Showing papers on "Receptive field published in 2000"

Sparse coding and decorrelation in primary visual cortex during natural vision.

Abstract: Theoretical studies suggest that primary visual cortex (area V1) uses a sparse code to efficiently represent natural scenes. This issue was investigated by recording from V1 neurons in awake behaving macaques during both free viewing of natural scenes and conditions simulating natural vision. Stimulation of the nonclassical receptive field increases the selectivity and sparseness of individual V1 neurons, increases the sparseness of the population response distribution, and strongly decorrelates the responses of neuron pairs. These effects are due to both excitatory and suppressive modulation of the classical receptive field by the nonclassical receptive field and do not depend critically on the spatiotemporal structure of the stimuli. During natural vision, the classical and nonclassical receptive fields function together to form a sparse representation of the visual world. This sparse code may be computationally efficient for both early vision and higher visual processing.

Coding of border ownership in monkey visual cortex.

Abstract: Areas V1 and V2 of the visual cortex have traditionally been conceived as stages of local feature representations. We investigated whether neural responses carry information about how local features belong to objects. Single-cell activity was recorded in areas V1, V2, and V4 of awake behaving monkeys. Displays were used in which the same local feature (contrast edge or line) could be presented as part of different figures. For example, the same light‐dark edge could be the left side of a dark square or the right side of a light square. Each display was also presented with reversed contrast. We found significant modulation of responses as a function of the side of the figure in .50% of neurons of V2 and V4 and in 18% of neurons of the top layers of V1. Thus, besides the local contrast border information, neurons were found to encode the side to which the border belongs (“border ownership coding”). A majority of these neurons coded border ownership and the local polarity of luminance‐chromaticity contrast. The others were insensitive to contrast polarity. Another 20% of the neurons of V2 and V4, and 48% of top layer V1, coded local contrast polarity, but not border ownership. The border ownership-related response differences emerged soon (,25 msec) after the response onset. In V2 and V4, the differences were found to be nearly independent of figure size up to the limit set by the size of our display (21°). Displays that differed only far outside the conventional receptive field could produce markedly different responses. When tested with more complex displays in which figure-ground cues were varied, some neurons produced invariant border ownership signals, others failed to signal border ownership for some of the displays, but neurons that reversed signals were rare. The influence of visual stimulation far from the receptive field center indicates mechanisms of global context integration. The short latencies and incomplete cue invariance suggest that the border-ownership effect is generated within the visual cortex rather than projected down from higher levels.

Spectral-Temporal Receptive Fields of Nonlinear Auditory Neurons Obtained Using Natural Sounds

Abstract: The stimulus-response function of many visual and auditory neurons has been described by a spatial-temporal receptive field (STRF), a linear model that for mathematical reasons has until recently been estimated with the reverse correlation method, using simple stimulus ensembles such as white noise. Such stimuli, however, often do not effectively activate high-level sensory neurons, which may be optimized to analyze natural sounds and images. We show that it is possible to overcome the simple-stimulus limitation and then use this approach to calculate the STRFs of avian auditory forebrain neurons from an ensemble of birdsongs. We find that in many cases the STRFs derived using natural sounds are strikingly different from the STRFs that we obtained using an ensemble of random tone pips. When we compare these two models by assessing their predictions of neural response to the actual data, we find that the STRFs obtained from natural sounds are superior. Our results show that the STRF model is an incomplete description of response properties of nonlinear auditory neurons, but that linear receptive fields are still useful models for understanding higher level sensory processing, as long as the STRFs are estimated from the responses to relevant complex stimuli.

Spatial distribution of contextual interactions in primary visual cortex and in visual perception.

Abstract: To examine the role of primary visual cortex in visuospatial integration, we studied the spatial arrangement of contextual interactions in the response properties of neurons in primary visual cortex of alert monkeys and in human perception. We found a spatial segregation of opposing contextual interactions. At the level of cortical neurons, excitatory interactions were located along the ends of receptive fields, while inhibitory interactions were strongest along the orthogonal axis. Parallel psychophysical studies in human observers showed opposing contextual interactions surrounding a target line with a similar spatial distribution. The results suggest that V1 neurons can participate in multiple perceptual processes via spatially segregated and functionally distinct components of their receptive fields.

Visual behaviour mediated by retinal projections directed to the auditory pathway

Abstract: An unresolved issue in cortical development concerns the relative contributions of intrinsic and extrinsic factors to the functional specification of different cortical areas. Ferrets in which retinal projections are redirected neonatally to the auditory thalamus have visually responsive cells in auditory thalamus and cortex, form a retinotopic map in auditory cortex and have visual receptive field properties in auditory cortex that are typical of cells in visual cortex. Here we report that this cross-modal projection and its representation in auditory cortex can mediate visual behaviour. When light stimuli are presented in the portion of the visual field that is 'seen' only by this projection, 'rewired' ferrets respond as though they perceive the stimuli to be visual rather than auditory. Thus the perceptual modality of a neocortical region is instructed to a significant extent by its extrinsic inputs. In addition, gratings of different spatial frequencies can be discriminated by the rewired pathway, although the grating acuity is lower than that of the normal visual pathway.

Spatial sensitivity of macaque inferior temporal neurons

Abstract: Recent findings in dorsal visual stream areas and computational work raise the question whether neurons at the end station of the ventral visual stream can code for stimulus position. The authors provide the first detailed, quantitative data on the spatial sensitivity of neurons in the anterior part of the inferior temporal cortex (area TE) in awake, fixating monkeys. They observed a large variation in receptive field (RF) size (ranging from 2.8 degrees to 26 degrees ). TE neurons differed in their optimal position, with a bias toward the foveal position. Moreover, the RF profiles of most TE neurons could be fitted well with a two-dimensional Gaussian function. Most neurons had only one region of high sensitivity and showed a smooth decline in sensitivity toward more distal positions. In addition, the authors investigated some of the possible determinants of such spatial sensitivity. First, testing with low-pass filtered versions of the stimuli revealed that the general preference for the foveal position and the size of the RFs was not due simply to TE neurons receiving input with a lower spatial resolution at more eccentric positions. The foveal position was still preferred after intense low-pass filtering. Second, although an increase in stimulus size consistently broadened spatial sensitivity profiles, it did not change the qualitative features of these profiles. Moreover, size selectivity of TE neurons was generally position invariant. Overall, the results suggest that TE neurons can code for the position of stimuli in the central region of the visual field.

Modular Organization of Frequency Integration in Primary Auditory Cortex

Abstract: Two fundamental aspects of frequency analysis shape the functional organization of primary auditory cortex. For one, the decomposition of complex sounds into different frequency components is reflected in the tonotopic organization of auditory cortical fields. Second, recent findings suggest that this decomposition is carried out in parallel for a wide range of frequency resolutions by neurons with frequency receptive fields of different sizes (bandwidths). A systematic representation of the range of frequency resolution and, equivalently, spectral integration shapes the functional organization of the iso-frequency domain. Distinct subregions, or "mod- ules," along the iso-frequency domain can be demonstrated with various measures of spectral integration, including pure-tone tuning curves, noise masking, and electrical cochlear stimulation. This modularity in the representation of spectral integration is expressed by intrinsic cortical connections. This organization has implications for our understanding of psychophysical spectral integration measures such as the critical band and general cortical coding strategies.

Trigeminohypothalamic and Reticulohypothalamic Tract Neurons in the Upper Cervical Spinal Cord and Caudal Medulla of the Rat

Abstract: Sensory information that arises in orofacial organs facilitates exploratory, ingestive, and defensive behaviors that are essential to overall fitness and survival. Because the hypothalamus plays an important role in the execution of these behaviors, sensory signals conveyed by the trigeminal nerve must be available to this brain structure. Recent anatomical studies have shown that a large number of neurons in the upper cervical spinal cord and caudal medulla project directly to the hypothalamus. The goal of the present study was to identify the types of information that these neurons carry to the hypothalamus and to map the route of their ascending axonal projections. Single-unit recording and antidromic microstimulation techniques were used to identify 81 hypothalamic-projecting neurons in the caudal medulla and upper cervical (C(1)) spinal cord that exhibited trigeminal receptive fields. Of the 72 neurons whose locations were identified, 54 were in laminae I-V of the dorsal horn at the level of C(1) (n = 22) or nucleus caudalis (Vc, n = 32) and were considered trigeminohypothalamic tract (THT) neurons because these regions are within the main projection territory of trigeminal primary afferent fibers. The remaining 18 neurons were in the adjacent lateral reticular formation (LRF) and were considered reticulohypothalamic tract (RHT) neurons. The receptive fields of THT neurons were restricted to the innervation territory of the trigeminal nerve and included the tongue and lips, cornea, intracranial dura, and vibrissae. Based on their responses to mechanical stimulation of cutaneous or intraoral receptive fields, the majority of THT neurons were classified as nociceptive (38% high-threshold, HT, 42% wide-dynamic-range, WDR), but in comparison to the spinohypothalamic tract (SHT), a relatively high percentage of low-threshold (LT) neurons were also found (20%). Responses to thermal stimuli were found more commonly in WDR than in HT neurons: 75% of HT and 93% of WDR neurons responded to heat, while 16% of HT and 54% of WDR neurons responded to cold. These neurons responded primarily to noxious intensities of thermal stimulation. In contrast, all LT neurons responded to innocuous and noxious intensities of both heat and cold stimuli, a phenomenon that has not been described for other populations of mechanoreceptive LT neurons at spinal or trigeminal levels. In contrast to THT neurons, RHT neurons exhibited large and complex receptive fields, which extended over both orofacial ("trigeminal") and extracephalic ("non-trigeminal") skin areas. Their responses to stimulation of trigeminal receptive fields were greater than their responses to stimulation of non-trigeminal receptive fields, and their responses to innocuous stimuli were induced only when applied to trigeminal receptive fields. As described for SHT axons, the axons of THT and RHT neurons ascended through the contralateral brain stem to the supraoptic decussation (SOD) in the lateral hypothalamus; 57% of them then crossed the midline to reach the ipsilateral hypothalamus. Collateral projections were found in the superior colliculus, substantia nigra, red nucleus, anterior pretectal nucleus, and in the lateral, perifornical, dorsomedial, suprachiasmatic, and supraoptic hypothalamic nuclei. Additional projections (which have not been described previously for SHT neurons) were found rostral to the hypothalamus in the caudate-putamen, globus pallidus, and substantia innominata. The findings that nonnociceptive signals reach the hypothalamus primarily through the direct THT route, whereas nociceptive signals reach the hypothalamus through both the direct THT and the indirect RHT routes suggest that highly prioritized painful signals are transferred in parallel channels to ensure that this critical information reaches the hypothalamus, a brain area that regulates homeostasis and other humoral responses required for the survival of the organism.

Response Properties and Organization of Nociceptive Neurons in Area 1 of Monkey Primary Somatosensory Cortex

Abstract: The organization and response properties of nociceptive neurons in area 1 of the primary somatosensory cortex (SI) of anesthetized monkeys were examined. The receptive fields of nociceptive neurons...

Transduction Sites of Vagal Mechanoreceptors in the Guinea Pig Esophagus

Abstract: Extrinsic afferent neurons play an essential role in both sensation and reflex control of visceral organs, but their specialized morphological peripheral endings have never been functionally identified. Extracellular recordings were made from fine nerve trunks running between the vagus nerve and esophagus of the guinea pig. Mechanoreceptors, which responded to esophageal distension, fired spontaneously, had low thresholds to circumferential stretch, and were slowly adapting. Calibrated von Frey hairs (0.12 mN) were used to probe the serosal surface at 100–200 sites, which were mapped on a video image of the live preparation. Each stretch-sensitive unit had one to three highly localized receptive fields (“hot spots”), which were marked with Indian ink applied on the tip of the von Frey hair. Recorded nerve trunks were then filled anterogradely, using biotinamide in an artificial intracellular solution. Receptive fields were consistently associated with intraganglionic laminar endings (IGLEs) in myenteric ganglia, but not with other filled neuronal structures. The average distance of receptive fields to IGLEs was 73 ± 14 μm (24 receptive fields, from 12 units; n = 5), compared to 374 ± 17 μm for 240 randomly generated sites ( n = 5; p < 0.001). After maintained probing on a single receptive field, spontaneous discharge of units was inhibited, as were responses to distension. During adapted discharge to maintained distension, interspike intervals were distributed in a narrow range. This indicates that multiple receptive fields interact to encode mechanical distortion in a graded manner. IGLEs are specialized transduction sites of mechanosensitive vagal afferent neurons in the guinea pig esophagus.

Interspike Intervals, Receptive Fields, and Information Encoding in Primary Visual Cortex

Abstract: In the primate primary visual cortex (V1), the significance of individual action potentials has been difficult to determine, particularly in light of the considerable trial-to-trial variability of responses to visual stimuli. We show here that the information conveyed by an action potential depends on the duration of the immediately preceding interspike interval (ISI). The interspike intervals can be grouped into several different classes on the basis of reproducible features in the interspike interval histograms. Spikes in different classes bear different relationships to the visual stimulus, both qualitatively (in terms of the average stimulus preceding each spike) and quantitatively (in terms of the amount of information encoded per spike and per second). Spikes preceded by very short intervals (3 msec or less) convey information most efficiently and contribute disproportionately to the overall receptive-field properties of the neuron. Overall, V1 neurons can transmit between 5 and 30 bits of information per second in response to rapidly varying, pseudorandom stimuli, with an efficiency of approximately 25%. Although some (but not all) of our results would be expected from neurons that use a firing-rate code to transmit information, the evidence suggests that visual neurons are well equipped to decode stimulus-related information on the basis of relative spike timing and ISI duration.

Identification and localisation of auditory areas in guinea pig cortex

Abstract: The organisation of guinea pig auditory cortex was studied by combining histological methods with microelectrode mapping. This allowed the location of seven auditory areas to be determined in relation to the visual and primary somatosensory areas. The auditory areas were identified by single-unit recordings and their borders defined by evoked potential mapping. The visual areas were identified by their relatively high densities of myelinated fibres, while the primary somatosensory cortex was identified by its characteristic barrels of high cytochrome oxidase (CYO) activity in layer IV. The auditory region had moderate levels of CYO and myelin staining. When staining was optimal, there was a clear edge to the moderate CYO activity, which apparently corresponds to the dorsal border of the primary auditory area (AI) and the other core field that lies dorsocaudal to it (DC). Thus the primary somatosensory area and the visual and auditory regions were separated from each other by a region with lower levels of CYO and myelin staining. The ventral borders of AI and DC could not be determined histologically as there were no sharp transitions in the levels of CYO or myelin staining. The two core areas were partially surrounded by belt areas. The dorsorostral belt and most of the belt around DC responded more strongly to broad-band stimuli than pure tones, while the ventrorostral belt, small field and a belt zone ventral to the rostral part of DC responded better to pure tones. Units in the small field (S) typically had higher thresholds and broader tuning to pure tones than AI, while units in the ventrorostral belt typically had longer onset latencies and gave more sustained responses than units in AI.

Texture segregation in the human visual cortex: A functional MRI study.

Abstract: The segregation of visual scenes based on contour information is a fundamental process of early vision. Contours can be defined by simple cues, such as luminance, as well as by more complex cues, such as texture. Single-cell recording studies in monkeys suggest that the neural processing of complex contours starts as early as primary visual cortex. Additionally, lesion studies in monkeys indicate an important contribution of higher order areas to these processes. Using functional MRI, we have investigated the level at which neural correlates of texture segregation can be found in the human visual cortex. Activity evoked by line textures, with and without texture-defined boundaries, was compared in five healthy subjects. Areas V1, V2/VP, V4, TEO, and V3A were activated by both kinds of line textures as compared with blank presentations. Textures with boundaries forming a checkerboard pattern, relative to uniform textures, evoked significantly more activity in areas V4, TEO, less reliably in V3A, but not in V1 or V2/VP. These results provide evidence that higher order areas with large receptive fields play an important role in the segregation of visual scenes based on texture-defined boundaries.

Wide-field, motion-sensitive neurons and matched filters for optic flow fields

Abstract: The receptive field organization of a class of visual interneurons in the fly brain (vertical system, or VS neurons) shows a striking similarity to certain self-motion-induced optic flow fields. The present study compares the measured motion sensitivities of the VS neurons (Krapp et al. 1998) to a matched filter model for optic flow fields generated by rotation or translation. The model minimizes the variance of the filter output caused by noise and distance variability between different scenes. To that end, prior knowledge about distance and self-motion statistics is incorporated in the form of a “world model”. We show that a special case of the matched filter model is able to predict the local motion sensitivities observed in some VS neurons. This suggests that their receptive field organization enables the VS neurons to maintain a consistent output when the same type of self-motion occurs in different situations.

Fundamental GABAergic amacrine cell circuitries in the retina: nested feedback, concatenated inhibition, and axosomatic synapses.

Abstract: Presynaptic gamma-aminobutyrate-immunoreactive (GABA+) profiles were mapped in the cyprinid retina with overlay microscopy: a fusion of electron and optical imaging affording high-contrast ultrastructural and immunocytochemical visualization. GABA+ synapses, deriving primarily from amacrine cells (ACs), compose 92% of conventional synapses and 98% of the input to bipolar cells (BCs) in the inner plexiform layer. GABA+ AC synapses, the sign-inverting elements of signal processing, are deployed in micronetworks and distinctive synaptic source/target topologies. Nested feedback micronetworks are formed by three types of links (BC --> AC, reciprocal BC AC synapses) arranged as nested BC [AC --> AC] loops. Circuits using nested feedback can possess better temporal performance than those using simple reciprocal feedback loops. Concatenated GABA+ micronetworks of AC --> AC and AC --> AC --> AC chains are common and must be key elements for lateral spatial, temporal, and spectral signal processing. Concatenated inhibitions may represent exceptionally stable, low-gain, sign-conserving devices for receptive field construction. Some chain elements are GABA immunonegative (GABA-) and are, thus, likely glycinergic synapses. GABA+ synaptic baskets target the somas of certain GABA+ and GABA- cells, resembling cortical axosomatic synapses. Finally, all myelinated intraretinal profiles are GABA+, suggesting that some efferent systems are sources of GABAergic inhibition in the cyprinid retina and may comprise all axosomatic synapses. These micronetworks are likely the fundamental elements for receptive field shaping in the inner plexiform layer, although few receptive field models incorporate them as functional components. Conversely, simple feedback and feedforward synapses may often be chimeras: the result of an incomplete view of synaptic topology.

Suppression outside the classical cortical receptive field.

Abstract: The important visual stimulus parameters for a given cell are defined by the classical receptive field (CRF). However, cells are also influenced by visual stimuli presented in areas surrounding the CRF. The experiments described here were conducted to determine the incidence and nature of CRF surround influences in the primary visual cortex. From extracellular recordings in the cat's striate cortex, we find that for over half of the cells investigated (56%, 153/271), the effect of stimulation in the surround of the CRF is to suppress the neuron's activity by at least 10% compared to the response to a grating presented within the CRF alone. For the remainder of the cells, the interactions were minimal and a few were of a facilitatory nature. In this paper, we focus on the suppressive interactions. Simple and complex cell types exhibit equal incidences of surround suppression. Suppression is observed for cells in all layers, and its degree is strongly correlated between the two eyes for binocular neurons. These results show that surround suppression is a prevalent form of inhibition and may play an important role in visual processing.

Bilateral receptive field neurons and callosal connections in the somatosensory cortex.

Abstract: Earlier studies recording single neuronal activity with bilateral receptive fields in the primary somatosensory cortex of monkeys and cats agreed that the bilateral receptive fields were related exclusively to the body midline and that the ipsilateral information reaches the cortex via callosal connections since they are dense in the cortical region representing the midline structures of the body while practically absent in the regions representing the distal extremities. We recently found a substantial number of neurons with bilateral receptive fields on hand digits, shoulders–arms or legs–feet in the caudalmost part (areas 2 and 5) of the postcentral gyrus in awake Japanese monkeys ( Macaca fuscata ). I review these results, discuss the functional implications of this bilateral representation in the postcentral somatosensory cortex from a behavioural standpoint and give a new interpretation to the midline fusion theory.

Local Disparity Not Perceived Depth Is Signaled by Binocular Neurons in Cortical Area V1 of the Macaque

Abstract: Binocular neurons that are closely related to depth perception should respond selectively for stimuli eliciting an appropriate depth sensation. To separate perceived depth from local disparity within the receptive field, sinusoidal luminance gratings were presented within a circular aperture. The disparity of the aperture was coupled to that of the grating, thereby rendering unambiguous the psychophysical matching between repeating cycles of the grating. In cases in which the stimulus disparity differs by one horizontal period of the grating, the portion of the grating that locally covers a receptive field is binocularly identical, but the depth sensation is very different because of the aperture. For 117 disparity-selective V1 neurons tested in two monkeys, the overwhelming majority responded equally well to configurations that were locally identical but led to different perceptions of depth. Because the psychophysical sensation is not reflected in the firing rate of V1 neurons, the signals that make stereo matches explicit are most likely elaborated in extrastriate cortex.

Striate cortex increases contrast gain of macaque LGN neurons.

Abstract: Recurrent projections comprise a universal feature of cerebral organization. Here, we show that the corticofugal projections from the striate cortex (VI) to the lateral geniculate nucleus (LGN) robustly and multiplicatively enhance the responses of parvocellular neurons, stimulated by gratings restricted to the classical receptive field and modulated in luminance, by over two-fold in a contrast-independent manner at all but the lowest contrasts. In the equiluminant plane, wherein stimuli are modulated in chromaticity with luminance held constant, such enhancement is strongly contrast dependent. These projections also robustly enhance the responses of magnocellular neurons but contrast independently only at high contrasts. Thus, these results have broad functional significance at both network and neuronal levels by providing the experimental basis and quantitative constraints for a wide range of models on recurrent projections and the control of contrast gain.

Neural Responses in the Retinotopic Representation of the Blind Spot in the Macaque V1 to Stimuli for Perceptual Filling-In

Abstract: When visual stimuli that cover the entire blind spot are presented monocularly, the color and brightness of the surrounding field are seen within the blind spot, although it receives no retinal input. Important questions about such perceptual filling-in are whether neurons in the visual system representing visual field locations within the blind spot are activated when filling-in occurs and, if so, what the properties of these neurons are. To address these questions, we recorded the activities of single neurons in the primary visual cortex (V1) of the awake monkey. We first identified the area of V1 representing the region of the blind spot and then assessed neural responses to stationary visual stimuli of various size. We found that there are neurons in layer 4 and deeper laminae, particularly layer 6, that respond to large stimuli covering the blind spot which induces perceptual filling-in. Most of these neurons had very large binocular receptive fields that extended outside the blind spot. These neurons also preferred relatively large stimuli and exhibited color selectivity. These results indicate that when a large uniform surface is presented on the blind spot, neurons at the V1 region representing the blind spot transmit signals essential for filling-in that inform of the presence of a large surface as well as the absence of smaller stimuli at the blind spot.

Diverse receptive fields in the lateral geniculate nucleus during thalamocortical development.

Abstract: Most models of thalamocortical development in the visual system assume a homogeneous population of thalamic inputs to the cortex, each with concentric on- or off-center receptive fields. To test this, we made high-resolution spatial maps of receptive fields in the developing ferret lateral geniculate nucleus (LGN). Developing receptive fields (RFs), had a variety of shapes: some concentric, others elongated (like adult cortical receptive fields) and some with 'hot spots' of sensitivity. These receptive fields seemed to arise from convergence of multiple retinal afferents onto LGN neurons. We present a Hebbian model whereby imprecise retinogeniculate connections help refine geniculocortical connections, sharpening both thalamocortical topography and perhaps orientation selectivity.

Endogenous dopaminergic regulation of horizontal cell coupling in the mammalian retina

Abstract: Horizontal cells in an isolated wholemount preparation of the mouse retina were injected with Lucifer yellow and neurobiotin to characterize both the pattern of gap junctional connectivity and its regulation by dopamine. The injected horizontal cells had a uniform morphology of a round cell body, a compact dendritic tree, and an axon, which could sometimes be traced to an expansive terminal system. The dendro-dendritic gap junctions between neighboring cells mediated both weak Lucifer yellow dye coupling and strong neurobiotin tracer coupling. The extent of the tracer coupling was decreased by either exogenous dopamine (100 mu M) or cyclic adenosine monophosphate (cAMP) analogs and was significantly increased by the D(1) antagonist SCH 23390 (1D mu M). These results provide the first evidence in the mammalian retina that the gap junctions between horizontal cells are endogenously regulated by dopamine, which acts through D(1) receptors to increase the intracellular cAMP. It has been proposed that the gap junctional coupling between horizontal cells is mediated by connexin 32 (Cx32), but the pattern and dopaminergic regulation of horizontal cell coupling were unaffected in Cx32-knockout mice, ruling out the possible involvement of Cx32. Every tracer-coupled horizontal cell showed calbindin immunoreactivity, and vice versa, providing strong evidence that the horizontal cells in the mouse retina comprise a single cell type. Like the axonless horizontal cells in other mammalian retinas, the axon-bearing horizontal cells in the mouse retina are coupled by gap junctions that are permeable to Lucifer yellow and dopamine sensitive, suggesting that the mouse horizontal cells have hybrid properties to compensate for the absence of axonless horizontal cells. (C) 2000 Wiley-Liss, Inc.