- Home
- A-Z Publications
- Annual Review of Vision Science
- Previous Issues
- Volume 6, 2020
Annual Review of Vision Science - Volume 6, 2020
Volume 6, 2020
-
-
Fifty Years Exploring the Visual System
Vol. 6 (2020), pp. 1–23More LessWe as a couple spent 50 years working in visual psychophysics of color vision, temporal vision, and luminance adaptation. We sought collaborations with ophthalmologists, anatomists, physiologists, physicists, and psychologists, aiming to relate visual psychophysics to the underlying physiology of the primate retina. This review describes our journey and reflections in exploring the visual system.
-
-
-
Genetic and Environmental Risk Factors for Keratoconus
Vol. 6 (2020), pp. 25–46More LessKeratoconus, a progressive corneal ectasia, is a complex disease with both genetic and environmental risk factors. The exact etiology is not known and is likely variable between individuals. Conditions such as hay fever and allergy are associated with increased risk, while diabetes may be protective. Behaviors such as eye rubbing are also implicated, but direct causality has not been proven. Genetics plays a major role in risk for some individuals, with many large pedigrees showing autosomal inheritance patterns. Several genes have been implicated using linkage and follow-up sequencing in these families. Genome-wide association studies for keratoconus and for quantitative traits such as central corneal thickness have identified several genetic loci that contribute to a cumulative risk for keratoconus, even in people without a family history of the disease. Identification of risk genes for keratoconus is improving our understanding of the biology of this complex disease.
-
-
-
Minimally Invasive Glaucoma Surgery: A Critical Appraisal of the Literature
Vol. 6 (2020), pp. 47–89More LessMicro- or minimally invasive glaucoma surgeries (MIGS) have been the latest addition to the glaucoma surgical treatment paradigm. This term refers not to a single surgery, but rather to a group of distinct procedures and devices that aim to decrease intraocular pressure. Broadly, MIGS can be categorized into surgeries that increase the trabecular outflow [Trabectome, iStent (first and second generations), Hydrus microstent, Kahook Dual Blade and gonioscopy-assisted transluminal trabeculotomy], surgeries that increase suprachoroidal outflow (Cypass microstent and iStent Supra), and conjunctival bleb-forming procedures (Xen gel stent and InnFocus microshunt). Compared to traditional glaucoma surgeries, such as trabeculectomy and glaucoma drainage device implantation (Ahmed, Baerveldt, and Molteno valves), MIGS are touted to have less severe complications and shorter surgical time. MIGS represent an evolving field, and the efficacy and complications of each procedure should be considered independently, giving more importance to high-quality and longer-term studies.
-
-
-
Human Organoids for the Study of Retinal Development and Disease
Vol. 6 (2020), pp. 91–114More LessRecent advances in stem cell engineering have led to an explosion in the use of organoids as model systems for studies in multiple biological disciplines. Together with breakthroughs in genome engineering and the various omics, organoid technology is making possible studies of human biology that were not previously feasible. For vision science, retinal organoids derived from human stem cells allow differentiating and mature human retinal cells to be studied in unprecedented detail. In this review, we examine the technologies employed to generate retinal organoids and how organoids are revolutionizing the fields of developmental and cellular biology as they pertain to the retina. Furthermore, we explore retinal organoids from a clinical standpoint, offering a new platform with which to study retinal diseases and degeneration, test prospective drugs and therapeutic strategies, and promote personalized medicine. Finally, we discuss the range of possibilities that organoids may bring to future retinal research and consider their ethical implications.
-
-
-
Cellular-Scale Imaging of Transparent Retinal Structures and Processes Using Adaptive Optics Optical Coherence Tomography
Vol. 6 (2020), pp. 115–148More LessHigh-resolution retinal imaging is revolutionizing how scientists and clinicians study the retina on the cellular scale. Its exquisite sensitivity enables time-lapse optical biopsies that capture minute changes in the structure and physiological processes of cells in the living eye. This information is increasingly used to detect disease onset and monitor disease progression during early stages, raising the possibility of personalized eye care. Powerful high-resolution imaging tools have been in development for more than two decades; one that has garnered considerable interest in recent years is optical coherence tomography enhanced with adaptive optics. State-of-the-art adaptive optics optical coherence tomography (AO-OCT) makes it possible to visualize even highly transparent cells and measure some of their internal processes at all depths within the retina, permitting reconstruction of a 3D view of the living microscopic retina. In this review, we report current AO-OCT performance and its success in visualizing and quantifying these once-invisible cells in human eyes.
-
-
-
Microglia Activation and Inflammation During the Death of Mammalian Photoreceptors
Vol. 6 (2020), pp. 149–169More LessPhotoreceptors are highly specialized sensory neurons with unique metabolic and physiological requirements. These requirements are partially met by Müller glia and cells of the retinal pigment epithelium (RPE), which provide essential metabolites, phagocytose waste, and control the composition of the surrounding microenvironment. A third vital supporting cell type, the retinal microglia, can provide photoreceptors with neurotrophic support or exacerbate neuroinflammation and hasten neuronal cell death. Understanding the physiological requirements for photoreceptor homeostasis and the factors that drive microglia to best promote photoreceptor survival has important implications for the treatment and prevention of blinding degenerative diseases like retinitis pigmentosa and age-related macular degeneration.
-
-
-
Reprogramming Müller Glia to Regenerate Retinal Neurons
Vol. 6 (2020), pp. 171–193More LessIn humans, various genetic defects or age-related diseases, such as diabetic retinopathies, glaucoma, and macular degeneration, cause the death of retinal neurons and profound vision loss. One approach to treating these diseases is to utilize stem and progenitor cells to replace neurons in situ, with the expectation that new neurons will create new synaptic circuits or integrate into existing ones. Reprogramming non-neuronal cells in vivo into stem or progenitor cells is one strategy for replacing lost neurons. Zebrafish have become a valuable model for investigating cellular reprogramming and retinal regeneration. This review summarizes our current knowledge regarding spontaneous reprogramming of Müller glia in zebrafish and compares this knowledge to research efforts directed toward reprogramming Müller glia in mammals. Intensive research using these animal models has revealed shared molecular mechanisms that make Müller glia attractive targets for cellular reprogramming and highlighted the potential for curing degenerative retinal diseases from intrinsic cellular sources.
-
-
-
Axon Regeneration in the Mammalian Optic Nerve
Vol. 6 (2020), pp. 195–213More LessThe damage or loss of retinal ganglion cells (RGCs) and their axons accounts for the visual functional defects observed after traumatic injury, in degenerative diseases such as glaucoma, or in compressive optic neuropathies such as from optic glioma. By using optic nerve crush injury models, recent studies have revealed the cellular and molecular logic behind the regenerative failure of injured RGC axons in adult mammals and suggested several strategies with translational potential. This review summarizes these findings and discusses challenges for developing clinically applicable neural repair strategies.
-
-
-
Retinal Ganglion Cell Axon Wiring Establishing the Binocular Circuit
Carol Mason, and Nefeli SlaviVol. 6 (2020), pp. 215–236More LessBinocular vision depends on retinal ganglion cell (RGC) axon projection either to the same side or to the opposite side of the brain. In this article, we review the molecular mechanisms for decussation of RGC axons, with a focus on axon guidance signaling at the optic chiasm and ipsi- and contralateral axon organization in the optic tract prior to and during targeting. The spatial and temporal features of RGC neurogenesis that give rise to ipsilateral and contralateral identity are described. The albino visual system is highlighted as an apt comparative model for understanding RGC decussation, as albinos have a reduced ipsilateral projection and altered RGC neurogenesis associated with perturbed melanogenesis in the retinal pigment epithelium. Understanding the steps for RGC specification into ipsi- and contralateral subtypes will facilitate differentiation of stem cells into RGCs with proper navigational abilities for effective axon regeneration and correct targeting of higher-order visual centers.
-
-
-
Topographic Variations in Retinal Encoding of Visual Space
Vol. 6 (2020), pp. 237–259More LessA retina completely devoid of topographic variations would be homogenous, encoding any given feature uniformly across the visual field. In a naive view, such homogeneity would appear advantageous. However, it is now clear that retinal topographic variations exist across mammalian species in a variety of forms and patterns. We briefly review some of the more established topographic variations in retinas of different mammalian species and focus on the recent discovery that cells belonging to a single neuronal subtype may exhibit distinct topographic variations in distribution, morphology, and even function. We concentrate on the mouse retina—originally viewed as homogenous—in which genetic labeling of distinct neuronal subtypes and other advanced techniques have revealed unexpected anatomical and physiological topographic variations. Notably, different subtypes reveal different patterns of nonuniformity, which may even be opposite or orthogonal to one another. These topographic variations in the encoding of visual space should be considered when studying visual processing in the retina and beyond.
-
-
-
Organization, Function, and Development of the Mouse Retinogeniculate Synapse
Liang Liang, and Chinfei ChenVol. 6 (2020), pp. 261–285More LessVisual information is encoded in distinct retinal ganglion cell (RGC) types in the eye tuned to specific features of the visual space. These streams of information project to the visual thalamus, the first station of the image-forming pathway. In the mouse, this connection between RGCs and thalamocortical neurons, the retinogeniculate synapse, has become a powerful experimental model for understanding how circuits in the thalamus are constructed to process these incoming lines of information. Using modern molecular and genetic tools, recent studies have suggested a more complex circuit organization than was previously understood. In this review, we summarize the current understanding of the structural and functional organization of the retinogeniculate synapse in the mouse. We discuss a framework by which a seemingly complex circuit can effectively integrate and parse information to downstream stations of the visual pathway. Finally, we review how activity and visual experience can sculpt this exquisite connectivity.
-
-
-
Signals Related to Color in the Early Visual Cortex
Vol. 6 (2020), pp. 287–311More LessVisual images can be described in terms of the illuminants and objects that are causal to the light reaching the eye, the retinal image, its neural representation, or how the image is perceived. Respecting the differences among these distinct levels of description can be challenging but is crucial for a clear understanding of color vision. This article approaches color by reviewing what is known about its neural representation in the early visual cortex, with a brief description of signals in the eye and the thalamus for context. The review focuses on the properties of single neurons and advances the general theme that experimental approaches based on knowledge of feedforward signals have promoted greater understanding of the neural code for color than approaches based on correlating single-unit responses with color perception. New data from area V1 illustrate the strength of the feedforward approach. Future directions for progress in color neurophysiology are discussed: techniques for improved single-neuron characterization, for investigations of neural populations and small circuits, and for the analysis of natural image statistics.
-
-
-
Role of Feedback Connections in Central Visual Processing
Vol. 6 (2020), pp. 313–334More LessThe physiological response properties of neurons in the visual system are inherited mainly from feedforward inputs. Interestingly, feedback inputs often outnumber feedforward inputs. Although they are numerous, feedback connections are weaker, slower, and considered to be modulatory, in contrast to fast, high-efficacy feedforward connections. Accordingly, the functional role of feedback in visual processing has remained a fundamental mystery in vision science. At the core of this mystery are questions about whether feedback circuits regulate spatial receptive field properties versus temporal responses among target neurons, or whether feedback serves a more global role in arousal or attention. These proposed functions are not mutually exclusive, and there is compelling evidence to support multiple functional roles for feedback. In this review, the role of feedback in vision will be explored mainly from the perspective of corticothalamic feedback. Further generalized principles of feedback applicable to corticocortical connections will also be considered.
-
-
-
Linking Neuronal Direction Selectivity to Perceptual Decisions About Visual Motion
Vol. 6 (2020), pp. 335–362More LessPsychophysical and neurophysiological studies of responses to visual motion have converged on a consistent set of general principles that characterize visual processing of motion information. Both types of approaches have shown that the direction and speed of target motion are among the most important encoded stimulus properties, revealing many parallels between psychophysical and physiological responses to motion. Motivated by these parallels, this review focuses largely on more direct links between the key feature of the neuronal response to motion, direction selectivity, and its utilization in memory-guided perceptual decisions. These links were established during neuronal recordings in monkeys performing direction discriminations, but also by examining perceptual effects of widespread elimination of cortical direction selectivity produced by motion deprivation during development. Other approaches, such as microstimulation and lesions, have documented the importance of direction-selective activity in the areas that are active during memory-guided direction comparisons, area MT and the prefrontal cortex, revealing their likely interactions during behavioral tasks.
-
-
-
Visual Functions of Primate Area V4
Vol. 6 (2020), pp. 363–385More LessArea V4—the focus of this review—is a mid-level processing stage along the ventral visual pathway of the macaque monkey. V4 is extensively interconnected with other visual cortical areas along the ventral and dorsal visual streams, with frontal cortical areas, and with several subcortical structures. Thus, it is well poised to play a broad and integrative role in visual perception and recognition—the functional domain of the ventral pathway. Neurophysiological studies in monkeys engaged in passive fixation and behavioral tasks suggest that V4 responses are dictated by tuning in a high-dimensional stimulus space defined by form, texture, color, depth, and other attributes of visual stimuli. This high-dimensional tuning may underlie the development of object-based representations in the visual cortex that are critical for tracking, recognizing, and interacting with objects. Neurophysiological and lesion studies also suggest that V4 responses are important for guiding perceptual decisions and higher-order behavior.
-
-
-
Coding Perceptual Decisions: From Single Units to Emergent Signaling Properties in Cortical Circuits
Vol. 6 (2020), pp. 387–409More LessSpiking activity in single neurons of the primate visual cortex has been tightly linked to perceptual decisions. Any mechanism that reads out these perceptual signals to support behavior must respect the underlying neuroanatomy that shapes the functional properties of sensory neurons. Spatial distribution and timing of inputs to the next processing levels are critical, as conjoint activity of precursor neurons increases the spiking rate of downstream neurons and ultimately drives behavior. I set out how correlated activity might coalesce into a micropool of task-sensitive neurons signaling a particular percept to determine perceptual decision signals locally and for flexible interarea transmission depending on the task context. As data from more and more neurons and their complex interactions are analyzed, the space of computational mechanisms must be constrained based on what is plausible within neurobiological limits. This review outlines experiments to test the new perspectives offered by these extended methods.
-
-
-
Anatomy and Function of the Primate Entorhinal Cortex
Vol. 6 (2020), pp. 411–432More LessThe entorhinal cortex (EC) is a critical element of the hippocampal formation located within the medial temporal lobe (MTL) in primates. The EC has historically received attention for being the primary mediator of cortical information going into and coming from the hippocampus proper. In this review, we highlight the significance of the EC as a major player in memory processing, along with other associated structures in the primate MTL. The complex, convergent topographies of cortical and subcortical input to the EC, combined with short-range intrinsic connectivity and the selective targeting of EC efferents to the hippocampus, provide evidence for subregional specialization and integration of information beyond what would be expected if this structure were a simple conduit of information for the hippocampus. Lesion studies of the EC provide evidence implicating this region as critical for memory and the flexible use of complex relational associations between experienced events. The physiology of this structure's constituent principal cells mirrors the complexity of its anatomy. EC neurons respond preferentially to aspects of memory-dependent paradigms including object, place, and time. EC neurons also show striking spatial representations as primates explore visual space, similar to those identified in rodents navigating physical space. In this review, we highlight the great strides that have been made toward furthering our understanding of the primate EC, and we identify paths forward for future experiments to provide additional insight into the role of this structure in learning and memory.
-
-
-
Tuning the Senses: How the Pupil Shapes Vision at the Earliest Stage
Vol. 6 (2020), pp. 433–451More LessThe pupil responds reflexively to changes in brightness and focal distance to maintain the smallest pupil (and thus the highest visual acuity) that still allows sufficient light to reach the retina. The pupil also responds to a wide variety of cognitive processes, but the functions of these cognitive responses are still poorly understood. In this review, I propose that cognitive pupil responses, like their reflexive counterparts, serve to optimize vision. Specifically, an emphasis on central vision over peripheral vision results in pupil constriction, and this likely reflects the fact that central vision benefits most from the increased visual acuity provided by small pupils. Furthermore, an intention to act with a bright stimulus results in preparatory pupil constriction, which allows the pupil to respond quickly when that bright stimulus is subsequently brought into view. More generally, cognitively driven pupil responses are likely a form of sensory tuning: a subtle adjustment of the eyes to optimize their properties for the current situation and the immediate future.
-
-
-
Can We See with Melanopsin?
Vol. 6 (2020), pp. 453–468More LessA small fraction of mammalian retinal ganglion cells are directly photoreceptive thanks to their expression of the photopigment melanopsin. These intrinsically photosensitive retinal ganglion cells (ipRGCs) have well-established roles in a variety of reflex responses to changes in ambient light intensity, including circadian photoentrainment. In this article, we review the growing evidence, obtained primarily from laboratory mice and humans, that the ability to sense light via melanopsin is also an important component of perceptual and form vision. Melanopsin photoreception has low temporal resolution, making it fundamentally biased toward detecting changes in ambient light and coarse patterns rather than fine details. Nevertheless, melanopsin can indirectly impact high-acuity vision by driving aspects of light adaptation ranging from pupil constriction to changes in visual circuit performance. Melanopsin also contributes directly to perceptions of brightness, and recent data suggest that this influences the appearance not only of overall scene brightness, but also of low-frequency patterns.
-
-
-
Retinal Image Formation and Sampling in a Three-Dimensional World
Vol. 6 (2020), pp. 469–489More LessIn this review, I develop an empirically based model of optical image formation by the human eye, followed by neural sampling by retinal ganglion cells, to demonstrate the perceptual effects of blur, aliasing, and distortion of visual space in the brain. The optical model takes account of ocular aberrations and their variation across the visual field, in addition to variations of defocus due to variation of target vergence in three-dimensional scenes. Neural sampling by retinal ganglion cells with receptive field size and spacing that increases with eccentricity is used to visualize the neural image carried by the optic nerve to the brain. Anatomical parameters are derived from psychophysical studies of sampling-limited visual resolution of sinusoidal interference fringes. Retinotopic projection of the neural image onto brainstem nuclei reveals features of the neural image in a perceptually uniform brain space where location and size of visual objects may be measured by counting neurons.
-