CHARLES DARWIN TO ASA GRAY. June 8 [1860].
... I have been making some little trifling observations which have interested and perplexed me much. I find with primroses and cowslips, that about an equal number of plants are thus characterised.
SO-CALLED (by me) MALE plant. Pistil much shorter than stamens; stigma rather smooth,--POLLEN GRAINS LARGE, throat of corolla short.
SO-CALLED FEMALE plant. Pistil much longer than stamens, stigma rougher, POLLEN-GRAINS SMALLER,--throat of corolla long.
I have marked a lot of plants, and expected to find the so-called male plant barren; but judging from the feel of the capsules, this is not the case, and I am very much surprised at the difference in the size of the pollen... If it should prove that the so-called male plants produce less seed than the so-called females, what a beautiful case of gradation from hermaphrodite to unis.e.xual condition it will be! If they produce about equal number of seed, how perplexing it will be.
CHARLES DARWIN TO J.D. HOOKER. Down, December 17 [1860?].
... I have just been ordering a photograph of myself for a friend; and have ordered one for you, and for heaven"s sake oblige me, and burn that now hanging up in your room.--It makes me look atrociously wicked.
... In the spring I must get you to look for long pistils and short pistils in the rarer species of Primula and in some allied Genera. It holds with P. Sinensis. You remember all the fuss I made on this subject last spring; well, the other day at last I had time to weigh the seeds, and by Jove the plants of primroses and cowslip with short pistils and large grained pollen (Thus the plants which he imagined to be tending towards a male condition were more productive than the supposed females.) are rather more fertile than those with long pistils, and small-grained pollen. I find that they require the action of insects to set them, and I never will believe that these differences are without some meaning.
Some of my experiments lead me to suspect that the large-grained pollen suits the long pistils and the small-grained pollen suits the short pistils; but I am determined to see if I cannot make out the mystery next spring.
How does your book on plants brew in your mind? Have you begun it?...
Remember me most kindly to Oliver. He must be astonished at not having a string of questions, I fear he will get out of practice!
[The Primula-work was finished in the autumn of 1861, and on November 8th he wrote to Sir J.D. Hooker:--
"I have sent my paper on dimorphism in Primula to the Linn. Soc. I shall go up and read it whenever it comes on; I hope you may be able to attend, for I do not suppose many will care a penny for the subject."
With regard to the reading of the paper (on November 21st), he wrote to the same friend:--
"I by no means thought that I produced a "tremendous effect" in the Linn. Soc., but by Jove the Linn. Soc. produced a tremendous effect on me, for I could not get out of bed till late next evening, so that I just crawled home. I fear I must give up trying to read any paper or speak; it is a horrid bore, I can do nothing like other people."
To Dr. Gray he wrote, (December 1861):--
"You may rely on it, I will send you a copy of my Primula paper as soon as I can get one; but I believe it will not be printed till April 1st, and therefore after my Orchid Book. I care more for your and Hooker"s opinion than for that of all the rest of the world, and for Lyell"s on geological points. Bentham and Hooker thought well of my paper when read; but no one can judge of evidence by merely hearing a paper."
The work on Primula was the means of bringing my father in contact with the late Mr. John Scott, then working as a gardener in the Botanic Gardens at Edinburgh,--an employment which he seems to have chosen in order to gratify his pa.s.sion for natural history. He wrote one or two excellent botanical papers, and ultimately obtained a post in India.
(While in India he made some admirable observations on expression for my father.) He died in 1880.
A few phrases may be quoted from letters to Sir J.D. Hooker, showing my father"s estimate of Scott:--
"If you know, do please tell me who is John Scott of the Botanical Gardens of Edinburgh; I have been corresponding largely with him; he is no common man."
"If he had leisure he would make a wonderful observer; to my judgment I have come across no one like him."
"He has interested me strangely, and I have formed a very high opinion of his intellect. I hope he will accept pecuniary a.s.sistance from me; but he has. .h.i.therto refused." (He ultimately succeeded in being allowed to pay for Mr. Scott"s pa.s.sage to India.)
"I know nothing of him excepting from his letters; these show remarkable talent, astonishing perseverance, much modesty, and what I admire, determined difference from me on many points."
So highly did he estimate Scott"s abilities that he formed a plan (which however never went beyond an early stage of discussion) of employing him to work out certain problems connected with intercrossing.
The following letter refers to my father"s investigations on Lythrum (He was led to this, his first case of trimorphism by Lecoq"s "Geographie Botanique," and this must have consoled him for the trick this work played him in turning out to be so much larger than he expected. He wrote to Sir J.D. Hooker: "Here is a good joke: I saw an extract from Lecoq, "Geograph. Bot.," and ordered it and hoped that it was a good sized pamphlet, and nine thick volumes have arrived!"), a plant which reveals even a more wonderful condition of s.e.xual complexity than that of Primula. For in Lythrum there are not merely two, but three castes, differing structurally and physiologically from each other:]
CHARLES DARWIN TO ASA GRAY. Down, August 9 [1862].
My dear Gray,
It is late at night, and I am going to write briefly, and of course to beg a favour.
The Mitch.e.l.la very good, but pollen apparently equal-sized. I have just examined Hottonia, grand difference in pollen. Echium vulgare, a humbug, merely a case like Thymus. But I am almost stark staring mad over Lythrum (On another occasion he wrote (to Dr. Gray) with regard to Lythrum: "I must hold hard, otherwise I shall spend my life over dimorphism."); if I can prove what I fully believe, it is a grand case of TRIMORPHISM, with three different pollens and three stigmas; I have castrated and fertilised above ninety flowers, trying all the eighteen distinct crosses which are possible within the limits of this one species! I cannot explain, but I feel sure you would think it a grand case. I have been writing to Botanists to see if I can possibly get L.
hyssopifolia, and it has just flashed on me that you might have Lythrum in North America, and I have looked to your Manual. For the love of heaven have a look at some of your species, and if you can get me seed, do; I want much to try species with few stamens, if they are dimorphic; Nesaea verticillata I should expect to be trimorphic. Seed! Seed! Seed!
I should rather like seed of Mitch.e.l.la. But oh, Lythrum!
Your utterly mad friend, C. DARWIN.
P.S.--There is reason in my madness, for I can see that to those who already believe in change of species, these facts will modify to a certain extent the whole view of Hybridity. (A letter to Dr. Gray (July, 1862) bears on this point: "A few days ago I made an observation which has surprised me more than it ought to do--it will have to be repeated several times, but I have scarcely a doubt of its accuracy. I stated in my Primula paper that the long-styled form of Linum grandiflorum was utterly sterile with its own pollen; I have lately been putting the pollen of the two forms on the stigma of the SAME flower; and it strikes me as truly wonderful, that the stigma distinguishes the pollen; and is penetrated by the tubes of the one and not by those of the other; nor are the tubes exserted. Or (which is the same thing) the stigma of the one form acts on and is acted on by pollen, which produces not the least effect on the stigma of the other form. Taking s.e.xual power as the criterion of difference, the two forms of this one species may be said to be generically distinct.")
[On the same subject he wrote to Sir Joseph Hooker in August 1862:--
"Is Oliver at Kew? When I am established at Bournemouth I am completely mad to examine any fresh flowers of any Lythraceous plant, and I would write and ask him if any are in bloom."
Again he wrote to the same friend in October:--
"If you ask Oliver, I think he will tell you I have got a real odd case in Lythrum, it interests me extremely, and seems to me the strangest case of propagation recorded amongst plants or animals, viz. a necessary triple alliance between three hermaphrodites. I feel sure I can now prove the truth of the case from a mult.i.tude of crosses made this summer."
In an article, "Dimorphism in the Genitalia of Plants" ("Silliman"s Journal," 1862, volume x.x.xiv. page 419), Dr. Gray pointed out that the structural difference between the two forms of Primula had already been defined in the "Flora of North America," as DIOECIO-DIMORPHISM. The use of this term called forth the following remarks from my father. The letter also alludes to a review of the "Fertilisation of Orchids" in the same volume of "Silliman"s Journal."]
CHARLES DARWIN TO ASA GRAY. Down, November 26 [1862].
My dear Gray,
The very day after my last letter, yours of November 10th, and the review in "Silliman," which I feared might have been lost, reached me.
We were all very much interested by the political part of your letter; and in some odd way one never feels that information and opinions painted in a newspaper come from a living source; they seem dead, whereas all that you write is full of life. The reviews interested me profoundly; you rashly ask for my opinion, and you must consequently endure a long letter. First for Dimorphism; I do not AT PRESENT like the term "Dioecio-dimorphism;" for I think it gives quite a false notion, that the phenomena are connected with a separation of the s.e.xes.
Certainly in Primula there is unequal fertility in the two forms, and I suspect this is the case with Linum; and, therefore I felt bound in the Primula paper to state that it might be a step towards a dioecious condition; though I believe there are no dioecious forms in Primulaceae or Linaceae. But the three forms in Lythrum convince me that the phenomenon is in no way necessarily connected with any tendency to separation of s.e.xes. The case seems to me in result or function to be almost identical with what old C.K. Sprengel called "dichogamy," and which is so frequent in truly hermaphrodite groups; namely, the pollen and stigma of each flower being mature at different periods. If I am right, it is very advisable not to use the term "dioecious," as this at once brings notions of separation of s.e.xes.
... I was much perplexed by Oliver"s remarks in the "Natural History Review" on the Primula case, on the lower plants having s.e.xes 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 s.e.xual 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 cla.s.ses of Mono and Dioecia; whilst of terrestrial plants (the aquatic genera being subtracted) only one-thirteenth of the genera belong to these two cla.s.ses. 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 s.e.xes? 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 s.e.xes 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 pa.s.sage 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."]