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- More Letters of Charles Darwin Volume II - 50/133 -

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

More Letters of Charles Darwin Volume II - 50/133

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