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1.  Tool Manipulation Knowledge is Retrieved by way of the Ventral Visual Object Processing Pathway 
Here we find, using functional Magnetic Resonance Imaging (fMRI), that object manipulation knowledge is accessed by way of the ventral object processing pathway. We exploit the fact that parvocellular channels project to the ventral but not the dorsal stream, and show that increased neural responses for tool stimuli are observed in the inferior parietal lobule when those stimuli are visible only to the ventral object processing stream. In a control condition, tool-preferences were observed in a superior and posterior parietal region for stimuli titrated so as to be visible by the dorsal visual pathway. Functional connectivity analyses confirm the dissociation between sub-regions of parietal cortex according to whether their principal afferent input is via the ventral or dorsal visual pathway. These results challenge the ‘Embodied Hypothesis of Tool Recognition’, according to which tool identification critically depends on simulation of object manipulation knowledge. Instead, these data indicate that retrieval of object-associated manipulation knowledge is contingent on accessing the identity of the object, a process that is subserved by the ventral visual pathway.
PMCID: PMC3795789  PMID: 23810714
2.  The Specialization of Function: Cognitive and Neural Perspectives 
Cognitive neuropsychology  2011;28(0):147-155.
A unifying theme that cuts across all research areas and techniques in the cognitive and brain sciences is whether there is specialization of function at levels of processing that are ‘abstracted away’ from sensory inputs and motor outputs. Any theory that articulates claims about specialization of function in the mind/brain confronts the following types of interrelated questions, each of which carries with it certain theoretical commitments. What methods are appropriate for decomposing complex cognitive and neural processes into their constituent parts? How do cognitive processes map onto neural processes, and at what resolution are they related? What types of conclusions can be drawn about the structure of mind from dissociations observed at the neural level, and vice versa? The contributions that form this Special Issue of Cognitive Neuropsychology represent recent reflections on these and other issues from leading researchers in different areas of the cognitive and brain sciences.
PMCID: PMC3964597  PMID: 22185234
3.  A right visual field advantage for visual processing of manipulable objects 
Information about object-associated manipulations is lateralized to left parietal regions, while information about the visual form of tools is represented bilaterally in ventral occipito-temporal cortex. It is unknown how lateralization of motor-relevant information in left hemisphere dorsal regions may affect the visual processing of manipulable objects. We used a lateralized masked priming paradigm to test for a Right Visual Field (RVF) advantage in tool processing. Target stimuli were tools and animals, and briefly presented primes were identical to, or scrambled versions of the targets. In Experiment 1, primes were presented either to the left or right of the centrally presented target, while in Experiment 2 primes were presented in one of 8 locations arranged radially around the target. In both experiments there was a RVF advantage in priming effects for tool but not for animal targets. Control experiments showed that participants were at chance for matching the identity of the lateralized primes in a picture-word matching experiment, and also ruled out a general RVF speed-of-processing advantage for tool images. These results indicate that the overrepresentation of tool knowledge in the left hemisphere affects visual object recognition, and suggest that interaction between the dorsal and ventral streams occurs during object categorization.
PMCID: PMC3508329  PMID: 22864955
4.  What is in a tool concept? Dissociating manipulation knowledge from function knowledge 
Memory & cognition  2012;40(8):1303-1313.
Research on patients with apraxia, a deficit in skilled action, has shown that the ability to use objects may be differentially impaired relative to knowledge about object function. Here we show, using a modified neuropsychological test, that similar dissociations can be observed in response times in healthy adults. Participants were asked to decide which two of three presented objects shared the same manipulation or the same function; triads were presented in picture- and word-format, and responses were made manually (button press) or with a basic level naming response (verbally). For manual responses (Experiment 1) participants were slower to make manipulation judgments for word stimuli compared to picture stimuli, while there was no difference between word and picture stimuli for function judgments. For verbal naming responses (Experiment 2) participants were again slower for manipulation judgments over word stimuli compared to picture stimuli; however, and in contrast to Experiment 1, function judgments over word stimuli were faster than function judgments over picture stimuli. These data support the hypotheses that knowledge of object function and knowledge of object manipulation correspond to dissociable types of object knowledge, and that simulation over motor information is not necessary in order to retrieve knowledge of object function.
PMCID: PMC3767427  PMID: 22898926
5.  Spatial Frequency Tuning Reveals Interactions between the Dorsal and Ventral Visual Systems 
Journal of cognitive neuroscience  2013;25(6):862-871.
It is widely argued that the ability to recognize and identify manipulable objects depends on the retrieval and simulation of action-based information associated with using those objects. Evidence for that view comes from fMRI studies that have reported differential BOLD contrast in dorsal visual stream regions when participants view manipulable objects compared with a range of baseline categories. An alternative interpretation is that processes internal to the ventral visual pathway are sufficient to support the visual identification of manipulable objects and that the retrieval of object-associated use information is contingent on analysis of the visual input by the ventral stream. Here, we sought to distinguish these two perspectives by exploiting the fact that the dorsal stream is largely driven by magnocellular input, which is biased toward low spatial frequency visual information. Thus, any tool-selective responses in parietal cortex that are driven by high spatial frequencies would be indicative of inputs from the ventral visual pathway. Participants viewed images of tools and animals containing only low, or only high, spatial frequencies during fMRI. We find an internal parcellation of left parietal “tool-preferring” voxels: Inferior aspects of left parietal cortex are driven by high spatial frequency information and have privileged connectivity with ventral stream regions that show similar category preferences, whereas superior regions are driven by low spatial frequency information. Our findings suggest that the automatic activation of complex object-associated manipulation knowledge is contingent on analysis of the visual input by the ventral visual pathway.
PMCID: PMC3767423  PMID: 23410033
6.  Watching the brain in action 
eLife  2013;2:e00866.
Functional magnetic resonance imaging has been used to identify the different networks in the brain that underpin the use of tools by humans.
PMCID: PMC3667576  PMID: 23741621
fMRI; tool use; intentions; action; neuroscience; motor; Human
7.  What drives the organization of object knowledge in the brain? The distributed domain-specific hypothesis 
Trends in cognitive sciences  2011;15(3):97-103.
Various forms of category-specificity have been described at both the cognitive and neural levels, inviting the inference that different semantic domains are processed by distinct, dedicated mechanisms. Here we argue for an extension of a Domain-Specific interpretation to these phenomena that is based on network-level analyses of functional coupling among brain regions. On this view, domain-specificity in one region of the brain emerges because of innate connectivity with a network of regions that also process information about that domain. Recent findings are reviewed that converge with this framework, and a new direction is outlined for understanding the neural principles that shape the organization of conceptual knowledge.
PMCID: PMC3056283  PMID: 21317022
8.  Factors Determining Semantic Facilitation and Interference in the Cyclic Naming Paradigm 
The cyclic naming paradigm, in which participants are slower to name pictures blocked by semantic category than pictures in an unrelated context, offers a window into the dynamics of the mapping between lexical concepts and words. Here we provide evidence for the view that incremental adjustments to the connection weights from semantics to lexical items provides an elegant explanation of a range of observations within the cyclic naming paradigm. Our principal experimental manipulation is to vary the within-category semantic distance among items that must be named together in a block. In the first set of experiments we find that naming latencies are, if anything, faster for within-category semantically close blocks compared to within-category semantically far blocks, for the first presentation of items. This effect can be explained by the fact that there will be more spreading activation, and thus greater priming at the lexical level, for within-category semantically close blocks than within-category semantically far blocks. We test this explanation by inserting intervening filler items (geometric shapes), and show as predicted, that while intervening unrelated trials abolish short-lived semantic priming effects, the long-lag interference effect that is characteristic of this paradigm is unaffected. These data place new constraints on explanations of the cyclic naming effect, and related phenomena, within a model of language production.
PMCID: PMC3283118  PMID: 22363309
lexical access; speech production; semantic interference; semantic facilitation; semantic distance; cyclic naming paradigm
9.  Judging semantic similarity: An event-related fMRI study with auditory word stimuli 
Neuroscience  2010;169(1):279-286.
Much of mental life consists in thinking about object concepts that are not currently within the scope of perception. The general system that enables multiple representations to be maintained and compared is referred to as ‘working memory’ (Repovš and Baddeley, 2006), and involves regions in medial and lateral parietal and frontal cortex (e.g., Smith and Jonides, 1999). It has been assumed that the contents of working memory index information in regions of the brain that are critical for processing and storing object knowledge. To study the processes involved in thinking about common object concepts, we used event related fMRI to study BOLD activity while participants made judgments of conceptual similarity over pairs of sequentially presented auditory words. Through a combination of conventional fMRI analysis approaches and multi-voxel pattern analysis (MVPA), we show that the brain responses associated with the second word in a pair carry information about the conceptual similarity between the two members of the pair. This was the case in frontal and parietal regions involved in the working memory and decision components of the task for both analysis approaches. However, in other regions of the brain, including early visual regions, MVPA permitted classification of semantic distance relationships where conventional averaging approaches failed to show a difference. These findings suggest that diffuse and statistically sub-threshold ‘scattering’ of BOLD activity in some regions may carry substantial information about the contents of mental representations.
PMCID: PMC2908262  PMID: 20412836
10.  Category-specific organization in the human brain does not require visual experience 
Neuron  2009;63(3):397-405.
The ventral stream refers to a neural pathway that projects from early visual areas through to anterior temporal cortex, and comprises regions in ventral and lateral occipital-temporal cortex. The ventral stream is critical for recognizing visually presented objects. Functional imaging studies of the human brain have shown that different regions within the ventral stream show differential activation to nonliving (tools, houses) and living stimuli (animals, faces). The causes of these category preferences are widely debated. Using functional magnetic resonance imaging, we find that the same regions of the ventral stream that show category preferences for nonliving stimuli and animals in sighted adults, show the same category preferences in adults who are blind since birth. Both blind and sighted participants had larger blood oxygen-level dependent (BOLD) responses in the medial fusiform gyrus for nonliving stimuli compared to animal stimuli, and differential BOLD responses in lateral occipital cortex for animal stimuli compared to nonliving stimuli. These findings demonstrate that the medial-to-lateral bias by conceptual domain in the ventral stream does not require visual experience in order to develop, and suggest the operation of innately determined domain-specific constraints on the organization of object knowledge.
PMCID: PMC2743253  PMID: 19679078
11.  The Representation of Tools in Left Parietal Cortex Is Independent of Visual Experience 
Psychological science  2010;21(6):764-771.
Tool use depends on processes represented in distinct regions of left parietal cortex. We studied the role of visual experience in shaping neural specificity for tools in parietal cortex by using functional magnetic resonance imaging with sighted, late-blind, and congenitally blind participants. Using a region-of-interest approach in which tool-specific areas of parietal cortex were identified in sighted participants viewing pictures, we found that specificity in blood-oxygen-level-dependent responses for tools in the left inferior parietal lobule and the left anterior intraparietal sulcus is independent of visual experience. These findings indicate that motor- and somatosensory-based processes are sufficient to drive specificity for representations of tools in regions of parietal cortex. More generally, some aspects of the organization of the dorsal object-processing stream develop independently of the visual information that forms the major sensory input to that pathway in sighted individuals.
PMCID: PMC2908275  PMID: 20483823
tools; blind humans; conceptual knowledge; dorsal visual pathway; fMRI
12.  Action-related properties of objects shape object representations in the ventral stream 
Neuron  2007;55(3):507-520.
The principles driving the organization of the ventral object-processing stream remain unknown. Here we show that stimulus-specific repetition suppression (RS) in one region of the ventral stream is biased according to motor-relevant properties of objects. Quantitative analysis confirmed that this result was not confounded with similarity in visual shape. A similar pattern of biases in RS according to motor-relevant properties of objects was observed in dorsal stream regions in the left hemisphere. These findings suggest that neural specificity for ‘tools’ in the ventral stream is driven by similarity metrics computed over motor-relevant information represented in dorsal structures. Support for this view is provided by converging results from functional connectivity analyses of the fMRI data and a separate neuropsychological study. More generally, these data suggest that a basic organizing principle giving rise to ‘category-specificity’ in the ventral stream may involve similarity metrics computed over information represented elsewhere in the brain.
PMCID: PMC2000824  PMID: 17678861
13.  Gender agreement and multiple referents 
Rivista di linguistica  2008;20(2):285-307.
We report a new pattern of usage in current, spoken Italian that has implications for both psycholinguistic models of language production and linguistic theories of language change. In Italian, gender agreement is mandatory for both singular and plural nouns. However, when two or more nouns of different grammatical gender appear in a conjoined noun phrase (NP), masculine plural agreement is required. In this study, we combined on-line and off-line methodologies in order to assess the mechanisms involved in gender marking in the context of multiple referents. The results of two pronoun production tasks showed that plural feminine agreement was significantly more difficult than plural masculine agreement. In a separate study using offline judgements of acceptability, we found that agreement violations in Italian are tolerated more readily in the case of feminine conjoined noun phrases (e.g., la mela e la banana ‘the:fem apple:fem and the: fem banana: fem’) than masculine conjoined noun phrases (e.g., il fiore e il libro ‘the:mas flower: mas and the:mas book:mas’). Implications of these results are discussed both at the level of functional architecture within the language production system and at the level of changes in language use.*
PMCID: PMC2964885  PMID: 21037930
14.  The word class effect in the picture–word interference paradigm 
The word class effect in the picture–word interference paradigm is a highly influential finding that has provided some of the most compelling support for word class constraints on lexical selection. However, methodological concerns called for a replication of the most convincing of those effects. Experiment 1 was a direct replication of Pechmann and Zerbst (2002; Experiment 4). Participants named pictures of objects in the context of noun and adverb distractors. Naming took place in bare noun and sentence frame contexts. A word class effect emerged in both bare noun and sentence frame naming conditions, suggesting a semantic origin of the effect. In Experiment 2, participants named objects in the context of noun and verb distractors whose word class relationship to the target and imageability were orthogonally manipulated. As before, naming took place in bare noun and sentence frame naming contexts. In both naming contexts, distractor imageability but not word class affected picture naming latencies. These findings confirm the sensitivity of the picture–word interference paradigm to distractor imageability and suggest the paradigm is not sensitive to distractor word class. The results undermine the use of the word class effect in the picture–word interference paradigm as supportive of word class constraints during lexical selection.
PMCID: PMC2908256  PMID: 19998070
Lexical access; Lexical selection; Grammatical class; Imageability; Picture–word interference
15.  Concepts and Categories: A Cognitive Neuropsychological Perspective 
One of the most provocative and exciting issues in cognitive science is how neural specificity for semantic categories of common objects arises in the functional architecture of the brain. More than two decades of research on the neuropsychological phenomenon of category-specific semantic deficits has generated detailed claims about the organization and representation of conceptual knowledge. More recently, researchers have sought to test hypotheses developed on the basis of neuropsychological evidence with functional imaging. From those two fields, the empirical generalization emerges that object domain and sensory modality jointly constrain the organization of knowledge in the brain. At the same time, research within the embodied cognition framework has highlighted the need to articulate how information is communicated between the sensory and motor systems, and processes that represent and generalize abstract information. Those developments point toward a new approach for understanding category specificity in terms of the coordinated influences of diverse regions and cognitive systems.
PMCID: PMC2908258  PMID: 18767921
category-specific semantic deficits; apraxia; semantic organization; domain specific; sensory motor
16.  Semantic Interference in a Delayed Naming Task: Evidence for the Response Exclusion Hypothesis 
In 2 experiments participants named pictures of common objects with superimposed distractor words. In one naming condition, the pictures and words were presented simultaneously on every trial, and participants produced the target response immediately. In the other naming condition, the presentation of the picture preceded the presentation of the distractor by 1,000 ms, and participants delayed production of their naming response until distractor word presentation. Within each naming condition, the distractor words were either semantic category coordinates of the target pictures or unrelated. Orthogonal to this manipulation of semantic relatedness, the frequency of the pictures’ names was manipulated. The authors observed semantic interference effects in both the immediate and delayed naming conditions but a frequency effect only in the immediate naming condition. These data indicate that semantic interference can be observed when target picture naming latencies do not reflect the bottleneck at the level of lexical selection. In the context of other findings from the picture–word interference paradigm, the authors interpret these data as supporting the view that the semantic interference effect arises at a postlexical level of processing.
PMCID: PMC2908266  PMID: 18194068
semantic interference effect; lexical selection by competition; picture-word interference; delayed naming; response exclusion hypothesis
17.  The role of the dorsal visual processing stream in tool identification 
Psychological science  2010;21(6):772-778.
The dorsal visual processing stream subserves object directed action, while the ventral visual processing stream subserves visual object recognition. Little is known about how information computed by dorsal stream structures influences object recognition. We used Continuous Flash Suppression to functionally isolate the information computed by the dorsal stream from that computed by the ventral stream. We show that the information originating from the dorsal stream influences not only decisions requiring superordinate category knowledge, but also decisions that entail the selection of a basic-level object. We further show that the information computed by the dorsal stream does not carry specific functional information about objects. Our results indicate that the dorsal stream, in isolation from the ventral stream, is agnostic as to the identity of the objects that it processes. Instead, we suggest that structures within the dorsal stream compute motor-relevant information (e.g., graspability) that influences the identification of manipulable objects.
PMCID: PMC2908271  PMID: 20483820
18.  The cumulative semantic cost does not reflect lexical selection by competition☆ 
Acta psychologica  2010;134(3):279-289.
The cumulative semantic cost describes a phenomenon in which picture naming latencies increase monotonically with each additional within-category item that is named in a sequence of pictures. Here we test whether the cumulative semantic cost requires the assumption of lexical selection by competition. In Experiment 1 participants named a sequence of pictures, while in Experiment 2 participants named words instead of pictures, preceded by a gender marked determiner. We replicate the basic cumulative semantic cost with pictures (Exp. 1) and show that there is no cumulative semantic cost for word targets (Exp. 2). This pattern was replicated in Experiment 3 in which pictures and words were named along with their gender marked definite determiner, and were intermingled within the same experimental design. In addition, Experiment 3 showed that while picture naming induces a cumulative semantic cost for subsequently named words, word naming does not induce a cumulative semantic cost for subsequently named pictures. These findings suggest that the cumulative semantic cost arises prior to lexical selection and that the effect arises due to incremental changes to the connection weights between semantic and lexical representations.
PMCID: PMC2908277  PMID: 20347062
Cumulative semantic cost; Semantic interference; Lexical access; Picture naming; Semantic access

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