I see that Rhexia grows in Ma.s.sachusetts; 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 pa.s.sage 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 Exac.u.m 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 book; 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 book 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 novelty 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 novel 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 ill.u.s.trating the encouragement and friendly a.s.sistance 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 a.s.sume to be unis.e.xual, and so far as known represented by male individuals only. Theoretically you have no doubt a.s.signed 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 book, 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 a.s.surance 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-ma.s.ses 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-ma.s.s 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 dest.i.tute of ovules, their place being subst.i.tuted 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.