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Biomedical subjects

Semir Zeki

Publications and source records attributed to Semir Zeki.

15 recordsLinked to original sources
PubMed2006

Perceptual compression of space through position integration.

The mechanism of positional localization has recently been debated due to interest in the flash-lag effect, which occurs when a briefly flashed stationary stimulus is perceived to lag behind a spatially aligned moving stimulus. Here we report positional localization observed at motion offsets as well as at onsets. In the 'flash-lead' effect, a moving object is perceived to be behind a spatially concurrent stationary flash before the two disappear. With 'reverse-repmo', subjects mis-localize the final position of a moving bar in the direction opposite to the trajectory of motion. Finally, we demonstrate that simultaneous onset and offset effects lead to a perceived compression of visual space. By characterizing illusory effects observed at motion offsets as well as at onsets, we provide evidence that the perceived position of a moving object is the result of an averaging process over a short time period, weighted towards the most recent positions. Our account explains a variety of motion illusions, including the compression of moving shapes when viewed through apertures.

Cues↗
PubMed2006

Seeing invisible motion: a human FMRI study.

A common view about visual consciousness is that it could arise when and where activity reaches some higher level of processing along the cortical hierarchy. Reports showing that activity in striate cortex can be dissociated from awareness , whereas the latter modulates activity in higher areas , point in this direction. In the specific case of visual motion, a central, "perceptual" role has been assigned to area V5: several human and monkey studies have shown V5 activity to correlate with the motion percept. Here we show that activity in this and other higher cortical areas can be also dissociated from perception and follow the physical stimulus instead. The motion information in a peripheral grating modulated fMRI responses, despite being invisible to human volunteers: under crowding conditions , areas V3A, V5, and parietal cortex still showed increased activity when the grating was moving compared to when it was flickering. We conclude that stimulus-specific activation of higher cortical areas does not necessarily result in awareness of the underlying stimulus.

Adult↗
PubMed2006

Effect of background colors on the tuning of color-selective cells in monkey area V4.

When objects are viewed in different illuminants, their color does not change or changes little in spite of significant changes in the wavelength composition of the light reflected from them. In previous studies, we have addressed the physiology underlying this color constancy by recording from cells in areas V1, V2, and V4 of the anesthetized monkey. Truly color-coded cells, ones that respond to a patch of a given color irrespective of the wavelength composition of the light reflected from it, were only found in area V4. In the present study, we have used a different approach to test the responses of V4 cells in both anesthetized and awake behaving monkeys. Stimuli of different colors, embedded within a Mondrian-type multicolored background, were used to identify the chromatic selectivity of neurons. The illumination of the background was then varied, and the tuning of V4 neurons was tested again for each background illumination. With anesthetized monkeys, the psychophysical effect of changing background illumination was inferred from our own experience, whereas in the awake behaving animal, it was directly reported by the monkey. We found that the majority of V4 neurons shifted their color-tuning profile with each change in the background illumination: each time the color of the background on the computer screen was changed so as to simulate a change in illumination, cells shifted their color-tuning function in the direction of the chromaticity component that had been increased. A similar shift was also observed in colored match-to-sample psychometric functions of both human and monkey. The shift in monkey psychometric functions was quantitatively equivalent to the shift in the responses of the corresponding population of cells. We conclude that neurons in area V4 exhibit the property of color constancy and that their response properties are thus able to reflect color perception.

Action Potentials↗
PubMed2005

The Ferrier Lecture 1995 behind the seen: the functional specialization of the brain in space and time.

The visual brain consists of many different visual areas, which are functionally specialized to process and perceive different attributes of the visual scene. However, the time taken to process different attributes varies; consequently, we see some attributes before others. It follows that there is a perceptual asynchrony and hierarchy in visual perception. Because perceiving an attribute is tantamount to becoming conscious of it, it follows that we become conscious of different attributes at different times. Visual consciousness is therefore distributed in time. Given that we become conscious of different visual attributes because of activity at different, functionally specialized, areas of the visual brain, it follows that visual consciousness is also distributed in space. Therefore, visual consciousness is not a single unified entity, but consists of many microconsciousnesses.

Animals↗
PubMed2005

The chronoarchitecture of the cerebral cortex.

We review here a new approach to mapping the human cerebral cortex into distinct subdivisions. Unlike cytoarchitecture or traditional functional imaging, it does not rely on specific anatomical markers or functional hypotheses. Instead, we propose that the unique activity time course (ATC) of each cortical subdivision, elicited during natural conditions, acts as a temporal fingerprint that can be used to segregate cortical subdivisions, map their spatial extent, and reveal their functional and potentially anatomical connectivity. We argue that since the modular organisation of the brain and its connectivity evolved and developed in natural conditions, these are optimal for revealing its organisation. We review the concepts, methodology and first results of this approach, relying on data obtained with functional magnetic resonance imaging (fMRI) when volunteers viewed traditional stimuli or a James Bond movie. Independent component analysis (ICA) was used to identify voxels belonging to distinct functional subdivisions, based on their differential spatio-temporal fingerprints. Many more regions could be segregated during natural viewing, demonstrating that the complexity of natural stimuli leads to more differential responses in more functional modules. We demonstrate that, in a single experiment, a multitude of distinct regions can be identified across the whole brain, even within the visual cortex, including areas V1, V4 and V5. This differentiation is based entirely on the differential ATCs of different areas during natural viewing. Distinct areas can therefore be identified without any a priori hypothesis about their function or spatial location. The areas we identified corresponded anatomically across subjects, and their ATCs showed highly area-specific inter-subject correlations. Furthermore, natural conditions led to a significant de-correlation of interregional ATCs compared to rest, indicating an increase in regional specificity during natural conditions. In contrast, the correlation between ATCs of distant regions of known substantial anatomical connections increased and reflected their known anatomical connectivity pattern. We demonstrate this using the example of the language network involving Broca's and Wernicke's area and homologous areas in the two hemispheres. In conclusion, this new approach to brain mapping may not only serve to identify novel functional subdivisions, but to reveal their connectivity as well.

Brain Mapping↗
PubMed2005

Brain dynamics during natural viewing conditions--a new guide for mapping connectivity in vivo.

We describe here a new way of obtaining maps of connectivity in the human brain based on interregional correlations of blood oxygen level-dependent (BOLD) signal during natural viewing conditions. We propose that anatomical connections are reflected in BOLD signal correlations during natural brain dynamics. This may provide a powerful approach to chart connectivity, more so than that based on the 'resting state' of the human brain, and it may complement diffusion tensor imaging. Our approach relies on natural brain dynamics and is therefore experimentally unbiased and independent of hypothesis-driven, specialized stimuli. It has the advantage that natural viewing leads to considerably stronger cortical activity than rest, thus facilitating detection of weaker connections. To validate our technique, we used functional magnetic resonance imaging (fMRI) to record BOLD signal while volunteers freely viewed a movie that was interrupted by resting periods. We used independent component analysis (ICA) to segregate cortical areas before characterizing the dynamics of their BOLD signal during free viewing and rest. Natural viewing and rest each revealed highly specific correlation maps, which reflected known anatomical connections. Examples are homologous regions in visual and auditory cortices in the two hemispheres and the language network consisting of Wernicke's area, Broca's area, and a premotor region. Correlations between regions known to be directly connected were always substantially higher than between nonconnected regions. Furthermore, compared to rest, natural viewing specifically increased correlations between anatomically connected regions while it decreased correlations between nonconnected regions. Our findings therefore demonstrate that natural viewing conditions lead to particularly specific interregional correlations and thus provide a powerful environment to reveal anatomical connectivity in vivo.

Acoustic Stimulation↗
PubMed2004

Law and the brain: introduction.

Combining law and the brain as a matter for study requires the integration not just of two apparently remote fields of study but also of two profoundly different orientations towards research and study. We believe that, in spite of the difficulties, such a combination, perhaps even emerging in a new specialized discipline in the future, will not only enrich both fields but is the ineluctable consequence of the current assault on the secrets of the brain. The effort to bring the fields together is therefore a worthy task, and this issue is the first systematic effort to test this expectation.

Behavior↗
PubMed2004

Functional brain mapping during free viewing of natural scenes.

Previous imaging studies have used mostly perceptually abstracted, idealized, or static stimuli to show segregation of function in the cerebral cortex. We wanted to learn whether functional segregation is maintained during more natural, complex, and dynamic conditions when many features have to be processed simultaneously, and identify regions whose activity correlates with the perception of specific features. To achieve this, we used functional magnetic resonance imaging (fMRI) to measure brain activity when human observers viewed freely dynamic natural scenes (a James Bond movie). The intensity with which they perceived different features (color, faces, language, and human bodies) was assessed psychometrically in separate sessions. In all subjects different features were perceived with a high degree of independence over time. We found that the perception of each feature correlated with activity in separate, specialized areas whose activity also varied independently. We conclude that even in natural conditions, when many features have to be processed simultaneously, functional specialization is preserved. Our method thus opens a new way of brain mapping, which allows the localization of a multitude of brain areas based on a single experiment using uncontrolled, natural stimuli. Furthermore, our results show that the intensity of activity in a specialized area is linearly correlated with the intensity of its perceptual experience. This leads us to suggest that each specialized area is directly responsible for the creation of a feature-specific conscious percept (a microconsciousness). Hum. Brain Mapp. 21:75-83, 2004.

Adult↗
PubMed2004

The neurology of ambiguity.

One of the primordial functions of the brain is the acquisition of knowledge. The apparatus that it has evolved to do so is flexible enough to allow it to acquire knowledge about unambiguous conditions on the one hand (colour vision being a good example), and about situations that are capable of two or more interpretations, each one of which has equal validity with the others. However, in the latter instance, we can only be conscious of one interpretation at any given moment. The study of ambiguity thus gives us some insights into how activity at different stations of the brain can result in a micro-consciousness for an attribute, and also tell us something about interactions between different cerebral areas that result in several potential micro-conscious correlates, though only one predominates at any given time. Finally, the study of ambiguity also gives us insights into the neurological machinery that artists have tapped to create the ambiguity that is commonly a hallmark of great works of art.

Art↗
PubMed2004

The neural correlates of maternal and romantic love.

Romantic and maternal love are highly rewarding experiences. Both are linked to the perpetuation of the species and therefore have a closely linked biological function of crucial evolutionary importance. Yet almost nothing is known about their neural correlates in the human. We therefore used fMRI to measure brain activity in mothers while they viewed pictures of their own and of acquainted children, and of their best friend and of acquainted adults as additional controls. The activity specific to maternal attachment was compared to that associated to romantic love described in our earlier study and to the distribution of attachment-mediating neurohormones established by other studies. Both types of attachment activated regions specific to each, as well as overlapping regions in the brain's reward system that coincide with areas rich in oxytocin and vasopressin receptors. Both deactivated a common set of regions associated with negative emotions, social judgment and 'mentalizing', that is, the assessment of other people's intentions and emotions. We conclude that human attachment employs a push-pull mechanism that overcomes social distance by deactivating networks used for critical social assessment and negative emotions, while it bonds individuals through the involvement of the reward circuitry, explaining the power of love to motivate and exhilarate.

Adolescent↗
PubMed2004

The chronoarchitecture of the human brain--natural viewing conditions reveal a time-based anatomy of the brain.

A dominant tendency in cerebral studies has been the attempt to locate architecturally distinct parts of the cortex and assign special functions to each, through histological, clinical or hypothesis-based imaging experiments. Here we show that the cerebral cortex can also be subdivided into different components temporally, without any a priori hypotheses, based on the principle of functional independence. This states that distinct functional subdivisions have activity time courses (ATCs) that are, if not independent, at least characteristic to each when the brain is exposed to natural conditions. To approach a time-based anatomy experimentally, we recorded whole-brain activity using functional magnetic resonance imaging (fMRI) and analyzed the data with independent component analysis (ICA). Our results show that a multitude of cortical areas can be identified based purely on their characteristic ATCs during natural conditions. We demonstrate that a more "rich" stimulation (free viewing of a movie) leads to more areas being activated in a specific way than conventional stimuli, allowing for a more detailed dissection of the cortex into its subdivisions. We show that stimulus-driven functionally specialized areas can be identified by intersubject correlation even if their function is unknown. Chronoarchitectonic mapping thus opens the prospect of identifying previously unknown cortical subdivisions based on natural viewing conditions by exploiting the characteristic temporal "fingerprint" that is unique to each.

Artifacts↗
PubMed2004

Neural correlates of beauty.

We have used the technique of functional MRI to address the question of whether there are brain areas that are specifically engaged when subjects view paintings that they consider to be beautiful, regardless of the category of painting (that is whether it is a portrait, a landscape, a still life, or an abstract composition). Prior to scanning, each subject viewed a large number of paintings and classified them into beautiful, neutral, or ugly. They then viewed the same paintings in the scanner. The results show that the perception of different categories of paintings are associated with distinct and specialized visual areas of the brain, that the orbito-frontal cortex is differentially engaged during the perception of beautiful and ugly stimuli, regardless of the category of painting, and that the perception of stimuli as beautiful or ugly mobilizes the motor cortex differentially.

Adult↗
PubMedDate not supplied

The functional organization of area V2, I: specialization across stripes and layers.

We used qualitative tests to assess the sensitivity of 1043 V2 neurons (predominantly multiunits) in anesthetised macaque monkeys to direction, length, orientation, and color of moving bar stimuli. Spectral sensitivity was additionally tested by noting ON or OFF responses to flashed stimuli of varied size and color. The location of 649 units was identified with respect to cycles of cytochrome oxidase stripes (thick-inter-thin-inter) and cortical layer. We used an initial 8-way stripe classification (4 stripes, and 4 "marginal" zones at interstripes boundaries), and a 9-way layer classification (5 standard layers (2-6), and 4 "marginal" strata at layer boundaries). These classes were collapsed differently for particular analyses of functional distribution; the main stripe-by-layer analysis was performed on 18 compartments (3 stripes x 6 layers). We found direction sensitivity only within thick stripes, orientation sensitivity mainly in thick stripes and interstripes, and spectral sensitivity mainly in thin stripes. Positive length summation was relatively more frequent in thick stripes and interstripes, and negative length/size summation in thin stripes. All these "majority" characteristics of stripes were most prominent in layers 3A and 3B. By contrast, "minority" characteristics (e.g. spectral sensitivity in thick stripes; positive size summation in thin stripes) tended to be most frequent in the outer layers, that is, layers 2 and 6. In consequence, going by the four functions tested, the distinctions between stripes were maximal in layer 3, moderate in layer 2, and minimal in layer 6. Pooling all layers, there was some indication of asymmetry in the stripe cycle, in that thin stripe characteristics (spectral sensitivity, orientation insensitivity, and negative size summation) were also evident in the marginal zone and interstripe immediately lateral to a thin stripe, but less so medially. Within thin stripes, spectral and orientation selectivities were negatively correlated; this was still more accentuated amongst the minority spectrally tuned cells of thick stripes, but absent from interstripes, where these two properties were randomly assorted. Directional and spectral sensitivities were each coupled to negative size summation, but not to each other. We conclude that these functional characteristics of stripes are consistent with segregated, specialized pathways ascending through their middle layers, whilst the outer layers, 1, 2, and 6, utilize feedback from higher areas to adopt a more integrative role.

Animals↗
PubMedDate not supplied

The functional organization of area V2, II: the impact of stripes on visual topography.

We have examined the visuotopic organization of area V2 of macaque monkeys in relation to its modular construction, comprising repetitive cycles of stripes running perpendicular to the border with area V1. Receptive fields were plotted in anesthetised animals, mainly using long penetrations parallel to the V1 border crossing several stripes in dorsal V2 within the representation of paracentral, inferior visual field. We confirm that each set of modules (thick, thin, and interstripes) mounts an unbroken coverage of the visual field, since there is almost invariably some overlap between the aggregate fields recorded in successive stripes of the same class, at intervals of one cycle. Also as expected, penetrations perpendicular to the stripes record changes in eccentricity along an isopolar visual meridian. We measured the size of the point image along such an isopolar meridian in nine cases, and showed that on average it exceeds the length of a typical cycle; again, this implies that no point in space escapes analysis by any of the functional modules. The representation of eccentricity across a cycle of stripes resembles a "ratchet" model, in which the gradient of eccentricity across a single stripe exceeds the gradient across the full cycle, leading to discontinuities ("switchbacks") at the borders between stripes. The shift in eccentricity across the width of a stripe is sufficient to maintain a virtually continuous map across successive stripes of the same class; when coupled to receptive field scatter about the mean trend, this creates the overlap of aggregate fields. The presence of topographic discontinuities, in addition to functional ones, at stripe boundaries, reflects autonomy of function at the local level. At a global level, however, the organization of V2 may promote integration across stripes, since the range of visual overlap, and intrinsic connections, exceeds a single cycle. Furthermore, the reduplication of modules along the isopolar axis has the effect of increasing the isopolar magnification factor (mm/deg); the ratio of isopolar to iso-eccentric magnification is approximately 1.5:1, and this elongation abets the external configuration of V2 that, stretching concentrically around V1, has long been hypothesized to facilitate functional interaction between the two areas.

Animals↗