Spinal fusion surgery use among adults with low back pain enrolled in a digital musculoskeletal program: an observational study.

OBJECTIVES: To compare 12-month spinal fusion surgery rates in the setting of low back pain among digital musculoskeletal (MSK) program participants versus a comparison cohort who only received usual care. STUDY DESIGN: Retrospective cohort study with propensity score matched comparison cohort using commercial medical claims data representing over 100 million commercially insured lives. METHODS: All study subjects experienced low back pain between January 2020 and December 2021. Digital MSK participants enrolled in the digital MSK low back program between January 2020 and December 2021. Non-participants had low back pain related physical therapy (PT) between January 2020 and December 2021. Digital MSK participants were matched to non-participants with similar demographics, comorbidities and baseline MSK-related medical care use. Spinal fusion surgery rates at 12 months post participation were compared. RESULTS: Compared to non-participants, digital MSK participants had lower rates of spinal fusion surgery in the post-period (0.7% versus 1.6%; p < 0.001). Additionally, in the augmented inverse probability weighting (AIPW) model, digital MSK participants were found to have decreased odds of undergoing spinal fusion surgery (adjusted odds ratio: 0.64, 95% CI: 0.51-0.81). CONCLUSIONS: This study provides evidence that participation in a digital MSK program is associated with a lower rate of spinal fusion surgery.

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 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.

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.

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.

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.

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.

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.

Neuronal nonlinearity explains greater visual spatial resolution for darks than lights.

Astronomers and physicists noticed centuries ago that visual spatial resolution is higher for dark than light stimuli, but the neuronal mechanisms for this perceptual asymmetry remain unknown. Here we demonstrate that the asymmetry is caused by a neuronal nonlinearity in the early visual pathway. We show that neurons driven by darks (OFF neurons) increase their responses roughly linearly with luminance decrements, independent of the background luminance. However, neurons driven by lights (ON neurons) saturate their responses with small increases in luminance and need bright backgrounds to approach the linearity of OFF neurons. We show that, as a consequence of this difference in linearity, receptive fields are larger in ON than OFF thalamic neurons, and cortical neurons are more strongly driven by darks than lights at low spatial frequencies. This ON/OFF asymmetry in linearity could be demonstrated in the visual cortex of cats, monkeys, and humans and in the cat visual thalamus. Furthermore, in the cat visual thalamus, we show that the neuronal nonlinearity is present at the ON receptive field center of ON-center neurons and ON receptive field surround of OFF-center neurons, suggesting an origin at the level of the photoreceptor. These results demonstrate a fundamental difference in visual processing between ON and OFF channels and reveal a competitive advantage for OFF neurons over ON neurons at low spatial frequencies, which could be important during cortical development when retinal images are blurred by immature optics in infant eyes.

Neuronal mechanisms underlying differences in spatial resolution between darks and lights in human vision.

Artists and astronomers noticed centuries ago that humans perceive dark features in an image differently from light ones; however, the neuronal mechanisms underlying these dark/light asymmetries remained unknown. Based on computational modeling of neuronal responses, we have previously proposed that such perceptual dark/light asymmetries originate from a luminance/response saturation within the ON retinal pathway. Consistent with this prediction, here we show that stimulus conditions that increase ON luminance/response saturation (e.g., dark backgrounds) or its effect on light stimuli (e.g., optical blur) impair the perceptual discrimination and salience of light targets more than dark targets in human vision. We also show that, in cat visual cortex, the magnitude of the ON luminance/response saturation remains relatively constant under a wide range of luminance conditions that are common indoors, and only shifts away from the lowest luminance contrasts under low mesopic light. Finally, we show that the ON luminance/response saturation affects visual salience mostly when the high spatial frequencies of the image are reduced by poor illumination or optical blur. Because both low luminance and optical blur are risk factors in myopia, our results suggest a possible neuronal mechanism linking myopia progression with the function of the ON visual pathway.

Chromatic and Achromatic Spatial Resolution of Local Field Potentials in Awake Cortex.

Local field potentials (LFPs) have become an important measure of neuronal population activity in the brain and could provide robust signals to guide the implant of visual cortical prosthesis in the future. However, it remains unclear whether LFPs can detect weak cortical responses (e.g., cortical responses to equiluminant color) and whether they have enough visual spatial resolution to distinguish different chromatic and achromatic stimulus patterns. By recording from awake behaving macaques in primary visual cortex, here we demonstrate that LFPs respond robustly to pure chromatic stimuli and exhibit ∼2.5 times lower spatial resolution for chromatic than achromatic stimulus patterns, a value that resembles the ratio of achromatic/chromatic resolution measured with psychophysical experiments in humans. We also show that, although the spatial resolution of LFP decays with visual eccentricity as is also the case for single neurons, LFPs have higher spatial resolution and show weaker response suppression to low spatial frequencies than spiking multiunit activity. These results indicate that LFP recordings are an excellent approach to measure spatial resolution from local populations of neurons in visual cortex including those responsive to color.

Properties of lateral interaction in color and brightness induction.

In a visual scene, when objects are surrounded by other components, neural mechanisms increase the perceived color and brightness difference between an object and its surround, potentially enhancing an observer's ability to segment objects. Despite almost two centuries of empirical investigations, the nature of induction mechanisms remains elusive. To elucidate the nature of these mechanisms, we introduce a new method for measuring color and brightness induction that allows separate manipulation of lateral interactions and adaptation, and controls for eye-movement-related effects. We use the method to examine the function relating induction magnitude to contrast change in the surround, the symmetry of induction in complementary directions for the three cardinal color axes, and the effect of blur between the test and the surround. On average, brightness induction was more linear than chromatic induction. The induction magnitude was similar for surrounds of complementary colors on average and for many conditions, and when individual observers deviated from symmetry it could be on either side. Edge blur did not change the induction magnitude. For slower presentations, light/dark induction increased to further reduce asymmetry, suggesting that previously found light/dark induction asymmetry is not due to lateral interactions or prolonged adaptation. Lateral interactions underlying induction are thus mostly symmetric for color and brightness axes and involve spatially opponent filters of modest widths, rather than edge extraction.

Segregating animals in naturalistic surroundings: interaction of color distributions and mechanisms.

Humans have been shown to rapidly detect animals in naturalistic scenes, but the role of color in this task is unclear. We first analyze the color information contained in a large number of images of salient and camouflaged animals in generic backgrounds. We found that color distributions of most animals and of their immediate backgrounds were oriented along other than the cardinal directions of color space. In addition, the maximum distances between animals and background distributions also tended to be along noncardinal directions, suggesting a role for higher-order cortical color mechanisms whose preferred axes are distributed widely in color space. We measured temporal thresholds for segmenting animal color distributions from background distributions in the absence of spatial cues. Combined over all observers and all images in our sample, thresholds for segmenting isoluminant projections of these distributions were lower than for segmenting the original distributions and considerably lower than for segmenting achromatic projections. Color information is thus likely to be useful in segregating animals in generic views, i.e., views not purposely chosen by the photographer to enhance the visibility of the animal. However, a comparison of thresholds with distances between distributions failed to reveal any advantage conferred by higher-order color mechanisms.

Eye movements and the neural basis of context effects on visual sensitivity.

The effects of context on visual sensitivity are well established (e.g., sensitivity to luminance flicker is substantially higher on mean-gray surrounds than on white or black surrounds). The neural mechanisms generating context effects, however, remain unresolved. In the absence of direct tests, some theories invoke enhancement of edges by lateral inhibition, whereas others rely on transients caused by miniature eye movements that maintain fixation. We first replicated the luminance results on human observers and found unexpectedly that sensitivity to red-green flicker is also affected by surround color, being substantially higher on mean-gray surrounds than on red or green surrounds. To identify the neural bases of both context effects, we used in vivo electrophysiological recordings of primate magnocellular and parvocellular ganglion cell responses to luminance and red-green modulations, respectively. To test neuronal sensitivity to stationary edge contrast, neuronal responses were measured at various distances from the modulation edge against various surrounds. We found no evidence of enhanced responses to stationary edges on any surrounds, ruling out lateral inhibition-type explanations. To simulate the effects of eye movements, target patches were abruptly displaced while measuring responses. Abruptly displaced edges evoked vigorous transient responses that were selective for modulation-phase on mean-gray surrounds, but were phase-invariant on other surrounds. Eye movements could thus enhance detection of flicker on mean-gray surrounds, and neurometric analyses supported a primary role for eye movements in enhancing sensitivity. In addition, the transformation of spatial edges to transient neuronal responses by eye movements provides the signals for detecting luminance and color edges in natural scenes.

Salience of unique hues and implications for color theory.

The unique hues--blue, green, yellow, red--form the fundamental dimensions of opponent-color theories, are considered universal across languages, and provide useful mental representations for structuring color percepts. However, there is no neural evidence for them from neurophysiology or low-level psychophysics. Tapping a higher prelinguistic perceptual level, we tested whether unique hues are particularly salient in search tasks. We found no advantage for unique hues over their nonunique complementary colors. However, yellowish targets were detected faster, more accurately, and with fewer saccades than their complementary bluish targets (including unique blue), while reddish-greenish pairs were not significantly different in salience. Similarly, local field potentials in primate V1 exhibited larger amplitudes and shorter latencies for yellowish versus bluish stimuli, whereas this effect was weaker for reddish versus greenish stimuli. Consequently, color salience is affected more by early neural response asymmetries than by any possible mental or neural representation of unique hues.