Chapter 28

PLANT RESPONSE--ON THE INFLUENCE OF TEMPERATURE

Effect of very low temperature--Influence of high temperature--Determination of death-point--Increased response as after-effect of temperature variation--Death of plant and abolition of response by the action of steam.

For every plant there is a range of temperature most favourable to its vital activity. Above this optimum, the vital activity diminishes, till a maximum is reached, when it ceases altogether, and if this point be maintained for a long time the plant is apt to be killed. Similarly, the vital activity is diminished if the temperature be lowered below the optimum, and again, at a minimum point it ceases, while below this minimum the plant may be killed. We may regard these maximum and minimum temperatures as the death-points. Some plants can resist these extremes better than others. Length of exposure, it should however be remembered, is also a determining factor in the question as to whether or not the plant shall survive unfavourable conditions of temperature. Thus we have hardy plants, and plants that are affected by excessive variations of temperature. Within the characteristic power of the species, there may be, again, a certain amount of individual difference.

These facts being known, I was anxious to determine whether the undoubted changes induced by temperature in the vital activity of plants would affect electrical response.

#Effect of very low temperature.#--As regards the influence of very low temperature, I had opportunities of studying the question on the sudden appearance of frost. In the previous week, when the temperature was about 10° C., I had obtained strong electric response in radishes whose value varied from ·05 to ·1 volt. But two or three days later, as the effect of the frost, I found electric response to have practically disappeared. A few radishes were, however, found somewhat resistant, but the electric response had, even in these cases, fallen from the average value of ·075 V. under normal temperature to ·003 V. after the frost. That is to say, the average sensitiveness had been reduced to about 1/25th. On warming the frost-bitten radish to 20° C. there was an appreciable revival, as shown by increase in response. In specimens where the effect of frost had been very great, i.e. in those which showed little or no electric response, warming did not restore responsiveness. From this it would appear that frost killed some, which could not be subsequently revived, whereas others were only reduced to a condition of torpidity, from which there was revival on warming.

I now tried the effect of artificial lowering of temperature on various plants. A plant which is very easily affected by cold is a certain species of Eucharis lily. I first obtained responses with the leaf-stalk of this lily at the ordinary temperature of the room (17° C.). I then placed it for fifteen minutes in a cooling chamber, temperature -2° C., for only ten minutes, after which, on trying to obtain response, it was found to have practically disappeared. I now warmed the plant by immersing it for awhile in water at 20° C., and this produced a revival of the response (fig. 35). If the plant be subjected to low temperature for too long a time, there is then no subsequent revival.

I obtained a similar marked diminution of response with the flower-stalk of Arum lily, on lowering the temperature to zero.

My next attempt was to compare the sensibility of different plants to the effect of lowered temperatures. For this purpose I chose three specimens: (1) Eucharis lily; (2) Ivy; and (3) Holly. I took their normal response at 17° C., and found that, generally speaking, they attained a fairly constant value after the third or fourth response. After taking these records of normal response, I placed the specimens in an ice-chamber, temperature 0° C., for twenty-four hours, and afterwards took their records once more at the ordinary temperature of the room. From these it will be seen that while the responsiveness of Eucharis lily, known to be susceptible to the effect of cold, had entirely disappeared, that of the hardier plants, Holly and Ivy, showed very little change (fig. 36).

Another very curious effect that I have noticed is that when a plant approaches its death-point by reason of excessively high or low temperature, not only is its general responsiveness diminished almost to zero, but even the slight response occasionally becomes reversed.

#Influence of high temperature, and determination of death-point.#--I next tried to find out whether a rise of temperature produced a depression of response, and whether the response disappeared at a maximum temperature--the temperature of death-point. For this purpose I took a batch of six radishes and obtained from them responses at gradually increasing temperatures. These specimens were obtained late in the season, and their electric responsiveness was much lower than those obtained earlier. The plant, previously kept for five minutes in water at a definite temperature (say 17° C.), was mounted in the vibration apparatus and responses observed. The plant was then dismounted, and replaced in the water-bath at a higher temperature (say 30° C.) again, for five minutes. A second set of responses was now taken. In this way observations were made with each specimen till the temperature at which response almost or altogether ceased was reached. I give below a table of results obtained with six specimens of radish, from which it would appear that response begins to be abolished in these cases at temperatures varying from 53° to 55° C.

TABLE SHOWING EFFECT OF HIGH TEMPERATURE IN ABOLISHING RESPONSE

Temperature Galvanometric response (100 dns. = ·07 V.)

(1) {17° C 70 dns. {53° " 4 "

(2) {17° " 160 " {53° " 1 "

(3) {17° " 100 " {50° " 2 "

(4) {17° " 80 " {55° " 0 "

(5) {17° " 40 " {60° " 0 "

(6) {17° " 60 " {55° " 0 "

#Electric heating.#--The experiments just described were, however, rather troublesome, inasmuch as, in order to produce each variation of temperature, the specimen had to be taken out of the apparatus, warmed, and remounted. I therefore introduced a modification by which this difficulty was obviated. The specimen was now enclosed in a glass chamber (fig. 37), which also contained a spiral of German-silver wire, through which electric currents could be sent, for the purpose of heating the chamber. By varying the intensity of the current, the temperature could be regulated at will. The specimen chosen for experiment was the leaf-stalk of celery. It was kept at each given temperature for ten minutes, and two records were taken during that time. It was then raised by 10° C., and the same process was repeated. It will be noticed from the record (fig. 38) that in this particular case, as the temperature rose from 20° C. to 30° C., there was a marked diminution of response. At the same time, in this case at least, recovery was quicker. At 20° C., for example, the response was 21 dns., and the recovery was not complete in the course of a minute. At 30° C., however, the response had been reduced to 7·5 divisions, but there was almost complete recovery in twelve seconds. As the temperature was gradually increased, a continuous decrease of response occurred. This diminution of response with increased temperature appears to be universal, but the quickening of recovery may be true of individual cases only.

TABLE SHOWING DIMINUTION OF RESPONSE WITH INCREASING TEMPERATURE

(·01 Volt = 35 divisions)

Temperature Response

20° 21 30° 7·5 40° 5·5 50° 4 65° 3

In radishes response disappeared completely at 55° C., but with celery, heated in the manner described, I could not obtain its entire abolition at 60° C. or even higher. A noticeable circumstance, however, was the prolongation of the period of recovery at these high temperatures. I soon understood the reason of this apparent anomaly. The method adopted in the present case was that of dry heating, whereas the previous experiments had been carried on by the use of hot water. It is well known that one can stand a temperature of 100° C. without ill effects in the hot-air chamber of a Turkish bath, while immersion in water at 100° C. would be fatal.

In order to find out whether subjection to hot water would kill the celery-stalk, I took it out and placed it for five minutes in water at 55° C. This, as will be seen from the record taken afterwards, effectively killed the plant (fig. 38, w).

#Increased sensitiveness as after-effect of temperature variation.#--A very curious effect of temperature variation is the marked increase of sensitiveness which often appears as its after-effect. I noticed this first in a series of observations where records were taken during the rise of temperature and continued while the temperature was falling (fig. 39). The temperature was adjusted by electric heating. It was found that the responses were markedly enhanced during cooling, as compared with responses given at the same temperatures while warming (see table). Temperature variation thus seems to have a stimulating effect on response, by increasing molecular mobility in some way. The second record (fig. 40) shows the variation of response in Eucharis lily (1) during the rise, and (2) during the fall of temperature. Fig. 41 gives a curve of variation of response during the rise and fall of temperature.

TABLE SHOWING THE VARIATION OF RESPONSE IN SCOTCH KALE DURING THE RISE AND FALL OF TEMPERATURE

Temperature Response Response [Temperature rising] [Temperature falling]

19° C. 47 dns. -- 25° " 24 " -- ^ 30° " ¦ 11 " 23 dns. ¦ 50° " ¦ 8 " 16 " ¦ 70° " v 7 " -- ----->

#Point of temperature maximum.#--We have seen how, in cases of lowered temperature, response is abolished earlier in plants like Eucharis, which are affected by cold, than in the hardier plants such as Holly and Ivy. Plants again are unequally affected as regards the upper range. In the case of Scotch kale, for instance, response disappears after ten minutes of water temperature of about 55° C., but with Eucharis fairly marked response can still be obtained after such immersion and does not disappear till it has been subjected for ten minutes to hot water, at a temperature of 65° C. or even higher. The reason of this great power of resistance to heat is probably found in the fact that the Eucharis is a tropical plant, and is grown, in this country, in hot-houses where a comparatively high temperature is maintained.

#The effect of steam.#--I next wished to obtain a continuous record by which the effects of suddenly increased temperatures, culminating in the death of the plant, might be made evident. For this purpose I mounted the plant in the glass chamber, into which steam could be introduced. I had chosen a specimen which gave regular response. On the introduction of steam, with the consequent sudden increase of temperature, there was a transitory augmentation of excitability. But this quickly disappeared, and in five minutes the plant was effectively killed, as will be seen graphically illustrated in the record (fig. 42).

It will thus be seen that those modifications of vital activity which are produced in plants by temperature variation can be very accurately gauged by electric response. Indeed it may be said that there is no other method by which the moment of cessation of vitality can be so satisfactorily distinguished. Ordinarily, we are able to judge that a plant has died, only after various indirect effects of death, such as withering, have begun to appear. But in the electric response we have an immediate indication of the arrest of vitality, and we are thereby enabled to determine the death-point, which it is impossible to do by any other means.

It may be mentioned here that the explanation suggested by Kunkel, of the response being due to movement of water in the plant, is inadequate. For in that case we should expect a definite stimulation to be under all conditions followed by a definite electric response, whose intensity and sign should remain invariable. But we find, instead, the response to be profoundly modified by any influence which affects the vitality of the plant. For instance, the response is at its maximum at an optimum temperature, a rise of a few degrees producing a profound depression; the response disappears at the maximum and minimum temperatures, and is revived when brought back to the optimum. Anæsthetics and poisons abolish the response. Again, we have the response undergoing an actual reversal when the tissue is stale. All these facts show that mere movement of water could not be the effective cause of plant response.