Second-order Cues Research Articles

Formalising prestige bias: Differences between models with first-order and second-order cues.

Knowledge and behaviour are transmitted from one individual to another through social learning and eventually disseminated across the population. People often learn useful behaviours socially through selective bias rather than random selection of targets. Prestige bias, or the tendency to selectively imitate prestigious individuals, has been considered an important factor in influencing human behaviour. Although its importance in human society and culture has been recognised, the formulation of prestige bias is less developed than that of other social learning biases. To examine the effects of prestige bias on cultural evolution theoretically, it is imperative to formulate prestige and investigate its basic properties. We reviewed two definitions: one based on first-order cues, such as the demonstrator's appearance and job title, and the other based on second-order cues, such as people's behaviour towards the demonstrator (e.g. people increasingly pay attention to prestigious individuals). This study builds a computational model of prestige bias based on these two definitions and compares the cultural evolutionary dynamics they generate. Our models demonstrate the importance of distinguishing between the two types of formalisation, because they can have different influences on cultural evolution.

Connectivity-Defined Subdivisions of the Intraparietal Sulcus Respond Differentially to Abstraction during Decision-Making.

The intraparietal sulcus (IPS) has been implicated in numerous functions that range from representation of visual stimuli to action planning, but its role in abstract decision-making has been unclear, in part because low-level functions often act as confounds. Here, we address this problem using a task that dissociates abstract decision-making from sensory salience, attentional control, motor planning, and motor output. Functional MRI data were collected from healthy female and male human subjects while they performed a policy abstraction task requiring use of a more abstract (second-order) rule to select between two less abstract (first-order) rules that informed the motor response. By identifying IPS subdivisions with preferential connectivity to prefrontal regions that are differentially responsive to task abstraction, we found that a caudal IPS (cIPS) subregion with strongest connectivity to the pre-premotor cortex was preferentially active for second-order cues, whereas a rostral IPS subregion (rIPS) with strongest connectivity to the dorsal premotor cortex was active during attentional control over first-order cues. These effects for abstraction were seen in addition to cIPS activity that was specific to sensory salience, and rIPS activity that was specific to motor output. Notably, topographic responses to the second-order cue were detected along the caudal-rostral axis of IPS, mirroring the broader organization seen in lateral prefrontal cortex. Together, these data demonstrate that subregions within IPS exhibit activity responsive to policy abstraction, and they suggest that IPS may be organized into frontoparietal subnetworks that support hierarchical cognitive control.SIGNIFICANCE STATEMENT Abstract decision-making allows us to flexibly adapt our behavior to new contexts. Although much previous work has focused on the role of lateral prefrontal cortex in such decisions, the contributions of parietal cortex have been relatively understudied. Here, we demonstrate that spatially segregated subregions of human IPS with strong functional connections to lateral prefrontal cortex demonstrate activity selective for abstract decisions. This activity can be distinguished from responses resulting from cognitive processes related to sensory salience, attentional control, motor planning, and movement. Together, these findings indicate that different task demands are reflected in the topography of IPS, and they explicitly reveal a role in abstract decision-making.

Second-order texture gratings produce overestimation of height in depictions of rectangles and steps

The horizontal-vertical illusion (HVI) has been proposed as a method to increase the perceived height of steps, increase toe clearance and prevent falls. High contrast vertical stripes are placed on the step riser abutting a horizontal edge-highlighter creating ‘T’ junctions which are thought to promote the illusion. Various configurations of the HVI were tested including luminance gratings (L) and second-order modulations of contrast (CM), spatial frequency (FM) and orientation (OM). Observers were asked to compare the apparent height of gratings with that of either filled, unmodulated rectangles or unfilled rectangles. Rectangles were presented alone or as part of a step with a highlighter. In some conditions highlighters matched the properties of the grating; in others or not. In one critical experiment, the HVI was compared for steps with highlighters that were separated from the riser by a thin line and those where the risers and highlighters were continuous. All gratings except FM appeared taller when presented in the step configuration with a continuous, matching highlighter. This effect was greatly reduced when a thin line separated the grating from the highlighter and abolished for mis-matched highlighters and risers. In the rectangle conditions, all cues appeared taller than blank rectangles and L and CM appeared taller than filled-unmodulated rectangles. In conclusion, second-order cues may be useful for inducing the HVI onto steps. However, the ability of vertical stripes and edge-highlighters to accentuate perceived step height may be due to aggregation of the highlighter into the grating rather than the normal horizontal-vertical illusion.

Second-order cues to figure motion enable object detection during prey capture by praying mantises

Detecting motion is essential for animals to perform a wide variety of functions. In order to do so, animals could exploit motion cues, including both first-order cues-such as luminance correlation over time-and second-order cues, by correlating higher-order visual statistics. Since first-order motion cues are typically sufficient for motion detection, it is unclear why sensitivity to second-order motion has evolved in animals, including insects. Here, we investigate the role of second-order motion in prey capture by praying mantises. We show that prey detection uses second-order motion cues to detect figure motion. We further present a model of prey detection based on second-order motion sensitivity, resulting from a layer of position detectors feeding into a second layer of elementary-motion detectors. Mantis stereopsis, in contrast, does not require figure motion and is explained by a simpler model that uses only the first layer in both eyes. Second-order motion cues thus enable prey motion to be detected, even when perfectly matching the average background luminance and independent of the elementary motion of any parts of the prey. Subsequent to prey detection, processes such as stereopsis could work to determine the distance to the prey. We thus demonstrate how second-order motion mechanisms enable ecologically relevant behavior such as detecting camouflaged targets for other visual functions including stereopsis and target tracking.

Multivariate Analysis of BOLD Activation Patterns Recovers Graded Depth Representations in Human Visual and Parietal Cortex.

Navigating through natural environments requires localizing objects along three distinct spatial axes. Information about position along the horizontal and vertical axes is available from an object’s position on the retina, while position along the depth axis must be inferred based on second-order cues such as the disparity between the images cast on the two retinae. Past work has revealed that object position in two-dimensional (2D) retinotopic space is robustly represented in visual cortex and can be robustly predicted using a multivariate encoding model, in which an explicit axis is modeled for each spatial dimension. However, no study to date has used an encoding model to estimate a representation of stimulus position in depth. Here, we recorded BOLD fMRI while human subjects viewed a stereoscopic random-dot sphere at various positions along the depth (z) and the horizontal (x) axes, and the stimuli were presented across a wider range of disparities (out to ∼40 arcmin) compared to previous neuroimaging studies. In addition to performing decoding analyses for comparison to previous work, we built encoding models for depth position and for horizontal position, allowing us to directly compare encoding between these dimensions. Our results validate this method of recovering depth representations from retinotopic cortex. Furthermore, we find convergent evidence that depth is encoded most strongly in dorsal area V3A.

Basolateral amygdala, nicotinic cholinergic receptors, and nicotine: Pharmacological effects and addiction in animal models and humans.

The amygdala is involved in processing incoming information about rewarding stimuli and emotions that denote danger such as anxiety and fear. Bi-directional neural connections between basolateral amygdala (BLA) and brain regions such as nucleus accumbens, prefrontal cortex, hippocampus, and hindbrain regions regulate motivation, cognition, and responses to stress. Altered local regulation of BLA excitability is pivotal to the behavioral disturbances characteristic of posttraumatic stress disorder, and relapse to drug use induced by stress. Herein, we review the physiological regulation of BLA by cholinergic inputs, emphasizing the role of BLA nicotinic receptors. We review BLA-dependent effects of nicotine on cognition, motivated behaviors, and emotional states, including memory, taking and seeking drugs, and anxiety and fear in humans and animal models. The alterations in BLA activity observed in animal studies inform human behavioral and brain imaging research by enabling a more exact understanding of altered BLA function. Converging evidence indicates that cholinergic signaling from basal forebrain projections to local nicotinic receptors is an important physiological regulator of BLA and that nicotine alters BLA function. In essence, BLA is necessary for behavioral responses to stimuli that evoke anxiety and fear; reinstatement of cue-induced drug seeking; responding to second-order cues conditioned to abused drugs; reacquisition of amplified nicotine self-administration due to chronic stress during abstinence; and to promote responding for natural reward.

Assessment of mid-level vision segregation in mice with radially expanding/contracting targets

Rodents' ability to segregate and recognize objects is still debated today. Our group showed that rats can distinguish figures on a background by focusing on luminance-based first-order cues. In a follow-up study, we demonstrated that rats can then learn to differentiate between figures defined by second-order texture-based cues. The aim of this current study was to design and test a novel discrimination task for mice to test whether they can differentiate between complex motion trajectories based upon second-order cues. 8 mice separated in two groups performed discrimination tasks in automated touch screen operant chambers for 28 testing days. Group 1 had an expanding white square on a black background as a target, group 2 a contracting one (Phase 1), which were then showed simultaneously during testing on the two touchscreens. In a second phase, both black and white squares on inverted backgrounds were randomly interleaved (Phase 2). For Phase 1, 3 out of the 4 mice in the contracting group reached threshold performance of 75% correct after 18.8±5.0 sessions. Interestingly, only 1 out of the 4 mice in the expanding group reached threshold level in 28 sessions. Of interest, mice in the expanding group were thrice as likely to fail to perform 30 trials per session (31.3±3.7% vs 9.8±5.5% of the time). This suggests a possible selection bias toward contracting targets, which will be further evaluated. The mice that passed threshold in step 1 had an average discrimination score of 82.8±4.6% during their first session when introduced to Step 2, indicating that they can generalize. Their score then all slightly dropped, before reaching back threshold in about 4 sessions. Our data indicate that mice can distinguish complex motion trajectories while ignoring luminance-based cues. Radially expanding/contracting targets are thus an interesting tool for probing figure-ground segregation capabilities of rodents. Meeting abstract presented at VSS 2017

Nonlinear Y-Like Receptive Fields in the Early Visual Cortex: An Intermediate Stage for Building Cue-Invariant Receptive Fields from Subcortical Y Cells.

A significant fraction of neurons in early visual cortex have specialized receptive fields that allow them to selectively respond to the orientation of boundaries that are invariant to the cue (luminance, contrast, texture, motion) that defines them. However, the neural mechanism to construct such versatile receptive fields remains unclear. Using multielectrode recording, we found a large fraction of neurons in early visual cortex with receptive fields not selective for orientation that have spatial nonlinearities like those of subcortical Y cells. These are strong candidates for building cue-invariant orientation-selective neurons; we present a neural circuit model that pools such neurons in an imbalanced "push-pull" manner, to generate orientation-selective cue-invariant receptive fields.

In unison: First- and second-order information combine for integration of shape information.

The modulation of orientation around radial frequency (RF) patterns and RF textures is globally processed in both cases. This psychophysical study investigates whether the combination-a textured RF path obtained by applying an RF texture to an RF contour-is processed like a texture or a contour when making judgements about shape. Unlike RF textures, the impression of a closed flow was not required for global integration of textured RF paths, suggesting that these paths were processed as second-order, or contrast-defined contours. Luminance-defined (LD) RF paths were shown to globally integrate but with thresholds approximately half of those for the proposed second-order textured paths. The next experiment investigated whether this benefit was due to LD stimuli possessing double the amount of information (first- and second-order information). A mixed three-part contour composed of two different second-order texture components and an LD component was then employed to determine how the different cues combined. The mixed path thresholds matched predictions derived from a linear combination of first- and second-order cues. The conclusion is that the shape of isolated contours is processed using both first- and second-order information equally and that the contribution of texture is to carry additional second-order signal.

Cue-invariant shape recognition in rats as tested with second-order contours.

Nonhuman primates are the main animal model to investigate high-level properties of human cortical vision. For one property, transformation-invariant object recognition, recent studies have revealed interesting and unknown capabilities in rats. Here we report on the ability of rats to rely upon second-order cues that are important to structure the incoming visual images into figure and background. Rats performed a visual shape discrimination task in which the shapes were not only defined by first-order luminance information but also by a variety of second-order cues such as a change in texture properties. Once the rats were acquainted with a first set of second-order stimuli, they showed a surprising degree of generalization towards new second-order stimuli. The limits of these capabilities were tested in various ways, and the ability to extract the shapes broke down only in extreme cases where no local cues were available to solve the task. These results demonstrate how rats are able to make choices based on fairly complex strategies when necessary.

Individual variability in behavioral flexibility predicts sign-tracking tendency.

Sign-tracking rats show heightened sensitivity to food- and drug-associated cues, which serve as strong incentives for driving reward seeking. We hypothesized that this enhanced incentive drive is accompanied by an inflexibility when incentive value changes. To examine this we tested rats in Pavlovian outcome devaluation or second-order conditioning prior to the assessment of sign-tracking tendency. To assess behavioral flexibility we trained rats to associate a light with a food outcome. After the food was devalued by pairing with illness, we measured conditioned responding (CR) to the light during an outcome devaluation probe test. The level of CR during outcome devaluation probe test correlated with the rats' subsequent tracking tendency, with sign-tracking rats failing to suppress CR to the light after outcome devaluation. To assess Pavlovian incentive learning, we trained rats on first-order (CS+, CS−) and second-order (SOCS+, SOCS−) discriminations. After second-order conditioning, we measured CR to the second-order cues during a probe test. Second-order conditioning was observed across all rats regardless of tracking tendency. The behavioral inflexibility of sign-trackers has potential relevance for understanding individual variation in vulnerability to drug addiction.

Perception of second- and third-order orientation signals and their interactions

Orientation signals, which are crucial to many aspects of visual function, are more complex and varied in the natural world than in the stimuli typically used for laboratory investigation. Gratings and lines have a single orientation, but in natural stimuli, local features have multiple orientations, and multiple orientations can occur even at the same location. Moreover, orientation cues can arise not only from pairwise spatial correlations, but from higher-order ones as well. To investigate these orientation cues and how they interact, we examined segmentation performance for visual textures in which the strengths of different kinds of orientation cues were varied independently, while controlling potential confounds such as differences in luminance statistics. Second-order cues (the kind present in gratings) at different orientations are largely processed independently: There is no cancellation of positive and negative signals at orientations that differ by 45°. Third-order orientation cues are readily detected and interact only minimally with second-order cues. However, they combine across orientations in a different way: Positive and negative signals largely cancel if the orientations differ by 90°. Two additional elements are superimposed on this picture. First, corners play a special role. When second-order orientation cues combine to produce corners, they provide a stronger signal for texture segregation than can be accounted for by their individual effects. Second, while the object versus background distinction does not influence processing of second-order orientation cues, this distinction influences the processing of third-order orientation cues.

Increasing stimulus size impairs first- but not second-order motion perception

As stimulus size increases, the direction of high-contrast moving stimuli becomes increasingly difficult to perceive. This counterintuitive effect, termed spatial suppression, is believed to reflect antagonistic center-surround interactions--mechanisms that play key roles in tasks requiring sensitivity to relative motion. It is unknown, however, whether second-order motion also exhibits spatial suppression. To test this hypothesis, we measured direction discrimination thresholds for first- and second-order stimuli of varying sizes. The results revealed increasing thresholds with increasing size for first-order stimuli but demonstrated no spatial suppression of second-order motion. This selective impairment of first-order motion predicts increasing predominance of second-order cues as stimulus size increases. We confirmed this prediction by utilizing compound stimuli that contain first- and second-order information moving in opposite directions. Specifically, we found that for large stimuli, motion perception becomes increasingly determined by the direction of second-order cues. Overall, our findings show a lack of spatial suppression for second-order stimuli, suggesting that the second-order system may have distinct functional roles, roles that do not require high sensitivity to relative motion.

Visual deficits in amblyopia constrain normal models of second-order motion processing

It is well established that amblyopes exhibit deficits in processing first-order (luminance-defined) patterns. This is readily manifest by measuring spatiotemporal sensitivity (i.e. the “window of visibility”) to moving luminance gratings. However the window of visibility to moving second-order (texture-defined) patterns has not been systematically studied in amblyopia. To address this issue monocular modulation sensitivity (1/threshold) to first-order motion and four different varieties of second-order motion (modulations of either the contrast, flicker, size or orientation of visual noise) was measured over a five-octave range of spatial and temporal frequencies. Compared to normals amblyopes are not only impaired in the processing of first-order motion, but overall they exhibit both higher thresholds and a much narrower window of visibility to second-order images. However amblyopia can differentially impair the perception of some types of second-order motion much more than others and crucially the precise pattern of deficits varies markedly between individuals (even for those with the same conventional visual acuity measures). For the most severely impaired amblyopes certain second-order (texture) cues to movement in the environment are effectively invisible. These results place important constraints on the possible architecture of models of second-order motion perception in human vision.

Co-operative interactions between first- and second-order mechanisms in the processing of structure from motion

Structure from motion (SFM) is the ability to perceive three-dimensional structure from stimuli containing only two-dimensional motion signals and this ability seems to be a result of high-level cortical processes. It has long been thought that local motion signals defined by second-order cues only weakly contribute to perception of SFM since performance on purely second-order SFM tasks is poor, relative to first-order stimuli. We hypothesized that the mechanisms responsible for deriving SFM were insensitive to low-level stimulus attributes such as the first- or second-order nature of the dots composing the stimulus, in other words: that they were "cue-invariant", but that large differences in sensitivity to local first- and second-order motions were responsible for previous findings. By manipulating the relative strength of first-order dots in an SFM stimulus that combines first- and second-order dots, we show that the two types of motion can separately support SFM and co-operatively interact to produce vivid three-dimensional percepts. This provides strong support that the mechanisms underlying SFM are cue-invariant.

First- and second-order configural sensitivity for greeble stimuli in baboons

Previous studies on nonhuman primates have shown inconsistencies in their processing of first- and second-order relational properties of facial stimuli. Using greeble stimuli sharing configural properties with faces, this study assessed configural processing in baboons. Five baboons were trained to recognize a positive stimulus among pairs of greebles in a two-alternative forced choice task. They were then tested with new stimulus pairs involving either a first-order version, with modifications in global qualitative spatial relations, or a second-order version, with modifications of finer spatial relations. Performance remained above chance in all test conditions, including when only second-order cues were available, but it was higher for first-order trials. It is proposed that an extensive training with greebles led to the processing of second-order relational properties. These results demonstrate that configural sensitivity is not restricted to faces in baboons and suggest that a common mechanism may support configural processing for face and nonface stimuli.

What is second-order vision for? Discriminating illumination versus material changes

The human visual system is sensitive to second-order modulations of the local contrast (CM) or amplitude (AM) of a carrier signal. Second-order cues are detected independently of first-order luminance signals; however, it is not clear why vision should benefit from second-order sensitivity. Analysis of the first- and second-order contents of natural images suggests that these cues tend to occur together, but their phase relationship varies. We have shown that in-phase combinations of LM and AM are perceived as a shaded corrugated surface whereas the anti-phase combination can be seen as corrugated when presented alone or as a flat material change when presented in a plaid containing the in-phase cue. We now extend these findings using new stimulus types and a novel haptic matching task. We also introduce a computational model based on initially separate first- and second-order channels that are combined within orientation and subsequently across orientation to produce a shading signal. Contrast gain control allows the LM + AM cue to suppress responses to the LM - AM when presented in a plaid. Thus, the model sees LM - AM as flat in these circumstances. We conclude that second-order vision plays a key role in disambiguating the origin of luminance changes within an image.

Perceptual learning for second-order cues in a shape-from-shading task

Perceptual learning for second-order cues in a shape-from-shading task

Biological motion perception is cue-invariant

Previous work investigating whether biological motion is supported by local second-order motion has been contradictory, with different groups finding either a difference or no difference in performance compared to that obtained with first-order stimuli. Here we show psychophysically, using randomized-polarity and contrast-modulated stimuli, that detection of second-order biological motion walkers is worse for stimuli defined by second-order cues, but this difference is explained by a difference in visibility of the local motion in the stimuli. By mixing first-order and second-order dots within the same stimulus, we show that, when the two types of dot are equally visible, first-order noise dots can mask a second-order walker, and vice-versa. We also show that direction-discrimination of normal, inverted and scrambled walkers follow the same pattern for second-order as that obtained with first-order stimuli. These results are consistent with biological motion being processed by a mechanism that is cue-invariant.

Spatiotemporal templates for detecting orientation-defined targets

Using the classification image technique, the present experiments revealed several characteristics of human observers' spatiotemporal templates for the detection of orientation-defined targets. The stimulus consisted of a spatial 5 x 5 array of elements displayed in 5 (Experiments 1 and 2) or 15 (Experiment 3) temporal frames. A target was defined by the first- or the second-order characteristics of the textures. In Experiment 1, a target signal was presented across all five frames, and observers typically relied on the most reasonable cues in all five frames for detecting targets. In other words, they used the first-order cue for detecting the first-order target and used the second-order cue for detecting the second-order target. Moreover, the spatial profile for detecting the first-order sustained target was localized at the border of the target area, but that for the second-order sustained target showed broader spatial tuning. Presenting the target in just the third temporal frame, as was done in Experiment 2, changed the temporal profile of the observers' templates in the expected manner: Observers used the first-order cue for the first-order target detection and the second-order cue for the second-order target detection only in the third frame. However, changing the temporal characteristics also affected the kinds of spatial cues that were used to detect a target. For example, the classification images revealed that observers used second-order cues (as well as first-order cues) to detect a first-order target, and there was a trend toward increasing the extent of spatial information used when the temporal information was restricted. In Experiment 3, we found similar results for detecting the first-order flashed target with finer, 15-temporal-frame presentation. Lastly, we showed that the classification image is a useful way to reveal individual differences that are not shown with traditional psychophysical techniques.