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.

Charles Darwin

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