CHAPTER XII._The Power of Movement in Plants

CONCLUDING REMARKS.

Nature of the circumnutating movement—History of a germinating seed—The

radicle first protrudes and circumnutates—Its tip highly

sensitive—Emergence of the hypocotyl or of the epicotyl from the ground

under the form of an arch–Its circumnutation and that of the

cotyledons—The seedling throws up a leaf-bearing stem—The

circumnutation of all the parts or organs—Modified

circumnutation—Epinasty and hyponasty—Movements of climbing

plants—Nyctitropic movements—Movements excited by light and

gravitation—Localised sensitiveness—Resemblance between the movements

of plants and animals—The tip of the radicle acts like a brain.

It may be useful to the reader if we briefly sum up the chief

conclusions, which, as far as we can judge, have been fairly well

established by the observations given in this volume. All the parts or

organs in every plant whilst they continue to grow, and some parts

which are provided with pulvini after they have ceased to grow, are

continually circumnutating. This movement commences even before the

young seedling has broken through the ground. The nature of the

movement and its causes, as far as ascertained, have been briefly

described in the Introduction. Why every part of a plant whilst it is

growing, and in some cases after growth has ceased, should have its

cells rendered more turgescent and its cell-walls more extensile first

on one side and then on another, thus inducing circumnutation is not

known. It would appear as if the changes in the cells required periods

of rest.

In some cases, as with the hypocotyls of Brassica, the leaves of

Dionaea and the joints of the Gramineæ, the circumnutating movement

when viewed under the microscope is seen to consist of innumerable

small oscillations. The part under observation suddenly jerks forwards

for a length of .002 to .001 of an inch, and then slowly retreats for a

part of this distance; after a few seconds it again jerks forwards, but

with many intermissions. The retreating movement apparently is due to

the elasticity of the resisting tissues. How far this oscillatory

movement is general we do not know, as not many circumnutating plants

were observed by us under the microscope; but no such movement could be

detected in the case of Drosera with a 2-inch object-glass which we

used. The phenomenon is a remarkable one. The whole hypocotyl of a

cabbage or the whole leaf of a Dionaea could not jerk forwards unless a

very large number of cells on one side were simultaneously affected.

Are we to suppose that these cells steadily become more and more

turgescent on one side, until the part suddenly yields and bends,

inducing what may be called a microscopically minute earthquake in the

plant; or do the cells on one side suddenly become turgescent in an

intermittent manner; each forward movement thus caused being opposed by

the elasticity of the tissues?

Circumnutation is of paramount importance in the life of every plant;

for it is through its modification that many highly beneficial or

necessary movements have been acquired. When light strikes one side of

a plant, or light changes into darkness, or when gravitation acts on a

displaced part, the plant is enabled in some unknown manner to increase

the always varying turgescence of the cells on one side; so that the

ordinary circumnutating movement is

modified, and the part bends either to or from the exciting cause; or

it may occupy a new position, as in the so-called sleep of leaves. The

influence which modifies circumnutation may be transmitted from one

part to another. Innate or constitutional changes, independently of any

external agency, often modify the circumnutating movements at

particular periods of the life of the plant. As circumnutation is

universally present, we can understand how it is that movements of the

same kind have been developed in the most distinct members of the

vegetable series. But it must not be supposed that all the movements of

plants arise from modified circumnutation; for, as we shall presently

see, there is reason to believe that this is not the case.

Having made these few preliminary remarks, we will in imagination take

a germinating seed, and consider the part which the various movements

play in the life-history of the plant. The first change is the

protrusion of the radicle, which begins at once to circumnutate. This

movement is immediately modified by the attraction of gravity and

rendered geotropic. The radicle, therefore, supposing the seed to be

lying on the surface, quickly bends downwards, following a more or less

spiral course, as was seen on the smoked glass-plates. Sensitiveness to

gravitation resides in the tip; and it is the tip which transmits some

influence to the adjoining parts, causing them to bend. As soon as the

tip, protected by the root-cap, reaches the ground, it penetrates the

surface, if this be soft or friable; and the act of penetration is

apparently aided by the rocking or circumnutating movement of the whole

end of the radicle. If the surface is compact, and cannot easily be

penetrated, then

the seed itself, unless it be a heavy one, is displaced or lifted up by

the continued growth and elongation of the radicle. But in a state of

nature seeds often get covered with earth or other matter, or fall into

crevices, etc., and thus a point of resistance is afforded, and the tip

can more easily penetrate the ground. But even with seeds lying loose

on the surface there is another aid: a multitude of excessively fine

hairs are emitted from the upper part of the radicle, and these attach

themselves firmly to stones or other objects lying on the surface, and

can do so even to glass; and thus the upper part is held down whilst

the tip presses against and penetrates the ground. The attachment of

the root-hairs is effected by the liquefaction of the outer surface of

the cellulose walls, and by the subsequent setting hard of the

liquefied matter. This curious process probably takes place, not for

the sake of the attachment of the radicles to superficial objects, but

in order that the hairs may be brought into the closest contact with

the particles in the soil, by which means they can absorb the layer of

water surrounding them, together with any dissolved matter.

After the tip has penetrated the ground to a little depth, the

increasing thickness of the radicle, together with the root-hairs, hold

it securely in its place; and now the force exerted by the longitudinal

growth of the radicle drives the tip deeper into the ground. This

force, combined with that due to transverse growth, gives to the

radicle the power of a wedge. Even a growing root of moderate size,

such as that of a seedling bean, can displace a weight of some pounds.

It is not probable that the tip when buried in compact earth can

actually circumnutate and thus aid its downward passage, but the

circumnutating movement will facilitate the tip entering any lateral

or oblique fissure in the earth, or a burrow made by an earth-worm or

larva; and it is certain that roots often run down the old burrows of

worms. The tip, however, in endeavouring to circumnutate, will

continually press against the earth on all sides, and this can hardly

fail to be of the highest importance to the plant; for we have seen

that when little bits of card-like paper and of very thin paper were

cemented on opposite sides of the tip, the whole growing part of the

radicle was excited to bend away from the side bearing the card or more

resisting substance, towards the side bearing the thin paper. We may

therefore feel almost sure that when the tip encounters a stone or

other obstacle in the ground, or even earth more compact on one side

than the other, the root will bend away as much as it can from the

obstacle or the more resisting earth, and will thus follow with

unerring skill a line of least resistance.

The tip is more sensitive to prolonged contact with an object than to

gravitation when this acts obliquely on the radicle, and sometimes even

when it acts in the most favourable direction at right angles to the

radicle. The tip was excited by an attached bead of shellac weighing

less than 1/200th of a grain (0.33 mg.); it is therefore more sensitive

than the most delicate tendril, namely, that of Passiflora gracilis,

which was barely acted on by a bit of wire weighing 1/50th of a grain.

But this degree of sensitiveness is as nothing compared with that of

the glands of Drosera, for these are excited by particles weighing only

1/78740 of a grain. The sensitiveness of the tip cannot be accounted

for by its being covered by a thinner layer of tissue than the other

parts, for it is protected by the relatively thick root-cap. It is

remarkable that although the radicle bends away, when one side of the

tip is slightly touched

with caustic, yet if the side be much cauterised the injury is too

great, and the power of transmitting some influence to the adjoining

parts causing them to bend, is lost. Other analogous cases are known to

occur.

After a radicle has been deflected by some obstacle, geotropism directs

the tip again to grow perpendicularly downwards; but geotropism is a

feeble power, and here, as Sachs has shown, another interesting

adaptive movement comes into play; for radicles at a distance of a few

millimeters from the tip are sensitive to prolonged contact in such a

manner that they bend towards the touching object, instead of from it

as occurs when an object touches one side of the tip. Moreover, the

curvature thus caused is abrupt; the pressed part alone bending. Even

slight pressure suffices, such as a bit of card cemented to one side.

therefore a radicle, as it passes over the edge of any obstacle in the

ground, will through the action of geotropism press against it; and

this pressure will cause the radicle to endeavour to bend abruptly over

the edge. It will thus recover as quickly as possible its normal

downward course.

Radicles are also sensitive to air which contains more moisture on one

side than the other, and they bend towards its source. It is therefore

probable that they are in like manner sensitive to dampness in the

soil. It was ascertained in several cases that this sensitiveness

resides in the tip, which transmits an influence causing the adjoining

upper part to bend in opposition to geotropism towards the moist

object. We may therefore infer that roots will be deflected from their

downward course towards any source of moisture in the soil.

Again, most or all radicles are slightly sensitive to light, and

according to Wiesner, generally bend a little

from it. Whether this can be of any service to them is very doubtful,

but with seeds germinating on the surface it will slightly aid

geotropism in directing the radicles to the ground.[1] We ascertained

in one instance that such sensitiveness resided in the tip, and caused

the adjoining parts to bend from the light. The sub-aërial roots

observed by Wiesner were all apheliotropic, and this, no doubt, is of

use in bringing them into contact with trunks of trees or surfaces of

rock, as is their habit.

[1] Dr. Karl Richter, who has especially attended to this subject (‘K.

Akad. der Wissenschaften in Wien,’ 1879, p. 149), states that

apheliotropism does not aid radicles in penetrating the ground.

We thus see that with seedling plants the tip of the radicle is endowed

with diverse kinds of sensitiveness; and that the tip directs the

adjoining growing parts to bend to or from the exciting cause,

according to the needs of the plant. The sides of the radicle are also

sensitive to contact, but in a widely different manner. Gravitation,

though a less powerful cause of movement than the other above specified

stimuli, is ever present; so that it ultimately prevails and determines

the downward growth of the root.

The primary radicle emits secondary ones which project

sub-horizontally; and these were observed in one case to circumnutate.

Their tips are also sensitive to contact, and they are thus excited to

bend away from any touching object; so that they resemble in these

respects, as far as they were observed, the primary radicles. If

displaced they resume, as Sachs has shown, their original

sub-horizontal position; and this apparently is due to diageotropism.

The secondary radicles emit tertiary ones, but these, in the case of

the bean, are not affected by gravitation; consequently they protrude

in all directions. Thus the general

arrangement of the three orders of roots is excellently adapted for

searching the whole soil for nutriment.

Sachs has shown that if the tip of the primary radicle is cut off (and

the tip will occasionally be gnawed off with seedlings in a state of

nature) one of the secondary radicles grows perpendicularly downwards,

in a manner which is analogous to the upward growth of a lateral shoot

after the amputation of the leading shoot. We have seen with radicles

of the bean that if the primary radicle is merely compressed instead of

being cut off, so that an excess of sap is directed into the secondary

radicles, their natural condition is disturbed and they grow downwards.

Other analogous facts have been given. As anything which disturbs the

constitution is apt to lead to reversion, that is, to the resumption of

a former character, it appears probable that when secondary radicles

grow downwards or lateral shoots upwards, they revert to the primary

manner of growth proper to radicles and shoots.

With dicotyledonous seeds, after the protrusion of the radicle, the

hypocotyl breaks through the seed-coats; but if the cotyledons are

hypogean, it is the epicotyl which breaks forth. These organs are at

first invariably arched, with the upper part bent back parallel to the

lower; and they retain this form until they have risen above the

ground. In some cases, however, it is the petioles of the cotyledons or

of the first true leaves which break through the seed-coats as well as

the ground, before any part of the stem protrudes; and then the

petioles are almost invariably arched. We have met with only one

exception, and that only a partial one, namely, with the petioles of

the two first leaves of Acanthus candelabrum. With Delphinium nudicaule

the petioles of the two cotyledons are

completely confluent, and they break through the ground as an arch;

afterwards the petioles of the successively formed early leaves are

arched, and they are thus enabled to break through the base of the

confluent petioles of the cotyledons. In the case of Megarrhiza, it is

the plumule which breaks as an arch through the tube formed by the

confluence of the cotyledon-petioles. With mature plants, the

flower-stems and the leaves of some few species, and the rachis of

several ferns, as they emerge separately from the ground, are likewise

arched.

The fact of so many different organs in plants of many kinds breaking

through the ground under the form of an arch, shows that this must be

in some manner highly important to them. According to Haberlandt, the

tender growing apex is thus saved from abrasion, and this is probably

the true explanation. But as both legs of the arch grow, their power of

breaking through the ground will be much increased as long as the tip

remains within the seed-coats and has a point of support. In the case

of monocotyledons the plumule or cotyledon is rarely arched, as far as

we have seen; but this is the case with the leaf-like cotyledon of the

onion; and the crown of the arch is here strengthened by a special

protuberance. In the Gramineæ the summit of the straight, sheath-like

cotyledon is developed into a hard sharp crest, which evidently serves

for breaking through the earth. With dicotyledons the arching of the

epicotyl or hypocotyl often appears as if it merely resulted from the

manner in which the parts are packed within the seed; but it is

doubtful whether this is the whole of the truth in any case, and it

certainly was not so in several cases, in which the arching was seen to

commence after the parts had wholly

escaped from the seed-coats. As the arching occurred in whatever

position the seeds were placed, it is no doubt due to temporarily

increased growth of the nature of epinasty or hyponasty along one side

of the part.

As this habit of the hypocotyl to arch itself appears to be universal,

it is probably of very ancient origin. It is therefore not surprising

that it should be inherited, at least to some extent, by plants having

hypogean cotyledons, in which the hypocotyl is only slightly developed

and never protrudes above the ground, and in which the arching is of

course now quite useless. This tendency explains, as we have seen, the

curvature of the hypocotyl (and the consequent movement of the radicle)

which was first observed by Sachs, and which we have often had to refer

to as Sachs’ curvature.

The several foregoing arched organs are continually circumnutating, or

endeavouring to circumnutate, even before they break through the

ground. As soon as any part of the arch protrudes from the seed-coats

it is acted upon by apogeotropism, and both the legs bend upwards as

quickly as the surrounding earth will permit, until the arch stands

vertically. By continued growth it then forcibly breaks through the

ground; but as it is continually striving to circumnutate this will aid

its emergence in some slight degree, for we know that a circumnutating

hypocotyl can push away damp sand on all sides. As soon as the faintest

ray of light reaches a seedling, heliotropism will guide it through any

crack in the soil, or through an entangled mass of overlying

vegetation; for apogeotropism by itself can direct the seedling only

blindly upwards. Hence probably it is that sensitiveness to light

resides in the tip of the cotyledons of the Gramineæ, and in

the upper part of the hypocotyls of at least some plants.

As the arch grows upwards the cotyledons are dragged out of the ground.

The seed-coats are either left behind buried, or are retained for a

time still enclosing the cotyledons. These are afterwards cast off

merely by the swelling of the cotyledons. But with most of the

Cucurbitaceæ there is a curious special contrivance for bursting the

seed-coats whilst beneath the ground, namely, a peg at the base of the

hypocotyl, projecting at right angles, which holds down the lower half

of the seed-coats, whilst the growth of the arched part of the

hypocotyl lifts up the upper half, and thus splits them in twain. A

somewhat analogous structure occurs in Mimosa pudica and some other

plants. Before the cotyledons are fully expanded and have diverged, the

hypocotyl generally straightens itself by increased growth along the

concave side, thus reversing the process which caused the arching.

Ultimately not a trace of the former curvature is left, except in the

case of the leaf-like cotyledons of the onion.

The cotyledons can now assume the function of leaves, and decompose

carbonic acid; they also yield up to other parts of the plant the

nutriment which they often contain. When they contain a large stock of

nutriment they generally remain buried beneath the ground, owing to the

small development of the hypocotyl; and thus they have a better chance

of escaping destruction by animals. From unknown causes, nutriment is

sometimes stored in the hypocotyl or in the radicle, and then one of

the cotyledons or both become rudimentary, of which several instances

have been given. It is probable that the extraordinary manner of

germination of _Megarrhiza Californica_,

_Ipomœa leptophylla_ and _pandurata_, and of _Quercus virens_, is

connected with the burying of the tuber-like roots, which at an early

age are stocked with nutriment; for in these plants it is the petioles

of the cotyledons which first protrude from the seeds, and they are

then merely tipped with a minute radicle and hypocotyl. These petioles

bend down geotropically like a root and penetrate the ground, so that

the true root, which afterwards becomes greatly enlarged, is buried at

some little depth beneath the surface. Gradations of structure are

always interesting, and Asa Gray informs us that with Ipomœa Jalappa,

which likewise forms huge tubers, the hypocotyl is still of

considerable length, and the petioles of the cotyledons are only

moderately elongated. But in addition to the advantage gained by the

concealment of the nutritious matter stored within the tubers, the

plumule, at least in the case of Megarrhiza, is protected from the

frosts of winter by being buried.

With many dicotyledonous seedlings, as has lately been described by De

Vries, the contraction of the parenchyma of the upper part of the

radicle drags the hypocotyl downwards into the earth; sometimes (it is

said) until even the cotyledons are buried. The hypocotyl itself of

some species contracts in a like manner. It is believed that this

burying process serves to protect the seedlings against the frosts of

winter.

Our imaginary seedling is now mature as a seedling, for its hypocotyl

is straight and its cotyledons are fully expanded. In this state the

upper part of the hypocotyl and the cotyledons continue for some time

to circumnutate, generally to a wide extent relatively to the size of

the parts, and at a rapid rate. But seedlings profit by this power of

movement only when it is modified, especially by the action of light

and

gravitation; for they are thus enabled to move more rapidly and to a

greater extent than can most mature plants. Seedlings are subjected to

a severe struggle for life, and it appears to be highly important to

them that they should adapt themselves as quickly and as perfectly as

possible to their conditions. Hence also it is that they are so

extremely sensitive to light and gravitation. The cotyledons of some

few species are sensitive to a touch; but it is probable that this is

only an indirect result of the foregoing kinds of sensitiveness, for

there is no reason to believe that they profit by moving when touched.

Our seedling now throws up a stem bearing leaves, and often branches,

all of which whilst young are continually circumnutating. If we look,

for instance, at a great acacia tree, we may feel assured that every

one of the innumerable growing shoots is constantly describing small

ellipses; as is each petiole, sub-petiole, and leaflet. The latter, as

well as ordinary leaves, generally move up and down in nearly the same

vertical plane, so that they describe very narrow ellipses. The

flower-peduncles are likewise continually circumnutating. If we could

look beneath the ground, and our eyes had the power of a microscope, we

should see the tip of each rootlet endeavouring to sweep small ellipses

or circles, as far as the pressure of the surrounding earth permitted.

All this astonishing amount of movement has been going on year after

year since the time when, as a seedling, the tree first emerged from

the ground.

Stems are sometimes developed into long runners or stolons. These

circumnutate in a conspicuous manner, and are thus aided in passing

between and over surrounding obstacles. But whether the circumnutating

movement has been increased for this special purpose is doubtful.

We have now to consider circumnutation in a modified form, as the

source of several great classes of movement. The modification may be

determined by innate causes, or by external agencies. Under the first

head we see leaves which, when first unfolded, stand in a vertical

position, and gradually bend downwards as they grow older. We see

flower-peduncles bending down after the flower has withered, and others

rising up; or again, stems with their tips at first bowed downwards, so

as to be hooked, afterwards straightening themselves; and many other

such cases. These changes of position, which are due to epinasty or

hyponasty, occur at certain periods of the life of the plant, and are

independent of any external agency. They are effected not by a

continuous upward or downward movement, but by a succession of small

ellipses, or by zigzag lines,—that is, by a circumnutating movement

which is preponderant in some one direction.

Again, climbing plants whilst young circumnutate in the ordinary

manner, but as soon as the stem has grown to a certain height, which is

different for different species, it elongates rapidly, and now the

amplitude of the circumnutating movement is immensely increased,

evidently to favour the stem catching hold of a support. The stem also

circumnutates rather more equally to all sides than in the case of

non-climbing plants. This is conspicuously the case with those tendrils

which consist of modified leaves, as these sweep wide circles; whilst

ordinary leaves usually circumnutate nearly in the same vertical plane.

Flower-peduncles when converted into tendrils have their circumnutating

movement in like manner greatly increased.

We now come to our second group of

circumnutating movements—those modified through external agencies. The

so-called sleep or nyctitropic movements of leaves are determined by

the daily alternations of light and darkness. It is not the darkness

which excites them to move, but the difference in the amount of light

which they receive during the day and night; for with several species,

if the leaves have not been brightly illuminated during the day, they

do not sleep at night. They inherit, however, some tendency to move at

the proper periods, independently of any change in the amount of light.

The movements are in some cases extraordinarily complex, but as a full

summary has been given in the chapter devoted to this subject, we will

here say but little on this head. Leaves and cotyledons assume their

nocturnal position by two means, by the aid of pulvini and without such

aid. In the former case the movement continues as long as the leaf or

cotyledon remains in full health; whilst in the latter case it

continues only whilst the part is growing. Cotyledons appear to sleep

in a larger proportional number of species than do leaves. In some

species, the leaves sleep and not the cotyledons; in others, the

cotyledons and not the leaves; or both may sleep, and yet assume widely

different positions at night.

Although the nyctitropic movements of leaves and cotyledons are

wonderfully diversified, and sometimes differ much in the species of

the same genus, yet the blade is always placed in such a position at

night, that its upper surface is exposed as little as possible to full

radiation. We cannot doubt that this is the object gained by these

movements; and it has been proved that leaves exposed to a clear sky,

with their blades compelled to remain horizontal, suffered much more

from the cold than others which were allowed to assume

their proper vertical position. Some curious facts have been given

under this head, showing that horizontally extended leaves suffered

more at night, when the air, which is not cooled by radiation, was

prevented from freely circulating beneath their lower surfaces; and so

it was, when the leaves were allowed to go to sleep on branches which

had been rendered motionless. In some species the petioles rise up

greatly at night, and the pinnae close together. The whole plant is

thus rendered more compact, and a much smaller surface is exposed to

radiation.

That the various nyctitropic movements of leaves result from modified

circumnutation has, we think, been clearly shown. In the simplest cases

a leaf describes a single large ellipse during the 24 h.; and the

movement is so arranged that the blade stands vertically during the

night, and reassumes its former position on the following morning. The

course pursued differs from ordinary circumnutation only in its greater

amplitude, and in its greater rapidity late in the evening and early on

the following morning. Unless this movement is admitted to be one of

circumnutation, such leaves do not circumnutate at all, and this would

be a monstrous anomaly. In other cases, leaves and cotyledons describe

several vertical ellipses during the 24 h.; and in the evening one of

them is increased greatly in amplitude until the blade stands

vertically either upwards or downwards. In this position it continues

to circumnutate until the following morning, when it reassumes its

former position. These movements, when a pulvinus is present, are often

complicated by the rotation of the leaf or leaflet; and such rotation

on a small scale occurs during ordinary circumnutation. The many

diagrams showing the movements of sleeping and non-sleeping leaves and

cotyledons should be compared, and it will be seen that they are

essentially alike. Ordinary circumnutation is converted into a

nyctitropic movement, firstly by an increase in its amplitude, but not

to so great a degree as in the case of climbing plants, and secondly by

its being rendered periodic in relation to the alternations of day and

night. But there is frequently a distinct trace of periodicity in the

circumnutating movements of non-sleeping leaves and cotyledons. The

fact that nyctitropic movements occur in species distributed in many

families throughout the whole vascular series, is intelligible, if they

result from the modification of the universally present movement of

circumnutation; otherwise the fact is inexplicable.

In the seventh chapter we have given the case of a Porlieria, the

leaflets of which remained closed all day, as if asleep, when the plant

was kept dry, apparently for the sake of checking evaporation.

Something of the same kind occurs with certain Gramineæ. At the close

of this same chapter, a few observations were appended on what may be

called the embryology of leaves. The leaves produced by young shoots on

cut-down plants of Melilotus Taurica slept like those of a Trifolium,

whilst the leaves on the older branches on the same plants slept in a

very different manner, proper to the genus; and from the reasons

assigned we are tempted to look at this case as one of reversion to a

former nyctitropic habit. So again with Desmodium gyrans, the absence

of small lateral leaflets on very young plants, makes us suspect that

the immediate progenitor of this species did not possess lateral

leaflets, and that their appearance in an almost rudimentary condition

at a somewhat more advanced age is the result of reversion to a

trifoliate predecessor. However this may be, the rapid circumnutating

gyrating movements of the little lateral leaflets, seem to be due

proximately to the pulvinus, or organ of movement, not having been

reduced nearly so much as the blade, during the successive

modifications through which the species has passed.

We now come to the highly important class of movements due to the

action of a lateral light. When stems, leaves, or other organs are

placed, so that one side is illuminated more brightly than the other,

they bend towards the light. This heliotropic movement manifestly

results from the modification of ordinary circumnutation; and every

gradation between the two movements could be followed. When the light

was dim, and only a very little brighter on one side than on the other,

the movement consisted of a succession of ellipses, directed towards

the light, each of which approached nearer to its source than the

previous one. When the difference in the light on the two sides was

somewhat greater, the ellipses were drawn out into a strongly-marked

zigzag line, and when much greater the course became rectilinear. We

have reason to believe that changes in the turgescence of the cells is

the proximate cause of the movement of circumnutation; and it appears

that when a plant is unequally illuminated on the two sides, the always

changing turgescence is augmented along one side, and is weakened or

quite arrested along the other sides. Increased turgescence is commonly

followed by increased growth, so that a plant which has bent itself

towards the light during the day would be fixed in this position were

it not for apogeotropism acting during the night. But parts provided

with pulvini bend, as Pfeffer has shown, towards the light; and here

growth does not come into play any more than in the ordinary

circumnutating movements of pulvini.

Heliotropism prevails widely throughout the vegetable kingdom, but

whenever, from the changed habits of life of any plant, such movements

become injurious or useless, the tendency is easily eliminated, as we

see with climbing and insectivorous plants.

Apheliotropic movements are comparatively rare in a well-marked degree,

excepting with sub-aërial roots. In the two cases investigated by us,

the movement certainly consisted of modified circumnutation.

The position which leaves and cotyledons occupy during the day, namely,

more or less transversely to the direction of the light, is due,

according to Frank, to what we call diaheliotropism. As all leaves and

cotyledons are continually circumnutating, there can hardly be a doubt

that diaheliotropism results from modified circumnutation. From the

fact of leaves and cotyledons frequently rising a little in the

evening, it appears as if diaheliotropism had to conquer during the

middle of the day a widely prevalent tendency to apogeotropism.

Lastly, the leaflets and cotyledons of some plants are known to be

injured by too much light; and when the sun shines brightly on them,

they move upwards or downwards, or twist laterally, so that they direct

their edges towards the light, and thus they escape being injured.

These paraheliotropic movements certainly consisted in one case of

modified circumnutation; and so it probably is in all cases, for the

leaves of all the species described circumnutate in a conspicuous

manner. This movement has hitherto been observed only with leaflets

provided with pulvini, in which the increased turgescence on opposite

sides is not followed by growth; and we can understand why this should

be so, as the movement is required only for a temporary purpose. It

would manifestly be

disadvantageous for the leaf to be fixed by growth in its inclined

position. For it has to assume its former horizontal position, as soon

as possible after the sun has ceased shining too brightly on it.

The extreme sensitiveness of certain seedlings to light, as shown in

our ninth chapter, is highly remarkable. The cotyledons of Phalaris

became curved towards a distant lamp, which emitted so little light,

that a pencil held vertically close to the plants, did not cast any

shadow which the eye could perceive on a white card. These cotyledons,

therefore, were affected by a difference in the amount of light on

their two sides, which the eye could not distinguish. The degree of

their curvature within a given time towards a lateral light did not

correspond at all strictly with the amount of light which they

received; the light not being at any time in excess. They continued for

nearly half an hour to bend towards a lateral light, after it had been

extinguished. They bend with remarkable precision towards it, and this

depends on the illumination of one whole side, or on the obscuration of

the whole opposite side. The difference in the amount of light which

plants at any time receive in comparison with what they have shortly

before received, seems in all cases to be the chief exciting cause of

those movements which are influenced by light. Thus seedlings brought

out of darkness bend towards a dim lateral light, sooner than others

which had previously been exposed to daylight. We have seen several

analogous cases with the nyctitropic movements of leaves. A striking

instance was observed in the case of the periodic movements of the

cotyledons of a Cassia; in the morning a pot was placed in an obscure

part of a room, and all the cotyledons rose up closed; another pot had

stood in the sunlight, and

the cotyledons of course remained expanded; both pots were now placed

close together in the middle of the room, and the cotyledons which had

been exposed to the sun, immediately began to close, while the others

opened; so that the cotyledons in the two pots moved in exactly

opposite directions whilst exposed to the same degree of light.

We found that if seedlings, kept in a dark place, were laterally

illuminated by a small wax taper for only two or three minutes at

intervals of about three-quarters of an hour, they all became bowed to

the point where the taper had been held. We felt much surprised at this

fact, and until we had read Wiesner’s observations, we attributed it to

the after-effects of the light; but he has shown that the same degree

of curvature in a plant may be induced in the course of an hour by

several interrupted illuminations lasting altogether for 20 m., as by a

continuous illumination of 60 m. We believe that this case, as well as

our own, may be explained by the excitement from light being due not so

much to its actual amount, as to the difference in amount from that

previously received; and in our case there were repeated alternations

from complete darkness to light. In this, and in several of the above

specified respects, light seems to act on the tissues of plants, almost

in the same manner as it does on the nervous system of animals. There

is a much more striking analogy of the same kind, in the sensitiveness

to light being localised in the tips of the cotyledons of Phalaris and

Avena, and in the upper part of the hypocotyls of Brassica and Beta;

and in the transmission of some influence from these upper to the lower

parts, causing the latter to bend towards the light. This influence is

also

transmitted beneath the soil to a depth where no light enters. It

follows from this localisation, that the lower parts of the cotyledons

of Phalaris, etc., which normally become more bent towards a lateral

light than the upper parts, may be brightly illuminated during many

hours, and will not bend in the least, if all light be excluded from

the tip. It is an interesting experiment to place caps over the tips of

the cotyledons of Phalaris, and to allow a very little light to enter

through minute orifices on one side of the caps, for the lower part of

the cotyledons will then bend to this side, and not to the side which

has been brightly illuminated during the whole time. In the case of the

radicles of Sinapis alba, sensitiveness to light also resides in the

tip, which, when laterally illuminated, causes the adjoining part of

the root to bend apheliotropically.

Gravitation excites plants to bend away from the centre of the earth,

or towards it, or to place themselves in a transverse position with

respect to it. Although it is impossible to modify in any direct manner

the attraction of gravity, yet its influence could be moderated

indirectly, in the several ways described in the tenth chapter; and

under such circumstances the same kind of evidence as that given in the

chapter on Heliotropism, showed in the plainest manner that

apogeotropic and geotropic, and probably diageotropic movements, are

all modified forms of circumnutation.

Different parts of the same plant and different species are affected by

gravitation in widely different degrees and manners. Some plants and

organs exhibit hardly a trace of its action. Young seedlings which, as

we know, circumnutate rapidly, are eminently sensitive; and we have

seen the hypocotyl of Beta bending

upwards through 109° in 3 h. 8 m. The after-effects of apogeotropism

last for above half an hour; and horizontally-laid hypocotyls are

sometimes thus carried temporarily beyond an upright position. The

benefits derived from geotropism, apogeotropism, and diageotropism, are

generally so manifest that they need not be specified. With the

flower-peduncles of Oxalis, epinasty causes them to bend down, so that

the ripening pods may be protected by the calyx from the rain.

Afterwards they are carried upwards by apogeotropism in combination

with hyponasty, and are thus enabled to scatter their seeds over a

wider space. The capsules and flower-heads of some plants are bowed

downwards through geotropism, and they then bury themselves in the

earth for the protection and slow maturation of the seeds. This burying

process is much facilitated by the rocking movement due to

circumnutation.

In the case of the radicles of several, probably of all seedling

plants, sensitiveness to gravitation is confined to the tip, which

transmits an influence to the adjoining upper part, causing it to bend

towards the centre of the earth. That there is transmission of this

kind was proved in an interesting manner when horizontally extended

radicles of the bean were exposed to the attraction of gravity for 1 or

1½ h., and their tips were then amputated. Within this time no trace of

curvature was exhibited, and the radicles were now placed pointing

vertically downwards; but an influence had already been transmitted

from the tip to the adjoining part, for it soon became bent to one

side, in the same manner as would have occurred had the radicle

remained horizontal and been still acted on by geotropism. Radicles

thus treated continued to grow out horizontally for two or three days,

until a new tip was

re-formed; and this was then acted on by geotropism, and the radicle

became curved perpendicularly downwards.

It has now been shown that the following important classes of movement

all arise from modified circumnutation, which is omnipresent whilst

growth lasts, and after growth has ceased, whenever pulvini are

present. These classes of movement consist of those due to epinasty and

hyponasty,—those proper to climbing plants, commonly called revolving

nutation,—the nyctitropic or sleep movements of leaves and

cotyledons,—and the two immense classes of movement excited by light

and gravitation. When we speak of modified circumnutation we mean that

light, or the alternations of light and darkness, gravitation, slight

pressure or other irritants, and certain innate or constitutional

states of the plant, do not directly cause the movement; they merely

lead to a temporary increase or diminution of those spontaneous changes

in the turgescence of the cells which are already in progress. In what

manner, light, gravitation, etc., act on the cells is not known; and we

will here only remark that, if any stimulus affected the cells in such

a manner as to cause some slight tendency in the affected part to bend

in a beneficial manner, this tendency might easily be increased through

the preservation of the more sensitive individuals. But if such bending

were injurious, the tendency would be eliminated unless it was

overpoweringly strong; for we know how commonly all characters in all

organisms vary. Nor can we see any reason to doubt, that after the

complete elimination of a tendency to bend in some one direction under

a certain stimulus, the power to bend in a directly

opposite direction might gradually be acquired through natural

selection.[2]

[2] See the remarks in Frank’s ‘Die wagerechte Richtung von

Pflanzentheilen’ (1870, pp. 90, 91, etc.), on natural selection in

connection with geotropism, heliotropism, etc.

Although so many movements have arisen through modified circumnutation,

there are others which appear to have had a quite independent origin;

but they do not form such large and important classes. When a leaf of a

Mimosa is touched it suddenly assumes the same position as when asleep,

but Brucke has shown that this movement results from a different state

of turgescence in the cells from that which occurs during sleep; and as

sleep-movements are certainly due to modified circumnutation, those

from a touch can hardly be thus due. The back of a leaf of Drosera

rotundifolia was cemented to the summit of a stick driven into the

ground, so that it could not move in the least, and a tentacle was

observed during many hours under the microscope; but it exhibited no

circumnutating movement, yet after being momentarily touched with a bit

of raw meat, its basal part began to curve in 23 seconds. This curving

movement therefore could not have resulted from modified

circumnutation. But when a small object, such as a fragment of a

bristle, was placed on one side of the tip of a radicle, which we know

is continually circumnutating, the induced curvature was so similar to

the movement caused by geotropism, that we can hardly doubt that it is

due to modified circumnutation. A flower of a Mahonia was cemented to a

stick, and the stamens exhibited no signs of circumnutation under the

microscope, yet when they were lightly touched they suddenly moved

towards the pistil. Lastly, the curling of the extremity of a tendril

when

touched seems to be independent of its revolving or circumnutating

movement. This is best shown by the part which is the most sensitive to

contact, circumnutating much less than the lower parts, or apparently

not at all.[3]

[3] For the evidence on this head, see the ‘Movements and Habits of

Climbing Plants,’ 1875, pp. 173, 174.

Although in these cases we have no reason to believe that the movement

depends on modified circumnutation, as with the several classes of

movement described in this volume, yet the difference between the two

sets of cases may not be so great as it at first appears. In the one

set, an irritant causes an increase or diminution in the turgescence of

the cells, which are already in a state of change; whilst in the other

set, the irritant first starts a similar change in their state of

turgescence. Why a touch, slight pressure or any other irritant, such

as electricity, heat, or the absorption of animal matter, should modify

the turgescence of the affected cells in such a manner as to cause

movement, we do not know. But a touch acts in this manner so often, and

on such widely distinct plants, that the tendency seems to be a very

general one; and if beneficial, it might be increased to any extent. In

other cases, a touch produces a very different effect, as with Nitella,

in which the protoplasm may be seen to recede from the walls of the

cell; in Lactuca, in which a milky fluid exudes; and in the tendrils of

certain Vitaceae, Cucurbitaceæ, and Bignoniaceae, in which slight

pressure causes a cellular outgrowth.

Finally it is impossible not to be struck with the resemblance between

the foregoing movements of plants and many of the actions performed

unconsciously by the lower animals.[4] With plants an

astonishingly small stimulus suffices; and even with allied plants one

may be highly sensitive to the slightest continued pressure, and

another highly sensitive to a slight momentary touch. The habit of

moving at certain periods is inherited both by plants and animals; and

several other points of similitude have been specified. But the most

striking resemblance is the localisation of their sensitiveness, and

the transmission of an influence from the excited part to another which

consequently moves. Yet plants do not of course possess nerves or a

central nervous system; and we may infer that with animals such

structures serve only for the more perfect transmission of impressions,

and for the more complete intercommunication of the several parts.

[4] Sachs remarks to nearly the same effect: “Dass sich die lebende

Pflanzensubstanz derart innerlich differenzirt, dass einzelne Theile

mit specifischen Energien ausgerüstet sind, ähnlich, wie die

verschiedenen Sinnesnerven des Thiere” (‘Arbeiten des Bot. Inst. in

Würzburg,’ Bd. ii. 1879, p. 282).

We believe that there is no structure in plants more wonderful, as far

as its functions are concerned, than the tip of the radicle. If the tip

be lightly pressed or burnt or cut, it transmits an influence to the

upper adjoining part, causing it to bend away from the affected side;

and, what is more surprising, the tip can distinguish between a

slightly harder and softer object, by which it is simultaneously

pressed on opposite sides. If, however, the radicle is pressed by a

similar object a little above the tip, the pressed part does not

transmit any influence to the more distant parts, but bends abruptly

towards the object. If the tip perceives the air to be moister on one

side than on the other, it likewise transmits an influence to the upper

adjoining part, which bends towards the source of moisture. When the

tip is excited by light (though

in the case of radicles this was ascertained in only a single instance)

the adjoining part bends from the light; but when excited by

gravitation the same part bends towards the centre of gravity. In

almost every case we can clearly perceive the final purpose or

advantage of the several movements. Two, or perhaps more, of the

exciting causes often act simultaneously on the tip, and one conquers

the other, no doubt in accordance with its importance for the life of

the plant. The course pursued by the radicle in penetrating the ground

must be determined by the tip; hence it has acquired such diverse kinds

of sensitiveness. It is hardly an exaggeration to say that the tip of

the radicle thus endowed, and having the power of directing the

movements of the adjoining parts, acts like the brain of one of the

lower animals; the brain being seated within the anterior end of the

body, receiving impressions from the sense-organs, and directing the

several movements.

INDEX.

Abies communis, effect of killing or injuring the leading shoot, 187

— pectinata, effect of killing or injuring the leading shoot, 187

—, affected by Æcidium elatinum, 188

Abronia umbellata, its single, developed cotyledon, 78

—, rudimentary cotyledon, 95

—, rupture of the seed coats, 105

Abutilon Darwinii, sleep of leaves and not of cotyledons, 314

—, nocturnal movement of leaves, 323

Acacia Farnesiana, state of plant when awake and asleep, 381, 382

—, appearance at night, 395

—, nyctitropic movements of pinnae, 402

—, the axes of the ellipses, 404

— lophantha, character of first leaf, 415

— retinoides, circumnutation of young phyllode, 236

Acanthosicyos horrida, nocturnal movement of cotyledon 304

Acanthus candelabrum, inequality in the two first leaves, 79

—, petioles not arched, 553

— latifolius, variability in first leaves 79

— mollis, seedling, manner of breaking through the ground, 78, 79

—, circumnutation of young leaf, 249, 269

— spinosus, 79

—, movement of leaves, 249

Adenanthera pavonia, nyctitropic movements of leaflets, 374

Æcidium elatinum, effect on the lateral branches of the silver fir, 188

Æsculus hippocastanum, movements of radicle, 28, 29

—, sensitiveness of apex of radicle, 172–174

Albizzia lophantha, nyctitropic movements of leaflets, 383

—, of pinnae, 402

Allium cepa, conical protuberance on arched cotyledon, 59

—, circumnutation of basal half of arched cotyledon, 60

—, mode of breaking through ground, 87

—, straightening process, 101

— porrum, movements of flower-stems, 226

Alopecurus pratensis, joints affected by apogeotropism, 503

Aloysia citriodora, circumnutation of stem, 210

Amaranthus, sleep of leaves, 387

— caudatus, nocturnal movement of cotyledons, 307

Amorpha fruticosa, sleep of leaflets, 354

Ampelopsis tricuspidata, hyponastic movement of hooked tips, 272–275

Amphicarpoea monoica, circumnutation and nyctitropic movements of leaves,

365

—, effect of sunshine on leaflets, 445

—, geotropic movements of, 520

Anoda Wrightii, sleep of cotyledons, 302, 312

—, of leaves, 324

—, downward movement of cotyledons, 444

Apheliotropism, or negative heliotropism, 5, 419, 432

Apios graveolens, heliotropic movements of hypocotyl, 422–424

— tuberosa, vertical sinking of leaflets at night, 368

Apium graveolens, sleep of cotyledons, 305

—, petroselinum, sleep of cotyledons, 304

Apogeotropic movements effected by joints or pulvini, 502

Apogeotropism, 5, 494; retarded by heliotropism, 501; concluding remarks

on, 507

Arachis hypogoea, circumnutation of gynophore, 225

—, effects of radiation on leaves, 289, 296

—, movements of leaves, 357

— rate of movement, 404

—, circumnutation of vertically dependent young gynophores, 519

—, downward movement of the same, 519

Arching of various organs, importance of, to seedling plants, 87, 88;

emergence of hypocotyls or epicotyls in the form of an, 553

Asparagus officinalis, circumnutation of plumules, 60–62.

—, effect of lateral light, 484

Asplenium trichomanes, movement in the fruiting fronds, 257, n.

Astragalus uliginosus, movement of leaflets, 355

Avena sativa, movement of cotyledons, 65, 66.

—, sensitiveness of tip of radicle to moist air, 183

—, heliotropic movement and circumnutation of cotyledon, 421, 422

—, sensitiveness of cotyledon to a lateral light, 477

—, young sheath-like cotyledons strongly apogeotropic, 499

Avena sativa, movements of oldish cotyledons, 499, 500

Averrhoa bilimbi, leaf asleep, 330

—, angular movements when going to sleep, 331–335

—, leaflets exposed to bright sunshine, 447

Azalea Indica, circumnutation of stem, 208

Bary, de, on the effect of the Æcidium on the silver fir, 188

Batalin, Prof., on the nyctitropic movements of leaves, 283; on the sleep

of leaves of Sida napoea, 322; on Polygonum aviculare, 387; on the effect

of sunshine on leaflets of Oxalis acetosella, 447

Bauhinia, nyctitropic movements, 373

—, movements of petioles of young seedlings, 401

—, appearance of young plants at night, 402

Beta vulgaris, circumnutation of hypocotyl of seedlings, 52

—, movements of cotyledons, 52, 53

—, effect of light, 124

—, nocturnal movement of cotyledons, 307

—, heliotropic movements of, 420

—, transmitted effect of light on hypocotyl, 482

—, apogeotropic movement of hypocotyl, 496

Bignonia capreolata, apheliotropic movement of tendrils, 432, 450

Bouché on Melaleuca ericaefolia, 383

Brassica napus, circumnutation of flower-stems, 226

Brassica oleracea, circumnutation of seedling, 10

—, of radicle, 11

—, geotropic movement of radicle, 11

—, movement of buried and arched hypocotyl, 13, 14, 15

—, conjoint circumnutation of hypocotyl and cotyledons, 16, 17, 18

—, of hypocotyl in darkness, 19

—, of a cotyledon with hypocotyl secured to a stick, 19, 20

—, rate of movement, 20

—, ellipses described by hypocotyls when erect, 105

—, movements of cotyledons, 115

—, — of stem, 202

—, — of leaves at night, 229, 230

—, sleep of cotyledons, 301

—, circumnutation of hypocotyl of seedling plant, 425

—, heliotropic movement and circumnutation of hypocotyls, 426

—, effect of lateral light on hypocotyls, 479–482

—, apogeotropic movement of hypocotyls, 500, 501

Brassica rapa, movements of leaves, 230

Brongniart, A., on the sleep of Strephium floribundum, 391

Bruce, Dr., on the sleep of leaves in Averrhoa, 330

Bryophyllum (vel Calanchoe) calycinum, movement of leaves, 237

Camellia Japonica, circumnutation of leaf, 231, 232

Candolle, A. de, on Trapa natans, 95; on sensitiveness of cotyledons, 127

Canna Warscewiczii, circumnutation of plumules, 58, 59

—, of leaf, 252

Cannabis sativa, movements of leaves, 250

—, nocturnal movements of cotyledons, 307

Cannabis sativa, sinking of the young leaves at night, 444

Cassia, nyctitropic movement of leaves, 369

Cassia Barclayana, nocturnal movement of leaves, 372

—, slight movement of leaflets, 401

— calliantha, uninjured by exposure at night, 289, n.

—, nyctitropic movement of leaves, 371

— circumnutating movement of leaves, 372

— corymbosa, cotyledons sensitive to contact, 126

—, nyctitropic movement of leaves, 369

— floribunda, use of sleep movements, 289

—, effect of radiation on the leaves at night, 294

—, circumnutating and nyctitropic movement of a terminal leaflet, 372, 373

—, movements of young and older leaves, 400

— florida, cotyledons sensitive to contact, 126

—, sleep of cotyledons, 308

— glauca, cotyledons sensitive to contact, 126

—, sleep of cotyledons, 308

— laevigata, effect of radiation on leaves, 289, n.

— mimosoides, movement of cotyledons. 116

—, sensitiveness of, 126

—, sleep of, 308

—, nyctitropic movement of leaves, 372

—, effect of bright sunshine on cotyledons, 446

— neglecta, movements of, 117

—, effect of light, 124

—, sensitiveness of cotyledons, 126

— nodosa, non-sensitive cotyledons, 126

—, do not rise at night, 308

— pubescens, non-sensitive cotyledons, 126

Cassia pubescens, uninjured by exposure at night, 293

—, sleep of cotyledons, 308

—, nyctitropic movement of leaves, 371

—, circumnutating movement of leaves, 372

—, nyctitropic movement of petioles, 400

—, diameter of plant at night, 402

— sp. (?) movement of cotyledons, 116

— tora, circumnutation of cotyledons and hypocotyls, 34, 35, 109, 308

—, effect of light, 124, 125

—, sensitiveness to contact, 125

—, heliotropic movement and circumnutation of hypocotyl, 431

—, hypocotyl of seedling slightly heliotropic, 454

—, apogeotropic movement of old hypocotyl, 497

—, movement of hypocotyl of young seedling, 510

Caustic (nitrate of silver), effect of, on radicle of bean, 150, 156; on

the common pea, 160.

Cells, table of the measurement of, in the pulvini of Oxalis corniculata,

120; changes in, 547

Centrosema, 365

Ceratophyllum demersum, movements of stem, 211

Cereus Landbeckii, its rudimentary cotyledons, 97

— speciossimus, circumnutation of stem, 206, 207

Cerinthe major, circumnutation of hypocotyl, 49

—, of cotyledons, 49

—, ellipses described by hypocotyls when erect, 107

— effect of darkness, 124

Chatin, M., on Pinus Nordmanniana, 389

Chenopodium album, sleep of leaves but not of cotyledons, 314, 319

Chenopodium album, movement of leaves, 387

Chlorophyll injured by bright light, 446

Ciesielski, on the sensitiveness of the tip of the radicles, 4, 523

Circumnutation, meaning explained, 1; modified, 263–279; and heliotropism,

relation between, 435; of paramount importance to every plant, 547

Cissus discolor, circumnutation of leaf, 233

Citrus aurantium, circumnutation of epicotyl, 28

—, unequal cotyledons, 95

Clianthus Dampieri, nocturnal movement of leaves, 297

Cobœa scandens, circumnutation of, 270

Cohn, on the water secreted by Lathraea squamaria, 86, n.; on the movement

of leaflets of Oxalis, 447

Colutea arborea, nocturnal movement of leaflets, 355

Coniferæ, circumnutation of, 211

Coronilla rosea, leaflets asleep, 355

Corylus avellana, circumnutation of young shoot, emitted from the epicotyl,

55, 56

—, arched epicotyl, 77

Cotyledon umbilicus, circumnutation of stolons, 219, 220

Cotyledons, rudimentary, 94–98; circumnutation of, 109–112; nocturnal

movements, 111, 112; pulvini or joints of, 112–122; disturbed periodic

movements by light, 123; sensitiveness of, to contact, 125; nyctitropic

movements of, 283, 297; list of cotyledons which rise or sink at night,

300; concluding remarks on their movements, 311

Crambe maritima, circumnutation of leaves, 228, 229

Crinum Capense, shape of leaves, 253

—, circumnutation of, 254

Crotolaria (sp.?), sleep of leaves, 340

Cryptogams, circumnutation of, 257–259

Cucumis dudaim, movement of cotyledons, 43, 44

—, sleep of cotyledons, 304

Cucurbita aurantia, movement of hypocotyl, 42

—, cotyledons vertical at night, 304

—, ovifera, geotropic movement of radicle, 38, 39

—, circumnutation of arched hypocotyl, 39

—, of straight and vertical hypocotyl, 40

—, movements of cotyledons, 41, 42, 115, 124

—, position of radicle, 89

—, rupture of the seed-coats, 102

—, circumnutation of hypocotyl when erect, 107, 108

—, sensitiveness of apex of radicle, 169–171

—, cotyledons vertical at night, 304

—, not affected by apogeotropism, 509

—, tips cauterised transversely, 537

Curvature of the radicle, 193

Cycas pectinata, circumnutation of young leaf, whilst emerging from the

ground, 58

—, first leaf arched, 78

—, circumnutation of terminal leaflets, 252

Cyclamen Persicum, movement of cotyledon, 46

—, undeveloped cotyledons, 78, 96

—, circumnutation of peduncle, 225

—, —, of leaf, 246, 247

—, downward apheliotropic movement of a flower-peduncle, 433–435

Cyclamen Persicum, burying of the pods, 433

Cyperus alternifolius, circumnutation of stem, 212

—, movement of stem, 509

Cytisus fragrans, circumnutation of hypocotyl, 37

—, sleep of leaves, 344, 397

—, apogeotropic movement of stem, 494–496

Dahlia, circumnutation of young leaves, 244–246

Dalea alopecuroides, leaflets depressed at night, 354

Darkness, effect of, on the movement of leaves, 407

Darlingtonia Californica, its leaves or pitchers apheliotropic, 450, n.

Darwin, Charles, on Maurandia semperflorens, 225; on the Swedish turnip,

230, n.; movements of climbing plants, 266, 271; the heliotropic movement

of the tendrils of Bignonia capreolata, 433; revolution of climbing plants,

451; on the curling of a tendril, 570

—, Erasmus, on the peduncles of Cyclamens, 433

—, Francis, on the radicle of Sinapis alba, 486; on Hygroscopic seeds,

489, n.

Datura stramonium, nocturnal movement of cotyledons, 298

Delpino, on cotyledons of Chaerophyllum and Corydalis, 96, n.

Delphinium nudicaule, mode of breaking through the ground, 80

—, confluent petioles of two cotyledons, 553

Desmodium gyrans, movement of leaflets, 257, n.

—, position of leaves at night, 285

—, sleep of leaves, not of cotyledons, 314

—, circumnutation and nyctitropic movement of leaves, 358–360

—, movement of lateral leaflets, 361

—, jerking of leaflets, 362

— nyctitropic movement of petioles, 400, 401

—, diameter of plant at night, 402

—, lateral movement of leaves, 404

—, zigzag movement of apex of leaf, 405

—, shape of lateral leaflet, 416

—, vespertilionis, 364, n.

Deutzia gracilis, circumnutation of stem, 205

Diageotropism, 5; or transverse-geotropism, 520

Diaheliotropism, 5; or Transversal-Heliotropismus of Frank, 419; influenced

by epinasty, 439; by weight and apogeotropism, 440

Dianthus caryophyllus, 230

—, circumnutation of young leaf, 231, 269

Dicotyledons, circumnutation widely spread among, 68

Dionoea, oscillatory movements of leaves, 261, 271

Dionoea muscipula, circumnutation of young expanding leaf, 239, 240

—, closure of the lobes and circumnutation of a full-grown leaf, 241

—, oscillations of, 242–244

Diurnal sleep, 419

Drosera Capensis, structure of first-formed leaves, 414

— rotundifolia, movement of young leaf, 237, 238

—, of the tentacles, 239

—, sensitiveness of tentacles, 261

—, shape of leaves, 414

—, leaves not heliotropic, 450

—, leaves circumnutate largely, 454

—, sensitiveness of 570

Duchartre on Trephrosia cariboea, 354; on the nyctitropic movement of the

Cassia, 369

Duval-Jouve, on the movements of Bryophyllum calycinum, 237; of the narrow

leaves of the Gramineæ, 413

Dyer, Mr. Thiselton, on the leaves of Crotolaria, 340; on Cassia

floribunda, 369, n., on the absorbent hairs on the buried flower-heads of

Trifolium subterraneum, 517

Echeveria stolonifera, circumnutation of leaf, 237

Echinocactus viridescens, its rudimentary cotyledons, 97

Echinocystis lobata, movements of tendrils, 266

—, apogeotropism of tendrils, 510

Elfving, F., on the rhizomes of Sparganium ramosum, 189; on the

diageotropic movement in the rhizomes of some plants, 521

Elymus arenareus, leaves closed during the day, 413

Embryology of leaves, 414

Engelmann, Dr., on the Quercus virens, 85

Epinasty, 5, 267

Epicotyl, or plumule, 5; manner of breaking through the ground, 77; emerges

from the ground under the form of an arch, 553

Erythrina caffra, sleep of leaves, 367

— corallodendron, movement of terminal leaflet, 367

— crista-galli, effect of temperature on sleep of leaves, 318

—, circumnutation and nyctitropic movement of terminal leaflets, 367

Eucalyptus resinifera, circumnutation of leaves, 244

Euphorbia jacquineaeflora, nyctitropic movement of leaves, 388

Flahault, M., on the rupture of seed-coats, 102–104, 106

Flower-stems, circumnutation of, 223–226

Fragaria Rosacea, circumnutation of stolon, 214–218

Frank, Dr. A. B., the terms Heliotropism and Geotropism, first used by him,

5, n.; radicles acted on by geotropism, 70, n.; on the stolons of Fragaria,

215; periodic and nyctitropic movements of leaves, 284; on the root-leaves

of plants kept in darkness, 443; on pulvini, 485; on natural selection in

connection with geotropism, heliotropism, etc., 570

—, on Transversal-Heliotropismus, 419

Fuchsia, circumnutation of stem, 205, 206

Gazania ringens, circumnutation of stem, 208 Genera containing sleeping

plants, 320, 321

Geotropism, 5; effect of, on the primary radicle, 196; the reverse of

apogeotropism, 512: effect on the tips of radicles, 543

Geranium cinereum, 304

— Endressii, 304

— Ibericum, nocturnal movement of cotyledons, 298

— Richardsoni, 304

— rotundifolium, nocturnal movement of cotyledon, 304, 312

— subcaulescens, 304

Germinating seed, history of a, 548

Githago segetum, circumnutation of hypocotyl, 21, 108

—, burying of hypocotyl, 109

—, seedlings feebly illuminated, 124, 128

—, sleep of cotyledon, 302

—, — leaves 321

Glaucium luteum, circumnutation of young leaves, 228

Gleditschia, sleep of leaves, 368

Glycine hispida, vertical sinking of leaflets, 366

Glycyrrhiza, leaflets depressed at night, 355

Godlewski, Emil, on the turgescence of the cells, 485

Gooseberry, effect of radiation, 284

Gossypium (var. Nankin cotton), circumnutation of hypocotyl, 22

—, movement of cotyledon, 22, 23

—, sleep of leaves, 324

—, arboreum (?), sleep of cotyledons, 303

—, Braziliense, nocturnal movement of leaves, 324

—, sleep of cotyledons, 303

— herbaceum, sensitiveness of apex of radicle, 168

—, radicles cauterised transversely, 537

— maritimum, nocturnal movement of leaves, 324

Gravitation, movements excited by, 567

Gray, Asa, on Delphinium nudicaule, 80; on Megarrhiza Californica, 81; on

the movements in the fruiting fronds of Aesplenium trichomanes, 257; on the

Amphicarpoea monoica, 520; on the Ipomœa Jalappa, 557

Grease, effect of, on radicles and their tips, 182, 185

Gressner, Dr. H., on the cotyledons of Cyclamen Persicum, 46, 77; on

hypocotyl of the same, 96

Gymnosperms, 389

Haberlandt, Dr., on the protuberance on the hypocotyl of Allium, 59; the

importance of the arch to seedling plants, 87; sub-aërial and subterranean

cotyledons, 110, n.; the arched hypocotyl, 554

Haematoxylon Campechianum, nocturnal movement of leaves, 368, 369

Hedera helix, circumnutation of stem, 207

Hedysarum coronarium, nocturnal movements of leaves, 356

Helianthemum prostratum, geotropic movement of flower-heads, 518

Helianthus annuus, circumnutation of hypocotyl, 45

—, arching of hypocotyl, 90

—, nocturnal movement of cotyledons, 305

Heliotropism, 5; uses of, 449; a modified form of circumnutation, 490

Helleborus niger, mode of breaking through the ground, 86

Hensen, Prof., on roots in worm-burrows, 72

Henslow, Rev. G., on the cotyledons of Phalaris Canariensis, 62

Hofmeister, on the curious movement of Spirogyra, 3, 259, n.; of the leaves

of Pistia stratiotes, 255; of cotyledons at night, 297; of petals, 414

— and Batalin on the movements of the cabbage, 229

Hooker, Sir J., on the effect of light on the pitchers of Sarracenia, 450

Hypocotyl, 5; manner of breaking through the ground, 77; emerges under the

form of an arch, 553

Hypocotyls and Epicotyls, circumnutation and other movements when arched,

98; power of straightening themselves, 100; rupture of the seed-coats,

102–106; illustration of, 106; circumnutation when erect, 107; when in

dark, 108

Hyponasty, 6, 267

Iberis umbellata, movement of stem, 202.

Illumination, effect of, on the sleep of leaves, 398

Imatophyllum vel Clivia (sp.?), movement of leaves, 255

Indigofera tinctoria, leaflets depressed at night, 354

Inheritance in plants, 407, 491

Insectivorous and climbing plants not heliotropic, 450; influence of light

on, 488

Ipomœa bona nox, arching of hypocotyl, 90

—, nocturnal position of cotyledons, 306, 312

— coerulea vel Pharbitis nil, circumnutation of seedlings, 47

—, movement of cotyledons, 47–49, 109

—, nocturnal movements of cotyledons, 305

—, sleep of leaves, 386

—, sensitiveness to light, 451

—, the hypocotyledonous stems heliotropic, 453

— coccinea, position of cotyledons at night, 306, 312

— leptophylla, mode of breaking through the ground, 83, 84

—, arching of the petioles of the cotyledons, 90

—, difference in sensitiveness to gravitation in different parts, 509

—, extraordinary manner of germination, 557

Ipomœa pandurata, manner of germination, 84, 557

— purpurea (vel Pharbitis hispida), nocturnal movement of cotyledons, 305,

312

—, sleep of leaves, 386

—, sensitiveness to light, 451

—, the hypocotyledonous stems heliotropic, 453

Iris pseudo-acorus, circumnutation of leaves, 253

Irmisch, on cotyledons of Ranunculus Ficaria, 96

Ivy, its stems heliotropic, 451

Kerner on the bending down of peduncles, 414

Klinostat, the, an instrument devised by Sachs to eliminate geotropism, 93

Kraus, Dr. Carl, on the underground shoots of Triticum repens, 189; on

Cannabis sativa, 250, 307, 312; on the movements of leaves, 318

Lactuca scariola, sleep of cotyledons, 305

Lagenaria vulgaris, circumnutation of seedlings, 42

—, of cotyledons, 43

—, cotyledons vertical at night, 304

Lathraea squamaria, mode of breaking through the ground, 85

—, quantity of water secreted, 85, 86, n.

Lathyrus nissolia, circumnutation of stem of young seedling, 33

—, ellipses described by, 107, 108

Leaves, circumnutation of, 226–262; dicotyledons, 226–252; monocotyledons,

252–257; nyctitropism of, 280; their temperature affected by their position

at night, 294; nyctitropic or sleep movements, 315, 394; periodicity of

their movements inherited, 407; embryology of, 414; so-called diurnal

sleep, 445

Leguminosae, sleep of cotyledons, 308; sleeping species, 340

Le Maout and Decaisne, 67

Lepidium sativum, sleep of cotyledons, 302

Light, movements excited by 418, 563; influence on most vegetable tissues,

486; acts on plant as on the nervous system of animals, 487

Lilium auratum, circumnutation of stem, 212

—, apogeotropic movement of stem, 498, 499

Linnæus, ‘Somnus Plantarum’, 280; on plants sleeping, 320; on the leaves

of Sida abutilon, 324; on Œnothera mollissima, 383

Linum Berendieri, nocturnal movement of cotyledons, 298

— usitatissimum, circumnutation of stem, 203

Lolium perenne, joints affected by apogeotropism, 502

Lonicera brachypoda, hooking of the tip, 272

—, sensitiveness to light, 453

Loomis, Mr., on the movements in the fruiting fronds of Asplenium

trichomanes, 257

Lotus aristata, effect of radiation on leaves, 292

— Creticus, leaves awake and asleep, 354

— Gebelii, nocturnal movement of cotyledons, 308

—, leaflets provided with pulvini, 353

— Jacobæus, movements of cotyledons, 35, 109

—, pulvini of, 115

Lotus Jacobæus, movements at night, 116, 121, 124

—, development of pulvini, 122

—, sleep of cotyledons, 308, 313

—, nyctitropic movement of leaves, 353

— major, sleep of leaves, 353

— perigrinus, movement of leaflets, 353

Lunularia vulgaris, circumnutation of fronds, 258

Lupinus, 340

— albifrons, sleep of leaves, 344

— Hartwegii, sleep of leaves, 341

— luteus, circumnutation of cotyledons, 38, 110

—, effect of darkness, 124

Lupinus, position of leaves when asleep, 341

—, different positions of leaves at night, 343

—, varied movements of leaves and leaflets, 395

— Menziesii, sleep of leaves, 343

— mutabilis, sleep of leaves, 343

— nanus, sleep of leaves, 343

— pilosus, sleep of leaves, 340, 341

— polyphyllus, sleep of leaves, 343

— pubescens, sleep of leaves by day and night, 342

—, position of petioles at night, 343

—, movements of petioles, 401

— speciosus, circumnutation of leaves, 236

Lynch, Mr. R., on Pachira aquatica, 95, n.; sleep movements of Averrhoa,

330

Maranta arundinacea, nyctitropic movement of leaves, 389–391

—, after much agitation do not sleep, 319

Marsilia quadrifoliata, effect of radiation at night, 292

—, circumnutation and nyctitropic movement of leaflets, 392–394

—, rate of movement, 404

Martins, on radiation at night, 284, n.

Masters, Dr. Maxwell, on the leading shoots of the Coniferæ, 211

Maurandia semperflorens, circumnutation of peduncle, 225

Medicago maculata, nocturnal position of leaves, 345

— marina, leaves awake and asleep, 344

Meehan, Mr., on the effect of an Æcidium on Portulaca oleracea, 189

Megarrhiza Californica, mode of breaking through the ground, 81

—, germination described by Asa Gray, 82

—, singular manner of germination, 83, 556

Melaleuca ericaefolia, sleep of leaves, 383

Melilotus, sleep of leaves, 345

— alba, sleep of leaves, 347

— coerulea, sleep of leaves, 347

— dentata, effect of radiation at night, 295

— elegans, sleep of leaves, 347

— gracilis, sleep of leaves, 347

— infesta, sleep of leaves, 347

— Italica, leaves exposed at night, 291

—, sleep of leaves, 347

— macrorrhiza, leaves exposed at night, 292

—, sleep of leaves, 347

— messanensis, sleep of leaves on full-grown and young plants, 348, 416

— officinalis, effect of exposure of leaves at night, 290, 296

—, nocturnal movement of leaves, 346, 347

—, circumnutation of leaves, 348

—, movement of petioles, 401

Melilotus parviflora, sleep of leaves, 347

— Petitpierreana, leaves exposed at night, 291, 296

—, sleep of leaves, 347

— secundiflora, sleep of leaves, 347

— suaveolens, leaves exposed at night, 291

—, sleep of leaves, 347

— sulcata, sleep of leaves, 347

— Taurica, leaves exposed at night, 291

—, sleep of leaves, 347, 415

Methods of observation, 6

Mimosa albida, cotyledons vertical at night, 116

—, not sensitive to contact, 127

—, sleep of cotyledons, 308

—, rudimentary leaflets, 364

—, nyctitropic movements of leaves, 379, 380

—, circumnutation of the main petiole of young leaf, 381

—, torsion, or rotation of leaves and leaflets, 400

—, first true leaf, 416

—, effect of bright sunshine on basal leaflets, 445

— marginata, nyctitropic movements of leaflets, 381

— pudica, movement of cotyledons, 105

—, rupture of the seed-coats, 105

—, circumnutation of cotyledons, 109

—, pulvini of, 113, 115

—, cotyledons vertical at night, 116

—, hardly sensitive to contact, 127

—, effect of exposure at night, 293

—, nocturnal movement of leaves, 297

—, sleep of cotyledons, 308

—, circumnutation and nyctitropic movement of main petiole, 374–378

—, of leaflets, 378

Mimosa albida, circumnutation and nyctitropic movement of pinnae, 402

—, number of ellipses described in given time, 406

—, effect of bright sunshine on leaflets, 446

Mirabilis jalapa and longiflora, nocturnal movements of cotyledons, 307

—, nyctitropic movement of leaves, 387

Mohl, on heliotropism in tendrils, stems, and twining plants, 451

Momentum-like movement, the accumulated effects of apogeotropism, 508

Monocotyledons, sleep of leaves, 389

Monotropa hypopitys, mode of breaking through the ground, 86

Morren, on the movements of stamens of Sparmannia and Cereus, 226

Müller, Fritz, on Cassia tora, 34; on the circumnutation of Linum

usitatissimum, 203; movements of the flower-stems of an Alisma, 226

Mutisia clematis, movement of leaves, 246

—, leaves not heliotropic, 451

Natural selection in connection with geotropism, heliotropism, etc., 570

Nephrodium molle, circumnutation of very young frond, 66

—, of older frond, 257

—, slight movement of fronds, 509

Neptunia oleracea, sensitiveness to contact, 128

—, nyctitropic movement of leaflets, 374

—, of pinnae, 402

Nicotiana glauca, sleep of leaves, 385, 386

—, circumnutation of leaves, 386

Nobbe, on the rupture of the seed-coats in a seedling of Martynia, 105

Nolana prostrata, movement of seedlings in the dark, 50

—, circumnutation of seedling, 108

Nyctitropic movement of leaves, 560

Nyctitropism, or sleep of leaves, 281; in connection with radiation, 286;

object gained by it, 413

Observation, methods of, 6

Œnothera mollissima, sleep of leaves, 383

Opuntia basilaris, conjoint circumnutation of hypocotyl and cotyledon, 44

—, thickening of the hypocotyl, 96

—, circumnutation of hypocotyl when erect, 107

—, burying of, 109

Orange, seedling, circumnutation of, 510

Orchis pyramidalis, complex movement of pollinia, 489

Oxalis acetosella, circumnutation of flower-stem, 224

—, effects of exposure to radiation at night, 287, 288, 296

—, circumnutation and nyctitropic movement in full-grown leaf, 326

—, circumnutation of leaflet when asleep, 327

—, rate of circumnutation of leaflets, 404

—, effect of sunshine on leaflets, 447

—, circumnutation of peduncle, 506

Oxalis acetosella, seed-capsules, only occasionally buried, 518

— articulata, nocturnal movements of cotyledons, 307

— (Biophytum) sensitiva, rapidity of movement of cotyledons during the

day, 26

—, pulvinus of, 113

—, cotyledons vertical at night, 116, 118

— bupleurifolia, circumnutation of foliaceous petiole, 328

—, nyctitropic movement of terminal leaflet, 329

— carnosa, circumnutation of flower-stem, 223

—, epinastic movements of flower-stem, 504

—, effect of exposure at night, 288, 296

—, movements of the flower-peduncles due to apogeotropism and other

forces, 503–506

— corniculata (var. cuprea), movements of cotyledons, 26

—, rising of cotyledons, 116

—, rudimentary pulvini of cotyledons, 119

—, development of pulvinus, 122

—, effect of dull light, 124

—, experiments on leaves at night, 288

— floribunda, pulvinus of cotyledons, 114

—, nocturnal movement, 118, 307, 313

— fragrans, sleep of leaves, 324

— Ortegesii, circumnutation of flower-stems, 224

—, sleep of large leaves, 327

—, diameter of plant at night, 402

—, large leaflets affected by bright sunshine, 447

— Plumierii, sleep of leaves, 327

— purpurea, exposure of leaflets at night, 293

— rosea, circumnutation of cotyledons, 23, 24

Oxalis rosea, pulvinus of, 113

—, movement of cotyledons at night, 117, 118, 307

—, effect of dull light, 124

—, non-sensitive cotyledons, 127

— sensitiva, movement of cotyledons, 109, 127, 128

—, circumnutation of flower-stem, 224

—, nocturnal movement of cotyledons, 307, 312

—, sleep of leaves, 327

— tropoeoloides, movement of cotyledons at night, 118, 120

— Valdiviana, conjoint circumnutation of cotyledons and hypocotyl, 25

—, cotyledons rising vertically at night, 114, 115, 117, 118

—, non-sensitive cotyledons, 127

—, nocturnal movement of cotyledon, 307, 312

—, sleep of leaves and not of cotyledons, 315

—, movements of leaves, 327

Pachira aquatica, unequal cotyledons, 95, n.

Pancratium littorale, movement of leaves, 255

Paraheliotropism, or diurnal sleep of leaves, 445

Passiflora gracilis, circumnutation and nyctitropic movement of leaves,

383, 384

—, apogeotropic movement of tendrils, 510

—, sensitiveness of tendrils, 550

Pelargonium zonale, circumnutation of stem, 203

—, and downward movement of young leaf, 232, 233, 269

Petioles, the rising of beneficial to plant at night, 402

Petunia violacea, downward movement and circumnutation of very young leaf,

248, 249, 269.

Pfeffer, Prof., on the turgescence of the cells, 2; on pulvini of leaves,

113, 117; sleep movements of leaves, 280, 283, 284; nocturnal rising of

leaves of Malva, 324; movements of leaflets in Desmodium gyrans, 358; on

Phyllanthus Niruri, 388; influence of a pulvinus on leaves, 396; periodic

movements of sleeping leaves, 407, 408; movements of petals, 414; effect of

bright sunshine on leaflets of Robinia, 445; effect of light on parts

provided with pulvini, 363

Phalaris Canariensis, movements of old seedlings, 62

—, circumnutation of cotyledons, 63, 64, 108

—, heliotropic movement and circumnutation of cotyledon towards a dim

lateral light, 427

—, sensitiveness of cotyledon to light, 455

—, effect of exclusion of light from tips of cotyledons, 456

—, manner of bending towards light, 457

—, effects of painting with Indian ink, 467

—, transmitted effects of light, 469

—, lateral illumination of tip, 470

—, apogeotropic movement of the sheath-like cotyledons, 497

—, change from a straight upward apogeotropic course to circumnutation,

499

—, apogeotropic movement of cotyledons, 500

Phaseolus Hernandesii, nocturnal movement of leaves and leaflets, 368

—, caracalla, 93

—, nocturnal movement of leaves, 368

—, effect of bright sunshine on leaflets, 446

Phaseolus multiflorus, movement of radicles, 29

—, of young radicle, 72

—, of hypocotyl, 91, 93

—, sensitiveness of apex of radicle, 163–167

—, to moist air, 181

—, cauterisation and grease on the tips, 535

—, nocturnal movement of leaves, 368

—, nyctitropic movement of the first unifoliate leaves, 397

— Roxburghii, effect of bright sunshine on first leaves, 445

—, vulgaris, 93

—, sleep of leaves, 318

—, vertical sinking of leaflets at night, 368

Phyllanthus Niruri, sleep of leaflets, 388

— linoides, sleep of leaves, 387

Pilocereus Houlletii, rudimentary cotyledons, 97

Pimelia spectabilis, sleep of leaves, 387

Pincers, wooden, through which the radicle of a bean was allowed to grow,

Pinus austriaca, circumnutation of leaves, 251, 252

— Nordmanniana, nyctitropic movement of leaves, 389

— pinaster, circumnutation of hypocotyl, 56

—, movement of two opposite cotyledons, 57

—, circumnutation of young leaf, 250, 251

—, epinastic downward movement of young leaf, 270

Pistia stratiotes, movement of leaves, 255

Pisum sativum, sensitiveness of apex of radicle, 158

—, tips of radicles cauterised transversely, 534

Plants, sensitiveness to light, 449; hygroscopic movements of, 489

Plants, climbing, circumnutation of, 264; movements of, 559

—, mature, circumnutation of, 201–214

Pliny on the sleep-movements of plants, 280

Plumbago Capensis, circumnutation of stem, 208, 209

Poinciana Gilliesii, sleep of leaves, 368

Polygonum aviculare, leaves vertical at night, 387

— convolvulus, sinking of the leaves at night, 318

Pontederia (sp.?), circumnutation of leaves, 256

Porlieria hygrometrica, circumnutation and nyctitropic movements of petiole

of leaf, 335, 336

—, effect of watering, 336–338

—, leaflets closed during the day, 413

Portulaca oleracea, effect of Æcidium on, 189

Primula Sinensis, conjoint circumnutation of hypocotyl and cotyledon, 45,

Pringsheim on the injury to chlorophyll, 446

Prosopis, nyctitropic movements of leaflets, 374

Psoralea acaulis, nocturnal movements of leaflets, 354

Pteris aquilina, rachis of, 86

Pulvini, or joints; of cotyledons, 112–122; influence of, on the movements

of cotyledons, 313; effect on nyctitropic movements, 396

Quercus (American sp.), circumnutation of young stem, 53, 54

— robur, movement of radicles, 54, 55

—, sensitiveness of apex of radicle, 174–176

Quercus virens, manner of germination, 85, 557

Radiation at night, effect of, on leaves, 284–286

Radicles, manner in which they penetrate the ground, 69–77; circumnutation

of 69; experiments with split sticks, 74; with wooden pincers, 75;

sensitiveness of apex to contact and other irritants, 129; of Vicia faba,

132–158; various experiments, 135–140; summary of results, 143–151; power

of an irritant on, compared with geotropism, 151–154; sensitiveness of tip

to moist air, 180; with greased tips, 185; effect of killing or injuring

the primary radicle, 187–191; curvature of, 193; affected by moisture, 198;

tip alone sensitive to geotropism, 540; protrusion and circumnutation in a

germinating seed, 548; tip highly sensitive, 550; the tip acts like the

brain of one of the lower animals, 573

—, secondary, sensitiveness of the tips in the bean, 154; become

vertically geotropic, 186–191

Ramey on the movements of the cotyledons of Mimosa pudica, and Clianthus

Dampieri at night, 297

Ranunculus Ficaria, mode of breaking through the ground, 86, 90

—, single cotyledon, 96

—, effect of lateral light, 484

Raphanus sativa, sensitiveness of apex of radicle, 171

—, sleep of cotyledons, 301

Rattan, Mr., on the germination of the seeds of Megarrhiza Californica, 82

Relation between circumnutation and heliotropism, 435

Reseda odorata, hypocotyl of seedling slightly heliotropic, 454

Reversion, due to mutilation, 190

Rhipsalis cassytha, rudimentary cotyledons, 97

Ricinus Borboniensis, circumnutation of arched hypocotyl, 53

Robinia, effect of bright sunshine on its leaves, 445

— pseudo-acacia, leaflets vertical at night, 355

Rodier, M., on the movements of Ceratophyllum demersum, 211

Royer, Ch., on the sleep-movements of plants, 281, n.; on the sleep of

leaves, 318; the leaves of Medicago maculata, 345; on Wistaria Sinensis,

354

Rubus idæus (hybrid) circumnutation of stem, 205

—, apogeotropic movement of stem, 498

Ruiz and Pavon, on Porlieria hygrometrica, 336

SACHS on “revolving nutation,” 1; intimate connection between turgescence

and growth, 2, n.; cotyledon of the onion, 59; adaptation of root-hairs,

69; the movement of the radicle, 70, 72, 73; movement in the hypocotyls of

the bean, etc., 91; sensitiveness of radicles, 131, 145, 198;

sensitiveness of the primary radicle in the bean, 155; in the common pea,

156; effect of moist air, 180; of killing or injuring the primary radicle,

186, 187; circumnutation of flower-stems, 225; epinasty, 268; movements of

leaflets of Trifolium incarnatum, 350; action of light in modifying the

periodic movements of leaves, 418; on geotropism and heliotropism, 436,

n.; on Tropaeolum majus, 453; on the hypocotyls slightly heliotropic, and

stems strongly apheliotropic of the ivy, 453; heliotropism of radicles,

482; experiments on tips of radicles of bean, 523, 524; curvature of the

hypocotyl, 555; resemblance between plants and animals, 571

Sarracenia purpurea, circumnutation of young pitcher, 227

Saxifraga sarmentosa, circumn utation of an inclined stolon, 218

Schrankia aculeata, nyctitropic movement of the pinnae, 381, 403

— uncinata, nyctitropic movements of leaflets, 381

Securigera coronilla, nocturnal movements of leaflets, 352

Seed-capsules, burying of, 513

Seed-coats, rupture of, 102–106

Seedling plants, circumnutating movements of, 10

Selaginella, circumnutation of 258

— Kraussii (?), circumnutation of young plant, 66

Sida napoea, depression of leaves at night, 322

—, no pulvinus, 322

— retusa, vertical rising of leaves, 322

— rhombifolia, sleep of cotyledons, 308

—, sleep of leaves, 314

—, vertical rising of leaves, 322

—, no pulvinus, 322

—, circumnutation and nyctitropic movements of leaf of young plant, 322

—, nyctitropic movement of leaves, 397

Siegesbeckia orientalis, sleep of leaves, 319, 384

Sinapis alba, hypocotyl bending towards the light, 461

—, transmitted effect of light on radicles, 482, 483, 567

—, growth of radicles in darkness, 486

Sinapis nigra, sleep of cotyledons, 301

Smilax aspera, tendrils apheliotropic, 451

Smithia Pfundii, non-sensitive cotyledons, 127

—, hyponastic movement of the curved summit of the stem, 274–276

—, cotyledons not sleeping at night, 308

—, vertical movement of leaves, 356

— sensitiva, sensitiveness of cotyledons to contact, 126

—, sleep of cotyledons, 308

Sophora chrysophylla, leaflets rise at night, 368

Solanum dulcamara, circumnutating stems, 266

— lycopersicum, movement of hypocotyl, 50

—, of cotyledons, 50

—, effect of darkness, 124

—, rising of cotyledons at night, 306

—, heliotropic movements of hypocotyl, 421

—, effect of an intermittent light, 457

—, rapid heliotropism, 461

— palinacanthum, circumnutation of arched hypocotyl, 51, 100

—, of cotyledon, 51

—, ellipses described by hypocotyl when erect, 107

—, nocturnal movement of cotyledons, 306

Sparganium ramosum, rhizomes of, 189

Sphaerophysa salsola, rising of leaflets, 355

Spirogyra princeps, movements of, 259, n.

Stahl, Dr., on the effect of Æcidium on shoot, 189; on the influence of

light on swarm-spores, 488, n.

Stapelia sarpedon, circumnutation of hypocotyl, 46, 47

—, minute cotyledons, 97

Stellaria media, nocturnal movement of leaves, 297

Stems, circumnutation of, 201–214

Stolons, or Runners, circumnutation of, 214–222, 558

Strasburger, on the effect of light on spores of Haematococcus, 455, n.;

the influence of light on the swarm-spores, 488.

Strawberry, stolons of the, circumnutate, but not affected by moderate

light, 454

Strephium floribundum, circumnutation and nyctitropic movement of leaves,

391, 392

Tamarindus Indica, nyctitropic movement of leaflets, 374

Transversal–heliotropismus (of Frank) or diaheliotropism, 438

Trapa natans, unequal cotyledons, 95, n.

Tecoma radicans, stems apheliotropic, 451

Tephrosia caribaea, 354

Terminology, 5

Thalia dealbata, sleep of leaves, 389

—, lateral movement of leaves, 404

Trichosanthes anguina, action of the peg on the radicle, 104

—, nocturnal movement of cotyledons, 304

Trifolium, position of terminal leaflets at night, 282

— globosum, with hairs protecting the seed-bearing flowers, 517

— glomeratum, movement of cotyledons, 309

— incarnatum, movement of cotyledons, 309

— Pannonicum, shape of first true leaf, 350, 415

Trifolium pratense, leaves exposed at night, 293

— repens, circumnutation of flower-stem, 225

—, circumnutating and epinastic movements of flower-stem, 276–279

—, nyctitropic movement of leaves, 349

—, circumnutation and nyctitropic movements of terminal leaflets, 352, 353

—, sleep movements, 349

— resupinatum, no pulvini to cotyledons, 118

—, circumnutation of stem, 204

—, effect of exposure at night, 295

—, cotyledons not rising at night, 118, 309

—, circumnutation and nyctitropic movements of terminal leaflets, 351, 352

— strictum, movements of cotyledons at night, 116, 118

—, nocturnal and diurnal movements of cotyledons, 309–311, 313

—, movement of the left-hand cotyledon, 316

— subterraneum, movement of flower-heads, 71

—, of cotyledons at night, 116, 118, 309

—, circumnutation of flower-stem, 224, 225

—, circumnutation and nyctitropic movements of leaves, 350

—, number of ellipses in 24 hours, 405

—, burying its flower-heads, 513, 514

—, downward movement of peduncle, 515

—, circumnutating movement of peduncle, 516

Trigonella Cretica, sleep of leaves, 345

Triticum repens, underground shoots of, become apogeotropic, 189

Triticum vulgare, sensitiveness of tips of radicle to moist air, 184

Tropaeolum majus (?), sensitiveness of apex of radicle to contact, 167

—, circumnutation of stem, 204

—, influence of illumination on nyctitropic movements, 338–340, 344

—, heliotropic movement and circumnutation of epicotyl of a young

seedling, 428, 429

—, of an old internode towards a lateral light, 430

—, stems of very young plants highly heliotropic, of old plants slightly

apheliotropic, 453

—, effect of lateral light, 484

— minus (?), circumnutation of buried and arched epicotyl, 27

Ulex, or gorse, first-formed leaf of, 415

Uraria lagopus, vertical sinking of leaflets at night, 365

Vaucher, on the burying of the flower-heads of Trifolium subterraneum, 513;

on the protection of seeds, 517

Verbena melindres (?), circumnutation of stem, 210

—, apogeotropic movement of stem, 495

Vicia faba, circumnutation of radicle, 29, 30

—, of epicotyl, 31–33

—, curvature of hypocotyl, 92

—, sensitiveness of apex of radicle, 132–134

—, of the tips of secondary radicles, 154

—, of the primary radicle above the apex, 155–158

—, various experiments, 135–143

—, summary of results, 143–151

—, power of an irritant on, compared with that of geotropism, 151–154

Vicia faba, circumnutation of leaves, 233–235

—, circumnutation of terminal leaflet, 235

—, effect of apogeotropism, 444

—, effect of amputating the tips of radicles, 523

—, regeneration of tips, 526

—, short exposure to geotropic action, 527

—, effects of amputating the tips obliquely, 528

—, of cauterising the tips, 529

—, of grease on the tips, 534

Vines, Mr., on cell growth, 3

Vries, De, on turgescence, 2; on epinasty and hyponasty, 6, 267, 268; the

protection of hypocotyls during winter, 557; stolons apheliotropic, 108;

the nyctitropic movement of leaves, 283; the position of leaves influenced

by epinasty, their own weight and apogeotropism, 440; apogeotropism in

petioles and midribs, 443; the stolons of strawberries, 454; the joints or

pulvini of the Gramineæ, 502

Watering, effect of, on Porlieria hygrometrica, 336–338

Wells, ‘Essay on Dew,’ 284, n.

Wiesner, Prof., on the circumnutation of the hypocotyl, 99, 100; on the

hooked tip of climbing stems, 272; observations on the effect of bright

sunshine on chlorophyll in leaves, 446; the effects of an intermittent

light, 457; on aërial roots, 486; on special adaptations, 490

Wigandia, movement of leaves, 248

Williamson, Prof., on leaves of Drosera Capensis, 414

Wilson, Mr. A. S., on the movements of Swedish turnip leaves, 230, 298

Winkler on the protection of seedlings, 108

Wistaria Sinensis, leaflets depressed at night, 354

—, circumnutation with lateral light, 452

Zea mays, circumnutation of cotyledon, 64

Zea mays, geotropic movement of radicles, 65

—, sensitiveness of apex of radicle to contact, 177–179

—, secondary radicles, 179

—, heliotropic movements of seedling, 64, 421

—, tips of radicles cauterised, 539

Zukal, on the movements of Spirulina, 259, n.