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... I was much perplexed by Oliver's remarks in the 'Natural History Review' on the Primula case, on the lower plants having sexes more often separated than in the higher plants, — so exactly the reverse of what takes place in animals. Hooker in his review of the 'Orchids' repeats this remark. There seems to be much truth in what you say ("Forms which are low in the scale as respects morphological completeness may be high in the scale of rank founded on specialisation of structure and function." — Dr. Gray, in 'Silliman's Journal.'), and it did not occur to me, about no improbability of specialisation in CERTAIN lines in lowly organised beings. I could hardly doubt that the hermaphrodite state is the aboriginal one. But how is it in the conjugation of Confervae — is not one of the two individuals here in fact male, and the other female? I have been much puzzled by this contrast in sexual arrangements between plants and animals. Can there be anything in the following consideration: By ROUGHEST calculation about one-third of the British GENERA of aquatic plants belong to the Linnean classes of Mono and Dioecia; whilst of terrestrial plants (the aquatic genera being subtracted) only one-thirteenth of the genera belong to these two classes. Is there any truth in this fact generally? Can aquatic plants, being confined to a small area or small community of individuals, require more free crossing, and therefore have separate sexes? But to return to our point, does not Alph. de Candolle say that aquatic plants taken as a whole are lowly organised, compared with terrestrial; and may not Oliver's remark on the separation of the sexes in lowly organised plants stand in some relation to their being frequently aquatic? Or is this all rubbish?

... What a magnificent compliment you end your review with! You and Hooker seem determined to turn my head with conceit and vanity (if not already turned) and make me an unbearable wretch.

With most cordial thanks, my good and kind friend, Farewell, C. DARWIN.

[The following passage from a letter (July 28, 1863), to Prof. Hildebrand, contains a reference to the reception of the dimorphic work in France: —

"I am extremely much pleased to hear that you have been looking at the manner of fertilisation of your native Orchids, and still more pleased to hear that you have been experimenting on Linum. I much hope that you may publish the result of these experiments; because I was told that the most eminent French botanists of Paris said that my paper on Primula was the work of imagination, and that the case was so improbable they did not believe in my results."]

CHARLES DARWIN TO ASA GRAY. April 19 [1864].

... I received a little time ago a paper with a good account of your Herbarium and Library, and a long time previously your excellent review of Scott's 'Primulaceae,' and I forwarded it to him in India, as it would much please him. I was very glad to see in it a new case of Dimorphism (I forget just now the name of the plant); I shall be grateful to hear of any other cases, as I still feel an interest in the subject. I should be very glad to get some seed of your dimorphic Plantagos; for I cannot banish the suspicion that they must belong to a very different class like that of the common Thyme. (In this prediction he was right. See 'Forms of Flowers,' page 307.) How could the wind, which is the agent of fertilisation, with Plantago, fertilise "reciprocally dimorphic" flowers like Primula? Theory says this cannot be, and in such cases of one's own theories I follow Agassiz and declare, "that nature never lies." I should even be very glad to examine the two dried forms of Plantago. Indeed, any dried dimorphic plants would be gratefully received...

Did my Lythrum paper interest you? I crawl on at the rate of two hours per diem, with 'Variation under Domestication.'

CHARLES DARWIN TO J.D. HOOKER. Down, November 26 [1864].

... You do not know how pleased I am that you have read my Lythrum paper; I thought you would not have time, and I have for long years looked at you as my Public, and care more for your opinion than that of all the rest of the world. I have done nothing which has interested me so much as Lythrum, since making out the complemental males of Cirripedes. I fear that I have dragged in too much miscellaneous matter into the paper.

... I get letters occasionally, which show me that Natural Selection is making GREAT progress in Germany, and some amongst the young in France. I have just received a pamphlet from Germany, with the complimentary title of "Darwinische Arten-Enstehung-Humbug"!

Farewell, my best of old friends, C. DARWIN.

CHARLES DARWIN TO ASA GRAY. September 10, [1867?].

... The only point which I have made out this summer, which could possibly interest you, is that the common Oxlip found everywhere, more or less commonly in England, is certainly a hybrid between the primrose and cowslip; whilst the P. elatior (Jacq.), found only in the Eastern Counties, is a perfectly distinct and good species; hardly distinguishable from the common oxlip, except by the length of the seed-capsule relatively to the calyx. This seems to me rather a horrid fact for all systematic botanists...

CHARLES DARWIN TO F. HILDEBRAND. Down, November 16, 1868.

My dear Sir,

I wrote my last note in such a hurry from London, that I quite forgot what I chiefly wished to say, namely to thank you for your excellent notices in the 'Bot. Zeitung' of my paper on the offspring of dimorphic plants. The subject is so obscure that I did not expect that any one would have noticed my paper, and I am accordingly very much pleased that you should have brought the subject before the many excellent naturalists of Germany.

Of all the German authors (but they are not many) whose works I have read, you write by far the clearest style, but whether this is a compliment to a German writer I do not know.

[The two following letters refer to the small bud-like "Cleistogamic" flowers found in the violet and many other plants. They do not open and are necessarily self-fertilised:]

CHARLES DARWIN TO J.D. HOOKER. Down, May 30 [1862].

... What will become of my book on Variation? I am involved in a multiplicity of experiments. I have been amusing myself by looking at the small flowers of Viola. If Oliver (Shortly afterwards he wrote: "Oliver, the omniscient, has sent me a paper in the 'Bot. Zeitung,' with most accurate description of all that I saw in Viola.") has had time to study them, he will have seen the curious case (as it seems to me) which I have just made clearly out, viz. that in these flowers, the FEW pollen grains are never shed, or never leave the anther-cells, but emit long pollen tubes, which penetrate the stigma. To-day I got the anther with the included pollen grain (now empty) at one end, and a bundle of tubes penetrating the stigmatic tissue at the other end; I got the whole under a microscope without breaking the tubes; I wonder whether the stigma pours some fluid into the anther so as to excite the included grains. It is a rather odd case of correlation, that in the double sweet violet the small flowers are double; i.e., have a multitude of minute scales representing the petals. What queer little flowers they are.

Have you had time to read poor dear Henslow's life? it has interested me for the man's sake, and, what I did not think possible, has even exalted his character in my estimation...

[The following is an extract from the letter given in part above, and refers to Dr. Gray's article on the sexual differences of plants:]

CHARLES DARWIN TO ASA GRAY. NOVEMBER 26 [1862].

... You will think that I am in the most unpleasant, contradictory, fractious humour, when I tell you that I do not like your term of "precocious fertilisation" for your second class of dimorphism [i.e. for cleistogamic fertilisation]. If I can trust my memory, the state of the corolla, of the stigma, and the pollen-grains is different from the state of the parts in the bud; that they are in a condition of special modification. But upon my life I am ashamed of myself to differ so much from my betters on this head. The TEMPORARY theory (This view is now generally accepted.) which I have formed on this class of dimorphism, just to guide experiment, is that the PERFECT flowers can only be perfectly fertilised by insects, and are in this case abundantly crossed; but that the flowers are not always, especially in early spring, visited enough by insects, and therefore the little imperfect self-fertilising flowers are developed to ensure a sufficiency of seed for present generations. Viola canina is sterile, when not visited by insects, but when so visited forms plenty of seed. I infer from the structure of three or four forms of Balsamineae, that these require insects; at least there is almost as plain adaptation to insects as in the Orchids. I have Oxalis acetosella ready in pots for experiment next spring; and I fear this will upset my little theory... Campanula carpathica, as I found this summer, is absolutely sterile if insects are excluded. Specularia speculum is fairly fertile when enclosed; and this seemed to me to be partially effected by the frequent closing of the flower; the inward angular folds of the corolla corresponding with the clefts of the open stigma, and in this action pushing pollen from the outside of the stigma on to its surface. Now can you tell me, does S. perfoliata close its flower like S. speculum, with angular inward folds? if so, I am smashed without some fearful "wriggling." Are the IMPERFECT flowers of your Specularia the early or the later ones? very early or very late? It is rather pretty to see the importance of the closing of flowers of S. speculum.

['Forms of Flowers' was published in July; in June, 1877, he wrote to Professor Carus with regard to the translation: —

"My new book is not a long one, viz. 350 pages, chiefly of the larger type, with fifteen simple woodcuts. All the proofs are corrected except the Index, so that it will soon be published.

"... I do not suppose that I shall publish any more books, though perhaps a few more papers. I cannot endure being idle, but heaven knows whether I am capable of any more good work."

The review alluded to in the next letter is at page 445 of the volume of 'Nature' for 1878:]

CHARLES DARWIN TO W. THISELTON DYER. Down, April 5, 1878.

My dear Dyer,

I have just read in 'Nature' the review of 'Forms of Flowers,' and I am sure that it is by you. I wish with all my heart that it deserved one quarter of the praises which you give it. Some of your remarks have interested me greatly... Hearty thanks for your generous and most kind sympathy, which does a man real good, when he is as dog-tired as I am at this minute with working all day, so good-bye.

C. DARWIN.

CHAPTER 2.XIII. — CLIMBING AND INSECTIVOROUS PLANTS

[My father mentions in his 'Autobiography' (volume i.) that he was led to take up the subject of climbing plants by reading Dr. Gray's paper, "Note on the Coiling of the Tendrils of Plants." ('Proc. Amer. Acad. of Arts and Sciences,' 1858.) This essay seems to have been read in 1862, but I am only able to guess at the date of the letter in which he asks for a reference to it, so that the precise date of his beginning this work cannot be determined.

In June 1863 he was certainly at work, and wrote to Sir J.D. Hooker for information as to previous publications on the subject, being then in ignorance of Palm's and H. v. Mohl's works on climbing plants, both of which were published in 1827.]

CHARLES DARWIN TO J.D. HOOKER. Down [June] 25 [1863].

My dear Hooker,

I have been observing pretty carefully a little fact which has surprised me; and I want to know from you and Oliver whether it seems new or odd to you, so just tell me whenever you write; it is a very trifling fact, so do not answer on purpose.

I have got a plant of Echinocystis lobata to observe the irritability of the tendrils described by Asa Gray, and which of course, is plain enough. Having the plant in my study, I have been surprised to find that the uppermost part of each branch (i.e. the stem between the two uppermost leaves excluding the growing tip) is CONSTANTLY and slowly twisting round making a circle in from one-half to two hours; it will sometimes go round two or three times, and then at the same rate untwists and twists in opposite directions. It generally rests half an hour before it retrogrades. The stem does not become permanently twisted. The stem beneath the twisting portion does not move in the least, though not tied. The movement goes on all day and all early night. It has no relation to light for the plant stands in my window and twists from the light just as quickly as towards it. This may be a common phenomenon for what I know, but it confounded me quite, when I began to observe the irritability of the tendrils. I do not say it is the final cause, but the result is pretty, for the plant every one and a half or two hours sweeps a circle (according to the length of the bending shoot and the length of the tendril) of from one foot to twenty inches in diameter, and immediately that the tendril touches any object its sensitiveness causes it immediately to seize it; a clever gardener, my neighbour, who saw the plant on my table last night, said: "I believe, Sir, the tendrils can see, for wherever I put a plant it finds out any stick near enough." I believe the above is the explanation, viz. that it sweeps slowly round and round. The tendrils have some sense, for they do not grasp each other when young.

Yours affectionately, C. DARWIN.

CHARLES DARWIN TO J.D. HOOKER. Down, July 14 [1863].

My dear Hooker,

I am getting very much amused by my tendrils, it is just the sort of niggling work which suits me, and takes up no time and rather rests me whilst writing. So will you just think whether you know any plant, which you could give or lend me, or I could buy, with tendrils, remarkable in any way for development, for odd or peculiar structure, or even for an odd place in natural arrangement. I have seen or can see Cucurbitaceae, Passion-flower, Virginian-creeper, Cissus discolor, Common-pea and Everlasting-pea. It is really curious the diversification of irritability (I do not mean the spontaneous movement, about which I wrote before and correctly, as further observation shows): for instance, I find a slight pinch between the thumb and finger at the end of the tendril of the Cucurbitaceae causes prompt movement, but a pinch excites no movement in Cissus. The cause is that one side alone (the concave) is irritable in the former; whereas both sides are irritable in Cissus, so if you excite at the same time both OPPOSITE sides there is no movement, but by touching with a pencil the two branches of the tendril, in any part whatever, you cause movement towards that point; so that I can mould, by a mere touch, the two branches into any shape I like...

CHARLES DARWIN TO ASA GRAY. Down, August 4 [1863].

My present hobby-horse I owe to you, viz. the tendrils: their irritability is beautiful, as beautiful in all its modifications as anything in Orchids. About the SPONTANEOUS movement (independent of touch) of the tendrils and upper internodes, I am rather taken aback by your saying, "is it not wel-known?" I can find nothing in any book which I have... The spontaneous movement of the tendrils is independent of the movement of the upper internodes, but both work harmoniously together in sweeping a circle for the tendrils to grasp a stick. So with all climbing plants (without tendrils) as yet examined, the upper internodes go on night and day sweeping a circle in one fixed direction. It is surprising to watch the Apocyneae with shoots 18 inches long (beyond the supporting stick), steadily searching for something to climb up. When the shoot meets a stick, the motion at that point is arrested, but in the upper part is continued; so that the climbing of all plants yet examined is the simple result of the spontaneous circulatory movement of the upper internodes. Pray tell me whether anything has been published on this subject? I hate publishing what is old; but I shall hardly regret my work if it is old, as it has much amused me...

CHARLES DARWIN TO ASA GRAY. May 28, 1864.

... An Irish nobleman on his death-bed declared that he could conscientiously say that he had never throughout life denied himself any pleasure; and I can conscientiously say that I have never scrupled to trouble you; so here goes. — Have you travelled South, and can you tell me whether the trees, which Bignonia capreolata climbs, are covered with moss or filamentous lichen or Tillandsia? (He subsequently learned from Dr. Gray that Polypodium incanum abounds on the trees in the districts where this species of Bignonia grows. See 'Climbing Plants,' page 103.) I ask because its tendrils abhor a simple stick, do not much relish rough bark, but delight in wool or moss. They adhere in a curious manner by making little disks, like the Ampelopsis... By the way, I will enclose some specimens, and if you think it worth while, you can put them under the simple microscope. It is remarkable how specially adapted some tendrils are; those of Eccremocarpus scaber do not like a stick, will have nothing to say to wool; but give them a bundle of culms of grass, or a bundle of bristles and they seize them well.

CHARLES DARWIN TO J.D. HOOKER. Down, June 10 [1864].

... 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.