On the neural correlates of object recognition awareness: relationship to computational activities and activities mediating perceptual awareness

Based on theoretical considerations of Aurell (1979) and Block (1995), we argue that object recognition awareness is distinct from purely sensory awareness and that the former is mediated by neuronal activities in areas that are separate and distinct from cortical sensory areas. We propose that two of the principal functions of neuronal activities in sensory cortex, which are to provide sensory awareness and to effect the computations that are necessary for object recognition, are dissociated. We provide examples of how this dissociation might be achieved and argue that the components of the neuronal activities which carry the computations do not directly enter the awareness of the subject. The results of these computations are sparse representations (i.e., vector or distributed codes) which are activated by the presentation of particular sensory objects and are essentially engrams for the recognition of objects. These final representations occur in the highest order areas of sensory cortex; in the visual analyzer, the areas include the anterior part of the inferior temporal cortex and the perirhinal cortex. We propose, based on lesion and connectional data, that the two areas in which activities provide recognition awareness are the temporopolar cortex and the medial orbitofrontal cortex. Activities in the temporopolar cortex provide the recognition awareness of objects learned in the remote past (consolidated object recognition), and those in the medial orbitofrontal cortex provide the recognition awareness of objects learned in the recent past. The activation of the sparse representation for a particular sensory object in turn activates neurons in one or both of these regions of cortex, and it is the activities of these neurons that provide the awareness of recognition of the object in question. The neural circuitry involved in the activation of these representations is discussed.     

Hemispheric asymmetries in semantic processing: evidence from false memories for ambiguous words

We used fMRI to investigate competition during language production in two word production tasks: object naming and color naming of achromatic line drawings. Generally, fMRI activation was higher for color naming. The line drawings were followed by a word (the distractor word) that referred to either the object, a related object, or an unrelated object. The effect of the distractor word on the BOLD response was qualitatively different for the two tasks. The activation pattern suggests two different kinds of competition during lexical retrieval: (1) Task-relevant responses (e.g., red in color naming) compete with task-irrelevant responses (i.e., the object’s name). This competition effect was dominant in prefrontal cortex. (2) Multiple task-relevant responses (i.e., target word and distractor word) compete for selection. This competition effect was dominant in ventral temporal cortex. This study provides further evidence for the distinct roles of frontal and temporal cortex in language production, while highlighting the effects of competition, albeit from different sources, in both regions.     

Large,colorful, or noisy? Attribute- and modality-specific activations during retrieval of perceptual attribute knowledge

Position emission tomography was used to investigate whether retrieval of perceptual knowledge from long-term memory activates unique cortical regions associated with the modality and/or attribute type retrieved. Knowledge about the typical color, size, and sound of common objects and animals was probed, in response to written words naming the objects. Relative to a nonsemantic control task, all the attribute judgments activated similar left temporal and frontal regions. Visual (color, size) knowledge selectively activated the right posterior inferior temporal (PIT) cortex, whereas sound judgments elicited selective activation in the left posterior superior temporal gyrus and the adjacent parietal cortex. All of the attribute judgments activated a left PIT region, but color retrieval generated more activation in this area. Size judgments activated the right medial parietal cortex. These results indicate that the retrieval of perceptual semantic information activates not only a general semantic network, but also cortical areas specialized for the modality and attribute type of the knowledge retrieved.     

Frontotemporal neural systems supporting semantic processing in Alzheimer’s disease

We hypothesized that semantic memory for object concepts involves both representations of visual feature knowledge in modality-specific association cortex and heteromodal regions that are important for integrating and organizing this semantic knowledge so that it can be used in a flexible, contextually appropriate manner. We examined this hypothesis in an fMRI study of mild Alzheimer’s disease (AD). Participants were presented with pairs of printed words and asked whether the words matched on a given visual–perceptual feature (e.g., guitar, violin: SHAPE). The stimuli probed natural kinds and manufactured objects, and the judgments involved shape or color. We found activation of bilateral ventral temporal cortex and left dorsolateral prefrontal cortex during semantic judgments, with AD patients showing less activation of these regions than healthy seniors. Moreover, AD patients showed less ventral temporal activation than did healthy seniors for manufactured objects, but not for natural kinds. We also used diffusion-weighted MRI of white matter to examine fractional anisotropy (FA). Patients with AD showed significantly reduced FA in the superior longitudinal fasciculus and inferior frontal-occipital fasciculus, which carry projections linking temporal and frontal regions of this semantic network. Our results are consistent with the hypothesis that semantic memory is supported in part by a large-scale neural network involving modality-specific association cortex, heteromodal association cortex, and projections between these regions. The semantic deficit in AD thus arises from gray matter disease that affects the representation of feature knowledge and processing its content, as well as white matter disease that interrupts the integrated functioning of this large-scale network.     

Orientation congruency effects for familiar objects: coordinate transformations in object recognition

How do observers recognize objects after spatial transformations? Recent neurocomputational models have proposed that object recognition is based on coordinate transformations that align memory and stimulus representations. If the recognition of a misoriented object is achieved by adjusting a coordinate system (or reference frame), then recognition should be facilitated when the object is preceded by a different object in the same orientation. In the two experiments reported here, two objects were presented in brief masked displays that were in close temporal contiguity; the objects were in either congruent or incongruent picture-plane orientations. Results showed that naming accuracy was higher for congruent than for incongruent orientations. The congruency effect was independent of superordinate category membership (Experiment 1) and was found for objects with different main axes of elongation (Experiment 2). The results indicate congruency effects for common familiar objects even when they have dissimilar shapes. These findings are compatible with models in which object recognition is achieved by an adjustment of a perceptual coordinate system.     

Neuroanatomical correlates of implicit and explicit memory for structurally possible and impossible visual objects

Implicit memory refers to nonconscious retrieval of past experience demonstrated by facilitation in test performance on tasks that do not require intentional recollection of previous experiences. Explicit memory, in contrast, refers to the conscious retrieval of prior information, as demonstrated during standard recall and recognition tasks. In this experiment, positron emission tomographic (PET) measurements of regional cerebral blood flow (CBF), a marker of local neuronal activity, were used to identify and contrast brain regions that participate in the perception, implicit memory, and explicit memory for structurally possible and impossible visual objects. Ten CBF images were acquired in 16 normal women as they made possible/impossible and old/new recognition decisions about previously studied (old) and nonstudied (new) structurally possible and impossible objects. As reported previously, object decisions for familiar possible objects were associated with increased CBF in the vicinity of the left inferior temporal and fusiform gyri and recognition memory for familiar possible objects was associated with increased CBF in the vicinity of the right hippocampus. In this report, we provide more extensive analyses of the roles of the inferior temporal cortex, the hippocampus, the parahippocampus, and the pulvinar in encoding and retrieval operations. Additionally, patterns of CBF increases and decreases provide information regarding the neural structures involved in implicit and explicit memory.     

Multichannel fNIRS assessment of overt and covert confrontation naming

Confrontation naming tasks assess cognitive processes involved in the main stage of word production. However, in fMRI, the occurrence of movement artifacts necessitates the use of covert paradigms, which has limited clinical applications. Thus, we explored the feasibility of adopting multichannel functional near-infrared spectroscopy (fNIRS) to assess language function during covert and overt naming tasks. Thirty right-handed, healthy adult volunteers underwent both naming tasks and cortical hemodynamics measurement using fNIRS. The overt naming task recruited the classical left-hemisphere language areas (left inferior frontal, superior and middle temporal, precentral, and postcentral gyri) exemplified by an increase in the oxy-Hb signal. Activations were bilateral in the middle and superior temporal gyri. However, the covert naming task recruited activation only in the left-middle temporal gyrus. The activation patterns reflected a major part of the functional network for overt word production, suggesting the clinical importance of fNIRS in the diagnosis of aphasic patients.     

Perceptual priming is not a necessary consequence of semantic classification of pictures

Four experiments investigated how repetition priming of object recognition is affected by the task performed in the prime and test phases. In Experiment 1 object recognition was tested using both vocal naming and two different semantic decision tasks (whether or not objects were manufactured, and whether or not they would be found inside the house). Some aspects of the data were inconsistent with contemporary models of object recognition. Specifically, object priming was eliminated with some combinations of prime and test tasks, and there was no evidence of perceptual (as opposed to conceptual or response) priming in either semantic classification task, even though perceptual identification of the objects is required for at least one of these tasks. Experiment 2 showed that even when perceptual demands were increased by brief presentation, the inside task showed no perceptual priming. Experiment 3 showed that the inside task did not appear to be based on conceptual priming either, as it was not primed significantly when the prime decisions were made to object labels. Experiment 4 showed that visual sensitivity could be restored to the inside task following practice on the task, supporting the suggestion that a critical factor is whether the semantic category is preformed or must be computed. The results show that the visual representational processes revealed by object priming depend crucially on the task chosen.     

Perceptual differentiation as a source of category effects in object processing: Evidence from naming and object decision

The locus of category effects in picture recognition and naming was examined in two experiments with normal subjects. Subjects carried out object decision (deciding whether the stimulus is a “real” object or not) and naming tasks with pictures of clothing, furniture, fruit, and vegetables. These categories are distinguished by containing either relatively many exemplars with similar perceptual structures (fruit and vegetables;structurally similar categories), or relatively few exemplars with similar perceptual structures (clothing and furniture;structurally dissimilar categories). In Experiment 1, responses to the stimuli from the structurally similar categories were slower than responses to stimuli from the structurally dissimilar categories, and this effect was larger in the naming than in the object decision task. Further, prior object decisions to stimuli from structurally similar categories facilitated their subsequent naming. In Experiment 2, we orthogonally manipulated object decision and naming as prime and target tasks, again with stimuli from the four categories. Category effects, with responses slower to objects from structurally similar categories, were again larger in naming than in object decision, and these category effects in naming were reduced by priming with both naming and object decision. We interpret the data to indicate that category effects in object naming can reflect visually based competition which is reduced by the preactivation of stored structural knowledge for objects.     

Domain-dependent activation during spatial and nonspatial auditory working memory

Visual system has been proposed to be divided into two, the ventral and dorsal, processing streams. The ventral pathway is thought to be involved in object identification whereas the dorsal pathway processes information regarding the spatial locations of objects and the spatial relationships among objects. Several studies on working memory (WM) processing have further suggested that there is a dissociable domain-dependent functional organization within the prefrontal cortex for processing of spatial and nonspatial visual information. Also the auditory system is proposed to be organized into two domain-specific processing streams, similar to that seen in the visual system. Recent studies on auditory WM have further suggested that maintenance of nonspatial and spatial auditory information activates a distributed neural network including temporal, parietal, and frontal regions but the magnitude of activation within these activated areas shows a different functional topography depending on the type of information being maintained. The dorsal prefrontal cortex, specifically an area of the superior frontal sulcus (SFS), has been shown to exhibit greater activity for spatial than for nonspatial auditory tasks. Conversely, ventral frontal regions have been shown to be more recruited by nonspatial than by spatial auditory tasks. It has also been shown that the magnitude of this dissociation is dependent on the cognitive operations required during WM processing. Moreover, there is evidence that within the nonspatial domain in the ventral prefrontal cortex, there is an across-modality dissociation during maintenance of visual and auditory information. Taken together, human neuroimaging results on both visual and auditory sensory systems support the idea that the prefrontal cortex is organized according to the type of information being maintained in WM.     

Cortical dynamics of contextually cued attentive visual learning and search: spatial and object evidence accumulation

How do humans use target-predictive contextual information to facilitate visual search? How are consistently paired scenic objects and positions learned and used to more efficiently guide search in familiar scenes? For example, humans can learn that a certain combination of objects may define a context for a kitchen and trigger a more efficient search for a typical object, such as a sink, in that context. The ARTSCENE Search model is developed to illustrate the neural mechanisms of such memory-based context learning and guidance and to explain challenging behavioral data on positive-negative, spatial-object, and local-distant cueing effects during visual search, as well as related neuroanatomical, neurophysiological, and neuroimaging data. The model proposes how global scene layout at a first glance rapidly forms a hypothesis about the target location. This hypothesis is then incrementally refined as a scene is scanned with saccadic eye movements. The model simulates the interactive dynamics of object and spatial contextual cueing and attention in the cortical What and Where streams starting from early visual areas through medial temporal lobe to prefrontal cortex. After learning, model dorsolateral prefrontal cortex (area 46) primes possible target locations in posterior parietal cortex based on goal-modulated percepts of spatial scene gist that are represented in parahippocampal cortex. Model ventral prefrontal cortex (area 47/12) primes possible target identities in inferior temporal cortex based on the history of viewed objects represented in perirhinal cortex.     

Neural correlates of viewing paintings: Evidence from a quantitative meta-analysis of functional magnetic resonance imaging data

Many studies involving functional magnetic resonance imaging (fMRI) have exposed participants to paintings under varying task demands. To isolate neural systems that are activated reliably across fMRI studies in response to viewing paintings regardless of variation in task demands, a quantitative meta-analysis of fifteen experiments using the activation likelihood estimation (ALE) method was conducted. As predicted, viewing paintings was correlated with activation in a distributed system including the occipital lobes, temporal lobe structures in the ventral stream involved in object (fusiform gyrus) and scene (parahippocampal gyrus) perception, and the anterior insula—a key structure in experience of emotion. In addition, we also observed activation in the posterior cingulate cortex bilaterally—part of the brain’s default network. These results suggest that viewing paintings engages not only systems involved in visual representation and object recognition, but also structures underlying emotions and internalized cognitions.     

FMRI of two measures of phonological processing in visual word recognition: ecological validity matters

Previous functional magnetic resonance imaging (fMRI) studies have investigated the role of phonological processing by utilizing nonword rhyming decision tasks (e.g., Pugh et al., 1996). Although such tasks clearly engage phonological components of visual word recognition, it is clear that decision tasks are more cognitively involved than the simple overt naming tasks, which more closely map onto normal reading behavior. Our research aim for this study was to examine the advantages of overt naming tasks for fMRI studies of word recognition processes. Process models are presented to highlight the similarities and differences between two cognitive tasks that are used in the word recognition literature, pseudohomophone naming (e.g., pronounce BRANE) and rhyming decision (e.g., do LEAT and JEAT rhyme?). An fMRI study identified several differences in cortical activation associated with the differences observed in the process models. Specifically, the results show that the overt naming task involved the insular cortex and inferior frontal gyrus, whereas the rhyming decision task engaged the temporal-parietal regions. It is argued that future fMRI research examining the neuroanatomical components of basic visual word recognition utilize overt naming tasks.     

The role of color information on object recognition: a review and meta-analysis

In this study, we systematically review the scientific literature on the effect of color on object recognition. Thirty-five independent experiments, comprising 1535 participants, were included in a meta-analysis. We found a moderate effect of color on object recognition (d = 0.28). Specific effects of moderator variables were analyzed and we found that color diagnosticity is the factor with the greatest moderator effect on the influence of color in object recognition; studies using color diagnostic objects showed a significant color effect (d = 0.43), whereas a marginal color effect was found in studies that used non-color diagnostic objects (d = 0.18). The present study did not permit the drawing of specific conclusions about the moderator effect of the object recognition task; while the meta-analytic review showed that color information improves object recognition mainly in studies using naming tasks (d = 0.36), the literature review revealed a large body of evidence showing positive effects of color information on object recognition in studies using a large variety of visual recognition tasks. We also found that color is important for the ability to recognize artifacts and natural objects, to recognize objects presented as types (line-drawings) or as tokens (photographs), and to recognize objects that are presented without surface details, such as texture or shadow. Taken together, the results of the meta-analysis strongly support the contention that color plays a role in object recognition. This suggests that the role of color should be taken into account in models of visual object recognition.     

Is naming faces different from naming objects? Semantic interference in a face- and object-naming task

A current debate regarding face and object naming concerns whether they are equally vulnerable to semantic interference. Although some studies have shown similar patterns of interference, others have revealed different effects for faces and objects. In Experiment 1, we compared face naming to object naming when exemplars were presented in a semantically homogeneous context (grouped by their category) or in a semantically heterogeneous context (mixed) across four cycles. The data revealed significant slowing for both face and object naming in the homogeneous context. This semantic interference was explained as being due to lexical competition from the conceptual activation of category members. When focusing on the first cycle, a facilitation effect for objects but not for faces appeared. This result permits us to explain the previously observed discrepancies between face and object naming. Experiment 2 was identical to Experiment 1, with the exception that half of the stimuli were presented as face/object names for reading. Semantic interference was present for both face and object naming, suggesting that faces and objects behave similarly during naming. Interestingly, during reading, semantic interference was observed for face names but not for object names. This pattern is consistent with previous assumptions proposing the activation of a person identity during face name reading.     

A selective deficit for living things after temporal lobectomy for relief of epileptic seizures.

Unilateral left and right temporal lobectomy patients and normal control subjects were tested on confrontation naming, speeded naming, category generation, and category and associate matching tasks. Both groups of patients were disproportionately impaired for living relative to nonliving things in confrontation naming, speeded naming, and category generation. We argue that damage to the temporal lobe impairs lexical retrieval most strongly for living things and that the anterior temporal cortices are convergence zones particularly necessary for retrieving the names of living things.     

Neural dynamics of object-based multifocal visual spatial attention and priming: object cueing, useful-field-of-view, and crowding

How are spatial and object attention coordinated to achieve rapid object learning and recognition during eye movement search? How do prefrontal priming and parietal spatial mechanisms interact to determine the reaction time costs of intra-object attention shifts, inter-object attention shifts, and shifts between visible objects and covertly cued locations? What factors underlie individual differences in the timing and frequency of such attentional shifts? How do transient and sustained spatial attentional mechanisms work and interact? How can volition, mediated via the basal ganglia, influence the span of spatial attention? A neural model is developed of how spatial attention in the where cortical stream coordinates view-invariant object category learning in the what cortical stream under free viewing conditions. The model simulates psychological data about the dynamics of covert attention priming and switching requiring multifocal attention without eye movements. The model predicts how “attentional shrouds” are formed when surface representations in cortical area V4 resonate with spatial attention in posterior parietal cortex (PPC) and prefrontal cortex (PFC), while shrouds compete among themselves for dominance. Winning shrouds support invariant object category learning, and active surface-shroud resonances support conscious surface perception and recognition. Attentive competition between multiple objects and cues simulates reaction-time data from the two-object cueing paradigm. The relative strength of sustained surface-driven and fast-transient motion-driven spatial attention controls individual differences in reaction time for invalid cues. Competition between surface-driven attentional shrouds controls individual differences in detection rate of peripheral targets in useful-field-of-view tasks. The model proposes how the strength of competition can be mediated, though learning or momentary changes in volition, by the basal ganglia. A new explanation of crowding shows how the cortical magnification factor, among other variables, can cause multiple object surfaces to share a single surface-shroud resonance, thereby preventing recognition of the individual objects.     

Object recognition in Williams syndrome: uneven ventral stream activation

Williams syndrome (WS) is a genetic disorder associated with severe visuospatial deficits, relatively strong language skills, heightened social interest, and increased attention to faces. On the basis of the visuospatial deficits, this disorder has been characterized primarily as a deficit of the dorsal stream, the occipitoparietal brain regions that subserve visuospatial processing. However, some evidence indicates that this disorder may also affect the development of the ventral stream, the occipitotemporal cortical regions that subserve face and object recognition. The present studies examined ventral stream function in WS, with the hypothesis that faces would produce a relatively more mature pattern of ventral occipitotemporal activation, relative to other objects that are also represented across these visual areas. Using functional magnetic imaging, we compared activation patterns during viewing of human faces, cat faces, houses and shoes in individuals with WS (age 14-27), typically developing 6-9-year-olds (matched approximately on mental age), and typically developing 14-26-year-olds (matched on chronological age). Typically developing individuals exhibited changes in the pattern of activation over age, consistent with previous reports. The ventral stream topography of individuals with WS differed from both control groups, however, reflecting the same level of activation to face stimuli as chronological age matches, but less activation to house stimuli than either mental age or chronological age matches. We discuss the possible causes of this unusual topography and implications for understanding the behavioral profile of people with WS.     

Colour specificity in episodic and in perceptual object recognition with enhanced colour impact

In three experiments we investigated the perceptual specificity of explicit (old-new object recognition) and implicit memory (word-picture matching) for colour. In order to enhance the impact of colour on processing, we increased the number of colours per object and we impaired shape information. We presented multicoloured pictures (Experiment 1), blurred and partially occluded pictures (Experiment 2) and coloured line drawings in visual noise (Experiment 3). Experiments 1 and 2 had an intentional study phase; the study phase of Experiment 3 was an incidental colour or category naming task. Changing colour from study to test always had negative effects on episodic recognition although colour was irrelevant. In contrast, in the matching task old pictures were generally matched faster than new ones independent of their colour congruence. In Experiment 3, an additional small advantage of congruent colours and of semantic processing occurred. We conclude that two different memory representations contribute to these tasks. Changes of an achromatic, more abstract representation, that is used in normal object recognition, and a representation of the specific exemplar that includes colour. This latter token is used in episodic recognition as well as in unusual perceptual tasks (Experiment 3).     

Cortical localisation of the visual and auditory word form areas: a reconsideration of the evidence

In this paper we examine the evidence for human brain areas dedicated to visual or auditory word form processing by comparing cortical activation for auditory word repetition, reading, picture naming, and environmental sound naming. Both reading and auditory word repetition activated left lateralised regions in the frontal operculum (Broca's area), posterior superior temporal gyrus (Wernicke's area), posterior inferior temporal cortex, and a region in the mid superior temporal sulcus relative to baseline conditions that controlled for sensory input and motor output processing. In addition, auditory word repetition increased activation in a lateral region of the left mid superior temporal gyrus but critically, this area is not specific to auditory word processing, it is also activated in response to environmental sounds. There were no reading specific activations, even in the areas previously claimed as visual word form areas: activations were either common to reading and auditory word repetition or common to reading and picture naming. We conclude that there is no current evidence for cortical sites dedicated to visual or auditory word form processing.