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J Kremers

Publications and source records attributed to J Kremers.

At least 37 records · Page 2Linked to original sources

Parallel pathways in the retina of Old and New World primates.

Old-world simians are all trichromats, but in most new-world primates there is a polymorphism; males are dichromats but most females are trichromats. In the old world simian, luminance and red-green chromatic channels defined by psychophysical experiments have as a basis parasol ganglion cells of the magnocellular (MC) pathway and midget ganglion cells of the parvocellular (PC) pathway respectively. Small bistratified ganglion cells provide a basis for a blue-yellow chromatic channel, which should probably be considered a separate entity. In both dichromatic and trichromatic new-world animals, the MC pathway and the small bistratified, blue-yellow system seem anatomically and physiologically similar to those in their old-world relatives. The midget ganglion cells of the parvocellular pathway in trichromats are anatomically and physiologically similar to the old-world pattern. In dichromatic animals, they are anatomically similar and physiologically resemble those of trichromatic animals, except for the lack of chromatic opponency. We conclude that these three systems may from a basic pattern for the visual pathway of primates. However, the results from dichromats indicate that the evolution of trichromacy may be found to be more complex than presently supposed.

Animals↗

The spatial precision of macaque ganglion cell responses in relation to vernier acuity of human observers.

Responses of parafoveal macaque ganglion cells were measured as a function of the contrast and position of an edge flashed within their receptive fields. The goal was to determine the ability of different cell types to signal edge location. For comparison, parafoveal vernier thresholds of human observers were measured with pairs of flashed edges. Cells of the magnocellular (MC-) pathway gave larger responses than cells of the parvocellular (PC-) pathway. Neurometric analyses comparing a cell's response at different edge positions were performed. The positional signal from single MC-pathway cells was more precise than from PC-pathway cells, especially at lower contrasts. In a second analysis, based on the neurophysiological results, responses from a matrix of ganglion cells were generated. Using a simple model, vernier performance expected from such a matrix was predicted as a function of edge length and contrast. Again, the MC-pathway gave a more precise positional signal than the PC-pathway despite the latter's numerical advantage. At contrasts of 20% and below, only the MC-pathway would appear capable of supporting vernier performance with our stimuli. At higher contrasts either the MC- or PC-pathway could provide an adequate signal.

Animals↗

Temporal response of ganglion cells of the macaque retina to cone-specific modulation.

The temporal response of cone inputs to macaque retinal ganglion cells were compared with cone-specific sinusoidal modulation used to isolate each cone type. For all cell types of the parvocellular (PC) pathway, temporal responsivity was similar for short (S)-, middle (M)-, and long (L)-wavelength-sensitive cone inputs, apart from small latency differences between inputs to center and surround. The temporal response resembled that expected from receptor physiology. Responses of cells of the magnocellular pathway to M- or L-cone modulation showed more complex properties indicative of postreceptoral processing. Human psychophysical temporal-sensitivity functions were acquired with S-cone modulation under conditions similar to those for the physiological measurements. Ratios of psychophysical to physiological data from S-cone cells (the only cells that respond to this stimulus) yielded an estimate of the central filter acting upon PC-pathway signals. The filter characteristic could be described by a four-stage low-pass filter with corner frequency 3-5 Hz.

Animals↗

Visual responses in the lateral geniculate nucleus of dichromatic and trichromatic marmosets (Callithrix jacchus).

New-world primates such as the marmoset (Callithrix jacchus) show polymorphism for the middle- to long-wavelength sensitive cone pigments. Each X-chromosome carries a gene for only one of three possible pigments. All males are thus dichromats, but some females will be trichromats. We have investigated the responses of cells of the parvocellular (PC) and magnocellular (MC) systems within animals from a single marmoset family. The middle- to long-wavelength pigment of dichromats was identified physiologically. Trichromats could readily be distinguished from dichromats by the presence of a high proportion of red-green opponent PC-cells. The physiological classification of phenotypes was confirmed with genetic analysis. The pattern of inheritance was consistent with current genetic models. In trichromatic females, the properties of cells resembled in detail those of cells from the PC- and MC-pathways of the macaque. In dichromats, cell responses resembled those of trichromats (except for the lack of opponency in PC-cells); PC-cells showed sustained and MC-cells transient responses, with a lower contrast gain for the former type. One difference was that a proportion of PC-cells in dichromats showed strong rod input even at high levels of retinal illuminance. Thus, in trichromatic marmosets the presence of two middle- to long-wave pigments appears to permit the elaboration of all the physiological properties associated with trichromacy.

Animals↗

Responses to pulses and sinusoids in macaque ganglion cells.

The goal of the study was to compare pulse responses with sinusoidal temporal responsivity. The response of macaque ganglion cells was measured to brief luminance and chromatic pulses and to luminance or chromatic sinusoidal modulation. To make both positive and negative lobes of the pulse response visible, responses to pulses of opposite polarity were combined to yield a linearized pulse response. Tests of superposition were used to evaluate the linearized pulse response to different combinations of pulse duration and Weber contrast. A prediction of the pulse response was derived using sinusoidal responsivity functions and Fourier synthesis. For ganglion cells of the parvocellular (PC) pathway, shape and absolute amplitude of linearized pulse responses corresponded well to the predicted responses over a range of pulse durations at 0.5 and 1.0 Weber contrast for both luminance and chromatic modulation. For ganglion cells of the magnocellular (MC) pathway, shape and amplitude of the linearized pulse responses and the predicted responses corresponded when the contrast-duration product was low. This correspondence held for luminance modulation over a thousand-fold range of retinal illuminance. For contrast-duration combinations that produced a more vigorous response, over 100 imp/sec, the linearized pulse responses of MC-pathway cells became larger and time-advanced relative to the linear prediction until saturation became apparent. Incorporation of high Michelson contrast responses in the Fourier synthesis captured the timing but not the amplitude of the linearized pulse response. The data suggest that a mechanism similar to a contrast gain control acts upon MC- but not PC-pathway-cells. The data confirm that use of linear modelling to describe temporal behaviour of retinal ganglion cells is appropriate for small signals.

Animals↗

The response of macaque ganglion cells and human observers to heterochromatically modulated lights: the effect of stimulus size.

Psychophysical sensitivity of human observers closely resembles responsivity of retinal ganglion cells of the magnocellular (MC-) pathway as a function of the relative phase of heterochromatically modulated lights. The MC-pathway phase effect is absent if the receptive field centre alone is stimulated. Here we confirm this physiological result, and show that the psychophysical phase shift is also abolished with small stimuli. The space constant of the psychophysical effect is consistent with a surround diameter for MC-pathway cells in the fovea of about 50 min arc, about 10 times estimated centre diameter. On changing retinal illuminance, the amplitude of the physiological and psychophysical phase shifts also changed in a parallel manner. These experiments support the hypothesis that the physiological origin of psychophysical phase shifts is in the MC-pathway, and indicate the spatial frequency (c. 2c/deg) below which the psychophysical phase shift should become apparent.

Animals↗

Responses of macaque ganglion cells and human observers to compound periodic waveforms.

We measured responses of macaque retinal ganglion cells to different periodic waveforms (sinusoidal, square, rapid-on and rapid-off sawtooth waveforms) for both luminance and equiluminant chromatic modulation. We analyzed the responses with a peak-to-trough detector. At low frequencies, on-center and off-center magnocellular (MC-) pathway cells showed a ten-fold higher responsivity to the rapid-on and rapid-off sawtooth respectively. Red-on (+L-M) and green-on (+M-L) parvocellular (PC-) pathway cells showed a four-fold greater responsivity to rapid red-on and rapid green-on equiluminant chromatic sawtooth waveforms respectively. At an equivalent retinal eccentricity, we measured psychophysical thresholds for luminance stimuli and chromatic stimuli. We concluded that luminance sawtooth sensitivities from psychophysics are consistent with selective detection through MC-pathway on- and off-center channels in the visual system. The differences between the compound periodic waveforms seen in the PC-pathway cell data did not occur in the psychophysics. In a second analysis, cell responses to sinusoidal modulation were used to predict the linear response to square-wave and sawtooth waveforms. PC-pathway cells showed linear temporal behavior over a wide range of contrasts, but MC-pathway cells displayed linear behavior only for low-contrast luminance modulation. Using these linear fits, we implemented a model incorporating central low-pass filtering in the MC- and PC-pathways before the peak-to-trough detector. This model captured better the time scale and relative sensitivity to periodic waveforms found in the psychophysical data.

Adult↗

Physiological mechanisms underlying psychophysical sensitivity to combined luminance and chromatic modulation.

If psychophysical detection thresholds are plotted in a middle-wavelength-sensitive (M) and long-wavelength-sensitive (L) cone coordinate system, the shape of the contour can be used to infer underlying detection mechanisms. We measured responses of macaque ganglion cells to combine chromatic and luminance modulation and expressed our results in such an M,L-cone space. Our aim was to test whether, with the use of this space, readily separable luminance and chromatic psychophysical mechanisms might be expected from physiological data. For parvocellular pathway cells, detection contours approximated elongated ellipses with maximum responsivity to chromatic modulation. The degree of elongation decreased as temporal frequency increased. Responses could be well described by linear subtraction of M- and L-cone signals, with a phase delay of 1-3 deg/Hz. For cells of the magnocellular pathway, detection contours were more complex. Orientation was variable between cells and temporal frequency dependent, and a frequency-doubled component was evoked by chromatic modulation. In relation to psychophysical detection thresholds plotted in such a space, the properties of parvocellular-pathway cells were sufficiently linear and homogeneous to make it plausible that this pathway might form the substrate for a linear chromatic mechanism. The properties of magnocellular-pathway cells, however, indicate that, insofar as a psychophysical luminance mechanism is based on their activity, its signature in the M,L-cone contrast space would be more difficult to identify.

Animals↗

Macaque ganglion cell responses to stimuli that elicit hyperacuity in man: detection of small displacements.

We measured responses of macaque ganglion cells as a function of contrast in a simple hyperacuity task, detection of displacement of an achromatic edge. Responses of ganglion cells of the magnocellular (MC) pathway were much more vigorous than those of cells of the parvocellular (PC) pathway. From the variability in the number of impulses in the response as compared with the distribution of impulses in maintained activity, it was possible to generate receiver operating characteristics for cells of the two pathways, and to predict individual cells' capability to detect a displacement with 75% probability. On comparing cell sensitivities to human psychophysical thresholds (75% probability of correct identification of displacement direction) at an equivalent retinal eccentricity (approximately 6 degrees), we found that one or two additional impulses in two MC pathway cells would suffice to support an ideal detector underlying psychophysical performance, at all contrast levels. Many more PC pathway cells would be required, especially at low contrasts. The much higher signal-to-noise ratio in the MC pathway relative to the PC pathway indicates that the MC pathway is likely to support this and other hyperacuity tasks.

Animals↗

Responses of macaque ganglion cells to movement of chromatic borders.

1. We have measured responses of macaque ganglion cells to moving borders under conditions designed to simulate the minimally distinct border (MDB) task. 2. Extending previous results, we show that minimization of responses of phasic ganglion cells of the magnocellular (MC)-pathway obey the photometric laws of transitivity and additivity. 3. To equal luminance borders, a residual response was present in MC-pathway cells analogous to the second harmonic response seen in these neurones with temporal chromatic modulation. It was proportional to the tritanopic purity difference (magnitude of delta Pt, the rectified middle- to long-wavelength cone opponent signal) between the two colours on either side of the border. For a delta Pt of one, the mean residual response was equivalent to the response evoked by achromatic borders of about 14% luminance contrast. Both these properties of the MC-pathway closely resemble psychophysical estimates as to the distinctness of equal luminance borders. 4. We show how MC-pathway cell responses could be used centrally to support the MDB task. It was difficult to generate a model from responses of tonic ganglion cells of the parvocellular (PC)-pathway which would support the task. 5. The MDB task is still possible psychophysically after blurring the retinal image. Although blurring the border spatially smeared the responses of MC-pathway ganglion cells and reduced their amplitude, responses still went through a minimum close to equal luminance. Thus, blurring the image did not affect the ability of MC-pathway cells to support the task. Blurring the retinal image decreased the 'sharpness' of the border response of tonic, PC-pathway ganglion cells, but response amplitude was unaffected. Response features indicative of centre-surround organization were attenuated. A central mechanism reliant on centre-surround field structure of PC-pathway cells would thus not be able to support the task after blurring. 6. Taken together, these results strongly suggest that the MC-pathway forms the sole physiological substrate of the MDB task, and any contribution of the PC-pathway is, indeed, minimal.

Animals↗

Sensitivity of macaque retinal ganglion cells and human observers to combined luminance and chromatic temporal modulation.

We measured the sensitivity of macaque retinal ganglion cells and human subjects to luminance and chromatic modulation and to two combined conditions as a function of temporal frequency. For both physiological and psychophysical data, we compared the sensitivities to luminance and chromatic modulation with the sensitivities in the combined conditions, using an additivity measure. When the physiological and the psychophysical data were taken together, the results suggested that under the combined conditions psychophysical sensitivity was the envelope of independent achromatic and chromatic mechanisms with physiological substrates in the magnocellular and the parvocellular pathways, respectively. In the combined conditions tested, sensitivity appeared to be set by a chromatic channel below 3 Hz and an achromatic channel above this frequency. This hypothesis was supported by a comparison of detection sensitivities with discrimination thresholds for the presence of chromatic alternation.

Adolescent↗

Angular velocity, not temporal frequency determines circular vection.

This paper shows that the experienced speed of circular vection depends on stimulus speed, not on stimulus temporal frequency. But why would anyone think the contrary? The point is that many modelers in the field of motion perception believe that perceived speed is determined by temporal frequency. Moreover, the optokinetic behaviour of the fly is said to be dependent on the temporal frequency, not the speed, of the stimulus pattern (Reichardt, 1987). It was the aim of the present experiment to test the notion that the experienced speed of circular vection is proportional to stimulus velocity information, which is carried by the temporal and the spatial characteristics of light.

Adult↗

Analytical stereophotogrammetric determination of three-dimensional knee-joint geometry.

An analytical stereophotogrammetric method is introduced to measure the three-dimensional geometry of articular surfaces in vitro. Information of this kind is particularly useful for mathematical joint models and anthropological studies. The method requires no specific equipment, such as a stereocomparator, contrarily to other techniques reported (e.g. Ghosh, 1983) and is relatively simple and inexpensive. The background of the method is outlined in the present paper, and results of accuracy and precision tests are presented. It is shown that an accuracy on the order of 0.2 mm (95% confidence interval) is well feasible in actual knee-joint evaluations, if the measuring procedure is conducted carefully. The method is illustrated by measuring and comparing the articular surface geometries of a bilateral pair of knee joints.

Biomechanical Phenomena↗

Photoreceptor topography and cone-specific electroretinograms.

It is implicit in many cone-specific ERG studies that the amplitude is proportional to the numbers of cones stimulated. The objective of these experiments was to test this idea by comparing ERGs obtained from different areas of the retina with histological data on cone-density distributions. The histology (Curcio et al., 1990) shows that the cumulative number of cones in the human retina increases exponentially with stimulus diameter between 0- and 40-deg eccentricity. L-, M-, and (L+M) cone-driven 30-Hz ERGs were obtained from a series of stimuli with one of the following configurations: (1) Circular stimuli of different angular subtense up to 70-deg diameter. (2) Annuli with 70-deg outer diameter but variable inner diameter. (3) Annuli of constant area but increasing eccentricity. Cone contrasts were equalized for each stimulus condition. The modulated and nonmodulated regions of the screen had the same mean hue and luminance. The data suggest that the L+M cone ERG amplitude increases with stimulus diameter in direct proportion to the estimated number of cones stimulated. Furthermore, the total L+M responses appear to be predicted from individual L and M responses by simple linear summation for both the disc and annular stimuli.

Electroretinography↗

Receptive fields of primate retinal ganglion cells studied with a novel technique.

We have reinvestigated receptive-field structure of ganglion cells of the macaque parafovea using counterphase modulation of a bipartite field. Receptive fields were mapped with luminance, chromatic, and cone-isolating stimuli. Center sizes of middle (M) and long (L) wavelength cone opponent cells of the parvocellular (PC) pathway were consistent with previous estimates (Gaussian radii of 2-4 min of arc, corresponding to center diameters of 6-12 min of arc). We calculate that a large factor of the enlargement relative to cone radius could be blur due to the eye's natural optics. Maps were consistent with cone selectivity in surround mechanisms, which had radii of 5-8 min of arc. For magnocellular (MC) cells, center size estimates were also consistent with grating measurements from the literature (also Gaussian radii of 2-4 min of arc). The surround mechanism contributing the MC-cell frequency-doubled response to chromatic modulation appears to possess a subunit structure, and we speculate it derives from nonlinear summation of signals from M,L-cone opponent subunits, such as midget bipolar cells.

Animals↗

Interaction between rod and cone signals in responses of lateral geniculate neurons in dichromatic marmosets (Callithrix jacchus).

Parvocellular (P-) and magnocellular (M-) cells in the marmoset LGN can receive prominent rod input up to relatively high illuminance levels (Kremers et al., 1997b). In the present paper, we quantify rod and cone input strengths under different retinal illuminance levels. The stimulus was based on the so-called "silent substitution" method. The activities of P- and M-cells of dichromatic animals were recorded extracellularly. We were able to adequately describe the response amplitudes and phases by a vector summation of rod and cone signals. At low retinal illuminance levels, the cells' responses were determined by rod and cone inputs. With increasing illuminances the strength of the cone input increased relative to the rod strength. But, we often found significant rod inputs up to illuminances equivalent to 700 td in the human eye or more. Rod input strength was more pronounced in cells with receptive fields at large retinal eccentricities. The phase differences between rod and cone inputs suggest that the rod signals lag about 45 ms behind the cone signals.

Animals↗

Ganglion cells of a short-wavelength-sensitive cone pathway in New World monkeys: morphology and physiology.

We have studied the morphology and physiology of retinal ganglion cells of a short-wavelength-sensitive cone (SWS-cone) pathway in dichromatic and trichromatic New World anthropoids, the capuchin monkey (Cebus apella) and tufted-ear marmoset (Callithrix jacchus). In Old World anthropoids, in which males and females are both trichromats, blue-ON/yellow-OFF retinal ganglion cells have excitatory SWS-cone and inhibitory middle- and long-wavelength-sensitive (MWS- and LWS-) cone inputs, and have been anatomically identified as small-field bistratified ganglion cells (SB-cells) (Dacey & Lee, 1994). Among retinal ganglion cells of New World monkeys, we find SB-cells which have very similar morphology to such cells in macaque and human; for example, the inner dendritic tree is larger and denser than the outer dendritic tree. We also find blue-on retinal ganglion cells of the capuchin to have physiological responses strongly resembling such cells of the macaque monkey retina; for example, responses were more sustained, with a gentler low frequency roll-off than MC-cells, and no evidence of contrast gain control. There was no difference between dichromatic and trichromatic individuals. The results support the view that SWS-cone pathways are similarly organized in New and Old World primates, consistent with the hypothesis that these pathways form a phylogenetically ancient color system.

Animals↗

Spatial and temporal response properties of the major retino-geniculate pathways of Old and New World monkeys.

Old World monkeys, apes and humans all enjoy trichromatic colour vision, and the absorption spectra of the photopigments are very similar in all species and all individuals. Colour vision in New World monkeys however, is very heterogeneous. In many species, the majority of individuals is dichromatic. Recently, anatomical and electrophysiological studies revealed that the retinal organisation in Old World monkeys and New World monkeys is very similar, although the cells belonging to the parvocellular pathway do not show any colour opponency and their spectral sensitivity is similar to that of the magnocellular cells. Apparently, the magnocellular and parvocellular pathways in the retina and the LGN have not developed as an adaptation to luminance and chromatic processing. It is more likely that the two pathways originally evolved to cover different ranges in the spatio-temporal domain. In the present paper, several spatial and temporal properties of parvo- and magnocellular cells (which are identical for dichromatic and trichromatic animals) are compared.

Animals↗