RECORD: Darwin, C. R. 1872.09.25-.10.30. [Drosera continued]. CUL-DAR54.143-176. Edited by John van Wyhe (Darwin Online, http://darwin-online.org.uk/)

REVISION HISTORY: Transcribed by Christine Chua and edited by John van Wyhe 5.2022. 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).


(56

Down Sept 25 I could perceive no difference in state of chitin of Geotropes from being clasped for nearly 6 days by Drosera & corresponding joint. Kept damp in blotting paper. The muscle apparently more disintegrated in Drosera specimen

Sept 26 (Yellow Stick) 8º 30' 1 gr to 1 oz extract of Hyosciamus Henbane on 7 leaves

(8º 31 on 2 leaves (with yellow sticks red Heads) put on large drop & at 9º 2'- 6' put on atoms of meat 9º 17' some have acted well on one leaf

(see Back) (a)

10º 30' no effect — (5º 10' some little inflection of the sub-marginals

(Sept 27 8º 20'— in 3 of leaves sub-marginals inflected, but I think solely result of drying & viscidity of the fluid: washed them)

(Sept 28' 8º A.m. no inflection, those which had been sub inflected there enough viscosity now almost expanded.)

Sept 26 8º 55' (Red & Black sticks) old [illeg] quite move in 48º atom [illeg] caused [movement] sol. of Belladonna 1 gr to 1 oz. on 6 leaves (9º 20' ie 25 some inflection) (10º 25' do)

(5º 10' 8º 15' a great deal of inflection in the sub-marginals; & one leaf still much inflected.)

Sept 27 8º 15. do the sub-marginals in the leaves fully reexpanded; the solid leaf which was closed only partially reexpanded, but quite healthy; but to a [illeg] & [few words illeg]

[56v]

(a) On the other leaf, I find the circumferential tent. which were not immersed in the Hyosciamus quite sensitive in a few minutes - whereas those on disc still immersed in the fluid did not move for 2º, but now one or two only have begun to move, so I think apt to make torpid.

12º 15' I think 3º 45' a little movement 1º 45; 4º 45' the short sub-central tentacles with meat have now all curved - Certainly immersion for 1º or so in Hyosciamus, make tentacles torpid.

 3º 15' 4º 15'

(57

Sept 27th 1872 11º 48' Yellow & red sticks 6 leaves with Phosphate of Potash 1 gr to 1 oz

5º. P.m. no more inflection than may be accounted for by drying of fluid viscid from glands—

Sept 28 8º Am. no true inflection.

(Sept 29th 8º 15 no effect whatever - leaves quite fresh.) quite Healthy

(Sept 30th 8º 15' do do] V

Sept 27 11. 57' Blue stick 6 leaves chloride of Potassium 1 gr to 1 oz

5º P.m hardly any inflection, & can be explained by drying of viscid fluid. V

Sept 28 8 A m no inflection.

Sept 29  8º 15 do - quite Healthy

— 30' do do do.

Chloride Carbonate Nitrate Sulphate Phosphate

[57v]

Raw meat acts quickly & energetically on the contents when the viscid matter removed as far as possible by stripe of blotting paper.

(58

Tentacles.

1872 Back

I cannot observe that rubbing with needle or with minute drop of alcohol, acetic acid, nitrous ether on the concave bending point, causes any movement: the gland must be excited.

The little hemispherical projection projects on all sides of tentacles: function not absorption — The ordinary point of bending is just above where purple fluid bleeds into the basal part, which is green with chlorophyll. In the longer tents', the bending extends rather higher up there than in the shorter ones; but a long distal portion always remain quite straight. tentacles are flattened; this is the section in an oval, with the upper side flat & the lower or outer convex — In the extreme marginal tentacles the flattened form of the tentacles follows from their forming prolongation of the leaf. the tentacle is formed of many cells, much elongated in the line of the tentacle, with spiral vessel in centre. I cannot decide whether it is exosmose with elasticity from cells on the upper side, or their true contraction, in accordance with the belief accepted in other case by the best observer Cohn. & Wyman.

(59

Sept 28' 8º 30' Yellow sticks 3 leaves Phosphate of ammonia 1 gr. to 1 oz. (Large drops)

8º 45' great inflection in 2 leaves: moderate in the 3d.

9º 5' astonishing inflection in all 3 leaves.

10º 10' leaf itself inflected 10º 45 2 leaves do

Sept 29th 8º 48' leaf in little glass with 10 Min of Phosphate of Ammonia. 1 gr. to 24 oz.

8º 57' ie in 9' plenty of inflection. 9º 11 immense inflection all except a few

10º 25 — all tents. except 25 inflected

11. 25 all but 7 inflected, on footstalk 1 or 2 injured

1º — all except 4.

2º 30' Everyone curved but 4 or 5 not closely

 In sport for counting - Pin

8º 55' 2 leaves with 20 minims of do

9º 11' 1 leaf with a good deal of inflection. ie in 16'

2º 30; 1 of the 2 leaves with all except 10 tents.—

5º — all except 15 inflected.

(Sept 30th 8º Am white thread in about same state)

The second leaf at least has all sub-marginal in a circle of Heads, but none of Marginals which seem too old.

Sept 8º All central & sub-marginals closely inflected)

— Strongly pronounced segregation whole length of tentacles in both these leaves.

(60

(Abstracted)

Down Sept 29th 9º 16'. small glass red mark

1 leaf in 10 minims of Phosphate of Ammonia 1 gr to 30 oz of water.

9º 38' [-] 16 [=] 22 some inflection (ie in 22)

10 30 a good deal of inflect: most of tents. upright.

1º great inflection

2º 30' all except 21.

5º all inflected, except 1, but 20 not closely inflected.

Sept 30th 8º Am outermost tents reexpanded. - all others still inflected

Sept 30' 8º 36 Jagged glass. 10 minim of Ph. of Am. 1 gr to 30 oz.

9º 18' only 3 not somewhat inflected - almost all other closely inflected

10º 30' about 10 not closely inflected - & 3 not at all inflected.

1º do do do —(safer to say 5 not inflected) 24 [+] 18 [=] 42

Sep 30 8º 44' do do Red. glass 1 gr to 30 oz

9º 20' only 14 not inflected - almost all others closely inflected

10º 30' - do not all inflected.

1º - 13 not at all inflected - do. do. do.

Black Thread

16 [-] 10 [=] 6

(60A

(Abstracted)

Oct I 10º 50' Jagged glass Phosphate of ammonia 1 gr to 40 oz— 10 minim

11º 55' almost all inflected - (ie 1º 5)

12º 50 all except about 7 [crossed]

12º 55' no about 25-30 not inflected seem old

Oct 1. 10º 55' Red glass same as last - repeated.

11º 50 all inflected except 3 at 55')

12 55' do do do

white wool

Oct 1. 11º 8' Blue glass - 1 gr to 36 oz 10 minim

11º 53' almost all inflected

12 55. no about 28 not inflected.

Oct 1. 11. 15' Yellow glass 1 gr to 36 oz 10 minims

11º 53 almost all inflected

12 50 all except about 7.

2º 35' only 4 not at least partially inflected.

Black wool

53 [-] 15 [=] 38

/over

[60Av]

Sol 1. put on of minute drop (ie 60th minim) of Phosphate of ammonia. 1 gr to 2 oz. to 3 glands, & all 3 moved a little. This is the weakest which has ever acted, whilst the drop held only for a few seconds to the glands.

With reference to all the experiments of weak sols of Ph. of ammonia & Nitrate, when leaf immersed, we must remember that when meat is put in centre, generally takes hour before marginal contract, & contraction spreads slowly outwards, here we have comparatively sudden contraction of all the tentacles. the sole doubt is whether the few which lag behind, & which are always extended are not aided by the excitement of all the others & more vigorous ones.

(60B

(Abstracted)

Oct 2d Down Phosphate of ammonia 1 gr to 50 oz 10 minims.

Red glass 9º 26' - (9º 35' great inflection)

10º 20' all but 3 strongly inflected - leaf itself inflected.

Jagged glass 9º 28' (9º 35' much great inflection)

10 9º 25 all but 4 inflected & 3 sub-inflected.

[in margin:] now bits put in wine glass

Oct 2d. 10º 30' jagged glass. 1 gr to 60 oz 10 minim Ph. of ammon.

10º 47 immense inflection, all but 4 or 5 not quite close

12º 5 all but 3 inflected, & 7 not very closely inflected. Leaf itself much inflected.

(This leaf had 235 tent less 3 = 232 ∴ each tent. got 1/668,160 of gr of Ph. of am

The purple fluid was segregated down nearly whole length of the longer tentacles.

Oct 2' 10' 45. Watch-glass 1 gr to 80 oz 20 minim. Ph. of 2 leaves 1 young & colourless, 1 red & older, both good.

(This red leaf had 177 tentacles - less 1 = 176 ∴ (George) each tentacle got 1/675,540 of gr of Ph. of Amm.)

Red leaf 10' 48' a good deal of inflection (10 52' all except 2, but 15 not closely inflected) (12º 5 every single one closely inflected except 1)

White leaf tender, opened in green House (10º 53 now inflecting considerably) (10º 57' [-] 45 [=] 12 wonderfully inflected)

(12º 5 20-40 not inflected) (3º all but 17 inflected)

(In the red leaf, the segregation had travelled down several times the length of glands.)

(60C

Oct 2' 11º 16' 2 leaves in watch-glass - red pin

20 minim of Phosphate of ammon 1 gr to 100 oz.

11º 26' both leaves considerably inflected 50 [-] 16 [=] 34

12º Redder leaf all inflected except 20 but (12º 50' all contracted except 11)

Other leaf all ― except about 29.

5º P.m. all inflected except 12 tentacles

second leaf --- 17 tentacles.

(see over)

Oct 3 8' A m all but 5 well inflected & leaf itself, subinflected

second leaf, all but 17 inflected & then the extreme outside tentacles, no doubt old - all the other tentacles most inflected.

Oct 3'. 8º 25 2 leaves (old Plate Red Sticks.)

with moderate drop on disc alone of Phosphate of ammonia 1 gr to 20 oz to see if absorption in centre quickly causes contraction of outer tentacles.

9º 20 no effect on smaller & pale leaf (10º 20' no) (3º 30' do)

Drops of Weak

9º 20' slight effect on larger leaf, but the drop spread a little at distal end.

10 20' do. 1º 30 sub-marginals slightly sub inflected all round. (2º 30 do)

[60Cv]

I do not think any of these experiments with Phosphate of Ammonia are trustworthy - Perhaps water not pure; but I pushed them too much about in immersing them - See p 60E about plain water

Always try plain leaves at the same time - I must try some of the weakest with Nitrate of Ammon again Try deep watch glasses.

[in margin:] I think watch-glasses wd be best, first turning leaf face downwards but this must be done with water specimen also

Try 2 leaves with 20 minim in watch glass & submerge by holding by forcep

60D

Oct 3d. 8º 40' small leaf jagged glass in 10 minims of Phosph. of ammonia. 1 gr to 200 oz.

8º 50' - strong inflection - all except 11 marginals.

9º 4' all inflected except 6 & 5 sub-inflected.

10 20 do do. do

Oct 3' 8º 48' Red glass. 10 M. of Ph. of amm 1 gr to 100 oz

― 9º 1 all inflected except 2 & 3 sub-inflected.

― 10 20' do do do

Oct 3d 9º 15 Yellow glass 10 m. of Ph. of amm 1 gr. to 400 oz.

― 9º 24' some inflection

10 24 all inflected except 19 & some of those sub-inflected. 24 [-] 15 [=] 9.

[60Dv]

N. B. The leaves which have been kept for a [words excised] in the very weak sol. of Ph. of ammonia are still inflected & do not differ in appearance from these water specimens.

(60E

1872 Oct 3d. Distilled water in same little saucers & treated in every way as last. specimens, 10 minims

Red glass. 10º 32' water from Bottle with pipe

5º P.m. none inflected: Oct 4' 8 A m. still no inflection

Down 1872

Blue glass 10º 35' Water from great bottle

10. 45' Plenty of inflection. 5º P.m all except a few long-Headed marginal inflected.

Oct 4 8º Am. still inflected:

Down 1872

10º 48' 2 leaves with 20 minim of Distilled water from Big Bottle

One of these rather young & tender considerably inflected at 11º - at 5º P m all the tent., except the long marginals, inflected

Oct 4 8º A m. still inflected & contents of cells segregated

The other & larger leaf. with many tents irregularly sub-inflected

I strongly suspect it was the immersing & pushing these in centre under the water which caused so much inflection.

(These cases may be neglected) (4 leaves) 3 strong almost

60F

1872 Down Oct 4th Put 8 leaves at 8º 30'-40', not very good ones, in distilled water in wine=glass (so water cd not have got viscid from secretion, which is a mistake) not a mistake if I use same quantity for salt & fluid

After interval of 35' some long-Head & a few round-Heads inflected How many

― of 1º 40' 4 of the leaves with a few round-Heads inflected & seven of the leaves with a very few long-Heads inflected.

After interval of 8º 25' - the round-Headed glands now very slightly inflected in the 4 leaves, & only those near pedicel of leaf.— the long-Heads in all the 7 leaves now rexpanding, so that they all stand vertical instead of closely inflected

After interval of 23º 30' - All the long-Heads perfectly reexpanded; but 2 of the leaves still have a few of the round-heads, very slightly sub-inflected.

I conclude that the inflection in my recent experiments with the ? very weak solution of Phosphate of ammonia was caused by my pushing the glands & whole leaf under water.

(61

1872 Bundle

Down. Sept 28 at 6º 50 P.m put on lots of atoms of glass & cork on glands on 3 leaves & atoms of dry meat on another leaf - Did this by light of single tallow candle, taking 3 or 4 minutes, & then covered up in absolute darkness. Next morning after 13º 15' all tentacles without exception has reached centre! So light no importance.— Darkness

When tentacles all closely inflected the glands form a perfect circle, owing to graduated length of tentacles, round an open circular space, studded with some of the small, short, upright glands, & which wd just hold a small fly, so that this wd be bathed all round & beneath. The Heads form a dark circle, fit very neatly. This is very sure before the leaf itself becomes inflected, for this action pushes glands to actual centre

This is when action on gland & this show straight inflection the normal movement

Tentacles

[61v]

Begin with centre

(62

Tentacles

Down Sept 29 When atom of meat put on one of longer tentacles it bends to near centre, but does not cause any other tentacle to bend. The influence not due to absorption, for too slow & touch suffices.

It does not appear probably that the influence passes down & across a multitude of cells, but along the matter surrounding the central spiral vessels. After the atom has reached the centre, if very small it send back influence to a few adjoining tentacles & these bend - if large several bend.

(If small but of meat does not reach, or is put on one side of middle of leaf, all the tentacles on same side clasp it after a time; but the influence does not extend across the leaf.

If put on centre the influence does extend. I can understand this, assuming that influence run along spiral vessels (How about Dionæa?) by supposing that influence runs one way at the 2 points of inosculation.

(63

Tentacles

Down Sept 29th - In footstalk, a little below the lowest tentacle on the pedicle. 3 bundles of vascular tissue diverge from a common mass. the middle one is the thickest, & run up middle of leaf, when it bifurcates, & both branches again bifurcate, again bifurcate & at last send branches to each margin & surface tentacle, & sub-branch to the other tentacles, in upper portion of leaf. Just where pedicel expands into disc of leaf, the central bundle sends out a branch on each side, which supplies the central tentacle of lower half of leaf. We now come to the main lateral bundles. These bifurcate a little way up & supply the sides of leaf of lower half.— But the inner main branch is remarkable by one of its upper & inner bifurcation, inosculating with the lower branches of the 3d bifurcation of the great central vessel. This inosculation I imagine is the line by which when meat put on exact centre, influence is sent to the lateral tentacles of lower half; & I imagine the influence will not flow upwards or outwards, otherwise meat on margin wd cause inflection of tent on opposite side of leaf

[in margin:] In different leaves, & even on opposite side of same leaf, good deal of variability in extent to some of vascular bundles

[63v]

Considering the [illeg] have [illeg] slight must the touch be

[63v2]

In D. Anglica easy to see at bifurcation that both vessel merely unite & run down joined

Influence travels centripetally & centrifugally — & radiates up & down centre of leaf. quicker this to extend laterally Travels quickly to gland to bending point.

[annotated sketch]

All on one side of leaf - contract & no contraction with opposite side of vessel lies near

(See George's Drawing)

(Heads of tentacles form a ring round centre which slowly inflected) — Heads form a dark ring

(64

Down Sept 30 — Cause of movement from inorganic atoms

There is no circulation in viscid matter (not worth mention) the viscid almost spherical drops extend a little way beneath gland down pedicel of do.

I find that astonishing small atoms, smaller than I have generally used, of Cork & thread & Carb. of Magnesia absorb movement of the fluid, & the particle is then brought into contact with the gland itself.

The viscid fluid also spreads a little over surface. Wherever & as long as atoms of cork & glass & Human hair really floated on viscid matter there was no movement; but when by moving these atoms a point was brought into contact, with gland movement commenced. I observed this with blue-glass - hair, & cork. The point of atom of glass wd be apt to touch, as would the clip of Hair.

The weight which I give are really too great, for they act, whilst floating if any point touches for a time the gland. By observing with rather strong lens under single Microscope, I see that astonishing minute atoms produce a very slight degree of movement & no more which movements cd be overlooked, as I have hitherto observed.

We cannot understand the capricious manner in which

[64v]

inorganic atoms act viz because they do not touch, even when the gland is sensitive.— The effect of repeated touch with needle harmonises with these results.

(I doubt about inorganic atoms causing more copious secretion. I cut off heads of some tentacles & put Carb. of ammonia 1 gr to 1 oz & Phosphate of amm 1 gr to 1 oz on cut edges did not cause any movement though adjoining glands when touched caused the tentacles to bend.

(65

1872 Down. Sept 27th at 8º 13' put fragment of fly on one side of middle of leaf. After interval of 32', seven tentacle inflected, opposite to fly: ie in a line drawn from near centre of leaf, or rather lower down than centre. After interval of 10º several more tentacles on each side inflected: next morning after interval of 23º, one half of solid leaf itself inflected so much as to stand at right angle to the remainder; more tentacles towards the basal half of leaf inflected then towards the distal end, move to lower [illeg]

The tentacles on the opposite side of leaf was not in the least affected; & so things remained for 3 whole days; after 4 days the tentacle were reexpanded, those on opposite side never having been affected.

Sept 28 8º 50' bit of raw meat in exact middle of leaf

After interval of 4º 10' all the tentacles, nearest to meat, all round, inflected. After 6º stil mor closely clasped on all sides, & distal end of leaf itself inflected. After 8º the whole leaf raised upwards or lifted towards the pedicel of the leaf. Next morning after 24º - all things in the same state whole solid leaf much inflected

(66

Down. Sept 28 - put atoms of meat on medial line of leaf, near junction with pedicel: after interval of 2º 30', some tentacles alone to base of pedicel inflected After 6º tentacle on both sides of pedicel & up both edges of leaf inflected

After 8º tentacles at distal end of leaf inflected but the lateral tentacles much less inflected: After 23º meat well clasped all round, except by extreme lateral tentacles

Sept 28 put atoms of meat on medial line at distal end of leaf - After interval of 2º 40', a few tentacles at distal end clasped. After 8º the tentacles near pedicel of leaf now somewhat inflected — the lateral tentacles much less inflected

After 23º meat well clasped, except by extreme lateral tentacles. repeated with same result

(67

1872 Sept 29 3 leaves with raw meat on one side in middle - to see whether stimulus transmitted right across leaf.

(1) 8º 35' large leaf - 5º P.m very little inflection

(Sept 30 8º 20; all on one side - perhaps rather more on pedicel end - but meat was a little pushed to centre.)

Sept 30 1º inflection down exactly 1/2 leaf

Oct I exactly 1/2 tentacles

― 2d 8 Am do do— but not the most external tentacles & showing size. none on on the pedicel of leaf

― 3' 8' Reopened again fully

(2) 8º 33'— (1º a few inflected ) 5º Pm very little inflection

(Sept 30' 8º 20' all opposite, but not more near pedicel than at summit)

(Sept 30 1º inflection down 1/2 leaf, not mor toward pedicel)

Oct I exactly half of tents — 1/2 of leaf closely inflected.

― 2 do (flooded)

 3 8 A.m exactly half of leaf.

Oct 4 8' A.m do

(3) 8º 33' very small leaf- atom rather nearer pedicel

1º -a few inflected. (5º many tentacles opposite raw meat not inflected, but none towards pedicel)

Sept 30 8.20' all on one side clasped: rather more near pedicel, 1º inflection down 1/2 leaf, not more towards pedicel.

Oct 1 exactly half of tentacles

Oct 2d exactly half – flooded

Oct 3d fully reopened again

General for all leaves

9º 40' no effect on any leaf (10º 30' no) (11º 35' no)

N.B. Oct I in some cases a few tentacles at very base move, & sometimes few are inflected. —ie sometimes a trace beyond medial line & sometimes a trace within medial line inflection

[unrelated material – draft]

(68

Down Oct. 4th Examined glands carefully - crushing them under compressor. Single spiral vessel in pedicel.

[sketch] orange tint

The oval head consists of an outer layer of small polygonal cells, forming an envelope which splits off under the compressor; & within this there is an oval Head formed of longitudinally elongated cells, with purple fluid of rather a different tint, & these cells enclose a bundle of parallel spiral vessels, which converge at the base to the one within the pedicel. Hence the core appears firmly striated transversely. I cd not ascertain where these vessels were united to the one spiral vessel in pedicel; whether the branches, or whether the contents consist of much elongated spiral cells, merely resting on the spiral vessel. I saw in Long-Head, likewise a number of similar spiral vessel forming a layer rather than a bundle.

Oct 30. By boiling in Potash, I have made out that they are long cylindrical cells, ending upwards in blunt point, some shorter than others, & ending downwards in abrupt truncated ends, & of larger diameter than the spiral vessel or vessels which enter in there midrib & run some way up amidst them. Central gland same structure — Long-Headed almost the same.

[68v]

Oct 2d 5º P.m. raw meat in centre (Blue & red stick)

I gave a mere prick in centre.

Oct 3d 8º 15' tentacles, except the extreme marginals, equally inflected all round the leaf, so I suppose I missed cutting nerve. See to this (Black wool in Spirits)

Results of last alone

Oct 28 I find that the medial nerve is certainly divided, & the division is plain to the right of the nerve, but some brown (after alcohol) pulpy matter is still in concretion, when viewed from both above & below: This is important, as bearing on means of communication. (It ought to be retested by repeated pricking close on one aide of medial nerve.)

This specimen shows well the marginal circumferential inosculation / At extreme margin the same bundle divides & sends minute branches into this 2 lateral tentacles,

[sketch] base of tentacles

run only to the point of bending no? In this case wd could understand how meat on glands causes only that tentacle to bend, whilst when brought to centre, an order is sent to several tentacles to bend. — Afterward I saw in spec' boiled in potash, that very fine vessels did contact some of the marginal tentacles laterally.

(I see some cases of trifurcation of vessels or tri-separation.)

(69

Down Oct 1 9º 50 (Red & black) raw meat on centre of Leaf

2º 30' (ie after interval of 4º 40') incipient inflection from all sides, chiefly opposite sides of leaf

Oct 2d 8º 5' equally clasped all round, but as yet only by submarginals

Oct 3' ― do do do do

Oct 4th I believe fully reexpanded

for result see p. 69A

Oct 1 9º 50 - (Match with Head) raw meat on centre of leaf & nerve divided just below the centre, 2º 30' (after interval of 4º 40') trace of inflection from distal end

Oct 2d 8º only slight inflection, at distal end alone

3d do do do do

4' 8 quite reexpanded

(see what has been cut) Spirits White Thread

Oct 1 9º 55' (Blue stick) meat as last. & nerve cut as in last.

2º 30 great inflection at distal end alone

(Oct 2d 8º strong inflection exactly across, transversely middle of leaf; edge of leaf itself slightly inflected. )

(Oct 3 8' do; but the shorter one tentacles on disc towards near the pedicel are a little inflected )

Black Thread Keep in spirit

Oct 1 9º (Yellow stick) --- as 2 last ---

2º 30 great inflection of distal end. (In all these cases the nearer tentacles to the meat moved

Oct 2d 8 strong inflection of submarginal at distal end.

Oct 3 do now almost rexpanded - None at base.

White Wool Keep in spirits

/over

(p. 69A)

Oct 28th (White Thread Spec') I find central nerve & the sub lateral (from central) divided. (These latter do not spring symmetrically in this specimen) but the true Lateral nerves are not divided.

I see George's inosculation between the 2 branches of the laterals

(Black Thread) central divided close at bifurcation, & on right side the sublateral & inner branch of true lateral. On left side only the nearer sub-laterals

As the tentacles were not inflected at base no influence can be sent by Georges inosculation.

There are inosculations all round margin of leaf! (see to this under compound)

George's inosculation plain.

(White wool) central divided, & on left side centro-lateral & inner branch of lateral m but on right the true lateral not touched., Yet no motor influences sent to base! (There must be radiation from 2 distant longitudinal space, near contracted nerve side & broad spinal cord— Hence a narrow prick, does not [rush] to paralyse the lower part.

It seems from these facts that a stimulus cannot be sent by the Great inosculation down to the base???

Can the transmission be sent only upwards. (wd atom of meat on one side very high up above inosculation cause all tents. on that side to close.??)

[unrelated material]

(69B

Oct 29 I saw what appeared to be an inosculation between the inner branches of great central nerve above the centre of leaf) but on close examination under compound this seemed due to a tentacle standing exactly over one branch & being supplied by a sub- branch from other great branch.

There are certainly in some cases loops near circumference Get George to draw & there certainly seem minute sub-branches from adjoining branches enterng the bases of marginal tentacles from adjoining branches (see same specimen). The Great Inosculation certainly contain spiral vessels.

In the large marginal tentacles certainly there are fine lateral spiral vessels, as well as the medial one. & this agrees with the 3 branches which I saw entering. into a single margin tentacle in some few places.

Hot Carbonate of ammonia does not dissolve, (after alcohol) the colourless grains of Chlorophyll - Hot Potash does do so

(69C

Oct 29th It is certain that at bifurcation the spiral vessels only unite

[annotated sketch]

Bifurcation

Spiral vessels figure

Chief Inosculation

bifurcation of great nerve contract

open triangular space

the great Bifurcation

Red lines nothing

main Lateral nerve

(Same structure in the circumferential inoscln.)

[annotated sketch]

2 spiral vessels terminating in points

Central nerve

[unrelated material]

(69C

(Order of giving case)

Oct 30 /72/ After finding that meat on one side did not cause movement on opposite side; whilst at base & distal end the whole leaf was affected, I looked to vessels to see whether the course threw any light on subject.— Their course is represented in Georges drawings, of a small leaf. The bifurcation & the occasional though rare trifurcation are not real, but merely consist of separation, as was shown by boiling in Potash. These vessels in the drawing are here seen to be distinct on opposite sides; the main branch of Central is united by a Grand inosculation with the lateral; & the 2 branches of the lateral are likewise united by an inosculation. Hence there is apparently a communication down each side of leaf, but not with opposite side.

However by examining larger leaves, I found, (always with much variation), that the branches inosculate in appearance, round the circumference, (see diagram) & even the 2 main branches of the great medial vessel continues these inosculation, at distal end of leaf Therefore if motor influence always followed

(68B

the course of the vascular bundles the whole leaf wd be in communication — Nevertheless there is a difference in the inosculations; those of circumference consist simply of a branch from the bundles on each side uniting, & moving up into one of the sub-marginal tentacles. — The vessels are much complex in their courses in the grand inosculation, which latter also is invariably present.— The course of the vessels is shown by the dotted lines in the diagram — I do not say that these vessels are absolutely continuous, for in several places I found one vessel ending in a point & cohering close to another which ends in a point — dove-tailed together & cohering, but apparently separated at one point of trifurcation— there were no less than 3 vessels thus cohering by these points see part oval. — In small leaves however the the vessels in the grand inosculation ar mor simple (see zz) & do not differs essentially from the circumferential inosculations, with, however, traces of the formation of the loops

(69D

ZZ

[sketch] union by contact of 3 vessels at their trifurcating point

Small leaf Red line shows course of vessel

[unrelated material]

(69E

From this greater complexity of the vessels in the grand inosculation, for its [branches] presence even in the small leaves it seems quite possible that the motor influence follows their line & not the simpler circumferential inosculations (If so we can understand the action of meat. — no. no.) There is no need to suppose that the motor influence is transmitted by the spiral vessel, but mor probably the pulpy matter which follow these vessels & is apparently enclosed with them in a membranous sheath. parenchyma tissue Give reasons viz the case there I cut right through the medial nerve, but not all pulpy matter, & communication was kept up; & from the inosculation of the vascular bundles not being perfect owing to the unions being effected by apparently closed blunt points in close contact.

In one leaf at point of Great bifurcation of

[in margin:] (I now think 2 distinct spinal marrows parallel; & near the great medial vascular bundle)

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Median vessel, 2 spiral vessels in centre terminated upwards in little point.

Certainly very fine branches of spiral vessels enter some of the marginal tentacles from the bundles which enter the adjoining tentacles. If these vessels were lines of communication of motor influence, 3 tentacles cd move together, which does not occur.

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Down. Oct, 1872. Meat on gland of extreme marginal tentacle influence nearly to bending point & sent quickly - The meat is curved to centre not touching other glands (if it touches other gland is rolled over) meat affect several glands in middle, so as to lie on several glands & their influence sent centrifugally to all the tentacles of all round; but affects this nearest first & than the more remote.

If meat put on medial line (in same sense) close to where pedicel joins leaf, then the adjoining marginal tentacles first contract & then those at opposite pole of leaf & ultimately the laterals & all clasp the meat.— So again if meat put on several glands in middle of leaf, towards the distal end, the adjoining marginal tentacles become inflected, then those near pedicel & then all round. But these movement are much slower than from gland to the bending point of tentacle. These facts show that motor influence sent centrifugally, & likewise either up or down the middle of leaf, mor readily than to the extreme marginals.

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Hence I at first supposed that the whole disc of leaf formed a single centre & transmitted its influence in radiating lines in all directions; but this is not the case. For when I put atoms of meat on 3 leaves on one side of central disc, so that it did not touch a single gland on opposite side, not a single tentacle was inflected on the opposite side. It is evident that the 2 lateral halfs of leaves are quite independent of each other in their movements (great contrast with Dionæa). This led me to examine the leaves carefully, after long-soaking in alcohol &c so as to be transparent, & I find that the vascular are distinct or do not inosculate on the two halfs. Low down in pedicel of leaf, 3 bundles of vessels are united into a single one, then diverge before they enter leaf (see diagram). The middle one, which is the

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thickest, sends off, just where leaf commence a small branch of each side, which supplies the lower & middle part of leaf. In centre of leaf it bifurcates, (that is divides, but I believe from the study of the veins in D. Anglica does not inosculate); & each main branch bifurcates several times, the branches sending a fine spiral vessel into everyone of the tentacles. Each of the 2 lateral main vessel, likewise bifurcate 2 or 3 times & supplies the tentacles on the lower side of leaf, (a little above the centre?) Now it is remarkable, that there is about 2/3 up the leaf one time inosculation (see to this) between main lateral crosses & one of bifurcation of the central vessel; (& I suspect a second bifurcation between the 2 main branches of the lateral vessels) Thus all the tentacles on each lateral half of leaf are put into communication by the course of the vascular

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D. Anglica support this [under]

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bundles, & there is no communication with the opposite side. This makes me believe (Dutrochet) that communication must by vascular bundles or by some tissue following the course. Hence I put meat on exact centre & cut with lancet the central (& lateral?) vessels, & in this case all the tentacles at distal end of leaf & none pedicel end contracted. — So that the appearance was like that when I put meat on one side, with the exception that the line of inflection & non-inflection was at right angles to each other. I think this proves that motor-influence follows definite lines, & not through all available tissues, but along the course of vascular bundles.

The motor-influence travels, as far as we can see, centripetally & centrifugally along the same lines: we see by meat placed on middle at base & distal end of leaf; & travelling from gland to bending point. & from central glands of oth tentacles to bending point, first

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of the nearer & then more distant tentacles.

It travels quickest in single tentacle to bending point from glands (In inosculation movement travel both ways??? but travel quicker down the central vessel) (Consider all this) No central organs like brain, for influence travels in all direction, from any gland - In this respect & apparently in travelling centripetally & centrifugally, differs much from nervous system; Various kinds of stimulation travel along one or different kinds of nerve— touch & reagents — That this is close similarity to nerves will be seen when we consider all substances paralyse or destroy the power of transmission

(The most probable a priori view (∗opinion of Cohn) that the motor influence or power of transmission wd be little specialised in Plants & wd be common to all the tissues, but this clearly is not the case with Drosera. There is polar channel, but not equatorial channel.

N. B Organic substances, as shown by fluids, act in 2 ways in causing inflection & keeping inflected position.


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

File last updated 25 September, 2022