... I have now read two German books, and all I believe that has been written on climbers, and it has stirred me up to find that I have a good deal of new matter. It is strange, but I really think no one has explained simple twining plants. These books have stirred me up, and made me wish for plants specified in them. I shall be very glad of those you mention. I have written to Veitch for young Nepenthes and Vanilla (which I believe will turn out a grand case, though a root creeper), if I cannot buy young Vanilla I will ask you. I have ordered a leaf-climbing fern, Lygodium. All this work about climbers would hurt my conscience, did I think I could do harder work. (He was much out of health at this time.)
[He continued his observations on climbing plants during the prolonged illness from which he suffered in the autumn of 1863, and in the following spring. He wrote to Sir J.D. Hooker, apparently in March 1864: —
"For several days I have been decidedly better, and what I lay much stress on (whatever doctors say), my brain feels far stronger, and I have lost many dreadful sensations. The hot-house is such an amusement to me, and my amusement I owe to you, as my delight is to look at the many odd leaves and plants from Kew... The only approach to work which I can do is to look at tendrils and climbers, this does not distress my weakened brain. Ask Oliver to look over the enclosed queries (and do you look) and amuse a broken-down brother naturalist by answering any which he can. If you ever lounge through your houses, remember me and climbing plants."
On October 29, 1864, he wrote to Dr. Gray: —
"I have not been able to resist doing a little more at your godchild, my climbing paper, or rather in size little book, which by Jove I will have copied out, else I shall never stop. This has been new sort of work for me, and I have been pleased to find what a capital guide for observations a full conviction of the change of species is."
On January 19, 1865, he wrote to Sir J.D. Hooker: —
"It is working hours, but I am trying to take a day's holiday, for I finished and despatched yesterday my climbing paper. For the last ten days I have done nothing but correct refractory sentences, and I loathe the whole subject."
A letter to Dr. Gray, April 9, 1865, has a word or two on the subject: —
"I have begun correcting proofs of my paper on 'Climbing Plants.' I suppose I shall be able to send you a copy in four or five weeks. I think it contains a good deal new and some curious points, but it is so fearfully long, that no one will ever read it. If, however, you do not SKIM through it, you will be an unnatural parent, for it is your child."
Dr. Gray not only read it but approved of it, to my father's great satisfaction, as the following extracts show: —
"I was much pleased to get your letter of July 24th. Now that I can do nothing, I maunder over old subjects, and your approbation of my climbing paper gives me VERY great satisfaction. I made my observations when I could do nothing else and much enjoyed it, but always doubted whether they were worth publishing. I demur to its not being necessary to explain in detail about the spires in CAUGHT tendrils running in opposite directions; for the fact for a long time confounded me, and I have found it difficult enough to explain the cause to two or three persons." (August 15, 1865.)
"I received yesterday your article (In the September number of 'Silliman's Journal,' concluded in the January number, 1866.) on climbers, and it has pleased me in an extraordinary and even silly manner. You pay me a superb compliment, and as I have just said to my wife, I think my friends must perceive that I like praise, they give me such hearty doses. I always admire your skill in reviews or abstracts, and you have done this article excellently and given the whole essence of my paper... I have had a letter from a good Zoologist in S. Brazil, F. Muller, who has been stirred up to observe climbers and gives me some curious cases of BRANCH-climbers, in which branches are converted into tendrils, and then continue to grow and throw out leaves and new branches, and then lose their tendril character." (October 1865.)
The paper on Climbing Plants was republished in 1875, as a separate book. The author had been unable to give his customary amount of care to the style of the original essay, owing to the fact that it was written during a period of continued ill-health, and it was now found to require a great deal of alteration. He wrote to Sir J.D. Hooker (March 3, 1875): "It is lucky for authors in general that they do not require such dreadful work in merely licking what they write into shape." And to Mr. Murray in September he wrote: "The corrections are heavy in 'Climbing Plants,' and yet I deliberately went over the MS. and old sheets three times." The book was published in September 1875, an edition of 1500 copies was struck off; the edition sold fairly well, and 500 additional copies were printed in June of the following year.]
INSECTIVOROUS PLANTS.
[In the summer of 1860 he was staying at the house of his sister-in-law, Miss Wedgwood, in Ashdown Forest, whence he wrote (July 29, 1860), to Sir Joseph Hooker; —
"Latterly I have done nothing here; but at first I amused myself with a few observations on the insect-catching power of Drosera; and I must consult you some time whether my 'twaddle' is worth communicating to the Linnean Society."
In August he wrote to the same friend: —
"I will gratefully send my notes on Drosera when copied by my copier: the subject amused me when I had nothing to do."
He has described in the 'Autobiography' (volume i.), the general nature of these early experiments. He noticed insects sticking to the leaves, and finding that flies, etc., placed on the adhesive glands were held fast and embraced, he suspected that the leaves were adapted to supply nitrogenous food to the plant. He therefore tried the effect on the leaves of various nitrogenous fluids — with results which, as far as they went, verified his surmise. In September, 1860, he wrote to Dr. Gray: —
"I have been infinitely amused by working at Drosera: the movements are really curious; and the manner in which the leaves detect certain nitrogenous compounds is marvellous. You will laugh; but it is, at present, my full belief (after endless experiments) that they detect (and move in consequence of) the 1/2880 part of a single grain of nitrate of ammonia; but the muriate and sulphate of ammonia bother their chemical skill, and they cannot make anything of the nitrogen in these salts! I began this work on Drosera in relation to GRADATION as throwing light on Dionaea."
Later in the autumn he was again obliged to leave home for Eastbourne, where he continued his work on Drosera. The work was so new to him that he found himself in difficulties in the preparation of solutions, and became puzzled over fluid and solid ounces, etc. etc. To a friend, the late Mr. E. Cresy, who came to his help in the matter of weights and measures, he wrote giving an account of the experiments. The extract (November 2, 1860) which follows illustrates the almost superstitious precautions he often applied to his researches: —
"Generally I have scrutinised every gland and hair on the leaf before experimenting; but it occurred to me that I might in some way affect the leaf; though this is almost impossible, as I scrutinised with equal care those that I put into distilled water (the same water being used for dissolving the carbonate of ammonia). I then cut off four leaves (not touching them with my fingers), and put them in plain water, and four other leaves into the weak solution, and after leaving them for an hour and a half, I examined every hair on all eight leaves; no change on the four in water; every gland and hair affected in those in ammonia.
"I had measured the quantity of weak solution, and I counted the glands which had absorbed the ammonia, and were plainly affected; the result convinced me that each gland could not have absorbed more than 1/64000 or 1/65000 of a grain. I have tried numbers of other experiments all pointing to the same result. Some experiments lead me to believe that very sensitive leaves are acted on by much smaller doses. Reflect how little ammonia a plant can get growing on poor soil — yet it is nourished. The really surprising part seems to me that the effect should be visible, and not under very high power; for after trying a high power, I thought it would be safer not to consider any effect which was not plainly visible under a two-thirds object glass and middle eye-piece. The effect which the carbonate of ammonia produces is the segregation of the homogeneous fluid in the cells into a cloud of granules and colourless fluid; and subsequently the granules coalesce into larger masses, and for hours have the oddest movements — coalescing, dividing, coalescing ad infinitum. I do not know whether you will care for these ill-written details; but, as you asked, I am sure I am bound to comply, after all the very kind and great trouble which you have taken."
On his return home he wrote to Sir J.D. Hooker (November 21, 1860): —
"I have been working like a madman at Drosera. Here is a fact for you which is certain as you stand where you are, though you won't believe it, that a bit of hair 1/78000 of one grain in weight placed on gland, will cause ONE of the gland-bearing hairs of Drosera to curve inwards, and will alter the condition of the contents of every cell in the foot-stalk of the gland."
And a few days later to Lyell: —
"I will and must finish my Drosera MS., which will take me a week, for, at the present moment, I care more about Drosera than the origin of all the species in the world. But I will not publish on Drosera till next year, for I am frightened and astounded at my results. I declare it is a certain fact, that one organ is so sensitive to touch, that a weight seventy-eight times less than that, viz., 1/1000 of a grain, which will move the best chemical balance, suffices to cause a conspicuous movement. Is it not curious that a plant should be far more sensitive to the touch than any nerve in the human body? Yet I am perfectly sure that this is true. When I am on my hobby-horse, I never can resist telling my friends how well my hobby goes, so you must forgive the rider."
The work was continued, as a holiday task, at Bournemouth, where he stayed during the autumn of 1862. The discussion in the following letter on "nervous matter" in Drosera is of interest in relation to recent researches on the continuity of protoplasm from cell to cell:]
CHARLES DARWIN TO J.D. HOOKER. Cliff Cottage, Bournemouth. September 26 [1862].
My dear Hooker,
Do not read this till you have leisure. If that blessed moment ever comes, I should be very glad to have your opinion on the subject of this letter. I am led to the opinion that Drosera must have diffused matter in organic connection, closely analogous to the nervous matter of animals. When the glands of one of the papillae or tentacles, in its natural position is supplied with nitrogenised fluid and certain other stimulants, or when loaded with an extremely slight weight, or when struck several times with a needle, the pedicel bends near its base in under one minute. These varied stimulants are conveyed down the pedicel by some means; it cannot be vibration, for drops of fluid put on quite quietly cause the movement; it cannot be absorption of the fluid from cell to cell, for I can see the rate of absorption, which though quick, is far slower, and in Dionaea the transmission is instantaneous; analogy from animals would point to transmission through nervous matter. Reflecting on the rapid power of absorption in the glands, the extreme sensibility of the whole organ, and the conspicuous movement caused by varied stimulants, I have tried a number of substances which are not caustic or corrosive... but most of which are known to have a remarkable action on the nervous matter of animals. You will see the results in the enclosed paper. As the nervous matter of different animals are differently acted on by the same poisons, one would not expect the same action on plants and animals; only if plants have diffused nervous matter, some degree of analogous action. And this is partially the case. Considering these experiments, together with the previously made remarks on the functions of the parts, I cannot avoid the conclusion, that Drosera possesses matter at least in some degree analogous in constitution and function to nervous matter. Now do tell me what you think, as far as you can judge from my abstract; of course many more experiments would have to be tried; but in former years I tried on the whole leaf, instead of on separate glands, a number of innocuous (This line of investigation made him wish for information on the action of poisons on plants; as in many other cases he applied to Professor Oliver, and in reference to the result wrote to Hooker: "Pray thank Oliver heartily for his heap of references on poisons.") substances, such as sugar, gum, starch, etc., and they produced no effect. Your opinion will aid me in deciding some future year in going on with this subject. I should not have thought it worth attempting, but I had nothing on earth to do.
My dear Hooker, Yours very sincerely, CH. DARWIN.
P.S. — We return home on Monday 28th. Thank Heaven!
[A long break now ensued in his work on insectivorous plants, and it was not till 1872 that the subject seriously occupied him again. A passage in a letter to Dr. Asa Gray, written in 1863 or 1864, shows, however, that the question was not altogether absent from his mind in the interim: —
"Depend on it you are unjust on the merits of my beloved Drosera; it is a wonderful plant, or rather a most sagacious animal. I will stick up for Drosera to the day of my death. Heaven knows whether I shall ever publish my pile of experiments on it."
He notes in his diary that the last proof of the 'Expression of the Emotions' was finished on August 22, 1872, and that he began to work on Drosera on the following day.]
CHARLES DARWIN TO ASA GRAY. [Sevenoaks], October 22 [1872].
... I have worked pretty hard for four or five weeks on Drosera, and then broke down; so that we took a house near Sevenoaks for three weeks (where I now am) to get complete rest. I have very little power of working now, and must put off the rest of the work on Drosera till next spring, as my plants are dying. It is an endless subject, and I must cut it short, and for this reason shall not do much on Dionaea. The point which has interested me most is tracing the NERVES! which follow the vascular bundles. By a prick with a sharp lancet at a certain point, I can paralyse one-half the leaf, so that a stimulus to the other half causes no movement. It is just like dividing the spinal marrow of a frog: — no stimulus can be sent from the brain or anterior part of the spine to the hind legs; but if these latter are stimulated, they move by reflex action. I find my old results about the astonishing sensitiveness of the nervous system (!?)of Drosera to various stimulants fully confirmed and extended...
[His work on digestion in Drosera and other points in the physiology of the plant soon led him into regions where his knowledge was defective, and here the advice and assistance which he received from Dr. Burdon Sanderson was of much value:]
CHARLES DARWIN TO J. BURDON SANDERSON. Down, July 25, 1873.
My dear Dr. Sanderson,
I should like to tell you a little about my recent work with Drosera, to show that I have profited by your suggestions, and to ask a question or two.
1. It is really beautiful how quickly and well Drosera and Dionaea dissolve little cubes of albumen and gelatine. I kept the same sized cubes on wet moss for comparison. When you were here I forgot that I had tried gelatine, but albumen is far better for watching its dissolution and absorption. Frankland has told me how to test in a rough way for pepsin; and in the autumn he will discover what acid the digestive juice contains.
2. A decoction of cabbage-leaves and green peas causes as much inflection as an infusion of raw meat; a decoction of grass is less powerful. Though I hear that the chemists try to precipitate all albumen from the extract of belladonna, I think they must fail, as the extract causes inflection, whereas a new lot of atropine, as well as the valerianate [of atropine], produce no effect.
3. I have been trying a good many experiments with heated water... Should you not call the following case one of heat rigor? Two leaves were heated to 130 deg, and had every tentacle closely inflected; one was taken out and placed in cold water, and it re-expanded; the other was heated to 145 deg, and had not the least power of re-expansion. Is not this latter case heat rigor? If you can inform me, I should very much like to hear at what temperature cold-blooded and invertebrate animals are killed.
4. I must tell you my final result, of which I am sure, [as to] the sensitiveness of Drosera. I made a solution of one part of phosphate of ammonia by weight to 218,750 of water; of this solution I gave so much that a leaf got 1/8000 of a grain of the phosphate. I then counted the glands, and each could have got only 1/1552000 of a grain; this being absorbed by the glands, sufficed to cause the tentacles bearing these glands to bend through an angle of 180 deg. Such sensitiveness requires hot weather, and carefully selected young yet mature leaves. It strikes me as a wonderful fact. I must add that I took every precaution, by trying numerous leaves at the same time in the solution and in the same water which was used for making the solution.
5. If you can persuade your friend to try the effects of carbonate of ammonia on the aggregation of the white blood corpuscles, I should very much like to hear the result.
I hope this letter will not have wearied you.
Believe me, yours very sincerely, CHARLES DARWIN.
CHARLES DARWIN TO W. THISELTON DYER. Down, 24 [December 1873?].