More Letters of Charles Darwin Volume II

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LETTER 632. TO J.D. HOOKER.
Down, 18th [November 1862].

Strange to say, I have only one little bother for you to-day, and that is
to let me know about what month flowers appear in Acropera Loddigesii and
luteola; for I want extremely to beg a few more flowers, and if I knew the
time I would keep a memorandum to remind you. Why I want these flowers is
(and I am much alarmed) that Mr. J. Scott, of Bot. Garden of Edinburgh (do
you know anything of him?) has written me a very long and clever letter, in
which he confirms most of my observations; but tells me that with much
difficulty he managed to get pollen into orifice, or as far as mouth of
orifice, of six flowers of A. Loddigesii (the ovarium of which I did not
examine), and two pods set; one he gathered, and saw a very few ovules, as
he thinks, on the large and mostly rudimentary placenta. I shall be most
curious to hear whether the other pod produces a good lot of seed. He says
he regrets that he did not test the ovules with chemical agents: does he
mean tincture of iodine? He suggests that in a state of nature the viscid
matter may come to the very surface of stigmatic chamber, and so pollen-
masses need not be inserted. This is possible, but I should think
improbable. Altogether the case is very odd, and I am very uneasy, for I
cannot hope that A. Loddigesii is hermaphrodite and A. luteola the male of
the same species. Whenever I can get Acropera would be a very good time
for me to look at Vanda in spirits, which you so kindly preserved for me.

LETTER 633. TO J. SCOTT.

(633/1. The following is Darwin's reply to the above letter from Scott.
In the first edition of "Fertilisation of Orchids" (page 209) he assumed
that the sexes in Acropera, as in Catasetum, were separate. In the second
edition (page 172) he writes: "I was, however, soon convinced of my error
by Mr. Scott, who succeeded in artificially fertilising the flowers with
their own pollen. A remarkable discovery by Hildebrand (633/2. "Bot.
Zeitung," 1863 and 1865.), namely, that in many orchids the ovules are not
developed unless the stigma is penetrated by the pollen-tubes...explains
the state of the ovarium in Acropera, as observed by me." In regard to
this subject see Letter 608.)

Down, November 12th, 1862.

I thank you most sincerely for your kindness in writing to me, and for
[your] very interesting letter. Your fact has surprised me greatly, and
has alarmed me not a little, for if I am in error about Acropera I may be
in error about Catasetum. Yet when I call to mind the state of the
placentae in A. luteola, I am astonished that they should produce ovules.
You will see in my book that I state that I did not look at the ovarium of
A. Loddigesii. Would you have the kindness to send me word which end of
the ovarium is meant by apex (that nearest the flower?), for I must try and
get this species from Kew and look at its ovarium. I shall be extremely
curious to hear whether the fruit, which is now maturing, produces a large
number of good and plump seed; perhaps you may have seen the ripe capsules
of other Vandeae, and may be able to form some conjecture what it ought to
produce. In the young, unfertilised ovaria of many Vandeae there seemed an
infinitude of ovules. In desperation it occurs to me as just possible, as
almost everything in nature goes by gradation, that a properly male flower
might occasionally produce a few seeds, in the same manner as female plants
sometimes produce a little pollen. All your remarks seem to me excellent
and very interesting, and I again thank you for your kindness in writing to
me. I am pleased to observe that my description of the structure of
Acropera seems to agree pretty well with what you have observed. Does it
not strike you as very difficult to understand how insects remove the
pollinia and carry them to the stigmas? Your suggestion that the mouth of
the stigmatic cavity may become charged with viscid matter and thus secure
the pollinia, and that the pollen-tubes may then protrude, seems very
ingenious and new to me; but it would be very anomalous in orchids, i.e. as
far as I have seen. No doubt, however, though I tried my best, I shall be
proved wrong in many points. Botany is a new subject to me. With respect
to the protrusion of pollen-tubes, you might like to hear (if you do not
already know the fact) that, as I saw this summer, in the little imperfect
flowers of Viola and Oxalis, which never open, the pollen-tubes always come
out of the pollen-grain, whilst still in the anthers, and direct themselves
in a beautiful manner to the stigma seated at some little distance. I hope
that you will continue your very interesting observations.

LETTER 634. TO J. SCOTT.
Down, November 19th [1862].

I am much obliged for your letter, which is full of interesting matter. I
shall be very glad to look at the capsule of the Acropera when ripe, and
pray present my thanks to Mr. MacNab. (634/1. See Letter 608 (Lindley,
December 15th, 1861). Also "Fertilisation of Orchids," Edition II., page
172, for an account of the observations on Acropera which were corrected by
Scott.) I should like to keep it till I could get a capsule of some other
member of the Vandeae for comparison, but ultimately all the seeds shall be
returned, in case you would like to write any notice on the subject. It
was, as I said (634/2. Letter 633.), only "in desperation" that I
suggested that the flower might be a male and occasionally capable of
producing a few seeds. I had forgotten Gartner's remark; in fact, I know
only odds and ends of Botany, and you know far more. One point makes the
above view more probable in Acropera than in other cases, viz. the presence
of rudimentary placentae or testae, for I cannot hear that these have been
observed in the male plants. They do not occur in male Lychnis dioica, but
next spring I will look to male holly flowers. I fully admit the
difficulty of similarity of stigmatic chamber in the two Acroperas. As far
as I remember, the blunt end of pollen-mass would not easily even stick in
the orifice of the chamber. Your view may be correct about abundance of
viscid matter, but seems rather improbable. Your facts about female
flowers occurring where males alone ought to occur is new to me; if I do
not hear that you object, I will quote the Zea case on your authority in
what I am now writing on the varieties of the maize. (634/3. See "Animals
and Plants," Edition II., Volume I., page 339: "Mr. Scott has lately
observed the rarer case of female flowers on a true male panicle, and
likewise hermaphrodite flowers." Scott's paper on the subject is in
"Trans. Bot. Soc. Edinburgh," Volume VIII. See Letter 151, Volume I.) I
am glad to hear that you are now working on the most curious subject of
parthenogenesis. I formerly fancied that I observed female Lychnis dioica
seeded without pollen. I send by this post a paper on Primula, which may
interest you. (634/4. "Linn. Soc. Journal," 1862.) I am working on the
subject, and if you should ever observe any analogous case I should be glad
to hear. I have added another very clever pamphlet by Prof. Asa Gray.
Have you a copy of my Orchis book? If you have not, and would like one, I
should be pleased to send one. I plainly see that you have the true spirit
of an experimentalist and good observer. Therefore, I ask whether you have
ever made any trials on relative fertility of varieties of plants (like
those I quote from Gartner on the varieties of Verbascum). I much want
information on this head, and on those marvellous cases (as some Lobelias
and Crinum passiflora) in which a plant can be more easily fertilised by
the pollen of another species than by its own good pollen. I am compelled
to write in haste. With many thanks for your kindness.

LETTER 635. TO J. SCOTT.
Down, 20th [1862?].

What a magnificent capsule, and good Heavens, what a number of seeds! I
never before opened pods of larger orchids. It did not signify a few seed
being lost, as it would be hopeless to estimate number in comparison with
other species. If you sow any, had you not better sow a good many? so I
enclose small packet. I have looked at the seeds; I never saw in the
British orchids nearly so many empty testae; but this goes for nothing, as
unnatural conditions would account for it. I suspect, however, from the
variable size and transparency, that a good many of the seeds when dry (and
I have put the capsule on my chimney-piece) will shrivel up. So I will
wait a month or two till I get the capsule of some large Vandeae for
comparison. It is more likely that I have made some dreadful blunder about
Acropera than that it should be male yet not a perfect male. May there be
some sexual relation between A. Loddigesii and luteola; they seem very
close? I should very much like to examine the capsule of the unimpregnated
flower of A. Loddigesii. I have got both species from Kew, but whether we
shall have skill to flower them I know not. One conjectures that it is
imperfect male; I still should incline to think it would produce by seed
both sexes. But you are right about Primula (and a very acute thought it
was): the long-styled P. sinensis, homomorphically fertilised with
own-form pollen, has produced during two successive homomorphic generations
only long-styled plants. (635/1. In "Forms of Flowers," Edition II., page
216, a summary of the transmission of forms in the "homomorphic" unions of
P. sinensis is given. Darwin afterwards used "illegitimate" for
homomorphic, and "legitimate" for "heteromorphic" ("Forms of Flowers,"
Edition i., page 24).) The short-styled the same, i.e. produced
short-styled for two generations with the exception of a single plant. I
cannot say about cowslips yet. I should like to hear your case of the
Primula: is it certainly propagated by seed?

LETTER 636. TO J. SCOTT.
Down, December 3rd, [1862?].

What a capital observer you are! and how well you have worked the primulas.
All your facts are new to me. It is likely that I overrate the interest of
the subject; but it seems to me that you ought to publish a paper on the
subject. It would, however, greatly add to the value if you were to cover
up any of the forms having pistil and anther of the same height, and prove
that they were fully self-fertile. The occurrence of dimorphic and non-
dimorphic species in the same genus is quite the same as I find in Linum.
(636/1. Darwin finished his paper on Linum in December 1862, and it was
published in the "Linn. Soc. Journal" in 1863.) Have any of the forms of
Primula, which are non-dimorphic, been propagated for some little time by
seed in garden? I suppose not. I ask because I find in P. sinensis a
third rather fluctuating form, apparently due to culture, with stigma and
anthers of same height. I have been working successive generations
homomorphically of this Primula, and think I am getting curious results; I
shall probably publish next autumn; and if you do not (but I hope you will)
publish yourself previously, I should be glad to quote in abstract some of
your facts. But I repeat that I hope you will yourself publish. Hottonia
is dimorphic, with pollen of very different sizes in the two forms. I
think you are mistaken about Siphocampylus, but I feel rather doubtful in
saying this to so good an observer. In Lobelia the closed pistil grows
rapidly, and pushes out the pollen and then the stigma expands, and the
flower in function is monoecious; from appearance I believe this is the
case with your plant. I hope it is so, for this plant can hardly require a
cross, being in function monoecious; so that dimorphism in such a case
would be a heavy blow to understanding its nature or good in all other
cases. I see few periodicals: when have you published on Clivia? I
suppose that you did not actually count the seeds in the hybrids in
comparison with those of the parent-forms; but this is almost necessary
after Gartner's observations. I very much hope you will make a good series
of comparative trials on the same plant of Tacsonia. (636/2. See Scott in
"Linn. Soc. Journal," VIII.) I have raised 700-800 seedlings from
cowslips, artificially fertilised with care; and they presented not a
hair's-breadth approach to oxlips. I have now seed in pots of cowslip
fertilised by pollen of primrose, and I hope they will grow; I have also
got fine seedlings from seed of wild oxlips; so I hope to make out the
case. You speak of difficulties on Natural Selection: there are indeed
plenty; if ever you have spare time (which is not likely, as I am sure you
must be a hard worker) I should be very glad to hear difficulties from one
who has observed so much as you have. The majority of criticisms on the
"Origin" are, in my opinion, not worth the paper they are printed on. Sir
C. Lyell is coming out with what, I expect, will prove really good remarks.
(636/3. Lyell's "Antiquity of Man" was published in the spring of 1863.
In the "Life and Letters," Volume III., pages 8, 11, Darwin's
correspondence shows his deep disappointment at what he thought Lyell's
half-heartedness in regard to evolution. See Letter 164, Volume I.) Pray
do not think me intrusive; but if you would like to have any book I have
published, such as my "Journal of Researches" or the "Origin," I should
esteem it a compliment to be allowed to send it. Will you permit me to
suggest one experiment, which I should much like to see tried, and which I
now wish the more from an extraordinary observation by Asa Gray on
Gymnadenia tridentata (in number just out of Silliman's N. American
Journal) (636/4. In Gymnadenia tridentata, according to Asa Gray, the
anther opens in the bud, and the pollen being somewhat coherent falls on
the stigma and on the rostellum which latter is penetrated by the pollen-
tubes. "Fertilisation of Orchids," Edition II., page 68. Asa Gray's
papers are in "American Journal of Science," Volume XXXIV., 1862, and
XXXVI., 1863.); namely, to split the labellum of a Cattleya, or of some
allied orchis, remove caudicle from pollen-mass (so that no loose grains
are about) and put it carefully into the large tongue-like rostellum, and
see if pollen-tubes will penetrate, or better, see if capsule will swell.
Similar pollen-masses ought to be put on true stigmas of two or three other
flowers of same plants for comparison. It is to discover whether rostellum
yet retains some of its primordial function of being penetrated by pollen-
tubes. You will be sorry that you ever entered into correspondence with
me. But do not answer till at leisure, and as briefly as you like. My
handwriting, I know, is dreadfully bad. Excuse this scribbling paper, as I
can write faster on it, and I have a rather large correspondence to keep
up.

LETTER 637. TO J. SCOTT.
Down, January 21st, 1863.

I thank you for your very interesting letter; I must answer as briefly as I
can, for I have a heap of other letters to answer. I strongly advise you
to follow up and publish your observations on the pollen-tubes of orchids;
they promise to be very interesting. If you could prove what I only
conjectured (from state of utriculi in rostellum and in stigma of Catasetum
and Acropera) that the utriculi somehow induce, or are correlated with,
penetration of pollen-tubes you will make an important physiological
discovery. I will mention, as worth your attention (and what I have
anxiously wished to observe, if time had permitted, and still hope to do)--
viz., the state of tissues or cells of stigma in an utterly sterile hybrid,
in comparison with the same in fertile parent species; to test these cells,
immerse stigmas for 48 hours in spirits of wine. I should expect in
hybrids that the cells would not show coagulated contents. It would be an
interesting discovery to show difference in female organs of hybrids and
pure species. Anyhow, it is worth trial, and I recommend you to make it,
and publish if you do. The pollen-tubes directing themselves to stigma is
also very curious, though not quite so new, but well worth investigation
when you get Cattleya, etc., in flower. I say not so new, for remember
small flowers of Viola and Oxalis; or better, see Bibliography in "Natural
History Review," No. VIII., page 419 (October, 1862) for quotation from M.
Baillon on pollen-tubes finding way from anthers to stigma in Helianthemum.
I should doubt gum getting solid from [i.e. because of] continued
secretion. Why not sprinkle fresh plaster of Paris and make impenetrable
crust? (637/1. The suggestion that the stigma should be covered with a
crust of plaster of Paris, pierced by a hole to allow the pollen-tubes to
enter, bears a resemblance to Miyoshi's experiments with germinating pollen
and fungal spores. See "Pringsheim's Jahrbucher," 1895; "Flora," 1894.)
You might modify experiment by making little hole in one lower corner, and
see if tubes find it out. See in my future paper on Linum pollen and
stigma recognising each other. If you will tell me that pollen smells the
stigma I will try and believe you; but I will not believe the Frenchman (I
forget who) who says that stigma of Vanilla actually attracts mechanically,
by some unknown force, the solid pollen-masses to it! Read Asa Gray in 2nd
Review of my Orchis book on pollen of Gymnadenia penetrating rostellum. I
can, if you like, lend you these Reviews; but they must be returned. R.
Brown, I remember, says pollen-tubes separate from grains before the lower
ends of tubes reach ovules. I saw, and was interested by, abstract of your
Drosera paper (637/2. A short note on the irritability of Drosera in the
"Trans. Bot. Soc. Edin." Volume VII.); we have been at very much the same
work.

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...