RECORD: Darwin, C. R. [1872-1873]. ['On the movement of the leaves of Drosera'], folios 52-68A / Draft of Expression. CUL-DAR61.77-89. (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 2.2026. RN1

NOTE: See record in the Darwin Online manuscript catalogue, enter its Identifier here. Reproduced with permission of the Syndics of Cambridge University Library and William Huxley Darwin. The volumes CUL-DAR 54-61 contain material for Darwin's book Insectivorous plants (1875).

The volume CUL-DAR61 contains Darwin's draft essay 'On the movement of the leaves of Drosera', written 1872-1873. He intended to publish this with a revised edition of Climbing plants before commencing a larger book draft in April 1874 which became Insectivorous plants. See the introduction to this essay by John van Wyhe.


[77]

(52

If a very darkened leaf has (p 12 This occurs with a pale leaf)

Selected, & of especially if this immersed in stronger solution, the first apparent change is that a layer of protoplasm, lining the whole whole inside   Bag lines good p. 5   of the elongated cell, shrivels simultaneously from both ends  Bg p. 5 & 12   & forms a larger elongated mess   The upper layer [illeg]  .  This soon divides into ten or several oval or spherical masses. ; or the larger or elongated mass still more c[illeg] it first becomes divided internally with a multitude of [illeg] spheres, which gradually coalesce into a few larger spheres, & these ultimately then separate from each other.   (a) The centre part first coalesces & then the [illeg] spheres are solid — current as large visible —Not absorption of C. of Ammonia — Perhaps not a carbonate at all.

[pencil insertions largely illeg]

(These masses of red protoplasm present infinitely diversified shapes forms, & undergo incessant changes. of form. The movements, however, are so slow that they would b might easily be overlook  but the results are conspicuous when useless to  the contents of the same cell are examined at intervals of from one to two or three minutes. Long as I watched several cells these slow movements never ceased. In fig [blank] we have a rough, but accurate drawing of the same cell, made at intervals of between two & three minutes; & if exhibiting [illeg] of the

[77v]

Sphere which was split had been immersed for 22° in dose of col. 1 ½ gr. to 1 oz of [illeg]

[paragraph in pencil largely illeg] both cannot be distinguished at all granular matter — p. 9. sphere going up & down on the same side.

[illeg] of currents. Not seen t killed gl. of [illeg]

From these various facts we see that c r. of c. ammon.  Cont

[line in pencil illeg]

Series of changes

In dark red leaf — exterior parts of cushion [illeg] with aggrate, & the content [illeg] & then the granular part of the [illeg] well c[illeg].  (Z)

This had better come at the end of C. of Amm.  p. 54

These [illeg] remarks relate to marginal tentacles

Some solids grow to tentacles of the central discs
p. 6 of new M.S  & other references

[line in ink illeg]

When I [words illeg] aggregation flashing down cells, namely arrested. I thought the C. of Ammonia was absorbed
That the carb absorbed by gland & [rest of line illeg]

But it is truly what it is the carbonate

When is do absorbed. [illeg] down to pedicels —

Not visibly [illeg] for Re[illeg] 24° ?

[line illeg] can effect —

Then what discrimination — no aggregation

Finally    The most remarkable facts met with to the C. of A. is the astonishing [remainder of ink note not transcribed]

[78]

(55

shaped atoms of the red protoplasm were rapidly driven up the longer side of the elongated cells, more slowly across the short ends, & so round & round. The rate of inculcation was extremely variable, & the circulation it often ceased. It probably accounts for a struggling movement which I continuous saw observed in a number of such particles of the protoplasm still cohering together & to larger masses, & likewise for a swinging swaying movement in separate, club-headed filaments of protoplasm. But this [illeg] circulation of the fluid contents obviously will not account for the many strange & diversified changes of form in the larger masses of protoplasm; & these must be due to some [illeg] inherent power is to living matter.

Altogether the vital few cases which may are actually visible in one of the [illeg] cells are as wonderful complex.

(The case of the segregation of the contents of the cells under the above circumstances into colour less fluid liquid & particular seemed to me so curious & [illeg] novel a fact that I tried some other experiments. I immersed a leaf in

[79]

(56

I move a lot

saliva & there was no segregation in 2° 30'; it was then placed in a solution of three grains of carbonate of Ammonia to one oz. of water, & the in 10' the contents of the cells was strongly segregated. The white of an egg did not act in 1° 30', but the same leaf was acted on in 5' by the above solution. With milk a leaf immersed in milk there was a little segregation after 1°. With one immersed in urine there was strong great segregation in 1° 40'; & I but in another leaf similarly immersed there was hardly any action after 7° 30'; what were this latter leaf was, however, on an inactive leaf condition, from the above solution of C. Ammonia did not produced a decided effect until 15' had elapsed. I also placed several leaves in a cold filtered infusion of raw meat;: with one there was no segregation in 5°; but on being placed in the above solution of C C. of Ammonia it was plainly acted on in 5'. With another leaf that first indication of segregation was observed in 2°. 35' 20'. In 3° 5' it elongated segregated masses of protoplasm were becoming beaded, & cover afterwards several distinct spheres were formed. With two other leaves there was some

[80]

(57

segregation in 1° 20' & 1° 50'; after immersion for 24° these two & other leaves presented an extraordinary degree of segregation & the tentacles even down to their base of tentacles, bases which presented a rubbed mottled appearance even to the naked eye. These masses [illeg] In all these cases, the several fewer spherical massed were formed than in the case of were formed than in the leaves treated with the carbonate of ammonia.— The same currents in the protoplasm were observed, as above observed seen. already described. These several observations show how much more rapidly a solution of C. of Ammonia acts than the above-named animal fluids; that the [time] result is nearly the same.)

Again I placed a leaf in a moderately thick solution of gum & then was no segregation after 1° 45'; but this leaf placed in the so treated as before with the solution of C. of Ammonia was acted on in 5'. I then placed immersed some leaves in a solution of three grains of sugar to one ounce of water (& therefore of the same strength as the solution of C. Ammonia) used in these experiments) & no change was caused in 4° 30', whereas the solution of C. of

[81]

(58

Ammonia acted in 5'. I explained Some atoms of some sugar, placed on these glands, it was this were dissolved by the secretion, which was largely increased in quantity, owing no doubt through the principle of to exosmose from the glands. The tentacles did not become inflected, but after 24° the fluid content of these cells had undergone cases a certain amount of segregation. Some leaves were immersed in thick 1 grain of sugar dense solution of sugar, gum & starch, & in the course of a few hours the content of [illeg] the cells had undergone great of the tentacles were greatly   segregated. This was without no doubt was due to exosmose, apparently from the whole surface of the leaves from these [illeg] leaf in the syrup had become quite flaccid; & those in the gum & starch were less flaccid & had their tentacles inflected in & bent in the most irregular manner; many of the long marginal tentacles being twisted like a corkscrew. These flaccid leaves being subsequently placed in distilled water for four days, had their tentacles partially reexpanded & the contents of mass of protoplasm in the cells were partially redissolved; & thus no doubt probably were due to

[82]

(64

solution, this remaining fluid could be purple instead of colourless; unless indeed it be assumed the to   masses of protoplasm the first precipitated  att[illeg] that which [2 words illeg] is in what [illeg].  (a)  The fact of segregation occurring in leaves left for many several hours immersed in distilled water, when some degree of endosmose instead of exosmose would naturally occur, seems at first is much opposed to the above conclusion; (B) At So again is the wonderful rapidity with which the contents of each the each successive cell, one about the size down the pedicel of the tentacles, one beneath the other, becomes cloudy, when to after the gland had absorbed carbonate of ammonia; for it seems hardly credible incredible the fluid could be all the less either could escape, either upward or downwards   though the wall of man cells so quickly as that. Would have been to occur on this view.

On the whole it seems the safest conclusion seems to be that the purple fluid is in so unstable a condition, like certain chemical compounds, that almost any disturbance suffices to cause a change in its character; the

[82v]

(a) There seems to be no close relation between the degree of exosmose & the degree of segregation : soft sugar added to placed on added to drops of viscid secretion of a gland causes much exos as far as I could judge much stronger exosmose that a little carbonate of C. of Ammonia, but causes far less segregation, after a very much longer interval of time.

[83]

(B)

 ; but it is not improbable that the glands of these leaves go on secreting, whilst the cut-off footstalks of the leaf leaves cannot about a sufficient supply of water; for when a plant furnished with roots is submerged there is no inflection of the tentacles & I believe no segregation.

[84]

(65

influence which induces this change being transmitted from the glands down the tentacles, through [illeg] a conducting tissue, which is rendered insufficient through the extension of when not in an oxygenated condition.

(one line open)

Before proceeding to describe the effects of dripping various solutions on to leaves, or with immersing them in it by drops on the leaves or their immersion, it will be necessary to make a few remarks on the action of distilled water, as some of the solutions tried were extremely weak. A growing plant with unimpaired roots may be submerged for days, as I have tried, for not without any effect in being produced. But submerged leaves plants after manner according to Dr Nitschke* are capable of [illeg] a fly if placed on the disc.

Abundant evidence has been given that distilled water dropped on the central discs of the leaves does not cause the marginal tentacles to become submerged inflected.

[85]

(66

When When however leaves are cut off & wholly submerged, the result is somewhat different. Some of the extreme marginal tentacles with elongated [erased pencil insertion above] glands very often become inflected & after a few hours reexpand   give function of time   their inflection went for   Therefore the inflection of their [illeg] tentacles must be disregarded in testing the power of any fluid. [pencil insertion not transcribed]

My usual plan was has been to immerse a certain number of leaves in the solution, & at the same time the same number in the same distilled water used in making the solution, testing both in every way exactly alike. Altogether leave were thus immersed in fresh water & were observed for several hours with the solution other leaves in the solution under trial were acted on. [insertion x] Of the above leaves      — had a few of the oval- leaves petal slightly [illeg]; & only — leaves had all.

[in margin:] Perhaps may state how long leaves were experimented on, but I ought to add that in some few cases these were [illeg]

[insertion x:] ; & until extreme weak solutions were employed there could be no doubt about the difference in the results

[86]

(67

(With respect to the act of segregation from the action of distilled water in the cells lying immediately beneath the oval glands, I first ca examined on successive days under the compound microscope 18 18 19 20 20 15 17° leaves, so as to & ascertained that the purple fluid in within the cells was perfectly homogenous. I then submerged them in the water for various period less than 24°, taking care not to touch the glands, & after tearing them under water for the following intervals times, again examined them under the same power, & found not a trace of segregation in the thirteen following cases. One leaf was left submerged for only 1° 42'; two leaves for 2°; two for 3°; four three for 3° 15'; three for 4° 30' ; one for 6° 15' one for 16° 30', & another for 21° 15'. In these two latter cases the glands were blackened, showing the segregation had commenced within them, but there was no trace of

[in margin:] There some cases to add

[87]

(68

Segregation in the cells of periods close   beneath the oval glands; & these cells have always afforded very [illeg] criterion in judging of the effects of various solutions. [illeg]

[new sheet of paper pasted on:]

In the following On the other hand in seven cases agregation was observed after immersion for the following periods periods here stated. In one unique instance (subject to some doubt) these a few spheres of protoplasm appeared to have been developed after an immersion of only 3° 30'; but I have come since to doubt whether very [illeg] this specimen was carefully examined before immersion. In a second instance after immersion for 5° 45' there was some little segregation ad in most of the tentacles, & this became strongly pronounced after 21°. In a third instance after an immersion of 5° 40' transparent lines could be seen in the homogenous purple fluid, but that which is the first which shows this to perhaps was commencing, but no distinct masses of protoplasm was developed formed : after 8° from the period of immersion there was fairly moderately well marked segregation in the uppermost part of the tentacles, but this did not increase during the 13 succeeding the next thirteen hours. In three other instances, after immersion for 6° 15', 16°, & 17°, the purple fluid in the cells beneath the glands no longer appeared quite homogenous, yet no spherical or elongated masses of protoplasm had been formed. Lastly

In the seventh instance after immersion for 15° 15' these strong segregation has arisen in all the tentacles.

[88]

(36

Ch. IV.

Aggregation

[illeg]

1 — 1'. 42

    2 2°

    —

    2 3°

8  3   3° 15

    3   4°  30'

    1    16  30

    1    21°

    —

    13 — [3 words illeg]

           Aggregation

⸮   1 — 3° 30' [illeg]?

    1  5°  45' – a little aggregation

    2  5°  40' [3 words illeg]

    3  6°  15  not quite [illeg]

    4  16°

    5  17

    15  15. [illeg] agren

[88v]

[Draft of Expression, p. 138, on the Cynopithecus niger.]

(a text)

niger, lately referred to, acting in this manner, & at the same time depresses the crest of hair on its forehead & shows its teeth; so that the movements of the feature are nearly the same when angered & pleased; & the two expressions can be distinguished only by those who are familiar with the animal.)

204 X

[89]

(68A

From these cases facts we see that it is extremely rare for [illeg] distinct segregation to occur after immersion in distilled water in less than five or six hours , for any period less than four or five hours;  (Z)  & that it often generally does not occur until a much longer period has elapsed.)

Previously used
p.62 & 63 new M.S

(I may add that many leaves were left in the distilled water for from 24° to 48°, & the glands in all these cases were blackened showing the segregation had come occurred within their cells; & in those specimens which were examined these was also fairly well marked segregation in the cells of the b pedicels of the tentacles.)

(Z) but when leaves been within the pure water I had [words illeg]


Return to homepage

Citation: John van Wyhe, ed. 2002-. The Complete Work of Charles Darwin Online. (http://darwin-online.org.uk/)

File last updated 23 February, 2026