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Haptic Perception: a Tutorial Attention, Perception, & Psychophysics 2009, 71 (7), 1439-1459 doi:10.3758/APP.71.7.1439 TU T ORIAL REVIEW Haptic perception: A tutorial S. J. LEDERMAN Queen’s University, Kingston, Ontario, Canada AND R. L. KLAT Z K Y Carnegie Mellon University, Pittsburgh, Pennsylvania This tutorial focuses on the sense of touch within the context of a fully active human observer. It is intended for graduate students and researchers outside the discipline who seek an introduction to the rapidly evolving field of human haptics. The tutorial begins with a review of peripheral sensory receptors in skin, muscles, tendons, and joints. We then describe an extensive body of research on “what” and “where” channels, the former dealing with haptic perception of objects, surfaces, and their properties, and the latter with perception of spatial layout on the skin and in external space relative to the perceiver. We conclude with a brief discussion of other significant issues in the field, including vision–touch interactions, affective touch, neural plasticity, and applications. Haptics is now commonly viewed as a perceptual sys- Most studies that focus on human sensations involve the tem, mediated by two afferent subsystems, cutaneous and application of various stimuli (hairs, sharp probes, warm kinesthetic, that most typically involves active manual and cool metal tips, etc.) to the skin of a passive observer, exploration (Lederman & Klatzky, 2009). Whereas vi- thereby limiting inputs to those of the cutaneous recep- sion and audition are recognized for providing highly tors. In his seminal 1962 paper on active touch, J. J. Gib- precise spatial and temporal information, respectively, son emphasized the polarity of one’s tactual experiences: the haptic system is especially effective at processing the Being passively touched tends to focus the observer’s at- material characteristics of surfaces and objects. Here we tention on his or her subjective bodily sensations, whereas concentrate on the behavioral research that has addressed contact resulting from active exploration tends to guide the phenomenology and functionality of haptic percep- the observer’s attention to properties of the external envi- tion. This excellent behavioral work stands on its own, ronment. Whereas the results of the passive-touch studies although where directly appropriate we relate it to work in clearly confirm that cutaneous inputs alone are sufficient neuroscience (for more general references, consult, e.g., to induce subjective sensations, they fail to recognize the Kandel, Schwartz, & Jessell, 2000; Squire, 2009). important role of cutaneous sensing when active explora- Because this tutorial is necessarily brief, for certain top- tion is permitted. ics we have also chosen to direct the reader to one or more Cutaneous receptors are found across the body surface, review chapters or books that offer further detailed discus- beneath both hairy and hairless skin. To date, the majority sion and extensive bibliographies containing important of human studies have focused on mechanoreceptors and original sources. The tutorial provides a comprehensive thermoreceptors located within the hairless (“glabrous”) bibliography followed by a list of other suggested review skin of the human hand (Jones & Lederman, 2006). Fig- articles, encyclopedia entries, and books about haptics and ure 1 shows the structure of palmar skin, together with the sense of touch that the reader may wish to consult. the specialized nerve endings of the four mechanorecep- tor populations that human neuroscience has shown are PERIPHERAL SENSORY MECHANISMS distributed within this region (see Johansson & Vallbo, 1983). The response characteristics of each population The haptic system uses sensory information derived are differentiated by both the relative size of its receptive from mechanoreceptors and thermoreceptors embedded field (small vs. large) and its relative adaptation rate (i.e., in the skin (“cutaneous” inputs) together with mechano- response to onset/offset of skin deformation vs. continued receptors embedded in muscles, tendons, and joints (“kin- response during sustained skin deformation), as outlined esthetic” inputs). in Table 1A. Table 1B shows the relatively optimal fea- S. J. Lederman, [email protected] 1439 © 2009 The Psychonomic Society, Inc. 1440 LEDERMAN AND KL ATZ K Y topic of debate), the somatosensory system is served by two subsystems, a “what” system that deals with percep- tual (and memory) functions, and a “where” system that deals with the perceptual guidance of action. Evidence that supports a “what/where” distinction for the somatosensory system include, for example, fMRI and behavioral studies by Reed, Klatzky, and Halgren (2005) and by Chan and Newell (2008), respectively. Reed et al. (2005) showed that haptic object recognition and object localization ac- tivated inferior and superior parietal areas, respectively, suggesting a correlation with the distinction between dor- sal and ventral visual streams made earlier by Ungerleider and Mishkin (1982). Chan and Newell showed behavioral evidence for a task-dependent what/ where distinction that transcends modalities by using a dual-task paradigm. Si- multaneous “what” or “where” tasks were found to mutu- ally interfere more than crossfunction tasks in both in- tramodal and crossmodal conditions, indicating resource Figure 1. Vertical section through the glabrous skin of the pools that depended on the task demands but not on the human hand. Schematic depiction of the two major layers of modality (vision, haptics) used to execute the task. Dijker- the skin (epidermis and dermis), and the underlying subcuta- man and De Haan (2007) have comprehensively evaluated neous tissue. The locations of the organized nerve terminals are also shown. Mr, Meissner corpuscle; Ml, Merkel cell complex; the neural and behavioral literatures for evidence of sepa- R, Ruffini ending; P, Pacinian corpuscle. From “Tactile Sensory rate processing streams used for somatosensory percep- Coding in the Glabrous Skin of the Human Hand,” by R. S. Jo- tion versus action (“what” vs. “how” systems), as well as hansson and A. B. Vallbo, 1983, Trends in Neurosciences, 6, p. 28. for distinguishing between haptic processing of external Copyright 1983 by Elsevier. Reprinted with permission. targets and sites on the body. An important issue that arises from this body of research is whether haptic processing of ture sensitivity, together with the primary functions with shape taps into a visual “what” pathway by invoking vi- which each mechanoreceptor population is associated. sual imagery, a topic we consider further below. The two additional peripheral receptor populations known For purposes of the present tutorial, we will organize as thermoreceptors (Stevens, 1991) respond to increases the following discussions of haptic perception in terms of or decreases in skin temperature, and mediate the human this functional distinction between “what” and “where” experiences of warmth and cold, respectively. systems. The kinesthetic inputs from mechanoreceptors in mus- cles, tendons, and joints contribute to the human percep- The “What” System tion of limb position and limb movement in space (see re- The “what” system in touch processes surfaces, objects, views by Gandevia, 1996; J. L. Taylor, 2009). Research in and their many different properties. The efficacy of this the motor-control field tends to treat kinesthetic feedback processing pathway is demonstrated by the finding that as sensory signals to be included in models (feedback, familiar objects are recognized quickly and with very high feedforward) of limb movement and grasping. Hence, we accuracy by touch alone (Klatzky, Lederman, & Metzger, will consider the contributions of kinesthesis and kines- 1985). The foundation for this ability lies in the sensory thetic inputs only where they are inextricably bound up primitives signaled by the peripheral receptors. A broad with human haptic processing and representation—that spectrum of properties results from further neural pro- is, for purposes of sensing, perceiving, and thinking about cessing of the receptor signals, with research providing objects, their properties, and the space within which they considerable insight into the computational nature of that reside. processing. Cutaneous and kinesthetic inputs are combined and To begin with, it is useful to divide haptically acces- weighted in different ways to serve various haptic func- sible object properties into two broad classes: material and tions. In the discussion that follows, we treat complex geometric. Material properties are defined as those inde- human haptic experience as being influenced by a variety pendent of the particular object sample being considered; of factors at multiple levels of processing. Accordingly, conversely, geometric properties describe the structure of it is neither possible nor particularly fruitful to separate that object sample. human haptic function into modular compartments as was once done (e.g., sensations, percepts, and cognitions). Spatial and Temporal Resolving Capacity of the Skin “WhaT” AND “Where” TOUCH SYSTEMS Before considering in the next section the haptic per- ception of object properties, it is important to be aware of Touch scientists have been recently and vigorously de- the extent to which the cutaneous system is limited by its bating whether, like vision (and audition, a more recent ability to resolve spatial and temporal details presented HAPTIC PERCEPTION 1441
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