More Letters of Charles Darwin — Volume 2

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 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 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 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 book that I state that I did not look at the ovarium of A. Loddigesii. Would you have the kindness to send me word which end of the ovarium is meant by apex (that nearest the flower?), for I must try and get this species from Kew and look at its ovarium. I shall be extremely curious to hear whether the fruit, which is now maturing, produces a large number of good and plump seed; perhaps you may have seen the ripe capsules of other Vandeae, and may be able to form some conjecture what it ought to produce. In the young, unfertilised ovaria of many Vandeae there seemed an infinitude of ovules. In desperation it occurs to me as just possible, as almost everything in nature goes by gradation, that a properly male flower might occasionally produce a few seeds, in the same manner as female plants sometimes produce a little pollen. All your remarks seem to me excellent and very interesting, and I again thank you for your kindness in writing to me. I am pleased to observe that my description of the structure of Acropera seems to agree pretty well with what you have observed. Does it not strike you as very difficult to understand how insects remove the pollinia and carry them to the stigmas? Your suggestion that the mouth of the stigmatic cavity may become charged with viscid matter and thus secure the pollinia, and that the pollen-tubes may then protrude, seems very ingenious and new to me; but it would be very anomalous in orchids, i.e. as far as I have seen. No doubt, however, though I tried my best, I shall be proved wrong in many points. Botany is a new subject to me. With respect to the protrusion of pollen-tubes, you might like to hear (if you do not already know the fact) that, as I saw this summer, in the little imperfect flowers of Viola and Oxalis, which never open, the pollen-tubes always come out of the pollen-grain, whilst still in the anthers, and direct themselves in a beautiful manner to the stigma seated at some little distance. I hope that you will continue your very interesting observations.