Should I stay or should I go? The cerebral bases of street-crossing decision.

An observer willing to cross a street must first estimate if the approaching cars offer enough time to safely complete the task. The brain areas supporting this perception, known as Time-To-Contact (TTC) perception, have been mainly studied through noninvasive correlational approaches. We carried out an experiment in which patients were tested during an awake brain surgery electrostimulation mapping to examine the causal implication of various brain areas in the street-crossing decision process. Forty patients were tested in a gap acceptance task before their surgery to establish a baseline performance. The task was individually adapted upon this baseline level and carried out during their surgery. We acquired and normalized to MNI space the coordinates of the functional areas that influenced task performance. A total of 103 stimulation sites were tested, allowing to establish a large map of the areas involved in the street-crossing decision. Multiple sites were found to impact the gap acceptance decision. A direct implication was however found mostly for sites within the right parietal lobe, while indirect implication was found for sites within the language, motor, or attentional networks. The right parietal lobe can be considered as causally influencing the gap acceptance decision. Other positive sites were all accompanied with dysfunction in other cognitive functions, and therefore should probably not be considered as the site of TTC estimation.

Functional characterization of macaque insula using task-based and resting-state fMRI.

Neurophysiological investigations over the past decades have demonstrated the involvement of the primate insula in a wide array of sensory, cognitive, affective and regulatory functions, yet the complex functional organization of the insula remains unclear. Here we examined to what extent non-invasive task-based and resting-state fMRI provides support for functional specialization and integration of sensory and motor information in the macaque insula. Task-based fMRI experiments suggested a functional specialization related to processing of ingestive/taste/distaste information in anterior insula, grasping-related sensorimotor responses in middle insula and vestibular information in posterior insula. Visual stimuli depicting social information involving conspecific`s lip-smacking gestures yielded responses in middle and anterior portions of dorsal and ventral insula, overlapping partially with the sensorimotor and ingestive/taste/distaste fields. Functional specialization/integration of the insula was further corroborated by seed-based whole brain resting-state analyses, showing distinct functional connectivity gradients across the anterio-posterior extent of both dorsal and ventral insula. Posterior insula showed functional correlations in particular with vestibular/optic flow network regions, mid-dorsal insula with vestibular/optic flow as well as parieto-frontal regions of the sensorimotor grasping network, mid-ventral insula with social/affiliative network regions in temporal, cingulate and prefrontal cortices and anterior insula with taste and mouth motor networks including premotor and frontal opercular regions.

Towards an optimization of functional localizers in non-human primate neuroimaging with (fMRI) frequency-tagging.

Non-human primate (NHP) neuroimaging can provide essential insights into the neural basis of human cognitive functions. While functional magnetic resonance imaging (fMRI) localizers can play an essential role in reaching this objective (Russ et al., 2021), they often differ substantially across species in terms of paradigms, measured signals, and data analysis, biasing the comparisons. Here we introduce a functional frequency-tagging face localizer for NHP imaging, successfully developed in humans and outperforming standard face localizers (Gao et al., 2018). FMRI recordings were performed in two awake macaques. Within a rapid 6 Hz stream of natural non-face objects images, human or monkey face stimuli were presented in bursts every 9 s. We also included control conditions with phase-scrambled versions of all images. As in humans, face-selective activity was objectively identified and quantified at the peak of the face-stimulation frequency (0.111 Hz) and its second harmonic (0.222 Hz) in the Fourier domain. Focal activations with a high signal-to-noise ratio were observed in regions previously described as face-selective, mainly in the STS (clusters PL, ML, MF; also, AL, AF), both for human and monkey faces. Robust face-selective activations were also found in the prefrontal cortex of one monkey (PVL and PO clusters). Face-selective neural activity was highly reliable and excluded all contributions from low-level visual cues contained in the amplitude spectrum of the stimuli. These observations indicate that fMRI frequency-tagging provides a highly valuable approach to objectively compare human and monkey visual recognition systems within the same framework.

Symmetry Processing in the Macaque Visual Cortex.

Symmetry is a highly salient feature of the natural world that is perceived by many species. In humans, the cerebral areas processing symmetry are now well identified from neuroimaging measurements. Macaque could constitute a good animal model to explore the underlying neural mechanisms, but a previous comparative study concluded that functional magnetic resonance imaging responses to mirror symmetry in this species were weaker than those observed in humans. Here, we re-examined symmetry processing in macaques from a broader perspective, using both rotation and reflection symmetry embedded in regular textures. Highly consistent responses to symmetry were found in a large network of areas (notably in areas V3 and V4), in line with what was reported in humans under identical experimental conditions. Our results suggest that the cortical networks that process symmetry in humans and macaques are potentially more similar than previously reported and point toward macaque as a relevant model for understanding symmetry processing.

Time-to-contact perception in the brain.

Time-to-contact (TTC) perception refers to the ability of an observer to estimate the remaining time before an object reaches a point in the environment, and is of crucial importance in daily life. Noninvasive correlational approaches have identified several brain areas sensitive to TTC information. Here we report the results of two studies, including one during an awake brain surgery, that aimed to identify the specific areas causally engaged in the TTC estimation process. In Study 1, we tested 40 patients with brain tumor in a TTC estimation task. The results showed that four of the six patients with impaired performance had tumors in right upper parietal cortex, although this tumoral location represented only six over 40 patients. In Study 2, 15 patients underwent awake brain surgery electrostimulation mapping to examine the implication of various brain areas in the TTC estimation process. We acquired and normalized to MNI space the coordinates of the functional areas that influenced task performance. Our results seem to demonstrate that the early stage of the TTC estimation process involved specific cortical territories in the ventral region of the right intraparietal sulcus. Downstream processing of TTC could also involve the frontal eye field (middle frontal gyrus) related to ocular search. We also found that deactivating language areas in the left hemisphere interfered with the TTC estimation process. These findings demonstrate a fine grained, cortical representation of TTC processing close to the ventral right intraparietal sulcus and complement those described in other human studies.

Stereomotion Processing in the Nonhuman Primate Brain.

The cortical areas that process disparity-defined motion-in-depth (i.e., cyclopean stereomotion [CSM]) were characterized with functional magnetic resonance imaging (fMRI) in two awake, behaving macaques. The experimental protocol was similar to previous human neuroimaging studies. We contrasted the responses to dynamic random-dot patterns that continuously changed their binocular disparity over time with those to a control condition that shared the same properties, except that the temporal frames were shuffled. A whole-brain voxel-wise analysis revealed that in all four cortical hemispheres, three areas showed consistent sensitivity to CSM. Two of them were localized respectively in the lower bank of the superior temporal sulcus (CSMSTS) and on the neighboring infero-temporal gyrus (CSMITG). The third area was situated in the posterior parietal cortex (CSMPPC). Additional regions of interest-based analyses within retinotopic areas defined in both animals indicated weaker but significant responses to CSM within the MT cluster (most notably in areas MSTv and FST). Altogether, our results are in agreement with previous findings in both human and macaque and suggest that the cortical areas that process CSM are relatively well preserved between the two primate species.

Functional architecture of the motor homunculus detected by electrostimulation.

KEY POINTS: We performed a prospective electrostimulation study of the motor homunculus in 100 patients without motor deficit or brain lesion in the precentral gyrus in order to acquire accurate Montreal Neurological Institute (MNI) coordinates of the functional areas. The analysis of 248 body coordinates in the precentral gyrus showed rare inter-individual variations in the medial-to-lateral somatotopic movement organization with quite similar intensity thresholds. Electrostimulation only induced basic and stereotyped movements. We detected a relative medial-to-lateral somatotopy of the wrist/hand/global/individual fingers, with sometimes different sites for an individual muscle or movement. We found some similarities to, but also substantial differences from, the seminal work of Penfield and colleagues. We propose an updated version of the human motor homunculus and of its correlation with the somatosensory homunculus, previously defined in MNI space with a similar brain mapping technique. ABSTRACT: In this prospective electrostimulation study, based on 100 operated patients without motor deficit or brain lesion in the precentral gyrus, we acquired coordinates of the functional areas of the motor homunculus and normalized them to standard MNI space. Among 608 sites stimulated in the precentral gyrus (and 1937 in gyri nearby), 248 positive points (40%) for motor response were detected - 245 in the precentral gyrus. Positive stimulations were detected through the 'on/off' outbreak effect, and only basic movements were detected. We found no significant difference in mean intensity threshold between the motor representations of the fingers (1.94 mA), tongue and lower limbs (both 2.0 mA), or face (2.25 mA). In the precentral gyrus, the evoked body movements displayed a medial-to-lateral somatotopy in very small (often <10 mm2 ) areas. The hand region displayed multiple areas for a specific movement, with areas inducing either global or single-finger movement (with a relative medial-to-lateral somatotopy). Among these tested patients, the somatotopic organization of the intact motor cortex showed little inter-individual variations. Unlike Penfield and collaborators, we evoked no sensations such as sense of movement or desire to move, and only 2% of motor responses outside the precentral gyrus. We propose a rationalization of the standard drawing of the motor homunculus according to MNI space. We found a somatotopic correlation perpendicular to the central sulcus when matching our motor data to those previously obtained for the somatosensory homunculus.

Asymmetry of pictorial space: A cultural phenomenon.

Art experts have argued that the mirror reversal of pictorial artworks produces an alteration of their spatial content. However, this putative asymmetry of the pictorial space remains to be empirically proved and causally explained. Here, we address these issues with the "corridor illusion," a size illusion triggered by the pictorial space of a receding corridor. We show that mirror-reversed corridors-receding respectively leftward and rightward-induce markedly different illusion strengths and thus convey distinct pictorial spaces. Remarkably, the illusion is stronger with the rightward corridor among native left-to-right readers (French participants, n = 40 males) but conversely stronger with the leftward corridor among native right-to-left readers (Syrian participants, n = 40 males). Together, these results demonstrate an asymmetry of the pictorial space and point to our reading/writing habits as a major cause of this phenomenon.

Functional architecture of the somatosensory homunculus detected by electrostimulation.

KEY POINTS: We performed a prospective electrostimulation study, based on 50 operated intact patients, to acquire accurate MNI coordinates of the functional areas of the somatosensory homunculus. In the contralateral BA1, the hand representation displayed not only medial-to-lateral, little-finger-to-thumb, but also rostral-to-caudal discrete somatotopy, with the tip of each finger located more caudally than the proximal phalanx. The analysis of the MNI body coordinates showed rare inter-individual variations in the medial-to-lateral somatotopic organization in these patients with rather different intensity thresholds needed to elicit sensations in different body parts. We found some similarities but also substantial differences with the previous, seminal works of Penfield and his colleagues. We propose a new drawing of the human somatosensory homunculus according to MNI space. ABSTRACT: In this prospective electrostimulation study, based on 50 operated patients with no sensory deficit and no brain lesion in the postcentral gyrus, we acquired coordinates in the standard MNI space of the functional areas of the somatosensory homunculus. The 3D brain volume of each patient was normalized to that space to obtain the MNI coordinates of the stimulation site locations. For 647 sites stimulated on Brodmann Area 1 (and 1025 in gyri nearby), 258 positive points for somatosensory response (40%) were found in the postcentral gyrus. In the contralateral BA1, the hand representation displayed not only medial-to-lateral and little-finger-to-thumb somatotopy, but also rostral-to-caudal discrete somatotopy, with the tip of each finger located more caudally than the proximal phalanx. We detected a medial-to-lateral, tip-to-base tongue organization but no rostral-to-caudal functional organization. The analysis of the MNI body coordinates showed rare inter-individual variations in the medial-to-lateral somatotopic organization in these patients with intact somatosensory cortex. Positive stimulations were detected through the 'on/off' outbreak effect and discriminative touch sensations were the sensations reported almost exclusively by all patients during stimulation. Mean hand (2.39 mA) and tongue (2.60 mA) positive intensity thresholds were lower (P < 0.05) than the intensities required to elicit sensations in the other parts of the body. Unlike the previous, seminal works of Penfield and colleagues, we detected no sensations such as sense of movement or desire to move, no somatosensory responses outside the postcentral gyrus, and no bilateral responses for face/tongue stimulations. We propose a rationalization of the standard drawing of the somatosensory homunculus according to MNI space.

Processing of Egomotion-Consistent Optic Flow in the Rhesus Macaque Cortex.

The cortical network that processes visual cues to self-motion was characterized with functional magnetic resonance imaging in 3 awake behaving macaques. The experimental protocol was similar to previous human studies in which the responses to a single large optic flow patch were contrasted with responses to an array of 9 similar flow patches. This distinguishes cortical regions where neurons respond to flow in their receptive fields regardless of surrounding motion from those that are sensitive to whether the overall image arises from self-motion. In all 3 animals, significant selectivity for egomotion-consistent flow was found in several areas previously associated with optic flow processing, and notably dorsal middle superior temporal area, ventral intra-parietal area, and VPS. It was also seen in areas 7a (Opt), STPm, FEFsem, FEFsac and in a region of the cingulate sulcus that may be homologous with human area CSv. Selectivity for egomotion-compatible flow was never total but was particularly strong in VPS and putative macaque CSv. Direct comparison of results with the equivalent human studies reveals several commonalities but also some differences.

Rewarding objects appear larger but not brighter.

Whether reward can accentuate the perception of visual objects, that is, makes them appear larger than they really are, is a long-standing and controversial question. Here, we revisit this issue with a novel two-alternative forced-choice paradigm combining asymmetric reward schedule and task reversal. In a first experiment, participants (n = 27) choose the larger of two unequally rewarded objects in some sessions and the smaller one in other sessions. Response biases toward the most rewarding object differ significantly between the reversed tasks, revealing an influence of reward on perceived sizes. In a second experiment, participants (n = 27) indicate either the brighter or darker object. In contrast with the first experiment, response biases are similar between those reversed tasks, indicating that the perceived luminance is immune to reward manipulation. Together, these results reveal that if two objects are associated with different amounts of reward, participants will perceive the more rewarded object to be slightly larger, but not brighter, than the less rewarded one.

New insights for estimating the genetic value of segregating apple progenies for irregular bearing during the first years of tree production.

Because irregular bearing generates major agronomic issues in fruit-tree species, particularly in apple, the selection of regular cultivars is desirable. Here, we aimed to define methods and descriptors allowing a diagnostic for bearing behaviour during the first years of tree maturity, when tree production is increasing. Flowering occurrences were collected at whole-tree and (annual) shoot scales on a segregating apple population. At both scales, the number of inflorescences over the years was modelled. Two descriptors were derived from model residuals: a new biennial bearing index, based on deviation around yield trend over years and an autoregressive coefficient, which represents dependency between consecutive yields. At the shoot scale, entropy was also considered to represent the within-tree flowering synchronicity. Clusters of genotypes with similar bearing behaviours were built. Both descriptors at the whole-tree and shoot scales were consistent for most genotypes and were used to discriminate regular from biennial and irregular genotypes. Quantitative trait loci were detected for the new biennial bearing index at both scales. Combining descriptors at a local scale with entropy showed that regular bearing at the tree scale may result from different strategies of synchronization in flowering at the local scale. The proposed methods and indices open an avenue to quantify bearing behaviour during the first years of tree maturity and to capture genetic variations. Their extension to other progenies and species, possible variants of descriptors, and their use in breeding programmes considering a limited number of years or fruit yields are discussed.

Default mode of brain function in monkeys.

Human neuroimaging has revealed a specific network of brain regions-the default-mode network (DMN)-that reduces its activity during goal-directed behavior. So far, evidence for a similar network in monkeys is mainly indirect, since, except for one positron emission tomography study, it is all based on functional connectivity analysis rather than activity increases during passive task states. Here, we tested whether a consistent DMN exists in monkeys using its defining property. We performed a meta-analysis of functional magnetic resonance imaging data collected in 10 awake monkeys to reveal areas in which activity consistently decreases when task demands shift from passive tasks to externally oriented processing. We observed task-related spatially specific deactivations across 15 experiments, implying in the monkey a functional equivalent of the human DMN. We revealed by resting-state connectivity that prefrontal and medial parietal regions, including areas 9/46d and 31, respectively, constitute the DMN core, being functionally connected to all other DMN areas. We also detected two distinct subsystems composed of DMN areas with stronger functional connections between each other. These clusters included areas 24/32, 8b, and TPOC and areas 23, v23, and PGm, respectively. Such a pattern of functional connectivity largely fits, but is not completely consistent with anatomical tract tracing data in monkeys. Also, analysis of afferent and efferent connections between DMN areas suggests a multisynaptic network structure. Like humans, monkeys increase activity during passive epochs in heteromodal and limbic association regions, suggesting that they also default to internal modes of processing when not actively interacting with the environment.

Effects of contrast and contrast adaptation on static receptive field features in macaque area V1.

The spatiotemporal features of the "static" receptive field (RF), as revealed with flashing bars or spots, determine other RF properties. We examined how some of these static RF features vary with contrast and contrast adaptation in area V1 of the anesthetized macaque monkey. RFs were mapped with light and dark flashing bars presented at three different contrasts, with the low and medium contrasts eliciting approximately 1/3 and 2/3 of the high-contrast response amplitude. The main results are as follows: 1) RF widths decreased when contrast decreased; however, the amount of decrease was less than that expected from an iceberg model and closer to the expectation of a contrast invariance of the RF width. 2) Area tuning experiments with drifting gratings showed an opposite effect of contrast: an increase in preferred stimulus diameter when contrast decreased. This implies that the effect of contrast on preferred stimulus size is not predictable from the static RF. 3) Contrast adaptation attenuated the effect of contrast on RF amplitude but did not significantly modify RF width. 4) RF subregion overlap was only marginally affected by changes in contrast and contrast adaptation; the classification of cells as simple and complex, when established from subregion overlap, appears to be robust with respect to changes in contrast and adaptation state. Previous studies have shown that the spatiotemporal features of the RF depend largely on the stimuli used to map the RF. This study shows that contrast is one elemental feature that contributes to the dynamics of the RF.

Privileged visual processing of the straight-ahead direction in humans.

At any moment, the objects we face are endowed with a special behavioral status, either as potential obstacles during navigation or as optimal targets for visually guided actions. Yet, the gaze frequently jumps from one location to another when exploring the visual surroundings, so that objects located straight-ahead are often seen from the corner of the eyes. In the present study, we tested the hypothesis that peripheral vision might nevertheless ensure a privileged processing of these behaviorally important objects. Human subjects were asked to respond as fast as possible to the appearance of visual objects in their peripheral field of view while gazing successively in different directions. The visual objects formed similar images on the retina and differed only with respect to their egocentric location: either straight-ahead or eccentric with respect to the head/body midline. We found that straight-ahead objects elicit consistently shorter behavioral responses than eccentric objects (median difference of at least 10 ms). Additional experiments indicate that neither binocular visual cues nor full attentional resources play a fundamental role in this mechanism, and that it cannot be resumed to a simple preference for objects contralateral to the direction of gaze. These results are in agreement with recent electrophysiological findings showing that the early integration of gaze-related signals in the visual cortex of macaque monkeys lead to a higher neuronal sensitivity to the straight-ahead direction.

Hidden Semi-Markov Models to Segment Reading Phases from Eye Movements.

Our objective is to analyze scanpaths acquired through participants achieving a reading task aiming at answering a binary question: Is the text related or not to some given target topic? We propose a data-driven method based on hidden semi-Markov chains to segment scanpaths into phases deduced from the model states, which are shown to represent different cognitive strategies: normal reading, fast reading, information search, and slow confirmation. These phases were confirmed using different external covariates, among which semantic information extracted from texts. Analyses highlighted some strong preference of specific participants for specific strategies and more globally, large individual variability in eye-movement characteristics, as accounted for by random effects. As a perspective, the possibility of improving reading models by accounting for possible heterogeneity sources during reading is discussed.

Anterior regions of monkey parietal cortex process visual 3D shape.

The intraparietal cortex is involved in the control of visually guided actions, like reach-to-grasp movements, which require extracting the 3D shape and position of objects from 2D retinal images. Using fMRI in behaving monkeys, we investigated the role of the intraparietal cortex in processing stereoscopic information for recovering the depth structure and the position in depth of objects. We found that while several areas (CIP, LIP, and AIP on the lateral bank; PIP and MIP on the medial bank) are activated by stereoscopic stimuli, AIP and an adjoining portion of LIP are sensitive only to depth structure. Furthermore, only these two regions are sensitive to both the depth structure and the 2D shape of small objects. These results indicate that extracting 3D spatial information from stereo involves several intraparietal areas, among which AIP and anterior LIP are more specifically engaged in extracting the 3D shape of objects.

An egocentric straight-ahead bias in primate's vision.

As we plan to reach or manipulate objects, we generally orient our body so as to face them. Other objects occupying the same portion of space will likely represent potential obstacles for the intended action. Thus, either as targets or as obstacles, the objects located straight in front of us are often endowed with a special behavioral status. Here, we review a set of recent electrophysiological, imaging and behavioral studies bringing converging evidence that the objects which lie straight-ahead are subject to privileged visual processing. More precisely, these works collectively demonstrate that when gaze steers central vision away from the straight-ahead direction, the latter is still prioritized in peripheral vision. Straight-ahead objects evoke (1) stronger neuronal responses in macaque peripheral V1 neurons, (2) stronger EEG and fMRI activations across the human visual cortex and (3) faster reactive hand and eye movements. Here, we discuss the functional implications and underlying mechanisms behind this phenomenon. Notably, we propose that it can be considered as a new type of visuospatial attentional mechanism, distinct from the previously documented classes of endogenous and exogenous attention.

Wide-field retinotopy reveals a new visuotopic cluster in macaque posterior parietal cortex.

We investigated the visuotopic organization of macaque posterior parietal cortex (PPC) by combining functional imaging (fMRI) and wide-field retinotopic mapping in two macaque monkeys. Whole brain blood-oxygen-level-dependent (BOLD) signal was recorded while monkeys maintained central fixation during the presentation of large rotating wedges and expending/contracting annulus of a "shaking" fruit basket, designed to maximize the recruitment of PPC neurons. Results of the surface-based population receptive field (pRF) analysis reveal a new cluster of four visuotopic areas at the confluence of the parieto-occipital and intra-parietal sulci, in a location previously defined histologically and anatomically as the posterior intra-parietal (PIP) region. This PIP cluster groups together two recently described areas (CIP1/2) laterally and two newly identified ones (PIP1/2) medially, whose foveal representations merge in the fundus of the intra-parietal sulcus. The cluster shares borders with other visuotopic areas: V3d posteriorly, V3A/DP laterally, V6/V6A medially and LIP anteriorly. Together, these results show that monkey PPC is endowed with a dense set of visuotopic areas, as its human counterpart. The fact that fMRI and wide-field stimulation allows a functional parsing of monkey PPC offers a new framework for studying functional homologies with human PPC.