Primate V2 Receptive Fields Derived from Anatomically Identified Large-Scale V1 Inputs.

In the primate visual system, visual object recognition involves a series of cortical areas arranged hierarchically along the ventral visual pathway. As information flows through this hierarchy, neurons become progressively tuned to more complex image features. The circuit mechanisms and computations underlying the increasing complexity of these receptive fields (RFs) remain unidentified. To understand how this complexity emerges in the secondary visual area (V2), we investigated the functional organization of inputs from the primary visual cortex (V1) to V2 by combining retrograde anatomical tracing of these inputs with functional imaging of feature maps in macaque monkey V1 and V2. We found that V1 neurons sending inputs to single V2 orientation columns have a broad range of preferred orientations, but are strongly biased towards the orientation represented at the injected V2 site. For each V2 site, we then constructed a feedforward model based on the linear combination of its anatomically-identified large-scale V1 inputs, and studied the response proprieties of the generated V2 RFs. We found that V2 RFs derived from the linear feedforward model were either elongated versions of V1 filters or had spatially complex structures. These modeled RFs predicted V2 neuron responses to oriented grating stimuli with high accuracy. Remarkably, this simple model also explained the greater selectivity to naturalistic textures of V2 cells compared to their V1 input cells. Our results demonstrate that simple linear combinations of feedforward inputs can account for the orientation selectivity and texture sensitivity of V2 RFs.

Perceptual transitions between object rigidity and non-rigidity: Competition and cooperation among motion energy, feature tracking, and shape-based priors.

Why do moving objects appear rigid when projected retinal images are deformed non-rigidly? We used rotating rigid objects that can appear rigid or non-rigid to test whether shape features contribute to rigidity perception. When two circular rings were rigidly linked at an angle and jointly rotated at moderate speeds, observers reported that the rings wobbled and were not linked rigidly, but rigid rotation was reported at slow speeds. When gaps, paint, or vertices were added, the rings appeared rigidly rotating even at moderate speeds. At high speeds, all configurations appeared non-rigid. Salient features thus contribute to rigidity at slow and moderate speeds but not at high speeds. Simulated responses of arrays of motion-energy cells showed that motion flow vectors are predominantly orthogonal to the contours of the rings, not parallel to the rotation direction. A convolutional neural network trained to distinguish flow patterns for wobbling versus rotation gave a high probability of wobbling for the motion-energy flows. However, the convolutional neural network gave high probabilities of rotation for motion flows generated by tracking features with arrays of MT pattern-motion cells and corner detectors. In addition, circular rings can appear to spin and roll despite the absence of any sensory evidence, and this illusion is prevented by vertices, gaps, and painted segments, showing the effects of rotational symmetry and shape. Combining convolutional neural network outputs that give greater weight to motion energy at fast speeds and to feature tracking at slow speeds, with the shape-based priors for wobbling and rolling, explained rigid and non-rigid percepts across shapes and speeds (R2 = 0.95). The results demonstrate how cooperation and competition between different neuronal classes lead to specific states of visual perception and to transitions between the states.

Perceptual Transitions between Object Rigidity & Non-rigidity: Competition and cooperation between motion-energy, feature-tracking and shape-based priors.

Why do moving objects appear rigid when projected retinal images are deformed non-rigidly? We used rotating rigid objects that can appear rigid or non-rigid to test whether shape features contribute to rigidity perception. When two circular rings were rigidly linked at an angle and jointly rotated at moderate speeds, observers reported that the rings wobbled and were not linked rigidly but rigid rotation was reported at slow speeds. When gaps, paint or vertices were added, the rings appeared rigidly rotating even at moderate speeds. At high speeds, all configurations appeared non-rigid. Salient features thus contribute to rigidity at slow and moderate speeds, but not at high speeds. Simulated responses of arrays of motion-energy cells showed that motion flow vectors are predominantly orthogonal to the contours of the rings, not parallel to the rotation direction. A convolutional neural network trained to distinguish flow patterns for wobbling versus rotation, gave a high probability of wobbling for the motion-energy flows. However, the CNN gave high probabilities of rotation for motion flows generated by tracking features with arrays of MT pattern-motion cells and corner detectors. In addition, circular rings can appear to spin and roll despite the absence of any sensory evidence, and this illusion is prevented by vertices, gaps, and painted segments, showing the effects of rotational symmetry and shape. Combining CNN outputs that give greater weight to motion energy at fast speeds and to feature tracking at slow, with the shape-based priors for wobbling and rolling, explained rigid and nonrigid percepts across shapes and speeds (R2=0.95). The results demonstrate how cooperation and competition between different neuronal classes leads to specific states of visual perception and to transitions between the states.

Clinical Significance of Lateral Pedicle Screw Malposition in Lumbar Spine Fusion.

STUDY DESIGN: Retrospective cohort study. OBJECTIVE: Our objective was to determine whether lateral pedicle screw breach affects fusion rates and patient-reported outcomes in lumbar fusion surgery. SUMMARY OF BACKGROUND DATA: Although lateral pedicle screw malposition is considered relatively benign, few studies have focused specifically on clinical outcomes or fusion rates associated with lateral screw malposition. METHODS: Twelve-month postoperative computed tomography scans were reviewed for lateral breach, severity of breach, and fusion status. Patients with lateral breach were compared with patients with no breach. Outcome measures included Numerical Pain Rating Scale for back and leg pain, Oswestry Disability Index, and SF-36 physical function (SF-36 PF). Multivariable linear and logistic regression and were adjusted for age, procedure, level, and/or baseline pain score. RESULTS: Forty-five patients (31%) demonstrated 1 or more lateral breaches as compared with 99 patients without breach. After adjusting for baseline scores and fusion level, patients with 2 or more screw breaches experienced SF-36 PF score improvements that were 3.43 points less ( P =0.016) than patients with no lateral breach. After adjusting for baseline Numerical Pain Rating Scale, there was also a significant decrease in the odds of achieving minimally clinical important difference in back pain relief in these patients. There was no observed effect of lateral breach on the odds of successful fusion. CONCLUSIONS: The current study did not observe an association between laterally malpositioned pedicle screws and nonunion. However, results are consistent with a negative effect on SF-36 PF scores and self-reported back pain at 12 months.

Predicting Pedicle Screw Pullout and Fatigue Performance: Comparing Lateral Dual-Energy X-Ray Absorptiometry, Anterior to Posterior Dual-Energy X-Ray Absorptiometry, and Computed Tomography Hounsfield Units.

BACKGROUND: As the prevalence and associated health care costs of osteoporosis continue to rise in our aging population, there is a growing need to continue to identify methods to predict spine construct integrity accurately and cost-effectively. Dual-energy x-ray absorptiometry (DEXA) in both anterior to posterior (AP) and lateral planes, as well as computed tomography (CT) Hounsfield units (HU), have all been investigated as potential preoperative predictive tools. The purpose of this study is to determine which of the 3 bone density analysis modalities has the highest potential for predicting pedicle screw biomechanics. METHODS: Lumbar spine specimens (L2, L3, and L4) from 6 fresh frozen cadavers were used for testing. AP-DEXA, lateral-DEXA, and CT images were obtained. Biomechanical testing of pedicle screws in each vertebrae was then performed including pullout strength and fatigue testing. Statistical analysis was performed. RESULTS: Pullout strength was best predicted by CT HU, followed by AP-DEXA, then lateral-DEXA (R 2 = 0.78, 0.70, and 0.40, respectively). Fatigue testing showed a significant correlation of relative rotation between HU value and AP-DEXA bone mineral density (R 2 = 0.54 and R 2 = 0.72, respectively), and there was a significant correlation between relative translation and HU value (R 2 = 0.43). There was a poor correlation between relative rotation and lateral-DEXA (R 2 = 0.13) as well as a poor correlation between relative translation and both AP- and lateral-DEXA (R 2 = 0.35 and R 2 = 0.02). CONCLUSIONS: CT is the only modality with a statistically significant correlation to all biomechanical parameters measured (pullout strength, relative angular rotation, and relative translation). AP-DEXA also predicts the biomechanical measures of screw pullout and relative angular rotation and is superior to lateral-DEXA. CT may provide an incremental benefit in assessing fatigue strength, but this should be weighed against the disadvantages of cost and radiation. CLINICAL RELEVANCE: The results of this study can help to inform clinicians on different bone density analyses and their implications on pedicle screw failure.

Perceptual scale for transparency: Common fate overrides geometrical and color cues.

Objects that pass light through are considered transparent, and we generally expect that the light coming out will match the color of the object. However, when the object is placed on a colored surface, the light coming back to our eyes becomes a composite of surface, illumination, and transparency properties. Despite that, we can often perceive separate overlaid and overlaying layers differing in colors. How neurons separate the information to extract the transparent layer remains unknown, but the physical characteristics of transparent filters generate geometrical and color features in retinal images, which could provide cues for separating layers. We estimated the relative importance of such cues in a perceptual scale for transparency, using stimuli in which X- or T-junctions, different relative motions, and consistent or inconsistent colors cooperated or competed in forced-preference psychophysics experiments. Maximum-likelihood Thurstone scaling revealed that motion increased transparency for X-junctions, but decreased transparency for T-junctions by creating the percept of an opaque patch. However, if the motion of a filter uncovered a dynamically changing but stationary pattern, sharing a common fate with the surround but forming T-junctions, the probability of seeing transparency was almost as high as for moving X-junctions, despite the stimulus being physically improbable. In addition, geometric cues overrode color inconsistency to a great degree. Finally, a linear model of transparency perception as a function of relative motions between filter, overlay, and surround layers, contour continuation, and color consistency, quantified a hierarchy of latent influences on when the filter is seen as a separate transparent layer.

A theory of cortical map formation in the visual brain.

The cerebral cortex receives multiple afferents from the thalamus that segregate by stimulus modality forming cortical maps for each sense. In vision, the primary visual cortex maps the multiple dimensions of the visual stimulus in patterns that vary across species for reasons unknown. Here we introduce a general theory of cortical map formation, which proposes that map diversity emerges from species variations in the thalamic afferent density sampling sensory space. In the theory, increasing afferent sampling density enlarges the cortical domains representing the same visual point, allowing the segregation of afferents and cortical targets by multiple stimulus dimensions. We illustrate the theory with an afferent-density model that accurately replicates the maps of different species through afferent segregation followed by thalamocortical convergence pruned by visual experience. Because thalamocortical pathways use similar mechanisms for axon segregation and pruning, the theory may extend to other sensory areas of the mammalian brain.

Spatial Induction in Color Scission.

An exception to the rule that only one color is seen at every retinotopic location happens when a bounded colored transparency or spotlight is seen on a differently colored surface. Despite the spectrum of the light from each retinotopic location being an inextricable multiplication of illumination, transmission, and reflectance spectra, we seem to be able to scission the information into background and transparency/spotlight colors. Visual cues to separating overlay and overlaid layers have been enumerated, but neural mechanisms that extract veridical colors for overlays have not been identified. Here, we demonstrate that spatial induction contributes to color scission by shifting the color of the overlay toward the actual color of the filter. By alternating filter and illumination spectra, we present naturalistic simulations where isomeric disks appear to be covered by filters/spotlights of near veridical colors, depending solely on the surrounding illumination. This previously unrecognized role for spatial induction suggests that color scission employs some general purpose neural mechanisms.

Mental geometry of perceiving 3D size in pictures.

We show that the classical problem of three-dimensional (3D) size perception in obliquely viewed pictures can be understood by comparing human performance to the optimal geometric solution. A photograph seen from the camera position, can form the same retinal projection as the physical 3D scene, but retinal projections of sizes and shapes are distorted in oblique viewing. For real scenes, we previously showed that size and shape inconstancy result despite observers using the correct geometric back-transform, because some retinal images evoke misestimates of object slant or viewing elevation. Now, we examine how observers estimate 3D sizes in oblique views of pictures of objects lying on the ground in different poses. Compared to estimates for real scenes, in oblique views of pictures, sizes were seriously underestimated for objects at frontoparallel poses, but there was almost no change for objects perceived as pointing toward the viewer. The inverse of the function relating projected length to pose, camera elevation and viewing azimuth, gives the optimal correction factor for inferring correct 3D lengths if the elevation and azimuth are estimated accurately. Empirical correction functions had similar shapes to optimal, but lower amplitude. Measurements revealed that observers systematically underestimated viewing azimuth, similar to the frontoparallel bias for object pose perception. A model that adds underestimation of viewing azimuth to the geometrical back-transform, provided good fits to estimated 3D lengths from oblique views. These results add to accumulating evidence that observers use internalized projective geometry to perceive sizes, shapes, and poses in 3D scenes and their pictures.

Mental geometry of three-dimensional size perception.

Judging the poses, sizes, and shapes of objects accurately is necessary for organisms and machines to operate successfully in the world. Retinal images of three-dimensional objects are mapped by the rules of projective geometry and preserve the invariants of that geometry. Since Plato, it has been debated whether geometry is innate to the human brain, and Poincare and Einstein thought it worth examining whether formal geometry arises from experience with the world. We examine if humans have learned to exploit projective geometry to estimate sizes and aspects of three-dimensional shape that are related to relative lengths and aspect ratios. Numerous studies have examined size invariance as a function of physical distance, which changes scale on the retina. However, it is surprising that possible constancy or inconstancy of relative size seems not to have been investigated for object pose, which changes retinal image size differently along different axes. We show systematic underestimation of length for extents pointing toward or away from the observer, both for static objects and dynamically rotating objects. Observers do correct for projected shortening according to the optimal back-transform, obtained by inverting the projection function, but the correction is inadequate by a multiplicative factor. The clue is provided by the greater underestimation for longer objects, and the observation that they seem to be more slanted toward the observer. Adding a multiplicative factor for perceived slant in the back-transform model provides good fits to the corrections used by observers. We quantify the slant illusion with two different slant matching measurements, and use a dynamic demonstration to show that the slant illusion perceptually dominates length nonrigidity. In biological and mechanical objects, distortions of shape are manifold, and changes in aspect ratio and relative limb sizes are functionally important. Our model shows that observers try to retain invariance of these aspects of shape to three-dimensional rotation by correcting retinal image distortions due to perspective projection, but the corrections can fall short. We discuss how these results imply that humans have internalized particular aspects of projective geometry through evolution or learning, and if humans assume that images are preserving the continuity, collinearity, and convergence invariances of projective geometry, that would simply explain why illusions such as Ames' chair appear cohesive despite being a projection of disjointed elements, and thus supplement the generic viewpoint assumption.

Geometrical structure of perceptual color space: Mental representations and adaptation invariance.

Similarity between percepts and concepts is used to accomplish many everyday tasks, e.g., object identification; so this similarity is widely used to construct geometrical spaces that represent stimulus qualities, but the intrinsic validity of the geometry, i.e., whether similarity operations support a particular geometry, is almost never tested critically. We introduce an experimental approach for equating relative similarities by setting perceived midpoints between pairs of stimuli. Midpoint settings are used with Varignon's Theorem to test the intrinsic geometry of a representation space, and its mapping to a physical space of stimuli. For perceptual color space, we demonstrate that geometrical structure depends on the mental representation used in judging similarity: An affine geometry was valid when observers used an opponent-color mental representation. Similarities based on a conceptual space of complementary colors thus power a geometric coordinate system. An affine geometry implies that similarity can be judged within straight lines and across parallel lines, and its neural coding could involve ratios of responses. We show that this perceptual space is invariant to changes in illumination color, providing a formal justification to generalize color constancy results measured for color categories, to all of color space. The midpoint measurements deviate significantly from midpoints in the extensively used "uniform" color spaces CIELAB and CIELUV, showing that these spaces do not provide adequate metric representation of perceived colors. Our paradigm can thus test for intrinsic geometrical assumptions underlying the representation space for many perceptual modalities, and for the extrinsic perceptual geometry of the space of physical stimuli.

Connectomic Identification and Three-Dimensional Color Tuning of S-OFF Midget Ganglion Cells in the Primate Retina.

In the trichromatic primate retina, the "midget" retinal ganglion cell is the classical substrate for red-green color signaling, with a circuitry that enables antagonistic responses between long (L)- and medium (M)-wavelength-sensitive cone inputs. Previous physiological studies showed that some OFF midget ganglion cells may receive sparse input from short (S)-wavelength-sensitive cones, but the effect of S-cone inputs on the chromatic tuning properties of such cells has not been explored. Moreover, anatomical evidence for a synaptic pathway from S cones to OFF midget ganglion cells through OFF midget bipolar cells remains ambiguous. In this study, we address both questions for the macaque monkey retina. First, we used serial block-face electron microscopy to show that every S cone in the parafoveal retina synapses principally with a single OFF midget bipolar cell, which in turn forms a private-line connection with an OFF midget ganglion cell. Second, we used patch electrophysiology to characterize the chromatic tuning of OFF midget ganglion cells in the near peripheral retina that receive combined input from L, M, and S cones. These "S-OFF" midget cells have a characteristic S-cone spatial signature, but demonstrate heterogeneous color properties due to the variable strength of L, M, and S cone input across the receptive field. Together, these findings strongly support the hypothesis that the OFF midget pathway is the major conduit for S-OFF signals in primate retina and redefines the pathway as a chromatically complex substrate that encodes color signals beyond the classically recognized L versus M and S versus L+M cardinal mechanisms.SIGNIFICANCE STATEMENT The first step of color processing in the visual pathway of primates occurs when signals from short (S)-, middle (M)-, and long (L)-wavelength-sensitive cone types interact antagonistically within the retinal circuitry to create color-opponent pathways. The midget (L versus M or "red-green") and small bistratified (S vs L+M, or "blue-yellow") ganglion cell pathways appear to provide the physiological origin of the cardinal axes of human color vision. Here we confirm the presence of an additional S-OFF midget circuit in the macaque monkey fovea with scanning block-face electron microscopy and show physiologically that a subpopulation of S-OFF midget cells combine S, L, and M cone inputs along noncardinal directions of color space, expanding the retinal role in color coding.

Amblyopia Affects the ON Visual Pathway More than the OFF.

Visual information reaches the cerebral cortex through parallel ON and OFF pathways that signal the presence of light and dark stimuli in visual scenes. We have previously demonstrated that optical blur reduces visual salience more for light than dark stimuli because it removes the high spatial frequencies from the stimulus, and low spatial frequencies drive weaker ON than OFF cortical responses. Therefore, we hypothesized that sustained optical blur during brain development should weaken ON cortical pathways more than OFF, increasing the dominance of darks in visual perception. Here we provide support for this hypothesis in humans with anisometropic amblyopia who suffered sustained optical blur early after birth in one of the eyes. In addition, we show that the dark dominance in visual perception also increases in strabismic amblyopes that have their vision to high spatial frequencies reduced by mechanisms not associated with optical blur. Together, we show that amblyopia increases visual dark dominance by 3-10 times and that the increase in dark dominance is strongly correlated with amblyopia severity. These results can be replicated with a computational model that uses greater luminance/response saturation in ON than OFF pathways and, as a consequence, reduces more ON than OFF cortical responses to stimuli with low spatial frequencies. We conclude that amblyopia affects the ON cortical pathway more than the OFF, a finding that could have implications for future amblyopia treatments.SIGNIFICANCE STATEMENT Amblyopia is a loss of vision that affects 2-5% of children across the world and originates from a deficit in visual cortical circuitry. Current models assume that amblyopia affects similarly ON and OFF visual pathways, which signal light and dark features in visual scenes. Against this current belief, here we demonstrate that amblyopia affects the ON visual pathway more than the OFF, a finding that could have implications for new amblyopia treatments targeted at strengthening a weak ON visual pathway.

Cortical Balance Between ON and OFF Visual Responses Is Modulated by the Spatial Properties of the Visual Stimulus.

The primary visual cortex of carnivores and primates is dominated by the OFF visual pathway and responds more strongly to dark than light stimuli. Here, we demonstrate that this cortical OFF dominance is modulated by the size and spatial frequency of the stimulus in awake primates and we uncover a main neuronal mechanism underlying this modulation. We show that large grating patterns with low spatial frequencies drive five times more OFF-dominated than ON-dominated neurons, but this pronounced cortical OFF dominance is strongly reduced when the grating size decreases and the spatial frequency increases, as when the stimulus moves away from the observer. We demonstrate that the reduction in cortical OFF dominance is not caused by a selective reduction of visual responses in OFF-dominated neurons but by a change in the ON/OFF response balance of neurons with diverse receptive field properties that can be ON or OFF dominated, simple, or complex. We conclude that cortical OFF dominance is continuously adjusted by a neuronal mechanism that modulates ON/OFF response balance in multiple cortical neurons when the spatial properties of the visual stimulus change with viewing distance and/or optical blur.

Measurement Techniques and Utility of Hounsfield Unit Values for Assessment of Bone Quality Prior to Spinal Instrumentation: A Review of Current Literature.

STUDY DESIGN: A systematic review. OBJECTIVE: The aim of this study was to evaluate the clinical utility of assessing bone quality using computed tomography (CT) attenuation in Hounsfield units (HU). SUMMARY OF BACKGROUND DATA: Assessing bone quality before spine instrumentation is an essential step of preoperative planning. Dual energy x-ray absorptiometry (DEXA) has been the gold standard for assessing bone mineral density (BMD); however, DEXA can result in spuriously elevated BMD measurements in patients with degenerative disease, compression fractures, and/or vascular calcifications. Measuring vertebral HU values has been proposed as an alternate method of assessing BMD and bone quality. METHODS: We searched MEDLINE, Cochrane Library, and EMBASE for studies correlating HU to BMD and to spine surgery outcomes. RESULTS: HU measurements correlate with success of lumbar interbody fusion (133.7 vs. 107.3 HU) and posterolateral fusion (167 vs. 139.8 HU), cage subsidence (112.4 vs. 140.2 HU), adjacent segment fractures (145.6 vs. 199.4 HU), pedicle screw loosening (116.4 vs. 132.7 HU) and (99.1 vs. 141.2 HU), and risk of incidental durotomy (149.2 vs. 177.0 HU). Intra and inter-rater reliability coefficients are 0.964 and 0.975. The correlation between HU values and BMD in nondegenerative patients is r = 0.52 compared with HU versus BMD in degenerative patients r = 0.18. CONCLUSION: HU value measurement is a simple and rapid technique to assess bone quality that should be performed in all patients with pre-existing CT scans. HU measurement has excellent inter and intra-rater reliability and can be performed on axial or sagittal images. L1 HU threshold values of 110 HU for detecting osteoporosis, and 135 HU for detecting osteopenia are 90% specific. In patients with significant degenerative disease, HU values should be given more credence. Additional high-quality prospective studies comparing HU and DEXA values to patient outcomes are necessary to validate the role of CT measurements in preoperative evaluation. LEVEL OF EVIDENCE: 4.

Motion changes response balance between ON and OFF visual pathways.

Humans are faster at detecting dark than light stationary stimuli, a temporal difference that originates early in the visual pathway. Here we show that this difference reverses when stimuli move, making detection faster for moving lights than darks. Human subjects judged the direction of moving edges and bars, and made faster and more accurate responses for light than for dark stimuli. This light/dark asymmetry is greatest at low speeds and disappears at high speeds. In parallel experiments, we recorded responses in the cat visual cortex for moving bars and again find that responses are faster for light bars than for dark bars moving at low speeds. We show that differences in the luminance-response function between ON and OFF pathways can reproduce these findings, and may explain why ON pathways are used for slow-motion image stabilization in many species.

Picture perception reveals mental geometry of 3D scene inferences.

Pose estimation of objects in real scenes is critically important for biological and machine visual systems, but little is known of how humans infer 3D poses from 2D retinal images. We show unexpectedly remarkable agreement in the 3D poses different observers estimate from pictures. We further show that all observers apply the same inferential rule from all viewpoints, utilizing the geometrically derived back-transform from retinal images to actual 3D scenes. Pose estimations are altered by a fronto-parallel bias, and by image distortions that appear to tilt the ground plane. We used pictures of single sticks or pairs of joined sticks taken from different camera angles. Observers viewed these from five directions, and matched the perceived pose of each stick by rotating an arrow on a horizontal touchscreen. The projection of each 3D stick to the 2D picture, and then onto the retina, is described by an invertible trigonometric expression. The inverted expression yields the back-projection for each object pose, camera elevation, and observer viewpoint. We show that a model that uses the back-projection, modulated by just two free parameters, explains 560 pose estimates per observer. By considering changes in retinal image orientations due to position and elevation of limbs, the model also explains perceived limb poses in a complex scene of two bodies lying on the ground. The inferential rules simply explain both perceptual invariance and dramatic distortions in poses of real and pictured objects, and show the benefits of incorporating projective geometry of light into mental inferences about 3D scenes.

Nonselective Wiring Accounts for Red-Green Opponency in Midget Ganglion Cells of the Primate Retina.

In primate retina, "red-green" color coding is initiated when signals originating in long (L) and middle (M) wavelength-sensitive cone photoreceptors interact antagonistically. The center-surround receptive field of "midget" ganglion cells provides the neural substrate for L versus M cone-opponent interaction, but the underlying circuitry remains unsettled, centering around the longstanding question of whether specialized cone wiring is present. To address this question, we measured the strength, sign, and spatial tuning of L- and M-cone input to midget receptive fields in the peripheral retina of macaque primates of either sex. Consistent with previous work, cone opponency arose when one of the cone types showed a stronger connection to the receptive field center than to the surround. We implemented a difference-of-Gaussians spatial receptive field model, incorporating known biology of the midget circuit, to test whether physiological responses we observed in real cells could be captured entirely by anatomical nonselectivity. When this model sampled nonselectively from a realistic cone mosaic, it accurately reproduced key features of a cone-opponent receptive field structure, and predicted both the variability and strength of cone opponency across the retina. The model introduced here is consistent with abundant anatomical evidence for nonselective wiring, explains both local and global properties of the midget population, and supports a role in their multiplexing of spatial and color information. It provides a neural basis for human chromatic sensitivity across the visual field, as well as the maintenance of normal color vision despite significant variability in the relative number of L and M cones across individuals.SIGNIFICANCE STATEMENT Red-green color vision is a hallmark of the human and nonhuman primate that starts in the retina with the presence of long (L)- and middle (M)-wavelength sensitive cone photoreceptor types. Understanding the underlying retinal mechanism for color opponency has focused on the broad question of whether this characteristic can emerge from nonselective wiring, or whether complex cone-type-specific wiring must be invoked. We provide experimental and modeling support for the hypothesis that nonselective connectivity is sufficient to produce the range of red-green color opponency observed in midget ganglion cells across the retina. Our nonselective model reproduces the diversity of physiological responses of midget cells while also accounting for systematic changes in color sensitivity across the visual field.