1. The organs are sensitive to the stimulus of gravity and the periodic movements are brought about by variation of geotropic curvature under change of temperature.
2. The movement is not confined to growing organs, but is also exhibited by organs which are fully grown and even by rigid trees.
3. The periodic movement is closely related to the diurnal variation of temperature. Fall of temperature from thermal-noon (about 2 p.m.) to thermal-dawn (about 6 a.m.) is attended by a movement of erection; rise of temperature from thermal-dawn to thermal-noon is followed, on the other hand, by a reverse movement of fall.
That the movement is primarily due to variation of temperature will be demonstrated in two different ways:
(_a_) by the change of normal rhythm of movement by artificial transpositions of periods of maximum and minimum temperature, and
(_b_) by the abolition of periodic movement through maintenance of constant temperature.
That the phenomenon is not nastic, but paratonic will be demonstrated:--
(_a_) by the reversal of closure into opening movement and _vice versa_, in consequence of inversion of the plant upside down, and
(_b_) by the diurnal variation of torsional movement, the direction of which is dependent on the directive action of the stimulus of gravity.
I shall now describe the diurnal movement of various geotropically curved plant-organs; the most striking example of this is furnished by the "Praying" Palm of Faridpore, already described. I shall here recapitulate some of the important features connected with the phenomenon.
DIURNAL MOVEMENT OF PALM TREES.
Movements similar to that of the Faridpore Palm (p. 12) are found in other Palm trees growing at an inclination from the vertical. I reproduce once more the diurnal curve given by the Sijberia Palm together with the curve of daily thermal variation (Fig. 199). It will be seen that the two curves resemble each other so closely that the curve of movement of the tree is practically a replica of the thermographic record. There can therefore be no doubt of the movement being brought about by variation of temperature; rise of temperature is attended by the movement of fall of the tree and _vice versa_. The record was commenced at noon; the temperature rose till the maximum was reached at about 3 p.m. and the tree also reached its lowest position at 3-45 p.m., the lag being 45 minutes. The temperature fell continuously after the maximum at 3 p.m., to the minimum at 6 a.m. next morning. In response to the falling temperature, the tree exhibited a movement of erection. The temperature rose after 6 a.m. and the movement of the tree became reversed from ascent to descent.
[Ill.u.s.tration: FIG. 199.--Diurnal record of the Sijberia Palm. Upper curve gives variation of temperature, and the lower curve the movement of the tree.]
I have already shown: (1) that the diurnal movement just described is due to physiological reaction, and that the movement is abolished at the death of the plant; (2) that light has little or no effect, since the thick bark and bases of leaves screen the living tissue from the action of light; (3) that transpiration has practically no effect on the periodic movement, since such movement takes place in other plants completely immersed under water; thus _Ipomoea aquatica_, a water plant, kept under water, gave the normal diurnal curve similar to that of the palm. The modifying effect of transpiration was in this case, completely excluded. I obtained similar effect with geotropically curved stem of _Basella cordifolia_ (p. 25); (4) that the weight of the plant-organ as such, has little effect on the diurnal curve, since an inverted plant continues for a few days to exhibit the periodic movement, in spite of the antagonistic effect of weight. A different experiment will be described (see p. 582) where the effect of weight was completely neutralised and the plant-organ gave, nevertheless, the normal diurnal curve.
[Ill.u.s.tration: FIG. 200.--Diurnal record of inclined palm tree, of geotropically curved proc.u.mbent stem of _Tropaeolum_ and the dia-geotropic leaf of palm. Note general similarity between diurnal curve of plants and the thermographic record.]
I have also shown that the diurnal movement is determined by the modifying influence of temperature on geotropic curvature. Rise of temperature opposes or neutralises the geotropic curvature; fall of temperature, on the other hand, accentuates it. The particular diurnal movement was not confined to the palm trees, but was exhibited by all plant-organs subjected to the stimulus of gravity.
DIURNAL MOVEMENT OF PROc.u.mBENT STEMS AND OF LEAVES.
[Ill.u.s.tration: FIG. 201.---Diurnal records of leaves of _Dahlia_, _Papaya_ and _Croton_.]
_Experiment 210._--In order to demonstrate the continuity of the phenomenon of diurnal movement I took various stems growing in water or land for my experiment. The plants were laid horizontally, till the stems bent up and a.s.sumed the stable position of geotropic equilibrium.
In figure 200 is given records of the inclined palm tree, of proc.u.mbent stem of _Tropaeolum_, and the leaf of the palm tree. The very close relation between the temperature-variation and the movement of different plant-organs is sufficiently obvious.
I shall next give a series of diurnal records of leaves of different plants such as those of _Dahlia_, _Papaya_ and _Croton_ (Fig. 201). In these also fall of temperature induces an up-movement while rise of temperature causes a fall of the leaf. I shall presently refer to the "personal equation" by which the record of one plant is distinguished from another.
CONTINUOUS DIURNAL RECORD FOR SUCCESSIVE THERMAL NOONS.
_Experiment 211._--The diurnal record given above, was taken from ordinary noon at 12 o"clock to noon next day. The diurnal curve becomes much simplified if the record be taken from _thermal-noon_ (at about 2 p.m.) to the thermal noon next day. The plant-organ becomes erected during falling temperature from thermal-noon to thermal-dawn next morning, and undergoes a fall during rise of temperature from thermal-dawn to thermal-noon. The subsequent diurnal records will therefore be given for 24 hours commencing with 2 p.m. In figure 202 is given diurnal records of geotropically curved stem of _Tropaeolum_ and the leaf of _Dahlia_ for two days in succession.
The thermal record shows that there was a continuous fall of temperature from thermal-noon at 2 p.m. to the thermal-dawn at 6 a.m. next morning, that is to say, for 16 hours. Rise of temperature through the same range occurred in 8 hours from 6 a.m. till 2 p.m. The average rate of rise of temperature was thus twice as quick as that of fall. This is clearly seen from the slopes of thermal curve during thermal ascent and descent.
The record of the movement of the plant shows a striking parallelism; the different plant-organs became erected from thermal-noon to thermal-dawn, and underwent a fall from thermal-dawn to thermal-noon.
The descent of the curve is, as in the case of thermal curve, relatively more abrupt. The records on two successive days are very similar, the slight difference being due to the physiological depression consequent on prolonged maintenance of the plants in a closed chamber.
[Ill.u.s.tration: FIG. 202.--Diurnal curve of the proc.u.mbent stem of _Tropaeolum majus_, and the leaf of _Dahlia_ for two successive days. In the thermographic record the up-curve represents fall, and down-curve rise of temperature.]
MODIFICATION OF THE DIURNAL CURVE.
I shall now proceed to explain the modifications that may occur in the standard thermo-geotropic curve.
_Turning points._--In the bulky Palm, the reversal of movement from fall to rise or _vice versa_ takes place about an hour after the thermal inversion. This lag is partly due to the time taken by a ma.s.s of tissue to a.s.sume the temperature of the surrounding air. There is, moreover, the question of physiological inertia which delays the reaction. In leaves this lag may be considerably less or even absent. In certain cases the reversal of movement may take place a little earlier than the temperature inversion. It should be remembered in this connection, that in response to temperature change, the leaf is often displaced to a considerable extent from its "mean position of equilibrium"; moreover the force of recovery is greatest at the two extreme positions. These considerations probably explain the quick return of the leaf to equilibrium position. The slow autonomous movement of the leaf may sometimes prove to be a contributory factor.
_Effect of irregular fluctuation of temperature._--In settled weather the diurnal rise and fall of temperature is very regular. But under less settled condition, owing to the change of direction of the wind, the temperature curve shows one or more fluctuations, specially in the forenoon. It was a matter of surprise to me to find the plant-record repeating the fluctuations of thermal record with astonishing fidelity.
This common twitch in the two records is seen in the record of the Sijberia Palm (Fig. 199). Certain plants are extremely sensitive to variation of temperature; so much so that these physiological indicators of thermal variation are far more delicate than ordinary thermometers.
_Effect of restricted pliability of the organ._--A leaf is more pliable in one direction than in the other. The pulvinus of _Mimosa_, for example, allows a greater amount of bending downwards than upwards; in consequence of this the leaf in its fall becomes almost parallel to the internode below; the up-movement is, however, far more restricted. The leaf in its most erect position still makes a considerable angle with the internode of the stem above it. If the leaf-stalk of a plant be restricted in its rise the erectile movement at night will reach a limit, and the top of the curve will remain flat. This is seen, ill.u.s.trated in the record of the leaf of _Croton_ (Fig. 202), which attains its maximum erection at 9 p.m. and the subsequent curve remains flattened till 7 a.m.; after this the leaf begins to execute its downward movement. In other cases, the range of up-movement is very great and the plant-organ erects itself continuously till morning. In certain cases the impulse of up-movement carries the organ beyond the stable position of equilibrium; after this the leaf begins to retrace its path slowly; the down-movement due to rise of temperature is, however, far more abrupt, and easily distinguishable from the previous slow return.
It will thus be seen that though the diurnal record consists of an alternating up and down curve, yet these minor characteristics or "personal equation" of the plant confers on the record a certain stamp of individuality.
_Effect of age._--In the floral leaves of _Nymphaea_ the thermonastic movement is of positive sign; that is to say, an erection of the petal during rise, and a fall during the lowering of temperature. The corresponding movement of leaves would therefore be an erection of the leaf in day-time, and a fall of the leaf at night. The periodic curve of such leaves would be of opposite sign to the standard thermo-geotropic curves given above. The leaf of _Nicotina_ is adduced as an example of a leaf which exhibits a movement of fall at night. But the fully grown and horizontally spread leaf I find that gives the normal record. The very young growing leaves give a different and somewhat erratic curve. The difference between growing and fully grown leaves is explained by the fact that the former would be affected by thermotropism, and the latter by thermo-geotropism. Young leaves exhibit moreover a p.r.o.nounced hyponasty or epinasty, which would naturally modify the diurnal curve.
Certain interesting variation is met with in the diurnal record of sprouting leaves of _Mimosa_ in spring. The movements of leaves grown later in the season, as will be explained in a later chapter, are very definite and characteristic. But the young leaves in spring exhibit no definite diurnal curve, but a series of automatic pulsations, the unsuspected presence of which in all leaves of _Mimosa_ will be demonstrated in a subsequent chapter. Later in the season, the leaf becomes tuned, as it were, to the periodic variation of the environment; the automatic movements become suppressed, and the diurnal periodicity becomes deeply impressed on the organism.
_Effect of season._--The diurnal curve may also be modified by the seasonal variation of any one of the effective factors. _Tropaeolum majus_, for example, exhibits positive phototropic action in one season and a negative reaction in a different season. These seasonal variations must necessarily modify the diurnal curve.
I shall now proceed to demonstrate the determining influence of thermal variation, and of stimulus of gravity on the thermo-geotropic movements.
The striking similarity of the thermograph, and the record of movement of plants demonstrate the causal relation between temperature variation and diurnal movement, of which the two additional tests described below offer further confirmation.
REVERSAL OF NORMAL RHYTHM.
The normal diurnal movement is, as we have seen, a fall during rise of temperature from morning to afternoon, and a rise from afternoon till next morning. I succeeded in reversing the normal rhythm of _Basella_ by reversing the normal variation of temperature at the two turning points, in the morning and in the afternoon. The plant was subjected to falling temperature in the morning and to rising temperature in the afternoon.
The normal movement now became reversed, _i.e._, an erection instead of fall in the forenoon and a fall instead of rise in the afternoon (p.
28).
EFFECT OF CONSTANT TEMPERATURE.
The second test which I shall employ is the effect of maintenance of constant temperature, which should wipe off, as it were, traces of periodic movement. It was necessary for this investigation to maintain the plant chamber at constant temperature throughout day and night. The usual thermostat is virtually a recess in a double-walled chamber filled with water, the chamber being covered with a heat insulating material.
But this contrivance is unsuitable for the plant chamber which is to contain good sized plants, and the recording apparatus. The problem of maintaining a large air-chamber at constant temperature presented many difficulties which were ultimately overcome by the device of an extremely sensitive thermal regulator.
_The Thermal Regulator._--I shall in a future paper give a complete account of the large thermostatic air-chamber. The important part of the apparatus is an electro-thermic regulator which interrupts the heating electric current as soon as the temperature of the chamber is raised a hundredth part of a degree above the predetermined temperature. The automatic make and break of the current takes place in rapid succession, and the temperature of the chamber is thus maintained constant within tenth of a degree, throughout day and night.
[Ill.u.s.tration: FIG. 203.--Abolition of diurnal movement in _Tropaeolum_ under constant temperature, and its restoration under normal daily fluctuation. The upper record is of temperature and the lower of plant movement.]
_Diurnal record of_ Tropaeolum _under constant temperature: Experiment 212._--The normal record of geotropically curved _Tropaeolum_ is already given in figure 202. In repeating the record I maintained the plant at constant temperature for 24 hours; the result of this is seen in the first part of the record (Fig. 203). The thermal record is practically horizontal, and the diurnal record of the plant shows no periodic movement. The thermal regulator was on the next day put out of operation, thus restoring the normal diurnal variation of temperature.
The record of the plant is seen to exhibit once more its normal periodic movement.
I have in the chapter on thermo-geotropism (p. 515) shown that the diurnal movement of a geotropically curved organ is determined in reference to the direction of force of gravity. This will be seen demonstrated in an interesting manner in the two following experiments on the effect of inversion of the plant on daily movement.