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We outline what we believe could be an improvement in future discussions of the brain acting as a Bayesian-Laplacian system. We do so by distinguishing between two broad classes of priors on which the brain's inferential systems operate: in one category are biological priors (β priors) and in the other artefactual ones (α priors). We argue that β priors, of which colour categories and faces are good examples, are inherited or acquired very rapidly after birth, are highly or relatively resistant to change through experience, and are common to all humans. The consequence is that the probability of posteriors generated from β priors having universal assent and agreement is high. By contrast, α priors, of which man-made objects are examples, are acquired post-natally and modified at various stages throughout post-natal life; they are much more accommodating of, and hospitable to, new experiences. Consequently, posteriors generated from them are less likely to find universal assent. Taken together, in addition to the more limited capacity of experiment and experience to alter the β priors compared with α priors, another cardinal distinction between the two is that the probability of posteriors generated from β priors having universal agreement is greater than that for α priors. The two categories are distinct at the extremes; there is, however, a middle range where they merge into one another to varying extents, resulting in posteriors that draw upon both categories., (© 2019 Federation of European Neuroscience Societies and John Wiley & Sons Ltd.)

We studied the neural mechanisms that are engaged during the experience of beauty derived from sorrow and from joy, two experiences that share a common denominator (beauty) but are linked to opposite emotional valences. Twenty subjects viewed and rerated, in a functional magnetic resonance imaging scanner, 120 images which each had classified into the following four categories: beautiful and sad; beautiful and joyful; neutral; ugly. The medial orbito-frontal cortex (mOFC) was active during the experience of both types of beauty. Otherwise, the two experiences engaged different parts of the brain: joyful beauty engaged areas linked to positive emotions while sorrowful beauty engaged areas linked to negative experiences. Separate regions of the cerebellum were engaged during experience of the two conditions. A functional connectivity analysis indicated that the activity within the mOFC was modulated by the supplementary motor area/middle cingulate cortex, known to be engaged during empathetic experiences provoked by other peoples' sadness. Hum Brain Mapp 38:4185-4200, 2017. © 2017 Wiley Periodicals, Inc., (© 2017 The Authors Human Brain Mapping Published by Wiley Periodicals, Inc.)

Whether the visual brain uses a parallel or a serial, hierarchical, strategy to process visual signals, the end result appears to be that different attributes of the visual scene are perceived asynchronously--with colour leading form (orientation) by 40 ms and direction of motion by about 80 ms. Whatever the neural root of this asynchrony, it creates a problem that has not been properly addressed, namely how visual attributes that are perceived asynchronously over brief time windows after stimulus onset are bound together in the longer term to give us a unified experience of the visual world, in which all attributes are apparently seen in perfect registration. In this review, I suggest that there is no central neural clock in the (visual) brain that synchronizes the activity of different processing systems. More likely, activity in each of the parallel processing-perceptual systems of the visual brain is reset independently, making of the brain a massively asynchronous organ, just like the new generation of more efficient computers promise to be. Given the asynchronous operations of the brain, it is likely that the results of activities in the different processing-perceptual systems are not bound by physiological interactions between cells in the specialized visual areas, but post-perceptually, outside the visual brain.

This study was undertaken to learn whether the principle of functional specialization that is evident at the level of the prestriate visual cortex extends to areas that are involved in grouping visual stimuli according to attribute, and specifically according to colour and motion. Subjects viewed, in an fMRI scanner, visual stimuli composed of moving dots, which could be either coloured or achromatic; in some stimuli the moving coloured dots were randomly distributed or moved in random directions; in others, some of the moving dots were grouped together according to colour or to direction of motion, with the number of groupings varying from 1 to 3. Increased activation was observed in area V4 in response to colour grouping and in V5 in response to motion grouping while both groupings led to activity in separate though contiguous compartments within the intraparietal cortex. The activity in all the above areas was parametrically related to the number of groupings, as was the prominent activity in Crus I of the cerebellum where the activity resulting from the two types of grouping overlapped. This suggests (a) that, the specialized visual areas of the prestriate cortex have functions beyond the processing of visual signals according to attribute, namely that of grouping signals according to colour (V4) or motion (V5); (b) that the functional separation evident in visual cortical areas devoted to motion and colour, respectively, is maintained at the level of parietal cortex, at least as far as grouping according to attribute is concerned; and (c) that, by contrast, this grouping-related functional segregation is not maintained at the level of the cerebellum., (Copyright © 2013 Elsevier Inc. All rights reserved.)

We here address the question of whether there is any correlation between subjective preference for simple configurations within a specific visual domain such as motion and strength of activity in visual areas in which that domain is emphasized. We prepared several distinctive patterns of dots in motion with various characteristics and asked humans to rate them according to their preference, before and while scanning the activity in their brains with functional magnetic resonance imaging. For simplicity, we restricted ourselves to motion in the fronto-parallel plane. Moving patterns produced activity in areas V1, V2, the V3 complex (V3, V3A, V3B) and V5, but only in areas V5, V3A/B and parietal cortex did the preferred kinetic patterns produce stronger activity when compared with the non-preferred ones. In addition, preferred patterns produced activity within field A1 of medial orbito-frontal cortex (mOFC), which is not otherwise activated by kinetic stimuli. Hence, for these areas, stronger neural activity correlated with subjective preference. We conclude that configurations of kinetic stimuli that are subjectively preferred correlate with stronger activity within early visual areas and within mOFC. This opens up the possibility of more detailed studies to relate subjective preferences to strength of activity in early visual areas and to relate activity in them to areas whose activity correlates with the subjective experience of beauty.

We wanted to learn whether activity in the same area(s) of the brain correlate with the experience of beauty derived from different sources. 21 subjects took part in a brain-scanning experiment using functional magnetic resonance imaging. Prior to the experiment, they viewed pictures of paintings and listened to musical excerpts, both of which they rated on a scale of 1-9, with 9 being the most beautiful. This allowed us to select three sets of stimuli--beautiful, indifferent and ugly--which subjects viewed and heard in the scanner, and rated at the end of each presentation. The results of a conjunction analysis of brain activity showed that, of the several areas that were active with each type of stimulus, only one cortical area, located in the medial orbito-frontal cortex (mOFC), was active during the experience of musical and visual beauty, with the activity produced by the experience of beauty derived from either source overlapping almost completely within it. The strength of activation in this part of the mOFC was proportional to the strength of the declared intensity of the experience of beauty. We conclude that, as far as activity in the brain is concerned, there is a faculty of beauty that is not dependent on the modality through which it is conveyed but which can be activated by at least two sources--musical and visual--and probably by other sources as well. This has led us to formulate a brain-based theory of beauty.

We pursued our functional magnetic resonance imaging (fMRI) studies of the neural correlates of romantic love in 24 subjects, half of whom were female (6 heterosexual and 6 homosexual) and half male (6 heterosexual and 6 homosexual). We compared the pattern of activity produced in their brains when they viewed the faces of their loved partners with that produced when they viewed the faces of friends of the same sex to whom they were romantically indifferent. The pattern of activation and de-activation was very similar in the brains of males and females, and heterosexuals and homosexuals. We could therefore detect no difference in activation patterns between these groups.

Most experiments on the "neural correlates of consciousness" employ stimulus reportability as an operational definition of what is consciously perceived. The interpretation of such experiments therefore depends critically on understanding the neural basis of stimulus reportability. Using a high volume of fMRI data, we investigated the neural correlates of stimulus reportability using a partial report object detection paradigm. Subjects were presented with a random array of circularly arranged disc-stimuli and were cued, after variable delays (following stimulus offset), to report the presence or absence of a disc at the cued location, using variable motor actions. By uncoupling stimulus processing, decision, and motor response, we were able to use signal detection theory to deconstruct the neural basis of stimulus reportability. We show that retinotopically specific responses in the early visual cortex correlate with stimulus processing but not decision or report; a network of parietal/temporal regions correlates with decisions but not stimulus presence, whereas classical motor regions correlate with report. These findings provide a basic framework for understanding the neural basis of stimulus reportability without the theoretical burden of presupposing a relationship between reportability and consciousness.

Several human and monkey studies have demonstrated a close relationship between motion perception and activation of area V5, leading to the general view that activity in this area correlates with the subjective experience of motion. In the present study, we investigate whether the responses of this area are still governed by the motion percept when the latter is in conflict with the reality of the physical visual stimulation. We simultaneously presented two different, specially designed random-dot kinematograms, one to each eye. These stimuli either both had a single direction of motion and worked in synergy, or had opposite motion directions and thus cancelled each other out perceptually. In this way, we were able to pit the visual stimulus (one vs. two stimulating directions) against the reported perception (directional motion vs. motion noise) of human volunteers during fMRI experiments. We found that a strong motion stimulus that is weakly perceived is more effective in activating V5 (as well as V3) than a weaker motion stimulus, which is nevertheless robustly perceived. Thus, contrary to the prevailing view of perception being the correlate of activity in higher visual areas, we show here that activity is instead dominated by the properties of the physical stimulus, raising the question of whether there is a subpopulation of cells in V5 whose activity is critical for generating the motion percept. In addition, our results provide the first robust evidence for the presence of directionally selective neuronal populations in human prestriate cortex.

In an event-related fMRI study, we scanned eighteen normal human subjects while they viewed three categories of pictures (events, objects and persons) which they classified according to desirability (desirable, indifferent or undesirable). Each category produced activity in a distinct part of the visual brain, thus reflecting its functional specialization. We used conjunction analysis to learn whether there is a brain area which is always active when a desirable picture is viewed, regardless of the category to which it belongs. The conjunction analysis of the contrast desirable > undesirable revealed activity in the superior orbito-frontal cortex. This activity bore a positive linear relationship to the declared level of desirability. The conjunction analysis of desirable > indifferent revealed activity in the mid-cingulate cortex and in the anterior cingulate cortex. In the former, activity was greater for desirable and undesirable stimuli than for stimuli classed as indifferent. Other conjunction analyses produced no significant effects. These results show that categorizing any stimulus according to its desirability activates three different brain areas: the superior orbito-frontal, the mid-cingulate, and the anterior cingulate cortices.

In the work reported here, we set out to study the neural systems that detect predictable temporal patterns and departures from them. We used functional magnetic resonance imaging (fMRI) to locate activity in the brains of subjects when they viewed temporally regular and irregular patterns produced by letters, numbers, colors and luminance. Activity induced by irregular sequences was located within dorsolateral prefrontal cortex, including an area that was responsive to irregular patterns regardless of the type of visual stimuli producing them. Conversely, temporally regular arrangements resulted in activity in the right frontal lobe (medial frontal gyrus), in the left orbito-frontal cortex and in the left pallidum. The results show that there is an abstractive system in the brain for detecting temporal irregularity, regardless of the source producing it.

Visual changes in feature movies, like in real-live, can be partitioned into global flow due to self/camera motion, local/differential flow due to object motion, and residuals, for example, due to illumination changes. We correlated these measures with brain responses of human volunteers viewing movies in an fMRI scanner. Early visual areas responded only to residual changes, thus lacking responses to equally large motion-induced changes, consistent with predictive coding. Motion activated V5+ (MT+), V3A, medial posterior parietal cortex (mPPC) and, weakly, lateral occipital cortex (LOC). V5+ responded to local/differential motion and depended on visual contrast, whereas mPPC responded to global flow spanning the whole visual field and was contrast independent. mPPC thus codes for flow compatible with unbiased heading estimation in natural scenes and for the comparison of visual flow with nonretinal, multimodal motion cues in it or downstream. mPPC was functionally connected to anterior portions of V5+, whereas laterally neighboring putative homologue of lateral intraparietal area (LIP) connected with frontal eye fields. Our results demonstrate a progression of selectivity from local and contrast-dependent motion processing in V5+ toward global and contrast-independent motion processing in mPPC. The function, connectivity, and anatomical neighborhood of mPPC imply several parallels to monkey ventral intraparietal area (VIP).

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.

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.

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.

We have recorded from wavelength-selective cells in macaque monkey visual area V2, interposed between areas V1 and V4 of the color-specialized pathway, to learn whether their responses correlate with perceived colors or are determined by the wavelength composition of light reflected from their receptive fields. All the cells we recorded from were unselective for the orientation and direction of motion of the stimulus, and all were histologically identified to be in the thin cytochrome oxidase stripes. Using multi-colored "Mondrian" scenes of the appropriate spatial configuration, areas of different color were placed in the receptive field of each cell and the entire scene illuminated by three projectors, passing long-, middle-, and short-wave light, respectively, in various combinations. Our results show that wavelength-selective cells in V2 respond to an area of any color depending on whether or not it reflects a sufficient amount of light of their preferred wavelength. In addition, the responses of a third of the cells tested were also influenced by the wavelength composition of their immediate surrounds, thus signaling the result of a local spatial comparison with respect to the amount of their preferred wavelength present. The responses of all also depended on the sequence with which their receptive fields were illuminated with light of the three different wavebands: cells were activated when there was an increase (and inhibited when there was a decrease) in the amount of their preferred wavelength with respect to the other two; the temporal route taken was therefore a determining factor, and, depending on it, cells would either respond or not to a particular combination of wavelengths. We conclude that although spatiotemporal wavelength comparisons are taking place in the color-specialized subdivisions of area V2, the determination of complete color-constant behavior at the neuronal level requires further processing, in other areas.

The primate visual brain consists of many separate, functionally specialized processing systems, each consisting of several apparently hierarchical stages or nodes. The evidence reviewed here leads me to speculate (a) that the processing systems are autonomous with respect to one another, (b) that activity at each node reaches a perceptual end point at a different time, resulting in a perceptual asynchrony in vision, and (c) that, consequently, activity at each node generates a microconsciousness. Visual consciousness is therefore distributed in space and time, with the universal organizing principle of abstraction applied separately within each processing system. The consequence of spatially and temporally distributed microconsciousnesses is that their integration is a multistage, nonhierarchical process that may involve a neural "glue."

Visual neglect occurs most frequently and persistently after lesions that include the right supramarginal gyrus (SMG), a part of the inferior parietal lobule. Patients with this syndrome make very few saccades to the left, and show abnormal performance on tasks in which they must covertly shift their attention to the left, suggesting that the right SMG is involved in the generation of saccades and attention shifts. Functional imaging studies of saccades and covert attention shifts in the normal brain, however, have shown weak or absent responses in both SMGs. We used event-related functional MRI to re-examine the responses to saccades and attention shifts within a single experiment, and to assess responses to left- and right-sided stimuli independently. When subjects made saccades to peripheral stimuli, the expected responses were seen in striate and prestriate cortex, the superior parietal lobules, the frontal eye fields, the supplementary motor area and the anterior insulae. In addition there was a response in the right SMG but not in the left SMG, as predicted from the clinical literature. When subjects made a covert visual assessment of the peripheral stimulus without any saccade, greater activity was seen in all of the areas in the frontoparietal network. Each area showed a bias towards contralateral stimuli, with two exceptions: the anterior insulae gave mainly ipsilateral responses, whilst the right SMG gave equal responses to right- and left-sided stimuli. These findings are discussed in the context of current theories pertaining to the clinical syndrome of neglect.

We have used simple psychophysical methods to determine the sites of color-generating mechanisms in the brain. In our first experiment, subjects viewed an abstract multicolored "Mondrian" display through one eye and an isolated patch from the display through the other. With normal binocular/monocular viewing, the patch has a different color when viewed on its own (void mode) or as part of the Mondrian display (natural mode) [Land, E. H. (1974) Proc. R. Inst. G. B. 49, 23-58]. When the two stimuli were viewed dichoptically, with the patch occupying the position that it would occupy in the Mondrian complex under normal viewing, the patch always appeared in its void color. In a second experiment, when subjects viewed multicolored displays through a different narrow-band filter placed over each eye, the information from the two eyes was combined to result in new colors, which were not seen through either of the two eyes alone. Taken together, these results dissect color-generating mechanisms into two stages, located at different sites of the brain: The first occurs before the appearance of binocular neurons in the cortex and compares wavelength information across space, whereas the second occurs after the convergence of the input from the two eyes and synthetically combines the results of the first.

We have used the technique of functional magnetic resonance imaging (fMRI) and a variety of colour paradigms to activate the human brain regions selective for colour. We show here that the region defined previously [Lueck et al. (1989) Nature, 340, 386-389; Zeki et al. (1991) J. Neurosci., 11, 641-649; McKeefry & Zeki (1997) Brain, 120, 2229-2242] as the human colour centre consists of two subdivisions, a posterior one, which we call V4 and an anterior one, which we refer to as V4alpha, the two together being part of the V4-complex. The posterior area is retinotopically organized while the anterior is not. We discuss our new findings in the context of previous studies of the cortical colour processing system in humans and monkeys. Our new insight into the organization of the colour centre in the human brain may also account for the variability in both severity and degree of recovery from lesions producing cerebral colour blindness (achromatopsia).

In this speculative essay, I examine two evolutionary developments underlying the enormous success of the human brain: its capacity to acquire knowledge and its variability across individuals. A feature of an efficient knowledge-acquiring system is, I believe, its capacity to abstract and to formulate ideals. Both attributes carry with them a clash between experience of the particular and what the brain has developed from experience of the many. Both therefore can lead to much disappointment in our daily lives. This disappointment is heightened by the fact that both abstraction and ideals are subject to variability in time within an individual and between individuals. Variability, which is a cherished source for evolutionary selection, can also be an isolating and individualizing feature in society. Thus the very features of the human brain which underlie our enormous evolutionary success can also be a major source of our misery.

We have studied patient PB, who, after an electric shock that led to vascular insufficiency, became virtually blind, although he retained a capacity to see colors consciously. For our psychophysical studies, we used a simplified version of the Land experiments [Land, E. (1974) Proc. R. Inst. G. B. 47, 23-58] to learn whether color constancy mechanisms are intact in him, which amounts to learning whether he can assign a constant color to a surface in spite of changes in the precise wavelength composition of the light reflected from that surface. We supplemented our psychophysical studies with imaging ones, using functional magnetic resonance, to learn something about the location of areas that are active in his brain when he perceives colors. The psychophysical results suggested that color constancy mechanisms are severely defective in PB and that his color vision is wavelength-based. The imaging results showed that, when he viewed and recognized colors, significant increases in activity were restricted mainly to V1-V2. We conclude that a partly defective color system operating on its own in a severely damaged brain is able to mediate a conscious experience of color in the virtually total absence of other visual abilities.

Looking at ambiguous figures results in rivalry with spontaneous alternation between two percepts. Using event-related functional magnetic resonance imaging, we localized transient human brain activity changes during perceptual reversals. Activation occurred in ventral occipital and intraparietal higher-order visual areas, deactivation in primary visual cortex and the pulvinar. Thus, without any physical stimulus changes, salient perceptual flips briefly engage widely separated specialized cortical areas, but are also associated with intermittent activity breakdown in structures putatively maintaining perceptual stability. Together, the dynamics of integrative perceptual experience are reflected in rapid spatially differentiated activity modulation within a cooperative set of neural structures.

We used the technique of functional magnetic resonance imaging to chart the colour pathways in the human brain beyond V4. We asked subjects to view objects that were dressed in natural and unnatural colours as well as their achromatic counterparts and compared the activity produced in the brain by each condition. The results showed that both naturally and unnaturally coloured objects activate a pathway extending from V1 to V4, though not overlapping totally the activity produced by viewing abstract coloured Mondrian scenes. Normally coloured objects activated, in addition, more anterior parts of the fusiform gyrus, the hippocampus and the ventrolateral frontal cortex. Abnormally coloured objects, by contrast, activated the dorsolateral frontal cortex. A study of the cortical covariation produced by these activations revealed that activity in large parts of the occipital lobe covaried with each. These results, considered against the background of previous physiological and clinical studies, allow us to discern three broad cortical stages of colour processing in the human brain. The first is based on V1 and possibly V2 and is concerned mainly with registering the presence and intensity of different wavelengths, and with wavelength differencing. The second stage is based on V4 and is concerned with automatic colour constancy operations, without regard to memory, judgement and learning. The third stage, based on the inferior temporal and frontal cortex, is more concerned with object colours. The results we report, as well as the schema that we suggest, also allow us to reconcile the computational theory of Land, implemented without regard to cognitive factors such as memory and learning, and the cognitive systems of Helmholtz and Hering, which view such factors as critical in the determination of colours.

We used a colour Mondrian--an abstract scene with no recognizable objects--and its achromatic version to image the change in blood oxygenation in the brains of 12 human subjects, with the aim of learning more about the position and variability of the colour centre in the human brain. The results showed a consistent association of colour stimulation with activation of an area that is distinct from the primary visual areas, and lies in the ventral occipitotemporal cortex; we refer to it as human V4. The position of human V4, as defined on functional grounds, varies between individuals in absolute terms but is invariably found on the lateral aspect of the collateral sulcus on the fusiform gyrus. There was no indication of lingual gyral activation. In further studies designed to reveal the topographic map within V4, we stimulated the superior and inferior visual fields separately, using the same stimuli. We found that human V4 contains a representation of both the superior and inferior visual fields. In addition, there appears to be retinotopic organization of V4 with the superior visual field being represented more medially on the fusiform gyrus and the inferior field more laterally, the two areas abutting on one another. We find no evidence that suggests the existence of a separate representation of the inferior hemifield for colour in more dorsolateral regions of the occipital lobe.

Methods for analysing functional imaging data have evolved rapidly over the last ten years. Standardized techniques based on formal mathematical and statistical theory and rigorous empirical validation have been proposed to facilitate comparisons of biological results between laboratories. This paper examines an image analysis technique that appears to identify unexpectedly large numbers of activated brain areas for the data collected. It concludes that the method may not adequately control for false positives, rendering interpretation of the functional anatomy difficult.

We have compared physiological data recorded from three alert macaque monkeys with separate observations of local connectivity, to locate and characterize the functional border between two related but distinct visual areas on the caudal face of the superior parietal gyrus. We refer to these areas as V6 and V6A. The occupy almost the entire extent of the anterior bank of the parieto-occipital sulcus, V6A being the more dorsal. These two areas are strongly interconnected. Anatomically, we have defined the border as the point at which labelled axon terminals first adopt a recognizably 'descending' pattern in their laminar characteristics, after injections of wheatgerm agglutinin-horseradish peroxidase into the dorsal half of the gyrus (in presumptive V6A). A similar principle was used to recognize the same border by the pattern of input from area V5, except that in this case the relevant transition in laminar characteristics is that between an 'intermediate' pattern (in V6) and an 'ascending' pattern (in V6A). V6A was found to be distinct from V6 in a number of its physiological properties. Unlike V6, it contains visually unresponsive cells as well as units with craniotopic receptive fields ('real-position' cells), units tuned to very slow stimulus speeds, units with complex visual selectivities and units with activity related to attention. V6A was also found to have a larger mean receptive field size and scatter than V6. By contrast, response properties related to the basic orientation and direction of moving bar stimuli were indistinguishable between V6 and V6A, as was the influence of gaze direction on cell activity in the two areas. Two-dimensional maps of the recording sites allowed reconstruction of the V6/V6A border. For comparison, the anatomical results were rendered on two-dimensional maps of identical format to those used to summarize the physiological data. After normalizing for relative size, the physiological and connectional estimates of the border between V6 and V6A were found to coincide, at least within the range of individual variation between hemispheres. An architectonic map in the same format was also made from a hemisphere stained for myelin and Nissl substance. Area PO, defined by its general density of myelination was not distinct in this material, but several architectural features were traceable and one of these was also found to approximate the V6/V6A border. The particular criteria that distinguish V6 from V6A differ from a recent description of areas PO and POd in the Cebus monkey; we believe it most likely that PO and POd together may correspond to V6.

We have used the technique of PET to chart the areas of human cerebral cortex specifically responsive to an optical flow stimulus simulating forward motion in depth over a flat horizontal surface. The optical flow display contained about 2000 dots accelerating in radial directions away from the focus of expansion, which subjects fixated at the centre of the display monitor. Dots remained of constant size, but their density decreased from the horizon, lying across the middle of the screen, to the foreground at the lower screen margin; the top half of the display was void. For the control stimulus the dot motions were randomized, removing any sensation of motion in depth and diminishing the impression of a flat terrain. Comparison of the regional cerebral blood flow (rCBF) elicited by the optical flow and control stimuli was thus intended to reveal any area selectively responsive to the radial velocity field that is characteristic of optical flow in its simplest natural form. Six subjects were scanned, and analysed as a group. Four subjects were analysed as individuals, their PET data being co-registered with MRIs of the cerebrum to localize rCBF changes to individual gyri and sulci. There were three main areas of activation associated with optical flow: the dorsal cuneus (area V3) and the latero-posterior precuneus (or superior parietal lobe) in the right hemisphere, and the occipito-temporal ventral surface, in the region of the fusiform gyrus, in both hemispheres. There was no significant activation of V1/V2, nor of V5. These results show that higher stages of motion take place in both the 'dorsal' and 'ventral' visual pathways, as these are commonly conceived, and that both may be fed by area V3. The information potentially derivable from optical flow concerns the direction of heading, and the layout of the visual environment, a form of three-dimensional structure-from-motion. The perceptual division of labour between the various activated areas cannot be directly inferred, though it is a reasonable supposition that the parietal activation reflects the utility of optic flow for guiding self-motion.

There are many instances in which human subjects perceive a component which is not physically present in a visual stimulus. To study the cerebral activity which correlates with the perception of such an illusory component, we chose Enigma, a static figure in which many subjects perceive illusory motion. By using the technique of positron emission tomography (PET) we recorded the relative regional cerebral blood flow (rCBF) in the brain of 13 subjects while they viewed it and reported seeing the illusory motion. We found that, when subjects perceived illusory motion, the increases in rCBF took place in regions of the brain closely related to, and perhaps identical with, area V5, as defined by the site of rCBF change that took place when the same subjects viewed a physically moving stimulus. In addition, there was activity in other cortical areas outside the visual cortex, not present when the subjects had been viewing objective motion. This suggests that the generation of illusory motion depends not only on a highly specific visual area but also on relative contributions from other parts of the brain that are not activated to the same extent when humans perceive objective motion.

In pursuing our work on the organization of human visual cortex, we wanted to specify more accurately the position of the visual motion area (area V5) in relation to the sulcal and gyral pattern of the cerebral cortex. We also wanted to determine the intersubject variation of area V5 in terms of position and extent of blood flow change in it, in response to the same task. We therefore used positron emission tomography (PET) to determine the foci of relative cerebral blood flow increases produced when subjects viewed a moving checkerboard pattern, compared to viewing the same pattern when it was stationary. We coregistered the PET images from each subject with images of the same brain obtained by magnetic resonance imaging, thus relating the position of V5 in all 24 hemispheres examined to the individual gyral configuration of the same brains. This approach also enabled us to examine the extent to which results obtained by pooling the PET data from a small group of individuals (e.g., six), chosen at random, would be representative of a much larger sample in determining the mean location of V5 after transformation into Talairach coordinates. After stereotaxic transformation of each individual brain, we found that the position of area V5 can vary by as much as 27 mm in the left hemisphere and 18 mm in the right for the pixel with the highest significance for blood flow change. There is also an intersubject variability in blood flow change within it in response to the same visual task. V5 nevertheless bears a consistent relationship, within each brain, to the sulcal pattern of the occipital lobe. It is situated ventrolaterally, just posterior to the meeting point of the ascending limb of the inferior temporal sulcus and the lateral occipital sulcus. In position it corresponds almost precisely with Flechsig's Feld 16, one of the areas that he found to be myelinated at birth.

This review is an enquiry into why the early clinical evidence for a colour centre in the cerebral cortex of man was so successfully dismissed for the best part of a century. The imperfection of this evidence cannot be the reason, for the same evidence that was rejected earlier is accepted today. Instead, it was because the prevalent concepts of vision as a function, and of the role of the cerebral cortex in it, dominated facts and prevented acceptance of evidence showing a specialization for colour in the visual cortex. It was only after those concepts were overthrown by the demonstration of functional specialization in the visual cortex of the primate that the evidence for a colour centre in the human brain became acceptable. Today, our new knowledge of the colour areas and pathways in the primate brain allows us to give a more complete account of the pathophysiology of cerebral achromatopsia in man.

Discusses V5, which consists of large myelinated fibers terminating in a zone of the posterior bank of the superior temporal sulcus. Specialization of V5; Location of V5 in the human brain; Relation of the level of activity in V5 with that in the primary visual cortex.