RECORD: Darwin, C. R. 1840-1841. Clarkia Pulchella — when in full flower, pistil longer than stamens. CUL-DAR49.3-15. (John van Wyhe ed., 2002-. The Complete Work of Charles Darwin Online, http://darwin-online.org.uk/)
REVISION HISTORY: Transcribed and edited by John van Wyhe 12.2025. RN1
NOTE: See record in the Darwin Online manuscript catalogue, enter its Identifier here. Reproduced with the permission of the Syndics of Cambridge University Library and William Huxley Darwin. The volume CUL-DAR49 contains notes for Natural selection chap. 3 on 'On...organic beings occasionally crossing' or dichogamy.
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Maer October /40/
Clarkia Pulchella (a)when in full flower pistil longer than stamens, but in half bud shorter & in this state (in flower kept in glass of water) the anthers had burst. (would it be impregnated before flower opened?) (in this state the petal-shaped head of stigma is closed whereas in full flower it is expanded??) There are four curious abortive stamens
In half opened flowers many light brown minute orthopterous (?) insects, which readily flew (same as in peas & beans) & I saw pollen adhering to their legs X likewise in flowers of Phlox — In Fucshia (out of door species) the anthers do not burst till flower has been some days open—
In Lobelia fulgens anthers united terminated by thin slit with brush— flower when although sometimes opened, the stigma does not protrude, but when in this state (& even before blossom opened) if the united anthers be
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(a) Reexamined some flowers same results.
In June 24th /41/. Pollen seems to be shed at extremity of anthers, when flower in full bloom
Thrips
is
The Minute insects haunt double stocks, where no trace of pollen. — also female Lychnis
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(a) greatly pressed or moved plenty of pollen is exuded. — Now if stigma at this period be capable of bug fertililizing 1 it must always be so by its own pollen, but if it is capable only when protruded (X in which state it has gained red tinge, is covered with adhesive fluid & is larger & expanded) then it must almost necessarily be fertilized by pollen of other & each later flowers, whose pollen is pushed out by growing stigma. Pollen very minute in this & all above-mentioned flowers not larger scarcely visible—
In white Flox Anthers do not shed pollen, until flowers fully expanded— pretty abundant, stigma of same height & just between anthers— but foreign pollen might easily be wafted to it— Pollen white in one variety & orange in other, varieties otherwise similar.—
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(a) I have reexamined Lobelia. (spec rather faded) the pollen certainly is profusely ready before stigma protrudes X
(& after it is protruded the pollen seems chiefly or entirely gone The stigma before it protrudes (ie when pollen of own flower most profuse) has its bilobed head closed, is green or greenish & small — whereas when Mature it is gradular, dark purplish adhesive & Lobes widely expanded—
In the unopended flowers of Phlox, I observe the segments of stigma are united, whereas they are expanded, with pollen grains adhering to them in the open ones. —
In some purple Phlox from Shrewsbury— I observed in those flowers, which had pollen burst, the stigma in some was expanded & grown above height of anthers in others only partially expanded & same height with anthers
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"Common Flowers" I mean those, in which apparently stigma is ready when anthers burst, & the impregnation might thus readily take place by breath of air (which I observe is effectual) & pollen from other flower might thus be most readily introduced—
Potentilla 2 spec. — In Goodesia judging from one bud. — Anthers partially burst. whilst before cleft head of stigma expanded. But pollen most abundant whilst stigma ready
Nasturtium in one branch anther of one stamen in each flower only had burst first one & other
Campanula— Oenothera in some of these pollen anthers burst partially before flower quite opened, & whilst stigma only partially expanded. —
Single Pink.—
Migniotte (anthers seem to burst one after other at longest interval; the upper one (in relation to flowers on stalk) bursting apparently always first. — (stigma must remain long open to impregnation. —)
Honeysuckle. Anthers burst before flower opens (damp autumn) I cannot see that stigma is not (a)
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In Fuchsia, in a scarlet sage Salvia (only a little longer) & in a Stylidium ((I do not believe it is Stylidium, for it was Pentand Monogynia)) pistil much longer than stamens— But in Fuchsia flowers depends.
Maer. June
in young flowers of Stylidium, the style is bent & stigma is covered with pollen so that Stylidium is hostile. There is some fact about pistil of this genus bending to anthers
In above flowers I know not at what period pollen is ripe—
⸮Can I not count in how many erect British flowers pistil much longer than stamens?
(a) then ready. When flower fully open, stamens & pistil fully exposed. — Pollen large. — Scabiosa arvensis. Flower like imposter— anthers & stigma fully exposed do Veronica. A foreign Tetradynam. flower) I expect nearly all of this class, Trollius, external rings of anthers mature before inner ones. A Polyand. Polygy plant. Convovulus stigma a god deal larger than anthers. Cistus. Polyand. Polyg. Rhododendron. Poppy Polyand. Monog. outer rings of anthers apparently mature after the interior just reverse of Trollius. I have examined many others— normal structure. — (Keg. Pent. Digyn. —) a Foreign Heath.
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Anchusa (a) anthers adhere to tube of corolla— at base of each segments or petal , a brush (of succulent spikes as in many flowers) covers each anther.) so that the anthers, even when burst are quite concealed— Stigma a little (not more than in very many flowers) taller than anthers, from which it is almost separated by brush.— stigma alone visible externally.—
During movement of plant, pollen would easily (as easily, perhaps, as in heartease reach stigma & by insects foreign pollen would most readily beintroduced: I have seen June (41) a small bee crawling in. & small Humble sucking. every flower on plant
Candy-tuft ( ) a Tetradyn: plant style as long again as stamens, as latter are somewhat concealed in little petals, as stigma quite exposed, as flower grows upright foreign pollen, I should think, would be introduced almost as readily as own—
Loasa tatentia. unready stamen at rt angle to pistil [sic] when pollen ready stand up & apparently one after other when erect longer than pistil. From these facts. foreign pollen (a)
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(a) might readily be introduced, as pollen is successively ready— Latter facts presumptive arguement of stigma remaining long subject to impregnation— As does fact of stigma bending first to one stamen & then to other—
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Lychnis Dioica— Stamens Anthers not particularly exposed.— rather so— pollen by no means excessively minute but decidedly minute nor very abundant spherical smooth
Four little upper stamens shed their pollen before four lower— & in some of the unopened bud the former were partially shed— P.P. The male flowers have short simple, pointed abortive pistil— which in its abortive state may be compared to the stigma &c &c in double flowers— I have not examined female flowers Stigmas rather long X & rather much exposed but not universally — One pod seemed to have mature seeds.—
Fox glove (a) Pollen very abundant & minute.— 2 longer stamens are ready first anthers lie on upper side of tube of corolla Lower side of mouth of corolla set with fine hairs, which necessarily catch (as I saw) pollen & upon bee entering must clog whole body. How often hairs are concerned with catching pollen— so then Bees Must bring either indifferently either pollen of own body flower or of some foreign flower—
In a Campanula— I remember lines or bands of hair on sides fronting anthers, covered with pollen— In pea on middle of pistil.
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(a) p.4 Anchusa=This is type of several analogous structures=viz of plant which produces small white fleshy balls—
P.P. I have twice seen young Thrips in Lychnis covered with pollen=never never saw Bees visiting this most common (June 41) flower=Saw Fly. visit several flowers & flew directly from male to female. directly afterwards saw other (name given in other place) with long legs & less adapted & all its legs & abdomen, base of sucker & hairy antenna profusely dusted with Lychnis pollen. I saw these 2 flies in 5 minutes— All pods in this group had seed vessels very full. In another group. do.— at last I saw Humble visit almost every flower almost in whole group.— sufficient to have impregnated that one can never say that Bees do not visit flowers till after very long experience.— I certainly do not believe there exists a flower, wh. in ten generations is not crossed.—
(a) Stigma does not expand or pollen burst till flower sometime fully expanded
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In Centaurea Jacea (a Composite) pollen bec mature before floret expands., stigma apparently not;— at base of stigma a little collar of bristles, which carries upwards in its growth pollen from the anthers before segments of corolla are expanded high above them.—
Is this not analogous to brush on middle of stigma of pea, carrying out pollen? (The pollen seems long to adhere to this collar) The pistil when thus grown long appears mature & in this state it must be more likely impreg by pollen of other florets rather than its own— Some Several other or most Composites appear to have projecting stigma & good stock of pollen.— Mem one order is in fact monoœcious:—
Kolreuter says all pollen aculeate & requires insects.—
Verbena. var. (hybrid??) of Melindres. Small tuft at mouth of tube of corolla partially hiding (almost, as, but less than in Anchusa) the anthers which hardly project above. Nevertheless the base of each corolla, inside this collar, was scattered with pollen as if anthers, had burst violently. stigma projecting a little above. "common flower"
(a)
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Didynam.
(a) in a pale–flesh–coloured species of Verbena with large leaves–the knotted papillae at mouth of corolla, project entirely over tube & quite hide anthers & generally quite stigma. Stigma reaches to between the pairs of anthers (as general in Didynam) In a pink species— stigma holds same relative position (wi both with had pollen adhering) but the papillae make mere fringe .— Stigma does not reach above— This brush in former species closes Mouth of corolla, as perfectly. as that, together with anthers themselves do in Vinca
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Mallope (a mallow fam:) many divided stigma so surrounded by the monadelph. anthers, that if mature (& I see no reason against) would probably be always impregnated at some time by its own pollen, except by action of insects.
Mem Kohlreuter!!
Examine more.—
In Lupine
I examined only one lower flower stalk pistil bent with stamens bent nearly at right / pr closely covered by sheath which forms tube towards end, open at extremity slit on underside but quite closed on Within this tube the extreme part of this sheath has stigma a little beyond anthers— Motion would send perhaps shake pollen (it appears however rather adhesive) in this tube sheath so as infallibly to reach stigma. (Sheath also protected by 2 folding petals).
Minute insects, I think, alone could carry pollen from flower to flower— I saw larva of Orthopt (1/4 of louse in size) crawl out of orifice at very apex of shield sheath, covered with pollen it must have pushed over little bristly stigma. X — NB In a Convolvulus I saw winged do crawls over anther, & 4 gr of pollen adhered to head which were left on stigma
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Before flower fully expanded — anthers burst.— In others even fading, there is also plenty of pollen:— In the early stages, even in flowers fully expanded— the many-threaded stigma is enclosed in the tube formed by anthers, & their summits externally invisible.— I should think not fit for impregnation in this state, in further advanced state the threads of stigma diverge.— (B) [sketch]
(In any stage I can see no method by which the pollen of lower (B) anthers could reach stigma, except by such means as would equally well bring foreign pollen)—
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Auricula (a) Pistil very short at base of tube of corolla, anthers at mouth of tube of corolla— pollen minute plentiful— apparently not mature till flower expanded.— Except by insects, the fall of in the upright flowers, self impregnation certain.
Some of the flowers droop considerably, these from movement also might be self impregnated readily.—
Stachys. Didynam. The four anthers with plenty of pollen lie so close to stigma, that wonderful if own pollen does not impregnate.— flower very open.—
In little wild Geranium anthers rise round & between divisions of spreading stigma as to make self impregnation very probable. In another creeping species. might be ranked as "common flower" (a)(a)
Stock (cultivated) stigma just beneath 4 longer anthers, touching the short ones— pollen abundant minute.— Tube formed by base of 4 petals narrow & deep embracing closely 4 upper anthers, which moreover lie deepish in it. Self impreg hence almost unavoidable except by minute insects (a)
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(a) In the Chinese Primrose. Anthers buried deeply in tube of corolla, pris stigma reaching to mouth of corolla— Fully as easily impregnated, as flower is upright, by foreign pollen
In Heliotrope, structure almost as in Auricula— tube of corolla narrow, stigma at very bottom, anthers half way down
(a)(a) In two Greenhouse Geraniums; in one stigma just rises above stamen & in the other rises good height above
(a). for foreign pollen. (not easily blown out of other flower.) would not readily work down to stigma so
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Hautboy Sir J. E. Smith1 says wild Eng— partly Dioecious ie some flowers better furnished with stamens, some with pistils— Examined some flowers in late autumn— all had plentiful pollen— minute grained.— & in all stigmas appeared perfect as "Common Flower"— If the ripening fruit of Hautboy proves impregnation;— good case for me.— (a)
Lemon Thyme.. Stigma projecting considerably beyond flower—anthers excessively minute, so that I cannot see pollen, apparently fixed together with filaments to corolla & not reaching above half way up cor tube of corolla— Evidently would be fertilized by foreign pollen full as easily as by its own— I much suspect anthers abortive from domestication.—
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(a) Maer June 14 /41/ Noticed this Day. Common strawberry blossoms, each flower swarming with small Staphylinidae., wh took flight. dusted with pollen, & innumerable numerous very minute moths X also dusted with pollen.—
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Hore-Hound— Marrubium Stigma as long as anthers—placed a little way within genus near Thymus tube of corolla— pollen minute favourable to self impregnation— (outside of nearly all anthers there adhered pearlike globules, much bigger, which I thought was pollen
⸮ova?).—
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(a) In a monstrous Snap-Dragon I saw hairy organ very like this in Penstemon— Though it was not placed symmetrically
[excised] some foreign Heath Mouth.
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Passion Flower Shrewsbury Gynand? Pollen large-grained, rather adhesive tolerably abundant, five large anthers reaching very near
(⸮1/6? of inch) of three large stigma. As flower depends & stalk so pendulous I am astonished it requires artificial impregnation. It would be worth while to measure pollen. I think this tends to shows, how little chance movement has to do with impregnation.—
Shrewsbury July/41/ I found two species do keep nearly erect, & that [see drawing in Ms] anthers burst on lower & opposite, (or at right angles) side ) in reference to secreting surface of stigma— Pollen moreover adheres so that no ordinary shake will disturb it, hence impregnation a great chance without insects.—
X
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Maer June /41/—
Spinach— Dioecious— Pollen minute in great abundance from largeness of anthers— stigmas feathery, but not very large .— Plant when struck by stick, a cloud of pollen— not frequented by insects— flower small & unattractive [NB colour not necessary to attractiveness Mem. Rhubarb yet Rhubarb is conspicuous Cabbage] I wet a leaf on female Spinach & shook neighbouring male plants. it was dusted over— What prodigality, if we consider an absolute creator. —not so compared with seeds & ova— If our faculties permit us to admire. why not to wonder with doubt.— This plant has not succeeded in being attractive or in getting hooked pollen.=
In common Sorrell, female plants appear. tallest??— pollen very minute— about 2/3 of 1/1000 of inch apparently??? Not abundant, but nearly all the anthers were split, probably they burst at certain warm periods— flowers apparently not attractive.— Every seed in some plants seemed swelling, yet only few small number near, but adjoining Hay field so full that with wind in right direction I can fancy air charged with pollen=
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(July 23. 1841 on Woking common saw many hive Bees at Heath.—)
Erica cinerea. (Maer June 41/ a close approach to a shut-up flower. Mem. Antirrhinum!)— teeth of corolla close close round stigma. Which just projects— anthers united round style (a) beneath— shaking of plant would cause pollen most readily to fall on stigma.— Pollen seems ready, at least opening an unopened flower generally spills it, before full inflorescence.— Pollen not small. =Not frequented by Bees (a)=
Melampyrum pratense. Maer Jun 41. Didynam. One of the Scropulaceae in the Scrophular., nearly — upper petal a hood, always back of which style lies closely curls round inner concavity. & projects a little at upper upper edge of mouth of vault, on each side of which brush of pailla, wh. Partly separates the projecting stigma from anthers— insects entering corolla, would scarcely brush both organs. "common flower".—
[sketch]
stigma anthers lower petal
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a) Result of several days observation— But on another day & many subsequent ones I found heath-plantation abounding with many kinds of Bees all hard at work at two species of Heath— Bees certainly cry out & unite "now for Heath—" &c
Citation: John van Wyhe, ed. 2002-. The Complete Work of Charles Darwin Online. (http://darwin-online.org.uk/)
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