LETTER 638. TO J. SCOTT. Down, February 16th {1863}.
Absence from home has prevented me from answering you sooner. I should think that the capsule of Acropera had better be left till it shows some signs of opening, as our object is to judge whether the seeds are good; but I should prefer trusting to your better judgment. I am interested about the Gongora, which I hope hereafter to try myself, as I have just built a small hot-house.
Asa Gray's observations on the rostellum of Gymnadenia are very imperfect, yet worth looking at. Your case of Imatophyllum is most interesting (638/1. A sucker of Imatophyllum minatum threw up a shoot in which the leaves were "two-ranked instead of four-ranked," and showed other differences from the normal.—"Animals and Plants," Edition II., Volume I., page 411.); even if the sport does not flower it will be worth my giving. I did not understand, or I had forgotten, that a single frond on a fern will vary; I now see that the case does come under bud-variation, and must be given by me. I had thought of it only as proof {of} inheritance in cryptogams; I am much obliged for your correction, and will consult again your paper and Mr. Bridgeman's. (638/2. The facts are given in "Animals and Plants," Edition II., Volume I., page 408.) I enclose varieties of maize from Asa Gray. Pray do not thank me for trusting you; the thanks ought to go the other way. I felt a conviction after your first letter that you were a real lover of Natural History.
If you can advance good evidence showing that bisexual plants are more variable than unisexual, it will be interesting. I shall be very glad to read the discussion which you are preparing. I admit as fully as any one can do that cross-impregnation is the great check to endless variability; but I am not sure that I understand your view. I do not believe that the structure of Primula has any necessary relation to a tendency to a dioecious structure, but seeing the difference in the fertility of the two forms, I felt bound unwillingly to admit that they might be a step towards dioeciousness; I allude to this subject in my Linum paper. (638/3. "Linn. Soc. Journal," 1863.) Thanks for your answers to my other queries. I forgot to say that I was at Kew the other day, and I find that they can give me capsules of several Vandeae.
LETTER 639. TO J. SCOTT. Down, March 24th {1863}.
Your letter, as every one you have written, has greatly interested me. If you can show that certain individual Passifloras, under certain known or unknown conditions of life, have stigmas capable of fertilisation by pollen from another species, or from another individual of its own species, yet not by its own individual pollen (its own individual pollen being proved to be good by its action on some other species), you will add a case of great interest to me; and which in my opinion would be quite worth your publication. (639/1. Cases nearly similar to those observed by Scott were recorded by Gartner and Kolreuter, but in these instances only certain individuals were self-impotent. In "Animals and Plants," Edition II., Volume II., page 114, where the phenomenon is fully discussed, Scott's observations ("Trans. Bot. Soc. Edin." 1863) are given as the earliest, except for one case recorded by Lecoq ("Fecondation," 1862). Interesting work was afterwards done by Hildebrand and Fritz Muller, as illustrated in many of the letters addressed to the latter.) I always imagined that such recorded cases must be due to unnatural conditions of life; and think I said so in the "Origin." (639/2. See "Origin of Species," Edition I., page 251, for Herbert's observations on self-impotence in Hippeastrum. In spite of the uniformness of the results obtained in many successive years, Darwin inferred that the plants must have been in an "unnatural state.") I am not sure that I understand your result, {nor} whether it means what I have above obscurely expressed. If you can prove the above, do publish; but if you will not publish I earnestly beg you to let me have the facts in detail; but you ought to publish, for I may not use the facts for years. I have been much interested by what you say on the rostellum exciting pollen to protrude tubes; but are you sure that the rostellum does excite them? Would not tubes protrude if placed on parts of column or base of petals, etc., near to the stigma? Please look at the "Cottage Gardener" (or "Journal of Horticulture") (639/3. "Journal of Horticulture" and "Cottage Gardener," March 31st, 1863. A short note describing Cruger's discovery of self-fertilisation in Cattleya, Epidendrum, etc., and referring to the work of "an excellent observer, Mr. J. Scott." Darwin adds that he is convinced that he has underrated the power of tropical orchids occasionally to produce seeds without the aid of insects.) to be published to-morrow week for letter of mine, in which I venture to quote you, and in which you will see a curious fact about unopened orchid flowers setting seed in West Indies. Dr. Cruger attributes protrusion of tubes to ants carrying stigmatic secretion to pollen (639/4. In Cruger's paper ("Linn. Soc. Journ." VIII., 1865; read March 3rd 1864) he speaks of the pollen-masses in situ being acted on by the stigmatic secretion, but no mention is made of the agency of ants. He describes the pollen-tubes descending "from the {pollen} masses still in situ down into the ovarian canal."); but this is mere hypothesis. Remember, pollen-tubes protrude within anther in Neottia nidus-avis. I did think it possible or probable that perfect fertilisation might have been effected through rostellum. What a curious case your Gongora must be: could you spare me one of the largest capsules? I want to estimate the number of seed, and try my hand if I can make them grow. This, however, is a foolish attempt, for Dr. Hooker, who was here a day or two ago, says they cannot at Calcutta, and yet imported species have seeded and have naturally spread on to the adjoining trees! Dr. Cruger thinks I am wrong about Catasetum: but I cannot understand his letter. He admits there are three forms in two species; and he speaks as if the sexes were separate in some and that others were hermaphrodites (639/5. Cruger ("Linn. Soc. Journal," VIII., page 127) says that the apparently hermaphrodite form is always sterile in Trinidad. Darwin modified his account in the second edition of the orchid book.); but I cannot understand what he means. He has seen lots of great humble-bees buzzing about the flowers with the pollinia sticking to their backs! Happy man!! I have the promise, but not yet surety, of some curious results with my homomorphic seedling cowslips: these have not followed the rule of Chinese Primula; homomorphic seedlings from short-styled parent have presented both forms, which disgusts me.
You will see that I am better; but still I greatly fear that I must have a compulsory holiday. With sincere thanks and hearty admiration at your powers of observation...
My poor P. scotica looks very sick which you so kindly sent me. (639/6. Sent by Scott, January 6th, 1863.)
LETTER 640. TO J. SCOTT. April 12th {1863}.
I really hardly know how to thank you enough for your very interesting letter. I shall certainly use all the facts which you have given me (in a condensed form) on the sterility of orchids in the work which I am now slowly preparing for publication. But why do you not publish these facts in a separate little paper? (640/1. See Letter 642, note, for reference to Scott's paper.) They seem to me well worth it, and you really ought to get your name known. I could equally well use them in my book. I earnestly hope that you will experiment on Passiflora, and let me give your results. Dr. A. Gray's observations were made loosely; he said in a letter he would attend this summer further to the case, which clearly surprised him much. I will say nothing about the rostellum, stigmatic utriculi, fertility of Acropera and Catasetum, for I am completely bewildered: it will rest with you to settle these points by your excellent observations and experiments. I must own I never could help doubting Dr. Hooker's case of the poppy. You may like to hear what I have seen this morning: I found (640/2. See Letter 658.) a primrose plant with flowers having three pistils, which when pulled asunder, without any tearing, allowed pollen to be placed on ovules. This I did with three flowers—pollen-tubes did not protrude after several days. But this day, the sixteenth (N.B.—primulas seem naturally slowly fertilised), I found many tubes protruded, and, what is very odd, they certainly seemed to have penetrated the coats of the ovules, but in no one instance the foramen of the ovule!! I mention this because it directly bears on your explanation of Dr. Cruger's case. (640/3. Cruger's case here referred to is doubtless the cleistogamic fertilisation of Epidendrum, etc. Scott discusses the question of self-fertilisation at great length in a letter to Darwin dated April, and obviously written in 1863. In Epidendrum he observed a viscid matter extending from the stigmatic chamber to the anther: pollen-tubes had protruded from the anther not only where it was in contact with the viscid matter, but also from the central part, and these spread "over the anterior surface of the rostellum downward into the stigma." Cruger believed the viscid matter reaching the anther was a necessary condition for the germination of the pollen-grains. Scott points out that the viscid matter is produced in large quantity only after the pollen-grains have penetrated the stigma, and that it is, in fact, a consequence, not a preliminary to fertilisation. He finally explains Cruger's case thus: "The greater humidity and equability of temperature consequent on such conditions {i.e. on the flowers being closed} is, I believe, the probable cause of these abnormally conditioned flowers so frequently fertilising themselves." Scott also calls attention to the danger of being deceived by fungal hyphae in observations on germination of pollen.) I believe that your explanation is right; I should never have thought of it; yet this was stupid of me, for I remember thinking that the almost closed imperfect flowers of Viola and Oxalis were related to the protrusion of the pollen-tubes. My case of the Aceras with the aborted labellum squeezed against stigma supports your view. (640/4. See "Fertilisation of Orchids," Edition II., page 258: the pollen germinated within the anther of a monstrous flower.) Dr. Cruger's notion about the ants was a simple conjecture. About cryptogamic filaments, remember Dr. C. says that the unopened flowers habitually set fruit. I think that you will change your views on the imperfect flowers of Viola and Oxalis...