LETTER 590. TO J.D. HOOKER.

(590/1. The genera Scaevola and Leschenaultia, to which the following letter refers, belong to the Goodeniaceae (Goodenovieae, Bentham & Hooker), an order allied to the Lobeliaceae, although the mechanism of fertilisation resembles rather more nearly that of Campanula. The characteristic feature of the flower in this order is the indusium, or, as Delpino (590/2. Delpino's observations on Dichogamy, summarised by Hildebrand in "Bot. Zeitung," 1870, page 634.) calls it, the "collecting cup": this cuplike organ is a development of the style, and serves the same function as the hairs on the style of Campanula, namely, that of taking the pollen from the anthers and presenting it to the visiting insect. During this stage the immature stigma is at the bottom of the cup, and though surrounded by pollen is incapable of being pollinated. In most genera of the order the pollen is pushed out of the indusium by the growth of the style or stigma, very much as occurs in Lobelia or the Compositae. Finally the style emerges from the indusium (590/3. According to Hamilton ("Proc. Linn. Soc. N. S. Wales," X., 1895, page 361) the stigma rarely grows beyond the indusium in Dampiera. In the same journal (1885-6, page 157, and IX., 1894, page 201) Hamilton has given a number of interesting observations on Goodenia, Scaevola, Selliera, Brunonia. There seem to be mechanisms for cross- and also for self-fertilisation.), the stigmas open out and are pollinated from younger flowers. The mechanism of fertilisation has been described by F. Muller (590/4. In a letter to Hildebrand published in the "Bot. Zeitung," 1868, page 113.), and more completely by Delpino (loc. cit.).

Mr. Bentham wrote a paper (590/5. "Linn. Soc. Journal," 1869, page 203.) on the style and stigma in the Goodenovieae, where he speaks of Mr. Darwin's belief that fertilisation takes place outside the indusium. This statement, which we imagine Mr. Bentham must have had from an unpublished source, was incomprehensible to him as long as he confined his work to such genera as Goodenia, Scaevola, Velleia, Coelogyne, in which the mechanism is much as above described; but on examining Leschenaultia the meaning became clear. Bentham writes of this genus:—"The indusium is usually described as broadly two-lipped, without any distinct stigma. The fact appears to be that the upper less prominent lip is stigmatic all over, inside and out, with a transverse band of short glandular hairs at its base outside, while the lower more prominent lip is smooth and glabrous, or with a tuft of rigid hairs. Perhaps this lower lip and the upper band of hairs are all that correspond to the indusium of other genera; and the so-called upper lip, outside of which impregnation may well take place, as observed by Mr. Darwin, must be regarded as the true stigma."

Darwin's interest in the Goodeniaceae was due to the mechanism being apparently fitted for self-fertilisation. In 1871 a writer signing himself F.W.B. made a communication to the "Gardeners' Chronicle" (590/6. 1871, page 1103.), in which he expresses himself as "agreeably surprised" to find Leschenaultia adapted for self-fertilisation, or at least for self-pollinisation. This led Darwin to publish a short note in the same journal, in which he describes the penetration of pollen-tubes into the viscid surface on the outside of the indusium. (590/7. 1871, page 1166. He had previously written in the "Journal of Horticulture and Cottage Gardener," May 28th, 1861, page 151:—"Leschenaultia formosa has apparently the most effective contrivance to prevent the stigma of one flower ever receiving a grain of pollen from another flower; for the pollen is shed in the early bud, and is there shut up round the stigma within a cup or indusium. But some observations led me to suspect that nevertheless insect agency here comes into play; for I found by holding a camel-hair pencil parallel to the pistil, and moving it as if it were a bee going to suck the nectar, the straggling hairs of the brush opened the lip of the indusium, entered it, stirred up the pollen, and brought out some grains. I did this to five flowers, and marked them. These five flowers all set pods; whereas only two other pods set on the whole plant, though covered with innumerable flowers...I wrote to Mr. James Drummond, at Swan River in Australia,...and he soon wrote to me that he had seen a bee cleverly opening the indusium and extracting pollen.") He also describes how a brush, pushed into the flower in imitation of an insect, presses "against the slightly projecting lower lip of the indusium, opens it, and some of the hairs enter and become smeared with pollen." The yield of pollen is therefore differently arranged in Leschenaultia; for in the more typical genera it depends on the growth of the style inside the indusium. Delpino, however (see Hildebrand's version, loc. cit.), describes a similar opening of the cup produced by pressure on the hairs in some genera of the order.)

Down, June 7th [1860].

Best and most beloved of men, I supplicate and entreat you to observe one point for me. Remember that the Goodeniaceae have weighed like an incubus for years on my soul. It relates to Scaevola microcarpa. I find that in bud the indusium collects all the pollen splendidly, but, differently from Leschenaultia, cannot be afterwards easily opened. Further, I find that at an early stage, when the flower first opens, a boat-shaped stigma lies at the bottom of the indusium, and further that this stigma, after the flower has some time expanded, grows very rapidly, when the plant is kept hot, and pushes out of the indusium a mass of pollen; and at same time two horns project at the corners of the indusium. Now the appearance of these horns makes me suppose that these are the stigmatic surfaces. Will you look to this? for if they be by the relative position of the parts (with indusium and stigma bent at right angles to style) [I am led to think] that an insect entering a flower could not fail to have [its] whole back (at the period when, as I have seen, a whole mass of pollen is pushed out) covered with pollen, which would almost certainly get rubbed on the two horns. Indeed, I doubt whether, without this aid, pollen would get on to the horns. What interests me in the case is the analogy in result with the Lobelia, but by very different means. In Lobelia the stigma, before it is mature, pushes by its circular brush of hairs the pollen out of the conjoined anthers; here the indusium collects pollen, and then the growth of the stigma pushes it out. In the course of about 1 1/2 hour, I found an indusium with hairs on the outer edge perfectly clogged with pollen, and horns protruded, which before the 1 1/2 hour had not one grain of pollen outside the indusium, and no trace of protruding horns. So you will see how I wish to know whether the horns are the true stigmatic surfaces. I would try the case experimentally by putting pollen on the horns, but my greenhouse is so cold, and my plant so small, and in such a little pot, that I suppose it would not seed...

The little length of stigmatic horns at the moment when pollen is forced out of the indusium, compared to what they ultimately attain, makes me fancy that they are not then mature or ready, and if so, as in Lobelia, each flower must be fertilised by pollen from another and earlier flower.

How curious that the indusium should first so cleverly collect pollen and then afterwards push it out! Yet how closely analogous to Campanula brushing pollen out of the anther and retaining it on hairs till the stigma is ready. I am going to try whether Campanula sets seed without insect agency.

LETTER 591. TO J.D. HOOKER.

(591/1. The following letters are given here rather than in chronological order, as bearing on the Leschenaultia problem. The latter part of Letter 591 refers to the cleistogamic flowers of Viola.)

Down, May 1st [1862].

If you can screw out time, do look at the stigma of the blue Leschenaultia biloba. I have just examined a large bud with the indusium not yet closed, and it seems to me certain that there is no stigma within. The case would be very important for me, and I do not like to trust solely to myself. I have been impregnating flowers, but it is rather difficult...

I have just looked again at Viola canina. The case is odder: only 2 stamens which embrace the stigma have pollen; the 3 other stamens have no anther-cells and no pollen. These 2 fertile anthers are of different shape from the 3 sterile others, and the scale representing the lower lip is larger and differently shaped from the 4 other scales representing 4 other petals.

In V. odorata (single flower) all five stamens produce pollen. But I daresay all this is known.

LETTER 592. TO J.D. HOOKER. November 3rd [1862].

Do you remember the scarlet Leschenaultia formosa with the sticky margin outside the indusium? Well, this is the stigma—at least, I find the pollen-tubes here penetrate and nowhere else. What a joke it would be if the stigma is always exterior, and this by far the greatest difficulty in my crossing notions should turn out a case eminently requiring insect aid, and consequently almost inevitably ensuring crossing. By the way, have you any other Goodeniaceae which you could lend me, besides Leschenaultia and Scaevola, of which I have seen enough?

I had a long letter the other day from Crocker of Chichester; he has the real spirit of an experimentalist, but has not done much this summer.

LETTER 593. TO F. MULLER. Down, April 9th and 15th [1866].

I am very much obliged by your letter of February 13th, abounding with so many highly interesting facts. Your account of the Rubiaceous plant is one of the most extraordinary that I have ever read, and I am glad you are going to publish it. I have long wished some one to observe the fertilisation of Scaevola, and you must permit me to tell you what I have observed. First, for the allied genus of Leschenaultia: utterly disbelieving that it fertilises itself, I introduced a camel-hair brush into the flower in the same way as a bee would enter, and I found that the flowers were thus fertilised, which never otherwise happens; I then searched for the stigma, and found it outside the indusium with the pollen-tubes penetrating it; and I convinced Dr. Hooker that botanists were quite wrong in supposing that the stigma lay inside the indusium. In Scaevola microcarpa the structure is very different, for the immature stigma lies at the base within the indusium, and as the stigma grows it pushes the pollen out of the indusium, and it then clings to the hairs which fringe the tips of the indusium; and when an insect enters the flower, the pollen (as I have seen) is swept from these long hairs on to the insect's back. The stigma continues to grow, but is not apparently ready for impregnation until it is developed into two long protruding horns, at which period all the pollen has been pushed out of the indusium. But my observations are here at fault, for I did not observe the penetration of the pollen-tubes. The case is almost parallel with that of Lobelia. Now, I hope you will get two plants of Scaevola, and protect one from insects, leaving the other uncovered, and observe the results, both in the number of capsules produced, and in the average number of seeds in each. It would be well to fertilise half a dozen flowers under the net, to prove that the cover is not injurious to fertility.

With respect to your case of Aristolochia, I think further observation would convince you that it is not fertilised only by larvae, for in a nearly parallel case of an Arum and a Aristolochia, I found that insects flew from flower to flower. I would suggest to you to observe any cases of flowers which catch insects by their probosces, as occurs with some of the Apocyneae (593/1. Probably Asclepiadeae. See H. Muller, "Fertilisation of Flowers," page 396.); I have never been able to conceive for what purpose (if any) this is effected; at the same time, if I tempt you to neglect your zoological work for these miscellaneous observations I shall be guilty of a great crime.

To return for a moment to the indusium: how curious it is that the pollen should be thus collected in a special receptacle, afterwards to be swept out by insects' agency!

I am surprised at what you tell me about the fewness of the flowers of your native orchids which produce seed-capsules. What a contrast with our temperate European species, with the exception of some species of Ophrys!—I now know of three or four cases of self-fertilising orchids, but all these are provided with means for an occasional cross.

I am sorry to say Dr. Cruger is dead from a fever.

I received yesterday your paper in the "Botanische Zeitung" on the wood of climbing plants. (593/2. Fritz Muller, "Ueber das Holz einiger um Desterro wachsenden Kletterpflanzen." "Botanische Zeitung," 1866, pages 57, 65.) I have read as yet only your very interesting and curious remarks on the subject as bearing on the change of species; you have pleased me by the very high compliments which you pay to my paper. I have been at work since March 1st on a new English edition (593/3. The 4th Edition.) of my "Origin," of which when published I will send you a copy. I have much regretted the time it has cost me, as it has stopped my other work. On the other hand, it will be useful for a new third German edition, which is now wanted. I have corrected it largely, and added some discussions, but not nearly so much as I wished to do, for, being able to work only two hours daily, I feared I should never get it finished. I have taken some facts and views from your work "Fur Darwin"; but not one quarter of what I should like to have quoted.

LETTER 594. TO A.G. MORE. Down, June 24th, 1860.

I hope that you will forgive the liberty which I take in writing to you and requesting a favour. Mr. H.C. Watson has given me your address, and has told me that he thought that you would be willing to oblige me. Will you please to read the enclosed, and then you will understand what I wish observed with respect to the bee-orchis. (594/1. Ophrys apifera.) What I especially wish, from information which I have received since publishing the enclosed, is that the state of the pollen-masses should be noted in flowers just beginning to wither, in a district where the bee-orchis is extremely common. I have been assured that in parts of Isle of Wight, viz., Freshwater Gate, numbers occur almost crowded together: whether anything of this kind occurs in your vicinity I know not; but, if in your power, I should be infinitely obliged for any information. As I am writing, I will venture to mention another wish which I have: namely, to examine fresh flowers and buds of the Aceras, Spiranthes, marsh Epipactis, and any other rare orchis. The point which I wish to examine is really very curious, but it would take too long space to explain. Could you oblige me by taking the great trouble to send me in an old tin canister any of these orchids, permitting me, of course, to repay postage? It would be a great kindness, but perhaps I am unreasonable to make such a request. If you will inform me whether you have leisure so far to oblige me, I would tell you my movements, for on account of my own health and that of my daughter, I shall be on the move for the next two or three weeks.

I am sure I have much cause to apologise for the liberty which I have taken...

LETTER 595. TO A.G. MORE. Down, August 3rd, 1860.

I thank you most sincerely for sending me the Epipactis [palustris]. You can hardly imagine what an interesting morning's work you have given me, as the rostellum exhibited a quite new modification of structure. It has been extremely kind of you to take so very much trouble for me. Have you looked at the pollen-masses of the bee-Ophrys? I do not know whether the Epipactis grows near to your house: if it does, and any object takes you to the place (pray do not for a moment think me so very unreasonable as to ask you to go on purpose), would you be so kind [as] to watch the flowers for a quarter of an hour, and mark whether any insects (and what?) visit these flowers.

I should suppose they would crawl in by depressing the terminal portion of the labellum; and that when within the flower this terminal portion would resume its former position; and lastly, that the insect in crawling out would not depress the labellum, but would crawl out at back of flower. (595/1. The observations of Mr. William Darwin on Epipactis palustris given in the "Fertilisation of Orchids," Edition II., 1877, page 99, bear on this point. The chief fertilisers are hive-bees, which are too big to crawl into the flower. They cling to the labellum, and by depressing it open up the entrance to the flower. Owing to the elasticity of the labellum and its consequent tendency to spring up when released, the bees, "as they left the flower, seemed to fly rather upwards." This agrees with Darwin's conception of the mechanism of the flower as given in the first edition of the Orchid book, 1862, page 100, although at that time he imagined that the fertilising insect crawled into the flower. The extreme flexibility and elasticity of the labellum was first observed by Mr. More (see first edition, page 99). The description of the flower given in the above letter to Mr. More is not quite clear; the reader is referred to the "Fertilisation of Orchids," loc. cit.) An insect crawling out of a recently opened flower would, I believe, have parts of the pollen-masses adhering to the back or shoulder. I have seen this in Listera. How I should like to watch the Epipactis.

If you can it any time send me Spiranthes or Aceras or O. ustulata, you would complete your work of kindness.

P.S.—If you should visit the Epipactis again, would you gather a few of the lower flowers which have been opened for some time and have begun to wither a little, and observe whether pollen is well cleared out of anther-case. I have been struck with surprise that in nearly all the lower flowers sent by you, though much of the pollen has been removed, yet a good deal of pollen is left wasted within the anthers. I observed something of this kind in Cephalanthera grandiflora. But I fear that you will think me an intolerable bore.

LETTER 596. TO A.G. MORE. Down, August 5th, 1860.

I am infinitely obliged for your most clearly stated observations on the bee-orchis. It is now perfectly clear that something removes the pollen-masses far more with you than in this neighbourhood. But I am utterly puzzled about the foot-stalk being so often cut through. I should suspect snails. I yesterday found thirty-nine flowers, and of them only one pollen-mass in three flowers had been removed, and as these were extremely much-withered flowers I am not quite sure of the truth of this. The wind again is a new element of doubt. Your observations will aid me extremely in coming to some conclusion. (596/1. Mr. More's observations on the percentage of flowers in which the pollinia were absent are quoted in "Fertilisation of Orchids," Edition I., page 68.) I hope in a day or two to receive some day-moths, on the probosces of which I am assured the pollen-masses of the bee-orchis still adhere (596/2. He was doomed to disappointment. On July 17th, 1861, he wrote to Mr. More:—"I found the other day a lot of bee-Ophrys with the glands of the pollinia all in their pouches. All facts point clearly to eternal self-fertilisation in this species; yet I cannot swallow the bitter pill. Have you looked at any this year?")...

I wrote yesterday to thank you for the Epipactis. For the chance of your liking to look at what I have found: take a recently opened flower, drag gently up the stigmatic surface almost any object (the side of a hooked needle), and you will find the cap of the hemispherical rostellum comes off with a touch, and being viscid on under-surface, clings to needle, and as pollen-masses are already attached to the back of rostellum, the needle drags out much pollen. But to do this, the curiously projecting and fleshy summits of anther-cases must at some time be pushed back slightly. Now when an insect's head gets into the flower, when the flap of the labellum has closed by its elasticity, the insect would naturally creep out by the back-side of the flower. And mark when the insect flies to another flower with the pollen-masses adhering to it, if the flap of labellum did not easily open and allow free ingress to the insect, it would surely rub off the pollen on the upper petals, and so not leave it on stigma. It is to know whether I have rightly interpreted the structure of this whole flower that I am so curious to see how insects act. Small insects, I daresay, would crawl in and out and do nothing. I hope that I shall not have wearied you with these details.

If you would like to see a pretty and curious little sight, look to Orchis pyramidalis, and you will see that the sticky glands are congenitally united into a saddle-shaped organ. Remove this under microscope by pincers applied to foot-stalk of pollen-mass, and look quickly at the spontaneous movement of the saddle-shaped organs and see how beautifully adapted to seize proboscis of moth.

LETTER 597. TO J.D. HOOKER December 4th [1860].

Many thanks about Apocynum and Meyen.

The latter I want about some strange movements in cells of Drosera, which Meyen alone seems to have observed. (597/1. No observations of Meyen are mentioned in "Insectivorous Plants.") It is very curious, but Trecul disbelieves that Drosera really clasps flies! I should very much wish to talk over Drosera with you. I did chloroform it, and the leaves which were already expanded did not recover thirty seconds of exposure for three days. I used the expression weight for the bit of hair which caused movement and weighed 1/78000 of a grain; but I do not believe it is weight, and what it is, I cannot after many experiments conjecture. (597/2. The doubt here expressed as to whether the result is due to actual weight is interesting in connection with Pfeffer's remarkable discovery that a smooth object in contact with the gland produces no effect if the plant is protected from all vibration; on an ordinary table the slight shaking which reaches the plant is sufficient to make the body resting on the gland tremble, and thus produce a series of varying pressures—under these circumstances the gland is irritated, and the tentacle moves. See Pfeffer, "Untersuchungen aus d. bot. Institut zu Tubingen," Volume I., 1885, page 483; also "Insectivorous Plants," Edition II., page 22.) The movement in this case does not depend on the chemical nature of substance. Latterly I have tried experiments on single glands, and a microscopical atom of raw meat causes such rapid movement that I could see it move like hand of clock. In this case it is the nature of the object. It is wonderful the rapidity of the absorption: in ten seconds weak solution of carbonate of ammonia changes not the colour, but the state of contents within the glands. In two minutes thirty seconds juice of meat has been absorbed by gland and passed from cell to cell all down the pedicel (or hair) of the gland, and caused the sap to pass from the cells on the upper side of the pedicel to the lower side, and this causes the curvature of the pedicel. I shall work away next summer when Drosera opens again, for I am much interested in subject. After the glandular hairs have curved, the oddest changes take place—viz., a segregation of the homogeneous pink fluid and necessary slow movements in the thicker matter. By Jove, I sometimes think Drosera is a disguised animal! You know that I always so like telling you what I do, that you must forgive me scribbling on my beloved Drosera. Farewell. I am so very glad that you are going to reform your ways; I am sure that you would have injured your health seriously. There is poor Dana has done actually nothing—cannot even write a letter—for a year, and it is hoped that in another YEAR he may quite recover.

After this homily, good night, my dear friend. Good heavens, I ought not to scold you, but thank you, for writing so long and interesting a letter.

LETTER 598. TO E. CRESY. Down, December 12th [1860?].

After writing out the greater part of my paper on Drosera, I thought of so many points to try, and I wished to re-test the basis of one large set of experiments, namely, to feel still more sure than I am, that a drop of plain water never produces any effect, that I have resolved to publish nothing this year. For I found in the record of my daily experiments one suspicious case. I must wait till next summer. It will be difficult to try any solid substances containing nitrogen, such as ivory; for two quite distinct causes excite the movement, namely, mechanical irritation and presence of nitrogen. When a solid substance is placed on leaf it becomes clasped, but is released sooner than when a nitrogenous solid is clasped; yet it is difficult (except with raw meat and flies) to be sure of the result, owing to differences in vigour of different plants. The last experiments which I tried before my plants became too languid are very curious, and were tried by putting microscopical atoms on the gland itself of single hairs; and it is perfectly evident that an atom of human hair, 1/76000 of a grain (as ascertained by weighing a length of hair) in weight, causes conspicuous movement. I do not believe (for atoms of cotton thread acted) it is the chemical nature; and some reasons make me doubt whether it is actual weight; it is not the shadow; and I am at present, after many experiments, confounded to know what the cause is. That these atoms did really act and alter the state of the contents of all the cells in the glandular hair, which moved, was perfectly clear. But I hope next summer to make out a good deal more...

LETTER 599. TO J.D. HOOKER. Down, May 14th [1861].

I have been putting off writing from day to day, as I did not wish to trouble you, till my wish for a little news will not let me rest...

I have no news to tell you, for I have had no interesting letters for some time, and have not seen a soul. I have been going through the "Cottage Gardener" of last year, on account chiefly of Beaton's articles (599/1. Beaton was a regular contributor to the "Cottage Gardener," and wrote various articles on cross breeding, etc., in 1861. One of these was in reply to a letter published in the "Cottage Gardener," May 14th, 1861, page 112, in which Darwin asked for information as to the Compositae and the hollyhock being crossed by insect visitors. In the number for June 8th, 1861, page 211, Darwin wrote on the variability of the central flower of the carrot and the peloria of the central flower in Pelargonium. An extract from a letter by Darwin on Leschenaultia, "Cottage Gardener," May 28th, 1861, page 151, is given in Letter 590, note.); he strikes me as a clever but d—d cock-sure man (as Lord Melbourne said), and I have some doubts whether to be much trusted. I suspect he has never recorded his experiment at the time with care. He has made me indignant by the way he speaks of Gartner, evidently knowing nothing of his work. I mean to try and pump him in the "Cottage Gardener," and shall perhaps defend Gartner. He alludes to me occasionally, and I cannot tell with what spirit. He speaks of "this Mr. Darwin" in one place as if I were a very noxious animal.

Let me have a line about poor Henslow pretty soon.

(599/2. In a letter of May 18th, 1861, Darwin wrote again:—)

By the way, thanks about Beaton. I have now read more of his writings, and one answer to me in "Cottage Gardener." I can plainly see that he is not to be trusted. He does not well know his own subject of crossing.

LETTER 600. TO J.D. HOOKER.

(600/1. Part of this letter has been published in "Life and Letters," III., page 265.)

2, Hesketh Crescent, Torquay [1861].

...The beauty of the adaptation of parts seems to me unparalleled. I should think or guess [that] waxy pollen was most differentiated. In Cypripedium, which seems least modified, and a much exterminated group, the grains are single. In all others, as far as I have seen, they are in packets of four; and these packets cohere into many wedge-formed masses in Orchis, into eight, four, and finally two. It seems curious that a flower should exist which could, at most, fertilise only two other flowers, seeing how abundant pollen generally is; this fact I look at as explaining the perfection of the contrivance by which the pollen, so important from its fewness, is carried from flower to flower. By the way, Cephalanthera has single pollen-grains, but this seems to be a case of degradation, for the rostellum is utterly aborted. Oddly, the columns of pollen are here kept in place by very early penetration of pollen-tubes into the edge of the stigma; nevertheless, it receives more pollen by insect agency. Epithecia [Dichaea] has done me one good little turn. I often speculated how the caudicle of Orchis had been formed. (600/2. The gradation here suggested is thoroughly worked out in the "Fertilisation of Orchids," Edition I., page 323, Edition II., page 257.) I had noticed slight clouds in the substance half way down; I have now dissected them out, and I find they are pollen-grains fairly embedded and useless. If you suppose the pollen-grains to abort in the lower half of the pollinia of Epipactis, but the parallel elastic threads to remain and cohere, you have the caudicle of Orchis, and can understand the few embedded and functionless pollen-grains. I must not look at any more exotic orchids: hearty thanks for your offer. But if you would make one single observation for me on Cypripedium, I should be glad. Asa Gray writes to me that the outside of the pollen-masses is sticky in this genus; I find that the whole mass consists of pollen-grains immersed in a sticky brownish thick fluid. You could tell by a mere lens and penknife. If it is, as I find it, pollen could not get on the stigma without insect aid. Cypripedium confounds me much. I conjecture that drops of nectar are secreted by the surface of the labellum beneath the anthers and in front of the stigma, and that the shield over the anthers and the form of labellum is to compel insects to insert their proboscis all round both organs. (600/3. This view was afterwards given up.) It would be troublesome for you to look at this, as it is always bothersome to catch the nectar secreting, and the cup of the labellum gets filled with water by gardener's watering.

I have examined Listera ovata, cordata, and Neottia nidus avis: the pollen is uniform; I suspect you must have seen some observation founded on a mistake from the penetration and hardening of sticky fluid from the rostellum, which does penetrate the pollen a little.

It is mere virtue which makes me not wish to examine more orchids; for I like it far better than writing about varieties of cocks and hens and ducks. Nevertheless, I have just been looking at Lindley's list in the "Vegetable Kingdom," and I cannot resist one or two of his great division of Arethuseae, which includes Vanilla. And as I know so well the Ophreae, I should like (God forgive me) any one of the Satyriadae, Disidae and Corycidae.

I fear my long lucubrations will have wearied you, but it has amused me to write, so forgive me.

LETTER 601. TO J.D. HOOKER.

(601/1. Part of the following letter is published in the "Life and Letters," the remainder, with the omission of part bearing on the Glen Roy problem, is now given as an example of the varied botanical assistance Darwin received from Sir Joseph Hooker. For the part relating to Verbascum see the "Variation of Animals and Plants," Edition II., 1875, Volume II., page 83. The point is that the white and yellow flowered plants which occur in two species of Verbascum are undoubted varieties, yet "the sterility which results from the crossing of the differently coloured varieties of the same species is fully as great as that which occurs in many cases when distinct species are crossed."

The sterility of the long-styled form (B) of Linum grandiflorum, with its own pollen is described in "Forms of Flowers," Edition II., page 87: his conclusions on the short-styled form (A) differ from those in the present letter.)

September 28th [1861].

I am going to beg for help, and I will explain why I want it.

You offer Cypripedium; I should be very glad of a specimen, and of any good-sized Vandeae, or indeed any orchids, for this reason: I never thought of publishing separately, and therefore did not keep specimens in spirits, and now I should be very glad of a few woodcuts to illustrate my few remarks on exotic orchids. If you can send me any, send them by post in a tin canister on middle of day of Saturday, October 5th, for Sowerby will be here.

Secondly: Have you any white and yellow varieties of Verbascum which you could give me, or propagate for me, or LEND me for a year? I have resolved to try Gartner's wonderful and repeated statement, that pollen of white and yellow varieties, whether used on the varieties or on DISTINCT species, has different potency. I do not think any experiment can be more important on the origin of species; for if he is correct we certainly have what Huxley calls new physiological species arising. I should require several species of Verbascum besides the white and yellow varieties of the same species. It will be tiresome work, but if I can anyhow get the plants, it shall be tried.

Thirdly: Can you give me seeds of any Rubiaceae of the sub-order Cinchoneae, as Spermacoce, Diodia, Mitchella, Oldenlandia? Asa Gray says they present two forms like Primula. I am sure that this subject is well worth working out. I have just almost proved a very curious case in Linum grandiflorum which presents two forms, A and B. Pollen of A is perfectly fertile on stigma of A. But pollen of B is absolutely barren on its own stigma; you might as well put so much flour on it. It astounded me to see the stigmas of B purple with its own pollen; and then put a few grains of similar-looking pollen of A on them, and the germen immediately and always swelled; those not thus treated never swelling.

Fourthly: Can you give me any very hairy Saxifraga (for their functions) [i.e. the functions of the hairs]?

I send you a resume of my requests, to save you trouble. Nor would I ask for so much aid if I did not think all these points well worth trying to investigate.

My dear old friend, a letter from you always does me a world of good. And, the Lord have mercy on me, what a return I make.

LETTER 602. TO J.D. HOOKER. Down, October 4th [1861].

Will you have the kindness to read the enclosed, and look at the diagram. Six words will answer my question. It is not an important point, but there is to me an irresistible charm in trying to make out homologies. (602/1. In 1880 he wrote to Mr. Bentham: "It was very kind of you to write to me about the Orchideae, for it has pleased me to an extreme degree that I could have been of the least use to you about the nature of the parts."—"Life and Letters," III., page 264.) You know the membranous cup or clinandrum, in many orchids, behind the stigma and rostellum: it is formed of a membrane which unites the filament of the normal dorsal anther with the edges of the pistil. The clinandrum is largely developed in Malaxis, and is of considerable importance in retaining the pollinia, which as soon as the flower opens are quite loose.

The appearance and similarity of the tissues, etc., at once gives suspicion that the lateral membranes of the clinandrum are the two other and rudimentary anthers, which in Orchis and Cephalanthera, etc., exist as mere papillae, here developed and utilised.

Now for my question. Exactly in the middle of the filament of the normal anther, and exactly in the middle of the lateral membrane of the clinandrum, and running up to the same height, are quite similar bundles of spiral vessels; ending upwards almost suddenly. Now is not this structure a good argument that I interpret the homologies of the sides of clinandrum rightly? (602/2. Though Robert Brown made use of the spiral vessels of orchids, yet according to Eichler, "Bluthendiagramme," 1875, Volume I., page 184, Darwin was the first to make substantial additions to the conclusions deducible from the course of the vessels in relation to the problem of the morphology of these plants. Eichler gives Darwin's diagram side by side with that of Van Tieghem without attempting to decide between the differences in detail by which they are characterised.)

I find that the great Bauer does not draw very correctly! (602/3. F. Bauer, whom Pritzel calls "der grosste Pflanzenmaler." The reference is to his "Illustrations of Orchidaceous Plants, with Notes and Prefatory Remarks by John Lindley," London, 1830-38, Folio. See "Fertilisation of Orchids," Edition II., page 82.) And, good Heavens, what a jumble he makes on functions.

LETTER 603. TO J.D. HOOKER. Down, October 22nd. [1861].

Acropera is a beast,—stigma does not open, everything seems contrived that it shall NOT be anyhow fertilised. There is something very odd about it, which could only be made out by incessant watching on several individual plants.

I never saw the very curious flower of Canna; I should say the pollen was deposited where it is to prevent inevitable self-fertilisation. You have no time to try the smallest experiment, else it would be worth while to put pollen on some stigmas (supposing that it does not seed freely with you). Anyhow, insects would probably carry pollen from flower to flower, for Kurr states the tube formed by pistil, stamen and "nectarblatt" secretes (I presume internally) much nectar. Thanks for sending me the curious flower.

Now I want much some wisdom; though I must write at considerable length, your answer may be very brief.

(FIGURE 8.—FLORAL DIAGRAM OF AN ORCHID. The "missing bundle" could not be found in some species.)

In R. Brown's admirable paper in the "Linnean Transacts." (603/4. Volume XVI., page 685.) he suggests (and Lindley cautiously agrees) that the flower of orchids consists of five whorls, the inner whorl of the two whorls of anthers being all rudimentary, and when the labellum presents ridges, two or three of the anthers of both whorls [are] combined with it. In the ovarium there are six bundles of vessels: R. Brown judged by transverse sections. It occurred to me, after what you said, to trace the vessels longitudinally, and I have succeeded well. Look at my diagram [Figure 8] (which please return, for I am transported with admiration at it), which shows the vessels which I have traced, one bundle to each of fifteen theoretical organs, and no more. You will see the result is nothing new, but it seems to confirm strongly R. Brown, for I have succeeded (perhaps he did, but he does not say so) in tracing the vessels belonging to each organ in front of each other to the same bundle in the ovarium: thus the vessels going to the lower sepal, to the side of the labellum, and to one stigma (when there are two) all distinctly branch from one ovarian bundle. So in other cases, but I have not completely traced (only seen) that going to the rostellum. But here comes my only point of novelty: in all orchids as yet looked at (even one with so simple a labellum as Gymnadenia and Malaxis) the vessels on the two sides of the labellum are derived from the bundle which goes to the lower sepal, as in the diagram. This leads me to conclude that the labellum is always a compound organ. Now I want to know whether it is conceivable that the vessels coming from one main bundle should penetrate an organ (the labellum) which receives its vessels from another main bundle? Does it not imply that all that part of the labellum which is supplied by vessels coming from a lateral bundle must be part of a primordially distinct organ, however closely the two may have become united? It is curious in Gymnadenia to trace the middle anterior bundle in the ovarium: when it comes to the orifice of the nectary it turns and runs right down it, then comes up the opposite side and runs to the apex of the labellum, whence each side of the nectary is supplied by vessels from the bundles, coming from the lower sepals. Hence even the thin nectary is essentially, I infer, tripartite; hence its tendency to bifurcation at its top. This view of the labellum always consisting of three organs (I believe four when thick, as in Mormodes, at base) seems to me to explain its great size and tripartite form, compared with the other petals. Certainly, if I may trust the vessels, the simple labellum of Gymnadenia consists of three organs soldered together. Forgive me for writing at such length; a very brief answer will suffice. I am desperately interested in the subject: the destiny of the whole human race is as nothing to the course of vessels of orchids...

What plant has the most complex single stigma and pistil? The most complex I, in my ignorance, can think of is in Iris. I want to know whether anything beats in modification the rostellum of Catasetum. To-morrow I mean to be at Catasetum. Hurrah! What species is it? It is wonderfully different from that which Veitch sent me, which was C. saccatum.

According to the vessels, an orchid flower consists of three sepals and two petals free; and of a compound organ (its labellum), consisting of one petal and of two (or three) modified anthers; and of a second compound body consisting of three pistils, one normal anther, and two modified anthers often forming the sides of the clinandrum.

LETTER 604. TO JOHN LINDLEY.

(604/1. It was in the autumn of 1861 that Darwin made up his mind to publish his Orchid work as a book, rather than as a paper in the Linnean Society's "Journal." (604/2. See "Life and Letters," III., page 266.) The following letter shows that the new arrangement served as an incitement to fresh work.)

Down, October 25th [1861?]

Mr. James Veitch has been most generous. I did not know that you had spoken to him. If you see him pray say I am truly grateful; I dare not write to a live Bishop or a Lady, but if I knew the address of "Rucker"? and might use your name as introduction, I might write. I am half mad on the subject. Hooker has sent me many exotics, but I stopped him, for I thought I should make a fool of myself; but since I have determined to publish I much regret it.

(FIGURE 9.—HABENARIA CHLORANTHA (Longitudinal course of bundles).)

(605/1. The three upper curved outlines, two of which passing through the words "upper sepal," "upper petal," "lower sepal," were in red in the original; for explanation see text.)

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 book (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 book on plants. (606/4. Possibly a book similar to that described in Letter 696.)

LETTER 607. TO G. BENTHAM. Down, November 26th [1861].

Our notes have crossed on the road. 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 book, 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 book 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 abroad this 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 books? 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 book in the "Chronicle" by the very broadest 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 book. 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 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.