Stages Of Cortical Processing Research Articles

Informational Masking Effects of Similarity and Uncertainty on Early and Late Stages of Auditory Cortical Processing

Understanding speech in a background of other people talking is a difficult listening situation for hearing-impaired individuals, and even for those with normal hearing. Speech-on-speech masking is known to contribute to increased perceptual difficulty over nonspeech background noise because of informational masking provided over and above the effects of energetic masking. While informational masking research has identified factors of similarity and uncertainty between target and masker that contribute to reduced behavioral performance in speech background noise, critical gaps in knowledge including the underlying neural-perceptual processes remain. By systematically manipulating aspects of acoustic similarity and uncertainty in the same auditory paradigm, the current study examined the time course and objectively quantified these informational masking effects at both early and late stages of auditory processing using auditory evoked potentials (AEPs). Thirty participants were included in a cross-sectional repeated measures design. Target-masker similarity was manipulated by varying the linguistic/phonetic similarity (i.e., language) of the talkers in the background. Specifically, four levels representing hypothesized increasing levels of informational masking were implemented: (1) no masker (quiet); (2) Mandarin; (3) Dutch; and (4) English. Stimulus uncertainty was manipulated by task complexity, specifically presentation of target-to-target interval (TTI) in the auditory evoked paradigm. Participants had to discriminate between English word stimuli (/bæt/ and /pæt/) presented in an oddball paradigm under each masker condition pressing buttons to either the target or standard stimulus. Responses were recorded simultaneously for P1-N1-P2 (standard waveform) and P3 (target waveform). This design allowed for simultaneous recording of multiple AEP peaks, as well as accuracy, reaction time, and d' behavioral discrimination to button press responses. Several trends in AEP components were consistent with effects of increasing linguistic/phonetic similarity and stimulus uncertainty. All babble maskers significantly affected outcomes compared to quiet. In addition, the native language English masker had the largest effect on outcomes in the AEP paradigm, including reduced P3 amplitude and area, as well as decreased accuracy and d' behavioral discrimination to target word responses. AEP outcomes for the Mandarin and Dutch maskers, however, were not significantly different across any measured component. Latency outcomes for both N1 and P3 also supported an effect of stimulus uncertainty, consistent with increased processing time related to greater task complexity. An unanticipated result was the absence of the interaction of linguistic/phonetic similarity and stimulus uncertainty. Observable effects of both similarity and uncertainty were evidenced at a level of the P3 more than the earlier N1 level of auditory cortical processing suggesting that higher-level active auditory processing may be more sensitive to informational masking deficits. The lack of significant interaction between similarity and uncertainty at either level of processing suggests that these informational masking factors operated independently. Speech babble maskers across languages altered AEP component measures, behavioral detection, and reaction time. Specifically, this occurred when the babble was in the native/same language as the target, while the effects of foreign language maskers did not differ. The objective results from this study provide a foundation for further investigation of how the linguistic content of target and masker and task difficulty contribute to difficulty understanding speech-in-noise.

Neurophysiology: The Three-Dimensional Building Blocks of Object Vision

A new study of the macaque visual cortex has revealed that visual area V4 performs substantial analysis of solid shape structure. The findings draw new attention to the embedding of local three-dimensional shape analysis into the early cortical stages of visual processing.

Curvature domains in V4 of macaque monkey.

An important aspect of visual object recognition is the ability to perceive object shape. Two basic components of complex shapes are straight and curved contours. A large body of evidence suggests a modular hierarchy for shape representation progressing from simple and complex orientation in early areas V1 and V2, to increasingly complex stages of curvature representation in V4, TEO, and TE. Here, we reinforce and extend the concept of modular representation. Using intrinsic signal optical imaging in Macaque area V4, we find sub-millimeter sized modules for curvature representation that are organized from low to high curvatures as well as domains with complex curvature preference. We propose a possible 'curvature hypercolumn' within V4. In combination with previous studies, we suggest that the key emergent functions at each stage of cortical processing are represented in systematic, modular maps.

Task-related activity in human visual cortex.

The brain exhibits widespread endogenous responses in the absence of visual stimuli, even at the earliest stages of visual cortical processing. Such responses have been studied in monkeys using optical imaging with a limited field of view over visual cortex. Here, we used functional MRI (fMRI) in human participants to study the link between arousal and endogenous responses in visual cortex. The response that we observed was tightly entrained to task timing, was spatially extensive, and was independent of visual stimulation. We found that this response follows dynamics similar to that of pupil size and heart rate, suggesting that task-related activity is related to arousal. Finally, we found that higher reward increased response amplitude while decreasing its trial-to-trial variability (i.e., the noise). Computational simulations suggest that increased temporal precision underlies both of these observations. Our findings are consistent with optical imaging studies in monkeys and support the notion that arousal increases precision of neural activity.

Breaking down the cocktail party: Attentional modulation of cerebral audiovisual speech processing

Recent studies utilizing electrophysiological speech envelope reconstruction have sparked renewed interest in the cocktail party effect by showing that auditory neurons entrain to selectively attended speech. Yet, the neural networks of attention to speech in naturalistic audiovisual settings with multiple sound sources remain poorly understood. We collected functional brain imaging data while participants viewed audiovisual video clips of lifelike dialogues with concurrent distracting speech in the background. Dialogues were presented in a full-factorial design, comprising task (listen to the dialogues vs. ignore them), audiovisual quality and semantic predictability. We used univariate analyses in combination with multivariate pattern analysis (MVPA) to study modulations of brain activity related to attentive processing of audiovisual speech. We found attentive speech processing to cause distinct spatiotemporal modulation profiles in distributed cortical areas including sensory and frontal-control networks. Semantic coherence modulated attention-related activation patterns in the earliest stages of auditory cortical processing, suggesting that the auditory cortex is involved in high-level speech processing. Our results corroborate views that emphasize the dynamic nature of attention, with task-specificity and context as cornerstones of the underlying neuro-cognitive mechanisms.

State-Dependent Changes in Perception and Coding in the Mouse Somatosensory Cortex

An animal's behavioral state is reflected in the dynamics of cortical population activity and its capacity to process sensory information. To better understand the relationship between behavioral states and information processing, mice are trained to detect varying amplitudes of whisker-deflection under two-photon calcium imaging. Layer 2/3 neurons in the vibrissal primary somatosensory cortex are imaged across different behavioral states, defined based on detection performance (low to high-state) and pupil diameter. The neurometric curve in each behavioral state mirrors the corresponding psychometric performance, with calcium signals predictive of the animal's choice. High behavioral states are associated with lower network synchrony, extending over shorter cortical distances. The decrease in correlation across neurons in high state results in enhanced information transmission capacity at the population level. The observed state-dependent changes suggest that the coding regime within the first stage of cortical processing may underlie adaptive routing of relevant information through the sensorimotor system.

The time‐course of speech perception revealed by temporally‐sensitive neural measures

Recent advances in cognitive neuroscience have provided a detailed picture of the early time-course of speech perception. In this review, we highlight this work, placing it within the broader context of research on the neurobiology of speech processing, and discuss how these data point us toward new models of speech perception and spoken language comprehension. We focus, in particular, on temporally-sensitive measures that allow us to directly measure early perceptual processes. Overall, the data provide support for two key principles: (a) speech perception is based on gradient representations of speech sounds and (b) speech perception is interactive and receives input from higher-level linguistic context at the earliest stages of cortical processing. Implications for models of speech processing and the neurobiology of language more broadly are discussed. This article is categorized under: Psychology > Language Psychology > Perception and Psychophysics Neuroscience > Cognition.

Neural representations of perceptual color experience in the human ventral visual pathway

Color is a perceptual construct that arises from neural processing in hierarchically organized cortical visual areas. Previous research, however, often failed to distinguish between neural responses driven by stimulus chromaticity versus perceptual color experience. An unsolved question is whether the neural responses at each stage of cortical processing represent a physical stimulus or a color we see. The present study dissociated the perceptual domain of color experience from the physical domain of chromatic stimulation at each stage of cortical processing by using a switch rivalry paradigm that caused the color percept to vary over time without changing the retinal stimulation. Using functional MRI (fMRI) and a model-based encoding approach, we found that neural representations in higher visual areas, such as V4 and VO1, corresponded to the perceived color, whereas responses in early visual areas V1 and V2 were modulated by the chromatic light stimulus rather than color perception. Our findings support a transition in the ascending human ventral visual pathway, from a representation of the chromatic stimulus at the retina in early visual areas to responses that correspond to perceptually experienced colors in higher visual areas.

Poor early cortical differentiation of speech predicts perceptual difficulties of severely hearing-impaired listeners in multi-talker environments

Hearing impairment disrupts processes of selective attention that help listeners attend to one sound source over competing sounds in the environment. Hearing prostheses (hearing aids and cochlear implants, CIs), do not fully remedy these issues. In normal hearing, mechanisms of selective attention arise through the facilitation and suppression of neural activity that represents sound sources. However, it is unclear how hearing impairment affects these neural processes, which is key to understanding why listening difficulty remains. Here, severely-impaired listeners treated with a CI, and age-matched normal-hearing controls, attended to one of two identical but spatially separated talkers while multichannel EEG was recorded. Whereas neural representations of attended and ignored speech were differentiated at early (~ 150 ms) cortical processing stages in controls, differentiation of talker representations only occurred later (~250 ms) in CI users. CI users, but not controls, also showed evidence for spatial suppression of the ignored talker through lateralized alpha (7–14 Hz) oscillations. However, CI users’ perceptual performance was only predicted by early-stage talker differentiation. We conclude that multi-talker listening difficulty remains for impaired listeners due to deficits in early-stage separation of cortical speech representations, despite neural evidence that they use spatial information to guide selective attention.

Unreliable feedback deteriorates information processing in primary visual cortex

It is well-established that increased sensory uncertainty impairs perceptual decision-making and leads to degraded neural stimulus representations. Recently, we also showed that providing unreliable feedback to choices leads to changes in perceptual decision-making similar to those of increased stimulus noise: A deterioration in objective task performance, a decrease in subjective confidence and a lower reliance on sensory information for perceptual inference. To investigate the neural basis of such feedback-based changes in perceptual decision-making, in the present study, two groups of healthy human participants (n = 15 each) performed a challenging visual orientation discrimination task while undergoing functional magnetic resonance imaging (fMRI). Critically, one group received reliable feedback regarding their task performance in an intervention phase, whereas the other group correspondingly received unreliable feedback - thereby keeping stimulus information constant. The effects of feedback reliability on performance and stimulus representation in the primary visual cortex (V1) were studied by comparing the pre- and post-intervention test phases between the groups. Compared to participants who received reliable feedback, those receiving unreliable feedback showed a decline in task performance that was paralleled by reduced distinctness of fMRI response patterns in V1. These results show that environmental uncertainty can affect perceptual inference at the earliest cortical processing stages.

Stereotactic electroencephalography in humans reveals multisensory signal in early visual and auditory cortices

Unequivocally demonstrating the presence of multisensory signals at the earliest stages of cortical processing remains challenging in humans. In our study, we relied on the unique spatio-temporal resolution provided by intracranial stereotactic electroencephalographic (SEEG) recordings in patients with drug-resistant epilepsy to characterize the signal extracted from early visual (calcarine and pericalcarine) and auditory (Heschl's gyrus and planum temporale) regions during a simple audio-visual oddball task. We provide evidences that both cross-modal responses (visual responses in auditory cortex or the reverse) and multisensory processing (alteration of the unimodal responses during bimodal stimulation) can be observed in intracranial event-related potentials (iERPs) and in power modulations of oscillatory activity at different temporal scales within the first 150 msec after stimulus onset. The temporal profiles of the iERPs are compatible with the hypothesis that MSI occurs by means of direct pathways linking early visual and auditory regions. Our data indicate, moreover, that MSI mainly relies on modulations of the low-frequency bands (foremost the theta band in the auditory cortex and the alpha band in the visual cortex), suggesting the involvement of feedback pathways between the two sensory regions. Remarkably, we also observed high-gamma power modulations by sounds in the early visual cortex, thus suggesting the presence of neuronal populations involved in auditory processing in the calcarine and pericalcarine region in humans.

Tactile Cross-Modal Acceleration Effects on Auditory Steady-State Response.

In the sensory cortex, cross-modal interaction occurs during the early cortical stages of processing; however, its effect on the speed of neuronal activity remains unclear. In this study, we used magnetoencephalography (MEG) to investigate whether tactile stimulation influences auditory steady-state responses (ASSRs). To this end, a 0.5-ms electrical pulse was randomly presented to the dorsum of the left or right hand of 12 healthy volunteers at 700 ms while a train of 25-ms pure tones were applied to the left or right side at 75 dB for 1,200 ms. Peak latencies of 40-Hz ASSR were measured. Our results indicated that tactile stimulation significantly shortened subsequent ASSR latency. This cross-modal effect was observed from approximately 50 ms to 125 ms after the onset of tactile stimulation. The somatosensory information that appeared to converge on the auditory system may have arisen during the early processing stages, with the reduced ASSR latency indicating that a new sensory event from the cross-modal inputs served to increase the speed of ongoing sensory processing. Collectively, our findings indicate that ASSR latency changes are a sensitive index of accelerated processing.

Mapping adaptation, deviance detection, and prediction error in auditory processing

Various studies have suggested that auditory deviance detection is organized in a hierarchical manner with ascending levels of complexity. Event-related potentials (ERP) are considered to reflect different cortical processing stages. In the current electroencephalographic study, we employed an auditory sequence oddball paradigm to investigate different levels of cortical auditory processing and the contribution of neuronal habituation and prediction error mechanism to N1 and Mismatch Negativity (MMN). Our findings suggest that N1 reflects a lower cortical process primarily involved in the encoding of simple physical features and is thus mainly modulated by neuronal attenuation and not complex top-down mechanisms. By analyzing within-sequence signal differences, we divided the MMN into distinct subcomponents reflecting different hierachical levels of auditory processing. We determined a “first-order” MMN that reflects the processing of simple deviant features (such as frequency) and “higher-order” MMNs that occur at regularity violation of complex patterns or unexpected inputs that do not allow further predictions. In our source localization analysis, both the primary auditory cortex and left IFG were primarily involved in the detection of simple, physically deviant features, while the right IFG was associated with the processing of novel, unexpected auditory inputs and the ACC with regularity violation of known patterns. Summarizing, our results might contribute to a better understanding of the different complexities of neuronal habituation and prediction error mechanisms at different levels of cortical auditory processing.

Somatosensory Cortex Efficiently Processes Touch Located Beyond the Body.

The extent to which a tool is an extension of its user is a question that has fascinated writers and philosophers for centuries [1]. Despite two decades of research [2-7], it remains unknown how this could be instantiated at the neural level. To this aim, the present study combined behavior, electrophysiology and neuronal modeling to characterize how the human brain could treat a tool like an extended sensory "organ." As with the body, participants localize touches on a hand-held tool with near-perfect accuracy [7]. This behavior is owed to the ability of the somatosensory system to rapidly and efficiently use the tool as a tactile extension of the body. Using electroencephalography (EEG), we found that where a hand-held tool was touched was immediately coded in the neural dynamics of primary somatosensory and posterior parietal cortices of healthy participants. We found similar neural responses in a proprioceptively deafferented patient with spared touch perception, suggesting that location information is extracted from the rod's vibrational patterns. Simulations of mechanoreceptor responses [8] suggested that the speed at which these patterns are processed is highly efficient. A second EEG experiment showed that touches on the tool and arm surfaces were localized by similar stages of cortical processing. Multivariate decoding algorithms and cortical source reconstruction provided further evidence that early limb-based processes were repurposed to map touch on a tool. We propose that an elementary strategy the human brain uses to sense with tools is to recruit primary somatosensory dynamics otherwise devoted to the body.

Representational untangling by the firing rate nonlinearity in V1 simple cells.

An important computational goal of the visual system is 'representational untangling' (RU): representing increasingly complex features of visual scenes in an easily decodable format. RU is typically assumed to be achieved in high-level visual cortices via several stages of cortical processing. Here we show, using a canonical population coding model, that RU of low-level orientation information is already performed at the first cortical stage of visual processing, but not before that, by a fundamental cellular-level property: the thresholded firing rate nonlinearity of simple cells in the primary visual cortex (V1). We identified specific, experimentally measurable parameters that determined the optimal firing threshold for RU and found that the thresholds of V1 simple cells extracted from in vivo recordings in awake behaving mice were near optimal. These results suggest that information re-formatting, rather than maximisation, may already be a relevant computational goal for the early visual system.

Bodily self-relatedness in vicarious touch is reflected at early cortical processing stages.

Studies have suggested that there is a strong link between the bodily self and the mechanisms underlying vicarious representations. Here, we used somatosensory ERPs to investigate the temporal dynamics of vicarious touch for stimuli that are more or less related to one's own body (human hands vs. rubber gloves). We found that vicarious touch effects were restricted to self-relatable events (human hands) at early implicit stages of somatosensory processing (P45). At later more cognitive stages of processing (late positive complex, LPC), the vicarious touch effect was stronger for self-relatable events (touch on human hands) than nonself-relatable events (touch on rubber gloves) but present for both. Both effects, but especially the vicarious touch effect for human hands at P45, were stronger in individuals with higher levels of interoceptive awareness. Our results confirm that there is a tight link between vicarious touch and the bodily self and characterize P45 effects of vicarious touch as its likely neural basis. We propose that vicarious processes and the embodied self may be representationally indistinct (linked in a common neural representation) at early implicit somatosensory processing stages.

Cortical encoding of speech enhances task-relevant acoustic information.

Speech is the most important signal in our auditory environment, and the processing of speech is highly dependent on context. However, it is unknown how contextual demands influence the neural encoding of speech. Here, we examine the context dependence of auditory cortical mechanisms for speech encoding at the level of the representation of fundamental acoustic features (spectrotemporal modulations) using model-based functional magnetic resonance imaging. We found that the performance of different tasks on identical speech sounds leads to neural enhancement of the acoustic features in the stimuli that are critically relevant to task performance. These task effects were observed at the earliest stages of auditory cortical processing, in line with interactive accounts of speech processing. Our work provides important insights into the mechanisms that underlie the processing of contextually relevant acoustic information within our rich and dynamic auditory environment.

Evolutionary Elongation of the Time Window of Integration in Auditory Cortex: Macaque vs. Human Comparison of the Effects of Sound Duration on Auditory Evoked Potentials.

The auditory cortex integrates auditory information over time to obtain neural representations of sound events, the time scale of which critically affects perception. This work investigated the species differences in the time scale of integration by comparing humans and monkeys regarding how their scalp-recorded cortical auditory evoked potentials (CAEPs) decrease in amplitude as stimulus duration is shortened from 100 ms (or longer) to 2 ms. Cortical circuits tuned to processing sounds at short time scales would continue to produce large CAEPs to brief sounds whereas those tuned to longer time scales would produce diminished responses. Four peaks were identified in the CAEPs and labeled P1, N1, P2, and N2 in humans and mP1, mN1, mP2, and mN2 in monkeys. In humans, the N1 diminished in amplitude as sound duration was decreased, consistent with the previously described temporal integration window of N1 (>50 ms). In macaques, by contrast, the mN1 was unaffected by sound duration, and it was clearly elicited by even the briefest sounds. Brief sounds also elicited significant mN2 in the macaque, but not the human N2. Regarding earlier latencies, both P1 (humans) and mP1 (macaques) were elicited at their full amplitudes even by the briefest sounds. These findings suggest an elongation of the time scale of late stages of human auditory cortical processing, as reflected by N1/mN1 and later CAEP components. Longer time scales of integration would allow neural representations of complex auditory features that characterize speech and music.

Topographic Organization of the 'Third-Tier' Dorsomedial Visual Cortex in the Macaque.

The boundaries of the visual areas located anterior to V2 in the dorsomedial region of the macaque cortex remain contentious. This region is usually conceptualized as including two functional subdivisions: the dorsal component of area V3 (V3d) laterally and another area named the parietooccipital area (PO) or V6 medially. However, the nature of the putative border between V3d and PO/V6 has remained undefined. We recorded the receptive fields of multiunit clusters in male macaques and reconstructed the locations of recording sites using histological sections and computer-generated maps. Immediately adjacent to dorsomedial V2, we observed a representation of the lower contralateral quadrant that represented the vertical meridian at its rostral border. This region formed a simple eccentricity gradient from ∼<5° in the annectant gyrus to >60° in the parietooccipital medial sulcus. There was no topographic reversal where one would expect to find the border between V3d and PO/V6. Rather, near the midline, this lower quadrant map continued directly into a representation of the peripheral upper visual field without an intervening lower quadrant representation. Therefore, cortex previously assigned to the medial part of V3d and to PO/V6 forms a single map that includes parts of both quadrants. Together with previous observations that V3d and PO/V6 are densely myelinated relative to adjacent cortex and share similar input from V1, these results suggest that they are parts of a single area (for which we suggest the designation V6), which is distinct from the one forming the ventral component of the third-tier complex.SIGNIFICANCE STATEMENT The primate visual cortex has a large number of areas. Knowing the extent of each visual area and how they can be distinguished from each other is essential for the interpretation of experiments aimed at understanding visual processing. Currently, there are conflicting models of the organization of the dorsomedial visual cortex rostral to area V2 (one of the earliest stages of cortical processing of vision). By conducting large-scale electrophysiological recordings, we found that what were originally thought to be distinct areas in this region (dorsal V3 and the parietooccipital area PO/V6), together form a single map of the visual field. This will help to guide future functional studies and the interpretation of the outcomes of lesions involving the dorsal visual cortex.

Using deep learning to probe the neural code for images in primary visual cortex.

Primary visual cortex (V1) is the first stage of cortical image processing, and major effort in systems neuroscience is devoted to understanding how it encodes information about visual stimuli. Within V1, many neurons respond selectively to edges of a given preferred orientation: These are known as either simple or complex cells. Other neurons respond to localized center–surround image features. Still others respond selectively to certain image stimuli, but the specific features that excite them are unknown. Moreover, even for the simple and complex cells—the best-understood V1 neurons—it is challenging to predict how they will respond to natural image stimuli. Thus, there are important gaps in our understanding of how V1 encodes images. To fill this gap, we trained deep convolutional neural networks to predict the firing rates of V1 neurons in response to natural image stimuli, and we find that the predicted firing rates are highly correlated (newcommand{bialpha}{boldsymbol{alpha}}newcommand{bibeta}{boldsymbol{beta}}newcommand{bigamma}{boldsymbol{gamma}}newcommand{bidelta}{boldsymbol{delta}}newcommand{bivarepsilon}{boldsymbol{varepsilon}}newcommand{bizeta}{boldsymbol{zeta}}newcommand{bieta}{boldsymbol{eta}}newcommand{bitheta}{boldsymbol{theta}}newcommand{biiota}{boldsymbol{iota}}newcommand{bikappa}{boldsymbol{kappa}}newcommand{bilambda}{boldsymbol{lambda}}newcommand{bimu}{boldsymbol{mu}}newcommand{binu}{boldsymbol{nu}}newcommand{bixi}{boldsymbol{xi}}newcommand{biomicron}{boldsymbol{micron}}newcommand{bipi}{boldsymbol{pi}}newcommand{birho}{boldsymbol{rho}}newcommand{bisigma}{boldsymbol{sigma}}newcommand{bitau}{boldsymbol{tau}}newcommand{biupsilon}{boldsymbol{upsilon}}newcommand{biphi}{boldsymbol{phi}}newcommand{bichi}{boldsymbol{chi}}newcommand{bipsi}{boldsymbol{psi}}newcommand{biomega}{boldsymbol{omega}}{overline {{bf{CC}}} _{{bf{norm}}}} = 0.556 ± 0.01) with the neurons' actual firing rates over a population of 355 neurons. This performance value is quoted for all neurons, with no selection filter. Performance is better for more active neurons: When evaluated only on neurons with mean firing rates above 5 Hz, our predictors achieve correlations of newcommand{bialpha}{boldsymbol{alpha}}newcommand{bibeta}{boldsymbol{beta}}newcommand{bigamma}{boldsymbol{gamma}}newcommand{bidelta}{boldsymbol{delta}}newcommand{bivarepsilon}{boldsymbol{varepsilon}}newcommand{bizeta}{boldsymbol{zeta}}newcommand{bieta}{boldsymbol{eta}}newcommand{bitheta}{boldsymbol{theta}}newcommand{biiota}{boldsymbol{iota}}newcommand{bikappa}{boldsymbol{kappa}}newcommand{bilambda}{boldsymbol{lambda}}newcommand{bimu}{boldsymbol{mu}}newcommand{binu}{boldsymbol{nu}}newcommand{bixi}{boldsymbol{xi}}newcommand{biomicron}{boldsymbol{micron}}newcommand{bipi}{boldsymbol{pi}}newcommand{birho}{boldsymbol{rho}}newcommand{bisigma}{boldsymbol{sigma}}newcommand{bitau}{boldsymbol{tau}}newcommand{biupsilon}{boldsymbol{upsilon}}newcommand{biphi}{boldsymbol{phi}}newcommand{bichi}{boldsymbol{chi}}newcommand{bipsi}{boldsymbol{psi}}newcommand{biomega}{boldsymbol{omega}}{overline {{bf{CC}}} _{{bf{norm}}}} = 0.69 ± 0.01 with the neurons' true firing rates. We find that the firing rates of both orientation-selective and non-orientation-selective neurons can be predicted with high accuracy. Additionally, we use a variety of models to benchmark performance and find that our convolutional neural-network model makes more accurate predictions.