More Letters of Charles Darwin — Volume 2

(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.)