sake catch me some of each and put in spirits and keep them separate. I am almost certain that they belong to two castes, with long and short proboscids. This is so curious a point that it seems worth making out. I cannot hear of a clover field near here.
LETTER 614. TO JOHN LUBBOCK (Lord Avebury). Cliff Cottage, Bournemouth, Wednesday, September 3rd {1862}.
I beg a million pardons. Abuse me to any degree, but forgive me: it is all an illusion (but almost excusable) about the bees. (614/1. H. Muller, "Fertilisation of Flowers," page 186, describes hive-bees visiting Trifolium pratense for the sake of the pollen. Darwin may perhaps have supposed that these were the variety of bees whose proboscis was long enough to reach the nectar. In "Cross and Self Fertilisation," page 361, Darwin describes hive-bees apparently searching for a secretion on the calyx. In the same passage in "Cross and Self Fertilisation" he quotes Muller as stating that hive-bees obtain nectar from red clover by breaking apart the petals. This seems to us a misinterpretation of the "Befruchtung der Blumen," page 224.) I do so hope that you have not wasted any time from my stupid blunder. I hate myself, I hate clover, and I hate bees.
(FIGURE 10.—DIAGRAM OF CRUCIFEROUS FLOWER. FIGURE 11.—DISSECTION OF CRUCIFEROUS FLOWER.
Laid flat open, showing by dotted lines the course of spiral vessels in all the organs; sepals and petals shown on one side alone, with the stamens on one side above with course of vessels indicated, but not prolonged. Near side of pistil with one spiral vessel cut away.)
LETTER 615. TO J.D. HOOKER. Cliff Cottage, Bournemouth, September 11th, 1862.
You once told me that Cruciferous flowers were anomalous in alternation of parts, and had given rise to some theory of dedoublement.
Having nothing on earth to do here, I have dissected all the spiral vessels in a flower, and instead of burning my diagrams {Figures 10 and 11}, I send them to you, you miserable man. But mind, I do not want you to send me a discussion, but just some time to say whether my notions are rubbish, and then burn the diagrams. It seems to me that all parts alternate beautifully by fours, on the hypothesis that two short stamens of outer whorl are aborted (615/1. The view given by Darwin is (according to Eichler) that previously held by Knuth, Wydler, Chatin, and others. Eichler himself believes that the flower is dimerous, the four longer stamens being produced by the doubling or splitting of the upper (i.e. antero-posterior) pair of stamens. If this view is correct, and there are good reasons for it, it throws much suspicion on the evidence afforded by the course of vessels, for there is no trace of the common origin of the longer stamens in the diagram (Figure 11). Again, if Eichler is right, the four vessels shown in the section of the ovary are misleading. Darwin afterwards gave a doubtful explanation of this, and concluded that the ovary is dimerous. See Letter 616.); and this view is perhaps supported by their being so few, only two sub-bundles in the two lateral main bundles, where I imagine two short stamens have aborted, but I suppose there is some valid objection against this notion. The course of the side vessels in the sepals is curious, just like my difficulty in Habenaria. (615/2. See Letter 605.) I am surprised at the four vessels in the ovarium. Can this indicate four confluent pistils? anyhow, they are in the right alternating position. The nectary within the base of the shorter stamens seems to cause the end sepals apparently, but not really, to arise beneath the lateral sepals.
I think you will understand my diagrams in five minutes, so forgive me for bothering you. My writing this to you reminds me of a letter which I received yesterday from Claparede, who helped the French translatress of the "Origin" (615/3. The late Mlle. Royer.), and he tells me he had difficulty in preventing her (who never looked at a bee's cell) from altering my whole description, because she affirmed that an hexagonal prism must have an hexagonal base! Almost everywhere in the "Origin," when I express great doubt, she appends a note explaining the difficulty, or saying that there is none whatever!! (615/4. See "Life and Letters," II., page 387.) It is really curious to know what conceited people there are in the world (people, for instance, after looking at one Cruciferous flower, explain their homologies).
This is a nice, but most barren country, and I can find nothing to look at. Even the brooks and ponds produce nothing. The country is like Patagonia. my wife is almost well, thank God, and Leonard is wonderfully improved ...Good God, what an illness scarlet fever is! The doctor feared rheumatic fever for my wife, but she does not know her risk. It is now all over.
(FIGURE 12.)
LETTER 616. TO J.D. HOOKER. Cliff Cottage, Bournemouth, Thursday Evening {September 18th, 1862}.
Thanks for your pleasant note, which told me much news, and upon the whole good, of yourselves. You will be awfully busy for a time, but I write now to say that if you think it really worth while to send me a few Dielytra, or other Fumariaceous plant (which I have already tried in vain to find here) in a little tin box, I will try and trace the vessels; but please observe, I do not know that I shall have time, for I have just become wonderfully interested in experimenting on Drosera with poisons, etc. If you send any Fumariaceous plant, send if you can, also two or three single balsams. After writing to you, I looked at vessels of ovary of a sweet-pea, and from this and other cases I believe that in the ovary the midrib vessel alone gives homologies, and that the vessels on the edge of the carpel leaf often run into the wrong bundle, just like those on the sides of the sepals. Hence I {suppose} in Crucifers that the ovarium consists of two pistils; AA {Figure 12} being the midrib vessels, and BB being those formed of the vessels on edges of the two carpels, run together, and going to wrong bundles. I came to this conclusion before receiving your letter.
I wonder why Asa Gray will not believe in the quaternary arrangement; I had fancied that you saw some great difficulty in the case, and that made me think that my notion must be wrong.
LETTER 617. TO J.D. HOOKER. Down, September 27th {1862}.
Masdevallia turns out nothing wonderful (617/1. This may refer to the homologies of the parts. He was unable to understand the mechanism of the flower.—"Fertilisation of Orchids," Edition II., page 136.); I was merely stupid about it; I am not the less obliged for its loan, for if I had lived till 100 years old I should have been uneasy about it. It shall be returned the first day I send to Bromley. I have steamed the other plants, and made the sensitive plant very sensitive, and shall soon try some experiments on it. But after all it will only be amusement. Nevertheless, if not causing too much trouble, I should be very glad of a few young plants of this and Hedysarum in summer (617/2. Hedysarum or Desmodium gyrans, the telegraph-plant.), for this kind of work takes no time and amuses me much. Have you seeds of Oxalis sensitiva, which I see mentioned in ? By the way, what a fault it is in Henslow's "Botany" that he gives hardly any references; he alludes to great series of experiments on absorption of poison by roots, but where to find them I cannot guess. Possibly the all-knowing Oliver may know. I can plainly see that the glands of Drosera, from rapid power (almost instantaneous) of absorption and power of movement, give enormous advantage for such experiments. And some day I will enjoy myself with a good set to work; but it will be a great advantage if I can get some preliminary notion on other sensitive plants and on roots.
Oliver said he would speak about some seeds of Lythrum hyssopifolium being preserved for me. By the way, I am rather disgusted to find I cannot publish this year on Lythrum salicaria; I must make 126 additional crosses. All that I expected is true, but I have plain indication of much higher complexity. There are three pistils of different structure and functional power, and I strongly suspect altogether five kinds of pollen all different in this one species! (617/3. See "Forms of Flowers," Edition II., page 138.)
By any chance have you at Kew any odd varieties of the common potato? I want to grow a few plants of every variety, to compare flowers, leaves, fruit, etc., as I have done with peas, etc. (617/4. "Animals and Plants," Edition II., Volume I., page 346. Compare also the similar facts with regard to cabbages, loc. cit., page 342. Some of the original specimens are in the Botanical Museum at Cambridge.)
LETTER 618. J.D. HOOKER TO CHARLES DARWIN.
(618/1. The following is part of Letter 144, Volume I. It refers to reviews of "Fertilisation of Orchids" in the "Gardeners' Chronicle," 1862, pages 789, 863, 910, and in the "Natural History Review," October, 1862, page 371.)
November 7th, 1862.
Dear old Darwin,
I assure you it was not my fault! I worried Lindley over and over again to notice your orchid in the "Chronicle" by the very brst hints man could give. (618/2. See "Life and Letters," III., page 273.) At last he said, "really I cannot, you must do it for me," and so I did—volontiers. Lindley felt that he ought to have done it himself, and my main effort was to write it "a la Lindley," and in this alone I have succeeded—that people all think it is exactly Lindley's style!!! which diverts me vastly. The fact is, between ourselves, I fear that poor L. is breaking up—he said that he could not fix his mind on your He works himself beyond his mental or physical powers.
And now, my dear Darwin, I may as well make a clean breast of it, and tell you that I wrote the "Nat. Hist. Review" notice too—to me a very difficult task, and one I fancied I failed in, comparatively. Of this you are no judge, and can be none; you told me to tell Oliver it pleased you, and so I am content and happy.
LETTER 619. TO W.E. DARWIN. Down, 4th {about 1862-3?}
I have been looking at the fertilisation of wheat, and I think possibly you might find something curious. I observed in almost every one of the pollen-grains, which had become empty and adhered to (I suppose the viscid) branching hairs of the stigma, that the pollen-tube was always (?) emitted on opposite side of grain to that in contact with the branch of the stigma. This seems very odd. The branches of the stigma are very thin, formed apparently of three rows of cells of hardly greater diameter than pollen-tube. I am astonished that the tubes should be able to penetrate the walls. The specimens examined (not carefully by me) had pollen only during few hours on stigma; and the mere SUSPICION has crossed me that the pollen-tubes crawl down these branches to the base and then penetrate the stigmatic tissue. (619/1. See Strasburger's "Neue Untersuchungen uber den Befruchtungsvorgang bei den Phanerogamen," 1884. In Alopecurus pratensis he describes the pollen as adhering to the end of a projection from the stigma where it germinates; the tube crawls along or spirally round this projection until it reaches the angle where the stigmatic branch is given off; here it makes an entrance and travels in the middle lamella between two cells.) The paleae open for a short period for stigma to be dusted, and then close again, and such travelling down would take place under protection. High powers and good adjustment are necessary. Ears expel anthers when kept in water in room; but the paleae apparently do not open and expose stigma; but the stigma could easily be artificially impregnated.
If I were you I would keep memoranda of points worth attending to.
2.X.II. MELASTOMACEAE, 1862-1881.
(620/1. The following series of letters (620-630) refers to the Melastomaceae and certain other flowers of analogous form. In 1862 Darwin attempted to explain the existence of two very different sets of stamens in these plants as a case of dimorphism, somewhat analogous to the state of things in Primula. In this view he was probably wrong, but this does not diminish the interest of the crossing experiments described in the letters. The persistence of his interest in this part of the subject is shown in the following passage from his Preface to the English translation of H. Muller's "Befruchtung der Blumen"; the passage is dated February, 1882, but was not published until the following year.
"There exist also some few plants the flowers of which include two sets of stamens, differing in the shape of the anthers and in the colour of the pollen; and at present no one knows whether this difference has any functional significance, and this is a point which ought to be determined."
It is not obvious why he spoke of the problem as if no light had been thrown on it, since in 1881 Fritz Muller had privately (see Letter 629) offered an explanation which Darwin was strongly inclined to accept. (620/2. H. Muller published ("Nature," August 4th, 1881) a letter from his brother Fritz giving the theory in question for Heeria. Todd ("American Naturalist," April 1882), described a similar state of things in Solanum rostratum and in Cassia: and H.O. Forbes ("Nature," August 1882, page 386) has done the same for Melastoma. In Rhexia virginica Mr. W.H. Leggett ("Bulletin Torrey Bot. Club, New York," VIII., 1881, page 102) describes the curious structure of the anther, which consists of two inflated portions and a tubular part connecting the two. By pressing with a blunt instrument on one of the ends, the pollen is forced out in a jet through a fine pore in the other inflated end. Mr. Leggett has seen bees treading on the anthers, but could not get near enough to see the pollen expelled. In the same journal, Volume IX., page 11, Mr. Bailey describes how in Heterocentron roseum, "upon pressing the bellows-like anther with a blunt pencil, the pollen was ejected to a full inch in distance." On Lagerstroemia as comparable with the Melastomads see Letter 689.) Fritz Muller's theory with regard to the Melastomads and a number of analogous cases in other genera are discussed in H. Muller's article in "Kosmos" (620/3. "Kosmos," XIII., 1883, page 241.), where the literature is given. F. Muller's theory is that in Heeria the yellow anthers serve merely as a means of attracting pollen-collecting bees, while the longer stamens with purple or crimson anthers supply pollen for fertilising purposes. If Muller is right the pollen from the yellow anthers would not normally reach the stigma. The increased vigour observed in the seedlings from the yellow anthers would seem to resemble the good effect of a cross between different individuals of the same species as worked out in "Cross and Self Fertilisation," for it is difficult to believe that the pollen of the purple anthers has become, by adaptation, less effective than that of the yellow anthers. In the letters here given there is some contradiction between the statements as to the position of the two sets of stamens in relation to the sepals. According to Eichler ("Bluthendiagramme, II., page 482) the longer stamens may be either epipetalous or episepalous in this family.
The work on the Melastomads is of such intrinsic importance that we have thought it right to give the correspondence in considerable detail; we have done so in spite of the fact that Darwin arrived at no definite conclusion, and in spite of an element of confusion and unsatisfactoriness in the series of letters. This applies also to Letter 629, written after Darwin had learned Fritz Muller's theory, which is obscured by some errors or slips of the pen.)
LETTER 620. TO G. BENTHAM. Down, February 3rd {1862?}
As you so kindly helped me before on dimorphism, will you forgive me begging for a little further information, if in your power to give it? The case is that of the Melastomads with eight stamens, on which I have been experimenting. I am perplexed by opposed statements: Lindley says the stamens which face the petals are sterile; Wallich says in Oxyspora paniculata that the stamens which face the sepals are destitute of pollen; I find plenty of apparently good pollen in both sets of stamens in Heterocentron {Heeria}, Monochoetum, and Centradenia. Can you throw any light on this? But there is another point on which I am more anxious for information. Please look at the enclosed miserable diagram. I find that the pollen of the yellow petal-facing stamens produce more than twice as much seed as the pollen of the purple sepal-facing stamens. This is exactly opposed to Lindley's statement—viz., that the petal-facing stamens are sterile. But I cannot at present believe that the case has any relation to abortion; it is hardly possible to believe that the longer and very curious stamens, which face the sepals in this Heterocentron, are tending to be rudimentary, though their pollen applied to their own flowers produces so much less seed. It is conformable with what we see in Primula that the {purple} sepal-facing anthers, which in the plant seen by me stood quite close on each side of the stigma, should have been rendered less fitted to fertilise the stigma than the stamens on the opposite side of the flower. Hence the suspicion has crossed me that if many plants of the Heterocentron roseum were examined, half would be found with the pistil nearly upright, instead of being rectangularly bent down, as shown in the diagram (620/4. According to Willis, "Flowering Plants and Ferns," 1897, Volume II., page 252, the style in Monochoetum, "at first bent downwards, moves slowly up till horizontal."); or, if the position of pistil is fixed, that in half the plants the petal-facing stamens would bend down, and in the other half of the plants the sepal-facing stamens would bend down as in the diagram. I suspect the former case, as in Centradenia I find the pistil nearly straight. Can you tell me? (620/5. No reply by Mr. Bentham to this or the following queries has been found.) Can the name Heterocentron have any reference to such diversity? Would it be asking too great a favour to ask you to look at dried specimens of Heterocentron roseum (which would be best), or of Monochoetum, or any eight-stamened Melastomad, of which you have specimens from several localities (as this would ensure specimens having been taken from distinct plants), and observe whether the pistil bends differently or stamens differently in different plants? You will at once see that, if such were the fact, it would be a new form of dimorphism, and would open up a large field of inquiry with respect to the potency of the pollen in all plants which have two sets of stamens—viz., longer and shorter. Can you forgive me for troubling you at such unreasonable length? But it is such waste of time to experiment without some guiding light. I do not know whether you have attended particularly to Melastoma; if you have not, perhaps Hooker or Oliver may have done so. I should be very grateful for any information, as it will guide future experiments.
P.S.—Do you happen to know, when there are only four stamens, whether it is the petal or sepal-facers which are preserved? and whether in the four-stamened forms the pistil is rectangularly bent or is straight?
LETTER 621. TO ASA GRAY. Down, February 16th {1862?}.
I have been trying a few experiments on Melastomads; and they seem to indicate that the pollen of the two curious sets of anthers (i.e. the petal-facers and the sepal-facers) have very different powers; and it does not seem that the difference is connected with any tendency to abortion in the one set. Now I think I can understand the structure of the flower and means of fertilisation, if there be two forms,—one with the pistil bent rectangularly out of the flower, and the other with it nearly straight.
Our hot-house and green-house plants have probably all descended by cuttings from a single plant of each species; so I can make out nothing from them. I applied in vain to Bentham and Hooker; but Oliver picked out some sentences from Naudin, which seem to indicate differences in the position of the pistil.
I see that Rhexia grows in Massachusetts; and I suppose has two different sets of stamens. Now, if in your power, would you observe the position of the pistil in different plants, in lately opened flowers of the same age? (I specify this because in Monochaetum I find great changes of position in the pistils and stamens, as flower gets old). Supposing that my prophecy should turn out right, please observe whether in both forms the passage into the flower is not {on} the upper side of the pistil, owing to the basal part of the pistil lying close to the ring of filaments on the under side of the flower. Also I should like to know the colour of the two sets of anthers. This would take you only a few minutes, and is the only way I see that I can find out whether these plants are dimorphic in this peculiar way—i.e., only in the position of the pistil (621/1. In Exacum and in Saintpaulia the flowers are dimorphic in this sense: the style projects to either the right or the left side of the corolla, from which it follows that a right-handed flower would fertilise a left-handed one, and vice versa. See Willis, "Flowering Plants and Ferns," 1897, Volume I., page 73.) and in its relation to the two kinds of pollen. I am anxious about this, because if it should prove so, it will show that all plants with longer and shorter or otherwise different anthers will have to be examined for dimorphism.
LETTER 622. TO ASA GRAY. March 15th {1862}.
...I wrote some little time ago about Rhexia; since then I have been carefully watching and experimenting on another genus, Monochaetum; and I find that the pistil is first bent rectangularly (as in the sketch sent), and then in a few days becomes straight: the stamens also move. If there be not two forms of Rhexia, will you compare the position of the part in young and old flowers? I have a suspicion (perhaps it will be proved wrong when the seed-capsules are ripe) that one set of anthers are adapted to the pistil in early state, and the other set for it in its later state. If bees visit the Rhexia, for Heaven's sake watch exactly how the anther and stigma strike them, both in old and young flowers, and give me a sketch.
Again I say, do not hate me.
LETTER 623. TO J.D. HOOKER. Leith Hill Place, Dorking, Thursday, 15th {May 1862}.
You stated at the Linnean Society that different sets of seedling Cinchona (623/1. Cinchona is apparently heterostyled: see "Forms of Flowers," Edition II., page 134.) grew at very different rate, and from my Primula case you attributed it probably to two sorts of pollen. I confess I thought you rash, but I now believe you were quite right. I find the yellow and crimson anthers of the same flower in the Melastomatous Heterocentron roseum have different powers; the yellow producing on the same plant thrice as many seeds as the crimson anthers. I got my neighbour's most skilful gardener to sow both kinds of seeds, and yesterday he came to me and said it is a most extraordinary thing that though both lots have been treated exactly alike, one lot all remain dwarfs and the other lot are all rising high up. The dwarfs were produced by the pollen of the crimson anthers. In Monochaetum ensiferum the facts are more complex and still more strange; as the age and position of the pistils comes into play, in relation to the two kinds of pollen. These facts seem to me so curious that I do not scruple to ask you to see whether you can lend me any Melastomad just before flowering, with a not very small flower, and which will endure for a short time a greenhouse or sitting-room; when fertilised and watered I could send it to Mr. Turnbull's to a cool stove to mature seed. I fully believe the case is worth investigation.
P.S.—You will not have time at present to read my orchid I never before felt half so doubtful about anything which I published. When you read it, do not fear "punishing" me if I deserve it.
Adios. I am come here to rest, which I much want.
Whenever you have occasion to write, pray tell me whether you have Rhododendron Boothii from Bhootan, with a smallish yellow flower, and pistil bent the wrong way; if so, I would ask Oliver to look for nectary, for it is an abominable error of Nature that must be corrected. I could hardly believe my eyes when I saw the pistil.
LETTER 624. TO ASA GRAY. January 19th {1863}.
I have been at those confounded Melastomads again; throwing good money (i.e. time) after bad. Do you remember telling me you could see no nectar in your Rhexia? well, I can find none in Monochaetum, and Bates tells me that the flowers are in the most marked manner neglected by bees and lepidoptera in Amazonia. Now the curious projections or horns to the stamens of Monochaetum are full of fluid, and the suspicion occurs to me that diptera or small hymenoptera may puncture these horns like they puncture (proved since my orchid was published) the dry nectaries of true Orchis. I forget whether Rhexia is common; but I very much wish you would next summer watch on a warm day a group of flowers, and see whether they are visited by small insects, and what they do.
LETTER 625. TO I.A. HENRY. Down, January 20th {1863}.
...You must kindly permit me to mention any point on which I want information. If you are so inclined, I am curious to know from systematic experiments whether Mr. D. Beaton's statement that the pollen of two shortest anthers of scarlet Pelargonium produce dwarf plants (625/1. See "Animals and Plants," Edition II., Volume II., page 150, for a brief account of Darwin's experiments on this genus. Also loc. cit., page 338 (note), for a suggested experiment.), in comparison with plants produced from the same mother-plant by the pollen of longer stamens from the same flower. It would aid me much in some laborious experiments on Melastomads. I confess I feel a little doubtful; at least, I feel pretty nearly sure that I know the meaning of short stamens in most plants. This summer (for another object) I crossed Queen of Scarlet Pelargonium with pollen of long and short stamens of multiflora alba, and it so turns out that plants from short stamens are the tallest; but I believe this to have been mere chance. My few crosses in Pelargonium were made to get seed from the central peloric or regular flower (I have got one from peloric flower by pollen of peloric), and this leads me to suggest that it would be very interesting to test fertility of peloric flowers in three ways,—own peloric pollen on peloric stigma, common pollen on peloric stigma, peloric pollen on common stigma of same species. My object is to discover whether with change of structure of flower there is any change in fertility of pollen or of female organs. This might also be tested by trying peloric and common pollen on stigma of a distinct species, and conversely. I believe there is a peloric and common variety of Tropaeolum, and a peloric or upright and common variation of some species of Gloxinia, and the medial peloric flowers of Pelargonium, and probably others unknown to me.
LETTER 626. TO I.A. HENRY. Hartfield, May 2nd {1863}.
In scarlet dwarf Pelargonium, you will find occasionally an additional and abnormal stamen on opposite and lower side of flower. Now the pollen of this one occasional short stamen, I think, very likely would produce dwarf plants. If you experiment on Pelargonium I would suggest your looking out for this single stamen.
I observed fluctuations in length of pistil in Phloxes, but thought it was mere variability.
If you could raise a bed of seedling Phloxes of any species except P. Drummondii, it would be highly desirable to see if two forms are presented, and I should be very grateful for information and flowers for inspection. I cannot remember, but I know that I had some reason to look after Phloxes. (626/1. See "Forms of Flowers," Edition II., page 119, where the conjecture is hazarded that Phlox subulata shows traces of a former heterostyled condition.)
I do not know whether you have used microscopes much yet. It adds immensely to interest of all such work as ours, and is indeed indispensable for much work. Experience, however, has fully convinced me that the use of the compound without the simple microscope is absolutely injurious to progress of N{atural} History (excepting, of course, with Infusoria). I have, as yet, found no exception to the rule, that when a man has told me he works with the compound alone his work is valueless.
LETTER 627. TO ASA GRAY. March 20th {1863}.
I wrote to him {Dr. H. Cruger, of Trinidad} to ask him to observe what the insects did in the flowers of Melastomaceae: he says not proper season yet, but that on one species a small bee seemed busy about the horn-like appendages to the anthers. It will be too good luck if my study of the flowers in the greenhouse has led me to right interpretation of these appendages.
LETTER 628. TO J.D. HOOKER. Down, November 28th {1871}.
If you had come here on Sunday I should have asked you whether you could give me seed or seedlings of any Melastomad which would flower soon to experiment on! I wrote also to J. Scott to ask if he could give me seed.
Several years ago I raised a lot of seedlings of a Melastomad greenhouse bush (Monochaetus or some such name) (628/1. Monochaetum.) from stigmas fertilised separately by the two kinds of pollen, and the seedlings differed remarkably in size, and whilst young, in appearance; and I never knew what to think of the case (so you must not use it), and have always wished to try again, but they are troublesome beasts to fertilise.
On the other hand I could detect no difference in the product from the two coloured anthers of Clarkia. (628/2. Clarkia has eight stamens divided into two groups which differ in the colour of the anthers.) If you want to know further particulars of my experiments on Monochaetum (?) and Clarkia, I will hunt for my notes. You ask about difference in pollen in the same species. All dimorphic and trimorphic plants present such difference in function and in size. Lythrum and the trimorphic Oxalis are the most wonderful cases. The pollen of the closed imperfect cleistogamic flowers differ in the transparency of the integument, and I think in size. The latter point I could ascertain from my notes. The pollen or female organs must differ in almost every individual in some manner; otherwise the pollen of varieties and even distinct individuals of same varieties would not be so prepotent over the individual plant's own pollen. Here follows a case of individual differences in function of pollen or ovules or both. Some few individuals of Reseda odorata and R. lutea cannot be fertilised, or only very rarely, by pollen of the same plant, but can by pollen of any other individual. I chanced to have two plants of R. odorata in this state; so I crossed them and raised five seedlings, all of which were self sterile and all perfectly fertile with pollen of any other individual mignonette. So I made a self sterile race! I do not know whether these are the kinds of facts which you require.
Think whether you can help me to seed or better seedlings (not cuttings) of any Melastomad.
LETTER 629. TO F. MULLER. Down, March 20th, 1881.
I have received the seeds and your most interesting letter of February 7th. The seeds shall be sown, and I shall like to see the plants sleeping; but I doubt whether I shall make any more detailed observations on this subject, as, now that I feel very old, I require the stimulus of some y to make me work. This stimulus you have amply given me in your remarkable view of the meaning of the two-coloured stamens in many flowers. I was so much struck with this fact with Lythrum, that I began experimenting on some Melastomaceae, which have two sets of extremely differently coloured anthers. After reading your letter I turned to my notes (made 20 years ago!) to see whether they would support or contradict your suggestion. I cannot tell yet, but I have come across one very remarkable result, that seedlings from the crimson anthers were not 11/20ths of the size of seedlings from the yellow anthers of the same flowers. Fewer good seeds were produced by the crimson pollen. I concluded that the shorter stamens were aborting, and that the pollen was not good. (629/1. "Shorter stamens" seems to be a slip of the pen for "longer,"—unless the observations were made on some genus in which the structure is unusual.) The mature pollen is incoherent, and must be {word illegible} against the visiting insect's body. I remembered this, and I find it said in my EARLY notes that bees would never visit the flowers for pollen. This made me afterwards write to the late Dr. Cruger in the West Indies, and he observed for me the flowers, and saw bees pressing the anthers with their mandibles from the base upwards, and this forced a worm-like thread of pollen from the terminal pore, and this pollen the bees collected with their hind legs. So that the Melastomads are not opposed to your views.
I am now working on the habits of worms, and it tires me much to change my subject; so I will lay on one side your letter and my notes, until I have a week's leisure, and will then see whether my facts bear on your views. I will then send a letter to "Nature" or to the Linn. Soc., with the extract of your letter (and this ought to appear in any case), with my own observations, if they appear worth publishing. The subject had gone out of my mind, but I now remember thinking that the imperfect action of the crimson stamens might throw light on hybridism. If this pollen is developed, according to your view, for the sake of attracting insects, it might act imperfectly, as well as if the stamens were becoming rudimentary. (629/2. As far as it is possible to understand the earlier letters it seems that the pollen of the shorter stamens, which are adapted for attracting insects, is the most effective.) I do not know whether I have made myself intelligible.
LETTER 630. TO W. THISELTON-DYER. Down, March 21st {1881}.
I have had a letter from Fritz Muller suggesting a and very curious explanation of certain plants producing two sets of anthers of different colour. This has set me on fire to renew the laborious experiments which I made on this subject, now 20 years ago. Now, will you be so kind as to turn in your much worked and much holding head, whether you can think of any plants, especially annuals, producing 2 such sets of anthers. I believe that this is the case with Clarkia elegans, and I have just written to Thompson for seeds. The Lythraceae must be excluded, as these are heterostyled.
I have got seeds from Dr. King of some Melastomaceae, and will write to Veitch to see if I can get the Melastomaceous genera Monochaetum and Heterocentron or some such name, on which I before experimented. Now, if you can aid me, I know that you will; but if you cannot, do not write and trouble yourself.
2.X.III. CORRESPONDENCE WITH JOHN SCOTT, 1862-1871.
"If he had leisure he would make a wonderful observer, to my judgment; I have come across no one like him."—Letter to J.D. Hooker, May 29th {1863}.
(631/1. The following group of letters to John Scott, of whom some account is given elsewhere (Volume I., Letters 150 and 151, and Index.) deal chiefly with experimental work in the fertilisation of flowers. In addition to their scientific importance, several of the letters are of special interest as illustrating the encouragement and friendly assistance which Darwin gave to his correspondent.)
LETTER 631. JOHN SCOTT TO CHARLES DARWIN. Edinburgh Botanic Gardens, November 11th, 1862.
I take the liberty of addressing you for the purpose of directing your attention to an error in one of your ingenious explanations of the structural adaptations of the Orchidaceae in your late work. This occurs in the genus Acropera, two species of which you assume to be unisexual, and so far as known represented by male individuals only. Theoretically you have no doubt assigned good grounds for this view; nevertheless, experimental observations that I am now making have already convinced me of its fallacy. And I thus hurriedly, and as you may think prematurely, direct your attention to it, before I have seen the final result of my own experiment, that you might have the longer time for reconsidering the structure of this genus for another edition of your interesting if indeed it be not already called for. I am furthermore induced to communicate the results of my yet imperfect experiments in the belief that the actuating principle of your late work is the elicitation of truth, and that you will gladly avail yourself of this even at the sacrifice of much ingenious theoretical argumentation.
Since I have had an opportunity of perusing your work on orchid fertilisation, my attention has been particularly directed to the curiously constructed floral organs of Acropera. I unfortunately have as yet had only a few flowers for experimental enquiry, otherwise my remarks might have been clearer and more satisfactory. Such as they are, however, I respectfully lay {them} before you, with a full assurance of their veracity, and I sincerely trust that as such you will receive them.
Your observations seem to have been chiefly directed to the A. luteola, mine to the A. Loddigesii, which, however, as you remark, is in a very similar constructural condition with the former; having the same narrow stigmatic chamber, abnormally developed placenta, etc. In regard to the former point—contraction of stigmatic chamber—I may remark that it does not appear to be absolutely necessary that the pollen-masses penetrate this chamber for effecting fecundation. Thus a raceme was produced upon a plant of A. Loddigesii in the Botanic Gardens here lately; upon this I left only six flowers. These I attempted to fertilise, but with two only of the six have I been successful: I succeeded in forcing a single pollen-mass into the stigmatic chamber of one of the latter, but I failed to do this on the other; however, by inserting a portion of a pedicel with a pollinium attached, I caused the latter to adhere, with a gentle press, to the mouth of the stigmatic chamber. Both of these, as I have already remarked, are nevertheless fertilised; one of them I have cut off for examination, and its condition I will presently describe; the other is still upon the plant, and promises fair to attain maturity. In regard to the other four flowers, I may remark that though similarly fertilised—part having pollinia inserted, others merely attached—they all withered and dropped off without the least swelling of the ovary. Can it be, then, that this is really an {andro-monoecious} species?—part of the flowers male, others truly hermaphrodite.
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 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 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 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.
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