Perception, action, and the body model.

In 1992, Goodale and Milner proposed to study the visual system based on function, thus dissociating vision for perception (ventral stream) and vision for action (dorsal stream). This became known as the Perception and Action model (PAM). Following the PAM in the visual system, a somatosensory PAM was proposed including a body representation for perception and a separate for action. This review explores the body model of the hand and how it relates to the PAM. The body model refers to the internal representation of the body that is responsible for position sense. Previous research has shown that the representation of the hand features systematic distortions: an overestimation of hand width and an underestimation of finger length. These distortions have been reported using different paradigms, different body parts, and in various settings. Thus, body model distortions appear to be a characteristic of human body representation. If the body model of the hand is distorted, how can actions like reaching and grasping be accurate? We review evidence that body model distortions may in fact provide a functional benefit to our actions, that cortical maps in the somatosensory and motor cortices reflect these distortions, and that actions rely on a distorted body model. We argue that the body model is a product of both the ventral and dorsal somatosensory streams. Further, we suggest that the body model is an example of the inextricable link between the two streams.

The distorted hand: systematic but 'independent' distortions in both explicit and implicit hand representations in young female adults.

It has long been assumed that an accurate representation of the size and shape of one's body is necessary to successfully interact with the environment. Previous research has shown accurate representations when healthy participants make overt judgments (i.e. explicit) about the size of their bodies. However, when body size is judged implicitly, studies have shown systematic distortions. One suggestion for these differences, is that explicit and implicit representations are informed by different sensory modalities. Explicit representations rely on vision whereas implicit representations are informed by haptics. We designed an experiment to investigate if explicit representations that are informed by haptics are more like implicit representation featuring systematic distortions. We asked female participants to estimate the size of their fingers and hands in three different tasks: an explicit-haptic, an implicit, and an explicit-vision task. The results showed that all three representations were distorted and furthermore, the distortions for each representation were different from one another. These results suggest that inaccurate finger and hand length are a stereotypical feature of body representation that is present in both visual and haptic domains. We discuss the results in relation to theories of body representation.

Growing into your hand: the developmental trajectory of the body model.

We rely on accurate body representations to successfully interact with the environment. As adults, we rely on many years of experience with a body that has stayed relatively the same size. Children, however, go through periods of rapid growth and whether or not their body representation matches this physical growth is unknown. To address this question, we examined the developmental trajectory of the body model of the hand. The body model is the representation of our bodies that underlies position sense. We recruited a group of children (8-16 years) and a control group of young adults (18-26 years) and asked them to complete the body model task. In this task, participants estimated the location of ten different landmarks (the tips and metacarpophalangeal joints of each of their five fingers). The position (XY location) of each estimate was tracked using an Optotrak camera. From the XY locations we derived hand width and finger length. Not surprisingly, children's physical hand width and finger length were smaller than adults but remarkably, the body model, was similar for both groups. This result indicates that children overestimate hand size and suggests that the body model is ahead of physical growth. This result contradicts the notion that body representation lags physical growth during puberty, accounting for the clumsy motor behaviour characteristic of teens. We discuss the results in relation to the different taxonomies of body representation and how an enlarged representation of the hand during childhood may influence action.

Chubby hands or little fingers: sex differences in hand representation.

Disturbed body representation is a condition defined by the perception that one's body size is different from their anatomical size. While equal amounts of males and females suffer from disturbed body representation, there appear to be differences in the direction of this distortion. Females will typically overestimate, whereas males will typically underestimate body size. One part of the body that has been consistently misperceived is the hands. This misrepresentation consists of two distinct characteristics: an overestimation of hand width, and an underestimation of finger length. Many of these studies, however, have used predominately female participants, allowing for the possibility that women are driving this distortion. The aim of the present study was to examine possible sex differences in hand perception. To this end, participants estimated the location of ten landmarks on their hands when their hands were hidden from view. Our results indicate that females follow the characteristic distortion, whereas males only underestimate finger length (albeit more than females). These findings are surprising, because the hands are not an area of concern for weight gain/loss. We discuss these findings in relation to body dysmorphia literature.

Long- but not short-term tool-use changes hand representation.

Tool-use has been found to change body representation. For example, participants who briefly used a mechanical grabber to pick up objects perceived their forearms to be longer immediately after its use (e.g., Cardinali et al., Curr Biol 19(12):R478-R479, 2009; they incorporated the tool into their perceived arm size). While some studies have investigated the long-term effects of tool-use on body representation, none of these studies have used a tool that encapsulates the entire body part (e.g., a glove). Moreover, the relationship between tool-use and the body model (the representation of the body's spatial characteristics) has yet to be explored. To test this, we recruited 19 elite baseball players (EBP) and 18 age-matched controls to participate in a hand representation task. We included EBP because of their many years (8+) of training with a tool (baseball glove). The task required participants to place their hands underneath a covered glass tabletop (no vision of their hands), and to point to where they believed 10 locations (the tips and bases of each finger) were on their hands (Coelho et al., Psychol Res 81(6):1224-1231, 2017). Each point's XY coordinates was tracked using an Optotrak camera. From these coordinates, we mapped out the participants perceived hand size. The results showed that when compared to the controls, EBP underestimated hand width and finger length of both hands. This indicates that long-term tool use produces changes in the body model for both, the trained and untrained hands. We conducted a follow-up study to examine if 15 min of glove use would change perceived hand size in control participants. Novice baseball players (participants without baseball experience: NBP) were recruited and hand maps were derived before and after 15 min of active catching with a glove. Results showed no significant differences between the pre and post hand maps. When we compared between the two experiments, the EBP showed smaller hand representation for both hand width and finger length, than the NBP. We discuss these results in relation to theories of altered body ownership.

The visual and haptic contributions to hand perception.

Previous research has found that the perception of our hands is distorted. The characteristics of this distortion are an overestimation of hand width and an underestimation of finger length. The present study examined the role that different sensory modalities (vision and/or haptics) play in the perception of our hands. Participants pointed to their concealed hand in one of three groups: Vision+Haptics, Vision-only, or Haptics-only. Participants in the Vision+Haptics group had vision (non-informative) of the experimental setup and of the pointing hand, but no vision of the hand being estimated. They also experienced haptic feedback as the palm of the hand was in contact with the undersurface of a tabletop, where the estimations were made. Participants in the Vision-only group, instead of placing the hand to be estimated underneath the tabletop, they placed it behind their backs. Participants in this group were asked to imagine as if the hand was under the table when making their estimations. In the Haptics-only group, participants completed the task with the hand underneath the tabletop (as in the Vision+Haptics group) but did so while wearing a blindfold (no vision). All participants estimated the position of ten landmarks on the hand: the fingertip and the metacarpophalangeal joint of each digit. Hand maps were constructed using a 3D motion capture system. Participants in the Haptics-only group produced the most accurate hand maps. We discuss the possibility that vision interferes with somatosensory processing.

A kinematic examination of hand perception.

Previous research has found that the perception of our hands is inaccurate. This distorted representation has several constant characteristics including an overestimation of hand width and an underestimation of finger length. In this study, we further investigate this phenomenon by exploring the boundaries of hand representation. Participants placed one hand underneath a table top so it was occluded from view. Using their free hand, participants were instructed to point to the location where they believed the tips and bases of each of their fingers were. These ten landmarks were recorded using a motion capture system. One group of participants pointed to the landmarks in a random order (as done in previous studies) while another group pointed to them in a systematic fashion (from the tip of the thumb sequentially through to the pinky). Furthermore, to explore if having a frame of reference facilitates hand perception, some participants initiated each of their estimations directly from the previous landmark while others initiated them from a home spot located outside the span of the hand. Results showed that the participants who pointed in the systematic order made numerous accurate judgments of hand size and were overall more precise than participants who pointed in a random order. Including a frame of reference however, had no effect on the judgments. The results also showed asymmetries in hand perception. These findings are discussed in relation to different possible internal body representations and hemispheric asymmetries in body perception.