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 pa.s.sage 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 h.o.m.ologies).
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 G.o.d, and Leonard is wonderfully improved ...Good G.o.d, 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 h.o.m.ologies, 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 h.o.m.ologies 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 amus.e.m.e.nt. 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 a.s.sure 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 a.n.a.logous 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 a.n.a.logous 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 pa.s.sage from his Preface to the English translation of H. Muller"s "Befruchtung der Blumen"; the pa.s.sage 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 Ca.s.sia: 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 a.n.a.logous 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 dest.i.tute 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.