INTERMITTENT STIMULATION BY LIGHT : V. THE RELATION BETWEEN INTENSITY AND CRITICAL FREQUENCY FOR DIFFERENT PARTS OF THE SPECTRUM

1. An optical system is described which furnishes large flickering fields whose brightness, even when reduced with monochromatic filters, is capable of covering the complete range of the relation between critical frequency and intensity. 2. For a centrally located test field of 19° diameter, with light from different parts of the spectrum, the data divide into a low intensity section identified with rod function, and a high intensity section identified with cone function. The transition between the two sections is marked by an inflection point which is sharp, except for 450 and 490 mµ where, though clearly present, it is somewhat rounded. 3. The intensity range covered by the flicker function is smallest in the red, and increases steadily as the wave-length decreases. The increase is due entirely to the extent of the low intensity, rod section which is smallest (non-existent for S. S.) in the red and largest in the violet. The high intensity cone portion for all colors is in the same position on the intensity axis, and the only effect of decreasing wave-length is to shift the rod section to lower intensities without changing its shape. 4. The measurements are faithfully described by two similar equations, one for the rods and one for the cones, both equations being derived from the general stationary state equation already used for various visual functions.     

INTERMITTENT STIMULATION BY LIGHT : II. THE MEASUREMENT OF CRITICAL FUSION FREQUENCY FOR THE HUMAN EYE

An apparatus and a procedure are described to measure the critical frequency of flicker using different portions of the eye. The observer, looking through a pupil of fixed dimensions, views a field of 2° whose illumination is periodically interrupted and which is surrounded by a field of 10° whose illumination is continuous but otherwise identical with the interrupted field. Various parts of the apparatus are concerned with controlling and recording the retinal position of the field, its intensity, its spectral composition, and the frequency of interruption of its illumination. The procedure is so simplified and regulated that a complete set of readings over the whole intensity range of vision can be made at one sitting without fatigue or strain.     

INTENSITY AND CRITICAL FREQUENCY FOR VISUAL FLICKER

Using the rotating striped cylinder device previously employed for determination of the flicker response function with lower animals, corresponding measurements have been made with human observers. The curves based upon the relation between critical flash frequency and critical intensity for the signalling of the recognition of flicker have the properties of human flicker fusion data as obtained by other methods. They also have the quantitative properties of the flicker curves provided by the motor responses of insects and fishes to the seen movement of flashes. This applies to the variation found in repeated measurements as well as to the nature of the analytical function describing the connection between flash frequency and intensity. The data for human visual flicker and those for the responses of lower animals are therefore essentially homologous.     

ON THE VARIABILITY OF CRITICAL ILLUMINATION FOR FLICKER FUSION AND INTENSITY DISCRIMINATION

From the data of experiments with bees in which threshold response is employed as a means of recognizing visual discrimination between stripes of equal width alternately illuminated by intensities I ₁ and I ₂, it is shown that the detectable increment of intensity ΔI, where ΔI = I ₂ - I ₁, is directly proportional to σ_I2 (I ₁ being fixed). From tests of visual acuity, where I ₁ = 0 and the width of the stripes is varied, σ_I2 = kI ₂ + const.; here I ₂ = ΔI, and ΔI/I ₂ = 1. When the visual excitability of the bee is changed by dark adaptation, λI ≡ kΔI (= k'' σ_ΔI) = k'''' I + const. For the measurements of critical illumination at threshold response to flicker, σ_I2 (= σ_ΔI) = k I ₂ = k'' ΔI + const. The data for critical illumination producing threshold response to flicker in the sun-fish Lepomis show for the rods σ_I2 = K I ₂ for the cones σ_I2 = K''(I ₂ + const.). The data thus indicate that in all these experiments essentially the same visual function is being examined, and that the recognition of the production of a difference in effect by alternately illuminated stripes takes place in such a way that d (ΔI)/d (σ_I2) = const., and that ΔI is directly proportional to I (or "I ₂," depending on the nature of the experiment). It is pointed out that the curve for each of the cases considered can be gotten equally well if mean I or σ_I is plotted as a function of the independent variable involved in the experiment. Certain consequences of these and related facts are important for the treatment of the general problem of intensity discrimination.     

PHOTOCHEMISTRY OF VISUAL PURPLE : III. THE RELATION BETWEEN THE INTENSITY OF LIGHT AND THE RATE OF BLEACHING OF VISUAL PURPLE.

It is shown that the velocity of bleaching of visual purple by light, under comparable conditions of concentration, volume, and surface exposed, is directly proportional to the intensity.     

ON CRITICAL FREQUENCY AND CRITICAL ILLUMINATION FOR RESPONSE TO FLICKERED LIGHT

The curve of mean critical flicker frequency as a function of illumination has been determined for the reaction of the sunfish Lepomis to flicker. It exhibits expected quantitative disagreements with the curve of mean critical illumination as a function of flicker frequency in the same organism. The form of the dependence of the variation of critical frequency of flicker upon illumination can be predicted from a knowledge of the way in which variation of critical illumination depends upon flicker frequency. It is pointed out that these findings have an important bearing upon the interpretation of the data of intensity discrimination.     

CRITICAL ILLUMINATION AND CRITICAL FREQUENCY FOR RESPONSE TO FLICKERED LIGHT,IN DRAGONFLY LARVAE

Curves relating flicker frequency (F) to mean critical illumination (I_m) for threshold response to flickered light, with equal durations of light and no light intervals, and relating illumination (I) to mean critical flicker frequency (F_m) for the same response, have been obtained from homogeneous data based upon the reactions of dragonfly larvae (Anax junius). These curves exhibit the properties already described in the case of the fish Lepomis. The curve for F_m lies above the curve of I_m by an amount which, as a function of I, can be predicted from a knowledge either of the variation of I_m or of F_m. The law of the observable connection between F and I is properly expressed as a band, not as a simple curve. The variation of I_m (and of F_m) is not due to "experimental error," but is an expression of the variable character of the organism''s capacity to exhibit the reaction which is the basis of the measurements. As in other series of measurements, P.E. _I is a rectilinear function of I_m; P.E. _F passes through a maximum as F (or I) increases. The form of P.E. _F as a function of I can be predicted from the measurements of P.E. _I. It is pointed out that the equations which have been proposed for the interpretation of curves of critical flicker frequency as a function of intensity, based upon the balance of light adaptation and dark adaptation, have in fact the character of "population curves;" and that their contained constants do not have the properties requisite for the consistent application of the view that the shape of the F - I curve is governed by the steady state condition of adaptation. These curves can, however, be understood as resulting from the achievement of a certain level of difference between the average effect of a light flash and its average after effect during the dark interval.     

TEMPERATURE AND CRITICAL ILLUMINATION FOR REACTION TO FLICKERING LIGHT : III. SUNFISH

For the sunfish Enneacanthus the mean value of the critical illumination for response to visual flicker at constant flash frequency (with light time = dark time) is related to temperature by the Arrhenius equation. The temperature characteristic for 1/I_m is different above and below 20°C. In each range (12° to 20°; 20° to 30°) the temperature characteristic is the same for rod and cone segments of the duplex flicker response contour: 8,200 and 14,400. This makes it difficult, if not impossible, to consider that the two groups of elements are organized in a significantly different way chemically. For the presumptively rod-connected elements implicated in response to flicker, the curve is markedly discontinuous, so that the high and low temperature parts are dislocated; whereas for the cones they are not. This is entirely consistent with other (e.g., genetic) evidence pointing to their separate physical substrata. The uncommon exhibition of a higher µ over a higher range of temperature, previously found, however, in a few cases, together with the different relations of rod and cone effects to the critical temperature, explain aspects of these data which in earlier incomplete measurements were found to be puzzling.     

羊草呼吸作用与温度、光照和土壤水分的关系

本文报道了两种土壤水分条件下羊草明呼吸速率与光照和温度的关系,以及暗呼吸速率与温度的关系。结果表明,羊草的明呼吸速率与光强呈非线性函数关系。在低光强下,明呼吸速率随光强升高而有较快的增加;随着光强的增高,其增加速度减慢。在温度低于羊草光合的高温补偿点的条件下,明呼吸速率在一定温度范围内随温度升高而增大;当温度达到一定限度时,有一个下降阶段,而后又回升。羊草的暗呼吸速率随温度增加而升高,且在一定限度内,其升高速度随温度增高而加快。当土壤干旱时,明呼吸速率显著降低,而暗呼吸速率仅略有减小。     

瓦氏黄颡鱼和鲢对光照强度和颜色的选择

以野生瓦氏黄颡鱼和人工养殖鲢为研究对象,设置4种光照强度梯度010、1030、100200、700800 lx及4种光照颜色红、白、蓝、绿,采用单因子实验法,研究了瓦氏黄颡鱼和鲢对光的趋避行为。结果表明:在光照强度选择的实验中,瓦氏黄颡鱼在010、1030、100200和700800 lx四个光照强度中出现的次数百分比分别是:78.33%21.51%、10.00%14.86%、7.33%8.27%和4.33%7.74%。瓦氏黄颡鱼在光照强度为010 lx的区域中出现的次数明显多于其他三种光照强度,且差异显著（P0.05）。而鲢在四种光照强度中出现的次数之间不存在显著性差异（P0.05）。在光照颜色选择实验中,瓦氏黄颡鱼在四种光照颜色中出现的次数百分比不存在显著性差异（P0.05）。鲢在红、白、蓝和绿四种光照颜色中出现的次数百分比分别是:12.67%13.63%、30.33%18.47%、35.67%24.73%、21.33%15.02%。鲢在白光、蓝光区域出现的次数显著高于其他两种区域,且差异显著（P0.05）。实验通过研究光照强度和光照颜色对瓦氏黄颡鱼和鲢行为的影响,为鱼类趋光性研究提供一定的参考。       

CRITICAL FREQUENCY OF FLICKER AS A FUNCTION OF INTENSITY OF ILLUMINATION FOR THE EYE OF THE BEE

The bee''s characteristic response to a movement of its visual field is used for the study of the relation between critical frequency of flicker and illumination. The critical flicker frequency varies with illumination in such a way that with increasing flicker frequency the intensity of illumination must be increased to produce a threshold response in the bee. The illuminations required to give a response in a bee at different flicker frequencies closely correspond to the intensities for threshold response in visual acuity tests. This is due to the different thresholds of excitability of the elements of the ommatidial mosaic. An analysis of the variation of the values for threshold intensities at the several flicker frequencies shows that the variation depends upon flicker frequency and upon the number of elements functioning at different intensities.     

瓦氏黄颡鱼和鲢对光照强度和颜色的选择简

以野生瓦氏黄颡鱼和人工养殖鲢为研究对象,设置4种光照强度梯度0—10、10—30、100—200、700—800 lx及4种光照颜色红、白、蓝、绿,采用单因子实验法,研究了瓦氏黄颡鱼和鲢对光的趋避行为。结果表明:在光照强度选择的实验中,瓦氏黄颡鱼在0—10、10—30、100—200和700—800 lx四个光照强度中出现的次数百分比分别是:78.33%±21.51%、10.00%±14.86%、7.33%±8.27%和4.33%±7.74%。瓦氏黄颡鱼在光照强度为0—10 lx的区域中出现的次数明显多于其他三种光照强度,且差异显著(P0.05)。而鲢在四种光照强度中出现的次数之间不存在显著性差异(P0.05)。在光照颜色选择实验中,瓦氏黄颡鱼在四种光照颜色中出现的次数百分比不存在显著性差异(P0.05)。鲢在红、白、蓝和绿四种光照颜色中出现的次数百分比分别是:12.67%±13.63%、30.33%±18.47%、35.67%±24.73%、21.33%±15.02%。鲢在白光、蓝光区域出现的次数显著高于其他两种区域,且差异显著(P0.05)。实验通过研究光照强度和光照颜色对瓦氏黄颡鱼和鲢行为的影响,为鱼类趋光性研究提供一定的参考。     

THRESHOLD INTENSITY OF ILLUMINATION AND FLICKER FREQUENCY FOR THE EYE OF THE SUN-FISH

The sun-fish Lepomis responds to a moving system of stripes by a motion of its body. By changing the velocity of motion of the stripe system different flicker frequencies can be produced and thus the relation of flicker frequency to critical intensity of illumination can be studied. Threshold illumination varies with flicker frequency in such a way that with increasing flicker frequency the intensity of illumination must be increased to produce a threshold response in the fish. The curve of critical illumination as a function of frequency is made up of two distinct parts. For an intensity range below 0.04 millilambert and flicker frequencies below 10 per second, the rods are in function. For higher intensities and flicker frequencies above 10, the cones come into play. The maximum frequency of flicker which can be perceived by the fish''s eye is slightly above 50 per second. The flicker curve for the eye of Lepomis can easily be compared with that for the human eye. The extent of the curve for the fish is greater at low illuminations, the fish being capable of distinguishing flicker at illuminations lower than can the human eye. The transition of rod vision to cone vision occurs for the fish and for the human eye at the same intensity and flicker frequency. The maximum frequency of flicker which can be perceived is for both about the same.