There are two subclasses of alpha cell in the mammalian retina, which are morphologically identical in plain view but have opposite responses to a luminance change: one is ON center and the other is OFF center. Recent studies have shown that the neural circuitries, which underlie light responses in such ON- and OFF-ganglion cell pairs, are not mirror symmetric with respect to the ON and OFF pathways (Pang et al., 2003; Zaghloul et al., 2003; Murphy & Rieke, 2006). This study examines alpha-cell homologues in the mouse retina and elucidates the synaptic mechanisms that generate their light responses. Morphological analysis of recorded cells revealed three subclasses that were essentially identical in plan view but had distinct vertical stratification levels. We refer to these cells as the sustained ON (ON-S), sustained OFF (OFF-S), and transient OFF (OFF-T) cells (Murphy & Rieke, 2006; Margolis & Detwiler, 2007). Both ON-S and OFF-S cells were largely driven through the ON pathway via changes in excitatory and inhibitory inputs, respectively. Light responses of OFF-T cells were driven by transient changes in excitatory and inhibitory inputs. Light responses of OFF-S cells were also measured in connexin 36 knockout mice in order to dissect glycinergic input arising from AII amacrine cells. At photopic/mesopic intensities, peak glycinergic input to OFF-S cells in the connexin 36 knockout mouse was reduced by ~85% compared to OFF-S cells in the wild-type retina. This is consistent with the idea that AII cells receive their input from ON-cone bipolar cells through gap junctions and in turn provide glycinergic inhibition to OFF-S cells.

The visual abilities of sharks show substantial interspecific variability. In addition, sharks may change their habitat and feeding strategy throughout life. As the eyes of sharks continue to grow throughout the animal’s lifetime, ontogenetic variability in visual ability may also occur. The topographic analysis of the photoreceptor and ganglion cell distributions can identify visual specializations and assess changes in visual abilities that may occur concurrently with eye growth. This study examines an ontogenetic series of whole-mounted retinas in two elasmobranch species, the sandbar shark, Carcharhinus plumbeus, and the shortspine spurdog, Squalus mitsukurii, to identify regional specializations mediating zones for improved spatial resolution. The study examines retinal morphology and presents data on summation ratios between photoreceptor and ganglion cell layers, anatomically determined peak spatial resolving power, and the angular extent of the visual field. Peak densities of photoreceptors and ganglion cells occur in similar retinal locations. The topographic distribution of neurons in the ganglion cell layer does not differ substantially with eye growth. However, predicted peak spatial resolution increases with eye growth from 4.3 to 8.9 cycles/deg in C. plumbeus and from 5.7 to 7.2 cycles/deg in S. mitsukurii. The topographic distribution of different-sized ganglion cells is also mapped in C. plumbeus, and a population of large ganglion cells (soma area 120–350 μm2) form a narrow horizontal streak across the retinal meridian, while the spatial distribution of ordinary-sized ganglion cells (soma area 30–120 μm2) forms an area in the central retina. Species-specific retinal specializations highlight differences in visually mediated behaviors and foraging strategies between C. plumbeus and S. mitsukurii.

Historically, the principal function of vision has been to provide the information needed to support action. Visually mediated actions rely on three systems: the gaze system responsible for locating and fixating task-relevant objects, the motor system of the limbs to carry out the task, and the visual system to supply information to the other two. All three systems are under the control of a fourth system, the schema system, which specifies the current task and plans the overall sequence of actions. These four systems have separate but interconnected cortical representations. The way these systems interact in time and space is discussed here in relation to two studies of the gaze changes and manipulations made during two ordinary food preparation tasks. The main conclusions are that complex action sequences consist of a succession of individual object-related actions, each of which typically involve a turn toward the object (if needed), followed by fixation and finally manipulation monitored by vision. Gaze often moves on to the next object just before manipulation is complete. Task-irrelevant objects are hardly ever fixated, implying that the control of fixation comes principally from top-down instructions from the schema system, not bottom-up salience. Single fixations have identifiable functions (locating, directing, guiding, and checking) related to the action to be taken. Several variants of the basic object-related action scheme are discussed, including single-action events in ball sports involving only one anticipatory gaze shift, continuous production loops in text and music reading, and storage–action alternation in copying tasks such as portrait sketching.

Bistratified (BS) ganglion cells have long been recognized in vertebrate retina. Thirty years ago, it became clear that bistratification allows the integration of ON and OFF retinal pathways to produce contrast-independent responses in ganglion cells. Best studied is the type 1 bistratified (BS1) ganglion cell of rabbit retina, the physiologically well-characterized ON-OFF directionally selective (DS) ganglion cell, which is co-stratified with the two types of starburst amacrine (SA) cells in sublaminae a and b of the inner plexiform layer (IPL). DS responses have recently been documented in the latter. In this report, BS1 cells are further studied and are used as “fiducials” to characterize a second type of BS ganglion cell. An example of a possible third type is shown to be distinct from examples of BS1 and BS2 cells. All three have two distinct, narrowly stratified arborizations, one in sublamina a and one in sublamina b. All have similar dimensions, except for their dendritic trees, differing also in branching pattern. BS1 cells have compact, regular, highly branched trees; BS2 cells have significantly larger, more sparsely branched, irregular, radiate trees; the proposed BS3 type is intermediate in field size, and its branching pattern is different from the first two. BS2 and BS3 cells are co-stratified, branching nearer to the margins of the IPL, out of range of SA cells. In a previous report by others, illustrating the morphology of intracellularly stained ganglion cells, one example each of both “orientation-selective” ganglion cells and “uniformity detectors” resembles the BS2 cell. A rationale is presented for correlating BS2 cells with uniformity detectors.

We investigated the effect of auditory–visual sensory integration on visual tasks that were predominantly dependent on parvocellular processing. These tasks were (i) detecting metacontrast-masked targets and (ii) discriminating orientation differences between high spatial frequency Gabor patch stimuli. Sounds that contained no information relevant to either task were presented before, synchronized with, or after the visual targets, and the results were compared to conditions with no sound. Both tasks used a two-alternative forced choice technique. For detecting metacontrast-masked targets, one interval contained the visual target and both (or neither) intervals contained a sound. Sound–target synchrony within 50 ms lowered luminance thresholds for detecting the presence of a target compared to when no sound occurred or when sound onset preceded target onset. Threshold angles for discriminating the orientation of a Gabor patch consistently increased in the presence of a sound. These results are compatible with sound-induced activity in the parvocellular visual pathway increasing the visibility of flashed targets and hindering orientation discrimination.

The pond turtle (Trachemys scripta elegans) exhibits a notably sluggish pupillary light reflex (PLR), with pupil constriction developing over several minutes following light onset. In the present study, we examined the dynamics of the efferent branch of the reflex in vitro using preparations consisting of either the isolated head or the enucleated eye. Stimulation of the oculomotor nerve (nIII) using 100-Hz current trains resulted in a maximal pupil constriction of 17.4% compared to 27.1% observed in the intact animal in response to light. When current amplitude was systematically increased from 1 to 400 μA, mean response latency decreased from 64 to 45 ms, but this change was not statistically significant. Hill equations fitted to these responses indicated a current threshold of 3.8 μA. Stimulation using single pulses evoked a smaller constriction (3.8%) with response latencies and threshold similar to that obtained using train stimulation. The response evoked by postganglionic stimulation of the ciliary nerve using 100-Hz trains was largely indistinguishable from that of train stimulation of nIII. However, application of single-pulse stimulation postganglionically resulted in smaller pupil constriction at all current levels relative to that of nIII stimulation, suggesting that there is amplification of efferent drive at the ganglion. Time constants for constrictions ranged from 88 to 154 ms with relaxations occurring more slowly at 174–361 ms. These values for timing from in vitro are much faster than the time constant 1.66 min obtained for the light response in the intact animal. The rapid dynamics of pupil constriction observed here suggest that the slow PLR of the turtle observed in vivo is not due to limitations of the efferent pathway. Rather, the sluggish response probably results from photoreceptive mechanisms or central processing.

Neurons in visual cortical area V1 typically respond well to lines or edges of specific orientations. There have been many studies investigating how the responses of these neurons to an oriented edge are affected by changes in luminance contrast. However, in natural images, edges vary not only in contrast but also in the degree of blur, both because of changes in focus and also because shadows are not sharp. The effect of blur on the response dynamics of visual cortical neurons has not been explored. We presented luminance-defined single edges in the receptive fields of parafoveal (1–6 deg eccentric) V1 neurons of two macaque monkeys trained to fixate a spot of light. We varied the width of the blurred region of the edge stimuli up to 0.36 deg of visual angle. Even though the neurons responded robustly to stimuli that only contained high spatial frequencies and 0.36 deg is much larger than the limits of acuity at this eccentricity, changing the degree of blur had minimal effect on the responses of these neurons to the edge. Primates need to measure blur at the fovea to evaluate image quality and control accommodation, but this might only involve a specialist subpopulation of neurons. If visual cortical neurons in general responded differently to sharp and blurred stimuli, then this could provide a cue for form perception, for example, by helping to disambiguate the luminance edges created by real objects from those created by shadows. On the other hand, it might be important to avoid the distraction of changing blur as objects move in and out of the plane of fixation. Our results support the latter hypothesis: the responses of parafoveal V1 neurons are largely unaffected by changes in blur over a wide range.

The visual system of birds includes an efferent projection from a visual area, the isthmo-optic nucleus in the midbrain, back to the retina. Using a combination of anterograde labeling of efferent fibers, reconstruction of dye-filled neurons, NADPH-diaphorase staining, and transmission electron microscopy, we have examined the distribution of efferent fibers and their synaptic structures in the chicken retina. We show that efferent fibers terminate strictly within the ventral retina. In two completely mapped retinas, only 2 fibers from a total of 15,359 terminated in the dorsal retina. The major synapse made by each efferent fiber is with a single efferent target amacrine cell (TC). This synapse consists of 5–25 boutons of 2 μm diameter, each with multiple active zones, pressed into the TC soma or synapsing with a basketwork of rudimentary TC dendrites in the inner nuclear layer (INL). This basketwork, which is sheathed by Muller cell processes, defines a private neuropil in the INL within which TCs were also seen to receive input from retinal neurons. In addition to the major synapse, efferent fibers typically produce several very thin processes that terminate nearby in single small boutons and for which the soma of a local amacrine cell is one of the likely postsynaptic partners. A minority of efferent fibers also give rise to a thicker process, terminating in a strongly diaphorase-positive ball about 5 μm in diameter.

Visual cue integration strategies are known to depend on cue reliability and how rapidly the visual system processes incoming information. We investigated whether these strategies also depend on differences in the information demands for different natural tasks. Using two common goal-oriented tasks, prehension and object placement, we determined whether monocular and binocular information influence estimates of three-dimensional (3D) orientation differently depending on task demands. Both tasks rely on accurate 3D orientation estimates, but 3D position is potentially more important for grasping. Subjects placed an object on or picked up a disc in a virtual environment. On some trials, the monocular cues (aspect ratio and texture compression) and binocular cues (e.g., binocular disparity) suggested slightly different 3D orientations for the disc; these conflicts either were present upon initial stimulus presentation or were introduced after movement initiation, which allowed us to quantify how information from the cues accumulated over time. We analyzed the time-varying orientations of subjects’ fingers in the grasping task and those of the object in the object placement task to quantify how different visual cues influenced motor control. In the first experiment, different subjects performed each task, and those performing the grasping task relied on binocular information more when orienting their hands than those performing the object placement task. When subjects in the second experiment performed both tasks in interleaved sessions, binocular cues were still more influential during grasping than object placement, and the different cue integration strategies observed for each task in isolation were maintained. In both experiments, the temporal analyses showed that subjects processed binocular information faster than monocular information, but task demands did not affect the time course of cue processing. How one uses visual cues for motor control depends on the task being performed, although how quickly the information is processed appears to be task invariant.

The aim of this study was to explore the plasticity of fixation in patients with central vision loss. Most of these patients use preferred retinal loci (PRLs) in the healthy eccentric part of the retina to fixate, but fixation stability and retinal location are not always optimal for best visual performance. This study examined whether fixation stability and a new PRL location can be trained and whether these changes in ocular motor control transfer into better reading performance. Six patients with age-related macular degeneration participated in the study. Fixation stability measurements, microperimetry, and auditory biofeedback training were performed with the MP-1 microperimeter. The auditory biofeedback was used during five 1-h long training sessions to improve fixation and relocate the PRL. Fixation location and stability were recorded while viewing four different targets: a cross, a letter, a word, and a nine-cycle radial grating. Visual acuity was assessed with the Early Treatment Diabetic Retinopathy Study (ETDRS) chart and reading performance with the MNRead test. The results showed that all patients developed a new PRL in an optimal location for reading, and they were able to use it consistently while viewing different targets. Fixation stability improved 53% after training. Learning transferred to the old PRL even though fixation stability at this location was not trained. All these improvements in ocular motor control translated into better reading performance: reading speed improved 38% and reading acuity and critical print size gained two lines. We conclude that the ability of the ocular motor system to fixate is flexible in patients with central vision loss: a new PRL can be trained, fixation stability can be improved, and learning transfers to an untrained location. These gains in ocular motor control result in better visual performance. This property can be successfully used to optimize the residual vision of patients with central vision loss.

Monocular deprivation early in development produces considerable change in the organization of connections within the central mammalian visual system. In the dorsal lateral geniculate nucleus, the soma, dendrites, and axon terminal fields of deprived cells become considerably smaller than nondeprived counterparts. We have examined the possibility that subcellular events enabling structural modification of deprived neurons include modification of proteins comprising the cytoskeleton. We examined the integrity of the cytoskeleton by measuring the response of a subset of its proteins to varying durations of monocular deprivation. Loss of all three neurofilament subunits (light, medium, and heavy) within deprived layers was observed to parallel changes in neuron gross structure. Monocular deprivation initiated beyond early life produced neither a change in structure nor a loss of neurofilament labeling.

We studied the correlation between the spatial frequency of complex stimuli and neuronal activity in the monkey inferotemporal (IT) cortex while performing a task that required visual recognition. Single-cell activity was recorded from the right IT cortex. The frequency components of the images used as stimuli were analyzed by using a fast Fourier transform, and a modulus was obtained for 40 spatial frequency ranges from 0.3 to 11.1 cycles/deg. We recorded 82 cells showing statistically significant responses (analysis of variance, P < 0.05) to at least one of the images used as a stimulus. Seventy-eight percent of these cells (n = 64) showed significant responses to at least three images, and in two thirds of them (n = 42), we found a statistically significant correlation (P < 0.05) between cell response and the modulus amplitude of at least one frequency range present in the images. Our results suggest that information about spatial frequency of the visual images is present in the IT cortex.

The retina displays numerous processes that follow a circadian rhythm. These processes are coordinated through the direct action of light on photoreceptive molecules and, in the absence of light, through autocrine/paracrine actions of extracellular neuromodulators. We previously described the expression of the genes encoding the secreted heparin-binding growth factors, midkine-a (mdka) and midkine-b (mdkb), in the retina of the zebrafish. Here, we provide evidence that the expression of mdka and mdkb follows a daily rhythm, which is independent of the presence or absence of light, and we propose that the expression of mdka is regulated by the circadian clock. Both qualitative and quantitative measures show that for mdka, the levels of mRNA and protein decrease during the night and increase during the subjective day. Qualitative measures show that the expression of mdkb increases during the second half of the subjective night and decreases during the second half of the subjective day. Within horizontal cells, the two midkine paralogs show asynchronous circadian regulation. Though intensely studied in the contexts of physiology and disease, this is the first study to provide evidence for the circadian regulation of midkines in the vertebrate nervous system.

The purpose of the present study was to compare standard multifocal visual evoked potential (mfVEP) pattern-reversal responses with those produced by pattern-onset, pattern-offset, and pulsed pattern stimuli. mfVEP recordings were obtained from five normal subjects using VERIS and a 4-electrode array. The standard reversal stimulus had 215 m-sequence steps (7.5-min duration). Pattern-onset and -offset responses were evaluated using sequences that all had 32 frames per m-step and 210 total steps (7.5 min); but the duration of the contrast step varied so that it was 1, 2, 4, 8, 12, or 16 of the 32 frames. The same series was also inverted so that adapting contrast was high and the stimulus step began with a contrast decrement. The effect of temporal sparseness was studied with positive contrast pulses (two-frame duration) within 16, 20, 24, 28, or 32 frames per m-step (all had 211 total steps). Four stimulus locations were isolated to study the effect of spatial sparseness. Standard mfVEP reversal responses were virtually identical to onset responses throughout the field, but ~3.5 times smaller. Responses to pattern onset were about twice as large as those for offset, especially in the lower hemifield, irrespective of adapting contrast level. Offset responses exhibited a different waveform compared with reversal, onset, or brief pulse responses. Though temporally sparse pattern-pulse responses were ~3.5 times larger than standard reversal responses, there was no improvement in signal-to-noise ratio (SNR). However, spatial isolation increased SNR by 22% for reversal responses and by 62% for temporally sparse pulses. Temporally sparse pattern-pulse stimuli do not improve mfVEP SNR unless they are also spatially sparse, suggesting that lateral inhibitory mechanisms have a greater impact than temporal contrast gain control mechanisms. The mfVEP response depends on the polarity of a contrast step, irrespective of the state of contrast adaptation.

Despite the extensive investigation of binocular and stereoscopic vision, relatively little is known about its importance in natural visually guided behavior. In this paper, we explored the role of binocular vision when walking over and around obstacles. We monitored eye position during the task as an indicator of the difference between monocular and binocular performances. We found that binocular vision clearly facilitates walking performance. Walkers were slowed by about 10% in monocular vision and raised their foot higher when stepping over obstacles. Although the location and sequence of the fixations did not change in monocular vision, the timing of the fixations relative to the actions was different. Subjects spent proportionately more time fixating the obstacles and fixated longer while guiding foot placement near an obstacle. The data are consistent with greater uncertainty in monocular vision, leading to a greater reliance on feedback in the control of the movements.

Theories of efficient sensory processing have considered the regularities of image properties due to the structure of the environment in order to explain properties of neuronal representations of the visual world. The regularities imposed on the input to the visual system due to the regularities of the active selection process mediated by the voluntary movements of the eyes have been considered to a much lesser degree. This is surprising, given that the active nature of vision is well established. The present article investigates statistics of image features at the center of gaze of human subjects navigating through a virtual environment and avoiding and approaching different objects. The analysis shows that contrast can be significantly higher or lower at fixation location compared to random locations, depending on whether subjects avoid or approach targets. Similarly, significant differences in the distribution of responses of model simple and complex cells between horizontal and vertical orientations are found over timescales of tens of seconds. By clustering the model simple cell responses, it is established that gaze was directed toward three distinct features of intermediate complexity the vast majority of time. Thus, this study demonstrates and quantifies how the visuomotor tasks of approaching and avoiding objects during navigation determine feature statistics of the input to the visual system through the combined influence on body and eye movements.

The purpose of this study was to test whether color–motion correlations carried by a pure color difference (S-cone component only) can be used to improve global motion extraction. We also examined the neural markers of color–motion correlation processing in event-related potentials. Color and motion information was dissociated using a two-colored random dot kinematogram, wherein coherent motion and motion noise differed from each other only in their S-cone component, with spatial and temporal parameters set so that global motion processing relied solely on a constant L-M component. Hence, when color and the local motion direction are correlated, more efficient segregation of coherent motion can only be brought about by the S-cone difference, and crucially, this S-cone component does not provide any effective input to a global motion mechanism but only changes the color appearance of the moving dots. The color contrasts (vector length in the S vs. L-M plane) of both the dots carrying coherent motion and the dots moving randomly were fixed at motion discrimination threshold to ensure equal effectiveness for motion extraction. In the behavioral experiment, participants were asked to discriminate between coherent and random motion, and d′ was determined for three different conditions: uncorrelated, uncued correlated, and cued correlated. In the electroencephalographic experiment, participants discriminated direction of motion for uncued correlated and cued correlated conditions. Color–motion correlations were found to improve performance. Cueing a specific color also modulated the N1 component of the event-related potential, with sources in visual area middle temporal. We conclude that S-cone signals “invisible” to the motion system can influence the analysis by direction-selective motion mechanisms through grouping of local motion signals by color. This grouping mechanism must precede motion processing and is likely to be under attentional control.

Recently, we showed that contrast sensitivity for color and high–spatial frequency luminance stimuli is enhanced during smooth pursuit eye movements (Schütz et al., 2008). In this study, we investigated the enhancement over a wide range of temporal and spatial frequencies. In Experiment 1, we measured the temporal impulse response function (TIRF) for colored stimuli. The TIRF for pursuit and fixation differed mostly with respect to the gain but not with respect to the natural temporal frequency. Hence, the sensitivity enhancement seems to be rather independent of the temporal frequency of the stimuli. In Experiment 2, we measured the spatial contrast sensitivity function for luminance-defined Gabor patches with spatial frequencies ranging from 0.2 to 7 cpd. We found a sensitivity improvement during pursuit for spatial frequencies above 2–3 cpd. Between 0.5 and 3 cpd, sensitivity was impaired by smooth pursuit eye movements, but no consistent difference was observed below 0.5 cpd. The results of both experiments are consistent with an increased contrast gain of the parvocellular retinogeniculate pathway.