Ifat Levy

Center-periphery organization of human object areas.

The organizing principles that govern the layout of human object-related areas are largely unknown. Here we propose a new organizing principle in which object representations are arranged according to a central versus peripheral visual field bias. The proposal is based on the finding that building-related regions overlap periphery-biased visual field representations, whereas face-related regions are associated with center-biased representations. Furthermore, the eccentricity maps encompass essentially the entire extent of object-related occipito-temporal cortex, indicating that most object representations are organized with respect to retinal eccentricity. A control experiment ruled out the possibility that the results are due exclusively to unequal feature distribution in these images. We hypothesize that brain regions representing object categories that rely on detailed central scrutiny (such as faces) are more strongly associated with processing of central information, compared to representations of objects that may be recognized by more peripheral information (such as buildings or scenes).

Eccentricity Bias as an Organizing Principle for Human High-Order Object Areas

We have recently proposed a center-periphery organization based on resolution needs, in which objects engaging in recognition processes requiring central-vision (e.g., face-related) are associated with center-biased representations, while objects requiring large-scale feature integration (e.g., buildings) are associated with periphery-biased representations. Here we tested this hypothesis by comparing the center-periphery organization with activations to five object categories: faces, buildings, tools, letter strings, and words. We found that faces, letter strings, and words were mapped preferentially within the center-biased representation. Faces showed a hemispheric lateralization opposite to that of letter strings and words. In contrast, buildings were mapped mainly to the periphery-biased representation, while tools activated both central and peripheral representations. The results are compatible with the notion that center-periphery organization allows the optimal allocation of cortical magnification to the specific requirements of various recognition processes.

The topography of high-order human object areas

Cortical topography is one of the most fundamental organizing principles of cortical areas. One such topography - eccentricity mapping - is present even in high-order, ventral stream visual areas. Within these areas, different object categories have specific eccentricity biases. In particular, faces, letters and words appear to be associated with central visual-field bias, whereas buildings are associated with a peripheral one. We propose that resolution needs are an important factor in organizing object representations: objects whose recognition depends on analysis of fine detail will be associated with central-biased representations, whereas objects whose recognition entails large-scale integration will be more peripherally biased.

From Fear to Safety and Back: Reversal of Fear in the Human Brain

Fear learning is a rapid and persistent process that promotes defense against threats and reduces the need to relearn about danger. However, it is also important to flexibly readjust fear behavior when circumstances change. Indeed, a failure to adjust to changing conditions may contribute to anxiety disorders. A central, yet neglected aspect of fear modulation is the ability to flexibly shift fear responses from one stimulus to another if a once-threatening stimulus becomes safe or a once-safe stimulus becomes threatening. In these situations, the inhibition of fear and the development of fear reactions co-occur but are directed at different targets, requiring accurate responding under continuous stress. To date, research on fear modulation has focused mainly on the shift from fear to safety by using paradigms such as extinction, resulting in a reduction of fear. The aim of the present study was to track the dynamic shifts from fear to safety and from safety to fear when these transitions occur simultaneously. We used functional neuroimaging in conjunction with a fear-conditioning reversal paradigm. Our results reveal a unique dissociation within the ventromedial prefrontal cortex between a safe stimulus that previously predicted danger and a “naive” safe stimulus. We show that amygdala and striatal responses tracked the fear-predictive stimuli, flexibly flipping their responses from one predictive stimulus to another. Moreover, prediction errors associated with reversal learning correlated with striatal activation. These results elucidate how fear is readjusted to appropriately track environmental changes, and the brain mechanisms underlying the flexible control of fear.

Large-Scale Mirror-Symmetry Organization of Human Occipito-Temporal Object Areas

We have combined functional maps of retinotopy (eccentricity and meridian mapping), object category, and motion in a group of subjects to explore the large-scale topography of higher-order object areas. Our results reveal seven consistent category-related entities situated in the occipito-temporal cortex adjoining early visual areas. These include two face-related regions, three object-related regions, and two building-related regions. Interestingly, this complex category-related pattern is organized in a large-scale dorso-ventral mirror symmetry of object category. Furthermore, correlating this pattern to the map of visual field eccentricity, we found that the entire network of areas could be related to a single and unified eccentricity map. We hypothesize that this large-scale organization points to a possible development of high-order object areas through extension and specialization of a single proto-representation.

Neural Representation of Subjective Value Under Risk and Ambiguity

Risk and ambiguity are two conditions in which the consequences of possible outcomes are not certain. Under risk, the probabilities of different outcomes can be estimated, whereas under ambiguity, even these probabilities are not known. Although most people exhibit at least some aversion to both risk and ambiguity, the degree of these aversions is largely uncorrelated across subjects, suggesting that risk aversion and ambiguity aversion are distinct phenomena. Previous studies have shown differences in brain activations for risky and ambiguous choices and have identified neural mechanisms that may mediate transitions from conditions of ambiguity to conditions of risk. Unknown, however, is whether the value of risky and ambiguous options is necessarily represented by two distinct systems or whether a common mechanism can be identified. To answer this question, we compared the neural representation of subjective value under risk and ambiguity. fMRI was used to track brain activation while subjects made choices regarding options that varied systematically in the amount of money offered and in either the probability of obtaining that amount or the level of ambiguity around that probability. A common system, consisting of at least the striatum and the medial prefrontal cortex, was found to represent subjective value under both conditions.

Widespread functional connectivity and fMRI fluctuations in human visual cortex in the absence of visual stimulation.

To what extent does the visual systems activity fluctuate when no sensory stimulation is present? Here, we studied this issue by examining spontaneous fluctuations in BOLD signal in the human visual system, while subjects were placed in complete darkness. Our results reveal widespread slow fluctuations during such rest periods. In contrast to stimulus-driven activity, during darkness, functionally distinct object areas were fluctuating in unison. These fMRI fluctuations became rapidly spatially de-correlated (39% drop in correlation level, P < 0.008) during visual stimulation. Functional connectivity analysis revealed that the slow spontaneous fluctuations during rest had consistent and specific neuro-anatomical distribution which argued against purely hemodynamic noise sources. Control experiments ruled out eye closure, low luminance and mental imagery as the underlying sources of the spontaneous fluctuations. These results demonstrate that, when no stimulus is present, sensory systems manifest a robust level of slow organized fluctuation patterns.

Choice from Non-Choice: Predicting Consumer Preferences from Blood Oxygenation Level-Dependent Signals Obtained during Passive Viewing

Decision-making is often viewed as a two-stage process, where subjective values are first assigned to each option and then the option of the highest value is selected. Converging evidence suggests that these subjective values are represented in the striatum and medial prefrontal cortex (MPFC). A separate line of evidence suggests that activation in the same areas represents the values of rewards even when choice is not required, as in classical conditioning tasks. However, it is unclear whether the same neural mechanism is engaged in both cases. To address this question we measured brain activation with functional magnetic resonance imaging while human subjects passively viewed individual consumer goods. We then sampled activation from predefined regions of interest and used it to predict subsequent choices between the same items made outside of the scanner. Our results show that activation in the striatum and MPFC in the absence of choice predicts subsequent choices, suggesting that these brain areas represent value in a similar manner whether or not choice is required.

Specificity of Human Cortical Areas for Reaches and Saccades

Electrophysiological studies in monkeys have identified effector-related regions in the posterior parietal cortex (PPC). The lateral intraparietal area, for example, responds preferentially for saccades, whereas the parietal reach region responds preferentially for arm movements. However, the degree of effector selectivity actually observed is limited; each area contains neurons selective for the nonpreferred effector, and many neurons in both areas respond for both effectors. We used functional magnetic resonance imaging to assess the degree of effector preference at the population level, focusing on topographically organized regions in the human PPC [visual area V7, intraparietal sulcus 1 (IPS1), and IPS2]. An event-related design adapted from monkey experiments was used. In each trial, an effector cue preceded the appearance of a spatial target, after which a Go signal instructed subjects to produce the specified movement with the specified effector. Our results show that the degree of effector specificity is limited in many cortical areas and transitions gradually from saccade to reach preference as one moves through the hierarchy of areas in the occipital, parietal, and frontal cortices. Saccade preference was observed in visual cortex, including early areas and V7. IPS1 exhibited balanced activation to saccades and reaches, whereas IPS2 showed a weak but significant preference for reaches. In frontal cortex, areas near the central sulcus showed a clear and absolute preference for reaches, whereas the frontal eye field showed little or no effector selectivity. Although these results contradict many theoretical conclusions about effector specificity, they are compatible with the complex picture arising from electrophysiological studies and also with previous imaging studies that reported mostly overlapping saccade- and arm-related activation. The results are also compatible with theories of efficient coding in cortex.

Like cognitive function, decision making across the life span shows profound age-related changes

Significance Although largely unstudied, behavioral changes in decision making across the life span have implications for problems associated with poor decision making at different life stages, such as careless driving in adolescents and disadvantageous medical or financial decision making in older adults. We examine age-based differences in individual decision-making characteristics—choice consistency, rationality, and preferences for known and unknown risks—in 12- to 90-y-olds. We found that even the healthiest of elders show profoundly compromised decision making, and that risk attitudes show systematic changes across the life span that have important policy implications. It has long been known that human cognitive function improves through young adulthood and then declines across the later life span. Here we examined how decision-making function changes across the life span by measuring risk and ambiguity attitudes in the gain and loss domains, as well as choice consistency, in an urban cohort ranging in age from 12 to 90 y. We identified several important age-related patterns in decision making under uncertainty: First, we found that healthy elders between the ages of 65 and 90 were strikingly inconsistent in their choices compared with younger subjects. Just as elders show profound declines in cognitive function, they also show profound declines in choice rationality compared with their younger peers. Second, we found that the widely documented phenomenon of ambiguity aversion is specific to the gain domain and does not occur in the loss domain, except for a slight effect in older adults. Finally, extending an earlier report by our group, we found that risk attitudes across the life span show an inverted U-shaped function; both elders and adolescents are more risk-averse than their midlife counterparts. Taken together, these characterizations of decision-making function across the life span in this urban cohort strengthen the conclusions of previous reports suggesting a profound impact of aging on cognitive function in this domain.