A THEORY OF VISUAL INTENSITY DISCRIMINATION

1. A theory of visual intensity discrimination is proposed in terms of the photochemical events which take place at the moment when a photosensory system already adapted to the intensity I is exposed to the just perceptibly higher intensity I+ΔI. Unlike previous formulations this theory predicts that the fraction ΔI/I, after rapidly decreasing as I increases, does not increase again at high intensities, but reaches a constant value which is maintained even at the highest intensities. 2. The theory describes quantitatively the intensity discrimination data of Drosophila, of the bee, and of Mya. 3. With some carefully considered exceptions the intensity discrimination data of the human eye fall into two classes: those with small test areas or with red light, which form a single continuous curve describing the function of the retinal cones alone, and those with larger areas, and with white, orange, and yellow light, which form a double curve showing a clear inflection point, and represent the separate function of the rods at intensities below the inflection point and of the cones at intensities above it. 4. The theory describes all these data quantitatively by treating the rods and cones as two independently functioning photosensory systems in accordance with the well established duplicity idea. 5. In terms of the theory the data of intensity discrimination give critical information about the order of both the photochemical and dark reactions in each photosensory system. The reactions turn out to be variously monomolecular and bimolecular for the different animals.     

THE RELATION BETWEEN VISUAL ACUITY AND ILLUMINATION

1. Visual acuity varies in a definite manner with the illumination. At low intensities visual acuity increases slowly in proportion to log I; at higher intensities it increases nearly ten times more rapidly in relation to log I; at the highest illuminations it remains constant regardless of the changes in log I. 2. These variations in visual acuity measure the variations in the resolving power of the retina. The retina is a surface composed of discrete rods and cones. Therefore its resolving power depends on the number of elements present in a unit area. The changes in visual acuity then presuppose that the number of elements in the retina is variable. This cannot be true anatomically; therefore it must be assumed functionally. 3. To explain on such a basis the variations of visual acuity, it is postulated that the thresholds of the cones and of the rods are distributed in relation to the illumination in a statistical manner similar to that of other populations. In addition the rods as a whole have thresholds lower than the cones. Then at low intensities the increase in visual acuity depends on the augmentation of the functional rod population which accompanies intensity increase; and at higher intensities the increase in visual acuity depends on the augmentation of the functional cone population. The number of cones per unit foveal area is much greater than the number of rods per unit peripheral area, which accounts for the relative rates of increase of rod and cone visual acuity with intensity. At the highest illuminations all the cones are functional and no increase in visual acuity is possible. 4. If this division into rod visual acuity and cone visual acuity is correct, a completely color-blind person should have only rod visual acuity. It is shown by a study of the data of two such individuals that this is true. 5. The rod and cone threshold distribution has been presented as a purely statistical assumption. It can be shown, however, that it is really a necessary consequence of a photochemical system which has already been used to describe other properties of vision. This system consists of a photosensitive material in reversible relation with its precursors which are its products of decomposition as well. 6. On the basis of these and other data it is shown that a minimal retinal area in the fovea, which can mediate all the steps in such functions as visual acuity, intensity discrimination, and color vision, contains about 540 cones. Certain suggestions with regard to a quantitative mechanism for color vision are then correlated with these findings, and are shown to be in harmony with accurately known phenomena in related fields of physiology.     

INTENSITY DISCRIMINATION IN THE HUMAN EYE : II. THE RELATION BETWEENΔI/IAND INTENSITY FOR DIFFERENT PARTS OF THE SPECTRUM

1. A new apparatus is described for measuring visual intensity discrimination over a large range of intensities, with white light and with selected portions of the spectrum. With it measurements were made of the intensity ΔI which is just perceptible when it is added for a short time to a portion of a field of intensity I to which the eye has been adapted. 2. For white and for all colors the fraction ΔI/I decreases as I increases and reaches an asymptotic minimum value at high values of I. In addition, with white light the relation between ΔI/I and I shows two sections, one at low intensities and the other at high intensities, the two being separated by an abrupt transition. These findings are contrary to the generally accepted measurements of Koenig and Brodhun; however, they confirm the recent work of Steinhardt, as well as the older work of Blanchard and of Aubert. The abrupt transition is in keeping with the Duplicity theory which attributes the two sections to the functions of the rods and cones respectively. 3. Measurements with five parts of the spectrum amplify these relationships in terms of the different spectral sensibilities of the rods and cones. With extreme red light the relation of ΔI/I to I shows only a high intensity section corresponding to cone function, while with other colors the low intensity rod section appears and increases in extent as the light used moves toward the violet end of the spectrum. 4. Like most of the previously published data from various sources, the present numerical data are all described with precision by the theory which supposes that intensity discrimination is determined by the initial photochemical and chemical events in the rods and cones.     

THE RELATION BETWEEN FOVEAL VISUAL ACUITY AND ILLUMINATION UNDER REDUCED OXYGEN TENSION

1. The foveal visual acuity of eleven subjects was studied in relation to illumination under normal atmospheric conditions and at simulated altitudes of 10,000 feet (14.3 per cent O₂) and 18,000 feet (10.3 per cent O₂). A mask was used to administer the desired mixtures of oxygen and nitrogen. At the end of each experiment, measurements were made while inhaling 100 per cent oxygen from a cylinder. A red filter (No. 70 Wratten) was used so as to study only the behavior of the cones of the retina. 2. The logarithm of illumination was plotted horizontally (abscissa) and the logarithm of visual acuity vertically (ordinate). The reduced oxygen tensions resulted in a shift of the curve to the right, along the intensity axis, the extent of the change being 0.24 of a log unit at 14.3 per cent O₂ and 0.47 of a log unit at 10.3 per cent O₂. These effects were completely counteracted within a few minutes by inhaling oxygen. 3. As a consequence of the shape of the curve, such a shift to the right resulted in a relatively large decrease of visual acuity at low illuminations. At increasing light intensities anoxia produced less and less change, until at very high illuminations the decrease was negligible. Thus with 10.34 per cent O₂ the visual acuity at 0.144 photons decreased an average of 0.344 of a log unit, to 45 per cent of its normal value. At 1320 photons, however, it decreased only 0.026 of a log unit, to 94 per cent of its normal value for that intensity.     

INTENSITY DISCRIMINATION IN THE HUMAN EYE : I. THE RELATION OFΔI/ITO INTENSITY

New measurements of the brightness difference sensibility of the eye corroborate the data of previous workers which show that ΔI/I decreases as I increases. Contrary to previous report, ΔI/I does not normally increase again at high intensities, but instead decreases steadily, approaching a finite limiting value, which depends on the area of the test-field and on the brightness of the surrounding field. On a logarithmic plot, the data of ΔI/I against I for test-fields below 2° are continuous, whereas those for test-fields above 2° show a sharp discontinuity in the region of intensity in which ΔI/I decreases rapidly. This discontinuity is shown to divide the data into predominantly rod function at low intensities, and predominantly cone function at high intensities. Fields below 2° give higher values of ΔI/I at all intensities, when compared with larger fields. Fields greater than one or two degrees differ from one another principally on the low intensity side of the break. Changes in area above this limit are therefore mainly effective by changing the number of rods concerned. This is confirmed by experiments controlling the relative numbers of rods and cones with lights of different wavelength and with different retinal locations. At high intensities ΔI/I is extremely sensitive to changes in brightness of surrounding visual fields, except for large test-fields which effectually furnish their own surrounds. This sensitivity is especially marked for fields of less than half a degree in diameter. Although the effect is most conspicuous for high intensities, the surround brightness seems to affect the relation between variables as a whole, except in very small fields where absence of a surround of adequate brightness results in the distortion of the theoretical relation otherwise found. The theoretical relationship for intensity discrimination derived by Hecht is shown to fit practically all of the data. Changes in experimental variables such as retinal image area, wavelength, fixation, and criterion may be described as affecting the numerical quantities of this relationship.     

The contribution of single and double cones to spectral sensitivity in budgerigars during changing light conditions

Bird colour vision is mediated by single cones, while double cones and rods mediate luminance vision in bright and dim light, respectively. In daylight conditions, birds use colour vision to discriminate large objects such as fruit and plumage patches, and luminance vision to detect fine spatial detail and motion. However, decreasing light intensity favours achromatic mechanisms and eventually, in dim light, luminance vision outperforms colour vision in all visual tasks. We have used behavioural tests in budgerigars (Melopsittacus undulatus) to investigate how single cones, double cones and rods contribute to spectral sensitivity for large (3.4°) static monochromatic stimuli at light intensities ranging from 0.08 to 63.5 cd/m². We found no influences of rods at any intensity level. Single cones dominate the spectral sensitivity function at intensities above 1.1 cd/m², as predicted by a receptor noise-limited colour discrimination model. Below 1.1 cd/m², spectral sensitivity is lower than expected at all wavelengths except 575 nm, which corresponds to double cone function. We suggest that luminance vision mediated by double cones restores visual sensitivity when single cone sensitivity quickly decreases at light intensities close to the absolute threshold of colour vision.     

THE VISUAL INTENSITY DISCRIMINATION OF THE HONEY BEE

1. Bees respond by a characteristic reflex to a movement in their visual field. By confining the field to a series of parallel stripes of different brightness it is possible to determine at any brightness of one of the two stripe systems the brightness of the second at which the bee will first respond to a displacement of the field. Thus intensity discrimination can be determined. 2. The discriminating power of the bee''s eye varies with illumination in much the same way that it does for the human eye. The discrimination is poor at low illumination; as the intensity of illumination increases the discrimination increases and seems to reach a constant level at high illuminations. 3. The probable error of See PDF for Equation decreases with increasing I exactly in the same way as does See PDF for Equation itself. The logarithm of the probable error of ΔI is a rectilinear function of log I for all but the very lowest intensities. Such relationships show that the measurements exhibit an internal self-consistency which is beyond accident. 4. A comparison of the efficiency of the bee''s eye with that of the human eye shows that the range over which the human eye can perceive and discriminate different brightnesses is very much greater than for the bee''s eye. When the discrimination power of the human eye has reached almost a constant maximal level the bee''s discrimination is still very poor, and at an illumination where as well the discrimination power of the human eye and the bee''s eye are at their best, the intensity discrimination of the bee is twenty times worse than in the human eye.     

THE VISIBILITY OF SINGLE LINES AT VARIOUS ILLUMINATIONS AND THE RETINAL BASIS OF VISUAL RESOLUTION

The visual resolution of a single opaque line against an evenly illuminated background has been studied over a large range of background brightness. It was found that the visual angle occupied by the thickness of the line when it is just resolved varies from about 10 minutes at the lowest illuminations to 0.5 second at the highest illuminations, a range of 1200 to 1. The relation between background brightness and just resolvable visual angle shows two sections similar to those found in other visual functions; the data at low light intensities represent rod vision while those at the higher intensities represent cone vision. With violet light instead of white the two sections become even more clearly defined and separated. The retinal image produced by the finest perceptible line at the highest brightness is not a sharp narrow shadow, but a thin broad shadow whose density distribution is described in terms of diffraction optics. The line of foveal cones occupying the center of this shadow suffers a decrease in the light intensity by very nearly 1 per cent in comparison either with the general retinal illumination or with that on the row of cones to either side of the central row. Since this percentage difference is near the limit of intensity discrimination by the retina, its retinal recognition is probably the limiting factor in the visual resolution of the line. The resolution of a line at any light intensity may also be limited by the just recognizable intensity difference, because this percentage difference varies with the prevailing light intensity. As evidence for this it is found that the just resolvable visual angle varies with the light intensity in the same way that the power of intensity discrimination of the eye varies with light intensity. It is possible that visual resolution of test objects like hooks and broken circles is determined by the recognition of intensity differences in their diffracted images, since the way in which their resolution varies with the light intensity is similar to the relation between intensity discrimination and light intensity.     

THE FLICKER RESPONSE CONTOUR FOR THE GECKO (ROD RETINA)

The flicker response contour for the gecko Sphaerodactylus (retina with only rods) agrees in all essential respects (intensity range, shape) with that for the turtle Pseudemys (cone retina), as determined under equivalent conditions with the same apparatus. With experimentally determined correction for the expansion of the iris at the very lowest intensities, the F - log I contour for the gecko is a simple probability integral. Its maximum F is lower than that for other animals; this means simply a smaller number of available sensory elements. The quantitative parallelism in the magnitudes of the intensities at the inflection of F - log I and the shape constants for rod and cone animals show that assumptions from comparative histological evidence concerning the properties of rods and cones in relation to visual performance may be quite misleading.     

INTERMITTENT STIMULATION BY LIGHT : III. THE RELATION BETWEEN INTENSITY AND CRITICAL FUSION FREQUENCY FOR DIFFERENT RETINAL LOCATIONS

When measurements of the critical fusion frequency for white light over a large range of intensities are made with the rod-free area of the fovea, the relation between critical frequency and log I is given by a single sigmoid curve, the middle portion of which approximates a straight line whose slope is 11.0. This single relation must be a function of the foveal cones. When the measurements are made with a retinal area placed 5° from the fovea, and therefore containing both rods and cones, the relation between critical frequency and log I shows two clearly separated sections. At the lower intensities the relation is sigmoid and reaches an upper level at about 10 cycles per second, which is maintained for 1.25 log units, and is followed by another sigmoid relationship at the higher intensities similar to the one given by the rod-free area alone. These two parts of the data are obviously separate functions of the rods at low intensities and of the cones at high intensities. This is further borne out by similar measurements made with retinal areas 15° and 20° from the fovea where the ratio of rods to cones is anatomically greater than at 5°. The two sections of the data come out farther apart on the intensity scale, the rod portion being at lower intensities and the cone portion at higher intensities than at 5°. The general form of the relation between critical frequency and intensity is therefore determined by the relative predominance of the cones and the rods in the retinal area used for the measurements.     

THE VISUAL DISCRIMINATION OF INTENSITY AND THE WEBER-FECHNER LAW

1. A study of the historical development of the Weber-Fechner law shows that it fails to describe intensity perception; first, because it is based on observations which do not record intensity discrimination accurately, and second, because it omits the essentially discontinuous nature of the recognition of intensity differences. 2. There is presented a series of data, assembled from various sources, which proves that in the visual discrimination of intensity the threshold difference ΔI bears no constant relation to the intensity I. The evidence shows unequivocally that as the intensity rises, the ratio See PDF for Equation first decreases and then increases. 3. The data are then subjected to analysis in terms of a photochemical system already proposed for the visual activity of the rods and cones. It is found that for the retinal elements to discriminate between one intensity and the next perceptible one, the transition from one to the other must involve the decomposition of a constant amount of photosensitive material. 4. The magnitude of this unitary increment in the quantity of photochemical action is greater for the rods than for the cones. Therefore, below a certain critical illumination—the cone threshold—intensity discrimination is controlled by the rods alone, but above this point it is determined by the cones alone. 5. The unitary increments in retinal photochemical action may be interpreted as being recorded by each rod and cone; or as conditioning the variability of the retinal cells so that each increment involves a constant increase in the number of active elements; or as a combination of the two interpretations. 6. Comparison with critical data of such diverse nature as dark adaptation, absolute thresholds, and visual acuity shows that the analysis is consistent with well established facts of vision.     

CRITICAL ILLUMINATION AND FLICKER FREQUENCY,AS A FUNCTION OF FLASH DURATION: FOR THE SUNFISH

From the relations between critical illumination in a flash (I_m) and the flash frequency (F) for response of the sunfish to visual flicker when the proportion of light time to dark time (t_L/t_D) in a flicker cycle is varied at one temperature (21.5°) the following results are obtained: At values of t_L/t_D between 1/9 and 9/1 the F - log I_m curves are progressively shifted toward higher intensities and lower F_max.. F_max. is a declining rectilinear function of the percentage of the flash cycle time occupied by light. The rod and the cone portions of the flicker curve are not shifted to the same extent. The cone portion and the rod region of the curve are each well described by a probability integral. In terms of F as 100 F/F_max. the standard deviation of the underlying frequency distribution of elemental contributions, summed to produce the effect proportional to F, is independent of t_L/t_D. The magnitude of log I_m at the inflection point (r''), however, increases rectilinearly with the percentage light time in the cycle. The proportionality between I_m and σ_II1 is independent of t_L/t_D. These effects are interpreted as consequences of the fact that the number of elements of excitation available for discrimination of flicker is increased by increasing the dark interval in a flash cycle. Decreasing the dark interval has therefore the same kind of effect as reducing the visual area, and not that produced by decreasing the temperature.     

THE INFLUENCE OF LIGHT ADAPTATION ON SUBSEQUENT DARK ADAPTATION OF THE EYE

The course of dark adaptation of the human eye varies with the intensity used for the light adaptation which precedes it. Preadaptation to intensities below 200 photons is followed only by rod adaptation, while preadaptation to intensities above 4000 photons is followed first by cone adaptation and then by rod adaptation. With increasing intensities of preadaptation, cone dark adaptation remains essentially the same in form, but covers an increasing range of threshold intensities. At the highest preadaptation the range of the subsequent cone dark adaptation covers more than 3 log units. Rod dark adaptation appears in two types—a rapid and a delayed. The rapid rod dark adaptation is evident after preadaptations to low intensities corresponding to those usually associated with rod function. The delayed rod dark adaptation shows up only after preadaptation to intensities which are hundreds of times higher than those which produce the maximal function of the rods in flicker, intensity discrimination, and visual acuity. The delayed form remains essentially constant in shape following different intensities of preadaptation. However, its time of appearance increases with the preadaptation intensity; after the highest preadaptation, it appears only after 12 or 13 minutes in the dark. These two modes of rod dark adaptation are probably the expression of two methods of formation of visual purple in the rods after its bleaching by the preadaptation lights.     

VISUAL ACUITY AND ILLUMINATION IN DIFFERENT SPECTRAL REGIONS

The relation between visual acuity and illumination was measured in red and blue light, using a broken circle or C and a grating as test objects. The red light data fall on single continuous curves representing pure cone vision. The blue light data fall on two distinct curves with a transition at about 0.03 photons. Values below this intensity represent pure rod vision. Those immediately above represent the cooperative activity of rods and cones, and yield higher visual acuities than either. Pure cone vision in this intensity region is given by central fixation (C test object). All the rest of the values above this transition region represent pure cone vision. In blue light the rod data with the C lie about 1.5 log units lower on the intensity axis (cone scale) than they do in white light, while with the grating they lie about 1.0 log unit lower than in white light. Both the pure rod and cone data with the C test object are precisely described by one form of the stationary state equation. With the grating test object and a non-limiting pupil, the pure rod and cone data are described by another form of the same equation in which the curve is half as steep. The introduction of a small pupil, which limits maximum visual acuity, makes the relation between visual acuity and illumination appear steeper. Determinations of maximum visual acuities under a variety of conditions show that for the grating the pupil has to be larger, the longer the wavelength of the light, in order for the pupil not to be the limiting factor. Similar measurements with the C show that when intensity discrimination at the retina is experimentally made the limiting factor in resolution, visual acuity is improved by conditions designed to increase image contrast. However, intensity discrimination cannot be the limiting factor for the ordinary test object resolution because the conditions designed to improve image contrast do not improve maximum visual acuity, while those which reduce image contrast do not produce proportional reductions of visual acuity.     

CRITICAL ILLUMINATION AND FLICKER FREQUENCY IN RELATED FISHES

Flicker response curves have been obtained at 21.5°C. for three genera of fresh water teleosts: Enneacanthus (sunfish), Xiphophorus (swordtail), Platypoecilius (Platy), by the determination of mean critical intensities for response at fixed flicker frequencies, and for a certain homogeneous group of backcross hybrids of swordtail x Platy (Black Helleri). The curves exhibit marked differences in form and proportions. The same type of analysis is applicable to each, however. A low intensity rod-governed section has added to it a more extensive cone portion. Each part is accurately described by the equation F = F_max./(1 + e ^{-p log-p logI/I_i}), where F = flicker frequency, I = associated mean critical intensity, and I_i is the intensity at the inflection point of the sigmoid curve relating F to log I. There is no correlation between quantitative features of the rod and cone portions. Threshold intensities, p, I_i, and F_max. are separately and independently determined. The hybrid Black Helleri show quantitative agreement with the Xiphophorus parental stock in the values of p for rods and cones, and in the cone F_max.; the rod F_max. is very similar to that for the Platy stock; the general level of effective intensities is rather like that of the Platy form. This provides, among other things, a new kind of support for the duplicity doctrine. Various races of Platypoecilius maculatus, and P. variatus, give closely agreeing values of I_m at different flicker frequencies; and two species of sunfish also agree. The effect of cross-breeding is thus not a superficial thing. It indicates the possibility of further genetic investigation. The variability of the critical intensity for response to flicker follows the rules previously found to hold for other forms. The variation is the expression of a property of the tested organism. It is shown that, on the assumption of a frequency distribution of receptor element thresholds as a function of log I, with fluctuation in the excitabilities of the marginally excited elements, it is to be expected that the dispersion of critical flicker frequencies in repeated measurements will pass through a maximum as log I is increased, whereas the dispersion of critical intensities will be proportional to I_m; and that the proportionality factor in the case of different organisms bears no relation to the form or position of the respective curves relating mean critical intensity to flicker frequency. These deductions agree with the experimental findings.     

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

The curve connecting mean critical illumination (I_m) and flicker frequency (F) for response of the sunfish Lepomis (Enneacanthus gloriosus) to flicker is systematically displaced toward lower intensities by raising the temperature. The rod and cone portions of the curve are affected in a similar way, so that (until maximum F is approached) the shift is a nearly constant fraction of I_m for a given change of temperature. These relationships are precisely similar to those found in the larvae of the dragonfly Anax. The modifications of the variability functions are also completely analogous. The effects found are consistent with the view that response to flicker is basically a matter of discrimination between effect of flashes of light and their after effects,—a form of intensity discrimination. They are not consistent with the stationary state formulation of the shape of the flicker curve. An examination of the relationships between the cone portion and the rod portion of the curves for the sunfish suggests a basis for their separation, and provides an explanation for certain "anomalous" features of human flicker curves. It is pointed out how tests of this matter will be made.     

THE EFFECTS OF VARIATIONS IN THE CONCENTRATION OF OXYGEN AND OF GLUCOSE ON DARK ADAPTATION

In this study we have analyzed the effects of variations in the concentrations of oxygen and of blood sugar on light sensitivity; i.e. dark adaptation. The experiments were carried out in an air-conditioned light-proof chamber where the concentrations of oxygen could be changed by dilution with nitrogen or by inhaling oxygen from a cylinder. The blood sugar was lowered by the injection of insulin and raised by the ingestion of glucose. The dark adaptation curves were plotted from data secured with an apparatus built according to specifications outlined by Hecht and Shlaer. During each experiment, observations were first made in normal air with the subject under basal conditions followed by one, and in most instances two, periods under the desired experimental conditions involving either anoxia or hyper- or hypoglycemia or variations in both the oxygen tension and blood sugar at the same time. 1. Dark adaptation curves were plotted (threshold against time) in normal air and compared with those obtained while inhaling lowered concentrations of oxygen. A decrease in sensitivity was observed with lowered oxygen tensions. Both the rod and cone portions of the curves were influenced in a similar way. These effects were counteracted by inhaling oxygen, the final rod thresholds returning to about the level of the normal base line in air or even below it within 2 to 3 minutes. The impairment was greatest for those with a poorer tolerance for low O₂. Both the inter- and intra-individual variability in thresholds increased significantly at the highest altitude. 2. In a second series of tests control curves were obtained in normal air. Then while each subject remained dark adapted, the concentrations of oxygen were gradually decreased. The regeneration of visual purple was apparently complete during the 40 minutes of dark adaptation, yet in each case the thresholds continued to rise in direct proportion to the degree of anoxia. The inhalation of oxygen from a cylinder quickly counteracted the effects for the thresholds returned to the original control level within 2 to 3 minutes. 3. In experiments where the blood sugar was raised by the ingestion of glucose in normal air, no significant changes in the thresholds were observed except when the blood sugar was rapidly falling toward the end of the glucose tolerance tests. However, when glucose was ingested at the end of an experiment in low oxygen, while the subject remained dark adapted, the effects of the anoxia were largely counteracted within 6 to 8 minutes. 4. The influence of low blood sugar on light sensitivity was then studied by injecting insulin. The thresholds were raised as soon as the effects of the insulin produced a fall in the blood sugar. When the subjects inhaled oxygen the thresholds were lowered. Then when the oxygen was withdrawn so that the subject was breathing normal air, the thresholds rose again within 1 to 2 minutes. Finally, if the blood sugar was raised by ingesting glucose, the average threshold fell to the original control level or even below it. 5. The combined effects of low oxygen and low blood sugar on light sensitivity were studied in one subject (W. F.). These effects appeared to be greater than when a similar degree of anoxia or hypoglycemia was brought about separately. 6. In a series of experiments on ten subjects the dark adaptation curves were obtained both in the basal state and after a normal breakfast. In nine of the ten subjects, the food increased the sensitivity of the subjects to light. 7. The experiments reported above lend support to the hypothesis that both anoxia and hypoglycemia produce their effects on light sensitivity in essentially the same way; namely, by slowing the oxidative processes. Consequently the effects of anoxia may be ameliorated by giving glucose and the effects of hypoglycemia by inhaling oxygen. In our opinion, the changes may be attributed directly to the effects on the nervous tissue of the visual mechanism and the brain rather than on the photochemical processes of the retina.     

Bat Eyes Have Ultraviolet-Sensitive Cone Photoreceptors

Mammalian retinae have rod photoreceptors for night vision and cone photoreceptors for daylight and colour vision. For colour discrimination, most mammals possess two cone populations with two visual pigments (opsins) that have absorption maxima at short wavelengths (blue or ultraviolet light) and long wavelengths (green or red light). Microchiropteran bats, which use echolocation to navigate and forage in complete darkness, have long been considered to have pure rod retinae. Here we use opsin immunohistochemistry to show that two phyllostomid microbats, Glossophaga soricina and Carollia perspicillata, possess a significant population of cones and express two cone opsins, a shortwave-sensitive (S) opsin and a longwave-sensitive (L) opsin. A substantial population of cones expresses S opsin exclusively, whereas the other cones mostly coexpress L and S opsin. S opsin gene analysis suggests ultraviolet (UV, wavelengths <400 nm) sensitivity, and corneal electroretinogram recordings reveal an elevated sensitivity to UV light which is mediated by an S cone visual pigment. Therefore bats have retained the ancestral UV tuning of the S cone pigment. We conclude that bats have the prerequisite for daylight vision, dichromatic colour vision, and UV vision. For bats, the UV-sensitive cones may be advantageous for visual orientation at twilight, predator avoidance, and detection of UV-reflecting flowers for those that feed on nectar.     

BRIGHTNESS DISCRIMINATION AS A FUNCTION OF THE DURATION OF THE INCREMENT IN INTENSITY

1. This investigation has been concerned with an analysis of brightness discrimination as it is influenced by the duration of ΔI. The durations used extend from 0.002 second to 0.5 second. 2. ΔI/I values at constant intensity are highest for the shortest duration and decrease with an increase in duration up to the limits of a critical exposure time. At durations longer than the critical duration the ratio ΔI/I remains constant. 3. The Bunsen-Roscoe law holds for the photolysis due to ΔI. This is shown by the fact that, within the limits of a critical duration, the product of ΔI and exposure time is constant for any value of prevailing intensity, I. 4. At durations greater than the critical duration the Bunsen-Roscoe law is superseded by the relation ΔI = Constant. This change of relation is considered in the light of Hartline''s discussion (1934). 5. The critical duration is a function of intensity. As intensity increases the critical duration decreases. 6. Hecht''s theory (1935) accounts for the data of this experiment if it be assumed that brightness discrimination is determined by a constant amount of photolysis.