有色光線ニ對スル鱗翅目幼蟲ノ反應ニ就テ = The Relative Stimulating Efficiency of the Different Wave Lengths of Light for the Phototropic Reactions of a Certain Lepidopterous Larva
저자
林泉 (京城帝國大學農學部昆蟲學硏究室)
발행기관
학술지명
권호사항
발행연도
1931
작성언어
Japanese
KDC
180
자료형태
학술저널
수록면
49-68(20쪽)
제공처
The problem as to precisely what wave lengths of light stimulate animal most is perhaps the most difficult of all the problems encountered in the investigation of animal conducts. It is easy to observe that animals react to colored lights, but it is quite difficult to determine whether such responses are brought about by the quality or quantity of the light, that is, by actual color or by effective intensity of light. There are much confusion in opinions concerning the problem and, as is expected in the case, great variations in results obtained as are shown by Gross '13, Mast '17, Loeb & Wasteneys '17, Washburn '17, Ku¨hn '27, etc. Many of those erroneous conclusions have been derived, first, because almost all experiments were carried out with animals which were brought in from different unknown sources and also without sufficient control experiments and, secondly, because the investigators, as a rule, have had d poor knowledge of the physics of colored lights.
The writer used in this investigation certain lepidopterous larvae, Malacosoma neustria testacea Motsch., which were reared in the laboratory under well defined conditions. Moreover, each of these animals was run through the experiment in total darkness either at the beginning or after the experiments with the colored lights so as to determine the specific behavior of individual animal. These experiments may be considered as controls of the light reactions of the animal.
The experiments with colored lights were conducted by a method essentially similar to that used in an investigation of orientation of the animal in a field of light where beams of two monochromatic lights crossed each other at a right angle. The detailed arrangements of one of the two sets of apparatus is shown in Fig. I and the light field so prepared is diagrammatically represented in Fig. II. The colored lights for the experiments were obtained by using a set of 7 Wratten filters Nos. 70-76 and in order to equalize the intensity of these colored lights a small concentrated filament lamp of 49.3mc., 81.0mc., 129.1mc. or 171.4mc. was used according to the need. The accurate adjustments of the intensities of two beams compared were accomplished by means of a flicker-photometer. Consequently, it is evident that we have a method here by means of which the stimulating effect of lights differing in wave lengths can be compared and their relative stimulating efficiency ascertained.
The results obtained in these experiments may be briefly summarized as follows.
I. The Behavior in a Dark Room.
As was already reported by the writer in 1928, some of these animals creeping on a smoked paper in the total absence of photic stimulation looped in a fairly constant fashion to the right or the left, a few were extremely variable, while still others moved in rather straight courses as is shown in Fig. III.
II. The Behavior in Illuminated Conditions.
Under the influence of an illumination most animals moved nearly in a straight course. Those several types of the behaviors as ascertained by the experiments in the total darkness have disappeared. Such a difference in the behavior as mentioned above has already been noted by several investigators e. g. Doflein '16, v. Buddenbrock '17, Minnich '19 and Hayashi '28, etc. Although most animals followed a nearly straight course in an illuminated field, the writer has found that there are great difference in the direction of movements, that is, some of them moved towards the source of light on a course approximately parallel to the rays of light while some animals deviated within some degrees from the direction of beams. The degrees of these deflections differed greatly by the species of animals used and by the factor of time for the same individual animal. For example, in the case of the larvae of Malacosoma neustria which are somewhat in a starved condition the deflection is not evident and moves more strictly on a straight line, but on the contrary, if they are fully fed or in such a stage as before the pupation the deflection may be more apparent than in the former condition. And again, in the case of the normal larvae of Malacosoma meustria, it is, as a rule, not so evident as those of Barathra brassicae L. which are in general not so sensitive to the photic stimulation. Thus the more insensitive the animals are the more deflections may appear.
III. The Difference of Behavior of the Same Individual Animal under the Conditions of both the Illuminated and Dark Fields.
In these cases the most significant discovery is the fact that the same animal which looped in a room of total darkness also curved with a striking similarity in an illuminated field, e. g. those which curved towards the same direction in the total darkness, although their curvatures were, as a rule, greatly reduced in an illuminated condition. Those which moved in a rather straight course in a total darkness oriented more exactly on the straight line. All these cases indicate the fact that there are thus some closely correlated relationships in the behaviors in the illuminated and dark conditions.
IV. The Kinetic Influence of Some Colored Lights.
Colored lights such as the green (530μμ), blue-green (460μμ), blue (450μμ) and yellow (578μμ) exert a kinetic influence upon the animal, that is, they tend to induce activity as is the case of the white light.
V. The Reaction to Two Sources of Light.
If the animal which moved in a nearly straight course towards a source of light is simultaneously exposed to two lights of the same quality and quantity, then the animal turns directly to neither light but orients itself so that its longitudinal axis is in a line which bisects the angle between the two lights, and if the lights are unequal in one of these factors, the intensity and the wave length, the animal deviates towards the stronger stimulating light and orients in such a position where the effective illumination on the two sides of the body is equal. Therefore the behavior of these animals is a typical "Tropotaxis" and the characteristic feature of the behavior is the point that the course of their movements is almost always a strictly straight one when their rate of locomotion is moderate. In the colored light experiments the writer used only these animals which oriented themselves in a tropotactic fashion as mentioned above.
VI. The Order of the Relative Stimulating Efficiency of the Different Wave Lengths of Equal Intensity.
For the larvae of Malacosoma neustria the order of the stimulating efficiency in the spectrum is determined experimentally as follows.
680 < 650 < 610 < 578 < 450 < 490 < 530μμ or,
Dark red < Red < Orange < Yellow < Blue < Blue-green < Green.
This conclusion is derived from the data presented in Table I. It is therefore evident that the light of mean wave length 530μμ (green) is the most effective one for the animal studied.
VII. The Photochemical Basis of the Reaction to Certain Colored Lights.
The fact that the green ray is the most efficient in causing phototropic reactions in the animal investigated suggests that in such an animal the photosensitive substance responsible for the reaction is most rapidly decomposed by light of λ=530μμ and that the following formula of light reactions which was proposed by Hecbt in the case of a white light is also applicable to this case in a more definite sense. That is,
Light → S → P+A; L ∥ P+A ∥ → T → Impulse
Here, S represents photosensitive substance, P and A precursors, L neutral substance, T stimulating substance and ∥P+A∥ a catalytic reaction by one or two of the precursors. It is assumed moreover that the reactions in the rectangle take place inside the sense cell.
VIII. The Relative Stimulating Efficiency of the Different Wave Lengths of Equal Intensity.
When Malacosoma larvae are subjected to two lights of different quality but of equal quantity crossing each other at a right angle the amount of turning of the body towards the more effective light, R, is expressed by the following equation :
R = k. t. log Q₁/ Q₂,
in which Q₁/ Q₂is the ratio between the specific activities of the two lights. R may be also expressed in this connection as follows :
R = k. tan α ,
where α is the angle of deflection (See Fig. II. α = A< B C′= 45˚-C< B C′). The values of tan α as shown in the relative stimulating effects of the different wave lengths were calculated and the result is expressed graphically in Fig. IV. As is evident in Fig. IV, the stimulating efficiency decreases from the region of the maximam rather rapidly in the direction of the more refrangeble rays and the blue is, in this animal, more effective than the yellow.
IX. The Behavior of the Animals which do not orient in the Tropotactic Fashion.
When animals with an innate tendency to deflect to one side of the body are subjected to the experiments, for example, under the influence of two lights of the blue and the yellow, they move either towards the yellow or the blue according to the innate direction of their deflection. These features are represented in Fig. V. As is evident from this figure, the courses of these animals are characterized by the curved paths and the degrees of the curvature increase as a rule when the larva moves into the light field of one light regardless of its nature.
X. The Problem of Color-Vision.
Phototropic reactions depend, however, not only upon the wave length of light but also upon the intensity of it. If, for instance, the light in the spectrum on either side of the maximum (green) be made sufficiently intense it becomes more effective than that at the maximum. There is therefore, so far as my experiments are concerned, no evidence indicating the presence of color-vision in the animal studied.
Acknowledgments:- The writer takes pleasure in expressing here his debt of gratitude to Prof. Dr. R. Kuroda for inviting him to publish this paper in this Journal and to Dr. T. Yamanouchi and Dr. M. Sawada for their advices in connection with the experiments.
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