Rapid integration of face detection and task set in visually guided reaching.

The superior colliculus (SC) has been increasingly implicated in the rapid processing of evolutionarily relevant stimuli like faces, but the behavioural relevance of such processing is unclear. The SC has also been implicated in the generation of express visuomotor responses (EVR), which are very short-latency (~80 ms) bursts of muscle activity time-locked to visual target presentation. These observations led us to investigate the influence of faces on EVRs. We recorded upper limb muscle activity from healthy participants as they reached toward targets in the presence of a distractor. In some experiments, faces were used as stimuli. Across blocks of trials, we varied the instruction as to which stimulus served as the target or distractor. Doing so allowed us to assess the impact of instruction on muscle recruitment given identical visual stimuli. We found that responses were uniquely modulated in tasks involving high-contrast faces, promoting reaches toward or away from a face depending on instruction. Follow-up experiments confirmed that the phenomenon required highly salient repeated faces and was not observed to non-facial stimuli nor to faces expressing different affects. This study extends the hypothesis that the SC mediates the EVR by demonstrating that faces impact muscle recruitment at short latencies that precede cortical activity for face perception. Our results constitute direct evidence for the behavioural relevance of face detection in the brainstem, and also implicate a role for top-down cortical pre-setting of the EVR depending on task context.

A priming study on naming real versus pictures of tools.

There is a growing body of literature demonstrating the relationship between the activation of sensorimotor processes in object recognition. It is unclear, however, if these processes are influenced by the differences in how real (3D) tools and two-dimensional (2D) images of tools are processed by the brain. Here, we examined if these differences could influence the naming of tools. Participants were presented with a prime stimulus that was either a picture of a tool, or a real tool, followed by a target stimulus that was always a real tool. They were then required to name each tool as they appeared. The functional use action required by the target tool was either the same (i.e., squeegee-paint roller) or different (i.e. knife-whisk) to the prime. We found that the format in which the prime tool was presented (i.e., a picture or real tool) had no influence on the participants' response times to naming the target tool. Furthermore, participants were faster at naming target tools relative to prime tools when the exact same tool was presented as both the prime and target. There was no difference in response times to naming the target tool relative to the prime when they were different tools, regardless of whether the tools' functional actions were the same or different. We also found more errors in naming target tools relative to the primes when different tools had a different functional action compared to when the same tool was presented as both the prime and the target. Taken together, our results highlight that the functional actions associated with tools do not facilitate or interfere with the recognition of tools for the purposes of naming. The theoretical implications of these results are discussed.

Transient visual pathway critical for normal development of primate grasping behavior.

An evolutionary hallmark of anthropoid primates, including humans, is the use of vision to guide precise manual movements. These behaviors are reliant on a specialized visual input to the posterior parietal cortex. Here, we show that normal primate reaching-and-grasping behavior depends critically on a visual pathway through the thalamic pulvinar, which is thought to relay information to the middle temporal (MT) area during early life and then swiftly withdraws. Small MRI-guided lesions to a subdivision of the inferior pulvinar subnucleus (PIm) in the infant marmoset monkey led to permanent deficits in reaching-and-grasping behavior in the adult. This functional loss coincided with the abnormal anatomical development of multiple cortical areas responsible for the guidance of actions. Our study reveals that the transient retino-pulvinar-MT pathway underpins the development of visually guided manual behaviors in primates that are crucial for interacting with complex features in the environment.

When perception intrudes on 2D grasping: evidence from Garner interference.

When participants reach out to pick up a real 3-D object, their grip aperture reflects the size of the object well before contact is made. At the same time, the classical psychophysical laws and principles of relative size and shape that govern visual perception do not appear to intrude into the control of such movements, which are instead tuned only to the relevant dimension for grasping. In contrast, accumulating evidence suggests that grasps directed at flat 2D objects are not immune to perceptual effects. Thus, in 2D but not 3D grasping, the aperture of the fingers has been shown to be affected by relative and contextual information about the size and shape of the target object. A notable example of this dissociation comes from studies of Garner interference, which signals holistic processing of shape. Previous research has shown that 3D grasping shows no evidence for Garner interference but 2D grasping does (Freud & Ganel, 2015). In a recent study published in this journal (Löhr-Limpens et al., 2019), participants were presented with 2D objects in a Garner paradigm. The pattern of results closely replicated the previously published results with 2D grasping. Unfortunately, the authors, who appear to be unaware the potential differences between 2D and 3D grasping, used their findings to draw an overgeneralized and unwarranted conclusion about the relation between 3D grasping and perception. In this short methodological commentary, we discuss current literature on aperture shaping during 2D grasping and suggest that researchers should play close attention to the nature of the target stimuli they use before drawing conclusions about visual processing for perception and action.

Pupil size is modulated by the size of equal-luminance gratings.

Pupil size changes with light. For this reason, researchers studying the effect of attention, contextual processing, and arousal on the pupillary response have matched the mean luminance of their stimuli across conditions to eliminate the contribution of differences in light levels. Here, we argue that the match of mean luminance is not enough. In Experiment 1, we presented a circular sinewave grating on a gray background for 2 seconds. The area of the grating could be 3°, 6°, or 9°. The mean luminance of each grating was equal to the luminance of the gray background, such that regardless of the size of the grating there was no change in mean luminance between conditions. Participants were asked to fixate the center of the grating and passively view it. We found that in all size conditions, there was a pupil constriction starting at about 300 ms after stimulus onset, and the pupil constriction increased with the size of the grating. In Experiment 2, when a small grating was presented immediately after the presentation of a large grating (or vice versa), the pupil constriction changed accordingly. In Experiment 3, we replicated Experiment 1 but had the subjects perform an attention-demanding fixation task in one session, and passively view the stimuli in the other. We found that the main effect of task was not significant. In sum, our results show that stimulus size can modulate pupil size robustly and steadily even when the luminance is matched across the different stimuli.

The material-weight illusion disappears or inverts in objects made of two materials.

The material-weight illusion (MWI) occurs when an object that looks heavy (e.g., stone) and one that looks light (e.g., Styrofoam) have the same mass. When such stimuli are lifted, the heavier-looking object feels lighter than the lighter-looking object, presumably because well-learned priors about the density of different materials are violated. We examined whether a similar illusion occurs when a certain weight distribution is expected (such as the metal end of a hammer being heavier), but weight is uniformly distributed. In experiment 1, participants lifted bipartite objects that appeared to be made of two materials (combinations of stone, Styrofoam, and wood) but were manipulated to have a uniform weight distribution. Most participants experienced an inverted MWI (i.e., the heavier-looking side felt heavier), suggesting an integration of incoming sensory information with density priors. However, a replication of the classic MWI was found when the objects appeared to be uniformly made of just one of the materials ( experiment 2). Both illusions seemed to be independent of the forces used when the objects were lifted. When lifting bipartite objects but asked to judge the weight of the whole object, participants experienced no illusion ( experiment 3). In experiment 4, we investigated weight perception in objects with a nonuniform weight distribution and again found evidence for an integration of prior and sensory information. Taken together, our seemingly contradictory results challenge most theories about the MWI. However, Bayesian integration of competing density priors with the likelihood of incoming sensory information may explain the opposing illusions. NEW & NOTEWORTHY We report a novel weight illusion that contradicts all current explanations of the material-weight illusion: When lifting an object composed of two materials, the heavier-looking side feels heavier, even when the true weight distribution is uniform. The opposite (classic) illusion is found when the same materials are lifted in two separate objects. Identifying the common mechanism underlying both illusions will have implications for perception more generally. A potential candidate is Bayesian inference with competing priors.

A pantomiming priming study on the grasp and functional use actions of tools.

It has previously been demonstrated that tool recognition is facilitated by the repeated visual presentation of object features affording actions, such as those related to grasping and their functional use. It is unclear, however, if this can also facilitate pantomiming. Participants were presented with an image of a prime followed by a target tool and were required to pantomime the appropriate action for each one. The grasp and functional use attributes of the target tool were either the same or different to the prime. Contrary to expectations, participants were slower at pantomiming the target tool relative to the prime regardless of whether the grasp and function of the tool were the same or different-except when the prime and target tools consisted of identical images of the same exemplar. We also found a decrease in accuracy of performing functional use actions for the target tool relative to the prime when the two differed in functional use but not grasp. We reconcile differences between our findings and those that have performed priming studies on tool recognition with differences in task demands and known differences in how the brain recognises tools and performs actions to make use of them.

What Role Does "Elongation" Play in "Tool-Specific" Activation and Connectivity in the Dorsal and Ventral Visual Streams?

Images of tools induce stronger activation than images of nontools in a left-lateralized network that includes ventral-stream areas implicated in tool identification and dorsal-stream areas implicated in tool manipulation. Importantly, however, graspable tools tend to be elongated rather than stubby, and so the tool-selective responses in some of these areas may, to some extent, reflect sensitivity to elongation rather than "toolness" per se. Using functional magnetic resonance imaging, we investigated the role of elongation in driving tool-specific activation in the 2 streams and their interconnections. We showed that in some "tool-selective" areas, the coding of toolness and elongation coexisted, but in others, elongation and toolness were coded independently. Psychophysiological interaction analysis revealed that toolness, but not elongation, had a strong modulation of the connectivity between the ventral and dorsal streams. Dynamic causal modeling revealed that viewing tools (either elongated or stubby) increased the connectivity from the ventral- to the dorsal-stream tool-selective areas, but only viewing elongated tools increased the reciprocal connectivity between these areas. Overall, these data disentangle how toolness and elongation affect the activation and connectivity of the tool network and help to resolve recent controversies regarding the relative contribution of "toolness" versus elongation in driving dorsal-stream "tool-selective" areas.

Sharpening vision by adapting to flicker.

Human vision is surprisingly malleable. A static stimulus can seem to move after prolonged exposure to movement (the motion aftereffect), and exposure to tilted lines can make vertical lines seem oppositely tilted (the tilt aftereffect). The paradigm used to induce such distortions (adaptation) can provide powerful insights into the computations underlying human visual experience. Previously spatial form and stimulus dynamics were thought to be encoded independently, but here we show that adaptation to stimulus dynamics can sharpen form perception. We find that fast flicker adaptation (FFAd) shifts the tuning of face perception to higher spatial frequencies, enhances the acuity of spatial vision-allowing people to localize inputs with greater precision and to read finer scaled text, and it selectively reduces sensitivity to coarse-scale form signals. These findings are consistent with two interrelated influences: FFAd reduces the responsiveness of magnocellular neurons (which are important for encoding dynamics, but can have poor spatial resolution), and magnocellular responses contribute coarse spatial scale information when the visual system synthesizes form signals. Consequently, when magnocellular responses are mitigated via FFAd, human form perception is transiently sharpened because "blur" signals are mitigated.

A TMS Investigation on the Role of Lateral Occipital Complex and Caudal Intraparietal Sulcus in the Perception of Object Form and Orientation.

We used TMS to assess the causal roles of the lateral occipital (LO) and caudal intraparietal sulcus (cIPS) areas in the perceptual discrimination of object features. All participants underwent fMRI to localize these areas using a protocol in which they passively viewed images of objects that varied in both form and orientation. fMRI identified six significant brain regions: LO, cIPS, and the fusiform gyrus, bilaterally. In a separate experimental session, we applied TMS to LO or cIPS while the same participants performed match-to-sample form or orientation discrimination tasks. Compared with sham stimulation, TMS to either the left or right LO increased RTs for form but not orientation discrimination, supporting a critical role for LO in form processing for perception- and judgment-based tasks. In contrast, we did not observe any effects when we applied TMS to cIPS. Thus, despite the clear functional evidence of engagement for both LO and cIPS during the passive viewing of objects in the fMRI experiment, the TMS experiment revealed that cIPS is not critical for making perceptual judgments about their form or orientation.

Time flies when we intend to act: temporal distortion in a go/no-go task.

Although many of our actions are triggered by sensory events, almost nothing is known about our perception of the timing of those sensory events. Here we show that, when people react to a sudden visual stimulus that triggers an action, that stimulus is perceived to occur later than an identical stimulus that does not trigger an action. In our experiments, participants fixated the center of a clock face with a rotating second hand. When the clock changed color, they were required to make a motor response and then to report the position of the second hand at the moment the clock changed color. In Experiment 1, in which participants made a target-directed saccade, the color change was perceived to occur 59 ms later than when they maintained fixation. In Experiment 2, in which we used a go/no-go paradigm, this temporal distortion was observed even when participants were required to cancel a prepared saccade. Finally, in Experiment 3, the same distortion in perceived time was observed for both go and no-go trials in a manual task in which no eye movements were required. These results suggest that, when a visual stimulus triggers an action, it is perceived to occur significantly later than an identical stimulus unrelated to action. Moreover, this temporal distortion appears to be related not to the execution of the action (or its effect) but rather to the programming of the action. In short, there seems to be a temporal binding between a triggering event and the triggered action.

Preserved Haptic Shape Processing after Bilateral LOC Lesions.

The visual and haptic perceptual systems are understood to share a common neural representation of object shape. A region thought to be critical for recognizing visual and haptic shape information is the lateral occipital complex (LOC). We investigated whether LOC is essential for haptic shape recognition in humans by studying behavioral responses and brain activation for haptically explored objects in a patient (M.C.) with bilateral lesions of the occipitotemporal cortex, including LOC. Despite severe deficits in recognizing objects using vision, M.C. was able to accurately recognize objects via touch. M.C.'s psychophysical response profile to haptically explored shapes was also indistinguishable from controls. Using fMRI, M.C. showed no object-selective visual or haptic responses in LOC, but her pattern of haptic activation in other brain regions was remarkably similar to healthy controls. Although LOC is routinely active during visual and haptic shape recognition tasks, it is not essential for haptic recognition of object shape. SIGNIFICANCE STATEMENT: The lateral occipital complex (LOC) is a brain region regarded to be critical for recognizing object shape, both in vision and in touch. However, causal evidence linking LOC with haptic shape processing is lacking. We studied recognition performance, psychophysical sensitivity, and brain response to touched objects, in a patient (M.C.) with extensive lesions involving LOC bilaterally. Despite being severely impaired in visual shape recognition, M.C. was able to identify objects via touch and she showed normal sensitivity to a haptic shape illusion. M.C.'s brain response to touched objects in areas of undamaged cortex was also very similar to that observed in neurologically healthy controls. These results demonstrate that LOC is not necessary for recognizing objects via touch.

Effects of material properties and object orientation on precision grip kinematics.

Successfully picking up and handling objects requires taking into account their physical properties (e.g., material) and position relative to the body. Such features are often inferred by sight, but it remains unclear to what extent observers vary their actions depending on the perceived properties. To investigate this, we asked participants to grasp, lift and carry cylinders to a goal location with a precision grip. The cylinders were made of four different materials (Styrofoam, wood, brass and an additional brass cylinder covered with Vaseline) and were presented at six different orientations with respect to the participant (0°, 30°, 60°, 90°, 120°, 150°). Analysis of their grasping kinematics revealed differences in timing and spatial modulation at all stages of the movement that depended on both material and orientation. Object orientation affected the spatial configuration of index finger and thumb during the grasp, but also the timing of handling and transport duration. Material affected the choice of local grasp points and the duration of the movement from the first visual input until release of the object. We find that conditions that make grasping more difficult (orientation with the base pointing toward the participant, high weight and low surface friction) lead to longer durations of individual movement segments and a more careful placement of the fingers on the object.

Decoding visual object categories in early somatosensory cortex.

Neurons, even in the earliest sensory areas of cortex, are subject to a great deal of contextual influence from both within and across modality connections. In the present work, we investigated whether the earliest regions of somatosensory cortex (S1 and S2) would contain content-specific information about visual object categories. We reasoned that this might be possible due to the associations formed through experience that link different sensory aspects of a given object. Participants were presented with visual images of different object categories in 2 fMRI experiments. Multivariate pattern analysis revealed reliable decoding of familiar visual object category in bilateral S1 (i.e., postcentral gyri) and right S2. We further show that this decoding is observed for familiar but not unfamiliar visual objects in S1. In addition, whole-brain searchlight decoding analyses revealed several areas in the parietal lobe that could mediate the observed context effects between vision and somatosensation. These results demonstrate that even the first cortical stages of somatosensory processing carry information about the category of visually presented familiar objects.

Equal-magnitude size-weight illusions experienced within and between object categories.

In the size-weight illusion (SWI), small objects feel heavier than larger objects of the same mass. This effect is typically thought to be a consequence of the lifter's expectation that the large object will outweigh the small object, because objects of the same type typically get heavier as they get larger. Here, we show that this perceptual effect can occur across object category, where there are no strong expectations about the correspondence between size and mass. One group of participants lifted same-colored large and small cubes with the same mass as one another, while another group lifted differently-colored large and small cubes with the same mass as one another. The group who lifted the same-colored cubes experienced a robust SWI and initially lifted the large object with more force than the small object. By contrast, the group who lifted the different-colored objects did so with equal initial forces on the first trial, but experienced just as strong an illusion as those who lifted the same-colored objects. These results demonstrate that color cues can selectively influence the application of fingertip force rates while not impacting at all upon the lifter's perception of object weight, highlighting a stark dissociation in how prior information affects perception and action.

Transient visual responses reset the phase of low-frequency oscillations in the skeletomotor periphery.

We recorded muscle activity from an upper limb muscle while human subjects reached towards peripheral targets. We tested the hypothesis that the transient visual response sweeps not only through the central nervous system, but also through the peripheral nervous system. Like the transient visual response in the central nervous system, stimulus-locked muscle responses (< 100 ms) were sensitive to stimulus contrast, and were temporally and spatially dissociable from voluntary orienting activity. Also, the arrival of visual responses reduced the variability of muscle activity by resetting the phase of ongoing low-frequency oscillations. This latter finding critically extends the emerging evidence that the feedforward visual sweep reduces neural variability via phase resetting. We conclude that, when sensory information is relevant to a particular effector, detailed information about the sensorimotor transformation, even from the earliest stages, is found in the peripheral nervous system.

The size-weight illusion induced through human echolocation.

Certain blind individuals have learned to interpret the echoes of self-generated sounds to perceive the structure of objects in their environment. The current work examined how far the influence of this unique form of sensory substitution extends by testing whether echolocation-induced representations of object size could influence weight perception. A small group of echolocation experts made tongue clicks or finger snaps toward cubes of varying sizes and weights before lifting them. These echolocators experienced a robust size-weight illusion. This experiment provides the first demonstration of a sensory substitution technique whereby the substituted sense influences the conscious perception through an intact sense.

A blind human expert echolocator shows size constancy for objects perceived by echoes.

Some blind humans make clicking noises with their mouth and use the reflected echoes to perceive objects and surfaces. This technique can operate as a crude substitute for vision, allowing human echolocators to perceive silent, distal objects. Here, we tested if echolocation would, like vision, show size constancy. To investigate this, we asked a blind expert echolocator (EE) to echolocate objects of different physical sizes presented at different distances. The EE consistently identified the true physical size of the objects independent of distance. In contrast, blind and blindfolded sighted controls did not show size constancy, even when encouraged to use mouth clicks, claps, or other signals. These findings suggest that size constancy is not a purely visual phenomenon, but that it can operate via an auditory-based substitute for vision, such as human echolocation.