Role of handedness-related vestibular cortical dominance upon the vestibular–ocular reflex

Letter to the Editors

Face segregation and recognition by cortical multi-scale line and edge coding

Models of visual perception are based on image representations in cortical area V1 and higher areas which contain many cell layers for feature extraction. Basic simple, complex and end-stopped cells provide input for line, edge and keypoint detection. In this paper we present an improved method for multi-scale line/edge detection based on simple and complex cells. We illustrate the line/edge representation for object reconstruction, and we present models for multi-scale face (object) segregation and recognition that can be embedded into feedforward dorsal and ventral data streams (the “what” and “where” subsystems) with feedback streams from higher areas for obtaining translation, rotation and scale invariance.

Multi-scale cortical keypoint representation for attention and object detection

Keypoints (junctions) provide important information for focus-of-attention (FoA) and object categorization/recognition. In this paper we analyze the multi-scale keypoint representation, obtained by applying a linear and quasi-continuous scaling to an optimized model of cortical end-stopped cells, in order to study its importance and possibilities for developing a visual, cortical architecture.We show that keypoints, especially those which are stable over larger scale intervals, can provide a hierarchically structured saliency map for FoA and object recognition. In addition, the application of non-classical receptive field inhibition to keypoint detection allows to distinguish contour keypoints from texture (surface) keypoints.

Face recognition by cortical multi-scale line and edge representations

Empirical studies concerning face recognition suggest that faces may be stored in memory by a few canonical representations. Models of visual perception are based on image representations in cortical area V1 and beyond, which contain many cell layers for feature extraction. Simple, complex and end-stopped cells provide input for line, edge and keypoint detection. Detected events provide a rich, multi-scale object representation, and this representation can be stored in memory in order to identify objects. In this paper, the above context is applied to face recognition. The multi-scale line/edge representation is explored in conjunction with keypoint-based saliency maps for Focus-of-Attention. Recognition rates of up to 96% were achieved by combining frontal and 3/4 views, and recognition was quite robust against partial occlusions.

Face normalization using multi-scale cortical keypoints

Empirical studies concerning face recognition suggest that faces may be stored in memory by a few canonical representations. Models of visual perception are based on image representations in cortical area V1 and beyond, which contain many cell layers for feature extractions. Simple, complex and end-stopped cells tuned to different spatial frequencies (scales) and/or orientations provide input for line, edge and keypoint detection. This yields a rich, multi-scale object representation that can be stored in memory in order to identify objects. The multi-scale, keypoint-based saliency maps for Focus-of-Attention can be explored to obtain face detection and normalization, after which face recognition can be achieved using the line/edge representation. In this paper, we focus only on face normalization, showing that multi-scale keypoints can be used to construct canonical representations of faces in memory.

Cortical object segregation and categorization by multi-scale line and edge coding

In this paper we present an improved scheme for line and edge detection in cortical area V1, based on responses of simple and complex cells, truly multi-scale with no free parameters. We illustrate the multi-scale representation for visual reconstruction, and show how object segregation can be achieved with coarse-to-finescale groupings. A two-level object categorization scenario is tested in which pre-categorization is based on coarse scales only, and final categorization on coarse plus fine scales. Processing schemes are discussed in the framework of a complete cortical architecture.

Recognition of facial expressions by cortical multi-scale line and edge coding

Empirical studies concerning face recognition suggest that faces may be stored in memory by a few canonical representations. Models of visual perception are based on image representations in cortical area V1 and beyond, which contain many cell layers for feature extraction. Simple, complex and end-stopped cells provide input for line, edge and keypoint detection. Detected events provide a rich, multi-scale object representation, and this representation can be stored in memory in order to identify objects. In this paper, the above context is applied to face recognition. The multi-scale line/edge representation is explored in conjunction with keypoint-based saliency maps for Focus-of-Attention. Recognition rates of up to 96% were achieved by combining frontal and 3/4 views, and recognition was quite robust against partial occlusions.

Cortical 3D Face Recognition Framework

Empirical studies concerning face recognition suggest that faces may be stored in memory by a few canonical representations. In cortical area V1 exist double-opponent colour blobs, also simple, complex and end-stopped cells which provide input for a multiscale line/edge representation, keypoints for dynamic routing and saliency maps for Focus-of-Attention. All these combined allow us to segregate faces. Events of different facial views are stored in memory and combined in order to identify the view and recognise the face including facial expression. In this paper we show that with five 2D views and their cortical representations it is possible to determine the left-right and frontal-lateral-profile views and to achieve view-invariant recognition of 3D faces.

Multi-scale cortical keypoints for realtime hand tracking and gesture recognition

Human-robot interaction is an interdisciplinary research area which aims at integrating human factors, cognitive psychology and robot technology. The ultimate goal is the development of social robots. These robots are expected to work in human environments, and to understand behavior of persons through gestures and body movements. In this paper we present a biological and realtime framework for detecting and tracking hands. This framework is based on keypoints extracted from cortical V1 end-stopped cells. Detected keypoints and the cells’ responses are used to classify the junction type. By combining annotated keypoints in a hierarchical, multi-scale tree structure, moving and deformable hands can be segregated, their movements can be obtained, and they can be tracked over time. By using hand templates with keypoints at only two scales, a hand’s gestures can be recognized.

An exploration of pre-attentive visual discrimination using event-related potentials

The Mismatch Negativity (MMN) has been characterised as a ‘pre-attentive’ component of an Event-Related Potential (ERP) that is related to discriminatory processes. Although well established in the auditory domain, characteristics of the MMN are less well characterised in the visual domain. The five main studies presented in this thesis examine visual cortical processing using event-related potentials. Novel methodologies have been used to elicit visual detection and discrimination components in the absence of a behavioural task. Developing paradigms in which a behavioural task is not required may have important clinical applications for populations, such as young children, who cannot comply with the demands of an active task. The ‘pre-attentive’ nature of visual MMN has been investigated by modulating attention. Generators and hemispheric lateralisation of visual MMN have been investigated by using pertinent clinical groups. A three stimulus visual oddball paradigm was used to explore the elicitation of visual discrimination components to a change in the orientation of stimuli in the absence of a behavioural task. Monochrome stimuli based on pacman figures were employed that differed from each other only in terms of the orientation of their elements. One such stimulus formed an illusory figure in order to capture the participant’s attention, either in place of, or alongside, a behavioural task. The elicitation of a P3a to the illusory figure but not to the standard or deviant stimuli provided evidence that the illusory figure captured attention. A visual MMN response was recorded in a paradigm with no task demands. When a behavioural task was incorporated into the paradigm, a P3b component was elicited consistent with the allocation of attentional resources to the task. However, visual discrimination components were attenuated revealing that the illusory figure was unable to command all attentional resources from the standard deviant transition. The results are the first to suggest that the visual MMN is modulated by attention. Using the same three stimulus oddball paradigm, generators of visual MMN were investigated by recording potentials directly from the cortex of an adolescent undergoing pre-surgical evaluation for resection of a right anterior parietal lesion. To date no other study has explicitly recorded activity related to the visual MMN intracranially using an oddball paradigm in the absence of a behavioural task. Results indicated that visual N1 and visual MMN could be temporally and spatially separated, with visual MMN being recorded more anteriorly than N1. The characteristic abnormality in retinal projections in albinism afforded the opportunity to investigate each hemisphere in relative isolation and was used, for the first time, as a model to investigate lateralisation of visual MMN and illusory contour processing. Using the three stimulus oddball paradigm, no visual MMN was elicited in this group, and so no conclusions regarding the lateralisation of visual MMN could be made. Results suggested that both hemispheres were equally capable of processing an illusory figure. As a method of presenting visual test stimuli without conscious perception, a continuous visual stream paradigm was developed that used a briefly presented checkerboard stimulus combined with masking for exploring stimulus detection below and above subjective levels of perception. A correlate of very early cortical processing at a latency of 60-80 ms (CI) was elicited whether stimuli were reported as seen or unseen. Differences in visual processing were only evident at a latency of 90 ms (CII) implying that this component may represent a correlate of visual consciousness/awareness. Finally, an oddball sequence was introduced into the visual stream masking paradigm to investigate whether visual MMN responses could be recorded without conscious perception. The stimuli comprised of black and white checkerboard elements differing only in terms of their orientation to form an x or a +. Visual MMN was not recorded when participants were unable to report seeing the stimulus. Results therefore suggest that behavioural identification of the stimuli was required for the elicitation of visual MMN and that visual MMN may require some attentional resources. On the basis of these studies it is concluded that visual MMN is not entirely independent of attention. Further, the combination of clinical and non-clinical investigations provides a unique opportunity to study the characterisation and localisation of putative mechanisms related to conscious and non-conscious visual processing.

Tracking the source of cerebellar epilepsy: hemifacial seizures associated with cerebellar cortical dysplasia.

Traditionally, subcortical structures such as the cerebellum are supposed to exert a modulatory effect on epileptic seizures, rather than being the primary seizure generator. We report a 14-month old girl presenting, since birth, with seizures symptomatic of a right cerebellar dysplasia, manifested as paroxystic contralateral hemifacial spasm and ipsilateral facial weakness. Multimodal imaging was used to investigate both anatomical landmarks related to the cerebellar lesion and mechanisms underlying seizure generation. Electric source imaging (ESI) supported the hypothesis of a right cerebellar epileptogenic generator in concordance with nuclear imaging findings; subsequently validated by intra-operative intralesional recordings. Diffusion spectrum imaging-related tractography (DSI) showed severe cerebellar structural abnormalities confirmed by histological examination. We suggest that hemispheric cerebellar lesions in cases like this are likely to cause epilepsy via an effect on the facial nuclei through ipsilateral and contralateral aberrant connections.

Effects of sodium arsenite on neurite outgrowth and glutamate AMPA receptor expression in mouse cortical neurons.

There has been broad concern that arsenic in the environment exerts neurotoxicity. To determine the mechanism by which arsenic disrupts neuronal development, primary cultured neurons obtained from the cerebral cortex of mouse embryos were exposed to sodium arsenite (NaAsO2) at concentrations between 0 and 2μM from days 2 to 4 in vitro and cell survival, neurite outgrowth and expression of glutamate AMPA receptor subunits were assessed at day 4 in vitro. Cell survival was significantly decreased by exposure to 2μM NaAsO2, whereas 0.5μM NaAsO2 increased cell survival instead. The assessment of neurite outgrowth showed that total neurite length was significantly suppressed by 1μM and 2μM NaAsO2, indicating that the lower concentration of NaAsO2 impairs neuritogenesis before inducing cell death. Immunoblot analysis of AMPA receptor subunit expression showed that the protein level of GluA1, a specific subunit of the AMPA receptor, was significantly decreased by 1μM and 2μM NaAsO2. When immunocytochemistry was used to confirm this effect by staining for GluA1 expression in neuropeptide Y neurons, most of which contain GluA1, GluA1 expression in neuropeptide Y neurons was found to be significantly suppressed by 1μM and 2μM NaAsO2 but to be increased at the concentration of 0.5μM. Finally, to determine whether neurons could be rescued from the NaAsO2-induced impairment of neuritogenesis by compensatory overexpression of GluA1, we used primary cultures of neurons transfected with a plasmid vector to overexpress either GluA1 or GluA2, and the results showed that GluA1/2 overexpression protected against the deleterious effects of NaAsO2 on neurite outgrowth. These results suggest that the NaAsO2 concentration inducing neurite suppression is lower than the concentration that induces cell death and is the same as the concentration that suppresses GluA1 expression. Consequently, the suppression of GluA1 expression by NaAsO2 seems at least partly responsible for neurite suppression induced by NaAsO2.

Sub-cortical and brainstem sites associated with chemo-stimulated increases in ventilation in humans.

We investigated the neural basis for spontaneous chemo-stimulated increases in ventilation in awake, healthy humans. Blood oxygen level dependent (BOLD) functional MRI was performed in nine healthy subjects using T2 weighted echo planar imaging. Brain volumes (52 transverse slices, cortex to high spinal cord) were acquired every 3.9 s. The 30 min paradigm consisted of six, 5-min cycles, each cycle comprising 45 s of hypoxic-isocapnia, 45 s of isooxic-hypercapnia and 45 s of hypoxic-hypercapnia, with 55 s of non-stimulatory hyperoxic-isocapnia (control) separating each stimulus period. Ventilation was significantly (p<0.001) increased during hypoxic-isocapnia, isooxic-hypercapnia and hypoxic-hypercapnia (17.0, 13.8, 24.9 L/min respectively) vs. control (8.4 L/min) and was associated with significant (p<0.05, corrected for multiple comparisons) signal increases within a bilateral network that included the basal ganglia, thalamus, red nucleus, cerebellum, parietal cortex, cingulate and superior mid pons. The neuroanatomical structures identified provide evidence for the spontaneous control of breathing to be mediated by higher brain centres, as well as respiratory nuclei in the brainstem.

The Na+-dependent chloride-bicarbonate exchanger SLC4A8 mediates an electroneutral Na+ reabsorption process in the renal cortical collecting ducts of mice.

Regulation of sodium balance is a critical factor in the maintenance of euvolemia, and dysregulation of renal sodium excretion results in disorders of altered intravascular volume, such as hypertension. The amiloride-sensitive epithelial sodium channel (ENaC) is thought to be the only mechanism for sodium transport in the cortical collecting duct (CCD) of the kidney. However, it has been found that much of the sodium absorption in the CCD is actually amiloride insensitive and sensitive to thiazide diuretics, which also block the Na-Cl cotransporter (NCC) located in the distal convoluted tubule. In this study, we have demonstrated the presence of electroneutral, amiloride-resistant, thiazide-sensitive, transepithelial NaCl absorption in mouse CCDs, which persists even with genetic disruption of ENaC. Furthermore, hydrochlorothiazide (HCTZ) increased excretion of Na+ and Cl- in mice devoid of the thiazide target NCC, suggesting that an additional mechanism might account for this effect. Studies on isolated CCDs suggested that the parallel action of the Na+-driven Cl-/HCO3- exchanger (NDCBE/SLC4A8) and the Na+-independent Cl-/HCO3- exchanger (pendrin/SLC26A4) accounted for the electroneutral thiazide-sensitive sodium transport. Furthermore, genetic ablation of SLC4A8 abolished thiazide-sensitive NaCl transport in the CCD. These studies establish what we believe to be a novel role for NDCBE in mediating substantial Na+ reabsorption in the CCD and suggest a role for this transporter in the regulation of fluid homeostasis in mice.

Oxygen tension controls the expression of the monocarboxylate transporter MCT4 in cultured mouse cortical astrocytes via a hypoxia-inducible factor-1α-mediated transcriptional regulation.

The monocarboxylate transporter MCT4 is a high capacity carrier important for lactate release from highly glycolytic cells. In the central nervous system, MCT4 is predominantly expressed by astrocytes. Surprisingly, MCT4 expression in cultured astrocytes is low, suggesting that a physiological characteristic, not met in culture conditions, is necessary. Here we demonstrate that reducing oxygen concentration from 21% to either 1 or 0% restored in a concentration-dependent manner the expression of MCT4 at the mRNA and protein levels in cultured astrocytes. This effect was specific for MCT4 since the expression of MCT1, the other astrocytic monocarboxylate transporter present in vitro, was not altered in such conditions. MCT4 expression was shown to be controlled by the transcription factor hypoxia-inducible factor-1α (HIF-1α) since under low oxygen levels, transfecting astrocyte cultures with a siRNA targeting HIF-1α largely prevented MCT4 induction. Moreover, the prolyl hydroxylase inhibitor dimethyloxalylglycine (DMOG) induced MCT4 expression in astrocytes cultured in presence of 21% oxygen. In parallel, glycolytic activity was enhanced by exposure to 1% oxygen as demonstrated by the increased lactate release, an effect dependent on MCT4 expression. Finally, MCT4 expression was found to be necessary for astrocyte survival when exposed for a prolonged period to 1% oxygen. These data suggest that a major determinant of astrocyte MCT4 expression in vivo is likely the oxygen tension. This could be relevant in areas of high neuronal activity and oxygen consumption, favouring astrocytic lactate supply to neurons. Moreover, it could also play an important role for neuronal recovery after an ischemic episode.