More Letters of Charles Darwin — Volume 2

(706A/1. We have found it convenient to place the two letters to Riley together, rather than separate them chronologically.)

Down, September 28th, 1881.

I must write half a dozen lines to say how much interested I have been by your "Further Notes" on Pronuba which you were so kind as to send me. (706A/2. "Proc. Amer. Assoc. Adv. Sci." 1880.) I had read the various criticisms, and though I did not know what answer could be made, yet I felt full confidence in your result, and now I see that I was right...If you make any further observation on Pronuba it would, I think, be well worth while for you to observe whether the moth can or does occasionally bring pollen from one plant to the stigma of a distinct one (706A/3. Riley discovered the remarkable fact that the Yucca moth (Pronuba yuccasella) lays its eggs in the ovary of Yucca flowers, which it has previously pollinated, thus making sure of a supply of ovules for the larvae.), for I have shown that the cross-fertilisation of the flowers on the same plant does very little good; and, if I am not mistaken, you believe that Pronuba gathers pollen from the same flower which she fertilises.

What interesting and beautiful observations you have made on the metamorphoses of the grasshopper-destroying insects.

LETTER 707. TO F. HILDEBRAND. Down, February 9th {1872}.

Owing to other occupations I was able to read only yesterday your paper on the dispersal of the seeds of Compositae. (707/1. "Ueber die Verbreitungsmittel der Compositenfruchte." "Bot. Zeitung," 1872, page 1.) Some of the facts which you mention are extremely interesting.

I write now to suggest as worthy of your examination the curious adhesive filaments of mucus emitted by the achenia of many Compositae, of which no doubt you are aware. My attention was first called to the subject by the achenia of an Australian Pumilio (P. argyrolepis), which I briefly described in the "Gardeners' Chronicle," 1861, page 5. As the threads of mucus dry and contract they draw the seeds up into a vertical position on the ground. It subsequently occurred to me that if these seeds were to fall on the wet hairs of any quadruped they would adhere firmly, and might be carried to any distance. I was informed that Decaisne has written a paper on these adhesive threads. What is the meaning of the mucus so copiously emitted from the moistened seeds of Iberis, and of at least some species of Linum? Does the mucus serve as a protection against their being devoured, or as a means of attachment. (707/2. Various theories have been suggested, e.g., that the slime by anchoring the seed to the soil facilitates the entrance of the radicle into the soil: the slime has also been supposed to act as a temporary water-store. See Klebs in Pfeffer's "Untersuchungen aus dem Bot. Inst. zu Tubingen," I., page 581.) I have been prevented reading your paper sooner by attempting to read Dr. Askenasy's pamphlet, but the German is too difficult for me to make it all out. (707/3. E. Askenasy, "Beitrage zur Kritik der Darwin'schen Lehre." Leipzig, 1872.) He seems to follow Nageli completely. I cannot but think that both much underrate the utility of various parts of plants; and that they greatly underrate the unknown laws of correlated growth, which leads to all sorts of modifications, when some one structure or the whole plant is modified for some particular object.

LETTER 708. TO T.H. FARRER. (Lord Farrer).

(708/1. The following letter refers to a series of excellent observations on the fertilisation of Leguminosae, made by Lord Farrer in the autumn of 1869, in ignorance of Delpino's work on the subject. The result was published in "Nature," October 10th and 17th, 1872, and is full of interesting suggestions. The discovery of the mechanism in Coronilla mentioned in a note was one of the cases in which Lord Farrer was forestalled.)

Down {1872}.

I declare I am almost as sorry as if I had been myself forestalled — indeed, more so, for I am used to it. It is, however, a paramount, though bothersome duty in every naturalist to try and make out all that has been done by others on the subject. By all means publish next summer your confirmation and a summary of Delpino's observations, with any new ones of your own. Especially attend about the nectary exterior to the staminal tube. (708/2. This refers to a species of Coronilla in which Lord Farrer made the remarkable discovery that the nectar is secreted on the outside of the calyx. See "Nature," July 2nd, 1874, page 169; also Letter 715.) This will in every way be far better than writing to Delpino. It would not be at all presumptuous in you to criticise Delpino. I am glad you think him so clever; for so it struck me.

Look at hind legs yourself of some humble and hive-bees; in former take a very big individual (if any can be found) for these are the females, the males being smaller, and they have no pollen-collecting apparatus. I do not remember where it is figured — probably in Kirby & Spence — but actual inspection better...

Please do not return any of my books until all are finished, and do not hurry.

I feel certain you will make fine discoveries.

LETTER 709. TO T.H. FARRER. (Lord Farrer). Sevenoaks, October 13th, 1872.

I must send you a line to say how extremely good your article appears to me to be. It is even better than I thought, and I remember thinking it very good. I am particularly glad of the excellent summary of evidence about the common pea, as it will do for me hereafter to quote; nocturnal insects will not do. I suspect that the aboriginal parent had bluish flowers. I have seen several times bees visiting common and sweet peas, and yet varieties, purposely grown close together, hardly ever intercross. This is a point which for years has half driven me mad, and I have discussed it in my "Var. of Animals and Plants under Dom." (709/1. In the second edition (1875) of the "Variation of Animals and Plants," Volume I., page 348, Darwin added, with respect to the rarity of spontaneous crosses in Pisum: "I have reason to believe that this is due to their stignas being prematurely fertilised in this country by pollen from the same flower." This explanation is, we think, almost certainly applicable to Lathyrus odoratus, though in Darwin's latest publication on the subject he gives reasons to the contrary. See "Cross and Self-Fertilisation," page 156, where the problem is left unsolved. Compare Letter 714 to Delpino. In "Life and Letters," III., page 261, the absence of cross-fertilisation is explained as due to want of perfect adaptation between the pea and our native insects. This is Hermann Muller's view: see his "Fertilisation of Flowers," page 214. See Letter 583, note.) I now suspect (and I wish I had strength to experimentise next spring) that from changed climate both species are prematurely fertilised, and therefore hardly ever cross. When artificially crossed by removal of own pollen in bud, the offspring are very vigorous.

Farewell. — I wish I could compel you to go on working at fertilisation instead of so insignificant a subject as the commerce of the country!

You pay me a very pretty compliment at the beginning of your paper.

LETTER 710. TO J.D. HOOKER.

(710/1. The following letters to Sir J.D. Hooker and the late Mr. Moggridge refer to Moggridge's observation that seeds stored in the nest of the ant Atta at Mentone do not germinate, though they are certainly not dead. Moggridge's observations are given in his book, "Harvesting Ants and Trap-Door Spiders," 1873, which is full of interesting details. The book is moreover remarkable in having resuscitated our knowledge of the existence of the seed-storing habit. Mr. Moggridge points out that the ancients were familiar with the facts, and quotes the well-known fable of the ant and the grasshopper, which La Fontaine borrowed from Aesop. Mr. Moggridge (page 5) goes on: "So long as Europe was taught Natural History by southern writers the belief prevailed; but no sooner did the tide begin to turn, and the current of information to flood from north to south, than the story became discredited."

In Moggridge's "supplement" on the same subject, published in 1874, the author gives an account of his experiments made at Darwin's suggestion, and concludes (page 174) that "the vapour of formic acid is incapable of rendering the seeds dormant after the manner of the ants," and that indeed "its influence is always injurious to the seeds, even when present only in excessively minute quantities." Though unable to explain the method employed, he was convinced "that the non-germination of the seeds is due to some direct influence voluntarily exercised by the ants, and not merely to the conditions found in the nest" (page 172). See Volume I., Letter 251.)

Down, February 21st {1873}.

You have given me exactly the information which I wanted.

Geniuses jump. I have just procured formic acid to try whether its vapour or minute drops will delay germination of fresh seeds; trying others at same time for comparison. But I shall not be able to try them till middle of April, as my despotic wife insists on taking a house in London for a month from the middle of March.

I am glad to hear of the Primer (710/2. "Botany" (Macmillan's Science Primers).); it is not at all, I think, a folly. Do you know Asa Gray's child book on the functions of plants, or some such title? It is very good in giving an interest to the subject.

By the way, can you lend me the January number of the "London Journal of Botany" for an article on insect-agency in fertilisation?

LETTER 711. TO J. TRAHERNE MOGGRIDGE. Down, August 27th, 1873.

I thank you for your very interesting letter, and I honour you for your laborious and careful experiments. No one knows till he tries how many unexpected obstacles arise in subjecting plants to experiments.

I can think of no suggestions to make; but I may just mention that I had intended to try the effects of touching the dampened seeds with the minutest drop of formic acid at the end of a sharp glass rod, so as to imitate the possible action of the sting of the ant. I heartily hope that you may be rewarded by coming to some definite result; but I fail five times out of six in my own experiments. I have lately been trying some with poor success, and suppose that I have done too much, for I have been completely knocked up for some days.

LETTER 712. TO J. TRAHERNE MOGGRIDGE. Down, March 10th, 1874.

I am very sorry to hear that the vapour experiments have failed; but nothing could be better, as it seems to me, than your plan of enclosing a number of the ants with the seeds. The incidental results on the power of different vapours in killing seeds and stopping germination appear very curious, and as far as I know are quite new.

P.S. — I never before heard of seeds not germinating except during a certain season; it will be a very strange fact if you can prove this. (712/1. Certain seeds pass through a resting period before germination. See Pfeffer's "Pflanzenphysiologie," Edition I., Volume II., page III.)

LETTER 713. TO H. MULLER. Down, May 30th, 1873.

I am much obliged for your letter received this morning. I write now chiefly to give myself the pleasure of telling you how cordially I admire the last part of your book, which I have finished. (713/1. "Die Befruchtung der Blumen durch Insekten": Leipzig, 1873. An English translation was published in 1883 by Prof. D'Arcy Thompson. The "Prefatory Notice" to this work (February 6th, 1882) is almost the last of Mr. Darwin's writings. See "Life and Letters," page 281.) The whole discussion seems to me quite excellent, and it has pleased me not a little to find that in the rough MS. of my last chapter I have arrived on many points at nearly the same conclusions that you have done, though we have reached them by different routes. (713/2. "The Effects of Cross and Self-Fertilisation in the Vegetable Kingdom": London, 1876.)

LETTER 714. TO F. DELPINO. Down, June 25th {1873}.

I thank you sincerely for your letter. I am very glad to hear about Lathyrus odoratus, for here in England the vars. never cross, and yet are sometimes visited by bees. (714/1. In "Cross and Self-Fertilisation," page 156, Darwin quotes the information received from Delpino and referred to in the present letter — namely, that it is the fixed opinion of the Italian gardeners that the varieties do intercross. See Letter 709.) Pisum sativum I have also many times seen visited by Bombus. I believe the cause of the many vars. not crossing is that under our climate the flowers are self-fertilised at an early period, before the corolla is fully expanded. I shall examine this point with L. odoratus. I have read H. Muller's book, and it seems to me very good. Your criticism had not occurred to me, but is, I think just — viz. that it is much more important to know what insects habitually visit any flower than the various kinds which occasionally visit it. Have you seen A. Kerner's book "Schutzmittel des Pollens," 1873, Innsbruck. (714/2. Afterwards translated by Dr. Ogle as "Flowers and their Unbidden Guests," with a prefatory letter by Charles Darwin, 1878.) It is very interesting, but he does not seem to know anything about the work of other authors.

I have Bentham's paper in my house, but have not yet had time to read a word of it. He is a man with very sound judgment, and fully admits the principle of evolution.

I have lately had occasion to look over again your discussion on anemophilous plants, and I have again felt much admiration at your work. (714/3. "Atti della Soc. Italiana di Scienze Nat." Volume XIII.)

(714/4. In the beginning of August, 1873, Darwin paid the first of several visits to Lord Farrer's house at Abinger. When sending copies of Darwin's letters for the "Life and Letters," Lord Farrer was good enough to add explanatory notes and recollections, from which we quote the following sketch.)

"Above my house are some low hills, standing up in the valley, below the chalk range on the one hand and the more distant range of Leith Hill on the other, with pretty views of the valley towards Dorking in one direction and Guildford in the other. They are composed of the less fertile Greensand strata, and are covered with fern, broom, gorse, and heath. Here it was a particular pleasure of his to wander, and his tall figure, with his broad-brimmed Panama hat and long stick like an alpenstock, sauntering solitary and slow over our favourite walks, is one of the pleasantest of the many pleasant associations I have with the place."

LETTER 715. TO T.H. FARRER (Lord Farrer).

(715/1. The following note by Lord Farrer explains the main point of the letter, which, however, refers to the "bloom" problem as well as to Coronilla: —

"I thought I had found out what puzzled us in Coronilla varia: in most of the Papilionaceae, when the tenth stamen is free, there is nectar in the staminal tube, and the opening caused by the free stamen enables the bee to reach the nectar, and in so doing the bee fertilises the plant. In Coronilla varia, and in several other species of Coronilla, there is no nectar in the staminal tube or in the tube of the corolla. But there are peculiar glands with nectar on the outside of the calyx, and peculiar openings in the tube of the corolla through which the proboscis of the bee, whilst entering the flower in the usual way and dusting itself with pollen, can reach these glands, thus fertilising the plant in getting the nectar. On writing this to Mr. Darwin, I received the following characteristic note.

The first postscript relates to the rough ground behind my house, over which he was fond of strolling. It had been ploughed up and then allowed to go back, and the interest was to watch how the numerous species of weeds of cultivation which followed the plough gradually gave way in the struggle for existence to the well-known and much less varied flora of an English common.")

Bassett, Southampton, August 14th, 1873.

You are the man to conquer a Coronilla. (715/2. In a former letter to Lord Farrer, Darwin wrote: "Here is a maxim for you, 'It is disgraceful to be beaten by a Coronilla.'") I have been looking at the half-dried flowers, and am prepared to swear that you have solved the mystery. The difference in the size of the cells on the calyx under the vexillum right down to the common peduncle is conspicuous. The flour still adhered to this side; I see little bracteae or stipules apparently with glandular ends at the base of the calyces. Do these secrete? It seems to me a beautiful case. When I saw the odd shape of the base of the vexillum, I concluded that it must have some meaning, but little dreamt what that was. Now there remains only the one serious point — viz.the separation of the one stamen. I daresay that you are right in that nectar was originally secreted within the staminal tube; but why has not the one stamen long since cohered? The great difference in structure for fertilisation within the same genus makes one believe that all such points are vary variable. (715/3. Coronilla emerus is of the ordinary papilionaceous type.) With respect to the non-coherence of the one stamen, do examine some flower-buds at a very early age; for parts which are largely developed are often developed to an unusual degree at a very early age, and it seems to me quite possible that the base of the vexillum (to which the single stamen adhered) might thus be developed, and thus keep it separate for a time from the other stamens. The cohering stamens to the right and left of the single one seem to me to be pushed out a little laterally. When you have finished your observations, you really ought to send an account with a diagram to "Nature," recalling your generalisation about the diadelphous structure, and now explaining the exception of Coronilla. (715/4. The observations were published in "Nature," Volume X., 1874, page 169.)

Do add a remark how almost every detail of structure has a meaning where a flower is well examined.

Your observations pleased me so much that I could not sit still for half an hour.

Please to thank Mr. Payne (715/5. Lord Farrer's gardener.) for his remarks, which are of value to me, with reference to Mimosa. I am very much in doubt whether opening the sashes can act by favouring the evaporation of the drops; may not the movement of the leaves shake off the drops, or change their places? If Mr. Payne remembers any plant which is easily injured by drops, I wish he would put a drop or two on a leaf on a bright day, and cover the plant with a clean bell-glass, and do the same for another plant, but without a bell-glass over it, and observe the effects.

Thank you much for wishing to see us again at Abinger, and it is very doubtful whether it will be Coronilla, Mr. Payne, the new garden, the children, E. {Lady Farrer}, or yourself which will give me the most pleasure to see again.