Life Movements in Plants Vol 1/The "Praying" Palm Tree

by Jagadish Chandra Bose

 

II.—THE "PRAYING" PALM TREE

By

Sir J. Bose,

Assisted by

Narendra Nath Neogi, M.Sc.

Perhaps no phenomenon is so remarkable and shrouded with greater mystery as the performances of a particular Date Palm near Faridpur in Bengal. In the evening, while the temple bells ring calling upon people to prayer, this tree bows down as if to prostrate itself. It erects its head again in the morning, and this process is repeated every day of the year. This extraordinary phenomenon has been regarded as miraculous, and pilgrims have been attracted in large numbers. It is alleged that offerings made to the tree have been the means of effecting marvellous cures. It is not necessary to pronounce any opinion on the subject; those cures may be taken as effective as other faith-cures now prevalent in the West.

This particular Date Palm, Phœnix dactylifera, is a full-grown rigid tree, its trunk being 5 metres in length and 25 cm. in diameter. It must have been displaced by storm from the vertical and is now at an inclination of about 60° to the vertical. In consequence of the diurnal movement, the trunk throughout its entire length is erected in the morning, and depressed in the afternoon. The highest point of the trunk thus moves up and down through one metre; the 'neck,' above the trunk, is concave to the sky in the morning; in the afternoon the curvature disappears, or is even slightly reversed. The large leaves which point high up against the sky in the morning are thus swung round in the afternoon through a vertical

Fig. 1. The Faridpur 'Praying' Palm: the upper photograph shows position in the morning; the lower position in the afternoon. The two fixed stakes are one metre in height. In front is seen erect trunk of a different Palm.

distance of about five metres. To the popular imagination the tree appears like a living giant, more than twice the height of a human being, which leans forward in the evening from its towering height and bends its neck till the crown of leaves press against the ground in an apparent attitude of devotion (Fig. 1). Two vertical stakes, each one metre high, give a general idea of the size of the tree and movements of the different parts of the trunk.

For an investigation in elucidation of this phenomenon it was necessary:—

1. To obtain an accurate record of the movement of the tree day and night, and determine the time of its maximum erection and fall.

2. To find whether this particular instance of movement was unique, or whether the phenomenon was universal.

3. To discover the cause of the periodic movement of the tree.

4. To find the reason of the remarkable similarity between the diurnal movement of the tree, and the diurnal variation of moto-excitability in Mimosa pudica.

5. To determine the relative effects of light and temperature on the movement.

6. To demonstrate the physiological character of the movement of the tree.

7. To discover the physiological factor whose variation determines the directive movement.

THE RECORDING APPARATUS.

I shall now describe the principle and construction of my recording apparatus (Fig. 2) seen attached to a horizontally growing stem of Mimosa pudica. When used to trace the movement of the palm tree, a reducing device is employed to keep the record within the plate. A lever, R¹, records the movement of the attached tree or plant on a moving plate of smoked glass. The plate is not in contact with the

Fig. 2. Apparatus for automatic record of movement of trees and plants; T, differential metallic thermometer; R, recording lever for temperature; R¹, for recording plant movement; C, clock-work for oscillation of recording plate. The same clock-work moves plate laterally in 24 hours.

tip of the recording lever, but separated from it by a distance of about 3 mm. A special oscillating device, actuated by clock-work, C, makes the plate move forwards and backwards. The forward movement brings about a momentary contact of the recording tip with the smoked plate inscribing a dot. These single dots are made at intervals of 15 minutes; at the expiration of the hour, however, contact is made three times in rapid succession, printing a thick dot. It is thus easy to determine the movement of the tree at all times of the day and night. A second lever, R, placed above, gives on the same plate, thermographic record of the diurnal variation of temperature. For this I use a differential thermometer, T, made of a compound strip of brass and steel. Curvature is induced by the differential expansion of the two pieces of metal. The up or down movement of the free end of the compound strip is further magnified by the recording lever. This arrangement was extremely sensitive and gave accurate record of variation of temperature. By the forward movement of the oscillating plate two dots are made at the same time,—one for the temperature and the other for the corresponding movement of the tree. As the two recorders do not move vertically up or down, but describe a circle, the dots vertically one above the other may not correspond as regards time. Any possibility of error in calculation is obviated by the fact that the thick dots in both the records are made every hour, and the subsequent thin dots at intervals of 15 minutes.

A difficulty arose at the beginning in obtaining sanction of the proprietor to attach the recorder to the tree. He was apprehensive that its miraculous power might disappear by profane contact with foreign-looking instruments. His misgivings were removed on the assurance that the instrument was made in my laboratory in India, and that it would be attached to the tree by one of my assistants, who was the son of a priest.

From results of observation it is found that the tree moves through its entire length; the fall of the highest point of the trunk is one metre. The movement is not

Fig. 3. Record of diurnal movement of the 'Praying' Palm (Phœnix dactylifera). Thermographic curve for 24 hours commencing at 9 in the evening is given in the upper record; the corresponding diurnal curve of movement of the tree is given in the lower. Successive dots at intervals of 15 minutes; thick dots at intervals of an hour.

passive, but an active force is exerted; the force necessary to counteract this movement is equivalent to the weight of 17 kilograms: in other words, the force is sufficient to lift a man off the ground. But far greater force would be required to restrain the change of curvature of the neck of the hard and rigid tree.

Before entering into the investigation of the cause of periodic movement I shall give a general account of its characteristics. A casual observation would lead one to conclude that the tree lifted itself at sunrise and prostrated at sunset. But continuous record obtained with my recorder attached to the upper part of the trunk shows that the tree was never at rest, but in a state of continuous movement, which underwent periodic reversals (Fig. 3). The tree attained its maximum erection at 7 in the morning, after which there is a rapid movement of fall. The down movement reached its maximum at 3-15 p.m., after which it was reversed and the tree erected itself to its greatest height at 7 next morning. This diurnal periodicity was maintained day after day.

UNIVERSALITY OF TREE MOVEMENT.

The next question which I wished to investigate was whether the movement of the particular Faridpur tree was a unique phenomenon. It appeared more likely that similar movement would, under careful observation, be detected in all trees. The particular palm tree was growing at a considerable inclination to the vertical; the movement of the tree and its leaves became easily noticeable, since the ground afforded a fixed and striking object of reference. In a tree growing more or less erect, the movement, if any, would escape notice, since such movements would be executed with only the empty space as the background.

Experiment 1.—Believing the phenomenon to be universal I experimented with a different Date Palm that was growing at my research station at Sijbaria on the

Fig. 4. Record of the Sijbaria Palm from noon for 24 hours. Successive dots at intervals of 15 minutes.

Ganges, situated at a distance of about 200 miles from Faridpur. The surrounding conditions were very different. The tree was much younger; it was 2 metres in height and inclined 20° to the vertical. The curve obtained with this tree (Fig. 4) was very similar to that of the Faridpur Palm, though the extent to the movement was much reduced. The tree attained the highest erect position at 7-15 a.m. and the lowest at 3-45 p.m. Hence the movement of the Faridpur Palm is not a solitary phenomenon.

THE CAUSE OF PERIODIC MOVEMENT.

The recurrent daily movement of the tree must be due to some diurnal changes in the environment, either the recurrent changes of light and darkness, or the diurnal changes of temperature. These changes synchronise to a certain extent; for, as the sun rises, light appears and the temperature begins to rise. It is therefore difficult to discriminate the effect of light from that of temperature. The only satisfactory method of discrimination would have been in the erection of a large structure with screens to cut off light. The effect of fluctuation of temperature under constant darkness would have demonstrated the effect of one agent without complication arising from the other. Unfortunately screening the tree was impracticable. I shall presently describe other experiments where the action of light was completely excluded.

The curve of movement of the tree, however, affords us material for correct inference as regards the relative effects of light and temperature. The experiment was commenced in March; light appeared at about 5 a.m., the sunrise being at 6-15 a.m.; the sun set at 6-15 p.m., and it became dark by 7 p.m. The incident light would be the most intense at about noon; after this it would decline continuously till night time. If the movement was due to light, its climax, either in up or down movement, would be reached at or about noon, and the opposite climax at midnight. But instead of this we find (Fig. 3) the up-movement reaching its highest point not at noon, but at 7 in the morning; after this the fall is rapid and continuous, and the lowest position was reached not in the evening but at 3-15 p.m. The fluctuation of light has, therefore, little to do with the movement of the tree.

Turning next to the element of variation of temperature we are at once struck by the fact that the curve of movement of the tree is practically a replica of the thermographic curve (Fig. 3). The fall of temperature is seen to induce a rise in the tree and vice versâ. There is a lag in the turning points of the two curves; thus while temperature began to rise at 6 a.m., the tree did not begin to fall till 7 a.m. There is in this case a lag of an hour; but the latent period may, sometimes, be as long as three hours. The delay is due to two reasons; it must take some time for the thick trunk of the tree to attain the temperature of the surrounding, and secondly, the physiological inertia will delay the reaction. As a result of other investigations, I find that the induced effect always lags behind the inducing cause. It is interesting in this connection to draw attention to the parallel phenomenon, which is described below, of lag in the variation of sensibility of Mimosa in response to variation of temperature. In this case the lag was found to be about three hours. Returning to the Palm, the tree continues to fall in the forenoon with rising temperature. At about 2-30 p.m. the temperature was at its maximum after which it began to decline; the movement of the tree was not reversed into erection till after 3-15 p.m., the lag being now 45 minutes nearly.

I may state here that the movement of the tree is not primarily affected by the periodicity of day and night, but by variation of temperature. In spring and in early summer the rise of temperature during the early part of the day and the fall of the temperature from afternoon to next morning, are regular and continuous; the corresponding movements of the tree are also regular. But at other seasons, owing to the sudden change of direction of the wind, the fluctuations of temperature are irregular. Thus at night there may be a sudden rise, and in the earlier part of the day sudden fall of temperature. And the record of movement of the tree is found to follow these fluctuations with astonishing fidelity, the rise of temperature being followed by a fall of the tree and vice versâ. That the movement is determined by the temperature variation is exhibited in a striking manner in Fig. 4, where, between 8 and 9 a.m., a common twitch will be noticed in the two curves.

While trying to obtain some clue to the mysterious movement of the tree, my attention was strongly attracted by certain striking similarities which the record of the movement of the tree showed to the curve of the diurnal variation of moto-excitability, of the pulvinus of Mimosa pudica, an account of which will be found in a subsequent Paper of the series.^[1]

PERIODIC MOVEMENT OF TREES AND DIURNAL VARIATION OF MOTO-EXCITABILITY IN MIMOSA PUDICA.

The excitability of the main pulvinus of Mimosa pudica I find does not remain constant during the 24 hours, but undergoes a striking periodic change. At certain hours of the day, the excitability is at its maximum; at a different period it practically disappears. The period of insensibility is about 7 a.m., which, strangely enough, is also the time when the palm tree attains its maximum height. At about 3 in the afternoon the excitability of Mimosa reaches its climax, and this is the time when the head of the palm tree bends down to its lowest position. For the determination of the periodic variation of excitability of Mimosa I devised a special apparatus by which an electric stimulus of constant intensity was automatically applied to the plant every hour of the day and night, the responsive moment being recorded at the same time. The amplitude of responsive fall of leaf under uniform stimulus gave a measure of excitability of

Fig. 5. Curve of variation of moto-excitability of Mimosa pudica. The upper curve gives variation of temperature and the lower, the corresponding variation of excitability.

the leaf at any particular moment. In the lower curve of Fig. 5 is given the record of diurnal variation of excitability of Mimosa. Comparison of this figure with Figs. 3 and 4, will show the remarkable resemblance between the curves of diurnal movement of the Palm tree, and of diurnal variation of moto-excitability of Mimosa. The excitability of Mimosa reached its maximum at about 3 in the afternoon, when the Palm was at its lowest position. After this hour excitability fell continuously till 7 or 8 next morning. Corresponding to this is the continuous erection of the Palm from its lowest position at 3 p.m. to the highest between 7 and 8 a.m. Still more remarkable is the modifying influence of variation of temperature on the diurnal curve of excitability in Mimosa, and the diurnal curve of movement of the Palm. This will be quite evident from the inspection of the temperature curves in Figs. 4 and 5.

I have shown elsewhere^[2] that the variation of moto-excitability of the pulvinus of Mimosa is a physiological function of temperature. The remarkable similarity between the diurnal variation of moto-excitability of Mimosa and diurnal movement of the Palm is due to the fact that both are determined by the physiological action of temperature. I shall presently describe experiments, which will establish the physiological character of the movement of the tree in response to changes of temperature.

The records that have been given show that it is the diurnal variation of temperature, and not of light that is effective in inducing the periodic movement of the tree. Further experiments will be given in support of this conclusion.

RELATIVE EFFECTS OF LIGHT AND TEMPERATURE.

As regards the possibility of light exerting any marked influence on the movement of the Palm tree, I have shown from study of time-relations of the movement, that this could not be the case. Moreover, it is impossible for light to reach the living tissue through the thick layer of bark that surrounds the tree. That the effect of light is negligible will appear from the accounts of following experiments, where the possibility of the effect of changing intensity of light is excluded by maintaining the plant in constant darkness, or in constant light.

The employment of the large Palm was obviously impracticable in these investigations. I, therefore, searched for other plant-organs in which the movement under variation of temperature was similar to that of the Date Palm. I found that the horizontally spread leaves of vigorous specimens of Arenga saccharifera growing in a flower pot executed movements which were practically the same as that of the Faridpur tree. The leaf moved downwards with rise of temperature and vice versâ.

There are many practical advantages in working with a small specimen. It can easily be placed under glass cover or taken to a glass house, thus completely eliminating the troublesome disturbance caused by the wind.

Diurnal movement in continued darkness: Experiment 2.—The plant was placed in a dark room and records taken continuously for three days. These did not differ in any way from the normal records taken in a glass house under daily variation of light and darkness. Exposure of plant to darkness for the very prolonged period of a week or more, undoubtedly interferes with the healthy photo-tonic condition of the plant. But such unhealthy condition did not make its appearance in the first few days.

PHYSIOLOGICAL CHARACTER OF THE MOVEMENT.

There may be a misgiving that the movement of the tree might be due to physical effect of temperature. If the upper strip of a differential thermometer be made of the more expansible brass and the lower of iron, the compound strip bends down with the rise of temperature. Similarly the movement of the tree might be due to the upper half being physically more expansible. It would have been possible to discriminate the physical from the physiological action by causing the death of the tree; in that case physical movement would have persisted, while the physiological action would have disappeared. As this test was not practicable, I tried the effect of physiological depression on the periodic movement of the leaf of Arenga saccharifera.

Effect of Drought: Experiment 3—In Fig. 6 is given a series of records of movement of the leaf-stalk of Arenga, first under normal condition, afterwards under increasing drought, brought about by withholding water. Tim uppermost is the thermographic record which remained practically the same for successive days. Below this are records of movement of the leaf (a) under normal condition, (b) after withholding water for three days, and (c) after deprivation for seven days. It will be noticed how the extent of movement is diminished under increasing physiological depression brought on by drought. On the seventh day, the responsive movement disappeared, there being now a mere fall of the leaf, which was slow and continuous. After this I supplied the plant with water and the periodic movement was in consequence nearly restored to its original vigour.

Effect of poison: Experiment 4.—In another experiment the normal diurnal record with the leaf was taken and the plant was afterwards killed by application of poisonous solution of potassium cyanide. The diurnal movement was found permanently abolished at the death of the plant.

Fig. 6. Effect of physiological depression on diurnal movement of the petiole of Arenga saccharifera. The uppermost curve exhibits variation of temperature, (a), normal diurnal curve, (b), modification after 3 days' and (c) after 7 days' withholding of water.

These experiments conclusively prove that the periodic movement of the leaf-stalk induced by variation of temperature is a physiological phenomenon, and from analogy we are justified in drawing the inference that the movement of the Faridpur tree is also physiological. The question, however, was finally settled by the unfortunate death of the tree which occurred the other day, nearly a year after I commenced my investigations. While presiding at my lecture on the subject, His Excellency Lord Ronaldshay, the Governor of Bengal, announced that a telegram had just reached him from his officer at Faridpur that "the palm tree was dead, and that its movements had ceased."

Since my investigation with the Faridpur 'Praying' Palm, I have received information regarding other Palms, which exhibit movements equally striking. One of the trees is growing by the side of a tank, the trunk of the tree being inclined towards it. The up-lifted leaves of this tree are swung round in the afternoon and dipped into the water of the tank.

The movement of the tree has been shown to be brought about by the physiological action of temperature variation; in other words the diurnal movement of the 'Praying' Palm is a THERMONASTIC PHENOMENON. I have found various creeping stems, branches and leaves of many trees, exhibit this particular movement of fall with a rise of temperature, and vice versâ. Such movements, I shall, for the sake of convenience, distinguish as belonging to the negative type.

Having found that the temperature is the modifying cause, the next point of inquiry relates to the discovery of the force, whose varying effects under changing temperature induces the periodic movement. I shall, in this connection, first discuss the various tentative theories that may be advanced in explanation of the movement.

TRANSPIRATION AND DIURNAL MOVEMENT.

It may be thought that the fall of the tree during rise of temperature may be due to passive yielding of the tree to its weight, there being increased transpiration and general loss of turgor at high temperature. I shall, however, show that the diurnal movement persists in the absence of transpiration.

Diurnal movement in absence of transpiration: Experiment 5.—In the leaf of Arenga saccharifera, I found that the petiole was the organ of movement. I cut off the transpiring lamina and covered the cut end with collodion flexile. The plant was now placed in a chamber saturated with moisture. The petiole continued to give records of its diurnal movement in every way similar to the record of the intact leaf. In another experiment with the water plant, Ipoemia reptans, immersed in water, the normal diurnal movement was given by the plant, where there could be no question of variation of turgor due to transpiration. (See also Expt. 7.)

In the diurnal movement of the 'Praying' Palm the concave curvature of the rigid neck in the morning, became flattened or slightly convex in the afternoon. The force necessary to cause this is enormously great, and could on no account result from the passive yielding to the weight of the upper part of the tree.

From the facts given above it will be seen that the diurnal movement is not brought about by variation in transpiration. I now turn to another phenomenon which appeared at first to have some connection with the movement of the tree. Kraus found that the tissue tensions of a shoot exhibit a daily periodicity. He, however, found that between 10°C. and 30°C., variation of temperature had no effect on the daily period. But as regards the diurnal movement of the tree, it is the temperature which is the principal factor. Kraus also found a daily variation of bulk in different plant-organs; this variation of bulk is connected with transpiration, for the removal of the transpiring leaves arrested this variation. But the periodic movement of the tree, as we have seen, is independent of transpiration.

Millardet observed a daily periodicity of tension in Mimosa pudica. He found that maximum tension occurs before dawn; the petiole becomes erected, the movement being upwards or towards the tip of the stem. Tension decreases during the day, and reaches a minimum early in the evening; in correspondence with this is the fall of the petiole, the movement being away from the tip of the stem.^[3] If the plant were placed upside down the periodic movement of the petiole in relation to the stem will evidently remain the same, but become reversed in space. Maximum tension in the morning will make the petiole approach the tip of the siem, i.e., the movement will be downwards instead of upwards as in the normal position. The experiment described below will show that the diurnal movement induced by variation of temperature is not reversed by placing the plant in an inverted position.

Diurnal movement in inverted position: Experiment 6.—I took a vigorous specimen of Arenga saccharifera growing in a pot, and took its normal record, which as explained before exhibited down-movement during rise, and an up-movement during fall of temperature. The plant was now held inverted, the upper side of the petiole now facing the earth. The diurnal curve of movement should now show an inversion, if that movement was solely determined by the anisotropy of the organ. But the record did not exhibit any such inversion. After being placed upside down, the leaf did not, on the first day, show any diurnal movement; there was, on the other hand, a continuous down-movement on account of the fall of the leaf by its own weight. But in the course of 24 hours the leaf readjusted itself to its unaccustomed position, and became somewhat erected under the action of geotropic stimulus. After the attainment of this new state of geotropic equilibrium, the leaf gave a very pronounced record of its diurnal movement which did not show any reversal; the inverted leaf continued to exhibit the same characteristic movements as in the normal position, that is to say, a down movement during rise, and an up-movement during fall of temperature. As the plant in the inverted position did not show any reversal of the periodic curve, it is clear that the diurnal movement is determined by the modifying influence of temperature on the physiological reaction of the plant to some external stimulus which is constant in direction. I shall presently show that it is the constant geotropic stimulus modified by the action of temperature, which determines the diurnal movement of the tree.

This will be better understood if I refer once more to certain characteristics in the movement of the "Praying" Palm. The neck of the tree was seen to be concave in the morning. The physiological effect of raising temperature is virtually to oppose or neutralise the geotropic curvature as seen in the flattening or slight reversal of curvature in the afternoon. Similarly, various plant organs, growing at an inclination to the vertical, are subjected to geotropic action, and thus assume different characteristic angles. This state of equilibrium is not static but may better be described as dynamic; for it will be shown that this state of geotropic balance is upset in a definite way, by variation of temperature.

That geotropism is an important factor in the diurnal movement is supported by the fact that the Sijbaria Palm with an inclination of 20° to the vertical exhibited a daily movement which was only moderate in extent. But the Faridpur Palm growing at an inclination of 60° was subjected more effectively to geotropic action, and exhibited movements which were far more pronounced. I shall now proceed to describe crucial experiments which will demonstrate the effect of change of temperature on geotropic curvature.

EFFECT OF VARIATION OF TEMPERATURE ON GEOTROPIC CURVATURE.

In the instances of diurnal movement already described the trees or their leaves were already at an inclination to the vertical. I now took a radial and erect shoot of Basella cordifolia growing in a pot and laid it horizontally for two weeks. The procumbent stem curved up and attained a state of equilibrium under the action of geotropic stimulus.

Diurnal curve of Basella cordifolia: Experiment 7.— The plant was completely immersed in a vessel of water, and its diurnal curve recorded. This resembled in all essentials the diurnal curve of the Palm; the slight deviation was due to the fact that owing to difference in the season (August) the temperature maximum was attained at 12-25 p. m., and the minimum at 6 a. m. The geotropic curvature was reduced to its minimum at the maximum temperature, and vice versâ. As in the case of the Palm so also in the procumbent stem of Basella there was a physiological lag, which was 50 minutes in the morning and about the same in the afternoon. The free end of the stem thus exhibited a diurnal movement up and down. The temperature, as stated before, began to rise from 6 a. m. and the down-movement commenced 50 minutes later, i.e., at 6-50 a. m. The temperature, after reaching

Fig. 7. Diurnal curve of movement of procumbent young stem of Mimosa pudica. Successive dots at intervals of 15 minutes.

the maximum, began to fall at 12-25 p. m., and the previous movement of fall of the stem was arrested and reversed into an erectile movement shortly after 1 p. m. There are thus two "turning points," one at 7 a.m., and the other at about 1 p.m.; at these periods the movement of the plant remains more or less arrested for more than half-an-hour.

I obtained records of similar diurnal movements with various procumbent or creeping stems. Figure 7 gives the diurnal record of the procumbent stem of a young specimen of Mimosa pudica.

The experiment that has just been described shows clearly that geotropic curvatures of stems is opposed, or neutralised to a greater or less extent, during rise of temperature, and this antagonistic reaction is removed during the fall of temperature. The diurnal movement of the plant completely immersed under water shows once more that transpiration has little to do with the diurnal movement.

REVERSAL OF NATURAL RHYTHM.

The diurnal rhythm of up and down movement in the particular specimen Basella had become established under the daily variation of temperature. I now attempted to reverse this rhythm by artificial variation of temperature. The plant was placed in water in a rectangular metallic vessel which was placed within a second outer vessel. The plant could thus be subjected, without any mechanical disturbance, to variation of temperature, by circulating warm or cold water in the outer vessel. In order to reverse the natural rhythm I subjected the plant to the action of falling temperature at the "turning" point at 7 a. m., at a time when the plant would have undergone a down-movement under the daily rise of temperature. Conversely the plant was subjected to the action of rising temperature at the second "turning" point at 1 p. m., when the movement under diurnal fall of temperature would have been one of erection.

Effect of fall of temperature: Experiment 8.—As stated before the experiment was carried out in the morning; ice cold water was circulated in the outer chamber, the fall of temperature was in this case sudden, and there was an almost immediate responsive movement. This appeared anomalous, since the latent period of response to slow variation of temperature was found from the diurnal curve to be as long as 50 minutes.

As a result of further investigations I found that variation of temperature produces two different effects which may he distinguished as transient and persistent. Sudden variation of temperature affects the superficial tissue, and gives rise to a transient reaction, while it takes a long time for temperature variation to react on the geotropically active tissue in the interior. The persistent effect therefore takes place after a latent period from one to three hours according to the thickness of the plant.

The persistent effect of rise of temperature is a movement downwards, that of fall of temperature is a movement upwards. These definite reactions will be seen exhibited in Figs. 8 and 9. The plant was stationary at the turning point in the morning hence the curve at first was horizontal. The temperature was gradually lowered through 5°C, from 29°C, to 24°C. in the course of five minutes and maintained at the lower temperature. There was no immediate effect, but after a latent period of 65 minutes the plant responded by a movement of erection. The natural movement at this period of the day would have been one of fall, but artificial change of temperature in the opposite direction effectively reversed the normal diurnal movement. The latent period for this reverse

Fig. 8.

Fig. 9.

Fig. 8. Reversal of normal rhythm: Erectile response Basella to gradual fall of temperature.
Fig. 9. Responsive fall of Basella to gradual rise of temperature.
(Dots at intervals of 5 minutes).

movement is, as stated before, 65 minutes as against 50 minutes in the normal diurnal movement. The increase in the latent period is probably due to the added physiological inertia in reversing the normal rhythm.

Effect of rise of temperature: Experiment 9.—The temperature was raised through 5°C at the second turning point at 1 p.m. After a latent period of 50 minutes the plant began to rise steadily (Fig. 9) thus exhibiting once more the reversal of its normal diurnal movement.

From the experiments described above it will be seen that the movement of the Palm, and of other organs growing at an inclination to the vertical, is brought about by the action of temperature in modifying the geotropic curvature. The ever present tendency of geotropic movement is opposed or helped by the physiological reaction induced by rise and fall of temperature respectively. The state of equilibrium is never permanent, but the dynamic balance is being constantly readjusted under changing conditions of the environment.

The movement of the tree furnishes an example of the negative type of THERMONASTIC MOVEMENT. Parallel phenomena are found in floral organs, where, in the well-known instance of Crocus, the perianth leaves open outwards during rise of temperature and close inwards during the onset of cold. Looked at from above, the opening outwards during rise of temperature is a movement downwards, and therefore belongs to the negative type. In such cases the changed rate of growth by variation of temperature is the most important factor in the movement. It may be asked whether all thermonastic movements must necessarily belong to the negative type, where rise of temperature is attended by a movement downwards. I shall in my Paper on "Thermonastic Phenomena" show that there is also a positive type where rise of temperature induces an up-movement or of closure.

SUMMARY.

The 'Praying' Palm of Faridpur, growing at an inclination of about 60° to the vertical, exhibited a diurnal movement by which its head became erected in the morning and depressed towards the afternoon, the outspread leaves pressing against the ground.

The record of the diurnal movement showed that the head was erected to the highest position between 7 and 8 in the morning, after which there was a continuous fall which reached its climax at 3-15 p.m.; after this the movement was reversed and the maximum erection was again reached next morning.

This phenomenon is not unique, but is found exhibited, more or less, by all trees and their branches and leaves.

Diurnal records of temperature, and movement of the tree showed, that the two curves closely resembled each other. Rise of temperature was attended by a fall of the tree, and vice versâ.

The movement is brought about by the physiological action of temperature; it may be arrested by artificially induced physiological depression, and is permanently abolished at death.

The movement is primarily determined by the modifying influence of temperature on geotropic curvature. Rise of temperature is found to oppose or neutralise geotropic curvature, the fall of temperature inducing the opposite effect. The ever present tendency of upwards geotropic movement is opposed or helped by the effects of rise and fall of temperature respectively.

The movement of the "Praying" Palm is a thermonastic phenomenon. The tree, apparently so rigid, responds as a gigantic pulvinoid to the changes of its environment.

 

1. See also Bose—Diurnal Variation of Moto-Excitability in Mimosa—Annals of Botany, Vol. XXVII, No. CVIII, October, 1913.
2. Bose—"Irritability of Plants," d. 60
3. Vines.—'Physiology of Plants,' 1886, pp. 405 and 543.