First-order Stimuli Research Articles

Early Auditory Event Related Potentials Distinguish Higher-Order From First-Order Aversive Conditioning.

Stimuli in reality rarely co-occur with primary reward or punishment to allow direct associative learning of value. Instead, value is thought to be inferred through complex higher-order associations. Rodent research has demonstrated that the formation and maintenance of first-order and higher-order associations are supported by distinct neural substrates. In this study, we explored whether this pattern of findings held true for humans. Participants underwent first-order and subsequent higher-order conditioning using an aversive burst of white noise or neutral tone as the unconditioned stimuli. Four distinct tones, initially neutral, served as first-order and higher-order conditioned stimuli. Autonomic and neural responses were indexed by pupillometry and evoked response potentials (ERPs) respectively. Conditioned aversive values of first-order and higher-order stimuli led to increased autonomic responses, as indexed by pupil dilation. Distinct temporo-spatial auditory evoked response potentials were elicited by first-order and high-order conditioned stimuli. Conditioned first-order responses peaked around 260 ms and source estimation suggested a primary medial prefrontal and amygdala source. Conversely, conditioned higher-order responses peaked around 120 ms with an estimated source in the medial temporal lobe. Interestingly, pupillometry responses to first-order conditioned stimuli were diminished after higher order training, possibly signifying concomitant incidental extinction, while responses to higher-order stimuli remained. This suggests that once formed, higher order associations are at least partially independent of first order conditioned representations. This experiment demonstrates that first-order and higher-order conditioned associations have distinct neural signatures, and like rodents, the medial temporal lobe may be specifically involved with higher-order conditioning.

Mechanisms of higher-order learning in the amygdala

Adaptive behaviour is under the potent control of environmental cues. Such cues can acquire value by virtue of their associations with outcomes of motivational significance, be they appetitive or aversive. There are at least two ways through which an environmental cue can acquire value, through first-order and higher-order conditioning. In first-order conditioning, a neutral cue is directly paired with an outcome of motivational significance. In higher-order conditioning, a cue is indirectly associated with motivational events via a directly conditioned first-order stimulus. The present article reviews some of the associations that support learning in first- and higher-order conditioning, as well as the role of the BLA and the molecular mechanisms involved in these two types of learning.

Contrast-modulated stimuli produce more superimposition and predominate perception when competing with comparable luminance-modulated stimuli during interocular grouping

Interocular grouping (IOG) is a binocular visual function that can arise during multi-stable perception. IOG perception was initiated using split-grating stimuli constructed from luminance (L), luminance-modulated noise (LM) and contrast-modulated noise (CM). In Experiment 1, three different visibility levels were used for L and LM (or first-order) stimuli, and compared to fixed-visibility CM (or second-order) stimuli. Eight binocularly normal participants indicated whether they perceived full horizontal or vertical gratings, superimposition, or other (piecemeal and eye-of-origin) percepts. CM stimuli rarely generated full IOG, but predominantly generated superimposition. In Experiment 2, Levelt’s modified laws were tested for IOG in nine participants. Split-gratings presented to each eye contained different visibility LM gratings, or LM and CM gratings. The results for the LM-vs-LM conditions mostly followed the predictions of Levelt’s modified laws, whereas the results for the LM-vs-CM conditions did not. Counterintuitively, when high-visibility LM and low-visibility CM split-gratings were used, high-visibility LM components did not predominate IOG perception. Our findings suggest that higher proportions of superimposition during CM-vs-CM viewing are due to binocular combination, rather than mutual inhibition. It implies that IOG percepts are more likely to be mediated at an earlier monocular, rather than a binocular stage. Our previously proposed conceptual framework for conventional binocular rivalry, which includes asymmetric feedback, visual saliency, or a combination of both (Skerswetat et al. Sci Rep 8:14432, 2018), might also account for IOG. We speculate that opponency neurons might mediate coherent percepts when dissimilar information separately enters the eyes.

Clicker Training Accelerates Learning of Complex Behaviors but Reduces Discriminative Abilities of Yucatan Miniature Pigs

Animal training is meant to teach specific behavioral responses to specific cues. Clicker training (CT) is a popular training method based on the use of a device that emits a sound of double-click to be associated as a first-order conditioned stimulus in contingency with positive reinforcements. After some repetitions, the clicker sound gains some incentive value and can be paired with the desired behavior. Animal trainers believed that CT can decrease training time compared to other types of training. Herein, we used two-month old miniature piglets to evaluate whether CT decreased the number of repetitions required to learn complex behaviors as compared with animals trained with voice instead of the clicker. In addition, we compared the number of correct choices of animals from both groups when exposed to object discriminative tests. Results indicated that CT decreased the number of repetitions required for pigs to learn to fetch an object but reduced the ability of animals to make correct choices during the discriminate trials. This suggests that CT is more efficient than voice to teach complex behaviors but reduces the ability of animals to use cognitive processes required to discriminate and select objects associated with reward.

Examination of Lower Level Motion Mechanisms That Provide Information to Object Tracking: An Examination Using Dichoptic Stimulation.

In this study, we examined the operation of first- and second-order motion mechanisms with respect to object tracking using dichoptic presentation. A bistable apparent motion stimulus composed of four rectangles arranged in square- and diamond-shapes in every other frame was presented binocularly, monocularly, or dichoptically using a stereoscope. Since past motion studies showed that the first-order motion mechanism cannot function under dichoptic stimulation, we evaluated the upper temporal frequency limits of object tracking with dichoptic presentation and compared these results with those obtained with ordinary binocular or monocular (nondichoptic) presentation. We found that the temporal limits were 4 -5 Hz, regardless of the viewing conditions. These limits are similar to those for within-attribute (first- and second-order) object tracking (4 -5 Hz) obtained in our previous study. Thus, this putative mechanism may be responsible for object tracking, based only on second-order components, even in the case of first-order stimuli.

Second-order visual sensitivity in the aging population.

Most visual and cognitive functions are affected by aging over the lifespan. In this study, our aim was to investigate the loss in sensitivity to different classes of second-order stimuli-a class of stimuli supposed to be mainly processed in extrastriate cortex-in the aging population. These stimuli will then allow one to identify specific cortical deficit independently of visibility losses in upstream parts of the visual pathway. For this purpose, we measured the sensitivity to first-order stimuli and second-order stimuli: orientation-modulated, motion-modulated or contrast-modulated as a function of spatial frequency in 50 aged participants. Overall, we observed a sensitivity loss for all classes of stimuli, but this loss differentially affects the three classes of second-order stimuli tested. It involves all modulation spatial frequencies in the case of motion modulation, but just high modulation spatial frequencies in the case of contrast- and orientation modulations. These observations imply that aging selectively affects the sensitivity to second-order stimuli depending on their type. Since there is evidence that these different second-order stimuli are processed in different regions of extrastriate cortex, this result may suggest that some visual cortical areas are more susceptible to aging effects than others.

Binocular combination of luminance profiles.

We develop and test a new two-dimensional model for binocular combination of the two eyes' luminance profiles. For first-order stimuli, the model assumes that one eye's luminance profile first goes through a luminance compressor, receives gain-control and gain-enhancement from the other eye, and then linearly combines the other eye's output profile. For second-order stimuli, rectification is added in the signal path of the model before the binocular combination site. Both the total contrast and luminance energies, weighted sums over both the space and spatial-frequency domains, were used in the interocular gain-control, while only the total contrast energy was used in the interocular gain-enhancement. To challenge the model, we performed a binocular brightness matching experiment over a large range of background and target luminances. The target stimulus was a dichoptic disc with a sharp edge that has an increment or decrement luminance from its background. The disk's interocular luminance ratio varied from trial to trial. To refine the model we tested three luminance compressors, five nested binocular combination models (including the Ding–Sperling and the DSKL models), and examined the presence or absence of total luminance energy in the model. We found that (1) installing a luminance compressor, either a logarithmic luminance function or luminance gain-control, (2) including both contrast and luminance energies, and (3) adding interocular gain-enhancement (the DSKL model) to a combined model significantly improved its performance. The combined model provides a systematic account of binocular luminance summation over a large range of luminance input levels. It gives a unified explanation of Fechner's paradox observed on a dark background, and a winner-take-all phenomenon observed on a light background. To further test the model, we conducted two additional experiments: luminance summation of discs with asymmetric contour information (Experiment 2), similar to Levelt (1965) and binocular combination of second-order contrast-modulated gratings (Experiment 3). We used the model obtained in Experiment 1 to predict the results of Experiments 2 and 3 and the results of our previous studies. Model simulations further refined the contrast space weight and contrast sensitivity functions that are installed in the model, and provide a reasonable account for rebalancing of imbalanced binocular vision by reducing the mean luminance in the dominant eye.

Functional magnetic resonance imaging (fMRI) - guided single unit recordings reveal higher order disparity selectivity in area PIP of the macaque brain.

Functional magnetic resonance imaging (fMRI) - guided single unit recordings reveal higher order disparity selectivity in area PIP of the macaque brain.

New insights into statistical learning and chunk learning in implicit sequence acquisition.

Implicit sequence learning is ubiquitous in our daily life. However, it is unclear whether the initial acquisition of sequences results from learning to chunk items (i.e., chunk learning) or learning the underlying statistical regularities (i.e., statistical learning). By grouping responses with or without a distinct chunk or statistical structure into segments and comparing these responses, previous studies have demonstrated both chunk and statistical learning. However, few studies have considered the response sequence as a whole and examined the temporal dependency of the entire sequence, where the temporal dependencies could disclose the internal representations of chunk and statistical learning. Participants performed a serial reaction time (SRT) task under different stimulus interval conditions. We found that sequence learning reflected by reaction time (RT) rather than motor improvements represented by movement time (MT). The temporal dependency of RT and MT revealed that both RT and MT displayed recursive patterns caused by biomechanical effects of response locations and foot transitions. Chunking was noticeable only in the presence of the recurring RT or MT but vanished after the recursive component was removed, implying that chunk formation may result from biomechanical constraints rather than learning itself. In addition, we observed notable first-order autocorrelations in RT. This trial-to-trial association enhanced as learning progressed regardless of stimulus intervals, reflecting the internal cognitive representation of the first-order stimulus contingencies. Our results suggest that initial acquisition of implicit sequences may arise from first-order statistical learning rather than chunk learning.

Optimized Parallel Coding of Second-Order Stimulus Features by Heterogeneous Neural Populations.

Efficient processing of sensory input is essential to ensure an organism's survival in its natural environment. Growing evidence suggests that sensory neurons can optimally encode natural stimuli by ensuring that their tuning opposes stimulus statistics, such that the resulting neuronal response contains equal power at all frequencies (i.e., is "white"). Such temporal decorrelation or whitening has been observed across modalities, but the effects of neural heterogeneities on determining tuning and thus responses to natural stimuli have not been investigated. Here, we investigate how heterogeneities in sensory pyramidal neurons organized in three parallel maps representing the body surface determine responses to second-order electrosensory stimulus features in the weakly electric fish Apteronotus leptorhynchus While some sources of heterogeneities such as ON- and OFF-type responses to first-order did not affect responses to second-order electrosensory stimulus features, other sources of heterogeneity within and across the maps strongly determined responses. We found that these cells effectively performed a fractional differentiation operation on their input with exponents ranging from zero (no differentiation) to 0.4 (strong differentiation). Varying adaptation in a simple model explained these heterogeneities and predicted a strong correlation between fractional differentiation and adaptation. Using natural stimuli, we found that only a small fraction of neurons implemented temporal whitening. Rather, a large fraction of neurons did not perform any significant whitening and thus preserved natural input statistics in their responses. We propose that this information is needed to properly decode optimized information sent in parallel through temporally whitened responses based on context. We demonstrate that heterogeneities in the same sensory neuron type can either have no or significant influence on their responses to second-order stimulus features. While an ON- or OFF-type response to first-order stimulus attributes has no significant influence on responses to second-order stimulus features, we found that only a small fraction of sensory neurons optimally encoded natural stimuli through high-pass filtering, thereby implementing temporal whitening. Surprisingly, a large fraction of sensory neurons performed little if no filtering of stimuli, thereby preserving natural stimulus statistics. We hypothesize that this pathway is necessary to properly decode optimized information contained in temporally whitened responses based on context.

Deficient Binocular Combination of Second-Order Stimuli in Amblyopia.

Sensory imbalances in humans with amblyopia have been well documented using luminance-modulated (first-order) stimuli. However, little is known regarding whether there is a deficient binocular combination in amblyopes for stimuli defined by modulations in contrast (second-order stimuli). To address this, we asked two questions: Does a sensory imbalance also exist in the binocular combination of second-order stimuli, and if so, is it more severe than that expected on the basis of the imbalance for first-order stimuli? The sensory imbalances of 14 adult amblyopes (mean age: 30.5 ± 11.5 years; 5 with strabismus and 9 without) were measured using a dichoptic phase combination task. Three types of second-order dichoptic stimulus pairs were used in the study, where the carriers in the two eyes were either correlated, anticorrelated, or uncorrelated. Results were compared with those obtained using first-order stimuli in all observers. We found that second-order binocular combination in amblyopes was not affected by the interocular carrier correlations. The amblyopic eye's contribution to the binocularly fused percept was much less than that of the nonamblyopic eye. The resulting sensory imbalance in binocular combination for second-order images was comparable to that for first-order images in 8 of the observers but was more severe in the other 6 amblyopes. These results indicate that amblyopia does not disrupt the normal architecture of binocular combination for second-order signals; however, there is an additional deficit in binocular combination of second-order image in some amblyopes that cannot be fully accounted for by the known first-order sensory imbalance.

First- and Second-Order Stimuli Reaction Time Measures Are Highly Sensitive to Mild Traumatic Brain Injuries.

Mild traumatic brain injury (mTBI) has subtle effects on several brain functions that can be difficult to assess and follow up. We investigated the impact of mTBI on the perception of sine-wave gratings defined by first- and second-order characteristics. Fifteen adults diagnosed with mTBI were assessed at 15 days, 3 months, and 12 months postinjury. Fifteen matched controls followed the same testing schedule. Reaction times (RTs) for flicker detection and motion direction discrimination were measured. Stimulus contrast of first- and second-order patterns was equated to control for visibility, and correct-response RT means, standard deviations (SDs), medians, and interquartile ranges (IQRs) were calculated. The level of symptoms was also evaluated to compare it to RT data. In general in mTBI, RTs were longer, and SDs as well as IQRs larger, than those of controls. In addition, mTBI participants' RTs to first-order stimuli were shorter than those to second-order stimuli, and SDs as well as IQRs larger for first- than for second-order stimuli in the motion condition. All these observations were made over the three sessions. The level of symptoms observed in mTBI was higher than that of control participants, and this difference did also persist up to 1 year after the brain injury, despite an improvement. The combination of RT measures with particular stimulus properties is a highly sensitive method for measuring mTBI-induced visuomotor anomalies and provides a fine probe of the underlying mechanisms when the brain is exposed to mild trauma.

Neural heterogeneities determine response characteristics to second-, but not first-order stimulus features.

Neural heterogeneities are seen ubiquitously, but how they determine neural response properties remains unclear. Here we show that heterogeneities can either strongly, or not at all, influence neural responses to a given stimulus feature. Specifically, we recorded from peripheral electroreceptor neurons, which display strong heterogeneities in their resting discharge activity, in response to naturalistic stimuli consisting of a fast time-varying waveform (i.e., first-order) whose amplitude (i.e., second-order or envelope) varied slowly in the weakly electric fish Apteronotus leptorhynchus. Although electroreceptors displayed relatively homogeneous responses to first-order stimulus features, further analysis revealed two subpopulations with similar sensitivities that were excited or inhibited by increases in the envelope, respectively, for stimuli whose frequency content spanned the natural range. We further found that a linear-nonlinear cascade model incorporating the known linear response characteristics to first-order features and a static nonlinearity accurately reproduced experimentally observed responses to both first- and second-order features for all stimuli tested. Importantly, this model correctly predicted that the response magnitude is independent of either the stimulus waveform's or the envelope's frequency content. Further analysis of our model led to the surprising prediction that the mean discharge activity can be used to determine whether a given neuron is excited or inhibited by increases in the envelope. This prediction was validated by our experimental data. Thus, our results provide key insight as to how neural heterogeneities can determine response characteristics to some, but not other, behaviorally relevant stimulus features.

Linear binocular combination of responses to contrast modulation: contrast-weighted summation in first- and second-order vision.

Binocular combination for first-order (luminance-defined) stimuli has been widely studied, but we know rather little about this binocular process for spatial modulations of contrast (second-order stimuli). We used phase-matching and amplitude-matching tasks to assess binocular combination of second-order phase and modulation depth simultaneously. With fixed modulation in one eye, we found that binocularly perceived phase was shifted, and perceived amplitude increased almost linearly as modulation depth in the other eye increased. At larger disparities, the phase shift was larger and the amplitude change was smaller. The degree of interocular correlation of the carriers had no influence. These results can be explained by an initial extraction of the contrast envelopes before binocular combination (consistent with the lack of dependence on carrier correlation) followed by a weighted linear summation of second-order modulations in which the weights (gains) for each eye are driven by the first-order carrier contrasts as previously found for first-order binocular combination. Perceived modulation depth fell markedly with increasing phase disparity unlike previous findings that perceived first-order contrast was almost independent of phase disparity. We present a simple revision to a widely used interocular gain-control theory that unifies first- and second-order binocular summation with a single principle--contrast--weighted summation-and we further elaborate the model for first-order combination. Second-order combination is controlled by first-order contrast.

Processing deficits of motion of contrast-modulated gratings in anisometropic amblyopia.

Several studies have indicated substantial processing deficits for static second-order stimuli in amblyopia. However, less is known about the perception of second-order moving gratings. To investigate this issue, we measured the contrast sensitivity for second-order (contrast-modulated) moving gratings in seven anisometropic amblyopes and ten normal controls. The measurements were performed with non-equated carriers and a series of equated carriers. For comparison, the sensitivity for first-order motion and static second-order stimuli was also measured. Most of the amblyopic eyes (AEs) showed reduced sensitivity for second-order moving gratings relative to their non-amblyopic eyes (NAEs) and the dominant eyes (CEs) of normal control subjects, even when the detectability of the noise carriers was carefully controlled, suggesting substantial processing deficits of motion of contrast-modulated gratings in anisometropic amblyopia. In contrast, the non-amblyopic eyes of the anisometropic amblyopes were relatively spared. As a group, NAEs showed statistically comparable performance to CEs. We also found that contrast sensitivity for static second-order stimuli was strongly impaired in AEs and part of the NAEs of anisometropic amblyopes, consistent with previous studies. In addition, some amblyopes showed impaired performance in perception of static second-order stimuli but not in that of second-order moving gratings. These results may suggest a dissociation between the processing of static and moving second-order gratings in anisometropic amblyopia.

A normative framework for the study of second-order sensitivity in vision.

While the contrast sensitivity approach has been successful in evaluating the processing of first-order stimuli, there is a need to develop comparable ways of assessing second-order vision. Our purpose here is to establish normative data on second-order contrast-, orientation-, and motion-modulation sensitivity in humans. We propose a unified framework, applying the quick contrast sensitivity function (qCSF) method, which was recently developed for the rapid measurement of contrast sensitivity across the full spatial-frequency range (Lesmes, Lu, Baek, & Albright, 2010), to measure both first- and second-order sensitivity functions. We first show that the qCSF methodology can be successfully adapted to different kinds of first- and second-order measurements. We provide a normative dataset for both first- and second-order sensitivity, and we show that the sensitivity to all these stimuli is equal in the two eyes. Our results confirm some strong differences between first- and second-order processing, in accordance with the classical filter-rectify-filter model. They suggest a common contrast detection mechanism but different second-order mechanisms.

Age-related changes in perception of movement in driving scenes.

Age-related changes in motion sensitivity have been found to relate to reductions in various indices of driving performance and safety. The aim of this study was to investigate the basis of this relationship in terms of determining which aspects of motion perception are most relevant to driving. Participants included 61 regular drivers (age range 22-87years). Visual performance was measured binocularly. Measures included visual acuity, contrast sensitivity and motion sensitivity assessed using four different approaches: (1) threshold minimum drift rate for a drifting Gabor patch, (2) Dmin from a random dot display, (3) threshold coherence from a random dot display, and (4) threshold drift rate for a second-order (contrast modulated) sinusoidal grating. Participants then completed the Hazard Perception Test (HPT) in which they were required to identify moving hazards in videos of real driving scenes, and also a Direction of Heading task (DOH) in which they identified deviations from normal lane keeping in brief videos of driving filmed from the interior of a vehicle. In bivariate correlation analyses, all motion sensitivity measures significantly declined with age. Motion coherence thresholds, and minimum drift rate threshold for the first-order stimulus (Gabor patch) both significantly predicted HPT performance even after controlling for age, visual acuity and contrast sensitivity. Bootstrap mediation analysis showed that individual differences in DOH accuracy partly explained these relationships, where those individuals with poorer motion sensitivity on the coherence and Gabor tests showed decreased ability to perceive deviations in motion in the driving videos, which related in turn to their ability to detect the moving hazards. The ability to detect subtle movements in the driving environment (as determined by the DOH task) may be an important contributor to effective hazard perception, and is associated with age, and an individuals' performance on tests of motion sensitivity. The locus of the processing deficits appears to lie in first-order, rather than second-order motion pathways.

A comparison of pedestal effects in first- and second-order patterns

The human visual system is sensitive to both luminance (first-order) and contrast (second-order) modulations in an image. A linear-nonlinear-linear model is commonly used to explain visual processing of second-order patterns. Here we used a pattern-masking paradigm to compare first-order and second-order visual mechanisms and to characterize the nonlinear properties underlying them. The carriers were either a high-frequency horizontal grating (8 c/°) or a binary random dot pattern; they were either added to a vertical low-frequency (2 c/°) sinusoidal grating (first-order stimuli) or multiplied by it (second-order stimuli). The incremental discrimination threshold of the target was measured with pedestals whose spatial properties matched those of the target, with the exception of contrast (in the first-order pedestal) or modulation depth (in the second-order pedestal). The threshold function showed a typical dipper shape for both first- and second-order stimuli. The results for the first-order stimuli with different types of carrier and the second-order stimuli with a grating carrier were well explained by a divisive inhibition model in which the facilitatory input was divided by the sum of broadband inhibitory inputs. The results for the second-order stimuli with a random-dot carrier were explained by a modified divisive inhibition model that operated on modulation depth. Our results suggest that divisive inhibition is required to explain visual discrimination in both first- and second-order patterns. However, the source and nonlinearity of the divisive inhibition may be different for these two types of patterns and carrier.

The company they keep: Background similarity influences transfer of aftereffects from second- to first-order stimuli

A wealth of studies has found that adapting to second-order visual stimuli has little effect on the perception of first-order stimuli. This is physiologically and psychologically troubling, since many cells show similar tuning to both classes of stimuli, and since adapting to first-order stimuli leads to aftereffects that do generalize to second-order stimuli. Focusing on high-level visual stimuli, we recently proposed the novel explanation that the lack of transfer arises partially from the characteristically different backgrounds of the two stimulus classes. Here, we consider the effect of stimulus backgrounds in the far more prevalent, lower-level, case of the orientation tilt aftereffect. Using a variety of first- and second-order oriented stimuli, we show that we could increase or decrease both within- and cross-class adaptation aftereffects by increasing or decreasing the similarity of the otherwise apparently uninteresting or irrelevant backgrounds of adapting and test patterns. Our results suggest that similarity between background statistics of the adapting and test stimuli contributes to low-level visual adaptation, and that these backgrounds are thus not discarded by visual processing but provide contextual modulation of adaptation. Null cross-adaptation aftereffects must also be interpreted cautiously. These findings reduce the apparent inconsistency between psychophysical and neurophysiological data about first- and second-order stimuli.

Functional Organization of Envelope-Responsive Neurons in Early Visual Cortex: Organization of Carrier Tuning Properties

It is well established that visual cortex neurons having similar selectivity for orientation, direction of motion, ocular dominance, and other properties of first-order (luminance-defined) stimuli are clustered into a columnar organization. However, the cortical architecture of neuronal responses to second-order (contrast/texture-defined) stimuli is poorly understood. A useful second-order stimulus is a contrast envelope, consisting of a finely detailed pattern (carrier) whose contrast varies on a coarse spatial scale (envelope). In this study, we analyzed the cortical organization of carrier tuning properties of neurons, which responded to contrast-modulated stimuli. We examined whether neurons tuned to similar carrier properties are clustered spatially and whether such spatial clusters are arranged in columns. To address these questions, we recorded single-unit activity, multiunit activity, and local field potentials simultaneously from area 18 of anesthetized cats, using single-channel microelectrodes and multielectrode arrays. Our data showed that neurons tuned to similar carrier spatial frequency are distributed in a highly clustered manner; neurons tuned to similar carrier orientation are also significantly clustered. Neurons along linear arrays perpendicular to the brain surface always exhibited similar optimal carrier spatial frequency, indicating a columnar organization. Multi-pronged tetrode recordings indicated that the diameter of these columns is ≥450 μm. Optimal carrier orientation was also significantly clustered but with finer-grain organization and greater scatter. These results indicate a fine anatomical structure of cortical organization of second-order information processing and suggest that there are probably more maps in cat area 18 than previously believed.