25-Haggard-Chap25 5/5/07 11:34 PM Page 567 25 Using conceptual knowledge in action and language M. van Elk, H. T. van Schie, O. Lindemann and H. Bekkering Action semantics refer to the conceptual knowledge that is necessary to guide our actions and can be defined in terms of action goals and action means. The present chapter outlines the concept of action semantics and presents a review of the existing literature dealing with functional mechanisms underlying conceptual knowledge and goal-directed behavior. Recent empirical findings are presented which indicate that semantic information is selected in a flexible manner in accordance with the action intention of the actor, whereby long-term conceptual associations may be overruled by current behavioral goals. These results extend the selection-for-action principle beyond perception to include the selection of semantic information that is relevant for the upcoming action. Findings are interpreted within the light of contemporary models of conceptual knowledge in action. Introduction Within the booming field of cognitive neuroscience, a growing number of studies underline the interrelatedness of perception, action and language. It is acknowledged that cognitive faculties do not operate in isolation, but rather share common neural mechanisms. This may come as no surprise, regarding the fact that the division of the mind in separate cognitive subsystems was originally inherited from intuitive folk-psychological concepts. The separate consideration of cognitive faculties has proven to be useful in acquiring a basic understanding of the workings of the brain. However, given the close interactions between different cognitive subsystems, it may be useful to think about adjusting our terminology to better cover the phenomena investigated. An exclusive focus on the uniqueness of each cognitive faculty may blind us for the elements that are common to different systems. In the present chapter we introduce the concept of ‘action semantics’ as a useful tool to bring together the closely related fields of language, action and conceptual knowledge. In short, action semantics refer to the conceptual knowledge that is necessary to guide our actions. Both long-term and short-term action semantics are important for a successful interaction with the surrounding world. For example, although most of the time we use everyday objects in an appropriate manner, sometimes objects acquire a new function when used in a different context. Imagine yourself reading a newspaper while being 25-Haggard-Chap25 568 5/5/07 11:34 PM Page 568 ACTION SEMANTICS disturbed by a mosquito. The folded newspaper will be an excellent device to get rid of the insect. Countless examples show that conceptual knowledge is selected and activated in a flexible way, according to the actor’s current action goals (e.g. using a screwdriver to open a paint-bucket). In the first part of this chapter we will discuss the nature of action semantics, the neural mechanisms supporting action semantics and its development over the course of life. It is suggested that action goals play a dominant role in understanding the behavior of others and in the selection of perceptual information for our own actions. In the second part we turn to the question of how conceptual knowledge might be represented within the brain. Different theories of conceptual knowledge in action will be discussed that disagree on the principles underlying the organization of conceptual knowledge. In the third section we will focus on the close relation between language and action. Whereas several studies report effects of language on action preparation, hardly any study investigated effects of action intention on the selection of semantic information. Recent experiments from our group show that semantics are automatically selected according to an actor’s present action goal, even when actions are in conflict with long-term knowledge of object use (Lindemann et al., 2006; M. Van Elk et al., unpublished data). In the final part of this chapter we will discuss theoretical implications of the findings presented for theories about conceptual knowledge in action. We suggest that the study of action semantics brings together several closely related fields and provides insight into the functional mechanisms underlying goal-directed behavior. Action semantics We start this section by introducing the concept of action semantics. In subsequent paragraphs we will then focus on (i) the neural mechanisms that mediate the use of conceptual knowledge in action, (ii) the acquisition of action semantics in young children and (iii) the selection of perceptual and semantic information for action. What are action semantics? Action semantics refer to the conceptual knowledge that is necessary to guide our actions. More specifically, the conceptual knowledge used for actions consists of our knowledge about the function and the use of concrete objects. Whenever we encounter a wellknown object, semantic information about object use will be retrieved automatically from memory. It is exactly this interplay between conceptual knowledge and action that is the topic of our investigation. The relation between action planning and conceptual knowledge has often been overlooked by theories in the domain of motor control. Likewise, theories of conceptual knowledge have paid little attention to the use of semantic knowledge for guiding behavior. For example, theoretical discussions on the modal or amodal nature of conceptual knowledge bypass the idea that our conceptual system in the first place evolved to enable us to cope with environmental demands. Being able to deal effectively with one’s environment is essential to survival and a conceptual system that enables organisms to successfully manipulate the surrounding world surely has great evolutionary advantages. 25-Haggard-Chap25 5/5/07 11:34 PM Page 569 ACTION SEMANTICS Action semantics capitalizes on the idea that conceptual knowledge is closely related to the control of action. Much of the information that is necessary to guide our actions is directly available to our senses. For example, we quickly pick up the natural object affordances that allow us to interact with categories of objects (e.g. seats) as well as specific instances of these categories (e.g. reclining chair, Gibson, 1986). However, not all information that is necessary to act successfully can be obtained directly through our senses. Within the first six months of their lives, children are able to grasp an object like a water faucet correctly, but the knowledge of how to turn a faucet is something that cannot be perceived directly upon mere observation of the object by itself. Most water faucets are round and can be acted upon in many ways in order to achieve the desired outcome. For most people, the knowledge that in most Western countries faucets are opened by means of a counter clockwise rotation will lead to the decision to grasp the tap with the thumb pointing upwards or away from the body allowing for a comfortable turning motion (Rosenbaum et al., 1995). In other words, the stored conceptual knowledge about water faucets is used for guiding our actions. Action semantics can be defined in terms of action goals (what to use an object for) and action means (how to use an object). With action means we typically denote the manner in which an object is handled (e.g. grasped), or more generally, the way in which we preferably interact with an object (e.g. a toothbrush is grasped at the handle, not at the brush). With action goals, on the other hand, we refer to the purpose or intent for which an object is used (e.g. a toothbrush is used for brushing teeth). Different levels of complexity can be discerned when describing action goals, ranging from high-level goalrepresentations to lower levels. For example, lessening one’s thirst may be a general action goal, whereas lifting a cup and bringing it to the mouth represents a lower level of action goals. A useful taxonomy to distinguish between different levels of goal-representations includes: (1) physical goals that refer to concrete objects, (2) action goals that refer to the functional use of objects and (3) mental goals that reflect a desired state of affairs in the world (Bekkering and Wohlschläger, 2000). Both action and mental goals are closely related to the intention or purpose of the actor. Physical goals reflect a lower level description of action goals and can be defined in terms of a movement’s goal-location. Still little is known about the functional processes through which selection and coordination of action goals and action means come about. At the behavioral level, the selection of action means (e.g. the way in which an object is grasped) often seems to fulfill the requirements of the action goal (what one is intending to do with the object; Rosenbaum et al., 1995), suggesting the possibility of a hierarchical organization supporting action semantics for goals and means. In accordance with this suggestion, previous research has implicated the importance of action goals for guiding the integrity and consistency of behavior over prolonged periods of time (Zalla et al., 2003; H. T. Van Schie et al., unpublished data). Neural mechanisms mediating action semantics The division between semantics for action goals and means is supported by the study of neuropsychological patients with semantic dementia, who sometimes display selective loss 569 25-Haggard-Chap25 570 5/5/07 11:34 PM Page 570 ACTION SEMANTICS of a particular category of semantic knowledge. Previous research has distinguished between different types of action deficits in patients with otherwise fully operational motor functions. Patients with ideational apraxia show a general loss of conceptual knowledge about the function of and the purpose for which they are used (Ochipa et al., 1992). These patients are able to perform simple motor-acts and to retrieve tool names, but fail to use objects in an appropriate way (e.g. they can use a toothbrush as a spoon). In contrast, ideomotor apraxia refers to the inability to execute specific motor commands after damage to the left parietal or fronto-parietal cortex (Buxbaum, 2001). Ideomotor apraxic patients show spatio-temporal errors when pantomiming object use or may grasp an object in an inappropriate way to find themselves at a loss as to which movements to make with it (grasp a stylus with a full grip). The behavior displayed by both groups of patients suggests a dissociation between knowledge about the functional use of objects (action goals) and knowledge of the usual interaction (action means) associated with objects (cf. Leiguarda and Marsden, 2000). Ideomotor deficits have been associated with damage to parietal cortex, which is involved in visuo-motor transformations necessary for grasping (e.g. Heilman et al., 1981; Buxbaum et al., 2003). In addition, damage to the left posterior parietal lobe is often associated with a deficit demonstrating the appropriate actions associated with particular objects (Ochipa et al., 1989). Ideational apraxia on the other hand has been found in association with lesions in premotor and prefrontal areas (Sirigu et al., 1998; Leiguarda and Marsden, 2000), consistent with recent evidence on the role of these areas in supporting goal-directed actions with objects after their prehension (H. T. Van Schie and H. Bekkering, unpublished data). Another important region that has been implicated in the retrieval of knowledge about actions is the posterior middle temporal region (MT) that is involved in the perception of motion (Tootell et al., 1995). Damage to the middle temporal region in neuropsychological patients is associated with a deficit in the recognition of tools (Tranel et al., 2003). The same area is also activated in association with verb generation reflecting actions to visually presented nouns (Fiez et al., 1996) or objects (Martin et al., 1995). Furthermore, Kable et al. (2002) found greater activity in bilateral MT to the presentation of action pictures than to pictures of objects, whereas words instead of pictures mainly activated left-sided MT. The reported activation of MT is consistent with the suggestion that accessing action knowledge involves incorporating motion features. In sum, these findings suggest that accessing action knowledge involves activating motor structures and areas supporting motor performance, consistent with the idea that action knowledge is stored or represented within the motor system (reviews in Martin and Chao, 2001; Tranel et al., 2003). Thus, having discussed the neural mechanisms involved in action semantics, we now turn to the question of how we acquire action semantics over the course of our lives. Insight into the acquisition of conceptual knowledge for action is gained from developmental studies, which will be the focus of the next section. Development of action semantics Piaget was among the first to suggest that the central role of action experience in cognitive development should be addressed more broadly (Piaget, 1953). The production of 25-Haggard-Chap25 5/5/07 11:34 PM Page 571 ACTION SEMANTICS goal-directed acts in the first year of life may not only refine the production of goal-directed acts in infants, it might also be crucial to the acquisition of conceptual knowledge about the world in general. Information gleaned from early action experiences may, for instance, facilitate sensitivity to the goal structure of infants’ own and others’ actions. Even at a very young age, children extract the meaningful structure of the surrounding world at multiple levels of analysis. Moreover, developmental theorists have considered this capacity a prerequisite both for having intentions and for understanding the intentions of others (Gergely et al., 1995; Meltzoff and Moore, 1995). Indeed, the ability to construe actions in terms of goals can be observed in preschoolers. Eighteen-month-old infants infer the goal of uncompleted human action sequences and infants as young as 14 months tend to imitate intended but not accidental acts (Meltzoff, 1995; Carpenter et al., 1998). In addition, 14-month-old infants re-enact the final goal of a modeled action, but do not always reproduce the means. Gergely et al. (2002) convincingly showed that children will follow a given course of action to achieve the given goal only if they consider it to be the most rational alternative. Their results indicate that imitation of goal-directed action by preverbal infants is a selective, conceptual knowledge-based process, rather than a simple re-enactment of the means used by a demonstrator, as was previously thought. This action knowledge may subsequently be extended to allow infants to interpret a broad range of events with respect to their meaning and goal structure (Sommerville and Woodward, 2005). For instance, after watching an actor reach for and grasp a toy, infants show a stronger novelty response to a change in the actor’s goal than to a change in the spatial location or trajectory of her reach (Woodward, 1998). Recent findings have stressed a correlation between the emergence of goal-directed behaviors and infants’ ability to detect goals in the acts of others (Sommerville and Woodward, 2005). In addition, a measurable impact of action experience on action interpretation was demonstrated. Developmental studies suggest that the ability to infer the intention of an observed actor is already present in children aged 12 months (e.g. Gergely et al., 1995). In a habituation study, 12-month-old infants were found to look longer at an object-directed action when an indirect reach-movement was made, but if no object was present no such preference was found (Phillips and Wellman, 2005). This suggests that children understand the observed action in terms of the actor’s goal. However, recent findings suggest that children’s understanding of actions is constrained by task context. While children tended automatically to imitate the observed action goal rather than the means by which it is performed (Bekkering and Wohlschläger, 2000), making the means more salient (e.g. putting an arrow on the movement-path) results in movement-related adjustments, although the goal of the action remains the same (Cox et al., unpublished data). Apparently, both goals and means play an important role in the acquisition of conceptual knowledge to interact with the surrounding world. The mechanism underlying the acquisition of action semantics seems to be the ability to imitate, which is probably mediated by the human mirror system. The discovery of mirror neurons in primates that respond selectively to both the execution of an action and the observation of actions being performed by another actor, led to the idea of an observation–execution matching system that is crucial to the understanding of observed 571 25-Haggard-Chap25 572 5/5/07 11:34 PM Page 572 ACTION SEMANTICS actions (Rizzolatti et al., 1996). In humans a comparable mirror system was identified, encompassing such brain areas as the superior temporal sulcus, the inferior parietal lobule and the inferior frontal gyrus (e.g. Grafton et al., 1996; Calvo-Merino et al., 2005). Several possible functions regarding the activity in mirror neurons have been put forward, of which the most important include (1) supporting a goal representation of intended action, (2) providing a mechanism to understand observed behavior and (3) supporting the acquisition of new behavior by imitation (e.g. Iacoboni et al., 1999; Umiltà et al., 2001; Rizzolatti et al., 2001). Mirror-related areas appear crucial to the understanding of observed actions and thereby provide the child with a mechanism to learn by observation. For example, recently it has been reported that the observation of errors performed by another subject led to an error-related brain response that was comparable to self-generated errors (Van Schie et al., 2004). This suggests that similar mechanisms are recruited during monitoring of one’s own or another’s action. Taken together, it seems that action understanding and acquisition of novel actions through observation are goal-directed activities, mediated by systems that support both the generation and observation of actions. The development of action semantics in young children enables them to make sense of observed actions and to perform goal-directed actions themselves. The acquisition and use of goal representations seems a critical aspect for the development of action semantics. In line with this suggestion are findings that underline the importance of action goals in dealing effectively with the enormous amount of information that enters our senses. Action goals in perception and action A guiding principle in perceptual processing is the ability to selectively attend to particular stimuli. According to the framework of selection-for-action (Allport, 1987) relevant information is selected in line with the action intention of the actor. For example, in the domain of visual perception it has repeatedly been found that specific visual information (e.g. orientation of an object) is selectively processed, depending on the action goal of the actor, e.g. grasping or pointing (Bekkering and Neggers, 2002; Hannus et al., 2005). In a recent study we found a selective enhancement of early visual ERP components when subjects intended to grasp as compared to when they intended to point towards visually presented objects (H. T. Van Schie, M. Van Elk and H. Bekkering, unpublished data). Because processing of orientation is relevant for grasping but not for pointing, the orientation of the object needs to be processed more actively in visual areas. Craighero et al. (1999) instructed participants to grasp objects of a particular orientation in response to stimuli showing a final hand position that could be congruent or incongruent with respect to the intended grasp. Response latencies were fastest to pictures with the greatest similarity to the final hand posture of the intended grasping movement, suggesting a close link between action preparation and perceptual processing. Interestingly, enhancement of visual processing has also been reported in relation to language comprehension. Upon reading a description of a nail being pounded either into the wall or into the floor, participants were faster in processing visual stimuli congruent with the implied orientation (Stanfield and Zwaan, 2001). 25-Haggard-Chap25 5/5/07 11:34 PM Page 573 ACTION SEMANTICS These and other studies suggest that in addition to bottom-up information that is elicited by the perception of an object, top-down effects of the action context and intention of an actor may selectively modulate visual processing. A similar principle may also support the selection of object semantics in accordance with the action intention of an individual. One hypothesis that plays a central part in the current chapter is that the selection of conceptual knowledge for action is guided by the same principle that is proposed to underlie the selection of perceptual information for action. Theories of motor control suggest that conceptual knowledge is involved in the selection of appropriate action plans (Rosenbaum et al., 2001). Functional imaging studies confirm that conceptual knowledge about the appropriate manner of grasping an object becomes activated automatically upon object identification (Chao and Martin, 2000; Grèzes and Decety, 2002; Grèzes et al., 2003). More specifically, it has been reported that action preparation and object perception are mutually dependent processes. For example, passive observation of tools has been reported to elicit motor activation in areas that are normally activated with tool use (e.g. Chao and Martin, 2000). In monkeys it has been found that neurons in grasping-related areas, namely the anterior intraparietal sulcus (AIP) and the ventral premotor cortex (F5), selectively respond to the passive observation of graspable objects (Murata et al., 1997). These canonical neurons are selectively responsive to objects of a particular size and shape and probably play an important role in specifying the motor properties that are required for directing an action towards a particular object. At the behavioral level it has been found that viewing an object automatically activates the appropriate hand shape for using it (Klatzky et al., 1989). The same automatic effect of appropriate handgrip aperture has also been reported in response to reading words of graspable objects that afforded either a full or a precision grip (Glover et al., 2004). Reading a word that represented a large object (e.g. ‘apple’) resulted in a larger grip aperture in the early part of the grasping movement as compared to reading a word that referred to a small object (e.g. ‘grape’). In a similar fashion, Tucker and Ellis (2001) presented subjects with pictures for a categorical judgment and found a compatibility effect between response type (making a precision or full grip with the hand) and object size (e.g. faster responding to the presentation of car keys when making a precision grip). Other studies suggest that action knowledge plays an important role in the retrieval of conceptual information. Chainay and Humphreys (2002) reported faster responding when making action decisions about the use of particular objects (e.g. do you perform a twisting or a pouring action with the object?) as compared to making contextual decisions (is the object typically found in the kitchen?). In addition, Yoon and Humphreys (2005) showed that activation of action knowledge plays a constitutive role in making functional judgments about object use. Participants observed objects either grasped with a congruent, an incongruent or without any handgrip and they verified the object name (‘Is this the name of the object?’) or the object action (‘Is the object used for …?’). A congruency effect of the observed action on making action decisions was found, while no such effect was present for naming the object. Together these studies support the view that the action and motor systems are involved in object identification. 573 25-Haggard-Chap25 574 5/5/07 11:34 PM Page 574 ACTION SEMANTICS Thus, perception of an object is found to evoke activation of an appropriate motorprogram that allows interaction with the object (e.g. facilitation of the appropriate handgrip). These findings are consistent with the theoretical notion of Gibson (1986) that whenever an actor perceives an object, affordances will be inferred that inform the actor about the possible actions that can be directed towards the object. However, in addition to automatic (bottom-up) effects resulting from object perception, in many cases the manner of object interaction is determined by the intention of an actor or the action goal. That is, although chairs typically afford sitting down, in the context of a different action goal (e.g. getting something from a top shelf) a different manner of interaction may be selected. The flexible use of action semantics will be investigated in the third part of this chapter. Before doing so, however, we turn to different views on the organization of conceptual knowledge, in order to provide a theoretical background against which empirical findings may be evaluated. Theories of conceptual knowledge In general, theories on conceptual knowledge may be classified into two broad categories. According to amodal theories, concepts are abstract, amodal and arbitrary entities that operate independently from the sensorimotor systems (e.g. Fodor, 1983). Concepts are considered as part of symbolic representational structures that function in an analogous way to language, (e.g. ‘language-of-thought’ hypothesis). An important argument in favor of amodal theories of representation concerns the ease with which different tokens of a particular category (type) are recognized, independent of perceptual differences. However, amodal theories of concepts also face important difficulties such as a lack of empirical validation, giving a specification of the transduction process from perceptual states to symbols and the grounding problem. If all knowledge is only represented as a set of interrelated symbols, the meaning of a particular symbol can only be specified by its relation to, and in terms of, other symbols. In contrast, modal theories of representation argue that concepts are grounded in modality-specific systems (e.g. Barsalou et al., 2003). Neuropsychological patients with damage to a particular sensory area and a specific deficit in categorical knowledge provide a powerful argument for the modal representation of concepts. Embodied cognition refers to different theories of conceptual representation that share the assumption of a multi-modal grounding of concepts across different sensorimotor areas (e.g. Clark, 1997; Gallese and Goldman, 1998; Lakoff and Johnson, 1999). However, embodied theories differ widely on the emphasis that is put on separate cognitive subsystems and on the way in which our conceptual knowledge is organized. It may be useful to think of modal theories on conceptual knowledge as differentiating along two different axes: ◆ the first axis covers whether conceptual knowledge is organized by principles internal to the organism or that it is rather determined by external, stimulus-driven processes; ◆ the second axis represents the theory’s relative focus on perceptual or action-related processes. 25-Haggard-Chap25 5/5/07 11:34 PM Page 575 THEORIES OF CONCEPTUAL KNOWLEDGE Accordingly, theories of conceptual knowledge may be represented in a two-dimensional conceptual space, as represented in Figure 25.1. Of course, the division of theories along two axes provides us with a simplified picture that overlooks finer nuances of different theories. Nevertheless, we suggest that the proposed division provides us with a useful tool to think about different approaches in studying the acquisition and organization of conceptual knowledge. The different axes represent the relative emphasis that is put on top-down or bottom-up processes on the one hand and on action- or perception-related systems on the other hand. In the remainder of this section we will discuss different proposals for the retrieval of conceptual knowledge in relation to the domain of action and focus on the strengths and weaknesses of embodied accounts of conceptual representation. Sensory functional hypothesis versus correlated structure principle The sensory–functional hypothesis, put forward by Warrington and Shallice (1984), argues that conceptual knowledge is stored in modality-specific sensory and motor structures. Hence, identification of instances of different semantic categories depends critically upon activating particular modality-specific semantic subsystems in the brain. Category-specific semantic deficits are the result of damage to modality-specific semantic subsystems. For example, damage to the visual semantic subsystem probably impairs semantic knowledge of categories, for which visual features are a defining characteristic. Borgo and Shallice (2001) report a patient who had suffered from herpetic encephalitis, showing a severe deficit in category-specific knowledge for living things. Interestingly, this patient also was impaired when tested for knowledge of other sensory quality categories Correlated structure principle Perceptual Symbols Theory Perceptual systems Gibson’s Affordances Associative Sequence Learning Sensory-functional hypothesis Ideomotor Theory of Action Action-semantics Motor-systems Two-route model to action Stimulus-driven Driven by organism Figure 25.1 Theories of conceptual knowledge represented in conceptual space. Axes represent the relative emphasis that is put on bottom-up or top-down processing on the one hand, and on perceptual or motor systems on the other hand. 575 25-Haggard-Chap25 576 5/5/07 11:34 PM Page 576 ACTION SEMANTICS (substances, materials and liquids), while performance with man-made artifacts was relatively spared. This suggests that retrieval of specific conceptual categories may be highly dependent upon the processing of sensory qualities. In contrast, Caramazza and Mahon (2003) propose that evolution favored domain-specific knowledge systems that resulted in a category-specific organization of semantic knowledge. According to the correlated structure principle, the organization of conceptual knowledge reflects the way in which different object properties are related to each other. Proponents of this alternative account usually point to the fine-graininess of the categories of category-specific deficits, e.g. independent impairment for the categories of fruits or animals (Caramazza and Shelton, 1998) and to the absence of a direct relation between severity of affected semantic modality and loss of category-specific semantic deficits. For example, Laiacona et al. (2003) report a patient with a good performance for ‘sensory quality’ categories, as assessed by verbal description compared to performance in a visual–verbal task, while showing a deficit in knowledge for living things. The assumption of a domain-specific organization of conceptual knowledge is consistent with the observation of fine-grained dissociations between different semantic categories. Perceptual symbols theory Perceptual symbols theory provides a perceptual theory of knowledge, according to which schematic representations are extracted from perceptual experience through a process of selective attention. A concept is represented by running a simulation of actual experiences with the concept and this process is referred to as re-enactment (Barsalou et al., 2003). For example, when viewing a visual stimulus, neurons in sensory areas are activated and conjunctive neurons in association areas store the activated sensory features in memory. In the absence of the stimulus, the stored representation can be retrieved through reenactment of the sensory areas by the association areas. Perceptual symbols theory has received empirical support for example from property verification studies in which subjects are asked to decide whether a given property is related to a particular concept (Pecher et al., 2004). Properties on subsequent trials could be either from the same or from a different modality. A modality-switching effect was found when a concept was previously presented with a property from a different modality. For example, reaction times were faster when verifying loud (auditory modality) for blender after previously verifying rustling (auditory) for leaves than after verifying tart (gustatory) for cranberries. These results suggest that subjects tend towards simulating a concept in a particular modality that was previously activated, thereby supporting a modal theory of conceptual representation. However, perceptual symbols theory focuses mainly on perceptual representations, leaving aside the organization of action knowledge, which is represented in Figure 25.1. Ideomotor theory of action We now turn to theories that explicitly focus on the representation of conceptual knowledge used in action. An intuitive and empirically validated idea dates back to William James, who stated that ‘every representation of a movement awakens in some degree the actual 25-Haggard-Chap25 5/5/07 11:34 PM Page 577 THEORIES OF CONCEPTUAL KNOWLEDGE movement which is its object’ (James, 1890). Although ideomotor theory initially emphasized the intentional and private mental causes of action, it was hypothesized that goal representations are crucially important in action control. According to the ideomotor theory, actions are represented in terms of their effects and these effect-representations are helpful in the selection of actions, by predicting the consequences of one’s actions (Hommel et al., 2001). The acquisition of effect-representations depends strongly upon the formation of response–effect associations, in which a given response becomes paired with its associated sensory effects. A preschooler pressing a button of the television’s remote control will soon find out the relation between a simple motor act and the switching on of the television. Consequently, this acquired response–effect association may guide future behavior. In a typical experiment, it is shown that after training participants that a left key response elicits a high tone, a response facilitation occurs for left responses upon hearing the high tone (e.g. Hommel, 1996). According to the ‘theory of event’ coding, actions and perceptual contents are coded in a common event-code (Hommel et al., 2001). In principle, event-codes contain stored representations of contingent response–effect couplings. Thereby, the main focus of ideomotor theory of action is on the automatic processes by which arbitrary connections are established. The selection process, whereby an actor chooses a particular action on the basis of its stored effect-representations, remains largely out of focus. Associative sequence learning A closely related view that was specifically intended to account for imitation of observed actions is called associative sequence learning (ASL). Instead of focusing on response–effect couplings, ASL emphasizes the coupling between stimuli and particular responses. In principle, any association could develop between a stimulus and a particular response if they are co-activated repeatedly. Therefore, the way in which much of our knowledge is organized consists of contingent stimulus–response associations. Heyes et al. (2005) have argued that imitation relies on learned associations between stimuli and actions. Participants had to respond to the observation of hand movements, by making a hand movement that was either congruent or incongruent to the observed movement (opening and closing). A stimulus–response compatibility effect was found for the observation of hand movements compatible with their own movement. However, when subjects were given a training 24 h prior to testing, in which they learned to close their hands in response to observed hand opening and vice versa, the stimulus–response compatibility of observed movement disappeared. Apparently, the automatic imitation of observed movements is mediated by experience and the authors suggest that imitation relies on processes of associative learning. According to this interpretation, neural connections involved in action observation and motor activation primarily reflect the learned association between observing and executing a particular action. Therefore, action representations consist of the learned association between stimuli and particular responses. Accordingly, ASL puts a big emphasis on bottom-up processing in the acquisition of conceptual knowledge for action. 577 25-Haggard-Chap25 578 5/5/07 11:34 PM Page 578 ACTION SEMANTICS Two-route model to action Although many of our everyday actions rely on stored long-term representations—as is the case for bringing a cup to your mouth, which you have probably done countless times—we sometimes need to bypass this semantic knowledge, for example when you intend to catch a spider with a cup. Rumiati and Humphreys (1998) propose a two-route model to action, with a semantic route accessing stored action representations and a second route that runs directly from perception to action. This two-route model has been proposed, partly based on the study of neuropsychological patients showing a dissociation between semantic knowledge and the ability to manipulate objects (e.g. Buxbaum and Saffran, 2002). In a behavioral study with neuropsychologically healthy adults, participants performed either gesturing responses (gesturing the use of an object) or naming responses (verbally giving names) in response to pictures of objects (Rumiati and Humphreys, 1998). Responses for both speech and gestures were distinguished according to whether a visually related error (e.g. razor instead of hammer which have a similar shape but a different function) or a semantically related error (e.g. hammer instead of saw which differ in shape but are semantically related) was made. More visual errors relative to semantically related errors for gesturing object use as compared to naming objects were found, suggesting that naming and using tools relies on different processing routes. Furthermore Rumiati and Tessari (2002) found that healthy volunteers imitated meaningful actions better than meaningless actions and this enhanced performance contributed to the availability of stored action goals in long-term memory. For meaningless actions, no long-term representation is available and imitation therefore relies on a direct route from vision to action, thereby placing more demands on action working memory. In a positron emission tomography study the neural mechanisms underlying imitation of both meaningful and meaningless actions were investigated (Rumiati et al., 2005). Imitation of meaningful actions activated the left inferior temporal gyrus, the angular gyrus and the parahippocampal gyrus (ventral stream), while the right parieto-occipital junction and superior parietal areas (dorsal stream) were mainly activated during imitation of meaningless actions, thereby supporting the view that both actions are mediated by different neural and functional systems. Imitation and execution of meaningful actions rely on accessing stored action semantic knowledge and online visual control, while the execution of meaningless actions in first instance relies on a dorsal perception–action system. Overview Theories discussed thus far disagree about the mechanisms underlying conceptual knowledge (e.g. perceptual simulation, learned association, direct or semantic routes to action). Each theory is based on specific neuropsychological findings and puts different emphasis on the importance of particular sensorimotor areas in the retrieval of conceptual knowledge. As represented in Figure 25.1, we propose that a theory of action semantics should take into account the important role played by the actor’s intentions. We argue that action goals play an important role in the acquisition and use of conceptual knowledge for action. 25-Haggard-Chap25 5/5/07 11:34 PM Page 579 SEMANTICS OF ACTION AND LANGUAGE Furthermore, we agree that conceptual knowledge for action is represented in modalityspecific brain areas, but that the organization and use of conceptual knowledge is guided by action goals. In the next section we will provide support for our position by presenting recent studies investigating effects of action on language processing. Before doing so, however, we will discuss the close relation between language and action in general. The topic of the organization of conceptual knowledge will be revisited in the final section, when we integrate empirical findings with contemporary theories of conceptual knowledge. Semantics of action and language Studies on the relation between language and action may give us more insight into the organization and selection of conceptual knowledge. Different experimental methods have established that action preparation can be affected by language and that action preparation involves accessing semantic knowledge. In the present section we will argue that action and language are highly dependent processes. Although experiments discussed might seem to suggest unidirectional effects from one subsystem to the other, we would like to emphasize the common functional and neural mechanisms that underlie reported effects. Links between language and action In recent years a couple of studies have suggested that action planning and semantic processing are mutually dependent processes. For instance, Creem and Proffitt (2001) asked subjects to grasp common household objects while at the same time they either performed a visuo-spatial or a semantic task. Concurrent semantic processing resulted in more inappropriate grasping of the objects as compared to concurrent visuo-spatial processing, suggesting the involvement of the semantic system when preparing to grasp a meaningful object. Other studies showed that semantic properties of distracting words or objects influenced the planning and execution of grasping movements (e.g. Jervis et al., 1999; Glover and Dixon, 2002). Glover and Dixon (2002) had participants grasping objects, which resulted in an enlargement of maximum grip aperture for objects with the word ‘large’ printed on top of them as compared to objects with the word ‘small’. In a similar way Gentilucci and Gangitano (1998) found that the words denoting ‘far’ and ‘near’ printed on objects affected movement kinematics similarly to the actual greater or shorter distances between hand position and object. These findings fit well with Glover’s model of action control, according to which action preparation involves semantic processing, while action execution operates relatively independently from semantic processes (Glover, 2004). Glenberg and Kaschak (2002) report the action sentence compatibility effect when participants judge the meaning of a sentence by making a movement towards or away from the own body. Participants showed difficulty when judging sentences that implied an action in the direction opposite to the response required for the sensibility judgment (e.g. responding ‘yes’ by making a movement towards the body while judging the 579 25-Haggard-Chap25 580 5/5/07 11:34 PM Page 580 ACTION SEMANTICS sentence ‘Close the drawer’, which implies movement away from the body). Interestingly, the action sentence compatibility effect was found for imperative sentences (e.g. ‘Close the drawer’), as well as for sentences describing the transfer of both concrete (e.g. ‘Liz gave you the book’) and abstract entities (e.g. ‘Liz told you the story’). A similar effect was found by Zwaan and Taylor (2006) who had subjects judge the sensibility of sentences describing actions implying a particular direction by rotating a manual device (e.g. ‘He turned down the volume’ implies movement in a counter-clockwise orientation). A congruency effect was found for direction of manual responses and implied direction by the sentences, suggesting that understanding action sentences relies on motor resonance. These findings support the idea that understanding of meaning is grounded in bodily activity. According to the principle of Hebbian learning, the efficacy of a neuronal ensemble onto another is strengthened whenever they become repeatedly co-activated (Hebb, 1949). This neuronal principle might underlie the close relation between language processing and action preparation. During language acquisition, for instance, children often develop specific links between a particular action and an associated word, probably resulting in a distributed functional network representing the meaning of words (Pulvermüller et al., 2005). Electrophysiological support for the involvement of the motor system in the understanding of action-related sentences comes from a study by Pulvermüller et al. (2005) that investigated the neural correlates of the processing of action verbs. Words that referred to specific body parts (e.g. ‘kick’ refers to foot, ‘lick’ to tongue etc.) elicited activity across the motor-strip close to the representation of the body parts to which the action words primarily referred. Tettamanti et al. (2005) identified a fronto-parietal circuit underlying listening to action-related sentences. This finding has been replicated several times and with different methods, including functional magnetic resonance imaging and transcranial magnetic stimulation (e.g. de Lafuente and Romo, 2004; Buccino et al., 2005) and suggests an underlying representation of action words, that consists of somatotopically organized neuronal assemblies. Semantic information in action preparation Taken together, recent findings suggest close relations between language and action systems. Although most studies discussed thus far investigated the influence of language on action, hardly any experiments have been conducted to explore the activation of semantic knowledge during action preparation. In a series of experiments we investigated whether the intention of an actor to use an object would involve selection of relevant semantic knowledge (Lindemann et al., 2006). Semantic activation in action preparation was investigated by using a language task in which priming effects of action context on the subsequent processing of words were measured. Semantic priming effects have reliably been reported when semantic context (e.g. a prime word or picture) facilitates processing of semantically related words, both for linguistic and nonlinguistic stimuli (e.g. Neely, 1991; Maxfield, 1997). For example, reading the word ‘doctor’ facilitates subsequent recognition of the word ‘nurse’, and no priming effect is expected if the 25-Haggard-Chap25 5/5/07 11:34 PM Page 581 SEMANTICS OF ACTION AND LANGUAGE subsequent word would be ‘fireman’. Considering the importance of goals in stored representations for action, we hypothesized that the preparation of an action would provide a semantic context for the processing of a subsequently presented word, resulting in a priming effect for words consistent with the goal of the upcoming action. The experimental set-up is represented in Figure 25.2. In front of the subject two objects (a magnifying glass and a cup) were placed at a table. A picture of one of the two objects appeared on the screen, and subjects were instructed to prepare the action associated with the object. Subjects were instructed to prepare the execution of a particular motor action (e.g. drink from a cup), but to delay their response until a word appeared on the computer screen. Subjects were instructed to withhold their response when a pseudoword was presented. Only when the word was a valid Dutch word were subjects allowed to execute the prepared movement. In the first experiment, presented words could be either congruent or incongruent with the action goal (e.g. for the cup ‘mouth’ is incongruent whereas ‘eye’ is incongruent). To control for the possibility of a simple picture-word priming effect, half of the subjects had to indicate by a simple finger-lifting response at which side of the table (left or right) the indicated object was present. If selection of semantic knowledge critically depends on the action intention of the actor, interactions between action planning and semantic processing were only expected in the grasping condition. Results are summarized in Figure 25.3 and, as can be seen, a word-congruency effect was found only when subjects intended to display + object 1 object 2 initial startposition Figure 25.2 Schematic representation of the experimental set-up. A cup (Object 1), a magnifying glass (Object 2) and a computer screen were placed in front of the subject. Grasping of the objects was initiated by releasing the right hand from the starting position. 581 25-Haggard-Chap25 11:34 PM Page 582 ACTION SEMANTICS grasp the object and to bring it to the associated goal. That is, subjects responded faster when the presented word was congruent with their present action intention. However, an alternative explanation for the absence of a congruency-effect in the finger-lifting condition might be that preparing a finger-lifting response is much easier than preparing a grasping movement. As a consequence, participants may have been better able to cognitively separate action preparation from word recognition, resulting in the absence of a behavioral interaction effect. In a second experiment this alternative explanation was ruled out by replacing words that represented the goal locations of the grasping movements by Dutch words denoting ‘left’ and ‘right’ that were more relevant to the finger-lifting task where subjects responded with left or right finger movements to indicate object position. Results showed a reversal of priming effects revealing priming for the finger-lifting condition but not for the grasping condition, confirming the hypothesis that action semantics are selectively activated in accordance with the action intended by the subject (see Figure 25.3). In the third experiment a semantic categorization task was used instead of a lexical decision task, to ensure deep semantic processing of the words and to exclude the possibility that subjects’ responses in the first two experiments only relied on visual word forms. Experiment 2 Experiment 1 Object-congruent words Object-incongruent words 540 520 RT (ms) RT (ms) Consistent words Inconsistent words 540 520 500 500 480 480 460 460 440 440 Object-grasping Finger-lifting Object-grasping Finger-lifting Experiment 3 & 4 Object-congruent words Object-incongruent words 540 520 RT (ms) 582 5/5/07 500 480 460 440 Object-grasping Object-grasping Figure 25.3 Mean response latencies arranged by action condition and word consistency. Subjects performed a lexical decision task with words representing goal locations in Experiment 1 and words representing spatial locations Experiment 2. Subjects performed a semantic categorization task in Experiment 3 and a final letter identification task in the Experiment 4. Error bars represent the 95% within-subject confidence intervals. RT, reaction time. 25-Haggard-Chap25 5/5/07 11:34 PM Page 583 SEMANTICS OF ACTION AND LANGUAGE Target words again included body parts that represented the goal locations of the actions associated with either the cup or the magnifying glass. In addition, action-unrelated words that referred to body parts were used as filler items and words that referred to animals were chosen as no-go stimuli. Subjects were instructed to execute the action associated with the object, only when the word referred to a body part. No response was to be executed when words denoted animals. Again, a reduction in response latencies was found for words that were consistent with the action goal of the subject (e.g. ‘eye’ for grasping the magnifying glass). Finally it was hypothesized that if the interaction between words and action intention critically depended on deep semantic processing of the words, the effect should disappear if subjects attended to words only at a superficial level. As can be seen in Figure 25.3 the action-priming effect indeed disappeared if subjects performed a simple final letter-identification task instead of semantic categorization. Together these experiments suggest that action goals form an integral part of the semantic representations that are activated during action preparation. Opposite to many of the automatic effects that were found in earlier studies investigating congruency effects between presented objects and required grips, when it comes to action goals, or functional knowledge of object use, action semantics are not automatically activated upon observation of an object, as is shown by the absence of the word priming effect in the finger-lifting condition in the first experiment, but are selected in accordance with the action intention of the actor. Semantic activation in preparing meaningful and meaningless actions Although in everyday life many objects are normally associated with typical goal locations (‘mouth’ for ‘cup’), it is often the case that one object can be used in many different ways, depending on the context within which the object is perceived (e.g. using a knife to spread butter or to cut your meat). An important aspect in the evolution and selection of human behavior seems to be the ability to flexibly adjust one’s strategies to environmental demands. Concerning the acquisition of action semantics, besides developing longterm representations of object use, it is equally important to be able to implement short-term action intentions (e.g. hitting a mosquito with a newspaper). To investigate the effects of short-term action goals upon the activation of semantic information, we provided subjects with the instruction to perform meaningful and meaningless actions with a cup and a magnifying glass (M. Van Elk et al., unpublished data). In half of all blocks subjects were instructed to bring both objects to their mouth, while in the other half they were encouraged to bring both objects to their right eye. A semantic categorization task was used and subjects had to delay the execution of the action until they recognized a word that represented a body part (similar to the third experiment in Lindemann et al., 2006). It was hypothesized that if subjects prepared to execute a meaningful action that is normally associated with the use of the object (e.g. bringing the cup to their mouth) a similar word-congruency effect could be expected as in the previous experiments (e.g. faster responding to ‘mouth’ as compared to ‘eye’ for intending to use the cup). However, with the intention to perform a meaningless action (e.g. bringing the 583 25-Haggard-Chap25 11:34 PM Page 584 ACTION SEMANTICS cup to the eye), long-term action knowledge has to be overruled in order to comply with the short-term action intention of the actor. Results are represented in Figure 25.4 and indicate that when subjects performed actions normally associated with the objects they were faster to respond to words that consistently described the goal-location of the action, replicating the results of Lindemann et al. (2006). Interestingly, when subjects intended to perform an incorrect action with an object, response latencies were sped up for words describing the short-term goal location of the intended action (e.g. faster responding to ‘eye’ compared to ‘mouth’, when intending to bring the cup to the eye). One possible confounding factor, however, may be that priming effects were not caused in accordance with the intended actions but rather via the task instruction that subjects received prior to each block. That is, subjects were instructed to move objects either to their mouth or to their eye in individual blocks, which may have facilitated the processing of the same words ‘mouth’ or ‘eye’ presented on the screen. In a second experiment Experiment 1 Experiment 2 640 620 600 640 Object-congruent words Object-incongruent words 620 600 Object-congruent words Object-incongruent words 580 RT (ms) RT (ms) 580 560 540 560 540 520 520 500 500 480 480 460 460 Meaningful Actions Meaningless Actions Meaningful Actions Meaningless Actions Experiment 3 Object-congruent words Object-incongruent words 640 620 600 580 RT (ms) 584 5/5/07 560 540 520 500 480 460 Meaningful Actions Meaningless Actions Figure 25.4 Mean response latencies arranged by action condition (meaningful or meaningless actions) and word consistency. Subjects conducted a semantic categorization task to words appearing on the screen. In the first experiment subjects performed meaningful and meaningless actions within the same block, while in the second experiment meaningful and meaningless actions were conducted in separate blocks. In the third experiment a concurrent articulatory suppression task was conducted. Error bars represent the 95% within-subject confidence intervals. RT, reaction time. 25-Haggard-Chap25 5/5/07 11:34 PM Page 585 THEORETICAL IMPLICATIONS we tried to rule out this explanation by having subjects prepare meaningful and meaningless actions in separate action blocks, resulting in both action goals being used within the same block. Results again revealed a priming effect for words congruent with the action goal in meaningful action blocks in which subjects used both objects in the usual fashion (see Figure 25.4). However, the effect was found to reverse in the meaningless action blocks, in which participants responded faster to words that were congruent to the shortterm action goal. Apparently, the action intention of the actor can overrule long-term conceptual associations. However, given the unusual task constraints it might be that subjects relied on a shortterm verbalization strategy in performing the task. When performing meaningless actions, subjects might have been silently repeating the goal location of the action, to avoid being confused by the words appearing on the screen. To prevent subjects from relying on a verbal strategy, we replicated the first experiment, but in addition subjects conducted a concurrent articulatory suppression task. Even when subjects were prevented from using a verbal strategy, they responded faster to words that were congruent with the short-term action goal, as can be seen in Figure 25.4. These findings convincingly show that the intention to perform a meaningless action with an object can overrule priming effects from long-term conceptual knowledge associated with the object. Instead, it appears that the present action goal of the actor determines the selection of semantic information in accordance with the intention of the actor. Theoretical implications of these findings will be discussed in the final part of this chapter. Theoretical implications As discussed in the section ‘Theories of conceptual knowledge’, a growing number of studies support the view that human cognition is deeply rooted in bodily activity (Gallese and Lakoff, 2005). Nowadays, discussion focuses on the way in which conceptual knowledge is organized and represented over different brain areas. In the present chapter we proposed the concept of action semantics as a useful way to think about conceptual knowledge for action. We argued that an important factor in the acquisition and use of action semantics is the action goal that is selected for a particular object. In the third section, two recent studies were discussed in more detail to support our view that semantics for action become activated selectively in accordance with the goal of the actor. When intending to grasp a cup, subjects responded faster if the target word consistently described the spatial goal of the intended action (Lindemann et al., 2006). However, when subjects were instructed to perform meaningless actions with the objects, the word-congruency effects reversed according to the present action intention of the actor (M. Van Elk et al., unpublished data). That is, subjects responded faster to the word ‘eye’ if they were intending to bring the cup to their eye, irrespective of the learned association in long-term memory to bring cups to the mouth. In this final section we will compare different approaches to conceptual knowledge for action in relation to the presented findings. First, dissociations between functional knowledge (‘what to use an object for’) and pragmatic knowledge (‘how to use an 585 25-Haggard-Chap25 586 5/5/07 11:34 PM Page 586 ACTION SEMANTICS object’) in neuropsychological patients are in line with our view that action goals and action means are two distinct aspects of action semantics (Leiguarda and Marsden, 2000). Exactly how action goals and means are represented in the brain is a matter of ongoing study. Different parts of the motor system appear to contribute to the retrieval of conceptual knowledge for action consistent with the sensory–functional hypothesis which argues for conceptual knowledge being organized according to modality (Warrington and Shallice, 1984). A similar view is proposed by perceptual symbols theory (Barsalou, 1999) according to which conceptual knowledge is represented in modality-specific brain systems. Although perceptual symbols theory does not explicitly focus on conceptual knowledge used in action, the retrieval of conceptual knowledge about object use is believed to involve sensorimotor simulation that includes all necessary features of a previously encountered object. For example, a conceptual representation of ‘cup’ is acquired by countless interactions with cups and related actions directed towards them. Therefore the conceptual representation of cup would probably include a perceptual simulation of a prototypical ‘cup’ and a motor simulation of bringing the cup towards the mouth. Perceptual symbols theory could account for priming effects resulting from preparation of meaningful actions for which a perceptual simulation is available (Lindemann et al., 2006). However, it is more difficult for the theory to explain the reversal of priming effects that were found with meaningless or unusual actions as presented in ‘Semantics of action and language’ (M. Van Elk et al., unpublished data). Perceptual symbols theory takes the prototypical concept as the cornerstone around which our conceptual knowledge is built. It would be interesting to think about what kind of simulations might underlie the flexible use of conceptual knowledge. Given the short-term priming effects in congruence with the immediate action intention it might be that a perceptual simulation also includes the short-term goal. Whenever a person intends to bring a magnifying glass to the mouth, the goal-related concept ‘mouth’ might form part of the action representation. However, the way in which perceptual simulations are flexibly selected by combining different representations into new nonexisting concepts or by preparing novel actions would need to be clarified. The two-route model to action proposes a direct route from vision to action that bypasses semantic information (Rumiati and Humphreys, 1998; Rumiati and Tessari, 2002). The two-route model accounts well for the failure of neuropsychological patients to imitate either meaningful or meaningless actions. Within this framework the findings of M. Van Elk et al. (unpublished data), in which participants performed either meaningful or meaningless actions with common objects, might be interpreted as reflecting the differential use of both pathways to action. When participants intend to perform meaningful actions with objects the semantic route is activated, in which long-term conceptual knowledge about the use of objects is represented. In contrast, the indirect route from perception to action might be used when participants perform meaningless actions with objects. However, considering that priming effects were found both when subjects made appropriate and inappropriate actions with objects, we suggest that semantics are involved for both meaningful and meaningless actions. That is, the two-route model for 25-Haggard-Chap25 5/5/07 11:34 PM Page 587 THEORETICAL IMPLICATIONS action cannot account for the involvement of semantics in meaningless actions. Rather than emphasizing the difference between meaningful and meaningless actions, we suggest that a crucial factor of all actions is the current goal of the actor. According to ideomotor theory of action, stored effect-representations enable people to select the appropriate action. Event-codes consist of contingent response–effect couplings that have been shaped over the course of our lives. Accordingly, preparing to grasp a cup will be associated with the typical goal of bringing it to the mouth. In a similar way the principle of associative learning states that stimulus–response associations reflect the amount of experience of acting in a particular way towards visual stimuli. However, ideomotor theory and associative learning face major difficulties when it comes to explaining short-term priming effects of action intention, as discussed in the present chapter. When subjects prepared meaningless actions with objects that they obviously had never conducted before, we found a selective activation of words that were congruent to the present short-term action goal. If priming-effects of action intention rely solely on associative learning we might have expected them to disappear when subjects prepared meaningless actions, just like automatic effects of imitation could be cancelled out by training participants to respond in a different way towards stimuli (Heyes et al., 2005). In contrast, we have shown that priming of words that are congruent to normal object use could be reversed when subjects had an action intention different from normal object use (M. Van Elk et al., unpublished data). This word-congruency effect reveals a counterstatistical association, as most participants probably will not have much experience bringing cups to their eye. However, it might be argued that while participants repeatedly brought objects towards incongruent action goals, new short-term associations between grasping the object and the incongruent goal indeed were established. According to the associative learning approach (Heyes et al., 2005), long-term stimulus–response associations were learned by practice and can be flexibly adjusted by training. Within the domain of imitation Heyes et al. (2005) convincingly showed that training participants to respond in an incongruent way to observed actions could cancel out the effects of automatic imitation. In a similar way, novel semantics for action might be established when subjects repeatedly conducted unusual actions towards objects. By allowing the flexible acquisition of novel action semantics, both ideomotor theory and the principle of associative learning do a better job than other theories of conceptual knowledge in action. However, in contrast with both theories we suggest that the acquisition of novel action semantics depends heavily on the goal of the actor. It is not the case that associations are arbitrarily consolidated according to incoming bottom-up stimulus information. Typically, the action intention of a person determines what kind of information is selected and attended. For example, it does not seem plausible that the simple observation of a cup automatically elicits a complete motor program that subsequently needs to be inhibited. A much simpler view is that what you intend to do with a cup determines the kind of information that is selected. To conclude, we propose that the selection of action semantics depends critically on the action intention of the actor. Whenever a usual action is conducted with an object, 587 25-Haggard-Chap25 588 5/5/07 11:34 PM Page 588 ACTION SEMANTICS associated semantic information is being activated accordingly, whereas when an object is used in an inappropriate manner, the short-term behavioral goal temporarily overrules the long-term associations. The selection processes underlying action semantics therefore bear resemblance to ‘selection-for-action’ in perception (Allport, 1987). Conclusions In sum, in the present chapter we sought to outline the concept of action semantics by presenting a review of existing literature on functional mechanisms underlying conceptual knowledge and goal-directed behavior supplied with recent empirical evidence supporting selection-for-action at the level of semantics. Our findings suggest a tight connection between semantic knowledge that is used in action and language, reflected by conceptual priming effects from action preparation to language processing. These findings suggest that the theory of selection-for-action (Allport, 1987) may be extended to include selection of semantic information that is relevant for the intended action and that the theory of selection-for-action may provide a useful framework for studying the associations between language, action and perception. We suggest that, comparable with the selection of perceptual information that is relevant to our current intentions and behavioral goals, the selection of action semantics may be equally important for smoothly moving throughout the world. Access to stored long-term conceptual action representations and acquisition of novel action semantics both enable us to adapt our behavior flexibly to continually changing environmental demands. The study of action semantics provides important insight into the organization of our conceptual knowledge and supports the idea that action knowledge is represented in sensorimotor systems. In addition, mechanisms of action, perception and language should not be studied in isolation, since the retrieval of semantic information involves action, visual as well as lexical semantics. Developmental studies are expected to provide important insight into the acquisition of action semantics and neuroimaging studies may help to understand the neural organization underlying knowledge of action goals and action means. Behavioral and electrophysiological approaches may contribute to generate further insight into the timing of activation and the flexible selection of conceptual knowledge that guides our behavior, providing new and exciting directions for future research. References Allport, A. (1987) Attention and selection-for-action. In Heuer, H. and Sanders, A. F. (eds), Perspectives on Perception and Action, pp. 395–420. Erlbaum, Hillsdale, NJ. Barsalou, L. W. (1999) Perceptual symbol systems. Behavioral and Brain Sciences, 22, 577–660. Barsalou, L. W., Simmons, W. K., Barbey, A. K. and Wilson C. D. (2003) Grounding conceptual knowledge in modality-specific systems. Trends in Cognitive Sciences, 7, 84–91. Bekkering, H. and Neggers, S. F.W. 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