The cone/horizontal cell network: a possible site for color constancy

Color vision is spectrally opponent, suggesting that spectrally opponent neurons, such as the horizontal cells in fish and turtle retinae, play a prominent role in color discrimination. In the accompanying paper (Kraaij et al., 1998), it was shown that the output signal of the horizontal cell system to the cones is not at all spectrally opponent. Therefore, a role for the spectrally opponent horizontal cells in color discrimination seems unlikely. In this paper, we propose that the horizontal cells play a prominent role in color constancy and simultaneous color contrast instead of in color discrimination. We have formulated a model of the cone/horizontal cell network based on measurements of the action spectra of the cones and of the feedback signal of the horizontal cell system to the various cone types. The key feature of the model is (1) that feedback is spectrally and spatially very broad and (2) that the gain of the cone synapse strongly depends on the feedback strength. This makes the synaptic gain of the cones strongly dependent on the spectral composition of the surround. Our model, which incorporates many physiological details of the outer retina, displays a behavior that can be interpreted as color constancy and simultaneous color contrast. We propose that the horizontal cell network modulates the cone synaptic gains such that the ratios of the cone outputs become almost invariant with the spectral composition of the global illumination. Therefore, color constancy appears to be coded in the retina.     

Distribution and development of short-wavelength cones differ between Macaca monkey and human fovea

Macaca monkey and humans have three cone types containing either long-wavelength (L), medium-wavelength (M), or short-wavelength (S)-specific opsin. The highest cone density is found in the fovea, which mediates high visual acuity. Most studies agree that the adult human fovea has a small S cone-free area, but data are conflicting concerning S-cone numbers in the adult Macaca monkey fovea, and little evidence exists for how either primate fovea develops its characteristic cone pattern. Single- and double-label in situ hybridization and immunocytochemistry have been used to determine the pattern of foveal S cones in both the fetal and adult Macaca and human. Both labels find a clear difference at all ages between monkey and human. Adult humans have a distinct but variable central zone about 100 microm wide that lacks S cones and is surrounded by a ring in which the S-cone density is 8%. This S cone-free zone is detectable at fetal week 15.5 (Fwk15.5), shortly after S opsin is expressed, and is similar to the adult by Fwk20.5. Adult monkey foveas have an overall S-cone foveal density of 10%, with several areas lacking a few S cones that are not coincident with the area of highest cone density. A surrounding zone at 200-microm eccentricity has an S-cone density averaging 25%, but, by 800 microm, this has decreased to 11%. Fetal day 77-135 monkeys all have a distribution and density of foveal S cones similar to adults, although the high-density ring is not obvious in fetal retinas. Estimates of the numbers of S cones missing in the fetal human fovea range from 234 to 328, whereas no more than 40 are missing in the fetal monkey. These results show that, in these two trichromatic primates, S-cone distribution and the developmental mechanisms determining S-cone topography are markedly different from the time that S cones are first detected.     

Identification, classification and anatomical segregation of cells with X-like and Y-like properties in the lateral geniculate nucleus of old-world primates

1. All the cells (158) that we studied in the lateral geniculate nuclei of Macaca nemestrina and Macaca irus could be distinguished as either X-like or Y-like on the basis of their responses to tests developed to classify cat retinal and lateral geniculate nucleus cells. These tests include responses to stationary spots, fast moving wands and moving gratings. 2. Response latencies to electrical stimulation of the optic chiasm were determined for 130 cells; no X-like cell showed a latency shorter than 1-7 ms, no Y-like cell showed a latency longer than 1-6 ms. Primate lateral geniculate nucleus cells with X-like properties thus receive their excitatory input from retinal cells with slowly conducting axons and these most probably include the tonic ganglion cells described by Gouras (1968, 1969); Y-like lateral geniculate nucleus cells are driven by retinal cells with faster conducting axons, most probably including the phasic ganglion cells described by Gouras. 3. Wiesel & Hubel (1966) classified monkey lateral geniculate nucleus cells into four main types based on their receptive-field properties, as revealed by spectrally and spatially distinct stimuli. We find that all Type I and Type II cells show X-like properties; all type IV cells show Y-like properties. Type III consists of a subtype that show X-like properties, here termed Type IIIx, and a subtype that show Y-like properties, here termed Type IIIy. 4. The first cells encountered as the micro-electrode reached the lateral geniculate nucleus were always X-like. In some penetrations only X-like cells were encountered as the electrode moved downward through the lateral geniculate nucleus. In the remaining penetrations, after recording X-like cells through most of the lateral geniculate nucleus, Y-like cells were then encountered. No X-like cells were found below Y-like cells. thus these two classes of cells are anatomically segregated within the primate lateral geniculate nucleus. Electrode marking showed the borger between X-like and Y-like cells to correspond to the border between the paro- and magnocellular layers of the lateral geniculate nucleus. Thus X-like cells (i.e. Types I, II and IIIx) occur in the parvocellular layers, Y-like cells (i.e. Types IIIy and IV)in the magnocellular layers.     

Absence of short-wavelength sensitive cones in the retinae of seals (Carnivora) and African giant rats (Rodentia)

Most non-primate mammals have two types of cone: short-wavelength sensitive (S) and middle-to-long-wavelength sensitive (M/L) cones. In two species of African giant rats, Cricetomys gambianus and C. emini, and in two species of earless seals, Phoca hispida and P. vitulina, the retinal cone types and cone distributions were assessed with antibodies specific for the M/L-cone opsin and the S-cone opsin, respectively. All four species were found to completely lack S-cones, while M/L-cones were present in low densities. M/L-cone densities, rod densities and cone/rod ratios were determined across the retina. Cone proportions are about 0.3-0. 5% in C. gambianus, 0.5-0.8% in C. emini, and 1.5-1.8% in P. hispida. An absence of S-cones has previously been reported in a few nocturnal mammals. As earless seals are visually active during night and day, we conclude that an absence of S-cones is not exclusively associated with nocturnality. The functional and comparative aspects are discussed.     

Human cone spectral sensitivities: a progress report

The spectral sensitivities of the short (S-), middle (M-) and long (L-) wave-sensitive cones have been measured in normal trichromats and in dichromats and monochromats of known genotype. For the S-cone sensitivities, three blue-cone monochromats and five normals were used; for the M-cone sensitivities, nine protanopes (three with a single L1M2 gene, three with a single L2M3 gene, one with both an L1M2 and an M gene, and two with both an L2M3 and an M gene); and for the L-cone sensitivities, 22 deuteranopes (five with a single L(ala180) gene and 17 with a single L(ser180) gene). We compare existing cone spectral sensitivity estimates with these results and with tritanopic color matches. The new findings are more consistent with the cone fundamentals of Stockman et al. (JOSA 1993(A10), 2491) than with those of Smith and Pokorny (Vision Research 1975(15), 161). The discrepancies that we find, however, are sufficient to warrant the replacement of both sets.     

Segregation of receptive field properties in the lateral geniculate nucleus of a New-World monkey, the marmoset Callithrix jacchus

The lateral geniculate nucleus (LGN) in humans and Old-World monkeys is dominated by the representation of the fovea in the parvocellular (PC) layers, and most PC cells in the foveal representation have red-green cone opponent receptive field properties. It is not known whether these features are both unique to trichromatic primates. Here we measured receptive field properties and the visuotopic organization of cells in the LGN of a New-World monkey, the marmoset Callithrix jacchus. The marmoset displays a polymorphism of cone opsins in the medium-long wavelength (ML) range, which allows the LGN of dichromatic ("red-green color blind") and trichromatic individuals to be compared. Furthermore, the koniocellular-interlaminar layers are segregated from the main PC layers in marmoset, allowing the functional role of this subdivision of the LGN to be assessed. We show that the representation of the visual field in the LGN is quantitatively similar in dichromatic and trichromatic marmosets and is similar to that reported for macaque; the vast majority of LGN volume is devoted to the central visual field. ON- and OFF-type responses are partially segregated in the PC layers so that responses are more commonly encountered near the external border of each layer. The red-green (ML) opponent cells in trichromatic animals were all located in the PC layers, and their receptive fields were within 16 degrees of the fovea. The koniocellular zone between the PC and magnocellular layers contained cells that receive excitatory input from short wavelength sensitive cones ("blue- cells") as well as other nonopponent cells. These results suggest that the basic organization of the LGN is common to dichromatic and trichromatic primates and provide further evidence that ML and SWS opponent signals are carried in distinct subdivisions of the retinogeniculocortical pathway.     

Separate colour-opponent mechanisms underlie the detection and discrimination of moving chromatic targets

Current opinion holds that human colour vision is mediated primarily via a colour-opponent pathway that carries information about both wavelength and luminance contrast (type I). However, some authors argue that chromatic sensitivity may be limited by a different geniculostriate pathway, which carries information about wavelength alone (type II). We provide psychophysical evidence that both pathways may contribute to the perception of moving, chromatic targets in humans, depending on the nature of the visual discrimination. In experiment 1, we show that adaptation to drifting, red-green stimuli causes reductions in contrast sensitivity for both the detection and direction discrimination of moving chromatic targets. Importantly, the effects of adaptation are not directionally specific. In experiment 2, we show that adaptation to luminance gratings results in reduced sensitivity for the direction discrimination, but not the detection of moving chromatic targets. We suggest that sensitivity for the direction discrimination of chromatic targets is limited by a colour-opponent pathway that also conveys luminance-contrast information, whereas the detection of such targets is limited by a pathway with access to colour information alone. The properties of these pathways are consistent with the known properties of type-I and type-II neurons of the primate parvocellular lateral geniculate nucleus and their cortical projections. These findings may explain the known differences between detection and direction discrimination thresholds for chromatic targets moving at low to moderate velocities.     

Convergence properties of solitary tract neurons responsive to cardiac receptor stimulation in the anesthetized cat

The convergence pattern of cardiac receptors, pulmonary C-fibers, carotid chemoreceptor, and baroreceptor afferents onto neurons within the nucleus of the solitary tract (NTS) was studied in the anesthetized (pentobarbitone sodium, 40 mg/kg,) paralyzed and artificially ventilated cat. Extra- and intracellular recordings were made from NTS neurons while stimulating both cardiac receptors by aortic root injections of veratridine (1-3 micrograms/kg) and pulmonary C-fibers by a right atrial injection of phenylbiguanide (10-20 micrograms/kg). The ipsilateral carotid body was stimulated by using arterial injection of CO2-saturated bicarbonate solution, whereas inflation of the ipsilateral carotid sinus was used to activate baroreceptors. The ipsilateral cardiac vagal branch, cervical vagus, and carotid sinus nerves were stimulated electrically (1 Hz, 0.2-1 ms, 1-35 V). In 78 NTS neurons recorded either extracellularly (n = 47) or intracellularly (n = 31), electrical stimulation of the cardiac branch of the vagus nerve evoked synaptic potentials (spikes and/or excitatory postsynaptic potentials) with an onset latency between 4 and 220 ms. Some neurons displayed both short and long latency inputs(15.5 +/- 1.8 and 160.0 +/- 8.5 ms; n = 14). Of these 78 neurons, 24 responded to veratridine stimulation of cardiac receptors (i.e., cardioreceptive neurons) by exhibiting an augmenting-decrementing discharge of 37 +/- 4 s in duration with a peak frequency of 30 +/- 5 Hz. Convergence from other cardiorespiratory receptors was noted involving either carotid chemoreceptors (n = 7) or pulmonary C-fibers (n = 4) or from both carotid chemoreceptors and pulmonary C-fibers (n = 6). In contrast, only one cardioreceptive NTS neuron was activated by distension of the carotid sinus. Recording sites recovered were confined to the medial NTS at the level of the area postrema and extended caudally into the commissural subnucleus. Our results indicate a convergence of carotid chemoreceptor and pulmonary C-fiber afferent inputs to cardioreceptive NTS neurons. With the paucity of baroreceptor inputs to these neurons it is suggested that sensory integration within the NTS may reflect regulatory versus defensive or protective reflex control.     

Effects of saccades on the activity of neurons in the cat lateral geniculate nucleus

Effects of saccades on individual neurons in the cat lateral geniculate nucleus (LGN) were examined under two conditions: during spontaneous saccades in the dark and during stimulation by large, uniform flashes delivered at various times during and after rewarded saccades made to small visual targets. In the dark condition, a suppression of activity began 200-300 ms before saccade start, peaked approximately 100 ms before saccade start, and smoothly reversed to a facilitation of activity by saccade end. The facilitation peaked 70-130 ms after saccade end and decayed during the next several hundred milliseconds. The latency of the facilitation was related inversely to saccade velocity, reaching a minimum for saccades with peak velocity >70-80 degrees /s. Effects of saccades on visually evoked activity were remarkably similar: a facilitation began at saccade end and peaked 50-100 ms later. When matched for saccade velocity, the time courses and magnitudes of postsaccadic facilitation for activity in the dark and during visual stimulation were identical. The presaccadic suppression observed in the dark condition was similar for X and Y cells, whereas the postsaccadic facilitation was substantially stronger for X cells, both in the dark and for visually evoked responses. This saccade-related regulation of geniculate transmission appears to be independent of the conditions under which the saccade is evoked or the state of retinal input to the LGN. The change in activity from presaccadic suppression to postsaccadic facilitation amounted to an increase in gain of geniculate transmission of approximately 30%. This may promote rapid central registration of visual inputs by increasing the temporal contrast between activity evoked by an image near the end of a fixation and that evoked by the image immediately after a saccade.     

Quantitative analysis of triphasic (H3) horizontal cell-cone connectivity in the cyprinid fish (roach) retina

Although horizontal cells encode chromatic information by means of a variety of spectrally opponent light-evoked response patterns, their synaptic connections with the different spectral classes of cone are not completely understood. In the cyprinid fish retina, where a hierarchical set of interactions between horizontal cells and cone types has been proposed, a particular type of horizontal cell generates light-evoked triphasic (red-hyperpolarizing/green depolarizing/blue-hyperpolarizing) responses. In the present study, we have studied the cone connectivity of these cells by intracellular recording and staining in the roach retina. The horizontal cells were first identified electrophysiologically using spectral stimuli, and then stained intracellularly with horseradish peroxidase. Light microscopy revealed that the cells had consistent H3-like morphologies. At an ultrastructural level, these horizontal cells were deduced to contact selectively blue-sensitive cones. Within the cone pedicles, the majority (approximately 80%) of the contacts were "central" to synaptic ribbons. Some 50% of the "lateral" processes were large and engulfed cone cytoplasm. Spinules were present within the contacted pedicles but not upon the dendrites of the stained horizontal cells, although previous work had suggested that horseradish peroxidase would not interfere with spinule dynamics. The results are discussed in terms of existing modes of horizontal cell-cone connectivity in cyprinid fish retinae.     

Functional organization of owl monkey lateral geniculate nucleus and visual cortex

The nocturnal, New World owl monkey (Aotus trivirgatus) has a rod-dominated retina containing only a single cone type, supporting only the most rudimentary color vision. However, it does have well-developed magnocellular (M) and parvocellular (P) retinostriate pathways and striate cortical architecture [as defined by the pattern of staining for the activity-dependent marker cytochrome oxidase (CO)] similar to that seen in diurnal primates. We recorded from single neurons in anesthetized, paralyzed owl monkeys using drifting, luminance-modulated sinusoidal gratings, comparing receptive field properties of M and P neurons in the lateral geniculate nucleus and in V1 neurons assigned to CO "blob," "edge," and "interblob" regions and across layers. Tested with achromatic stimuli, the receptive field properties of M and P neurons resembled those reported for other primates. The contrast sensitivity of P cells in the owl monkey was similar to that of P cells in the macaque, but the contrast sensitivities of M cells in the owl monkey were markedly lower than those in the macaque. We found no differences in eye dominance, orientation, or spatial frequency tuning, temporal frequency tuning, or contrast response for V1 neurons assigned to different CO compartments; we did find fewer direction-selective cells in blobs than in other compartments. We noticed laminar differences in some receptive field properties. Cells in the supragranular layers preferred higher spatial and lower temporal frequencies and had lower contrast sensitivity than did cells in the granular and infragranular layers. Our data suggest that the receptive field properties across functional compartments in V1 are quite homogeneous, inconsistent with the notion that CO blobs anatomically segregate signals from different functional "streams."     

Identification and distribution of photoreceptor subtypes in the neotenic tiger salamander retina

The neotenic tiger salamander retina is a major model system for the study of retinal physiology and circuitry, yet there are unresolved issues regarding the organization of the photoreceptors and the photoreceptor mosaic. The rod and cone subtypes in the salamander retina were identified using a combination of morphological and immunocytochemical markers for specific rod and cone opsin epitopes. Because the visual pigment mechanisms present in the tiger salamander retina are well characterized and the antibodies employed in these studies are specific for particular rod and cone opsin epitopes, we also were able to identify the spectral class of the various rod and cone subtypes. Two classes of rods corresponding to the "red" and "green" rods previously reported in amphibian retinas were identified. In serial semithin section analyses, rods and cones comprised 62.4+/-1.4% and 37.6+/-1.4% of all photoreceptors, respectively. One rod type comprising 98.0+/-0.7% of all rods showed the immunological and morphological characteristics of "red" rods, which are maximally sensitive to middle wavelengths. The second rod subtype comprised 2.0+/-0.7% of all rods and possessed the immunological and morphological characteristics of "green" rods, which are maximally sensitive to short wavelengths. By morphology four cone types were identified, showing three distinct immunological signatures. Most cones (84.8+/-1.5% of all cones), including most large single cones, the accessory and principal members of the double cone, and some small single cones, showed immunolabeling by antisera that recognize long wavelength-sensitive cone opsins. A subpopulation of small single cones (8.4+/-1.7% of all cones) showed immunolabeling for short wavelength-sensitive cone opsin. A separate subpopulation of single cones which included both large and small types (6.8+/-1.4% of all cones) was identified as the UV-Cone population and showed immunolabeling by antibodies that recognize rod opsin epitopes. Analysis of flatmounted retinas yielded similar results. All photoreceptor types appeared to be distributed in all retinal regions. There was no obvious crystalline organization of the various photoreceptor subtypes in the photoreceptor mosaic.     

Color from invisible flicker: a failure of the Talbot-Plateau law caused by an early 'hard' saturating nonlinearity used to partition the human short-wave cone pathway

The Talbot-Plateau law fails for flicker detected by the short-wavelength-sensitive (S) cones: a 30-40 Hz target, flickering too fast for the flicker to be resolved, looks more yellow than a steady target of the same average intensity. The color change, which is produced by distortion at an early compressive nonlinearity, was used to reveal a slightly bandpass S-cone temporal response before the distortion site and a lowpass response after it. The nonlinearity is probably a 'hard' nonlinearity that arises because the S-cone signal is limited by a response ceiling, which the mean signal level approaches and exceeds as the S-cone adaptation level increases. The nonlinearity precedes the combination of flicker signals from all three cone types.     

Spatial and temporal expression of cone opsins during monkey retinal development

The primate retina requires a coordinated series of developmental events to form its specialized photoreceptor topography. In this study, the temporal expression of cone photoreceptor opsin was determined in Macaca monkey retina. Markers for mRNA and protein that recognize short wavelength (S) and long/medium wavelength (L/M) opsin were used to determine (1) the temporal and spatial patterns of opsin expression, (2) the spatial relationship between S and L/M cones at the time of initial opsin expression, and (3) the relative time of cone and rod opsin expression (Dorn et al. [1995] Invest. Ophthalmol. Vis. Sci. 36:2634-2651). Adult cone outer segments were recognized by either L/M or S opsin antiserum. Of all adult cone inner segments, 88-90% contained L/M opsin mRNA, whereas 10-12% contained S opsin mRNA. Fetal cones initially showed cell membrane as well as outer segment labeling for opsin protein, but cell membrane labeling disappeared by birth. No cones at any age contained markers for both S and L/M opsin mRNA or protein. S and L/M opsin protein appeared in the fovea at fetal day 75. Once opsin expression progressed beyond the fovea, both mRNA and protein for S opsin were consistently detected more peripherally than L/M opsin. Cones at the peripheral edge of S opsin expression had basal telodendria that appeared to reach toward neighboring cones. Because interactions between cone populations could organize the cone mosaic, the spatial relationship between S cones and the first cones to express L/M protein was analyzed quantitatively by using double-label immunocytochemistry. No consistent relationship was found between these two cone populations. Cones are generated at least 1 week before rods across monkey retina. However, rod opsin protein appears in and around the fovea at fetal day 66, 1 week before cone opsin protein. This suggests that independent local factors control differentiation in these two photoreceptor populations.     

The human S-cone electroretinogram and its variation among subjects with and without L and M-cone function

PURPOSE: To examine the S-cone ERG in subjects with and without L and M-cone function. METHODS: Ganzfeld spectral flashes in the presence of strong Ganzfeld adapting fields are used to elicit S-cone ERGs. RESULTS: The S-cone ERG b-wave ranges from 0.2 to 4 mV in amplitude and 38-45 msec in implicit time. There is a progressive decrease in amplitude with age. The response is similar in subjects with or without L and M cone function. CONCLUSION: The S-cone ERG is detectable in subjects of all ages, but intersubject variability limits its diagnostic usefulness. The S-cone ERG is slightly later than but does not appear to be obviously influenced by the L and M-cone ERG.     

The enhanced S cone syndrome: an analysis of receptoral and post-receptoral changes

The purpose of the study was to test the hypothesis that the retinae of patients with enhanced S cone syndrome (ESCS) have more S cones than the normal retina and these cones have replaced some of the L and M cones. Standard and spectral full-field electroretinograms, measurements of L, M, and S cone system sensitivities and S cone acuity were obtained from three patients with ESCS. The results were qualitatively consistent with the presence of more S cones and more S cone ganglion cells. To test this hypothesis further, a model of the receptoral and post-receptoral components of the S cone system was used in conjunction with psychophysical measurements of S cone system sensitivity under flashed and steady-state adaptation conditions. Within the context of the model, the data were consistent with an increase in the number of S cones and S - (L + M) ganglion cells and with a decrease in the total L + M cone input to each S - (L + M) ganglion cell.     

Strobe rearing prevents the convergence of inputs with different response timings onto area 17 simple cells

Strobe rearing prevents the convergence of inputs with different response timings onto area 17 simple cells. J. Neurophysiol. 80: 3005-3020, 1998. The preceding paper showed that the loss of direction selectivity in simple cells induced by strobe rearing reflects the elimination of spatially ordered response timing differences across the receptive field that underlie spatiotemporal (S-T) inseparability. Here we addressed whether these changes reflected an elimination of certain timings or an alteration in how timings were associated in single cells. Timing in receptive fields was measured using stationary bars undergoing sinusoidal luminance modulation at different temporal frequencies (0.5-6 Hz). For each bar position, response phase versus temporal frequency data were fit by a line to obtain two measures: absolute phase and latency. In normal cats, many individual simple cells display a wide range of timings; in layer 4, the mean range for absolute phase and latency was 0.21 cycles and 39 ms, respectively. Strobe rearing compressed the mean timing ranges in single cells, to 0.08 cycles and 31 ms, respectively, and this compression accounted for the loss of inseparability. A similar compression was measured in layer 6 cells. In contrast, the range of timing values across the simple-cell population was relatively normal. Single cells merely sampled narrower than normal regions of the timing space. We sought to understand these cortical changes in terms of how inputs from the lateral geniculate nucleus (LGN) may have been affected by strobe rearing. In normal cats, a wide range of absolute phase and latency values exists among lagged and nonlagged LGN cells, and these thalamic timings account for most of the cortical timings. Also, S-T inseparability in many simple cells can be attributed to the convergence of lagged and/or nonlagged inputs. Strobe rearing did not change the sampling of lagged and nonlagged cells, and the geniculate timings continued to account for most of the cortical timings. However, strobe rearing virtually eliminated cortical receptive fields with mixed lagged and nonlagged timing, and it compressed the timing range in cells dominated by one or the other geniculate type. Thus strobe rearing did not eliminate certain timings in LGN or cortex, but prevented the convergence of different timings on single cells. To account for these results, we propose a developmental model in which strobe stimulation alters the correlational structure of inputs based on their response timing. Only inputs with similar timing become associated on single cortical cells, and this produces S-T separable receptive fields that lack the ability to confer a preferred direction of motion.     

Undetectable S cone electroretinogram b-wave in complete congenital stationary night blindness

AIMS: The short wavelength sensitive (S) cone electroretinograms (ERGs) were examined in two patients with the complete type of congenital stationary night blindness (CSNB). METHODS: Ganzfeld spectral flashes in the presence of strong white adapting fields were used to elicit the S cone ERGs. RESULTS: The S cone ERG b-wave was not detectable to short wavelength stimuli, while the mixed long (L) and middle (M) wavelength sensitive cone responses appeared normal in waveforms with normal amplitude in both patients. Both patients had normal colour vision on the Farnsworth Panel D-15. CONCLUSIONS: These ERG results indicated that the S cone system as well as rod system is more impaired in complete CSNB than the L and M cone system and that normal colour vision may not depend on a normal S cone ERG to full field stimuli.     

Medullary swallowing-related neurons in the anesthetized cat

Swallowing-related neurons (SRNs) were recorded systematically in the medulla oblongata of urethane-anesthetized cats. The SRNs received orthodromic inputs from the superior laryngeal nerve (SLN) and showed transient changes in their activity synchronous with swallowing. These neurons could be divided into three types. Type I SRNs are sensory-relay neurons from the SLN in the nucleus of the tractus solitarius (NTS), type II are interneurons located diffusely in the parvocellular reticular formation ventral to the NTS, which received oligosynaptic inputs from the SLN, and type III are motoneurons in the nucleus ambiguus. Some type II neurons still showed the swallowing activity after the animals were paralysed, which suggests that they could be involved in the generation of swallowing outputs.     

Dichromacy characterized by chrominance planes

Dichromacy is described in terms of dichromatic opponent colour spaces. By means of the perceptual criteria 'equally bright', 'neither blue nor yellow' and 'neither red nor green' and embedding in a three-dimensional colour space, it is possible for each type of dichromat to quantify a null-chrominance plane and a null-luminance plane, both of which intersect in the missing colour. These two null planes (or the trace of their intersection with the chromaticity chart) are the chromaticities of the dichromatic opponent primaries. Since a null-luminance plane contains only colour ('chrominance'), it is simply a chrominance plane. Under the assumption that the retinal short-wavelength cones do not contribute to luminance, the chrominance planes of the three types of dichromats intersect in a common straight line, the 'blue' fundamental primary vector. This constellation may serve as a general characterisation of dichromacy.