Modulation of V1 activity by shape: image-statistics or shape-based perception?

It is current dogma that neurons in primary visual cortex extract local edges from the scene from which later visual areas reconstruct more meaningful shapes. Recent neuroimaging studies, however, have shown V1 modulations by the degree of structure in the image (shape). These V1 modulations due to the level of shape coherence have been explained in one of two possible ways: due to changes in image statistics or shape-based perceptual influences from higher visual areas. Here we compare both hypotheses using stimuli composed of Gabor arrays constructed to form circular shapes that can be successively degraded by manipulating the orientations of individual Gabors while maintaining local and global statistics. In a first experiment, we confirm that V1 responses are inversely correlated with the degree of structure in the image. In a second experiment, stimulus predictions are compared based on the degree of circular shape or change in the image statistic varied (orientation variance) in the image. We find that these V1 modulations to shape change are correlated with low-level changes in orientation contrast rather than shape perception per se.     

Attentional capacity for processing concurrent stimuli is larger across sensory modalities than within a modality

One finding in attention research is that visual and auditory attention mechanisms are linked together. Such a link would predict a central, amodal capacity limit in processing visual and auditory stimuli. Here we show that this is not the case. Letter streams were accompanied by asynchronously presented streams of auditory, visual, and audiovisual objects. Either the letter streams or the visual, auditory, or audiovisual parts of the object streams were attended. Attending to various aspects of the objects resulted in modulations of the letter-stream-elicited steady-state evoked potentials (SSVEPs). SSVEPs were larger when auditory objects were attended than when either visual objects alone or when auditory and visual object stimuli were attended together. SSVEP amplitudes were the same in the latter conditions, indicating that attentional capacity between modalities is larger than attentional capacity within one and the same modality.     

Human primary visual cortex (V1) is selective for second-order spatial frequency

A variety of cues can differentiate objects from their surrounds. These include "first-order" cues such as luminance modulations and "second-order" cues involving modulations of orientation and contrast. Human sensitivity to first-order modulations is well described by a computational model involving spatially localized filters that are selective for orientation and spatial frequency (SF). It is widely held that first-order modulations are represented by the firing rates of simple and complex cells ("first-order" neurons) in primary visual cortex (V1) that, likewise, have spatially localized receptive fields that are selective for orientation- and SF. Human sensitivity to second-order modulations is well described by a filter-rectify-filter (FRF) model, with first- and second-order filters selective for orientation and SF. However, little is known about how neuronal activity in visual cortex represents second-order modulations. We tested the FRF model by using an functional (f)MRI-adaptation protocol to characterize the selectivity of activity in visual cortex to second-order, orientation-defined gratings of two different SFs. fMRI responses throughout early visual cortex exhibited selective adaptation to these stimuli. The low-SF grating was a more effective adapter than the high-SF grating, incompatible with the FRF model. To explain the results, we extended the FRF model by incorporating normalization, yielding a filter-rectify-normalize-filter model, in which normalization enhances selectivity for second-order SF but only for low spatial frequencies. We conclude that neurons in human visual cortex are selective for second-order SF, that normalization (surround suppression) contributes to this selectivity, and that the selectivity in higher visual areas is simply fed forward from V1.     

Task-related modulation of visual cortex

We performed a series of experiments to quantify the effects of task performance on cortical activity in early visual areas. Functional magnetic resonance imaging (fMRI) was used to measure cortical activity in several cortical visual areas including primary visual cortex (V1) and the MT complex (MT+) as subjects performed a variety of threshold-level visual psychophysical tasks. Performing speed, direction, and contrast discrimination tasks produced strong modulations of cortical activity. For example, one experiment tested for selective modulations of MT+ activity as subjects alternated between performing contrast and speed discrimination tasks. MT+ responses modulated in phase with the periods of time during which subjects performed the speed discrimination task; that is, MT+ activity was higher during speed discrimination than during contrast discrimination. Task-related modulations were consistent across repeated measurements in each subject; however, significant individual differences were observed between subjects. Together, the results suggest 1) that specific changes in the cognitive/behavioral state of a subject can exert selective and reliable modulations of cortical activity in early visual cortex, even in V1; 2) that there are significant individual differences in these modulations; and 3) that visual areas and pathways that are highly sensitive to small changes in a given stimulus feature (such as contrast or speed) are selectively modulated during discrimination judgments on that feature. Increasing the gain of the relevant neuronal signals in this way may improve their signal-to-noise to help optimize task performance.     

Vision enhances selective attention to body-related information

Viewing ones’ own hands while directing attention to one of the hands leads to earlier attentional modulations of somatosensory processing than when hands are not visible. This effect of vision on tactile-spatial selection could be explained by vision providing additional information about the location of the hands in external space. The present study investigated whether vision of the hands also affected tactile-attentional mechanisms when the relative locations of the hands were irrelevant. Participants silently counted infrequent tactile or auditory deviants in an alternating stream of tactile and auditory stimuli while ignoring stimuli in the other modality, when their hands were either visible or covered from view. Modality-selective attentional modulations of ERPs to tactile stimuli (when touches vs. tones were attended) were already present for the time range of the N80 component when hands were visible, but there were only later modulations (starting at N140) when hands were covered. This suggests that, rather than being restricted to tasks requiring spatial selection between body parts, vision of the hands can facilitate attention toward the body in far more general terms. In contrast to tactile stimuli, attentional modulations of ERPs to auditory stimuli (when tones vs. touches were attended) were not reliably affected by viewing the hands. This suggests that the primary purpose of visual facilitation may be to enhance the processing of body-related information only.     

Neural mechanisms of intermuscular coherence: implications for the rectification of surface electromyography

Oscillatory activity plays a crucial role in corticospinal control of muscle synergies and is widely investigated using corticospinal and intermuscular synchronization. However, the neurophysiological mechanisms that translate these rhythmic patterns into surface electromyography (EMG) are not well understood. This is underscored by the ongoing debate on the rectification of surface EMG before spectral analysis. Whereas empirical studies commonly rectify surface EMG, computational approaches have argued against it. In the present study, we employ a computational model to investigate the role of the motor unit action potential (MAUP) on the translation of oscillatory activity. That is, diverse MUAP shapes may distort the transfer of common input into surface EMG. We test this in a computational model consisting of two motor unit pools receiving common input and compare it to empirical results of intermuscular coherence between bilateral leg muscles. The shape of the MUAP was parametrically varied, and power and coherence spectra were investigated with and without rectification. The model shows that the effect of EMG rectification depends on the uniformity of MUAP shapes. When output spikes of different motor units are convolved with identical MUAPs, oscillatory input is evident in both rectified and nonrectified EMG. In contrast, a heterogeneous MAUP distribution distorts common input and oscillatory components are only manifest as periodic amplitude modulations, i.e., in rectified EMG. The experimental data showed that intermuscular coherence was mainly discernable in rectified EMG, hence providing empirical support for a heterogeneous distribution of MUAPs. These findings implicate that the shape of MUAPs is an essential parameter to reconcile experimental and computational approaches.     

Unitary haptic perception: integrating moving tactile inputs from anatomically adjacent and non-adjacent digits

How do we achieve unitary perception of an object when it touches two parts of the sensory epithelium that are not contiguous? We investigated this problem with a simple psychophysical task, which we then used in an fMRI experiment. Two wooden rods were moved over two digits positioned to be spatially adjacent. The digits were either from one foot (or hand) or one digit was from either foot (or hand). When the rods were moving in phase, one object was reliably perceived. By contrast, when the rods were moving out of phase, two objects were reliably perceived. fMRI revealed four cortical areas where activity was higher when the moving rods were perceived as one object relative to when they were perceived as two separate objects. Areas in the right inferior parietal lobule, the left inferior temporal sulcus and the left middle frontal gyrus were activated for this contrast regardless of the anatomical configuration of the stimulated sensory epithelia. By contrast, the left intraparietal sulcus was activated specifically when integration across the midline was required, irrespective of whether the stimulation was applied to the hands or feet. These results reveal a network of brain areas involved in generating a unified percept of the presence of an object that comes into contact with different parts of the body surface.     

The influence of target object shape on maximum grip aperture in human grasping movements

The shape of a target object could influence maximum grip aperture in human grasping movements in several different ways. Maximum grip aperture could be influenced by the required precision of digit placement, by the aim to avoid colliding with the wrong parts of the target objects, by the mass of the target objects, or by (mis)judging the width or the volume of the target objects. To examine the influence of these five factors, we asked subjects to grasp five differently shaped target objects with the same maximal width, height and depth and compared their maximum grip aperture with what one would expect for each of the five factors. The five target objects, a cube, a three-dimensional plus sign, a rectangular block, a cylinder and a sphere, were all grasped with the same final grip aperture. The experimentally observed maximum grip apertures correlated poorly with the maximum grip apertures that were expected on the basis of the required precision, the actual mass, the perceived width and the perceived volume. They correlated much better with the maximum grip apertures that were expected on the basis of avoiding unintended collisions with the target object. We propose that the influence of target object shape on maximum grip aperture might primarily be the result of the need to avoid colliding with the wrong parts of the target object.     

Temporal coherence sensitivity in auditory cortex

Natural sounds often contain energy over a broad spectral range and consequently overlap in frequency when they occur simultaneously; however, such sounds under normal circumstances can be distinguished perceptually (e.g., the cocktail party effect). Sound components arising from different sources have distinct (i.e., incoherent) modulations, and incoherence appears to be one important cue used by the auditory system to segregate sounds into separately perceived acoustic objects. Here we show that, in the primary auditory cortex of awake marmoset monkeys, many neurons responsive to amplitude- or frequency-modulated tones at a particular carrier frequency [the characteristic frequency (CF)] also demonstrate sensitivity to the relative modulation phase between two otherwise identically modulated tones: one at CF and one at a different carrier frequency. Changes in relative modulation phase reflect alterations in temporal coherence between the two tones, and the most common neuronal response was found to be a maximum of suppression for the coherent condition. Coherence sensitivity was generally found in a narrow frequency range in the inhibitory portions of the frequency response areas (FRA), indicating that only some off-CF neuronal inputs into these cortical neurons interact with on-CF inputs on the same time scales. Over the population of neurons studied, carrier frequencies showing coherence sensitivity were found to coincide with the carrier frequencies of inhibition, implying that inhibitory inputs create the effect. The lack of strong coherence-induced facilitation also supports this interpretation. Coherence sensitivity was found to be greatest for modulation frequencies of 16-128 Hz, which is higher than the phase-locking capability of most cortical neurons, implying that subcortical neurons could play a role in the phenomenon. Collectively, these results reveal that auditory cortical neurons receive some off-CF inputs temporally matched and some temporally unmatched to the on-CF input(s) and respond in a fashion that could be utilized by the auditory system to segregate natural sounds containing similar spectral components (such as vocalizations from multiple conspecifics) based on stimulus coherence.     

Common slow modulation of respiration, arterial blood pressure and cortical activity during sleep onset while napping.

Using healthy subjects, concomitant 30- to 60-s modulations of respiration, arterial blood pressure and EEG activity were found in 21 experiments about napping. Although mean arterial pressure (MAP) modulations above and below 1/30 Hz increased, in respiratory amplitude (RA) only the lower frequency components augmented significantly. This slow modulation of RA was found to be asymmetrical in time, the duration of RA decreasing parts in the modulation waves being 42% longer than the duration of RA increasing parts. The concomitance of the slow modulations in the different organ systems is accounted for by the influence of the common brainstem system (CBS), which regulates and integrates respiratory, cardiovascular and somatomotor systems and the adjustment of central nervous activity (vigilance). The described common, slow modulations outline the importance of dampening influences during sleep onset. They may provide an important tool for the investigation of the regulatory systems during sleep onset, as well as for investigations about sleep apnoea syndrome and Cheyne-Stokes breathing.     

Modulation of beta oscillations in the subthalamic area during action observation in Parkinson's disease

Mapping observed actions into the onlooker's motor system seems to provide the neurofunctional mechanisms for action understanding. Subthalamic nucleus (STN) local field potential (LFP) recordings in patients with movement disorders disclosed that network oscillations in the beta range are involved in conveying motor and non-motor information across the cortico-basal ganglia–thalamo-cortical loop. This evidence, together with the existence of connections between the STN and cortical areas active during observation of actions performed by other people, suggests that the STN oscillatory activity in specific frequency bands could encode not only motor information, but also information related to action observation. To test this hypothesis we directly recorded STN oscillations through electrodes for deep brain stimulation in patients with Parkinson's disease during observation of actions and of static objects. We found selective action-related oscillatory modulations in two functionally distinct beta bands: whereas low-beta oscillations (10–18 Hz) selectively desynchronized only during action-observation, high-beta oscillations (20–30 Hz) synchronized both during the observation of action and action-related objects. Low-beta modulations are therefore specific to action observation and high-beta modulations are related to the action scene. Our findings show that in the basal ganglia there are functional changes spreading to the action environment, probably presetting the motor system in relation to the motor context and suggesting that the dynamics of beta oscillations can contribute to action understanding mechanisms.     

Neural mechanisms of visual associative processing

This is a review of our work on multiple microelectrode recordings from the visual cortex of monkeys and subdural recordings from humans--related to the potential underlying neural mechanisms. The former hypothesis of object representation by synchronization in visual cortex (or more generally: of flexible associative processing) has been supported by our recent experiments in monkeys. They demonstrated local synchrony among rhythmic or stochastic gamma-activities (30-90 Hz) and perceptual modulation, according to the rules of figure-ground segregation. However, gamma-synchrony in primary visual cortex is restricted to few millimeters, challenging the synchronization hypothesis for larger cortical object representations. We found that the spatial restriction is due to gamma-waves, traveling in random directions across the object representations. It will be argued that phase continuity of these waves can support the coding of object continuity. Based on models with spiking neurons, potentially underlying neural mechanisms are proposed: (i) Fast inhibitory feedback loops can generate locally synchronized gamma-activities; (ii) Hebbian learning of lateral and feed forward connections with distance-dependent delays can explain the stabilization of cortical retinotopy, the limited size of synchronization, the occurrence of gamma-waves, and the larger receptive fields at successive levels; (iii) slow inhibitory feedback can support figure-ground segregation; (iv) temporal dispersion in far projections destroys coherence of fast signals but preserves slow amplitude modulations. In conclusion, it is proposed that the hypothesis of flexible associative processing by gamma-synchronization, including coherent representations of visual objects, has to be extended to more general forms of signal coupling.     

Brain state-dependency of coherent oscillatory activity in the cerebral cortex and basal ganglia of the rat

The nature of the coupling between neuronal assemblies in the cerebral cortex and basal ganglia (BG) is poorly understood. We tested the hypothesis that coherent population activity is dependent on brain state, frequency range, and/or BG nucleus using data from simultaneous recordings of electrocorticogram (ECoG) and BG local field potentials (LFPs) in anesthetized rats. The coherence between ECoG and LFPs simultaneously recorded from subthalamic nucleus (STN), globus pallidus (GP), and substantia nigra pars reticulata (SNr) was largely confined to slow- ( approximately 1 Hz) and spindle- (7-12 Hz) frequency oscillations during slow-wave activity (SWA). In contrast, during cortical activation, coherence was mostly restricted to high-frequency oscillations (15-60 Hz). The coherence between ECoG and LFPs also depended on BG recording site. Partial coherence analyses showed that, during SWA, STN and SNr shared the same temporal coupling with cortex, thereby forming a single functional axis. Cortex was also tightly, but independently, correlated with GP in a separate functional axis. During activation, STN, GP, and, to a lesser extent, SNr shared the same coherence with cortex as part of one functional axis. In addition, GP formed a second, independently coherent loop with cortex. These data suggest that coherent oscillatory activity is present at the level of LFPs recorded in cortico-basal ganglia circuits, and that synchronized population activity is dynamically organized according to brain state, frequency, and nucleus. These attributes further suggest that synchronized activity should be considered as one of a number of candidate mechanisms underlying the functional organization of these brain circuits.     

Assessment of across-muscle coherence using multi-unit vs. single-unit recordings

Coherence between electromyographic (EMG) signals has been used to identify correlated neural inputs to motor units (MUs) innervating different muscles. Simulations using a motor-unit model (Fuglevand et al. 1992) were performed to determine the ability of coherence between two multi-unit EMGs (mEMG) to detect correlated MU activity and the range of correlation strengths in which mEMG coherence can be usefully employed. Coherence between motor-unit and mEMG activities in two muscles was determined as we varied the strength of a 30-Hz periodic common input, the number of correlated MU pairs and variability of MU discharge relative to the common input. Pooled and mEMG coherence amplitudes positively and negatively accelerated, respectively, toward the strongest and most widespread correlating inputs. Furthermore, the relation between pooled and mEMG coherence was also nonlinear and was essentially the same whether correlation strength varied by changing common input strength or its distribution. However, the most important finding is that while the mEMG coherence saturates at the strongest common input strengths, this occurs at common input strengths greater than found in most physiological studies. Thus, we conclude that mEMG coherence would be a useful measure in many experimental conditions and our simulation results suggest further guidelines for using and interpreting coherence between mEMG signals.     

Antiphasic 40 Hz Oscillatory Current Stimulation Affects Bistable Motion Perception

When viewing ambiguous stimuli, conscious perception alternates spontaneously between competing interpretations of physically unchanged stimulus information. As one possible neural mechanism underlying the perceptual switches, it has been suggested that neurons dynamically change their pattern of synchronized oscillatory activity in the gamma band (30–80 Hz). In support of this hypothesis, there is correlative evidence from human electroencephalographic (EEG) studies for gamma band modulations during ambiguous perception. To establish a causal role of gamma band oscillations in the current study, we applied transcranial alternating current stimulation (tACS) at 40 Hz over occipital–parietal areas of both hemispheres during the presentation of bistable apparent motion stimuli that can be perceived as moving either horizontally or vertically. In this paradigm, the switch between horizontal and vertical apparent motion is likely to involve a change in interhemispheric functional coupling. We examined gamma tACS effects on the durations of perceived horizontal and vertical motion as well as on interhemispheric EEG coherence and found a decreased proportion of perceived horizontal motion together with an increase of interhemispheric gamma band coherence. In a control experiment using 6 Hz tACS, we did not observe any stimulation effects on behavior or coherence. Furthermore, external stimulation at 40 Hz was only effective when applied with 180° phase difference between hemispheres (anti-phase), as compared to in-phase stimulation with 0° phase difference. These findings suggest that externally desynchronizing gamma oscillations between hemispheres impairs interhemispheric motion integration and in turn biases conscious experience of bistable apparent motion.     

Improvement in the contrast of CEST MRI via intermolecular double quantum coherences

The tremendous potential of chemical exchange saturation transfer (CEST) agents as an emerging class of magnetic resonance imaging contrast media has been demonstrated in recent years. In a CEST experiment, a high CEST contrast is always welcome. However, when the exchange rate is low, which may happen in exchangeable solute protons of low concentration, it is usually hard to obtain an excellent CEST efficiency. Recently, we noted that the intermolecular multiple quantum coherence signal is more sensitive to the changes of the magnetization magnitude than a conventional single quantum coherence signal. Consequently, it may be easier when used in obtaining a CEST contrast. In this note, a modified COSY (two-dimensional correlated spectroscopy) revamped with an asymmetric Z-gradient echo detection (CRAZED) sequence combined with an off-resonance saturation pulse followed by a standard spin-echo imaging sequence was designed to obtain a better CEST contrast image based on the intermolecular double quantum coherence method. An analytical expression was derived from a dipolar field theory. Experiments were performed on an agar-glucose phantom, and the results demonstrate the feasibility of our method.     

Automated assessment of low contrast sensitivity for CT systems using a model observer

Purpose: Low contrast sensitivity of CT scanners is regularly assessed by subjective scoring of low contrast detectability within phantom CT images. Since in these phantoms low contrast objects are arranged in known fixed patterns, subjective rating of low contrast visibility might be biased. The purpose of this study was to develop and validate a software for automated objective low contrast detectability based on a model observer.Methods: Images of the low contrast module of the Catphan 600 phantom were used for the evaluation of the software. This module contains two subregions: the supraslice region with three groups of low contrast objects (each consisting of nine circular objects with diameter 2-15 mm and contrast 0.3, 0.5, and 1.0%, respectively) and the subslice region with three groups of four circular objects each (diameter 3-9 mm; contrast 1.0%). The software method offered automated determination of low contrast detectability using a NPWE (nonprewhitening matched filter with an eye filter) model observer for the supraslice region. The model observer correlated templates of the low contrast objects with the acquired images of the Catphan phantom and a discrimination index d' was calculated. This index was transformed into a proportion correct (PC) value. In the two-alternative forced choice (2-AFC) experiments used in this study, a PC ≥ 75% was proposed as a threshold to decide whether objects were visible. As a proof of concept, influence of kVp (between 80 and 135 kV), mAs (25-200 mAs range) and reconstruction filter (four filters, two soft and two sharp) on low contrast detectability was investigated. To validate the outcome of the software in a qualitative way, a human observer study was performed.Results: The expected influence of kV, mAs and reconstruction filter on image quality are consistent with the results of the proposed automated model. Higher values for d' (or PC) are found with increasing mAs or kV values and for the soft reconstruction filters. For the highest contrast group (1%), PC values were fairly above 75% for all object diameters >2 mm, for all conditions. For the 0.5% contrast group, the same behavior was observed for object diameters >3 mm for all conditions. For the 0.3% contrast group, PC values were higher than 75% for object diameters >6 mm except for the series acquired at the lowest dose (25 mAs), which gave lower PC values. In the human observer study similar trends were found.Conclusions: We have developed an automated method to objectively investigate image quality using the NPWE model in combination with images of the Catphan phantom low contrast module. As a first step, low contrast detectability as a function of both acquisition and reconstruction parameter settings was successfully investigated with the software. In future work, this method could play a role in image reconstruction algorithms evaluation, dose reduction strategies or novel CT technologies, and other model observers may be implemented as well.     

Pointing to others: How the target gender influences pointing performance

Pointing is a communicative gesture that allows individuals to share information about surrounding objects with other humans. Patients with heterotopagnosia are specifically impaired in pointing to other humans’ body parts but not in pointing to themselves or to objects. Here, we describe a female patient with heterotopagnosia who was more accurate in pointing to men’s body parts than to women’s body parts. We replicated this gender effect in healthy participants with faster reaction times for pointing to men’s body parts than to women’s body parts. We discuss the role of gender stereotypes in explaining why it is more difficult to point to women than to men.     

Control over response priming in visuomotor processing: a lateralized event-related potential study

In a typical Simon task responses are faster when the task-irrelevant stimulus location corresponds to the response location than when it does not. In the case of noncorrespondence it is assumed that externally triggered and internally selected responses are in conflict. Crucially, such conflict appears to be subject to contextual modulations as induced by the immediately preceding event, i.e., the Simon effect was found to be absent when a conflict trial preceded the current event (Stürmer et al. 2002, JEP:HPP). Here, we examined two possible accounts of this context effect in terms of early suppression of externally triggered S-R coding at a premotoric level versus late suppression at a motoric level. Lateralized event-related brain potentials (L-ERPs) were recorded in a Simon task and analyzed as a function of the correspondence sequence. L-ERP activity started earliest over occipito-parietal brain areas and revealed location-based S-R priming irrespective of the prior correspondence context. By contrast, when a noncorresponding trial preceded, such location-based priming was absent in L-ERP activity over the motor cortex (MC). Thus, in support of the late suppression view L-ERPs suggest a clear dissociation in function between externally triggered visuomotor functions within the dorsal stream and the MC reflecting context-controlled response activation.