Researches on Irritability of Plants
WORKS BY THE SAME AUTHOR.
RESPONSE IN THE LIVING AND NON-LIVING.
With 117 Illustrations, 8vo., 10s. 6d.
1902.
PLANT RESPONSE: AS A MEANS OF PHYSIOLOGICAL INVESTIGATION.
With 278 Illustrations, 8vo., 21s.
1906.
COMPARATIVE ELECTRO-PHYSIOLOGY: A PHYSICO-PHYSIOLOGICAL STUDY.
With 406 Illustrations, 8vo., 15s.
1907.
LONGMANS, GREEN, AND CO.
London, New York, Bombay, and Calcutta
RESEARCHES ON
IRRITABILITY OF PLANTS
BY
JAGADIS CHUNDER BOSE, M.A. D.Sc. C.S.I.
PROFESSOR, PRESIDENCY COLLEGE, CALCUTTA
WITH ILLUSTRATIONS
LONGMANS, GREEN, AND CO.
39 PATERNOSTER ROW, LONDON
NEW YORK, BOMBAY, AND CALCUTTA
1913
dedicated to
s. m.
PREFACE
I have in this work dealing with my researches on the irritability of plants introduced new methods by which the scope of investigation has been enlarged, and a very high degree of accuracy secured. In my previous treatise on Plant Response, the response recorder employed was a modification of the optical lever, automatic records being secured by the very inconvenient and tedious process of photography. The delay thus imposed retarded seriously the progress of the research. Those practically engaged in investigations on plants can realise the difficulties that arise from the too quick passage of the seasons. It thus frequently happened that by the time new instrumental appliances were rendered practicable the favourable season for the plant was over, involving the postponement of the experiment for another year. In spite of these difficulties, the long series of investigations that I then carried out gave many interesting results, which not only threw light on many obscure problems, but also led to the discovery of several important phenomena in plant physiology.
Some of these results, moreover, tended to cast doubt on certain conclusions that had found universal acceptance. It has, for example, been held that there was no transmission of true excitation in Mimosa, the propagated impulse being regarded as merely hydro-mechanical. The question whether the transmitted impulse was physical or physiological could only be satisfactorily decided if the plant could itself be made to record the velocity of its impulse and the changes induced in that velocity under physiological variations. This is but one out of several ideal methods of attacking problems in the life of plants, the realisation of which would make a great advance in physiological investigation.
It would also be desirable to discard, if possible, the troublesome method of obtaining record by photography, which necessitates work in a dark room; in this connection it should be remembered that subjection of the plant to darkness introduces complications by modifying its normal excitability. For these reasons, another requirement which it is necessary to fulfil is the devising of some simple and direct method of obtaining the record. And in order that the results obtained should not be influenced by any personal factor, it would be further desirable that the plant attached to the recording apparatus should be automatically excited by stimulus absolutely constant, should make its own responsive record, going through its own period of recovery, and embarking on the same cycle over again without assistance at any point on the part of the observer.
The difficulties encountered in realising these ideal requirements appeared at first to be insurmountable. In the records of response serious errors occurred as regards amplitude and time-relations, owing to the friction of the writing lever against the recording surface. As an extreme instance of this, in recording the rhythmic movement of the leaflets of Desmodium the very slight friction which the smoked-glass surface offered was enough to stop the pulse-record.
After many attempts, I was at last successful in overcoming all obstacles by the device of the Resonant and Oscillating Recorders. Taking the very difficult test of direct record of the rhythmic movements of Desmodium leaflets, it will be found that the pulsations recorded in this book not only gave accurate measure of the amplitude and period, but also the absolute rate of movement during any phase of their autonomous response. Again, in the matter of accurate measurement of short intervals of time required for the determination of the latent period and velocity of transmission of excitation, I have shown the possibility of recording time-intervals as short as a thousandth part of a second. A brief account of this is given in my paper "On an Automatic Method for the Investigation of the Velocity of Transmission of Excitation in Mimosa," read before the Royal Society. It will be recognised immediately in how many directions our power of inquiry has become extended by the elaboration of these new methods and the invention of several types of instrumental appliances described in this work.
In presenting the results of these investigations, it will be noted that the plant has been made to tell its own story, by means of its self-made records. Each experiment has been repeated at least a dozen times, in many cases as often as a hundred times. The results may therefore be accepted as fully attested. The establishment of the unity of responsive reactions in the plant and animal, which is the subject of this work, will be found highly significant, since it is only by the study of the simpler phenomena of irritability in the vegetal organisms that we can ever expect to elucidate the more complex physiological reactions in the animal tissues.
I take this opportunity to thank my research assistants, Messrs. Guruprasanna Das, L.M.S., and Surendra Chandra Das, M.A., for the very efficient help rendered by them in these researches.
J. C. BOSE.
Presidency College, Calcutta,
October 1912.
CONTENTS
Action of environment on plant—Revelation of internal condition by character of response—Problems to be solved—Electrical response—Mechanical response—Motile organ in Mimosa pudica—Response in plant and animal—Different phases of the responsive movement—Graphic record—Determination of absolute movement of leaf and its time-relations—Characteristic effects of different agencies on the response-curve—Specific difficulties in recording plant-response • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
1 |
THE RESONANT RECORDER
Advantages of intermittent contact in record—Two types of apparatus: the Oscillating Recorder, and the Resonant Recorder—Coercer and Vibrator—Perfect tuning—Recorders with standardised frequencies—Slide and clockwork—The record its own chronogram—Smoked surface and its fixation—Adjustments of the writer—Records with continuous and intermittent contacts • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
11 |
METHODS OF STIMULATION
Different methods of stimulating the plant: mechanical, chemical, thermal and electrical—Difficulties of securing quantitative stimuli—Direct and indirect stimulation—Ideal modes of stimulation—Electro-thermic stimulation—Stimulation by constant current—Stimulation by condenser-discharge—Non-polarisable electrodes—Direct, extra-electrodal, and intra-electrodal stimulation—Stimulation by induction-shock—Effects of make- and break-shock—Excitation by tetanising shock • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
23 |
TIME-RELATIONS OF THE RESPONSIVE MOVEMENT AND STANDARDISATION OF STIMULUS
Latent period of Mimosa—Apex time—Rate of responsive movement of leaf—Effect of intensity of stimulus, fatigue, and temperature—Periodic dot marker—Time relations of response and recovery—Effect of season—Response of Biophytum—Response of Neptunia—Arbitrary distinction between sensitive and ordinary plants—Differential response in Mimosa—Response of ordinary plants—Universal sensitiveness of plants—Standardisation of stimulus—Maximal and Minimal Stimuli—Extreme sensitiveness of Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
35 |
THE ADDITIVE EFFECT; INFLUENCE OF LOAD, TEMPERATURE, AND INTENSITY OF STIMULUS
Greater excitatory efficiency of the break-shock—Additive effect of stimulus—Quantitative relation of additive effect—Effect of load—Thermal chamber—Effect of temperature—Effect of increasing intensity of stimulus on response • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
52 |
VARIOUS TYPES OF RESPONSE
Necessity of uniform stimulation—The Periodic Starter—The Automatic Exciter—Electrolytic contact-maker—The complete Response-recorder—The factor of tonicity—Uniform
responses—Fatigue under shortened period of rest—Growing fatigue—Alternating fatigue—Staircase response—Explanation of erection of leaf under continuous stimulation—Fatigue-relaxation in plant and animal—Response under single stimulus and under tetanisation • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
64 |
EFFECTS OF DIFFERENT GASES ON EXCITABILITY OF MIMOSA
Induced change of excitability under sudden variation of light—Abolition of excitability by absorption of water—Restoration of excitability by application of glycerine—Stimulating, depressing, and toxic agents—Phenomenon of accommodation—Stimulating action of ozone—Effects of carbonic-acid gas, vapour of alcohol, ether, carbon disulphide, coal gas, chloroform, ammonia, sulphuretted hydrogen, laughing-gas, nitrogen dioxide, and sulphur dioxide • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
85 |
DEATH-SPASM IN PLANTS
Criterion of the death of plant—Abolition of electric response at death—Mechanical spasm of death—Water-bath for uniform rise of temperature—Excitatory effect of sudden cooling or heating—Erection of leaf with rising, and depression of leaf with falling, temperature—Thermo-mechanical inversion at the death-point—Necessity for specification of rate of rise of temperature—Death-record of Mimosa—Abolition
of response after death-spasm—Constancy of death-point exhibited by different specimens—Death-records of Desmodium gyrans and Vicia Fava—Death-spasm in ordinary plants—The electric-spasm of death—Lowering of death-point by fatigue and by poisonous solution • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
98 |
DETERMINATION OF THE LATENT PERIOD
Difficulties of accurate determination of Latent Period—Advantages of Resonant Recorder—Simultaneous tracings of tuning-fork exciter and Resonant Recorder—Automatic stimulation at a definite moment—Identical value of latent period in successive determinations—Accurate measurement of time-interval shorter than .005 second—Latent period little affected by inertia of recorder—Tabular statement of value of different specimens of Mimosa—Effect of season on latent period • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
108 |
INFLUENCE OF INTENSITY OF STIMULUS, FATIGUE, AND TEMPERATURE ON THE LATENT PERIOD
Diffuse stimulation under alternating-shock—Effect of intensity of stimulus on Latent Period—Influence of optimum condition—Effect of fatigue—Effect of temperature • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
122 |
VELOCITY OF TRANSMITTED IMPULSE IN PLANTS
Detection of transmitted excitation by means of electromotive variation—Specific tissue for conduction of excitation—Hydro-mechanical theory of transmission of stimulus—Propagation of excitatory protoplasmic change—Physiological test—Automatic record of transmission-period—Conditions for obtaining constant velocity—Determination of velocity of transmission in Mimosa—Differential method of determining velocity—Constancy of results—Tabular statement of different determinations of velocity—Effect of intensity of stimulus on velocity of transmission—Effects on sub-tonic tissue and on tissue in optimum condition—After-effect of stimulus in enhancing conductivity—Effect of optimum condition—Disturbing action of leakage of exciting current—Effect of fatigue—Effect of temperature—Velocity of transmission in Biophytum and Averrhoa—Direction of preferential conduction • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
132 |
EXCITATORY CHARACTER OF TRANSMITTED IMPULSE IN PLANTS
The hydro-mechanical theory—Inconclusive character of the anæsthetic experiment of Pfeffer and scalding experiment of Haberlandt—Kühne's experiment showing transmission of excitation under intense stimulation in a rigored nerve—Error introduced by employment of excessive intensities of stimulus—Discriminative polar effect of current in excitation—Block of transmission of excitation by local application of cold—Restoration of normal conductivity by tetanising shock in tissue paralysed by cold—Electro tonic arrest of excitatory impulse—Action of various poisons in inducing block of conduction • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
154 |
THE POSITIVE RESPONSE
Two opposite kinds of responses, negative and positive—Excitatory contraction, negative turgidity variation, fall of leaf, and concomitant negative electric variation—Positive electric response—Positive or erectile mechanical response—Dual impulses under different forms of stimuli—Exhibition of positive and negative impulses by different plants—Conditions for obtaining positive response—Characteristics of
positive impulse—Masking and unmasking of positive effect—Laws of Direct and Indirect effects of stimulus • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
176 |
POLAR EFFECTS OF ELECTRICAL CURRENT IN EXCITATION OF PLANTS
Polar excitation in animal tissues—Anomalous reactions in Protozoa—Mono-polar method—Current Reverser—Excitatory polar action in plant—Method of record—Effects of ascending and descending currents in animal and plant—Records giving time-relations—Potential slide—Measurement of e.m.f. and current—Potential keyboard • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
198 |
POLAR EFFECTS OF FEEBLE AND MODERATE CURRENTS ON VARIOUS SENSITIVE PLANTS
Polar effects of feeble and moderate currents on (1) leaflets of Mimosa, (2) leaflets of Biophytum, (3) leaflets of Neptunia, (4) leaflets of Averrhoa carambola, (5) leaflets of Averrhoa bilimbi, and (6) primary leaf of Mimosa—Excitation with feeble current only at kathode-make—Excitation at kathode-make and anode—break, under moderate current—Tabular statement of results • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
212 |
THE CONTRASTED EFFECTS OF ANODE AND KATHODE
Polar effects of currents on pulsation of Desmodium gyrans—Reduction of systolic contraction by anodic action—Diminution of diastolic expansion by kathodic action—Arrest at systole by make of kathode and diastolic expansion by break of kathode—Arrest at diastole by make of anode, and systolic contraction by break of anode—Effects of ascending and descending currents of feeble and strong intensity in nerve-and-muscle preparation—Parallel effects in petiole-and-pulvinus • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
234 |
EFFECT OF TEMPERATURE ON POLAR EXCITATION, AND MULTIPLE EXCITATION UNDER CONSTANT CURRENT
Excitability of nerve to induction-shock diminished by cooling—Nerve excitation by constant current enhanced by cooling—Excitation of conducting-tissue of Mimosa by constant current enhanced by cooling and depressed by warming—Ineffective stimulus becoming effective under cooling and vice versa—Multiple response induced in Biophytum by the passage of constant current—Comparison of sensitiveness of plant and animal—Minimum current for excitation of human tongue—Relatively higher sensitiveness of Biophytum • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
244 |
POLAR EFFECTS UNDER STRONG CURRENTS
Abnormal polar reactions in Protozoa—Transformation of polar reaction in leaf of Mimosa from Type II. to Type III. under strong current—Further transformation to Type IV. under stronger current—Exhibition of Type III. and Type IV. by leaflets of Mimosa, Biophytum, Averrhoa carambola—Law of polar action of strong currents • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
253 |
VARIATION OF POLAR REACTION UNDER TISSUE MODIFICATION
Modification of polar reaction under tissue-changes—Effect of age—After-effect of moderate stimulation—Modified polar effect; excitation at kathode-make and anode-make; excitation at kathode-make, kathode-break, and anode-make—General review of polar reactions • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
266 |
MULTIPLE AND AUTOMATIC RESPONSE
The Oscillating Recorder—Latent period of Biophytum—Refractory period—Response on 'all-or-none' principle—Multiple electrical response to a single strong stimulus—Multiple mechanical response to strong stimulus in Biophytum and Averrhoa—Continuity of multiple and automatic response—Ordinarily responding Biophytum converted into automatically responding condition by excess of stored energy—Automatically responding Desmodium converted to ordinarily responding condition by depletion of stored energy • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
278 |
THE AUTOMATIC PULSATIONS OF DESMODIUM GYRANS
Activity of detached leaflet of Desmodium—Pulsation maintained uniform under constant internal hydrostatic pressure—The plant-chamber—Time-relations of pulsating movement derived from dotted record—Significance of down and up movements—Systole and diastole—Table showing rates of movement of Desmodium leaflet at different phases • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
290 |
EFFECT OF HYDROSTATIC PRESSURE, LOAD, AND LIGATURE ON THE PULSATION OF DESMODIUM
Effect of internal hydrostatic pressure on the pulsation of Desmodium—Expansive erection of leaflet under increased pressure; diminution of the extent of systolic contraction—Effect of load: diminution of period—Stannius' ligature on heart-beat—Parallel effect of ligature on pulsation of Desmodium—Arrest of pulsation by a cut and revival by electric shock • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
300 |
EFFECT OF STIMULUS ON LEAFLET OF DESMODIUM AT STANDSTILL
Condition of standstill brought about by depletion of energy—Renewal of pulsation by the stimulus of light—Response to stimulus of induction-shock—Multiple response under tetanisation—Determination of the latent period and the apex time—Refractory period—Effect of stimulus on leaflets in sub-tonic condition—Effect of isolation on rhythmic activity—Gradual arrest of pulsation resulting from run-down of stored energy—Effect of fresh accession of energy • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
306 |
EFFECT OF ELECTRIC STIMULATION ON THE PULSATION OF DESMODIUM GYRANS
Effect of electric shock on Desmodium leaflet—Incapability of
tetanus—Extra pulsation induced by electric shock—Relative effectiveness of electric stimulus at diastolic phase—Effect of transmitted excitation on normal pulsation of heart and on pulsation of Desmodium leaflet—Effects of acceleration and inhibition • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
318 |
EFFECT OF TEMPERATURE ON RHYTHMIC PULSATION OF DESMODIUM GYRANS
Effect of lowering of temperature on rhythmic pulsation of cardiac tissue—Similar effect on the pulsation of Desmodium—Increase of systolic limit during cooling—Minimum temperature for arrest of pulsation—Arrest by cooling and subsequent revival by warming—Increase of diastolic limit during warming—Effect of rise of temperature on the pulsation of frog’s heart—Similar effect on the pulsation of Desmodium—Effect of rise above and return to normal temperature—Diminution of systolic contraction during rise of temperature—Increase of systolic contraction during fall of temperature—Permanent arrest due to heat-rigor • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
323 |
EFFECT OF CHEMICAL AGENTS ON THE AUTOMATIC PULSATION OF DESMODIUM GYRANS
Application of gaseous or liquid reagents—Modifying influence of tonic condition of specimen, strength, and duration of application—Effect of sugar solution—Effect of alcohol—Action of carbonic-acid gas—Effects of anæsthetics, ether, and chloroform—Action of carbon disulphide—Effect of copper sulphate solution—Effect of potassium cyanide solution—Antagonistic actions of acids and alkalis on the pulsations of the heart and of Desmodium—Similarities of reaction in rhythmic tissues animal and vegetal • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
332 |
GENERAL SURVEY
342
• • • • • • • • • • • • • • • • • • • • • • • • • • • • |
361 |
• • • • • • • • • • • • • • • • • • • • • • • • • • • • |
369 |
ILLUSTRATIONS
figure.
page
| 1. | Diagrammatic representation of Response Recorder • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
6 |
| 2. | Response curve of leaf of Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
7 |
| 3. | Resonant Recorder, upper part • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
17 |
| 4. | General View of the Resonant Recorder and accessories • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
20 |
| 5. | Record showing advantage of intermittent contact • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
21 |
| 6. | Electro-thermic stimulator • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
25 |
| 7. | Response of Mimosa to indirect thermal stimulation • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
26 |
| 8. | Response to stimulus of constant electric current • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
27 |
| 9. | Direct stimulation by condenser discharge • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
28 |
| 10. | Record of response to stimulation by condenser discharge • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
30 |
| 11. | Arrangement for applying single make- or break-shock • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
32 |
| 12. | Records giving apex time of response of Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
38 |
| 13. | The dot marker • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
40 |
| 14. | Response of Mimosa giving time-relations • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
41 |
| 15. | Record of response of leaflet of Biophytum • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
43 |
| 16. | Response of leaf of Neptunia • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
43 |
| 17. | Erectile response of leaf of Mimosa, due to local stimulation of upper half of pulvinus • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
45 |
| 18. | Responses of leaf of Desmodium gyrans • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
46 |
| 19. | The electric signal • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
52 |
| 20, 21. | Records showing greater efficiency of break-shock • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
53 |
| 22, 23, 24, 25. | Additive effects of stimulus • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
54-56 |
| 26. | Effect of load on response of Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
57 |
| 27, 28. | Effect of temperature on response of Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
60, 61 |
| 29. | Increasing response under increasing stimulation • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
62 |
| 30. | Periodic starter, and automatic exciter • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
66 |
| 31. | Electrolytic contact-maker • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
68 |
| 32. | Photograph of duplex type of Resonant Recorder • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
69 |
| 33. | Uniform responses of Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
72 |
| 34. | Fatigue under shortened period of rest • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
73 |
| 35. | Growing fatigue • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
74 |
| 36. | Periodic fatigue • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
74 |
| 37. | Staircase response in frog's muscle • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
77 |
| 38. | Staircase response in Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
77 |
| 39. | Effect of stimulus in modifying the tonic condition • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
79 |
| 40. | Alternating response • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
81 |
| 41. | Fatigue-decline in frog's muscle • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
81 |
| 42. | Different phases in the fatigue reversal in plant • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
83 |
| 43. | Fatigue reversal in Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
83 |
| 44. | Effect of sudden darkness on excitability of Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
| 45. | Abolition of motile excitability of pulvinus by absorption of water • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
86 |
| 46. | Stimulating action of ozone • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
90 |
| 47. | Effect of CO2 on excitability of Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
90 |
| 48. | Effect of vapour of alcohol • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
91 |
| 49. | Effect of ether • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
92 |
| 50. | Effect of carbon disulphide • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
93 |
| 51. | Effect of coal gas • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
93 |
| 52. | Abolition of excitability under chloroform • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
94 |
| 53. | Effect of ammonia • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
95 |
| 54. | Effect of sulphuretted hydrogen • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
95 |
| 55. | Effect of nitrogen dioxide • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
96 |
| 56. | Effect of sulphur dioxide • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
96 |
| 57. | Excitatory effect on Mimosa by sudden cooling or warming • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
100 |
| 58. | Death-curve of Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
102 |
| 59. | Abolition of response to warming or cooling after passing through the death-point • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
103 |
| 60. | Death-curve of Desmodium gyrans • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
104 |
| 61. | Thermo-mechanical inversion indicating death-point of leaf of bean • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
105 |
| 62. | Lowering of death-point under fatigue • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
106 |
| 63. | Effect of poison in lowering the death-point • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
106 |
| 64. | Latent period of hyoglossus muscle • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
109 |
| 65. | Simultaneous record of vibrating recorder and exciting tuning fork • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
110 |
| 66. | Apparatus for determination of latent period of Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
112 |
| 67. | Two successive records exhibiting identity of latent period of Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
115 |
| 68. | Record of latent period of highly excitable Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
116 |
| 69. | Record of latent period with a 200 D.V. recorder • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
117 |
| 70. | The same record magnified • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
118 |
| 71, 72. | Two records obtained with two different recorders • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
119 |
| 73. | Record of latent period of Neptunia • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
120 |
| 74. | Simultaneous excitation in interposed tract under alternating-shock • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
124 |
| 75, 76. | Effect of intensity of stimulus on latent period • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
126 |
| 77. | Constancy of latent period under stimuli above the maximal • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
127 |
| 78, 79. | Effect of fatigue • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
128 |
| 80, 81. | Effect of temperature • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
130 |
| 82. | Determination of velocity of transmission of excitation in Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
137 |
| 83, 84. | Determination of velocity by Differential Method • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
139, 140 |
| 85. | Effect of intensity of stimulus on velocity and after-effect • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
143 |
| 86. | Effect of optimum condition on velocity • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
145 |
| 87. | Effect of fatigue on velocity of transmission • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
147 |
| 88. | Effect of temperature on velocity • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
149 |
| 89. | Record giving transmission time in Biophytum • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
151 |
| 90. | Effect of cold in the retardation and arrest of transmission • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
162 |
| 91. | Arrangement for electrotonic block • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
165 |
| 92. | Record of effect of electrotonic block • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
166 |
| 93. | Records of transmitted excitation with the block off and on • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
167 |
| 94. | Effect of CuSO4 in abolishing conduction • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
172 |
| 95. | Abolition of conductivity by KCN • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
173 |
| 96. | Positive response followed by negative in Biophytum under indirect thermal stimulus • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
180 |
| 97. | Diphasic response in Biophytum under indirect chemical stimulation • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
182 |
| 98, 99. | Positive followed by negative in Averrhoa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
184 |
| 100. | Diphasic response in Mimosa due to indirect electric stimulation of petiole • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
185 |
| 101. | Diphasic response in Mimosa due to indirect thermal stimulation of stem • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
186 |
| 102. | Effect of intensity of stimulus in modifying the diphasic response • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
187 |
| 103. | Effect of diminishing distance in transforming positive into diphasic response in Biophytum • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
189 |
| 104. | Effect of diminishing distance on the response in Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
194 |
| 105. | Masking of the positive by the predominant negative • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
194 |
| 106. | • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
200 |
| 107. | Record of polar excitation under feeble current • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
202 |
| 108. | Excitation by ascending and descending current • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
204 |
| 109. | Records of responses to ascending and descending current in Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
205 |
| 110. | Records of responses to ascending and descending induction-shock • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
206 |
| 111. | The Potential Slide • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
208 |
| 112. | The Potential Keyboard • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
210 |
| 113. | The electrode holder • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
217 |
| 114. | Excitation induced in Biophytum by the make of kathode and break of anode • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
224 |
| 115. | Record of kathode-make and anode-break in Mimosa • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
230 |
| 116. | Effect of anode, opposing systolic contraction in Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
236 |
| 117. | Effect of kathode, opposing diastolic expansion in Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
236 |
| 118. | Alternate effect of anode and kathode • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
237 |
| 119. | Arrest at systole by make of kathode, and diastolic expansion at break of kathode • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
237 |
| 120. | Arrest at diastole by make of anode, and systolic contraction at break of anode • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
237 |
| 121. | Effects of descending and ascending currents at make and break on nerve-muscle and petiole-pulvinus • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
239 |
| 122. | Effect of cold on excitability to induction-shock • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
240 |
| 123. | Record showing abolition of polar excitation at high temperature • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
247 |
| 124. | Multiple excitation in Biophytum under constant current • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
249 |
| 125. | Record of polar excitation of Type III. • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
255 |
| 126. | Polar excitation Type IV. • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
256 |
| 127. | Transformation of Type II. to Type III. as after-effect of previous stimulation • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
268 |
| 128. | Abrupt transition from Type I. to Type III. • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
271 |
| 129. | Polar reactions, Km, Km Am, and Km Am Ab under gradually increasing current • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
273 |
| 130. | Record exhibiting modified response Km Kb Am • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
275 |
| 131. | The Oscillating Recorder • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
279 |
| 132. | Record giving latent period of Biophytum • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
281 |
| 133. | Record of responses of Biophytum to stimuli ·1 and 1 unit • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
282 |
| 134. | Responses of Biophytum to stimuli ·1, ·5, 1, and 2 units • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
284 |
| 135. | Multiple response in Biophytum under strong electric shock • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
284 |
| 136. | Multiple response in Averrhoa under a single strong electric shock • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
285 |
| 137. | Multiple response under constant stimulus of light • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
285 |
| 138. | Multiple response under single strong thermal shock • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
286 |
| 139. | Multiple response induced by strong chemical stimulation • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
287 |
| 140. | Leaf of Desmodium gyrans • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
290 |
| 141. | U-tube support and plant-chamber • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
292 |
| 142. | Continuous record of pulsations of Desmodium leaflet for four hours • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
294 |
| 143. | Record of a single pulsation of Desmodium giving time-relations • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
296 |
| 144. | Record of two successive pulsations • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
297 |
| 145. | Series of automatic pulsations of Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
298 |
| 146. | Effect of application of increased internal hydrostatic pressure on the pulsation of Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
301 |
| 147. | Effect of increasing load on Desmodium pulsation • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
302 |
| 148. | Effect of ligature in inducing arrest of pulsation of Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
303 |
| 149. | Arrest of pulsation by cut and revival by electric shock • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
304 |
| 150. | Action of light in renewing pulsation of Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
307 |
| 151. | Response of Desmodium leaflet, originally in a state of stand-still • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
308 |
| 152. | Multiple response under tetanisation • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
308 |
| 153. | Response of Desmodium exhibiting apex time • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
309 |
| 154. | Record exhibiting refractory period in Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
310 |
| 155. | After-effect of stimulation on pulsation of Desmodium in subtonic condition • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
312 |
| 156. | Effect of depletion of energy on pulsation of isolated Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
313 |
| 157. | Gradual stoppage of pulsation in isolated leaflet of Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
313 |
| 158. | Effect of stimulus in renewing pulsation of Desmodium, brought to standstill • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
315 |
| 159. | Response of Desmodium leaflet to stimulus of light • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
316 |
| 160. | Response of a depressed specimen of Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
316 |
| 161. | Effects of strong tetanisation on Desmodium pulsation • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
318 |
| 162, 163. | Extra pulsation induced by induction-shock applied at diastolic phase • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
319 |
| 164. | Inhibitory effect of transmitted excitation on the pulsation of vigorous leaflet of Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
320 |
| 165. | Augmentation of pulsation induced by transmitted excitation in less vigorous specimen of Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
321 |
| 166. | Effect of lowering of temperature on the pulsation of heart • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
324 |
| 167. | Effect of lowering of temperature on pulsation of Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
325 |
| 168, 169. | Effect of rapid cooling on Desmodium pulsation • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
326 |
| 170. | Effect of rise of temperature on amplitude and frequency of pulsation of the heart of frog • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
327 |
| 171, 172. | Effect of rise of temperature on the pulsation of Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
328 |
| 173. | Effect of continuous rise of temperature from 30° C. to 38·5° C. • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
329 |
| 174. | Effect of continuous rise from 30° C. to 42° C. and return to 30° C. • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
330 |
| 175. | Effect of rise of temperature and return • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
330 |
| 176. | Effect of dilute sugar solution • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
333 |
| 177. | Effect of alcohol on the pulsation of Desmodium • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
333 |
| 178. | Effect of internal application of strong alcohol • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
334 |
| 179. | Effect of dilute carbonic-acid gas • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
334 |
| 180. | Effect of strong carbonic-acid gas • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
335 |
| 181. | Effect of internal application of carbonic acid • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
335 |
| 182. | Effect of vapour of ether • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
336 |
| 183. | Effect of vapour of chloroform • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
337 |
| 184. | Effect of carbon disulphide • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
337 |
| 185. | Effect of copper sulphate solution • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
338 |
| 186. | Effect of potassium cyanide • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
339 |
| 187. | Arrest of pulsation of the heart of frog at diastole by the action of dilute lactic acid • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
339 |
| 188. | Arrest of pulsation of Desmodium at diastole by application of dilute lactic acid • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
340 |
| 189. | Arrest of pulsation of heart at systole, by the action of dilute sodium hydrate • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
340 |
| 190. | Arrest of pulsation of Desmodium at systole by the application of dilute solution of sodium hydrate • • • • • • • • • • • • • • • • • • • • • • • • • • • • |
340 |
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