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Biomedical subjects

M Alpern

Publications and source records attributed to M Alpern.

At least 37 records · Page 2Linked to original sources

Classical tritanopia.

1. A subject who has suffered from central serous chorio-retinopathy in his left eye noticed differences in the colour of a given light as perceived by each eye alone. Standard screening tests (colour order and colour matching) indicated a tritan defect in the left eye; the right eye was normal on these tests.2. The subject was dichromatic in his left eye, trichromatic in his right. The left-eye distimulus colour-matching functions, spectral luminosity, and wave-length discrimination functions were indistinguishable from corresponding data for congenital tritanopia. Comparable right-eye data were normal.3. Spectral dichromatic colour matches were invariant under changes of intensity and under addition of a common light to both halves of the field. (Grassmann's laws of linearity are satisfied.)4. Increment threshold versus intensity (t.v.i.) curves for a blue (481.9 nm) test on a yellow background yielded the normal three branches (for Pi(4)(mu), Pi(1)(mu) and Pi(3)(mu) respectively) in the trichromatic eye. In the dichromatic eye a single mechanism was found. It had the field sensitivity of Pi(4)(mu) whether measured with the blue, or with a violet (429.5 nm) test. No trace of Pi(3)(mu) or Pi(1)(mu) was ever discovered in the tritanopic eye. Both are normal in the trichromatic eye.5. The field sensitivities of Pi(4), Pi(5) and Pi(3) of the normal eye are well fitted by linear combinations of the spectral colour-matching functions of the trichromatic eye. Pi(4) and Pi(5) of the dichromatic eye are well fitted by linear combinations of the tritanopic matching functions.6. Colour matches made by the trichromatic eye do not match when viewed by the tritanopic eye, almost certainly because the ocular media of the two eyes have wave-length-dependent differences in absorption. For the largest difference (430 nm) the trichromatic eye transmits about 2.2 times more light than its fellow. When allowance is made for these differences, the field sensitivities of Pi(4) and Pi(5) of the two eyes do not differ. The field sensitivities of Pi(4) and Pi(5) of the normal eye, on the other hand, differ significantly from those of the average spectra obtained on four normal trichromats by Stiles, in a way that cannot be attributed to differences in transmittance of ocular media.7. It is concluded that classical (or acquired) tritanopia is not distinguishable in its manifestations from congenital tritanopia; furthermore, tritanopia can be regarded as a reduced form of normal trichromacy, once allowances are made for absorption of the ocular media and for variations among normal trichromats.8. Despite extensive search no evidence could be uncovered which might exclude the hypothesis that the colour vision in tritanopia depends exclusively upon absorption in only two foveal cone pigments, one long-wave-absorbing and one medium-wave-absorbing.

Adult↗

Perception of colour in unilateral tritanopia.

The unilateral tritanope described in the previous paper (Alpern, Kitahara & Krantz, 1983) was able to match every narrow-band light presented to his tritanopic eye with lights from a tristimulus colorimeter viewed in the adjacent field by the normal eye. In two regions of the spectrum (called isochromes) physically identical lights appeared identical to the observer's two eyes. One isochrome was close to 'blue' for the normal eye, the other was in the long-wave spectral region seen by the normal eye predominantly as 'red'. Between these isochromes the normal eye required less than spectral purity to match, dropping to near zero purity at 560-570 nm. A mixture of the two isochromes that appeared purple to the normal eye appeared neutral to the tritanopic eye. Hence dichoptic matches grossly violate Grassmann's additivity law. For the normal eye colour naming conformed to typical normal results. For the tritanopic eye the results were coherent with those found by dichoptic matching: the spectrum was divided into two regions by the achromatic neutral band. To the short-wave side, only the colour names 'blue' and 'white' were ever used. To the long-wave side the predominant colour names were 'red' and 'white' with some 'yellow'. Spectral lights appeared neither 'red-blue' nor greenish. Surrounding the test with an annulus either 430 nm, 650 nm, or a mixture of these, fails to induce any greenish appearance, although the achromatic band shifted in the expected directions. It is concluded that there must be exactly three functionally independent, essentially non-linear central codes for colour perception, and that these codes are different from those suggested in existing theories of colour perception.

Adaptation, Physiological↗

The directional sensitivities of the Stiles' colour mechanisms.

Field sensitivities of the three IIj (j = 3, 4, 5) mechanisms of Stiles were measured for monochromatic backgrounds of different wave numbers (mu)-1 traversing the eye through different points (r) displaced along a horizontal chord through the centre of the entrance pupil. Each mechanism shows an insensitivity to the direction of retinal incidence of short-wave backgrounds not previously described. The spectral densities of the centre-most part of the lens and of the macular pigment were measured on this eye. With reasonable assumptions the former allowed for correction at the receptor level of the directional sensitivity; together with the latter it allowed correction for the spectral sensitivity as well. No correction for the attenuation of the high spatial frequencies of the background as it traversed the pupil at different r was needed. The anomalies of section 2 (above) disappear after correction for losses in the eye media. After these corrections, for every mu and r, the results are well described by the parabola 'tentatively' suggested by Stiles (1939) for each mechanism, allowing only a small amount of variance attributable to experimental imprecision alone. Each mechanism is most sensitive to backgrounds going through essentially the same point of the pupil, independent of background. This result is inconsistent with a qualitative explanation of the 'hue shift' suggested by Safir, Hyams & Philpot (1971). The field sensitivity spectra for backgrounds traversing the pupil at this most effective point and at the 3.5 mm margin, are the data needed to predict this observer's brightness and colour matches of monochromatic lights passing through the entrance pupil at these two points according to a unified theory of the two Stiles-Crawford effects. In the following paper these predictions are quantified and confronted with results of the matching experiments (Alpern, Kitahara & Tamaki, 1982).

Adult↗

The dependence of the colour and brightness of a monochromatic light upon its angle of incidence on the retina.

The changes in brightness and colour of a monochromatic test light as its angle of incidence on the retina was changed from normal (pupil centre traverse) to oblique (3.5 mm temporal pupil traverse), was measured by matching it with three normally incident primaries. Results on two normal trichromats were generally in accord with published data on the Stiles-Crawford intensity and colour effects. One observer was also the subject of the preceding paper (Alpern & Kitahara, 1983) in which the field sensitivities of his foveal IIj(mu) (j = 3, 4, 5) mechanisms for normally, and obliquely, incident backgrounds were reported. For normal incidence, the colour matching functions are in rough accord with expectation if the action spectra of the three cone mechanisms, which provide the photoreceptor basis for his trichromacy, were the same IIj mechanisms for normal incidence. A unified theory is developed for both Stiles-Crawford intensity and colour effects, assuming that the same visual pigments in the same set of univariantly signalling cones absorbs both the normal incident primaries and the obliquely incident test. Given no free parameters for curve fitting, the Stiles-Crawford intensity effect data are in reasonable agreement with the theory if the photoreceptor basis of these matches were the normally and obliquely incident IIj(mu) mechanisms. The Stiles-Crawford colour effect data contradict the expectations of the unified theory applied with these same IIj(mu) mechanisms. Either II3(mu) is not a valid operational definition of the action spectrum of his short-wave sensitive photoreceptors or at least one assumption of the unified theory is false.

Color Perception↗

The saturation of monochromatic lights obliquely incident on the retina.

Foveal dark-adaptation undertaken to test the hypothesis that the excitation of rods causes the desaturation of 'yellow' lights in a 1 degree field traversing the margin of the pupil, fails to exclude that possibility. The desaturation is largest for a 1 degree outside diameter annular test, is still measurable with a 0.5 degree circular disk, but disappears for a 0.29 degree disk. The supersaturation of obliquely incident 501.2 nm test light follows the opposite pattern; it disappears with an annulus and is largest for a 0.29 degree circular field. It is unlikely that rods replace short-wave sensitive cones in the trichromatic match of an obliquely incident test with normally incident primaries. If rods as well as all three cones species are involved, the matches might not be trichromatic in the strong sense. Grassmann's law of scalar multiplication was tested and shown not to hold for the match of an obliquely incident test with normally incident primaries, though it remains valid whenever, both primaries and test strike the retina at the same angle of incidence (independent of that angle). The result in section 3 (above) cannot be due to rod intrusion. It persists (and becomes more conspicuous) on backgrounds (4.0 log scotopic td) which saturate rods. Moreover obliquely incident 'yellow' lights remain desaturated in intervals in the dark after a full bleach, whilst the test field is below rod threshold. The amount of desaturation does not differ appreciably from that normally found. The assumption of the unified theory of Alpern, Kitahara & Tamaki (1983) that the outer segments of only a single set of three cone species (with acceptance angles wide enough to include the entire exit pupil) contain the visual pigments absorbing both the normally incident primaries and the obliquely incident test is disproved by these results. Failure of Grassmann's law is most conspicuous under the conditions for which the changes in saturation upon changing from normal to oblique incidence are greatest and least when the saturation changes are the smallest. Either all unified theories of the Stiles-Crawford effects are wrong or all the effects of oblique incidence operate at a stage in the visual process at which the effects of radiation of different wave-lengths are no longer compounded by the simple linear laws.

Color Perception↗

The directional sensitivity of retinal rods.

Rod field sensitivity, 10-S(r) (i.e. the reciprocal of the radiance of a background required for 10-fold elevation of rod threshold) was measured for monochromatic backgrounds traversing the pupil at various points (r) on three subjects. The wave-length dependency of the directional sensitivities of the three foveal cone mechanisms of the principal subject have been reported previously (Alpern & Kitahara, 1983). Rods, as cones, are less sensitive to obliquely incident, than to normally incident backgrounds. At the pupil margin (4 mm) the effect is between 0.368 and 0.976 log10 units smaller for rods. After correction for losses by corneal reflexion and by absorption in the lens, S(r) for rods is reasonably described by the parabolic equation used by Stiles (1937) to quantify the directional sensitivity of cones. The small effect for rods precludes a description as consistently precise as this equation provides for cones. The steepness of the parabolic curve best fitting the directional sensitivity data of the rods of the principal subject was independent of background wave number. For a second subject, whose rods are supposed to be smaller, it was directly proportional to the square of that wave number. The latter is the expectation if the directional sensitivity of this subject's rods were determined by principles outlined in the diffraction theory of Simon (1970).

Humans↗

In search of the elusive long-wave fundamental.

Action spectra for threshold detection of flicker (30 Hz) were obtained on 11 deuteranopes under carefully controlled adaptation conditions. Individual differences were large, so that each one of the long-wave fundamentals proposed by different theorists finds reasonable justification in the spectrum measured on at least one deuteranope. Some deuteranopes' spectra are not described by any one of these "fundamentals". To a first approximation at least, trichromats' spectra show the property of linear additivity. One such trichromat's spectrum agreed well with that of a deuteranope with whom he shares a common erythrolabe, and appears to be uninfluenced by his chlorolabe-filled cones.

Color Perception↗

Visual pigment kinetics in abnormalities of the uvea-retinal epithelium interface in man.

In four patients with abnormalities of the interface between photoreceptors and their vascular supply, bleached visual pigments regenerated abnormally slowly, with double exponential time courses, although the time constant of the first exponential was abnormally fast. During therapy in one patient the slower exponential returned to normal early after treatment, whereas the fast component persisted even after clinically complete recovery. The results could be explained by the following hypothesis. In normal eyes, two pigment-regeneration routes exist, the more rapid (which depends on a short-lived intermediate) playing no role after a long full bleach. Uvea abnormalities cause slower regeneration in the normal main route and slower breakdown of the normally short-lived intermediate.

Adolescent↗

Lack of uniformity in colour matching.

1. The fraction of red in a red--green mixture matched to yellow increased as the intensities of the match constituents were increased sufficiently to bleach appreciable chlorolabe and erythrolabe. 2. All changes in matching found for a given normal trichromat, (i) with increase in the intensities of the matching components, (ii) as a function of time after the onset of very intense components, (iii) with change in the pupil region through which light enters the eye, and (iv) with change in the region of the retina under test, are consistent with the assumption that matching depends upon the absorption of light in three kinds of (individually colour blind) cones, each with its own visual pigment, provided that the lambda max densities of the latter can vary in the range 0.25--1.0 (common logarithmic units) depending upon the subject. 3. Individual differences in matching among normal (as well as among both varieties of red--green anomalous) trichromats, on the other hand, suggest that the extinction spectra of the cone pigments sensitive to long and medium wave lengths may differ from one trichromat to the next.

Adolescent↗

Cone pigments in human deutan colour vision defects.

1. The Nagel anomaloscope, neutral points and dichromatic matches to a spectral green light identified a population of seventy red-green dichromats. 2. The anomaloscope settings allow the calculation of the relative action spectrum of the match at the wave-length of the red (645 nm) and green (535 nm) primaries. The distribution of this ratio is bimodal; there are two clusters with a gap of about 0-75 long units between. Among the thirty-eight deuteranopes there are wide differences in anomaloscope matches; similar differences appear among the thirty-two protanopes. 3. Retinal densitometry of the foveas of fifteen of the deuteranopes is compared and contrasted with measurements on trichromats. In the former, only one photolabile pigment is found in the red-green region of the spectrum; normals always have two. The view of Rushton (1965a) that deuteranopes have erythrolabe but no measurable chlorolabe is confirmed for each member of this group. 4. Simple deuteranomalous show two red-green cone pigments. The difference spectra of extreme deuteranomalous are very similar to those found in deuteranopia. 5. Individual differnce in kinetics (photosensitivity, time constant of regeneration) and in the density and lambdamax of the difference spectrum of erythrolabe in deuteranopia are appreciable; the reasons for these differences are not clear.

Color Vision Defects↗

Variation in the action spectrum of erythrolabe among deuteranopes.

1. Eight deuteranopes matched a mixture of a monochromatic light on the long wave side of the neutral point and a violet (450 nm) primary to a fixed white as well as a monochromatic light on the short wave side of the neutral point mixed with a red (650 nm) primary, to the same white. For lambda greater than 530 nm, the former set of matches defined the action spectrum of the long wave sensitive foveal cones, and for lambda less than 480 nm, the latter that of the short wave sensitive cones. 2. Individual differences in the former matches were approximately correlated with the respective ratio of the sensitivities of the wave-length of the anomaloscope primaries, in a way that individual differences of the latter were not. 3. Assuming that eye media differences alone account for the differences in long wave sensitive foveal action spectra, the spectral reflectivity of the foveal fundus was predicted for these deuteranopes. The prediction is inconsistent with measurement. 4. Thirteen deuteranopes matched monochromatic spectral lights with a green (535 nm) and a blue (460 nm) primary. The result were analysed by von Kries' method in which differences in matching due to differences in eye media absorption are obviated. The matches of five differed significantly from one another when so analysed. It was concluded that at least one of two action spectra of the foveal cones of every one of these five differed from that of all of the others. 5. The canon that deuteranopes accept normal colour matches was evaluated by confronting a single normal with five deuternopes in the analytical anomaloscope of Baker & Rushton, set in the mode of each of the five in turn. Obvious differences existed between this normal's matches and those of four of five deuteranopes. 6. Explanations for differences in the spectrum of erythrolabe in different deuteranopes are evaluated. The possibilities that all have the identical visual pigment but (a) in cones with different optical funnelling properties or (b) in different optical densities are considered. Preliminary results are not in agreement with the expectations of either of these ideas. 7. It is suggested that the visual pigment in the foveal long wave sensitive cones of different deuternopes (and of different normals) may have different extinction spectra. The idea is consistent with micro-spectrophotometric measurements of rhodopsin in individual rods from different frogs (Bowmaker, Loew & Leibman, 1975).

Color Vision Defects↗

The red and green cone visual pigments of deuternomalous trichromacy.

1. Three "simple" deuteranomalous trichromats match with abnormally low "red" tristimulus values throughout the spectrum and abnormally high "green" tristimulus values in the long wave end of the spectrum which become normal (and then low) in the yellow-green. The spectrum locus of this transition differs from one anomalous to the other. Differences in the matches of two of these cannot be due to differences in eye media transmissivities alone. Therefore these two deuteranomalous have different cone visual pigments. 2. The analytical anomaloscope was used in the confrontation of one deuteranomalous with six deuteranopes in turn. In each confrontation the deuteranope set the anomaloscope in his mode and adjusted the intensity of the monochromatic light for a match. Deuteranomalous matches were rejected by four of these six deuteranopes. 3. They were accepted by two of the six. These two rejected each other's matches in a way not attributable to differences in eye media transmissivity. 4. Three different psychophysical techniques were used to measure the action spectra of the long wave cones of these two deuternopes. All three methods reveal small but systematic differences in lambdamax and shape of the curve for the one deuteranope compared with that of the other. 5. In red-green spectral range, these spectra are accurately described by different linear combinations of the color matching functions of the same deuteranomalous whose matches the two deuteranopes accept. Linear combinations of those of a second deuteranomalous, with at least one different kind of cone, fit less well. 6. The wave length discrimination curve of the former deuteranomalous was measured with a new method. The curves of two normals were also obtained for comparison. Wave-length discrimination predictions from the Stiles (1946) line element theory were compared to the anomalous curve. The deuteranopic action spectra were used in the line element to compute this deuteranomalous' discrimination. There is reasonable first order correspondence between prediction and observation, but the prediction is sensitive to small changes in the derivatives of the logarithms of the action spectra. 7. Line element prediction of the deuteranomalous step-by-step luminous efficiency curve is insensitive to such uncertainties. The agreement with expectation from the above assumptions and the measured step-by-step deuteranomalous luminous efficiency curve in the red-green part of the spectrum is therefore good. 8. It is concluded that the erythrolabe in one deuternope's long wave cones has the action spectrum of this deuteranomalous' long and the erythrolabe in the other deuternope's long wave sensitive cones has that of this deuternomalous' medium wave cones. This leads to a general hypothesis about the nature of all forms of red-green colour vision defects transmitted recessively on the X chromosome.

Color Vision Defects↗