Binocular Region Research Articles

M2 receptors are required for spatiotemporal sequence learning in mouse primary visual cortex.

Acetylcholine is a neurotransmitter that plays a variety of roles in the central nervous system. It was previously shown that blocking muscarinic receptors with a nonselective antagonist prevents a form of experience-dependent plasticity termed "spatiotemporal sequence learning" in the mouse primary visual cortex (V1). Muscarinic signaling is a complex process involving the combined activities of five different G protein-coupled receptors, M1-M5, all of which are expressed in the murine brain but differ from each other functionally and in anatomical localization. Here we present electrophysiological evidence that M2, but not M1, receptors are required for spatiotemporal sequence learning in mouse V1. We show in male mice that M2 is highly expressed in the neuropil in V1, especially in thalamorecipient layer 4, and colocalizes with the soma in a subset of somatostatin-expressing neurons in deep layers. We also show that expression of M2 receptors is higher in the monocular region of V1 than it is in the binocular region but that the amount of experience-dependent sequence potentiation is similar in both regions and that blocking muscarinic signaling after visual stimulation does not prevent plasticity. This work establishes a new functional role for M2-type receptors in processing temporal information and demonstrates that monocular circuits are modified by experience in a manner similar to binocular circuits.NEW & NOTEWORTHY Muscarinic acetylcholine receptors are required for multiple forms of plasticity in the brain and support perceptual functions, but the precise role of the five subtypes (M1-M5) are unclear. Here we show that the M2 receptor is specifically required to encode experience-dependent representations of spatiotemporal relationships in both monocular and binocular regions of mouse V1. This work identifies a novel functional role for M2 receptors in coding temporal information into cortical circuits.

The Development of Receptive Field Tuning Properties in Mouse Binocular Primary Visual Cortex.

The mouse primary visual cortex is a model system for understanding the relationship between cortical structure, function, and behavior (Seabrook et al., 2017; Chaplin and Margrie, 2020; Hooks and Chen, 2020; Saleem, 2020; Flossmann and Rochefort, 2021). Binocular neurons in V1 are the cellular basis of binocular vision, which is required for predation (Scholl et al., 2013; Hoy et al., 2016; La Chioma et al., 2020; Berson, 2021; Johnson et al., 2021). The normal development of binocular responses, however, has not been systematically measured. Here, we measure tuning properties of neurons to either eye in awake mice of either sex from eye opening to the closure of the critical period. At eye opening, we find an adult-like fraction of neurons responding to the contralateral-eye stimulation, which are selective for orientation and spatial frequency; few neurons respond to ipsilateral eye, and their tuning is immature. Fraction of ipsilateral-eye responses increases rapidly in the first few days after eye opening and more slowly thereafter, reaching adult levels by critical period closure. Tuning of these responses improves with a similar time course. The development and tuning of binocular responses parallel that of ipsilateral-eye responses. Four days after eye opening, monocular neurons respond to a full range of orientations but become more biased to cardinal orientations. Binocular responses, by contrast, lose their cardinal bias with age. Together, these data provide an in-depth accounting of the development of monocular and binocular responses in the binocular region of mouse V1 using a consistent set of visual stimuli and measurements.SIGNIFICANCE STATEMENT In this manuscript, we present a full accounting of the emergence and refinement of monocular and binocular receptive field tuning properties of thousands of pyramidal neurons in mouse primary visual cortex. Our data reveal new features of monocular and binocular development that revise current models on the emergence of cortical binocularity. Given the recent interest in visually guided behaviors in mice that require binocular vision (e.g., predation), our measures will provide the basis for studies on the emergence of the neural circuitry guiding these behaviors.

Chronic Monocular Deprivation Reveals MMP9-Dependent and -Independent Aspects of Murine Visual System Plasticity.

The deletion of matrix metalloproteinase MMP9 is combined here with chronic monocular deprivation (cMD) to identify the contributions of this proteinase to plasticity in the visual system. Calcium imaging of supragranular neurons of the binocular region of primary visual cortex (V1b) of wild-type mice revealed that cMD initiated at eye opening significantly decreased the strength of deprived-eye visual responses to all stimulus contrasts and spatial frequencies. cMD did not change the selectivity of V1b neurons for the spatial frequency, but orientation selectivity was higher in low spatial frequency-tuned neurons, and orientation and direction selectivity were lower in high spatial frequency-tuned neurons. Constitutive deletion of MMP9 did not impact the stimulus selectivity of V1b neurons, including ocular preference and tuning for spatial frequency, orientation, and direction. However, MMP9−/− mice were completely insensitive to plasticity engaged by cMD, such that the strength of the visual responses evoked by deprived-eye stimulation was maintained across all stimulus contrasts, orientations, directions, and spatial frequencies. Other forms of experience-dependent plasticity, including stimulus selective response potentiation, were normal in MMP9−/− mice. Thus, MMP9 activity is dispensable for many forms of activity-dependent plasticity in the mouse visual system, but is obligatory for the plasticity engaged by cMD.

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Mouse visual cortex contains a region of enhanced spatial resolution

The representation of space in mouse visual cortex was thought to be relatively uniform. Here we reveal, using population receptive-field (pRF) mapping techniques, that mouse visual cortex contains a region in which pRFs are considerably smaller. This region, the “focea,” represents a location in space in front of, and slightly above, the mouse. Using two-photon imaging we show that the smaller pRFs are due to lower scatter of receptive-fields at the focea and an over-representation of binocular regions of space. We show that receptive-fields of single-neurons in areas LM and AL are smaller at the focea and that mice have improved visual resolution in this region of space. Furthermore, freely moving mice make compensatory eye-movements to hold this region in front of them. Our results indicate that mice have spatial biases in their visual processing, a finding that has important implications for the use of the mouse model of vision.

Retinal and Callosal Activity-Dependent Chandelier Cell Elimination Shapes Binocularity in Primary Visual Cortex

In mammals with binocular vision, integration of the left and right visual scene relies on information in the center visual field, which are relayed from each retina in parallel and merge in the primary visual cortex (V1) through the convergence of ipsi- and contralateral geniculocortical inputs as well as transcallosal projections between two visual cortices. The developmental assembly of this binocular circuit, especially the transcallosal pathway, remains incompletely understood. Using genetic methods in mice, we found that several days before eye-opening, retinal and callosal activities drive massive apoptosis of GABAergic chandelier cells (ChCs) in the binocular region of V1. Blockade of ChC elimination resulted in a contralateral eye-dominated V1 and deficient binocular vision. As pre-vision retinal activities convey the left-right organization of the visual field, their regulation of ChC density through the transcallosal pathway may prime a nascent binocular territory for subsequent experience-driven tuning during the post-vision critical period.

Visual acuity in the flying snake, Chrysopelea paradisi.

Visual control during high-speed aerial locomotion requires a visual system adapted for such behaviors. Flying snakes (genus: Chrysopelea) are capable of gliding at speeds up to 11 m s-1 and perform visual assessments before take-off. Determining nuances of control requires a closed-loop experimental system, such as immersive virtual arenas. To characterize vision in the flying snake Chrysopelea paradisi, we used digitally reconstructed models of the head to determine a 3D field of vision. We also used optokinetic drum experiments and compared slowphase optokinetic nystagmus (OKN) speeds to calculate visual acuity and conducted preliminary experiments to determine whether snakes would respond to closed-loop virtual stimuli. Visual characterization showed that C. paradisi likely has a large field of view (308.5 ± 6.5° azimuthal range), with a considerable binocular region (33.0 ± 11.0° azimuthal width) that extends overhead. Their visual systems are broadly tuned and motion-sensitive, with peak OKN response gains of 0.50 ± 0.11 seen at 46.06 ± 11.08 Hz, and a low spatial acuity, with peak gain of 0.92 ± 0.41 seen at 2.89 ± 0.16 cpd (cycles per degree). These characteristics were used to inform settings in an immersive virtual arena, including framerate, brightness, and stimulus size. In turn, the immersive virtual arena was used to reproduce the optokinetic drum experiments. We elicited OKN in open-loop experiments, with a mean gain of 0.21 ± 0.9 seen at 0.019 ± 6x10-5 cpd and 1.79 ± 0.01 Hz. In closed-loop experiments, snakes did not exhibit OKN, but held the image fixed, indicating visual stabilization. These results demonstrate for that C. paradisi responds to visual stimuli in a digital virtual arena. The accessibility and adaptability of the virtual setup make it suitable for future studies of visual control in snakes and other animals in an unconstrained setting.

Mouse visual cortex contains a region of enhanced spatial resolution

The representation of space in mouse visual cortex was initially thought to be relatively uniform, with no strong biases towards any particular region of space. This contrasts with the primate visual cortex with its over-representation of the fovea, placing potential limits on the translation of research in mice to humans. Here we reveal a previously unsuspected organization of the visual cortex of mice that resembles that fovea-centric organization of human visual cortex. Using population receptive-field (pRF) mapping techniques, which allow estimates to be made of aggregate receptive field sizes, we found that mouse visual cortex contains a region in which pRFs are considerably smaller. This region, the ‘focea’, represents a location in space directly in front of, and slightly above, the mouse. Using two-photon imaging we show that the smaller pRFs are due to a more orderly representation of space and an over-representation of binocular regions of the visual scene. We also show that RFs of single neurons in areas LM, AL and AM are smaller at the focea. Mice have improved visual resolution in this region of space and freely-moving mice make compensatory eye-movements to hold this region in front of them. Our results indicate that the representation of space in mouse visual cortex is non-uniform and mice have spatial biases in their visual processing. The presence of a focea has important implications for the use of the mouse model of vision.

Influence of ocular dominance columns and patchy callosal connections on binocularity in lateral striate cortex: Long Evans versus albino rats

In albino rats, it has been reported that lateral striate cortex (V1) is highly binocular, and that input from the ipsilateral eye to this region comes through the callosum. In contrast, in Long Evans rats, this region is nearly exclusively dominated by the contralateral eye even though it is richly innervated by the callosum (Laing, Turecek, Takahata, & Olavarria, 2015). We hypothesized that the inability of callosal connections to relay ipsilateral eye input to lateral V1 in Long Evans rats is a consequence of the existence of ocular dominance columns (ODCs), and of callosal patches in register with ipsilateral ODCs in the binocular region of V1 (Laing et al., 2015). We therefore predicted that in albino rats input from both eyes intermix in the binocular region, without segregating into ODCs, and that callosal connections are not patchy. Confirming our predictions, we found that inputs from both eyes, studied with the transneuronal tracer WGA-HRP, are intermixed in the binocular zone of albinos, without segregating into ODCs. Similarly, we found that callosal connections in albino rats are not patchy but instead are distributed homogeneously throughout the callosal region in V1. We propose that these changes allow the transcallosal passage of ipsilateral eye input to lateral striate cortex, increasing its binocularity. Thus, the binocular region in V1 of albino rats includes lateral striate cortex, being therefore about 25% larger in area than the binocular region in Long Evans rats. Our findings provide insight on the role of callosal connections in generating binocular cells.

Retinal and Callosal Activity-Dependent Chandelier Cell Elimination Shapes Binocularity in Primary Visual Cortex

In mammals with binocular vision, integration of the left and right visual scene begins with information at the center visual field, which are relayed from each retina in parallel and merge in the primary visual cortex (V1) through the convergence of ipsi- and contralateral geniculocortical inputs as well as trans-callosal projections between two visual hemispheres. The developmental mechanisms that assemble this binocular circuit remain incompletely understood. Using genetic methods in mice, we found that during the days before eye-opening, retinal and callosal activity drives massive apoptosis of GABAergic chandelier cells (ChCs) in the binocular region of V1. Blockade of ChC elimination resulted in a contralateral-dominated V1 and deficient binocular vision. As activity patterns in the retina prior to vision convey the organization of the visual field, their regulation of ChC density through the trans-callosal pathway may prime a nascent binocular territory for subsequent experience-driven tuning during the post-vision critical period.

Lifestyle shapes bird of prey binocular visual field

Swooping toward a victim with its vicious talons extended, a bird of prey homes in for the kill with meticulous precision. ‘Birds, particularly birds of prey [raptors], are believed to forage primarily using visual cues’, says vision scientist Simon Potier from Lund University, Sweden, who was inspired by his father's love of birds of prey to take up falconry as a small child. Yet, Potier explains that it was not clear how the birds use the region where the views from the two eyes overlap (the binocular zone) during their rapier-like attacks. As the hunting strategies of birds of prey span low-speed scavenging to high-speed aerial and ground prey capture, Potier, Francesco Bonadonna and Olivier Duriez from Université de Montpellier, France, and an international team of collaborators wondered whether the binocular vision of these precision hunters might vary depending on their lifestyle.Potier turned to his own collection of almost 60 birds at Les Ailes de l'Urga theme park in France and other birds from Le Grand Parc du Puy du Fou to measure the binocular visual fields of birds ranging from carrion-feeding vultures to predatory Cooper's hawks and eagles that target victims on the ground. ‘We used a non-invasive procedure to measure visual field characteristics in alert birds’, says Potier, recalling the difficulty of holding the large eagles steady for 20 min while measuring the limit of the view from each eye in order to calculate the binocular region where the visual fields overlapped. ‘The experience of the falconers who helped was essential’, he says. In addition, he had to custom build individual bill holders for each species to keep their heads in place, ‘which took me quite some time to fit perfectly’, he laughs. However, Potier was relieved when the birds appeared unfazed after they were released: ‘Directly after the experiment, they still flew to me, which indicates that they were not stressed’, he smiles.Initially, Potier and his colleagues had anticipated that the binocular visual fields of the hunters would be wider than those of the scavengers, as the hunters have to precisely target a moving victim with their talons while the scavengers’ aim may not need to be as precise. However, the width of the binocular visual fields was remarkably similar. The contrasts were more apparent when the team plotted the full projections of the binocular region, showing that the zone above the beak where the view from the two eyes overlapped tapered more rapidly in the ground-hunting eagles than it did in Cooper's hawks (which target birds on the wing) and the scavenging vultures. There was also an extensive blind zone above the heads of the ground predators – which do not have to outmanoeuvre agile targets on the wing. Considering the various hunting strategies and differences in the head shapes of the 15 species, Potier and his colleagues suspect that the ridge of bone that shades the eyes of the ground-hunting species could partially obscure the upper portion of their binocular field to produce the large blind zone. In addition, their short, but wide, powerful beaks appear to shade the lower portion of their view.‘Our results show that the binocular field configuration is specialized to foraging tactics in raptors’, says Potier, who is keen to scrutinise the vision of the almost exclusively terrestrial secretary bird. ‘They have very large eyes… I would love to estimate their visual field’, he muses.

What Drives Bird Vision? Bill Control and Predator Detection Overshadow Flight.

Although flight is regarded as a key behavior of birds this review argues that the perceptual demands for its control are met within constraints set by the perceptual demands of two other key tasks: the control of bill (or feet) position, and the detection of food items/predators. Control of bill position, or of the feet when used in foraging, and timing of their arrival at a target, are based upon information derived from the optic flow-field in the binocular region that encompasses the bill. Flow-fields use information extracted from close to the bird using vision of relatively low spatial resolution. The detection of food items and predators is based upon information detected at a greater distance and depends upon regions in the retina with relatively high spatial resolution. The tasks of detecting predators and of placing the bill (or feet) accurately, make contradictory demands upon vision and these have resulted in trade-offs in the form of visual fields and in the topography of retinal regions in which spatial resolution is enhanced, indicated by foveas, areas, and high ganglion cell densities. The informational function of binocular vision in birds does not lie in binocularity per se (i.e., two eyes receiving slightly different information simultaneously about the same objects) but in the contralateral projection of the visual field of each eye. This ensures that each eye receives information from a symmetrically expanding optic flow-field centered close to the direction of the bill, and from this the crucial information of direction of travel and time-to-contact can be extracted, almost instantaneously. Interspecific comparisons of visual fields between closely related species have shown that small differences in foraging techniques can give rise to different perceptual challenges and these have resulted in differences in visual fields even within the same genus. This suggests that vision is subject to continuing and relatively rapid natural selection based upon individual differences in the structure of the optical system, retinal topography, and eye position in the skull. From a sensory ecology perspective a bird is best characterized as “a bill guided by an eye” and that control of flight is achieved within constraints on visual capacity dictated primarily by the demands of foraging and bill control.

Rapid plasticity of visually evoked responses in rat monocular visual cortex.

Sensory cortical circuits are shaped by experience during sensitive periods in development. In the primary visual cortex (V1) altered visual experience results in changes in visual responsiveness of cortical neurons. The experience-dependent refinement of the circuit in V1 is thought to rely on competitive interactions between feedforward circuits driven by the two eyes. However, recent data have provided evidence for an additional role of cortico-cortical circuits in this process. Indeed, experience-dependent changes in intracortical circuits can be induced rapidly and may result in rapid-onset functional changes. Unilateral occlusion of vision rapidly alters visual responsiveness, synaptic strength and connectivity of local circuits in the binocular region of V1 (V1b), where the inputs from the two eyes converge. In the monocular region of rodent V1 (V1m), where feedforward inputs from the ipsilateral eye are virtually absent, visual deprivation induces rapid plasticity in local circuits; however, functional changes seem to occur only after long periods of deprivation. In V1m there is currently no evidence for functional changes occurring within a time window compatible with that of local circuit plasticity. Here, we probed the visual responsiveness of neurons in rat V1m and assessed the effect of one day unilateral eye lid suture on single neuron visual responses. We report a novel form of plasticity within V1m that occurs on a timescale consistent with the earliest known changes in synaptic strength. Our data provide new insights into how sensory experience can rapidly modulate neuronal responses, even in the absence of direct competition between feedforward thalamocortical inputs.

Monocular deprivation induces dendritic spine elimination in the developing mouse visual cortex

It is well established that visual deprivation has a profound impact on the responsiveness of neurons in the developing visual cortex. The effect of visual deprivation on synaptic connectivity remains unclear. Using transcranial two-photon microscopy, we examined the effect of visual deprivation and subsequent recovery on dendritic spine remodeling of layer 5 pyramidal neurons in the mouse primary visual cortex. We found that monocular deprivation (MD), but not binocular deprivation (BD), increased dendritic spine elimination over 3 days in the binocular region of 4-week-old adolescent mice. This MD-induced dendritic spine elimination persisted during subsequent 2–4 days of binocular recovery. Furthermore, we found that average dendritic spine sizes were decreased and increased following 3-day MD and BD, respectively. These spine size changes induced by MD or BD tended to be reversed during subsequent binocular recovery. Taken together, these findings reveal differential effects of MD and BD on synaptic connectivity of layer 5 pyramidal neurons and underscore the persistent impact of MD on synapse loss in the developing visual cortex.

RETRACTED ARTICLE: Quality assessment for virtual reality technology based on real scene

Virtual reality technology is a new display technology, which provides users with real viewing experience. As known, most of the virtual reality display through stereoscopic images. However, image quality will be influenced by the collection, storage and transmission process. If the stereoscopic image quality in the virtual reality technology is seriously damaged, the user will feel uncomfortable, and this can even cause healthy problems. In this paper, we establish a set of accurate and effective evaluations for the virtual reality. In the preprocessing, we segment the original reference and distorted image into binocular regions and monocular regions. Then, the Information-weighted SSIM (IW-SSIM) or Information-weighted PSNR (IW-PSNR) values over the monocular regions are applied to obtain the IW-score. At the same time, the Stereo-weighted-SSIM (SW-SSIM) or Stereo-weighted-PSNR (SW-PSNR) can be used to calculate the SW-score. Finally, we pool the stereoscopic images score by combing the IW-score and SW-score. Experiments show that our method is very consistent with human subjective judgment standard in the evaluation of virtual reality technology.

PMFS: A Perceptual Modulated Feature Similarity Metric for Stereoscopic Image Quality Assessment

Stereoscopic image quality assessment (SIQA) is an important and challenging issue in three dimensional applications. In this letter, a perceptual modulated feature similarity (PMFS) metric for SIQA is proposed by considering the monocular and binocular perception properties. Specifically, stereoscopic image is first classified into monocular occlusion and binocular rivalry regions. Then, feature similarities between the original and distorted stereoscopic images are defined and measured for the monocular occlusion and binocular rivalry regions as the local monocular and binocular quality maps, respectively. Monocular and binocular just noticeable difference visual saliency models are presented to construct a modulation function to derive monocular and binocular quality scores. Finally, those scores are integrated into an overall quality score by support vector regression. Extensive experiments performed on LIVE phase II and MICT asymmetric databases demonstrate that the proposed PMFS metric can achieve much higher consistency with the subjective quality scores than some state-of-the-art SIQA metrics.

Identification of Eye-Specific Domains and Their Relation to Callosal Connections in Primary Visual Cortex of Long Evans Rats.

Ocular dominance columns (ODCs) exist in many primates and carnivores, but it is believed that they do not exist in rodents. Using a combination of transneuronal tracing, in situ hybridization for Zif268 and electrophysiological recordings, we show that inputs from both eyes are largely segregated in the binocular region of V1 in Long Evans rats. We also show that, interposed between this binocular region and the lateral border of V1, there lies a strip of cortex that is strongly dominated by the contralateral eye. Finally, we show that callosal connections colocalize primarily with ipsilateral eye domains in the binocular region and with contralateral eye input in the lateral cortical strip, mirroring the relationship between patchy callosal connections and specific sets of ODCs described previously in the cat. Our results suggest that development of cortical modular architecture is more conserved among rodents, carnivores, and primates than previously thought.

Perceptual integration across natural monocular regions

Natural scenes contain hidden regions, or occlusions, that differ in the two eyes, resulting in monocular regions that can only be seen by one eye. Such monocular regions appear to not be suppressed but seem to be integrated into the scene percept. Here we explore how the two eyes' views are combined to represent a scene that contains monocular regions, partially hidden behind a foreground occluding "fence." We measured performance in a density/numerosity discrimination task for scenes containing differing amounts of binocular and monocular information. We find that information from a number of separate monocular regions can be integrated into our overall percept of dot density/numerosity, although different observers use different strategies. If, however, both monocular and binocular information is present, observers appear to ignore the purely monocular regions, relying solely on the binocular information when making density/numerosity judgments. Our work suggests that binocular regions are favored over monocular regions, such that information from monocular regions is effectively ignored when binocular regions are present in a scene.

The subtlety of simple eyes: the tuning of visual fields to perceptual challenges in birds

Birds show interspecific variation both in the size of the fields of individual eyes and in the ways that these fields are brought together to produce the total visual field. Variation is found in the dimensions of all main parameters: binocular region, cyclopean field and blind areas. There is a phylogenetic signal with respect to maximum width of the binocular field in that passerine species have significantly broader field widths than non-passerines; broadest fields are found among crows (Corvidae). Among non-passerines, visual fields show considerable variation within families and even within some genera. It is argued that (i) the main drivers of differences in visual fields are associated with perceptual challenges that arise through different modes of foraging, and (ii) the primary function of binocularity in birds lies in the control of bill position rather than in the control of locomotion. The informational function of binocular vision does not lie in binocularity per se (two eyes receiving slightly different information simultaneously about the same objects from which higher-order depth information is extracted), but in the contralateral projection of the visual field of each eye. Contralateral projection ensures that each eye receives information from a symmetrically expanding optic flow-field from which direction of travel and time to contact targets can be extracted, particularly with respect to the control of bill position.

Enhanced iris recognition method based on multi-unit iris images

For the purpose of biometric person identification, iris recognition uses the unique characteristics of the patterns of the iris; that is, the eye region between the pupil and the sclera. When obtaining an iris image, the iris’s image is frequently rotated because of the user’s head roll toward the left or right shoulder. As the rotation of the iris image leads to circular shifting of the iris features, the accuracy of iris recognition is degraded. To solve this problem, conventional iris recognition methods use shifting of the iris feature codes to perform the matching. However, this increases the computational complexity and level of false acceptance error. To solve these problems, we propose a novel iris recognition method based on multi-unit iris images. Our method is novel in the following five ways compared with previous methods. First, to detect both eyes, we use Adaboost and a rapid eye detector (RED) based on the iris shape feature and integral imaging. Both eyes are detected using RED in the approximate candidate region that consists of the binocular region, which is determined by the Adaboost detector. Second, we classify the detected eyes into the left and right eyes, because the iris patterns in the left and right eyes in the same person are different, and they are therefore considered as different classes. We can improve the accuracy of iris recognition using this pre-classification of the left and right eyes. Third, by measuring the angle of head roll using the two center positions of the left and right pupils, detected by two circular edge detectors, we obtain the information of the iris rotation angle. Fourth, in order to reduce the error and processing time of iris recognition, adaptive bit-shifting based on the measured iris rotation angle is used in feature matching. Fifth, the recognition accuracy is enhanced by the score fusion of the left and right irises. Experimental results on the iris open database of low-resolution images showed that the averaged equal error rate of iris recognition using the proposed method was 4.3006%, which is lower than that of other methods.