Experiment 11.—I took a specimen of hooked tendril, and excited it by an electric shock. The response was by the greater contraction of the more excitable convex side, on account of which the curved specimen tended to open out. The record of this response is seen in Fig. 11; the apex-time was nearly two minutes, and the recovery was completed in the further course of 15 minutes.
From the responses of organs rendered anisotropic by the differential action of the environment we pass to others which show certain amount of anatomical and physiological differentiation between their upper and lower sides. I find that many petioles of leaves show movement in response to stimulus. Many pulvini, generally regarded as insensitive, are also found to exhibit responsive movements.
The most striking and familiar example of response is afforded by the main pulvinus of Mimosa pudica of which a record is given in Fig. 12. It is generally assumed that sensibility is confined to the lower half of the organ. It will be shown in a subsequent Paper that this is not the case. The upper half of the pulvinus is also sensitive though in a feeble degree, its excitability being about 80 times less than that of the lower half. On diffuse stimulation the predominant contraction of the lower half causes the fall of the leaf, the antagonistic reaction of the upper half being, in practice, negligible. In order to avoid unnecessary repetition, I shall ignore the feeble antagonistic reaction of the less excitable half of the organ, and shall use the word ‘contraction’ for ‘relatively greater contraction.’
It is interesting in this connection to refer to the response of the leaf of Water Mimosa (Neptunia oleracea). Here the reaction is very sluggish in comparison with that of Mimosa pudica. A tabular statement of contractile response of various radial, anisotropic and pulvinated organs will show a continuity in the contractile reaction; the difference exhibited is a question of degree and not of kind.
TABLE 1—PERIODS OF MAXIMUM CONTRACTION AND OF RECOVERY OF DIFFERENT PLANTS.
| Specimen | Period of maximum contraction | Period of recovery. |
| Radial organ: Tendril of Passiflora | 100 seconds | 4 minutes. |
| Anisotropic organ: Hooked tendril of Passiflora | 120 " | 13 " |
| Pulvinated organ: Pulvinus of Neptunia Oleracea | 180 " | 57 " |
| Pulvinus of Mimosa pudica | 3 " | 16 " |
As regards the excitatory fall of the leaf of Mimosa pudica, Pfeffer and Haberlandt are of opinion that this is due to the sudden diminution of turgor in the excited lower half of the pulvinus. The weight of the leaf, no longer supported by the distended lower cells, causes it to fall. This is accentuated by the expansion of the upper half of the pulvinus which is normally in a state of compression. According to this view the excitatory fall of the leaf is a passive, rather than an active, movement. I have, however, found that in determining the rapidity of the fall of Mimosa leaf the factors of expansive force of the upper half of the pulvinus and the weight of the leaf are negligible compared to the active force of contraction exerted by the lower half of the pulvinus (p. 87).
With regard to the fall of turgor, it is not definitely known whether excitation causes a sudden diminution in the osmotic strength of the cell-sap or an increase in the permeability of the ectoplast to the osmotic constituents of the cell. Pfeffer favours the former view, while others support the theory of variation of permeability.[F]
Whatever difference of opinion there may be in regard to the theories of osmotic and permeability variations, we have the indubitable fact of diminution of turgor and contractile fall of the pulvinus of Mimosa under excitation. The restoration of the original turgor brings about recovery and erection of the leaf. In connection with this the following experiments on responsive movements of the leaf under artificial variation of turgor will be found of interest:—
Effect of Increased Turgor: Experiment 12.—A young Mimosa plant was carefully transplanted and the root embedded in soil placed in a linen bag. This was held securely by a clamp, and one of the leaves of the plant attached to the recorder. Withholding of water for a day caused a general loss of turgor of the plant. A vessel full of water was now raised from below so that the linen bag containing the roots was now in water. The effect of increased turgor by suction of water by the roots became apparent by the upward movement of the leaf. The distance between the immersed portion of the plant and the leaf was 2 cm. and the up-movement of the leaf was indicated within 10 seconds of application of water (Fig. 13). The velocity with which the effect of increased turgor travelled was thus 2 mm. per second. The leaf exhibited increasing erection with absorption of water.
Effect of Diminution of Turgor: Experiment 13.—While the leaf in the above experiment was in process of erection, a quick change was made by substituting KNO3 solution for the water of the vessel in which the roots were immersed. The plasmolytic withdrawal of water at the roots gave rise to a wave of diminished turgor, the effect of which became perceptible within 40 seconds by the movement of fall of the leaf. (Fig. 13.)
In Mimosa excitation is manifested by the contraction of the pulvinus and the consequent movement of the leaf. But in most plants, excitatory movement cannot be realized on account of the rigidity of the plant structure, the thickness of the cell-wall and the want of facility for escape of water from the excited cells. I shall show later how excitation may be detected in the absence of mechanical movement.
As regards stimulation of vegetable tissues, there are various agencies besides electric shock, which induce excitatory contraction; these agencies I shall designate as stimuli. Excitation is detected in Mimosa by the downward movement of the leaf. It will be found that such excitatory movement is caused by a mechanical blow, by a prick or a cut, by the application of certain chemical agents, by the action of electric current and by the action of strong light. The study of the action of these stimuli will be given in greater detail in subsequent Papers.
I shall give below a general classification of different stimuli which cause excitation in vegetable tissues.
Electric Stimulus.—Induction shock, condenser discharge, the make of kathode and the break of anode.
Mechanical Stimulus.—Mechanical blow, friction, prick or cut.
Chemical Stimulus.—Effect of certain acids and of other chemical substances.
Thermal Stimulus.—Sudden variation of temperature; application of heated wire.
Radiation Stimulus.—Luminous radiation of the more refrangible portion of the spectrum; ultra-violet rays; thermal radiation in the infra-red region.
All these different forms of stimulus induce an excitatory contraction, a diminution of turgor, and a negative mechanical response or fall of a motile leaf.
A radial organ responds to stimulus by contraction in length; as all its flanks are equally excitable there is no lateral movement under diffuse stimulus.
Physiological anisotropy is induced in an organ, originally radial and isotropic, by the unequal action of the environment on its different sides. Diffuse stimulus induces a greater contraction of the more excitable side.
In a curved tendril the concave side is less excitable than the convex. Diffuse stimulus tends to straighten the curved tendril.
In the pulvinus of Mimosa pudica, the lower half is eighty times more excitable than the upper, and the fall of the leaf is due to the predominant contraction of the more excitable lower half.
A diminution of turgor takes place in the excited cells. Restoration of turgor brings about recovery of the leaf to its normal erect position. Independent experiments show that the fall of the leaf may be brought about by an artificial diminution of turgor, and the erection of the leaf by an increase of turgor.
BY
Sir J. C. Bose.
Several phenomena of daily periodicity are known, but the relations between the recurrent external changes and the resulting periodic variations are more or less obscure. As an example of this may be cited the periodic variation of growth. Here the daily periodicity exhibited by a plant is not only different in varying seasons, but it also differs in diverse species of plants. The complexity of the problem is very great, for not only are the direct effects of the changing environment to be taken into consideration but also their unknown after-effects. Even in the case of direct effect, different factors, such as light, temperature, turgor, and so on, are undergoing independent variations; it may thus happen that their reactions may sometimes be concordant and at other times discordant. The nyctitropic movement of plants affords another example of daily periodicity. The fanciful name of ‘sleep’ is often given to the closure of the leaflets of certain plants at night. The question whether plants sleep or not may be put in the form of the definite inquiry: Is the plant equally excitable throughout day and night? If not, is there any definite period at which it practically loses its excitability? Is there, again, another period at which the plant wakes up, as it were, to a condition of maximum excitability?
In the course of my investigations on the irritability of Mimosa pudica, I became aware of the existence of such a daily periodicity; that is to say, the moto-excitability of the pulvinus was found to be markedly diminished or even completely abolished at a certain definite period of the day; at another equally definite period, the excitability was observed to have attained its climax. The observations on the periodic variation of excitability appeared at first to be extremely puzzling. It might be thought, for example, that light would prove to be favourable for moto-excitability; in actual experiment the results apparently contradicted such a supposition: for the excitability of the plant was found much higher in the evening than in the morning. Favourable temperature, again, might be regarded as an important factor for the enhancement of the moto-excitability; it was, nevertheless, found that though the excitatory response was only moderate at that period of night when the temperature was at its minimum, yet the excitability was altogether abolished at another period when the temperature was several degrees higher. The obscurities which surrounded the subject were only removed as a result of protracted investigation and comparison of continuous automatic records made by the plant itself during several months, beginning with winter and ending in summer.
The question whether a plant like Mimosa exhibits diurnal variation of excitability can be experimentally investigated by subjecting the plant at every hour of the day and night to a test-stimulus of uniform intensity, and obtaining the corresponding mechanical responses. Under these circumstances the amplitude of response at any time will serve as a measure of the excitability of the plant at the particular time. Any periodic fluctuation of response will then demonstrate the periodic character of variation of excitability.
The investigation thus resolves itself into:—
The successful construction of a Response Recorder which will automatically record the response of the plant to uniform periodic stimulation at all hours of day or night;
the study of the effects of various external conditions on excitability;
the diurnal variation of excitability and its relation to the changes of external conditions.
I will first give a diagrammatic view of the different parts of the apparatus which I devised for this investigation.[G] The leaf of Mimosa is attached to one arm of a light aluminium lever, L, by means of thread. At right angles to the lever is the writing index W, which traces on a smoked glass plate allowed to fall at a definite rate by clockwork the responsive movement of the leaf. Under a definite stimulus of electric shock the leaf falls down, pulling the lever L, and moving the writer towards the left. (Fig. 14.) The amplitude of the response-curve measures the intensity of excitation. The leaf re-erects itself after a time, the corresponding record exhibiting recovery. A second stimulus is applied after a definite interval, say an hour, and the corresponding response shows whether the excitability of the plant has remained constant or undergone any variation.
Electric mode of excitation.—I find that one of the best methods of stimulating the plant is by means of tetanizing induction shock. The sensitiveness of Mimosa to electric stimulation is very great; the plant often responds to a shock which is quite imperceptible to a human subject. By the employment of a sliding induction coil, the intensity of the shock can be regulated with great accuracy; the secondary if gradually brought nearer the primary till a stimulus is found which is minimally effective. The intensity of stimulus actually employed is slightly higher than this, but within the sub-maximal range. When the testing stimulus is maintained constant and of sub-maximal intensity, then any variation of excitability is attended by a corresponding variation in the amplitude of response.
The exciting value of a tetanizing electric shock depends (1) on the intensity, (2) on the duration of shock. The intensity may be rendered uniform by placing the secondary at a fixed distance from the primary, and keeping the current in the primary circuit constant. The constancy of the current in primary circuit is secured by the employment of an accumulator or storage cell of definite electromotive force. It is far more difficult to secure the constant duration of the tetanizing shock in successive stimulations at intervals of, say, one hour during twenty-four hours. The duration of the induction shock given by the secondary coil depends on the length of time during which the primary circuit is completed in successive excitations. I have succeeded in overcoming the difficulty of securing uniformity of duration of shock by the employment of a special clockwork device.
The clockwork plunger.—The alarum clock can be so arranged that a wheel is suddenly released and allowed to complete one rapid revolution at intervals of, say, one hour. There is a fan-governor by which the speed of the revolution can be regulated and maintained constant. This will specially be the case when the alarum spring is long and fully wound. The succession of short releases twenty-four times during the day produce relatively little unwinding of the spring. On account of this and the presence of the fan-governor, the period of a single revolution of the wheel remains constant. By means of an eccentric the circular movement is converted into an up and down movement. The plunging rod R thus dips into a cup of mercury M, for a definite short interval and is then lifted off. The duration of closure can be regulated by raising or lowering the cup of mercury. In practice the duration of tetanizing shock is about 0.2 second.
The same clock performs three functions. The axis which revolves once in twelve hours has attached to it a wheel, and round this is wound a thread which allows the recording glass plate to fall through six inches in the course of twenty-four hours. A spoke attached to the minute hand releases the alarum at regular and pre-determined intervals of time, say once in an hour. The plunging rod R, actuated by the eccentric, causes a tetanizing shock of uniform intensity and duration to be given to the plant at specified times.
Constancy of resistance in the secondary circuit.—In order that the testing electric stimulus shall remain uniform, another condition has to be fulfilled, namely, the maintenance of constancy of resistance in the secondary circuit, including the plant. Electric connections have to be made with the latter by means of cloth moistened with dilute salt solution; drying of the salt solution, however, gives rise to a variation of resistance in the electrolytic contact. This difficulty is overcome by making the electrolytic resistance negligible compared to the resistance offered by the plant. Thin and flexible spirals of silver tinsel attached to the electrodes E, E′ are tied round the petiole and the stem, respectively. In order to secure better electric contact, a small strip of cloth moistened with dilute salt and glycerine is wound round the tinsel. As the resistance of contact is relatively small, and as drying is to a great extent retarded by glycerine, the total resistance of the secondary circuit undergoes practically no variation in the course of twenty-four hours. This will be seen from the following data. An experiment was commenced one day at 1 P.M., when the resistance offered by 8 cm. length of stem and 2 cm. length of petiole was found to be 1.5 million ohms. After twenty-four hours’ record, the resistance was measured the next day and was found unchanged. The fact that the stimulus remains perfectly uniform will be quite apparent when the records given in the course of this paper are examined in detail.
The amplitude of response affords, as we have seen, a measure of the excitability of the plant. In actual record friction of the writer against the glass surface becomes a source of error. This difficulty I have been able to overcome by the two independent devices, the Resonant Recorder and the Oscillating Recorder. In the former the writer is maintained by electric means in a state of continuous to and fro vibration, about ten times in a second. There is thus no continuous contact between the writer and the smoked glass surface, friction being thereby practically eliminated. The writer in this case taps a record, the successive dots occurring at intervals of 0.1 second. The responsive fall of the leaf is rapid, hence the successive dots in this part of the record are widely spaced; but the erection of the leaf during recovery takes place slowly, hence the recovery part of the curve appears continuous on account of the superposition of the successive dots. The advantage of the Resonant Recorder is that the curve exhibits both response and recovery. This apparatus is admirably suited for experiments which last for a few hours. There is, however, some drawback to its use in experiments which are continued for days together. This will be understood when we remember that for the maintenance of 10 vibrations of the writer in a second, 10 electric contacts have to be made; in other words, 36,000 intermittent electric currents have to be kept up per hour. This necessitates the employment of an electric accumulator having a very large capacity.
In the Oscillating Recorder the recording plate itself moves to and fro, making intermittent contact with the writer about once in a minute. The recording smoked glass plate is allowed to fall at a definite rate by the unwinding of a clock wheel. By an electromagnetic arrangement the holder of the smoked glass plate is made to oscillate to and fro, causing periodic contact with the writer.
The Oscillator is diagrammatically shown in Fig. 15. M, M′ are the two electromagnetic coils, the free ends of the horseshoe being pointed. Facing them are the conical holes of the soft iron armature A. This armature carries two rods which slide through hollow tubes. The distal ends of the rods support the holder H, carrying the smoked glass plate. Under normal conditions, the plate-holder is held by suitable springs, somewhat to the right of, and free from contact with, the writer. A clockwork C carries a rotating arm, which makes periodic contact with a pool of mercury contained in the vessel V, once in a minute. On the completion of the electromagnetic circuit, the armature A is attracted, the recording glass plate being thereby moved to the left making contact with the writer. The successive dots in the record thus take place at intervals of a minute. Only a moderate amount of electric current is thus consumed in maintaining the oscillation of the plate. A 4-volt storage cell of 20 amperes capacity is quite sufficient to work the apparatus for several days.
The responsive fall of the leaf of Mimosa is completed in the course of about two seconds. The leaf remains in the fallen or ‘contracted’ position for nearly fifteen seconds; it then begins to recover slowly. As the successive dots of the Oscillating Recorder are at intervals of a minute, the maximum fall of leaf is accomplished between two successive dots. The dotted response record here obtained exhibits the recovery from maximum fall under stimulation (cf. Fig. 23). The recovery of the leaf in one minute is less than one-tenth the total amplitude of the fall, and is proportionately the same in all the response records. Hence the successive amplitudes of response curves that are recorded at different hours of the day afford us measures of the relative variations of excitability of the plant at different times. This enables us to demonstrate the reality of diurnal variation of excitability. In my experimental investigations on the subject I have not been content to take my data from any particular method of obtaining response, but have employed both types of recorders, the Resonant and Oscillating. It will be shown that the results given by the different instruments are in complete agreement with each other.
Before giving the daily records of periodic variation of excitability, I will give my experimental results on the influence of various external conditions in modifying excitability. The conditions which are likely to affect excitability and induce periodicity are, first, the effects of light and darkness: under natural conditions the plant is subjected in the morning to the changing condition from darkness to light; then to the action of continued light during the day; and in the evening to the changing condition from light to darkness. A second periodic factor is the change in the condition of turgidity, which is at its maximum in the morning, as evidenced by the characteristic erect position of the petiole. Finally, the plant in the course of day and night is subjected to a great variation of temperature. I will now describe the effects of these various factors on excitability. It should be mentioned here that the experiments were carried out about the middle of the day, when the excitability, generally speaking, is found to remain constant.
I have frequently noticed that a depression of excitability occurred when the sky was darkened by passing clouds. This is clearly seen in the above records obtained with the Resonant Recorder. Uniform sub-maximal stimuli had been applied to a specimen of Mimosa at intervals of fifteen minutes. The dotted up-line represents the responsive fall, and the continuous down-line, the slow recovery. The first four are the normal uniform responses (Fig. 16). The next three show the depressing effect of relative darkness due to cloudy weather. The sky cleared after forty-five minutes, and we notice the consequent restoration of normal excitability.
Effect of sudden darkness and its continuation. Experiment 14.—In the next record (Fig. 17) is shown the immediate and continued action of darkness. The first two are the normal uniform responses in light. By means of screens, the plant was next subjected to sudden darkness; this brought about a marked depression of excitability. Subjection to sudden darkness thus acts as a stimulus inducing a marked but transient fall of excitability. Under the continuous action of darkness, however, the excitability is at first restored and then undergoes a persistent depression.
Effect of transition from darkness to light: Experiment 15.—Here we have to deal first with the immediate effect of sudden transition, and then with the persistent effect of continuous light. In the record given in Fig. 18 the plant had been kept in the dark and the responses taken in the usual manner. It was then subjected to light; the sudden change from darkness to light acted as a stimulus, inducing a transient depression of excitability. In this connection it is interesting to note that Godlewski found that in the phenomenon of growth, transition from darkness to light acted as a stimulus, causing a transient decrease in the normal rate. The effect of continued light on Mimosa is an enhancement of excitability.
I have often found that the moto-excitability is depressed under excessive turgor. Thus the “over-turgid” leaf of Biophytum sensitivum does not exhibit any mechanical response on rainy days.
Experiment 16.—The effect of excessive turgor on moto-excitability may be demonstrated in the case of Mimosa by allowing its main pulvinus to absorb water. The result is seen in the above record (Fig. 19), where water was applied on the pulvinus after the second response. It is seen how a depression of moto-excitability results from excessive turgor brought on by absorption of water. In such cases, however, the plant is found to accommodate itself to the abnormal condition and gradually regain its normal excitability in the course of one or two hours.
The moto-excitability of the pulvinus of Mimosa is greatly modified under the influence of temperature. For the purpose of this investigation I enclosed the plant in a glass chamber, raising the temperature to the desired degree by means of electric heating. Responses to identical stimuli were then taken at different temperatures. It was found that the effect of heightened temperature, up to an optimum, was to enhance the amplitude of response. Thus with a given specimen it was found that while at 22°C. the amplitude of response was 2.5 mm., it became 22 mm. at 27°C., and 52 mm. at 32°C. The excitability is enhanced under rising, and depressed under falling temperature. The moto-excitability of Mimosa is practically abolished at the minimum temperature of about 19°C.
Effect of lowering of temperature: Experiment 17.—A simple way of exhibiting the effect of lowering of temperature is by artificial cooling of the pulvinus. This cannot very well be done by application of a stream of cooled water, because, as we have seen, absorption of water by the pulvinus is attended by a loss of excitability: diluted glycerine has, however, no such drawback. This fluid at ordinary temperature was first applied on the pulvinus, and after an interval of half an hour records were taken in the usual manner. Cooled glycerine was then applied and the record taken once more; the results are seen in Figs. 20 and 21. In the former, the first response was normal at the temperature of the room, which was 32°C.; the next two exhibit depression of excitability under moderate cooling; the duration of application of moderately cooled glycerine is there indicated by the horizontal line below. On the cessation of application, the normal temperature was quickly restored, with the restoration of normal excitability.
In the next record (Fig. 21) is shown the effect of a more intense cold. It will be noticed that the first effect was a depression, and subsequently, a complete abolition of excitability. Thick dots in the record represent applications of stimulus which proved ineffective. It will also be noticed that even on the cessation of cooling, and the return of the tissue to normal temperature the induced abolition of excitability persisted as an after-effect for a considerable time. I have likewise found that the after-effect of cold in abolishing the conduction of excitation is also very persistent. These experiments show that owing to physiological inertia, the variations of excitability in the plant often lag considerably behind the external changes which induce them.
Effect of high temperature: Experiment 18.—It has been shown that the moto-excitability is enhanced by rising temperature; there is, however, an optimum temperature above which the excitability undergoes a depression. This is seen in the following record (Fig. 22), where the normal response at 32°C. was depressed on raising the temperature to 42°C.; the excitability was, however, gradually restored when the plant was allowed to regain the former temperature.
I may now briefly recapitulate some of the important results: darkness depresses and light exalts the moto-excitability. Excessive turgor depresses motility. Still more marked is the effect of temperature. Lowering of temperature depresses and finally abolishes the moto-excitability: rise of temperature enhances it up to an optimum temperature, but beyond this point the excitability undergoes depression. The change in excitability induced by the variation of external condition is not immediate; the induced effect, generally speaking, lags behind the inducing cause.
I will now give automatic records of responses taken once every hour for twenty-four hours. They prove conclusively the diurnal variation of excitability in Mimosa. After studying in detail the variations characteristic of particular times of the day, I will endeavour to correlate them with the effects brought on by the periodic changes of the environment.
Experiment 19.—As a typical example I will first give a record obtained in the month of February, that is, say, in spring. From this it will not be difficult to follow the variations which take place earlier in winter or later in summer.
The record given in Fig. 23 was commenced at 5 P.M. and continued to the same hour next day. The first thing noticeable is the periodic displacement of the base-line. This is due to the nyctitropic movements of the leaf. It should be remembered that the up-movement of the leaf is represented by down-curve, and vice versâ. After the maximum fall of the leaf, which in this case was attained at 9 P.M., there followed a reverse movement: the highest erection, indicative of maximum turgor, was reached at 6 A.M. The leaf then fell slowly and reached a middle position at noon. The extent of the nyctitropic movement varies in individual cases; in some it is slight, in others very large. The erectile movement began, as stated before, at about 9 P.M.; in some cases, however, it may occur as early as 6 P.M.
In following the characteristic variations of response occurring throughout the day, we find that while they are practically uniform between the hours of 5 and 6 P.M., a continuous decline is manifested after setting in of darkness (7 P.M.); the fall of excitability continues even after sunrise (6-30 A.M.), response being practically abolished at 8 A.M. The excitability is then gradually restored in a staircase manner, the maximum being reached after 12 noon. After attaining this, the excitability remains more or less constant till the evening. It will be noticed that the amplitude of response at 5 P.M. on the second day was the same as the corresponding response on the previous day.
The results of this and numerous other records taken in spring may be summarized as:—
1. The maximum excitability of Mimosa is attained between 1 and 3 P.M., and remains constant for several hours. In connection with the constancy of response at this period, it should be remembered that when the response is at its maximum a slight increase of excitability cannot further enhance the amplitude of response.
2. The excitability, generally speaking, undergoes a continuous decline from evening to morning, the response being practically abolished at or about 8 A.M.
3. From 8 A.M. to 12 noon, the excitability is gradually enhanced in a staircase manner, till the maximum excitability is reached after 1 P.M.
I have obtained numerous records in support of these conclusions, some of which are reproduced in the following figures. In these cases responses to uniform stimuli at intervals of half an hour were taken at different parts of the day, the recorder employed being of the Resonant type.
Mid-day record: Experiment 20.—The record of daily periodicity previously given shows that the excitability reaches its maximum after 12 noon, and that it remains constant at the maximum value for several hours. This fact is fully borne out in the following record obtained with a different specimen (Fig. 24). The responses were taken here from noon to 3 P.M., once every half-hour.
Evening record: Experiment 21.—The record given in Fig. 23 shows that the amplitude of response falls continuously after 6 P.M. It might be thought that the diminished amplitude in the first part may be due to the natural nyctitropic fall of the leaf. The range of the pulvinar movement being limited, it is clear that the extent of the responsive fall must become smaller on account of the natural fall of the leaf during the first part of the night. That this is not the whole explanation of the decline of response in the evening will be clear from certain facts which I will presently adduce. It was stated that the leaf of Mimosa exhibits nyctitropic fall from 6 to 9 P.M., after which there is a reverse movement of erection. In certain specimens, however, the erectile movement commenced as early as 6 P.M. It is obvious that in these latter cases diminution of amplitude of response cannot be due to the reduction of the range of movement of the leaf. In Fig. 25 is given a series of records from 6 to 10 P.M. obtained with a leaf in which erectile movement had commenced early in the evening. Though the full range of responsive movement was in this case available, yet the amplitude of successive responses is seen to undergo continuous diminution.
Record in the morning: Experiment 22.—The excitability is, as we have seen, nearly abolished about 8 A.M., after which there is a gradual restoration. This gradual enhancement of excitability to a maximum in the course of the forenoon is seen well illustrated in the record below (Fig. 26).
The record of daily periodicity given in Fig. 23 may be regarded as a typical example. Modifications may, however, be observed which are traceable to individual peculiarities. As an example of this, I give a record (Fig. 27) obtained with a specimen in which nyctitropic movement was very pronounced. The periodic variation of excitability exhibited here is practically the same as shown by other specimens. The interesting variation is in the character of the recovery from stimulus; the leaf was falling from 6 to 9 P.M.; owing to the shifting of the base-line upwards the recovery appears to be incomplete. After 9 P.M. the leaf was erected, at first slowly, then at a very rapid rate. The consequent fall of the base-line late at night is very abrupt; hence there is an apparent over-shooting in the line of recovery.