RECORD: Darwin, C. R. et al. 1873.10.07-12.12. Desmodium gyrans / Draft of Expression (fragment). CUL-DAR209.12.89-119. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/)

REVISION HISTORY: Transcribed by Christine Chua and edited by John van Wyhe 7.2023. 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 volume CUL-DAR209.12 contains material for Darwin's book Movement in plants (1880).

The Darwin family members who assisted in the observations and recorded notes include Emma, Francis and Leonard. Lettington, the gardener also shared his observations and assisted in the experiments. The draft fragment of Expression is in the hand of Ebenezer Norman and the text corresponds to Expression, p. 331.


(1

Oct 7 /73/

Hedysarum gyrans.

à Desmodium gyrans

Main leaf & minute leaflet beautifully silvery on both surfaces— very quickly removed by S Ether & surface thoroughly wetted.

Lower surface hairy & a ring of thick hairs round pedicel close to terminal leaflets

Consequently when I watered leaves with glass syringe only a few drop remained on & these are spherical.

The water did not cause movement, but the plant was moving slowly (temp. 76°)— I almost suspect rubbing fine pair above little leaflet causes movement.. & I am almost sure that violent shaking branch does.— Do leaflet depend like main leaf in sleep. (N.B I see old leaves a good deal spotted.

(Try water on these after removing with ether & at high temp.) Wax removed as easily almost as easily as in M. pudica from main & little leaflets.—

(The leaflets after being shaken moved up again very quickly— can the plant have acquired the habit of moving up & down, which it performs uselessly — like Expression??)

Do little leaflet go to sleep?

Try metallic squirt & violent stream

At what Temp is wax removed.

[1v]

Desmodium gyrans [illeg]— a vast number of spec judging by Loudons drawings of Hedysarum, the small & almost aborted lateral leaflets. (often only one many plants) quite an exception. in the genus.—

Plant have acquired habit, & when excited by touch expends superfluous nerves energy that is what we shd say with an animal. Uselessness goes for nothing how often must mimosa close with puffs of wind— blades of grass & insects.

2

Oct 7th 4° 15 20. (temp 83°) — I again shook plant & squirt squirted with Boss squirt.— I do not feel sure about leaflets, but great terminal leaflet most rapidly dipped shak themselves till they were inclined at very high-angle— Shaking stem & squirting did this (try gentle squirting at a distance)— as lateral shaking did it cannot be accounted for by weight of drop of water.

I left the plant with a vast number of leave now dependent which had been horizontal & took my walk, at about 4° 40' the leaves had much risen (though so late in evening) & I again shook them, & saw them rapidly sinking (so Henslow probably right. (see also Dic. Class. & Sachs) (Examine plant on hot day under Microscope shade glass)

(Put leaf on stand on cork with 2 crossed pins & clear portion with ether & put on large drop of water for 24°)

(3

Desmodium

Oct 8th 4° 30' P.m. 2 Branches with their upper large leaves horizontal — shook them rather strongly for 15s., & they soon in a minute or two began all to sink down. until they became inclined at about 45°; I think the movement was oscillating up & down— The little leaflets were not affected by this much shaking — after interval of 25m almost all had become again horizontal.

I find 4° 30' a vast number of leaves spotted with yellow which I feel sure was not the case yesterday.

They were watered yesterday about 10° to 11°— It (see Back)

I marked 2 of these leaves with bits of worsted to see how soon yellow spots disappear.—

Oct 9th. 9° 30' Repeated above experiment on 2 separate branches.— (temp 90°)— Several leaves horizontal, or a little inclined upwards— shook these for 15"— The main terminal leaflets all became inclined quickly but not so much as last night— The little lateral leaflets not affected. — I doubt about irritation of Hairy part, causing movement.—

[3v]

The temp. after plant was syringed was during the 29 1/2 hours, before I saw the spotting from 85° to 80° during the day & 74° to 75° during night— I used water which had been in hothouse & I suppose about at least 75° for syringing this latter fact will account for the spotting

(4

Desmodium. gyrans

Oct 9th 4° 15' P.m. I looked at leaves spotted with yellow & dipped them in water: whole surface silvery, & the yellow spots were not wetted as I expected; perhaps they wd have been so if I had tried them earlier.—

Oct 10. 10.45. Temp. of case 76° 1/2 fixed marks on opposite sides in exact line with tip of leaf, & observed 4 leaves for about an hour.

(1) rose great height above mark i.e. Oct. 11th (I today estimated the rise at 3/8' of an inch by tops & mark)/

1 or 2 tenths of an inch; then sank but not as low as mark; rose again to former heights, but during this time long remained stationary; sank a trifle, again rose a trifle.

2 moved but little, sank a little & again rose a little.

(5

Desmodium

3 Sank below mark & moved to left hand, but the latter perhaps only apparent; rose again, still keeping to left of mark, went on rising so as to be far to the left; sank down again far below mark & to left.

4 Sank a little below mark, rose to former position in the same vertical line; rose above mark; on level with mark; sank below mark.

The movement quite insensible, could only be detected by the marks; no jerking movement: may not the jerking m. of the rudimentary lat. leaflets be due to their much less weight & therefore less resistance to the same power. I observed no rotating movement, but may have overlooked it.

Leaves certainly stationary at intervals.

Oct 11th. Temp. of case 76°, I observed 1 leaf. rising; at 11° 33' still rising; at 11° 38. sinking at 12° 20' had risen. at 12° 50 risen an inch or two: I pinched base of main petiole & of main leaflet, there was slight downward movement, but not much, & afterwards the leaf became again horizontal.

(6

Desmodium

Oct 10th watered with metal squirt (Temp. of water 67°) 2 branchlets, bearing 12 leaflets most of them sub-horizontal & 3 nearly vertical. I watered them for several minutes; none of them became much inclined downwards, but I then observed that their laminae were at about an angle of 45° to the horizon: attend carefully to this

Oct 10th 8° 50' P.m. very dark. Temp of case 86°.—

At end of shoots petioles almost vertical & pressed to axis—

Main or terminal leaflet depending vertically. The little leaflets pointing vertically upwards & undergoing far stronger & more rapid movement than I have ever seen, from standing vertically upwards they would ie most parallel parallel to petiole, they would pass by rapid movement through nearly 90° & became horizontal or at right angles to petiole, rotating their laminæ whilst they did so.— I neglected to shake the plant to see effect. This is all wonderful & inexplicable!—

(7

Desmodium gyrans

Oct 11. 10° 30'

The petioles now stand at about 60° to 70° from axis so that it is clear that they rise towards axis when asleep.

At 10° 55' I syringed the plant from above & then laterally 5 times with large garden syringe with water at 67°–68°— After all was over the younger leave very little wet. Temp of case 76° The main leaflets were only a little depressed, I then shook plant & they became more depressed, 40° to 45° below horizon.— The little leaflets not in the least affected — they seem not subject to conscious or unconscious movements—

(Often only 1 or none to leaflets, is this not evidence of rudimentary condition ask Hooker how at Kew.)

The le plant was well shaken at 11°. 1' & it produced no effect on little leaflet at 11° 4' The main leaflets much more depressed. — At 11° 12' beginning, I think to rise at 11° 33' some of them now horizontal.

At above temp of 76° I did not notice any movement of little leaflets

(8

Desmodium

Oct 11th. (In all cases leaves were wound about close to bulb of flowers.)

(Temp. 80 1/2 - 79 1/2 — immersed for 30s, wax removed from borders of upper surface, 1 for additional 1m not from centre.)

(Same leaf temp. 81° - 80° 1/2 for 1m, the wetted borders wider, — under surface not all wetted.)

(same leaf 85° 84° 1/2 for 1m, all upper surface, & the lower except middle.)

(Lateral leaflet of same another leaf, 85° - 84 1/2 for 1m. imperfectly wetted, lower side hardly at all.)

(same lateral leaflet 86 1/4 - 85° 3/4 for 1m, both surfaces well wetted.

(Another leaf 85 - 84 1/2 main leaf for 1m upper surface all wetted, except small portion quite in middle; immersed it for additional 30s. all upper quite wetted, & all lower, except middle part.)—

(Basal part of last leaf. 88° - 87° for 1m. almost completely wetted, but not completely; lower surface after additional 30s. completely wetted.)Th Time seems important element— odd as ether acted so quickly.)

(Basal part of first leaf, 86 1/2 - 86° for 1m, upper surface not largely wetted— lower surface little wetted; little lateral leaflets of do; upper surface well wetted, but not the lower.) (over

[8v]

(A third leaf. 91° - 90 1/2 for 1m. all upper surface wetted, lower surface but little; after addition 30s at 90°, lower surface still but little wetted—

(So protective power variable.)— This same leaf immersed for 1m additional at 98° - 97°, even now lower surface not completely wetted

(8A 25

(Wax)

Desmodium

Oct 15. Put in lower portion of full-grown leaf for 4m. in water at 82° falling to 79°, upper surface hardly at all wetted.

[in margin:] say average 85°

Put in same portion for 3m at 86° to 84° borders broadly wetted; with additional 1m all except centre was wetted & centre portion became dry only by blowing hard on it.

(Upper half of same full-grown leaf: for 4m in water 86° to 83°, all upper surface wetted except central band close near to mid-rib & a broad border of lower surface wetted.

(About 3/4 of distal of younger leaf for 3m. in water 86° to 84°, borders of upper surface broadly wetted; with additional one minute all wetted except near mid-rib; lower surface well wetted.) (We here clearly see nearly the limit at which the bloom is removed by waving in water. It seems probable that warm rain acting for several hours wd produce same effect at a lower temp.

(p. 9

[diagram] angles which the leave stalk make with the stem

before — 27 3/4

after — 39

Imagine the leaves only at right angles are 39° to the horizon

(p 9

[diagram] after shaking each petiole diverged 11°

Lenny has made diagram.

(i.e directly [illeg] movement à to sleep)

[9v]

[diagram]

(9

Desmodium

Oct 12th. 12° — Temp. 80° 1/2 —the little leaflets were moving as energetically as at night—

NB only blades of young leaves of shake

(They are as rudimentary in structure as in function, for they do not sleep & do not move when branch shaken or syringed; but they retain the incessant slight movement of the large terminal leaf & being smaller are acted on more energetically.— Is there any common point which exerts all 3 leaflets.)

[sketch] I suppose 135°

(with Lenny who made sketch— petioles at about 45° to axis— blade at (90 + 45 =) 135° to axis? petiole, measured on under side — shook them for 30s & the blade was now at 90° to petiole; so that they moved down 45°. In one case L. thought from drawing that petiole had approached axis, a little, as in sleep., but this apparently did not occur with 2 other leaves. We now Know, at least that blade moves when shaken at junction with petiole above the 2 little leaflets.—

As on former occasion the latter leaflets not affected by the shaking. —

(10

Desmodium

Oct 12th Temp high & between about 80°

At about 3° 25' syringed plant 3 or 4 times violently then shook it for 1m & again shook it. The little leaflets, I believe were not affected, but the main ones depended at a greater ∟than 45° to horizon.

This may be, I think, accounted for by the vertical syringing having forced petioles apart.— I measured distance of one pair from base to base of leaf & the distance was considerably increased —(see diagram back)

It is very unlikely that pedicels approach each other, as if so, this wd require greater downward movement of blades to take angle of 45° or any proper inclination — On the contrary divergence of petioles wd add to inclination.— At 4° 15' the leave were all horizontal again. —

[sketch] this measured

(11

Desmodium

Oct 12th 10° 15' — 10° 45' P.m. Temp. 82.

Lenny observed plant asleep. — Leaves as in sketch on opposite side.

The little leaflets were in active movements, sometime insensibly slow, sometime by jerks, as much as he had observed in afternoon, when I saw that they were very active.—

Pedicels, he says, certainly much closer than during day — At top of shoot quite parallel as in fig. 2. — What wonderful movement the big leaflets are capable of— as they are often above a horizontal plane

Shaking plant had no effect— on position of leaflets.

à We think light of candle stimulated little leaflets??

He found one big leaf awake, but it appeared to go to sleep, whilst he was observing it.— The big leaflets by raising of pedicel are all crowded round axis, like pinnules of M. pudica together— Mutually protect each other.

(see Back for sketches)

(12

Desmodium

(Oct 13. Temp. 71°. No movement in little leaflets, when after 1 1/4 hours the temp. had risen to 77 1/2 I saw 2 leaflets move.)

At From 11°, when temp. was 72 to 11°. 15' when temp. was 74° there was no movement in tips of large leaflets. At 11. 20' the tip rose a little above the mark, at 11° 25' it was a little below the mark.

At 12°, 23' had risen to mark — at 12° 26m was still rising — at 12° 50' had risen greatly, & now temp. was 80°.

(At 10°. 45' I pricked with needle upper side of short hairy base of 2 main leaflets (marked with white thread), there was slight depression, but not more perhaps than the handling wd account for, & at 12°. 15' this leaf rose above horizontal line —

[12b]

Desmodium asleep. — Leonard

Oct 12' 10° 15 - 45' P.m

Fig I. leaf on one side of branch

He thinks ∟of petiole correct. [sketch]

leaves at summit of a shoot. Fig [sketch]

(13

Desmodium

Oct 13 the 10 ° 51 '. Pricked upper side of base of main petiole, close to axis, produced no effect — (Marked w. purple thread).

Pricked petioles at base of 2 or 3 leaflet— produced no effect.

10.54 — (Black thread) Pricked middle of petiole — no effect.

It shd be observed that all these prickings were done when temp. from 71 to 72; it is possible that when very hot they wd have produced an some effect. All these pricked leaves at 12° 15 ' when temp was 78 °, had risen above the horizontal.

(14

Oct 13th

before shaking ⨀ ⨀⨀ after shaking — (an upper pair of leaves measured from base of one to base of other

1st pair

Temp. about 74° so not very hot.

(N.B. see what fraction of increase of distance this is; as petioles all of about same length, if the increase of distance is nearly the same as that given at back of p. 11, increases of angle will be nearly the same viz. 11°

2d pair [sketch] ⨀ ⨀⨀ an upper pair of leaflets

3d pair [sketch] ⨀ ⨀⨀ a pair beneath the upper pair.

I shook the plant for 1m, & after a little interval gave another slight shaking.— Almost all the leaves became in a few minutes inclined, from being nearly horizontal to from 35° to 45° beneath the horizontal line.

(15

Desmodium

Oct 13th. 4° 45' dark rainy evening— leaves not so horizontal as during the day.

5° 18' well inclined at about 45° to horizon

(do M. pudica well asleep & many more of leaves of M. marginata.—)

6° Lettington observed & says well asleep, but leave not depending so completely as at 5° Am. next morning.

7° 40' P.m. thoroughly asleep. — I fixed needles opposite points of 3 large leaves, at 7° 40' & looked occasionally at them until 8° 10' & saw no trace of movement— one leaf point indeed changed a little, but I think it was the loose stick. [insertion:] temp. was at — first 81° & rose to 84°, & so ought to have been very active Next morning at 5° a.m. Let. observed them & found 2 of the leaves in exactly the same position with respect to the needles, but the thread was a little changed, as, however the plant was near awaking it might have moved a little from this cause. On the whole I may conclude no spont. movement in large leaves whilst these are asleep. (continued

(16

Desmodium

Oct 13th

With respect to little leaflets, Let -observed them carefully from 5° A.m. to 5° 30', whilst the temp. was 82° & saw no movement whatever; the first leaflet, which he saw move was 6° 55' a.m.; but the plant was then well awake for it began to wake at 5° 10' & was fully awake at 5°. 59', 6° am so that no leaflets were seen to move until an hour after awaking — The little leaflets at 5° A.m were "all sticking vertically upwards. in an inclined position" (i.e. inclined towards the main leaflet) & what what wd be forward in the day); but unfortunately L. did not observe whether all the leaflets held same position or inclination. He feels pretty sure that all the little leaflets were in the same position at 6° as at 5°. It is pretty clear that movement ceases, whether or not it is true sleep.

[16v]

The great heat made movements rapid—

(17

Desmodium

Oct 14. At 9' (about) Temp 92° to 93° I began tracing on the outside glass the movements of a terminal leaflet which almost faced the glass & was somewhat inclined upwards. I took the line from the Apex along the midrib. I also drew transverse lines at intervals shewing the inclination of the lamina. The tracing was finished at 9.35. See the tracing, & the Diagram, shows the actual distance viz horizontally .82 of an inch & vertically 1.2 of an inch in a horizontal & vertical direction over which the apex moved.

The lamina also changed its inclination. & the angle can be seen

All this was effected in 35°m. The course pursued was very irregular. The great heat made movement rapid. We thus see that the big leaf moves, (tho' never by jerks) exactly like the little leaflets: the latter however sweep, I believe, from right to left over nearly 180°, & far down & far above the plane of the terminal leaflet. Their laminæ I believe change their

18

Desmodium

inclination I believe as much as 90° 180 ie quite round. I cd see that their little sub-petioles which are hairy (longer actually & much relatively than that of the big leaflet) become much curved as they leaflets moved. They do not generally spring quite opposite each other, but sometimes they certainly do (due to their being rudimentary?

At 10. 45 (temp. 91 92) I fix gummed a minute pointed bit of paper on the common petiole of the same leaf & this rested on the stipules of the terminal leaflet, so that the sub-petiole of the latter cd move independently of the common petiole. I then traced, as before, during 31m. the movement of the main petiole, & at the same time the movement of the apex of the terminal leaflet.

(This gumming on of the paper retarded the movement)

(19

Desmodium

I shd add that the pointed bit of paper extended beyond the petiole to the middle of the terminal leaflet. See tracings of both. It is clear from this that both move, to a certain extent independently of each other; see the results p. 2 folio diagram. but the leaflet moves much more than the common petiole. These movements strikingly resemble those of tendril-bearing leaves together with their petioles. No doubt then vital movements afford the basis on which all kinds of movements have been developed.—

When speaking of the bending of the transverse direction of lamina, I can allude to case of Cassia, during sleep.—

See further on about very slight of axis, which will complicate & add to the result

[19v]

N. B. the greater length of sub-petioles of little leaflets will account for greater movements, together with less work to do.— Is greater length a remnant of former structure?

(Balfour suggests that wet injures leaves from preventing evaporation & consequently flows of saps & nourishment; but this principle wd apply to all leaves & only a few are specially protected. ∴ I think these must be in some way specially injured by wet.)

(Leaves shake like some Poplars— this may perhaps be one of uses of extremely flexibility of petioles)

(20

Desmodium.

Oct 15. Lett observed plant at 4° 47' A m. Temp only 77°; there was a small moon: could this have been cause. or was plant exhausted on the former occasion by prolonged high. temp.

However this may be L. says the main leaflets were awake, but not quite horizontal & the small leaflets were moving!!

Pulvinus

Structure of Leaf — (sub-petiole (is this right term?) of little leaflets either opposite or not so.— it is actually longer & much much thinner than the sub-petiole (ie part between stipules & base of leaflet) of the main leaflet.— This may partly account for greater movement. — is thickly covered with short thick white hair, whilst main petiole thinly covered with long thin hairs. — The sub-petiole of main leaflet, is rather thicker than the main – petiole.

(21

Desmodium

Oct 15'

I here see little leaflet bend inwards to opposite side beyond medial line — they will face in almost any direction— having such power of rotation — I have seen back of one facing to end of terminal leaflet. — I drove needle through sub-petioles, but caused no movement.— When I rubbed inner side with needle I thought became more active, but as the leaflets were very active hard to decide. — I rubbed upper side of sub-pet of main leaflet no effect— I drove needle through axis between intersection of 2 opposite main petioles no clear effect. —

Oct 15 I placed glass Plate over plant & observed end of 4 shoots to see if they moved.

stem

I observed at intervals from 9° to 2°. 15, & traced the line.

(Temp. high 89° to 92°)— One axis moved a little in 10' & 2 others in 20', but as this was early might have been towards light; but I think not for one moved to right from exactly opposed to course of sun.— The movement was very small & always

irregular, but certain: (1) was a very small & irregular

(22

Desmodium

oval or ellipse in line from nearly E. to W. — (2) a decided oval, with longer axis N. & S. (3) altogether an irregular movement. (4) of which the record began at 10° & was continued to 2° 15' formed a decided circle, but irregular.— These movement though very slight wd add slightly to movement of whole leaves & are just like what I have observed in the axes of many climbing plants & as Fritz Muller has described in Alisma. (see my paper.)

Oct 15. Put a rather young leaf in water at 70° & warmed it up to 145°; till 101°, no movement; but above this temp. the little leaflets rose vertically up with a slight deviation from the original horizontal angle {The leaf was killed & browned by the 145° as I saw next morning

I then took a fine large full-grown leaf & placed it in water at 95°, & slowly raised it slowly to 103°. During

(23

Desmodium

all this time i.e for 40m the little leaflets were in constant & rather quick movement. The tips described little circles or semi-circles.

I never saw but one jerking movement; at other times the m. was nearly steady, yet with some stationary pauses. This steadiness perhaps due to resistance of water. All this excellent proof of the effect of high temperature. If the leaflets moved as quickly for the whole 40 m, as whilst I watched them they must have made several dozen circlets. At By 12, 50m ie, after the 40' the water had fallen to 70°: but I warmed it to 100° & the leaflets again began to move. Afterwards I raised temp. to 122°; but next mg. leaflets appeared green & not killed. {It was pretty clear that it is sub-pet of leaflets which cause movement by spontaneous bending.

Early in experiment whilst the water was hot warm I cut off the main leaflet pretty close to the the sub-petiole, so that there was little

24

Desmodium

resistance to movement, & I cd easily have detected any spont. movement in the great cut-off leaflet. This absence of movement is singular & shews how much less easily excited it is than the leaflets. After water had fallen to 70° the hairy short sub-pet. of main leaflet became greatly curved downwards so that the remnant of the leaf held the position as if profoundly asleep.

I then immediately warmed water to 100°, & by accident it rose to 122°. After 30m from time when water was 70° the sub-pet. had become straight again, & the cut-off leaf was awake. In 1h time the water had fallen to 70°, but the cut-off leaf had not again gone to sleep; it was however only somewhat torpid, for next mg. after remaining in the cold water, it was again asleep.

(25

Desmodium Water on Leaves

Oct. 16th Temp. from 11° to 8° 30' P.m. from 86° - 89°. at night probably about 80° —(at 6° Am on 17th 76°, but rose by 7° 20' to 81°. at. 9° 15 87°, at 10°. 20' to 92 & at 11° to 101°

I fastened 3 nice leaves with needles on a bit board horizontally & put on large drops of distilled water.

(Removed water at 11° on 17th)

(Leaf 1) Black wool washed 2 or 3 times with ether & very soft camel brush middle surface on further side, & then washed surface with water & put on purest water. The wax was thoroughly removed. N.B. This leaf was thickly spotted with minute yellow spots, as other I suppose from former syringing

(Nov 4th I have been looking, most carefully at spiral vessels, & the utmost which I had better say is that the point & appearance made it probable that the one sometimes disappear by confluence.)

(Oct 25th I have just observed an interesting fact in a leaf which had no lateral leaflet; & in this case a ridge ran up each upper angle to the bases of the stipules of main leaflet, & one of these ended in a little free point, evidently the rudiment of one leaflet coherent with the main petiole.

It looked at first look a minute double stipule— I have no doubt my view is right.) th ie that the Cot. leaflet disappear by becoming confluent.

[in margin:] Why more vestiges did not occur suspended this much—

[25v]

Oct 18' 10° 30' Brown marks on midribs of the 2 leaves as yesterday— Water adheres to the cleaned surfaces of all 3 leaves—

Oct 20th same state.

(Oct 21' 8°-30' — water still adhere to surface of all 3 leaves.— Red wool midrib & side veins & cellular tissues on each side for narrow zone brown.— Black wool do, but not so extensive. All round the space on which the water rested a very narrow, & slight yet distinct brown line.— I think this must be due to water & not to ether; if any, as is possible is due to ether, for I have not seen other plants injured by a slight wash with ether. Yellow w. parts of space on which water rested with faintest trace of brown & perhaps the mid-rib also.)

Oct 22d (12° m.) same state; I think yellow wool a shade brown— surface of all 3 leaves easily wetted.

Oct 23d (12°) I certainly think a trace of yellow on midrib of yellow w.

— 25 8° all 6 leaves easily wetted

— 28. 12' all 5 (1 cut off) do. do. do.

— 31 do do. do perfectly adheres.

(26

Desmodium

Leaf 2. (red wool) washed spot with brush & ether & put on water towards base on both side of mid-rib.

Wax thoroughly removed. Also put on drop at distal end on normal surface— beautifully spherical & moved about like quicksilver & remained so for 24°.

not ethers

Leaf 3. (yellow w.) (nearest leaf) I put on large drop of water at 95°, & did this thrice leaving it each time for about 1/2m but this did seem to dissolve wax— I then gently brushed the surface with very soft camel brush 4 or 5 times, each time dipping brush in the water at 95°; & thus I removed most of the wax: yet the water, (which was put on near middle of leaf, rather towards base & on near side) did not adhere so closely to surface as in the 2 former ether leaves— These facts show, as I have observed in trying effect of hot warm water on leaves that agitation is very necessary to remove wax but hard warm rain wd give this. Dew wd not do so.— (over

[26v]

26 con. Also put drop at distal end on normal surface.

Oct 17th 11° the Black wool leaf had midrib when water rested freckled with brown.— Red wool has (i.e. ether) has mid-rib when water was & margin on each side brown.— The drops placed on leaf with secretion quite spherical & when sucked off surface not wetted, or trace of circle in one case but no mark left.

Evaporated to few drops of water in watch-glasses.

That from 2 leaves (yellow & red) (some snuff snuff fell on black) move mouth relatively to that from same parent water than can, I think be accounted for by dust falling as can during 24°. & the

Oct 18° I have now examined the mouth with weak compound m. & I can see in every ring in yell yellow & red wool glasses rings of very pale yellowish green matter, which crumble after drying over spirit lam[p]. There is no trace of such matter in the pure water evaporated. It is certain that water has absorbed a considerable quantity of matter from the leaves.— The only doubt is whether the ether may have injured the texture of leaves, hardly credible.

It is all moreover about delinquencies, it was my breath & so no doubt in all former cases.—

(26A

Desmodium

Oct 21st washed with camel-hair repeatedly, for this was necessary but very gently with water (97° - 94°) 3 leaves

(Temp of House & then 85°) Dried them carefully with blotting paper, & then put on at 11° large drop of pure water

Fastened the leaves on board as before.

White & Black & white red oldish leaves — (& White & red yellow (?) youngish leaf— looks tender.)— At 4° 10' Tem 77°, at 8° 40° 91°.—

Oct 22d 7° 20' temp 71°', at 9° 25' 81'° at 12° 15' 84°. — I then removed the fluid into 3 watch-glasses.— The leaves were wetted when I sucked off the water.

I rather think Red & White w. leaf & yellow are very little browned. — Day cloudy & rainy

[26v]

Oct 23d (12° midday) Yellow w. midrib with trace of brown & tissue close on each side pale. There are brown freckles here & there on each side where water rested. Red do. Black, more doubtful case.

(Oct 25th 8° there certainly is a trace of brown on midribs of Red & Black & W. leaves. — I examined carefully the young leaf yellow wool; the space on which water rested is rather slightly discoloured, & this is chiefly owing to mid-rib & lateral veins being slightly browned & to little patches or freckles quite brown, as if burnt. I have looked at these under Compound M. & the veins themselves see are brown, & much of the chlorophyll in the intermediate tissues has disappeared or been destroyed, & these portions are much more transparent. The mid-rib which has a light brown appearance owes it to minute freckles of brown, & the cellular tissues close on each side slightly discoloured. I now am inclined to believe that ether only removed wax more completely. I have now examined all 6 leaves & the brown of mid-rib extends right through to midrib on the under side, & at corresponding place where water lay.— This injury to the vascular bundles is curious, & it accords with the burnt freckles, where the vessels seemed browned all through.

(26 (B

Desmodium

(2) Red & white fine full-grown leaf

(9) watch-glass Black & white do leaf

(8) yellow & white rather young leaf, but full size.

(In all 3 glasses more sediment after evaporation than can be accounted for, I think, by dust during the 25°— on the successive rims. I cd see under weak power in compound, excessively-pale yellowish-green matter. — so this does not require ether to be extracted, Must be due to water alone & this must be injurious —at least in some degree.— This matter seem homogeneous when dry like dry gum or mucilage.

[26bv]

Oct 31' 8° A.m. I have been looking at the 2 leaves left i.e Black & red wool & there can be no doubt that they are considerably injured— several of the fine lateral veins are brown, & little freckles of decayed tissue where water rested.

Nov. 4th 8° I have been looking (& cut off) under compound at Red & White wool— well wetted when dipped it is clear that wax not secreted again. By scraping surface I can see that the browning of midrib is due on upper & lower surface to the less browning of the tissue which covers it & not to the browning of the fibro-spiral vessels themselves. There are now a vast number of spots, where the water lay, which are very pale green, & more transparent than other parts graduating into pale brown, owing to the decay of the chlorophyll & few other spots completely brown & decayed. It is odd surface of lower surface being of midrib being brown must be due to pressing very greatly for 24° on damp surface. No I have changed again, though the browning of midrib is chiefly due to surface it is not so exclusively; & the minute veins are likewise brown; but surface chiefly affected.

(27

Desmodium Effects of Cold

Oct 18th — put leaf out of Case which had been very hot about 90° into (at 8°h 48'm) into water at 46°: it had before petiole in same line with midrib of terminal leaflet: in 25m the petiole was curved a little inwards: in 1° 27m it was bent inwards at ∠'.— A second leaf let (with little leaflet) had petiole nearly but not quite straight with midrib:) & after 1° 27' had it bent inwards, & it had passed through about 70°. I then put both leaves for some minute into water about 60°, & soon (at 10° 50') put them into water at 79° & kept them for some time, till it fell to 75°; at 11° 12' the petiole of one leaf straightened itself considerably, but not that of the other: I then again & again raised water to 90° — At 11° 40' the petioles of both leaves, had bent inwards towards midrib, as if going asleep, in a quite abnormal manner !!!; & the petiole of one became bent laterally towards margin of leaf.— They remained at these angles & never straightened again till next morning. The above facts (with former facts) are enough to show that cold water causes sub-petioles to bend as if going asleep. It is the sub-petiole which bends.—

(28

Desmodium Cold water

In order to test cold water on other sleeping plants I put at 9° 8' a leaf of Mimosa marginata from hot Case into water at same temp viz 46°— & weighted it with lead at base so that it stood upright. — At 10° 30' (ie after 1h 22' all the leaflet were quite close, & the water had risen to temp 53°. At 10° 50' put it first into water at about 60°, & soon raised this to 79° letting it fall to 75°, at 11° 12' (ie in 22m) leaf much opened; at 12° 50' about half opened at 12° 50' completely open; & so it remained all day. At night it became in the water very slightly closed, but opened completely the following day morning.—)

Sensitivity (Keep)

(29

Desmodium Cold Water

Oxalis acetosella put kept in Cool greenhouse put into same water at temp 50°', at 8°h 57'm; at 9° i.e. in 3m partly closed, & fairly moderately well closed in 15' but not nearly so close as when naturally asleep.— & so it continued whilst was kept for hour or 2 in cold water.—

Put into warm water as before but did not open; on the next morning, however opened perfectly though still in water.—

The oxalis with copper foliage growing in hot-house & Ox. sensitiva from hot Case not in the least affected by the cold water.

The greenhouse Cassia from cool hothouse put left in the cold water for 3h 10', had its leaflets a little beveled, which is the first movement in going to sleep.

Cytisus [few words illeg]— Trifolium repens in greenhouse— Passiflora gracilis (which depresses leaves when asleep) Tobacco, (growing at out of doors, which slightly raises leaves) were none of them in the least affected.) Sensitivity

See Back

[29v]

Oct 21st I could generally place apex of leaflet leaves of Mimosa pudica very gently so as to rest on water of the same temp (viz about 75°), in which the plant was living; but when I similarly placed them on water, of temp. 43° - 44° the leaflets always closed, but not very closely or quickly — agrees with M. marginata.

(30

Desmodium

Oct 18. Veins of little leaflets have different appearance from greater simplicity & more irregularity— but essentially the same as in big leaflet.

Oct 18 a quite young plant., 9 inches high, when shaken, depressed its leaves.

Oct 19th 10° 30' temp of House 77 1/2° the little leaflets on this plant were moving. I repeatedly rubbed their sub-petioles, & almost every time they moved in a few seconds, though still before: I can hardly have any doubt. When I rubbed outside of petiole, they moved, but apparently not towards rubbed side but downwards.— I saw one pass under main leaflet, so that its midrib was parallel to that of latter — I do not doubt that they sweep through 180° in the same plane as main leaflet. they move in short in every conceivable direction.

(31

Desmodium

Oct 19th. Temp of House only 72°. at 12° syringed plant several time with water at 69° & then shook it strongly for some minutes— Leaves became depressed, but more slowly than usual, viz it took 4 or 5 3 or 4 m before fully depressed I finished shaking & brushing at about 12° 20'; at 12° 40' not much risen. but the leaves owing to low temp. were not quite horizontal until at 1°.— These plant in the 2 states were photographed by Lenny. Before shaking the young & upper leaves were horizontal & some several above horizontal.

The old & lower leaves always depend.—

I measured carefully inclination of 12 leaves, & (see Back) these varied from 35° to 57° to the perpendicular. ie. 55° to 33° with the horizon.

But most were nearer to the mean which was 48° to the perpendicular & 42° to the horizon.

On a former occasion, when plant had been kept very hot & it was syringed & shaken, the depression of the leaves was greater.

(32

Desmodium

It follows from above that the leaves moved downwards through an angle of 42° on an average.— It wd I shd think be advantageous as a protection against rain, if they were to assume the position of sleep, ie. perpendicularly down, but then this wd be at the expense of greater exertion & loss of time in recovering their horizontal position.

Seeing how well the leaves are protected by wax, so that water runs off them like quicksilver, it is surprising that they become depressed. No doubt the movement wd be superfluous in a cold climate, but long-continued warm rain wd remove much of the wax, & then it wd be highly important that all the water shd run off as quickly as possible, before the temperature rose much.

(p. 31

 

Before

After.

 

Petioles

Leaves

Petioles

Leaves

1

32

62

39

133

2

72

108

77

160

3

27

60

54

152

 

with axis above

with do

 

 

The above are the angles which the axis of the blade of leaves and the petioles make with the stalk (as seen on a photograph). before and after shaking. No 3 gives the best absolute angles because in this case the plane containing the leaves and stalks was parallel to the plane of the photograph.

Thus it will be seen that the petioles have moved down

[31v]

[diagram]

Leonard — Results of comparing photos of same plant in the same position— Several of the leaves before being shaken were above the horizontal & these have moved more.

[31b]

7° 5° 24° in the three cases and the leaves have moved down on the petioles 64° 47° 55°; that is these three shew the amount of movement between the leaf and the petiole.

So that the blade of the leaf, if the movement of the petiole & the blade be combined, has been 71°

No. 2 has similarly moved 52°

& No 3 has — 79°

[31bv]

The movement of the petiole from the axis, (now shown in 7 cases), is very important for me, as it is a directly opposite movement to sleep & therefor no doubt is a specialised one.

Oct 26 / 73 /

[unnumbered]

[data not transcribed]

to the Horizon.

[unnumbered verso]

to blush in the presence of strangers. The consci

of anything peculiar or even new in the dress

slight blemish on the person and more especial

the fact,─ points which are likely to attract

(33

Desmodium

Oct. 20' 11° House 76° 1/2 ─ had been below 70° ─ The leaflets were not moving.─ I rubbed several ─ put in sulphuric a. (2 per cent) & Chloroform, but no movement was excited. Nor when out on upper & under side of sub-petiole of main leaflet.─ at 4°15 case 78° & [illeg] some leaflets moving.

I looked at many leaves & when 1 or both leaflets absent, I could see no rudiment or vestige of them. —It is odd this sudden disappearance. I examined carefully 2 very young leaves, & the leaflets certainly have never been developed & then fallen off. But by tracing the vascular bundle, which as not symmetrical above the insertion of a simple leaflet, when one close is present, it is pretty clear that their sudden disappearance, at this present rudimentary stage, is due to their complete fusion of with the main petiole, as they can be traced up to the stipules of the main leaflet.—

(34

Desmodium

Oct 21st— 7° 30 Am. Temp. of case only 71° & leaflets in movement!! I suppose owing to increase of warmth for case had been down to 67° or 68°.

Oct 21. 8° 31— 9° — leaflets not very active & plant not been healthy. I cannot make out that rubbing the sub-petioles excites movement

Oct 27. When the leaflets are not moving & have not moved for some time, they diverge at about ∠' from petiole, but are directed a little forwards or towards main petiole. At 4° P.m old & young plant were brought to my Bedroom, the main leaflets from being shaken hang down & never rose again before night came on. The Temp of room during night varied from 60° to 56° & the petioles of neither plant rose at first towards axis on the contrary became more depressed as if they had been shaken.— But at 10° 5 the petiole of young plant had risen a little & at 2 A.m. (on morning of the 28th) the petioles of the young plant had certainly risen considerably; those

(35

Desmodium

of the old plant, which was in feeble state, never rose during whole night. At 6° 45' A.m. the main leaflets of young plant began to rise & awake. — The little leaflets of both plant never attained their position, from what they were on previous afternoon; & as the old terminal leaflet of young plant hung vertically down & went to sleep, & awoke in morning I think I may safely conclude that the little ones Leaflets do not change their position at night &.', & do not go to sleep)

(Oct 31. Spont. movements — if we suppose from widely extended & primordial nature, (like & of no more use than [illeg] movements) that these movements have been fully retained by D. whilst others & more recent lost, we can perhaps understand how with much reduced petiole in thickness & much less weight to carry, how the movements are so conspicuous.)

(36

Desmodium

Nov. 1. 7.40 A.m. put tuft of 5 leaves in water at 52° rising to 54° for during 1 hr & the angles of the petioles were not affected.

The leaves were left in the water which was a little warmed, & then allowed to cool to about 60°, until 11° 20', & I then found the petioles still unaffected; but the large terminal leaflets hung vertically down owing to the cold.

The tuft after staying so long in the cold water was then put in water at 92° which was quickly raised to 106°; the large terminal leaflets rose considerably; but the petioles were still in no way affected.

(I cut off a leaf with 2 little leaflets & heated the water to 99°, when I first saw a very slight movement. The leaf was left in the water for 3/4 hr. It was then quickly raised the to 99° & rather when I again first saw

37

Desmodium

slight movement. The water was now rather quickly raised to 105°. Now the leaflets began to move rapidly. I made a tracing of the movements of the apex of one on a glass plate above with a fixed sharp point as a guide lying beneath the leaflet: it made 2 small one irregular circles in 1 1/2 m., & about 2 1/2 circles in 5 m: the water had now fallen to 88°. I raised the temp. to 95° & made a fresh tracing; again it made a small irregular oval figure in 1 1/2m & about 2 ovals & a half in 3m 30'. Hence formerly during the 40m it the apex probably made about 13 circles. The movements were very difficult to trace on account of the small

38

Desmodium

size, irregularity & quickness of the movement. The figures are exactly like those, (but on a very small scale) which I have often traced of tips of tendrils, leaf-climbers & the axes of such plants whether or not they are climbers. There was no trace of any jerking movement.

(I also observed the tip of the main true leaflet in the hot water, but cd make out no movement except that of rising previously alluded to—

(Looked at petiole with only one lateral leaflet, but cd see no signs of confluence on opposite side—)

Mimosa pudica is the only plant I can remember which move as quickly as these little petioles — leaflets —

(40

Desmodium

Dec. 5' — Put 1 old— 1 moderate old & 1 qui young & fresh in water at 42, having first measured ∟ of blade & petiole— & left for 5m to 1/4 h.

In the young leaf for 5m & 1 of old no change of angle— in the 3d leaf it straightened itself— i.e. blade blade & petiole formed a nearly straight line.

Dec. 12th — 2° 30' — fairly warm day — little leaflets of young plant in steady slow movement— I tried repeatedly rubbing sub-petioles with needle & cd not decide whether it caused movement— when not moving, as I formerly ascertained does not cause movement.—

[40v]

Lettington looked again in winter & saw the main petioles were not depressed

p 25 Effects of Water on Leaves.

to 26 B


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

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