B RA I N RE SE A R CH RE V I EW S 54 ( 20 0 7 ) 2 1 9–2 3 2

a v a i l a b l e a t w w w. s c i e n c e d i r e c t . c o m

w w w. e l s e v i e r. c o m / l o c a t e / b r a i n r e s r e v


Central mechanisms in phantom limb perception: The past, present and future
Melita J. Giummarra a,b,⁎, Stephen J. Gibson b,c , Nellie Georgiou-Karistianis a , John L. Bradshaw a
Experimental Neuropsychology Research Unit, School of Psychology, Psychiatry and Psychological Medicine, Monash University, Clayton VIC 3800, Australia b National Ageing Research Institute, Parkville, VIC, Australia c Caulfield Pain Management and Research Centre, Caulfield, VIC, Australia

A R T I C LE I N FO Article history: Accepted 28 January 2007 Available online 1 February 2007 Keywords: Phantom sensation Phantom pain neuromatrix Mirror neuron Body schema

AB S T R A C T Phantom limbs provide valuable insight into the mechanisms underlying bodily awareness and ownership. This paper reviews the complexity of phantom limb phenomena (proprioception, form, position, posture and telescoping), and the various contributions of internal constructs of the body, or body schema, and neuromatrix theory in explaining these phenomena. Specific systems and processes that have received little attention in phantom limb research are also reviewed and highlighted as important future directions. These include prosthesis embodiment and extended physiological proprioception (i.e., the extension of the body's “area of influence” that thereby extends one's innate sense of proprioception), mirror neurons and cross-referencing of the phantom limb with the intact limb (and the related phenomena of perceiving referred sensations and mirrored movements in the phantom from the intact limb). The likely involvements of the body schema and the body–self neuromatrix, mirror neurons, and cross-callosal and ipsilateral mechanisms in phantom limb phenomena all suggest that the perception of a “normal” phantom limb (that is, a non-painful phantom that has the sensory qualities of an intact limb) is more than likely an epiphenomenon of normal functioning, action understanding and empathy, and potentially may even be evolutionarily adaptive and perhaps necessary. Phantom pain, however, may be a maladaptive failure of the neuromatrix to maintain global bodily constructs. © 2007 Elsevier B.V. All rights reserved.

1. 2. Introduction . . . . . . . . . . . . . . . . . . . . . Proprioception of the phantom limb . . . . . . . . 2.1. Distorted perception of the phantom limb . 2.2. Telescoping of the phantom limb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 220 220 221 221

⁎ Corresponding author. Experimental Neuropsychology Research Unit, School of Psychology, Psychiatry and Psychological Medicine, Monash University, Clayton VIC 3800, Australia. Fax: +61 3 9905 3948. E-mail address: melita.giummarra@med.monash.edu.au (M.J. Giummarra). 0165-0173/$ – see front matter © 2007 Elsevier B.V. All rights reserved. doi:10.1016/j.brainresrev.2007.01.009

. . . . . . . . . . 2001). . . . . eye (Sörös et al. . Mikulis et al. . . . . . . . . or throbbing. . . 1974.2. . . Jamison et al. . . . . 4. . . or under the amputee's volitional control (Roux et al. . and the evidence that the body schema (Section 4. . . . 2001. . . . The duration of phantom limb perception also varies between individuals. . . for example. .5. . . . . . . . Mirror neurons. . Cross referencing between the intact limb and the phantom limb . . . . . . . 2000). Phantom sensations are reported most commonly following the amputation of an arm or leg. . . 1979) and rectum (Cherng et al. . . Mirrored sensation and movement from the intact limb to the phantom limb . . This review proposes that – with the likely involvement of the body schema. . . . and potentially even evolutionarily adaptive and perhaps necessary. References. . . . . . . . .5. . . B RA I N R E SE A R CH RE V I EW S 54 ( 20 0 7 ) 2 1 9–2 3 2 Prosthesis use and embodiment . . . . . . .. . Weinstein. . .. . . Current theories of phantom limb perception are also reviewed. . . . . . ..220 3. or some part thereof (Ramachandran and Hirstein.g. . . . . .1. . . . 1994). Le Chapelain et al. . . . . . . Phantom sensations are perceived immediately after limb loss by most amputees (Ramachandran and Hirstein. . . . Acknowledgments . . .2. . . . 1992).1. . . . 2004). . 1998). . . . below). . . . . resting at the side of the body. 1998). . . 1968. . . .5. 2006a) (Section 4. . .. . . . . . nevertheless perceived by up to 98% of amputees following amputation (Ramachandran and Hirstein. . . . . . The body schema . and the more recently proposed roles of the fronto-parietal mirror neuron system (Brugger. . mirror systems. . if at all (Kooijman et al. below). The mechanisms of cross-referencing the phantom limb with the opposite limb are also considered (Section 4. . . . . . .. . 2001. . . . or in a posture that resembles the posture of the limb prior to amputation (Ramachandran and Hirstein.4. 4. . . .. . . . . . . . . . . . . . . . . amputees with phantom sensations may have a greater “postural empathy” for others such that they are better able to match their own body schema against the observed bodies of others. . . . . . teeth (Marbach. Spontaneous changes in posture of the phantom limb are also common in amputees (Ramachandran and Hirstein. limb . . 4. . however for some. . . . . . . . Introduction Phantom limbs are a seemingly curious phenomenon. . . . . . . . . 90). . . . . . . 1955. . Mirror neurons and action understanding . . bladder (Arcadi. .. . . . Farley and Smith. . . . . .. shape and posture – and may be perceived to be completely paralysed. Moore et al. . and “pain matrix” mirror system (Giummarra et al. and by about 20% of children with congenital limb aplasia (Melzack et al. . . . . . . . . . . . . .2. . . . . . . . . . . . Phantom phenomena problematic for current theory . Henderson and Smyth (1948) reported that the phantom tends to be “correctly aligned to the stump with which it moves” (p. . 1998) and in (normal) patients who are under anaesthesia (Bromage and Melzack. Proprioception of the phantom limb Phantom limbs are generally perceived to occupy veridical body space – being of a particular size. Often. 1997) and sometimes to . or to move spontaneously or reflexively (Ramachandran and Hirstein. . 2001. . . . non-painful phantom limb is very likely to be an epiphenomenon of normal functioning. . Ovesen et al. . . . . . . .1. Phantom sensations following removal of visceral organs may be painful in nature (for example. The cortical origins of phantom limb phenomena .. including Melzack’s (1990) neuromatrix theory (Section 4. . . . . . . . . . . 4. . . . . 2003). (b) cramping or squeezing. . 2000). . . . . . . . . . 4. . . . . . . . . 4. 1998). .. . . . . . . 1993). . . 1998). . . . 1977. 1999. and (c) shocking or shooting (Sherman. . below) plays an integral role in phantom limb perception. . . . . . the anterior cingulate cortex (ACC). . . . . . .. however. . years or even decades after limb loss before they fade completely. 1994). Brena and Sammons. . . . . . . . . . . . . .1. 2. . . . . . below). . 1979). . Machin and Williams. . they may emerge years or even decades after limb loss. . Through the mirror neuron system. . . . . although they have also been reported following removal of the breast (Aglioti et al. . 4. . . . . . . . . . . . . . . .. . . . . .4. . . . . . . . . .2. . . . . . . . . . . . . . . .. . Bressler et al. . . . .. or spinal cord injury (Bors. . . . . . insular regions. . . . and phantom sensations may be perceived for anything from a few days to weeks. . . . . . . . . . . . . . . . . . . . . action understanding and empathy. . . . . . . . . . The pain matrix refers to the pain-related network that primarily includes the secondary somatosensory cortex (SII). sensations of urination or erection following penis removal (Fisher.4. . . . . . . . . with pain usually characterised as either (a) burning. 2001..3. . . Biley. . .. Sherman. 1998). . . . . . . . tingling. . . 222 223 223 224 224 225 225 225 226 226 227 228 228 228 1. . . Melzack and Bromage. . . . . partially flexed at the elbow with the forearm pronated). . Conclusions . . penis (Fisher. 1998). . 1951. . The neuromatrix. . . . 2002. 4. . . . . 4. . . . . . . . . . . The phantom limb is generally described as adopting a “habitual” position and posture (e. Brugger et al.5. months. and cross-callosal and ipsilateral projections in phantom limb phenomena – the perception of a “normal”. . . Braun et al. .. . . . 2006. 1991). . and are thus potentially more likely to have a bodily experience that resembles that observed in others. . . . 1997). . 1999). 1998). . . . . . . . . . . . . . . and the movement-related areas such as the cerebellum and supplementary motor area (Singer et al. 1973). . . . . . . menstrual pain following hysterectomy or phantom pain that resembles presurgical pain) and tend to be characterised by functional sensations. . . . . . . . Experimentally induced sensory and motor coupling between the phantom and real 5. . . 2000. . . . Mirror neurons and empathy for pain in amputees . . Phantom pain is experienced by up to 80% of amputees (Kooijman et al. .. .4. . . . . . 1994. . . 2000. . . . . 4. . . The present paper reviews the literature on the perceived “body space” of phantom limbs. . . . . their interaction with prosthetic devices. . . nerve avulsion (Melzack. . . . . the phantom limb may be perceived to be stuck in a fixed position (Devor. .

1988). and that it is able to detect which muscle would be involved in the actual execution of the observed movement. 2002. 1983). Ettlin et al. 1973. The subject is instructed to fixate on the rubber limb. subjects report that they feel the touch on the rubber hand. feeling “like the toes are all turned down under the bottom of the foot”. Ramachandran and Hirstein. 1973. ranges. the subject's arm is hidden from view – either behind a screen or under the table – and a life-sized rubber model of the same arm is placed on the table in front of them. Henderson and Smyth (1948) also described a patient's anatomically unrealistic perception of phantom fingers that were “grossly twisted and interwoven”. and many reported that parts of the amputated limb were missing (61%). 1974). 1999. and another where the phantom arm was stuck behind the individual's back preventing him from sleeping on his back.. 2003). and is likely to be associated with the experience or anticipation of pain (Willoch et al. Other studies have described illusory limb displacement. Hunter et al. These findings suggest that while central representations of the body are indeed able to code for anatomically impossible limb positions. Empirical research on the mechanisms underlying distorted or disfigured phantom limbs is. Patients with spinal cord injury may also perceive their phantom sensations.. as if their arm has embodied the fake rubber limb (Botvinick and Cohen. Rather. as occupying anatomically unrealistic and unnatural postures. when the subjects own hidden finger was moved slightly. 1973). 1941).. (1980) reported that phantom limb illusions were only perceived by patients who retained consciousness during the spinal cord injury. sprouting). ephapses. 1973). but usually limited. André et al. 2. Halligan. 1973). the forearm remaining stationary while the hand continues to move downwards (Lackner and Taublieb. “it is being bent backwards”. that each digit was somehow twisted so that “each toe pointed in a different way” (Conomy. Ettlin et al. 1 . 1948). one where the phantom arm was extended from the shoulder at a 90° angle and the individual felt that he must walk through doorways sideways. or that the legs were “twisted”. for example. Fraser (2002) found that 63% of her sample felt that their limb remained in a fixed position when they moved other body parts. Wright et al. (2005) found that the mirror neuron system (see 4. or “blown up” (Bors. Telescoping of the phantom limb Phantom limbs may be perceived to be continuous and intact resembling a normal limb. however. (2001) suggest that phantom limbs may become seemingly deformed or dysmorphic as a result of ectopic activity in the stump (neuromas. or extending the nose with the Pinocchio effect (Lackner. Phantom limb posture in some patients with spinal cord injury also mirrors the position in which the limb was last seen during the spinal cord trauma (Bors. and was unlikely to occupy random or unusual positions. For example. these memories do not determine the constant or continuing posture of the phantom (Henderson and Smyth. 1951. including the (non-painful) perception that the “the arm is being broken”. including the hand being bent back towards the dorsal surface of the forearm (Craske. 2005). In one of the few studies to address phantom limb proprioception specifically. a small percentage report that they can move the phantom limb through anatomically impossible ranges or amplitudes (Price. the actual perception of such limb positions rarely occurs. for example. In normals. The perceived position of a limb in nonamputees (Gross and Melzack. however. 1951.4. had shrunken or were shortened (28%). Ehrsson et al. Some amputees report the experience of a general awareness of the presence of a phantom limb that is without positional or sensory qualities (André et al.1. Generally. distorted or disfigured. they experience phantom limbs that do not remain in the same position as that occupied by the limb at the time of anaesthesia. particularly when the latter is out of view (Fraser. 1998. “crossed”. Melzack and Bromage. Conomy. 2004). below) reacts to both biomechanically possible and impossible movements. 2001. Fraser. the limb was more likely to occupy one of a common repertoire of positions. 2000). the thalamus or the dorsal column nuclei. 1998. for example. Riddoch. and phantom limbs in paraplegics (Bors.. While memories of the limb immediately prior to amputation may endure in the phantom (Katz and Melzack. lacking. or of remapping in the somatosensory cortex. Conomy. Fraser (2002) found that the majority of amputees in her study indicated that their phantom limb took the general shape or form of the limb prior to amputation. while two small paint brushes are used to simultaneously stroke the rubber hand and the subject's hidden hand.2. particularly in the lower limbs.B RA I N RE SE A R CH RE V I EW S 54 ( 20 0 7 ) 2 1 9–2 3 2 221 dissociate from the stump. see Armel and Ramachandran's (2003) study in which the movement of a rubber finger (using the rubber limb paradigm1) into “painful” positions. Hill (1999) noted that distorted posture in a phantom limb may be more common following traumatic limb loss in which the limb had been distorted by the accident. or epidural block for lower limb surgery (Bromage and Melzack. Within minutes. some reported varying degrees of hyperextension or hyperflexion of the joint. when Craske (1977) vibrated and passively stretched the biceps brachii of his subjects. or telescoped so that the proximal In the rubber limb paradigm. amputees report that they can move their phantom limb through normal. Furthermore. the perception of anatomically impossible limb positions can be induced using muscle and tendon vibration. 2. not their hidden hand. 1980). the phantom limb takes on a posture that is abnormal... Melzack (1992) described two cases of phantom limbs in abnormal postures. 1974. 2002. 1999) and (normal) patients under anaesthesia (Bromage and Melzack. that would have been assumed by the limb outside of anaesthesia. Distorted perception of the phantom limb In rare instances. With respect to phantom limbs. 1976). or protruding down through a solid surface (Romani et al. or had become magnified (8%). in slight flexion. 1978). may also become dissociated from the real position of that limb when it is occluded from sight. 1951). Similarly. 1990. 1997). Alternatively. Hill. while patients are under brachial plexus nerve block for upper limb surgery (Melzack and Bromage. even in a phantom limb. or that “my hand is going through my shoulder”.. induced responses associated with pain. Romani et al. 1977).

. comments such as Melzack's (1990) report that “sometimes the limb is slowly telescoped into the stump” are ambiguous and could be interpreted either way.. activate areas remote from the limb representation (Flor et al.. 2006). 3.. and the endpoint of the amputated limb is perceived to be more distal than it actually is (McDonnell et al. discussion about telescoping of a phantom limb will refer to a phantom limb that has shortened over time with the distal portion gradually perceived to be closer to the stump. 2003. (1999) who found that upper limb amputees who use myoelectric prostheses do not show cortical reorganisation.. Yamamoto and Kitazawa.e. 2001. 1963). for example. the remaining parts of the limb appear to become magnified because they maintain their original size.. The upper limbs. Prosthesis use and embodiment Amputees commonly feel that their phantom limb embodies and thus becomes one with the prosthesis. While using a prosthesis. the more weakly represented regions of the limb may “fade from consciousness” while the distal portion of the limb becomes attached to the stump. as the phantom continues to provide sensory feedback to the area that previously represented the amputated limb. 2001). compared with those who use either a cosmetic prosthesis or no prosthesis. brachial plexus avulsion or in cases with preexisting peripheral nerve injury (Katz. and this disparity manifests in more rapid telescoping of the lower compared with upper limbs (Henderson and Smyth. cortical reorganisation or changes in stump sensitivity (Hill. Yamamoto et al. or whether it refers to active movement of the components of the phantom limb in accordance with the concept of actively retracting a telescope.. and such cortical reorganisation is strongly correlated with phantom pain. Ramachandran and Hirstein. compared with the lower limbs. and this process generally begins within the first few weeks post-amputation (Carlen et al. 1998). Also. Over time. 2006. it tends to be related to increased levels of phantom pain (Flor et al.. are more diffusely represented throughout the cortex as they are integral in fine unimanual and bimanual motor schemata. Katz (1992) suggested that the perceived distance between the phantom and the stump may be a function of the distance separating their respective representations in the somatosensory homunculus. In this paper.222 B RA I N R E SE A R CH RE V I EW S 54 ( 20 0 7 ) 2 1 9–2 3 2 portion of the limb is missing or has shrunken with just the more distal portion floating near. The term appears to have been used more often to refer to the former definition. 1999). Consequently. some authors have used the term either with the latter meaning or without clarifying the intended meaning.g. Flor et al. the definition of this phenomenon is lacking and it is ambiguous whether telescoping refers to the displacement of relative position of the components of the phantom limb over time. Essentially. for example. 2006. on the other hand. 2001). Solonen. or may change from moment to moment or day to day. The phantom limb is perceived to be telescoped in between 49 and 63% of cases. Yamamoto and Kitazawa. rake or screwdriver – in monkeys (Iriki et al. 1998). 1998). The distal portion of the phantom limb may subsequently disappear (Shukla et al. 1978). 1948. and they often confound their phantom limb with the prosthesis (André et al. 1996) and neurologically intact humans (Maravita et al.. Ramachandran and Hirstein. Jones. or “within” the stump (Flor et al. and not progressively. Murray. or remain “dangling” from the stump in about 50% of cases (Ramachandran and Hirstein. When a phantom limb becomes telescoped it usually does so in a “diffuse process. 2002). Weiss and Fishman.. 2005). Amputees who use a prosthesis extensively retain a different functional representation of the amputated limb compared to non-prosthesis users. 1992. 2006). the amputee's bodily experience is generally one of being whole. In some amputees telescoping may take place over a number of years. Phantom limbs apparently do not telescope in patients with spinal cord transection. and preserve the biological representation of the amputated limb. p.. nontelescoped) phantom limb might reverse maladaptive cortical changes. 2006). movement of a completely telescoped phantom arm corresponds to activity in the cortical region that represents the shoulder. While telescoping was originally thought to be a sign of adaptive plasticity.. however. affecting most of the limb simultaneously. such that the prosthesis is felt . Telescoped phantoms. in turn. 1998). 1982).. 1989). that is. Such embodiment may be associated with a twoway interaction between prosthesis use and phantom limb perception. the pre-existing representation of the amputated limb in the body schema – which also likely provides the template for phantom limb perception – may come to provide a valuable neural template for prosthesis use. from the stump towards the periphery” (Henderson and Smyth. This hypothesis is supported by Lotze et al. and this contributes to the relative facility and rapidity with which amputees can learn to control a prosthesis (André et al. Initially. 91). While telescoping is commonly reported in the literature. perhaps as a result of changes in receptive fields. Repeated use of the prosthetic limb may. which were congenitally missing both forearms and hands. reinforce the representation of the missing limb and thus reinforce the perception of phantom limb sensations. 1948. 1988. it appears that the prosthesis is incorporated into the body schema and becomes part of a coherent internal model of the body (Murray and Fox. It is widely accepted that phantom limbs become telescoped because the distal portion of any limb is more strongly represented in the cortex relative to the more proximal regions of the limb (Ramachandran and Hirstein. 1962. 2004). the bimodal neurons in the posterior parietal cortex that code peripersonal space appear to be recoded to represent space accessible to a handheld tool – e. Telescoping of the phantom appears to be associated with remapping of the distal portion of the phantom limb onto nearby regions of the cortex. (2006) proposes that the continued perception of an extended (i. These findings suggest that prostheses become embodied in the same way that a habitually employed tool does (Lewis. perhaps because there are conflicting representations of the deafferented body part according to visual and somatosensory afferents. see Poeck's (1964) case description of a child whose phantoms “gradually withdrew within the stump” when she approached the wall with her arms. 2001.. partially telescoped phantoms to activity in the region of the arm and non-telescoped phantoms to activity in the hand region (Flor et al. attached to. or phantom limb pain.

updating or evoking templates of the body schema can result in various perceptual aberrations. Recently. evolutionary design and function of the human body and a repertoire of motor templates (e. together with information about movements of the eyes.. The body schema consists of innate representations of the biological.. scratching the corresponding locus on the embodied prosthesis can relieve the itch (Giummarra et al. Future research. There is limited research to inform us of whether and how phantom sensations and phantom pain differ according to the use of a prosthesis. in somaesthetic aura (Kew et al. perceived absence of body parts in asomatognosia (Arzy et al.. 2005. or fused with. It also provides a neural platform for understanding and interacting with others throughout life (Brugger et al. neck.B RA I N RE SE A R CH RE V I EW S 54 ( 20 0 7 ) 2 1 9–2 3 2 223 to be a “part of them” (Murray. Rinehart et al. disembodiment and autoscopic phenomena in out-of-body experiences (Blanke. dynamic representation of the spatial and biomechanical properties of one's body. 2005) – together with inputs and outputs to various other cortical and subcortical regions. 2003. in press). Sakagami et al. 2002). Gow et al. the body schema. 2004). 2000. 2006). and they phenomenologically have a normal and intact body (André et al. These body part locations are monitored by visual input and defined by the boundaries of body parts and their proximity to other body parts and objects.. 1999).1. Brugger et al. 2003) and the temporo-parietal junction (Blanke and Arzy. Mazzoni et al. vestibular. is the internal. and Bonnier.. and (d) movement systems and efference copy—that is. 1996). 2004a). the phantom limb (Gow et al. Finally. had previously considered the concept of an organised spatial representation of the body with respect to the world. These studies could also investigate whether amputees' perceptions of prosthesis embodiment vary according to voluntary versus passive movement of the prosthesis. or Alice in Wonderland illusions... Phantom limb perception . Studies should examine prosthesis embodiment within a body schema framework. in 1905. when the phantom limb is itchy. Second. Lewis. suggesting that the prosthesis is represented in the contralateral somatosensory cortex in the same region as the phantom limb. Ramachandran. The body schema is fundamentally generated in the parietal cortex – particularly the multimodal cells in the superior parietal lobule (Lewis.. 2006. in brief. 2004. (c) visual system.g. 4. 1998) to explore prosthesis embodiment. Melzack. The relationship between phantom limb sensations. hand–mouth coordination) designed to promote survival in infancy (Gallagher et al. trunk and limbs (Stein. however. Our understanding of tool embodiment and extended physiological proprioception – that is. (b) vestibular system.. 2004). in 1890.. 2006b). The body schema Many authors have highlighted the distinction between the concepts of body image and the body schema (Gallagher. 2006b.. Blanke and Arzy. parieto-insular region (Ventre-Dominey et al. 2002). Miyazawa et al. the neural copy of a movement command that is sent to the parietal cortex to be mapped onto the body schema to generate expected sensory outcomes (McGonigle et al. the plastic and dynamic representation of the body that is constantly being modified and updated. Schiff and Pulver.. The cortical origins of phantom limb phenomena 4. 1998). and is derived from multiple sensory and motor inputs that interact with motor systems in the generation of actions. 2006). The prosthesis may be described as being embodied by.. and that the visual feedback that parts of the prosthesis are being stimulated is centrally processed and thus perceived as if the phantom limb is being stimulated. Schwoebel and Coslett (2005) highlighted that there are at least three distinct types of body representations. Maravita.. Investigation of the interaction between phantom sensations and a prosthesis may provide valuable insight into the interaction between internal constructs of the body and the embodiment of a tool as a functional extension of the body. coding egocentric frames of reference – based on the convergence of retinal. it is unknown whether prosthesis embodiment varies between upper limb and lower limb amputees. 1995b. Damage within the various sensory systems that contribute to maintaining. and to the premotor cortex in preparation for rapid corrective adjustment of movements when errors between the expected and actual sensory outcomes are detected (Helmholtz. Halligan et al. 1998)... The main systems that contribute to the properties of the body schema include: (a) proprioceptive and somatosensory systems. somaesthetic.. Additionally. had introduced the term schema to this concept. Maravita (2006). prosthesis use and embodiment is still not well understood. Neurons in the parietal cortex are involved in mapping “real space” relative to the body – that is. Ongoing bodily experience is brought about by the “body schema”.. First. and magnification and shrinking of various body parts. 1996. or the type of prosthesis used. including denial of limb ownership in somatoparaphrenia (Aglioti et al. Giummarra et al. The perceived convergence between phantom limb and prosthesis helps to coordinate movement and maintain fluid and natural motor control over the prosthesis (MacLachlan et al. and auditory signals relating to the self and space. proprioceptive. 1989).. whose missing limbs biologically do (upper limb) and do not (lower limb) habitually use tools. the body image is the lexical–semantic representation of the body including the names and functions of body parts and relations between body parts and external objects. the body structural description is a topological map of body part locations. 2001. 2004.. For some amputees. the extension of the body's “area of influence” that also extends one's innate sense of proprioception (Gow et al. It is the body schema that is implicated in providing a template for the perception of phantom limbs following amputation or deafferentation. and determine whether amputees with phantom sensations (particularly those with complex sensations) differ in adaptation to prosthesis use compared with other amputees without phantom sensations. 2004) – could benefit greatly from research on prosthesis embodiment. Experimental research could adopt the rubber limb paradigm (Botvinick and Cohen. 1995. 1997. 1992). 1997). 2004). and Holmes and Spence (2006) highlighted that Münk.. Head and Holmes (1911–1912) are often credited for coining the term “body schema”. 2006. 1986. supernumerary phantom limbs (Brugger.

immune and opioid systems (Melzack.g. emotional. 1961. further proposes that cramping pain and shooting phantom pain may arise from similar origins from spontaneous activity associated with intention to move muscles. amputation and deafferentation are commonly associated with cortical reorganisation. 1999). BIID (Bayne and Levy. 2006a. attempts to answer the telephone or fend off a blow with the missing limb) can trigger phantom sensations and pain (Giummarra et al. 1997.. 2005)) perceive a conflict in the body–self neuromatrix such that their physical body does not converge with their phenomenological experience? Would people with BIID perceive a phantom limb following amputation and if so. Weinstein and Sersen. These sensory systems are further accounted for in Melzack's (1989. such as electrical impulses (Flor et al. or vibration. Lotze et al....g. there is also evidence that cramping phantom pain is related to muscle tension in the residual limb (Sherman. including cytokines as well as the endocrine. unlike sources of pain triggered by injury or lesion. why do some amputees experience phantom pain when they observe another person in pain (i. why do some amputees . while for others phantom sensations diminish. These inputs thus generate the body–self neuromatrix primarily through the somatosensory. why do some congenital and childhood amputees feel phantom sensations as adults.. some only painless phantom sensations. 1996) neuromatrix theory of phantom limb perception.g. vestibular) in relation to phantom limb sensations? Third. 1999. autonomic.g.. how do internal representations of the self (that is.. The neuromatrix Phantom phenomena problematic for current theory Melzack's (1989. absence of all four limbs as reported by Brugger et al. 4. would this differ from other amputees without BIID? Seventh. however. why do phantom sensations continue to be perceived by some amputees. however. For example. 2005. a reduction in blood flow in the stump) (Sherman. 2004). 1996) neuromatrix theory of phantom limb perception extends theories of the body schema. Transcutaneous Electrical Nerve Stimulation (TENS). 2001). how and why do phantom limbs telescope actively? Fourth. 1995) and tactile or somatosensory inputs from the stump and/or remaining portion of the amputated limb (Melzack et al. or other sites) and environmental (e.. from the residual limb/stump. Melzack (1996) There are many aspects of phantom limb experience that current theory of phantom limb phenomena do not explain.. by providing continued sensory input via sensory discrimination training (the discrimination between the frequency and/or location of sensory inputs. telescope or disappear over time? Fifth.. changes in barometric pressure) experiences. (c) phasic and tonic cognitive and emotional inputs from other areas of the brain. why do people who desire amputation of a healthy body part (Body Integrity Identity Disorder.. may preserve a “normal”. updating and evoking internal representations of the body are responsible for maintaining templates of the phantom limb. Price. 1964). peripheral (e.. including stimulation of the vestibular system (André et al.3. motor.. below] (Ramachandran et al. 2000)? Sixth. A major limitation of the neuromatrix theory. Saadah and Melzack. as well as by spontaneous ectopic activity at central. (b) visual and other sensory inputs that influence the cognitive interpretation of the situation.. which. 1999). however. or which cannot be tested under current models. and some only phantom pain (as amputees in our studies have reported)? Second. why do some amputees with similar circumstances of limb loss experience no phantom sensations. 2001.. Continued sensory input through stimulation and functional sensitivity of the residual limb and stump.. 1994. The range of triggers that evoke or modify phantom sensations and pain provide evidence for the complex involvement of the various systems likely involved in bodily awareness and perception in phantom limb phenomena. non-painful representation of the amputated limb in the body schema (Bittar et al. limbic and thalamocortical systems.2.e. regardless of the correspondence or lack thereof between the ideal model and the actual body (Berlucchi and Aglioti. visual illusions of phantom limb embodiment using the mirror box [also see 4. 1999). The complex nature of phantom limbs suggests that the same systems responsible for maintaining. even when they have never experienced a limb (e.g. 4. “forgetting” the limb loss and the performance of automatic motor schemas with the phantom limb (e. These inputs primarily include (a) somatosensory inputs (cutaneous.e. 1992. visceral and other somatic receptors). (d) intrinsic neural inhibitory modulation inherent in all brain function. templates of the body schema) interact with specific sensory afferents (e. it is possibly too broad and difficult to be tested empirically. The neuromatrix theory provides a valuable model for the perception of phantom phenomena..224 B RA I N R E SE A R CH RE V I EW S 54 ( 20 0 7 ) 2 1 9–2 3 2 following congenital limb deficiency suggests that the brain may be genetically predisposed to represent a prototypical human body.5.. proprioceptive. is that while it can broadly account for the various aspects of phantom phenomena. 1992. and cognitive factors). visual. “empathic pain”). The innate body schema is then retained and modified throughout life experience (Melzack et al. 1990. and (e) the activity of the body's stress regulation systems. Ramachandran and Hirstein. and does this relate to how empathic one is? Finally. affective. Wilkins et al. 1997. 1994). 1990. and particularly phantom pain. Additionally.. 1997). Hill. First.. 1976.g. 2001)). for example. 1994). 1998). although there is evidence that burning pain is associated with vascular mechanisms (i. and spontaneous bursts of activity that produce output patterns that resemble activity associated with pain and thus lead to the conscious experience of phantom pain (Melzack.2. First. particularly with respect to painless phantom sensations. Le Chapelain et al. and proposes that conscious awareness and perception of the body and self are primarily generated within the brain via patterns of activity (or neurosignatures) that can be triggered or modulated by various perceptual inputs. 2005. according to Melzack (1996) high-frequency bursts of activity associated with deafferentation may be interpreted as hot or burning pain. While multiple sensory inputs are integrated to create the body–self neuromatrix. Weinstein et al. is often triggered by supraspinal (e. spinal or peripheral levels. myoelectric prosthesis use.

Brugger et al. 2003). 2006). similar mirror neuron systems were identified in humans and were found to result not only in action simulation in the motor cortex. and a group of control subjects.. 2006. (b) whether they perceive a “moveable” phantom limb compared with those who perceive a phantom limb that is “paralysed”. 2006. Morrison et al. Price. 2004). or the human experimenter. The level of activity in the ACC is strongly . research should examine the relationship between action understanding and phantom limb perception and movement. disgust (Wicker et al. in the secondary somatosensory cortex. at least in part. While these mechanisms may explain phantom limb perception following congenital limb absence (Brugger. but there was no evidence of such activation in the non-synaesthetic group. Rizzolatti et al. 2002). Singer et al. in order to understand the actions of others one must match the action against one's own motor system. This process may evoke phantom limb perception. or even the perceived embodiment of another's movements without apparent phantom limb perception (Frank. but also when it observed another monkey.. For example. Mirror neurons and empathy for pain in amputees In addition to action understanding. beyond the body schema. We propose that the mirror neuron systems for touch and pain may be involved in maintaining the representation of a phantom limb.. The latter include mirror neuron systems – particularly those involving movement (fronto-parietal mirror system).1. 2006).4. For example. Essentially. Brugger et al. Phantom limb perception in cases of congenital limb loss supports the possible role of mirror neurons in the retention of a representation of the missing limb within the body schema (Brugger. 4. or neither phantom pain or sensations. but also sub-threshold activity in muscles engaged in the observed action (Fadiga et al. non-painful phantom sensations are likely to also be related to other complex central processes involved in bodily perception... In particular. and much work is required to clarify its role in congenital and acquired limb loss. also process feelings of “emotional pain”. were identified by Keysers et al. 2005).. The role of mirror neurons in phantom limb perception is only just beginning to receive attention. we propose that they are also likely to be central to the perception of phantom limbs following acquired limb loss. but we also automatically interpret the experience throughout the same cortical networks that mediate personal experience of pain. we not only consciously comprehend that the other is in pain. perform the same goal-directed motor action. (1988) in the ventral premotor cortex in the macaque monkey. 2006). 2006. 2005. do amputees differ with respect to action perception and embodiment of another's motor system (i. There is strong evidence that the AIC and the ACC are involved in both the personal experience of pain and its empathic experience (Jackson et al. and in some cases triggering phantom pain. in which case activation of mirror neurons may reinforce the representation of the phantom limb.. Funk et al. 1988). mirror system activity) according to (a) whether they perceive phantom pain. Amputees may experience “somatic” or “postural” empathy during mirror neuron activity when observing others using their limbs (Price. For example. when she observed touch.B RA I N RE SE A R CH RE V I EW S 54 ( 20 0 7 ) 2 1 9–2 3 2 225 perceive sensations in the phantom limb that mirrors those in the intact limb? While the neuromatrix theory may be a very adequate model for the complexity of phantom limb pain.. Rizzolatti et al. 2000). 2003) or frustration (Abler et al. Fadiga et al. 2004). Mirror neurons were initially identified by Rizzolatti et al.. mirror neurons are involved in other domains of perception and perceptual understanding. An area in the left premotor cortex was activated in C to a greater extent than in the nonsynaesthetic group and the anterior insula cortex (AIC) was bilaterally activated in C. When observing another person in pain.. 2004. (2004) in people who were touched or observed another person or non-biological object being touched. 2005): using functional Magnetic Resonance Imaging (fMRI).. Mirror neurons Mirror neurons and action understanding Mirror neurons have an integral role in the observation and understanding of goal-directed actions between an agent and an object (Gallese. suggests that phantom limb perception may have evolved. Morrison et al. 2000. compared with controls. Activations in the somatosensory cortex were significantly higher in C. Mirror systems for touch. including emotion (Ruby and Decety. and pain (Jackson et al.. Mirror neuron activity may thus both require and reinforce the representation of the body and its functions within the body schema—even if a limb has been amputated.4.... Following the discovery of mirror neurons in monkeys. Singer et al. from the habitual observation of other people moving their limbs. and touch (somatosensory and posterior parietal cortices) – and mechanisms of cross-referencing between the phantom limb and intact limb (Giummarra et al. mirror neurons are likely to be responsible for the instinct to cringe or shudder when observing another person in pain (Miller. 1986). These neurons fired not only while the monkey executed a goal-directed action. pain (anterior cingulate cortex and anterior insula). 2000. associated with processing physical pain.. such as social rejection (Eisenberger et al. 2004. These same areas. 2005. 2005). (2005) investigated the neural networks involved in the perception of touch in their synaesthete subject (C). Price.. 2005. 2001. 4. 2006a). Synaesthesia for touch has also been described in a single case study (Blakemore et al. touch (Keysers et al. and the continued activation of the innate body schema.. 2004. The evidence suggests that in C the mirror system for touch is disinhibited and involves activity that exceeds the threshold for conscious tactile perception..4. Essentially. the type of prosthesis used and the perceived embodiment of the prosthesis.2. 4. but who perceived vivid phantom limbs (Brugger et al. the case of a 44-year-old amputee who was born missing all four limbs.. and (c) whether they use a prosthesis.e. Future research.. phantom sensations.. Amputees with phantom sensations may provide evidence that efficient action understanding requires a mirror neuron system that retains a representation of an intact and functioning body. 2005). activity in mirror neuron systems enables the observer to embody the other's motivational system. 2004). 2004). Blakemore et al. simulating cortical activity in the same neurons that are responsible for performing that action (Buccino et al.

. McGonigle et al. 2005). such as childbirth or third-degree burn victims. and a desire to exercise the limb. posture/form. itching and the ability to ease itch. These patients described sensations in the phantom that mirrored the RA symptoms perceived in the intact leg. 2006a). Empathically triggered pain is likely to be present in other populations who have experienced recent trauma. the sensory and affective qualities of the other's pain would be embodied and mapped onto the observer's body schema (Schwoebel et al. while the face – which is localised close to the upper limbs in the somatosensory homunculus – plays an emotionally expressive role. The present review will focus on the perception of mirrored sensation and movement from the intact to the phantom limb. resulting in emergency caesarean section delivery. which has been pre- sensitised. Giummarra et al. touch and exteroception. may be disinhibited.226 B RA I N R E SE A R CH RE V I EW S 54 ( 20 0 7 ) 2 1 9–2 3 2 correlated with an observer's ratings of the intensity of another's pain (Jackson et al. 2005. or understanding another's intentions.. 4. The ACC receives projections from the superior temporal areas. In amputees. 1995a). which seems to be a major role for mirror neurons. 2005). to pain.. see reviews by Flor. (2003) examined the phantom sensations in amputees who had been diagnosed with rheumatoid arthritis (RA) prior to amputation. clawing of the toes. (2006a) reported that 13% of their patients experienced phantom sensations that mirrored sensations from the intact limb. the role of transcallosal cross-referencing between the phantom limb and the intact limb has received somewhat less attention. Flor and Birbaumer. there is nevertheless some limited evidence for mirrorneuron-like activity in the lower limbs (Buccino et al. and mirror movements. 1998. temperature. while there may be mirror neuron activity in the pain matrix of non-amputees and other healthy people without actual perception of pain. 2005). or heightened activity in the limbic system due to emotional distress. we propose that it is also probable that empathically perceived pain in the phantom limb involves mirror system activity within the “pain matrix”. 2000). Through mirror system activity..... van der Velde and Everaerd. 2006a). discomfort or pain in the phantom by scratching or rubbing the contralateral limb. which maps the observed pain onto the body schema (Giummarra et al. or (c) whether the other's pain is generalised or specific to the amputee's past pain experiences. who experiences shooting pain from the groin that radiates down the legs when she is told of another person's traumatic experience (unpublished case). which play a role in semantic visual processing. 2002. leading to an empathic motivational state of readiness for action (Avenanti et al. Giummarra et al. These findings suggest that there is empathic inference of the sensory qualities of another's pain. by trauma. swelling in the ankle and/or knee. Cortical reorganisation of the lower limb onto the homuncular-adjacent genitals is also associated with heightened phantom pain (Flor. 2004). 2003. for example (a) aspects of traumatic experience. Haigh et al. (2006a) have described eight cases who reported that their phantom pain is triggered by observing. Cross referencing between the intact limb and the phantom limb 4. 2002. or inferring that another person is in pain. pain or injury. 2007). 2001). An important area of research includes investigating the role of mirror systems for pain in amputees who empathically feel phantom pain when observing another person in pain. This is a novel hypothesis and requires further research. Future research. and (b) the urogenital system through the increased reflex urge to empty the bladder or bowels for more efficient evasion of danger. Mirrored sensation and movement from the intact limb to the phantom limb While remapping and cortical reorganisation have received considerable attention in the phantom pain literature (for example.5.. and is probably involved in the affective components of the “pain matrix” during empathy for pain. 2002). (b) whether sensitisation to pain is generalised or specific to the amputated limb. However. automatic embodiment of the observer's motor system.1. thinking about. or observed to be experienced by another person (Avenanti et al. It is not clear whether amputees who have observed injuries incurred by another person are more likely to experience phantom pain in general. and there are similar motor action responses to a painful stimulus when it is either personally experienced. using the methodology of past studies that have examined mirror system activity for observed and perceived pain (Singer et al. particularly lower limb amputees.. This hyperalgesic man reported that he felt pain that was immediate and intense. 2002). In summary. It is of significance that only lower limb amputees have reported “mirror pain” (Giummarra et al. Flor et al. and in one patient with a supernumerary “alien” limb subsequent to callosal lesion during active movement only (Hari et al.. consider the case of a woman who experienced a long painful childbirth. The neuromatrix theory may suggest that this is due to triggering pain memories.5. 2004). leading to empathically perceived pain when another is in pain. for example. when he saw his wife hurt. The ACC has extensive outputs to the premotor and motor areas (Morrison et al. While the possible mirror system activity for the lower limbs is not well understood. including: ache. Experimental research should examine the factors that predispose amputees towards experiencing empathically triggered pain... The experience of mirror pain only in the lower limb suggests heightened activity of the autonomic nervous system. 2006). Mirror movements have been described in an upper limb amputee during passive and active movement of the intact hand (Halligan et al. and empathic activity in the autonomic nervous system's “flight” mechanism during empathy for pain. and appeared to be qualitatively similar to his own hypersensitivity to touch... mirrored pain may be perceived to be localised in the stump and/or phantom limb. it appears that the mirror neuron system in some people. which is more likely to involve (a) the lower limbs in readiness for flight from danger. including stiffness and limited ability to move the joints freely. The genitals are highly involved in emotional perceptual processes (Both et al. Bradshaw and Mattingley (2001) reported the first clinical case of feeling mirrored pain when observing another person in pain. For example. as a result of embodiment of the other's fight or flight dilemma. Bach and Tipper. despite the fact that they do not have a repertoire of meaningful or operant activities that could be involved in communicating. neuropathy or disability. Franz and Ramachandran (1998) provide evidence that the phantom hand remains .

Perceived illusions in the phantom limb are reported to be so realistic that amputees claim they can see and feel their phantom limb moving as if it was under their volitional control (Ramachandran.. a mirror is positioned vertically in the centre of a box.. During mirror therapy. these patients perceived “referred” sensation as pressure in the hidden. 2005). which included pain. Hamzei et al. The peripheral pathways involved in bimanual motor coupling endure in amputees. 2003). Following amputation or deafferentation. input which. Acerra and Moseley (2005) found that pain and paraesthesia could be induced in the affected limb of patients with CPRS (type 1) when corresponding areas of the unaffected limb were passively stimulated and observed in the mirror as if superimposed onto the affected limb. particularly during high-threshold somatosensory or motor afferents. Centrally. 1996) and lower limb (Brodie et al. and recruit local inhibitory interneurons during unimanual or antiphase movements (Carson. 1995). during moderately intense tactile stimulation. Kew et al.2. already modified by disease or pain. 2003)... 1998). either at the spinal or supraspinal level. 1996. 2006). together with evidence that the motor cortex ipsilateral to the deafferented limb undergoes cortical reorganisation (Schwenkreis et al. Ramachandran. Mirror box therapy has now been successfully used in both upper (Oakley et al. cross-referencing of the phantom limb to the intact. 1995). These findings suggest that pain – at least pain associated with amputation and CRPS (type 1) – may be related to (a) mapping observed stimulation onto the body schema. (1995) reported that upper limb amputees who used this technique feel their phantom arm move when they observe their own arm. Cortical reorganisation following deafferentation is thought to be based on various mechanisms. 2003.. 1996). (c) enhanced effects when mirror feedback was used to give the illusion of touching the phantom hand. such that movement of a phantom limb corresponds to the activation of motor neurons that once served the missing hand. and in some cases apparently causing the phantom limb to telescope (Ramachandran et al. Sanes et al. 1993. as long as the amputee can conduct voluntary movements in the phantom. (b) the referral of touch.. 2002. 2004b) amputees to assist in voluntary movement of the phantom limb and alleviation of phantom pain. inducing somatosensory and/or motor coupling between the phantom and real limb. including unmasking of preexisting synapses by removal of local inhibition. Cross-referencing mechanisms. 2001). 2005. Ramachandran et al. using the additional sense of vision with mirror feedback (Ramachandran. Ramachandran and Rogers-Ramachandran (1996) examined intermanual referral in 10 upper limb amputees using the mirror box paradigm. Ramachandran et al. move in the mirror. and axonal sprouting with the formation of new synapses (Schwenkreis et al. regardless of the type of stimulation of the good hand.B RA I N RE SE A R CH RE V I EW S 54 ( 20 0 7 ) 2 1 9–2 3 2 227 spatially coupled with the intact hand following amputation. Animal studies have shown that deafferentation is followed by rapid cortical reorganisation contralateral to the deafferented limb (Donoghue et al. clearly continue to function even after deafferentation... type 1) to reduce pain associated with movement of the affected limb (McCabe et al. Experimentally induced sensory and motor coupling between the phantom and real limb Illusions of touch and movement can be evoked in a phantom limb. and (b) a mismatch or reafference (efference copy) between intention to move the phantom limb and the lack of proprioceptive feedback from the deafferented limb.. the amputee observes the reflection of his normal limb moving in the mirror in order to induce the visual illusion that he has two intact limbs and that the intact limb is superimposed onto the felt position of his phantom limb (Ramachandran..... and the intact limb is placed so that the amputee can see its reflection. however. Sathian (2000) also reported the intermanual referral of sensation to a hand rendered anaesthetic by stroke or surgery using the mirror box paradigm. Ziemann et al. Sathian (2000) proposed that following decreased somatosensory input from the anaesthetic hand. the perception of “referred” or mirrored sensation from the intact to the phantom limb may be related to altered inhibitory mechanisms in these ipsilateral pathways. Amputees who use a functional prosthesis (Cruz et al. highlighting that cortical reorganisation may be use-dependent.. anaesthetic hand. These findings have been replicated in human studies (BrasilNeto et al.5. Creating the illusion of convergence between sensory systems . and enhanced interhemispheric interaction of the nondominant hemisphere onto the dominant hemisphere (Kobayashi et al.. contralateral limb may be related to the disinhibition and strengthening of pre-existing commissural connections between the cortical representation of the two limbs. but not temperature or pain.. vibration or joint movement. 1990. Conversely. and (e) referral was reported from the intact hand and forearm up to a level corresponding to the amputation of the other arm. 1994). cold. Unilateral movements – particularly movements that are rapid and repetitive or involve force and effort – are associated with an increase in the regional cerebral blood flow in the ipsilateral motor cortex (Carson. the amputated limb and “phantom” are placed on one side out of view. changes in neuronal membrane excitability. (d) movements of the real hand – either passively or actively – were referred to the phantom hand in six amputees. strengthening of existing synapses in processes such as long-term potentiation. these neural pathways are recruited to enhance performance of bimanual movements.. exceeds perceptual threshold resulting in the perception of referred sensation. Four out of five of the amputees tested who had “clenching” spasms in their phantom limbs found instant relief upon looking into the mirror and opening “both” hands simultaneously. particularly in amputees who can execute voluntary phantom limb movements. 2003). the neurons that correspond to the anaesthetic hand may become disinhibited and responsive to input from the ipsilateral hand.. 1996). In the mirror box paradigm. 2001). which reinnervate the stump muscles (Mercier et al. 2003). 1988). and found that (a) three amputees perceived referred sensation that was topographically organised such that touching the intact thumb elicited referred touch in the phantom thumb. Ordinarily. particularly a myoelectric prosthesis (Lotze et al. Mirror box therapy has also been used by patients in the early and intermediate stages of Complex Regional Pain Syndrome (CRPS. 2003. 4. or the arm of the experimenter. MacLachlan et al. exhibit less cortical reorganisation.

C. but could also trigger and restore neural activity in the deafferented motor cortex. Temporary phantom limbs evoked by vestibular caloric stimulation in amputees. and movementrelated pain in CRPS (type 1). (b) motor overflow from the intact limb to the phantom limb measured by fMRI.g. J. The influence of task characteristics on the intermanual asymmetry of motor overflow. Soc. O. Neural correlates of frustration. The effects of age and attention on motor overflow production—a review. Disownership of left hand and objects related to it in a patient with right brain damage. 1999. the potential mechanisms underlying the interaction between the phantom limb and the intact limb require clarification.. probably via reduced intracortical inhibition. Rev. G. 2005. Franchini.. Future research should investigate the relative role of internal global constructs of the body (the body schema). Rapid sensory remapping in the adult human brain as inferred from phantom breast perception. 270. in normals. H. 2004)..L. V. Bradshaw. B Biol. Neurol..S. affectivemotivational and evaluative-cognitive dimensions of bodily experience – provides a functional model for phantom limb perception.. S. the relative roles of the global body schema. S. Bueti. the nature of the triggers of phantom sensations and pain (e. We propose that specific central processes beyond the body schema and neuromatrix Abler.. J. The investigation of cross-referencing the phantom limb with other parts of the body is an area of research that has received little attention to date. Armatas. 557–567. vestibular stimulation. 354. Conclusions Research on non-painful phantom limbs provides valuable insight to the mechanisms underlying bodily awareness and embodiment.. Manfredi. and kinetics). and potentially even evolutionarily adaptive and perhaps necessary.. 2007. S. referred sensations. The body schema most likely provides the template for phantom limb perception. 1976. again suggesting that the visual capture phenomenon is related to the dominance of vision over somatic senses such as proprioception.M. 189–204. 54. B. Acerra. Brain Res.. N..A. the perception of a “normal” (non-painful) phantom limb is in the very least an epiphenomenon of normal functioning. Georgiou-Karistianis. Hoy.. 14.. Behav. T.E.. Galati. Moseley. the activation and disinhibition of various mirror neuron systems – including systems involved in action understanding.C. Urol. but could be both theoretically and clinically invaluable.. 2005. André. Ramachandran. 1996. Ramachandran and Rogers-Ramachandran. This technique has important treatment implications not only for reducing levels of phantom pain in amputees. Aglioti. McCabe et al.. N. Hoy et al. 2006. 1994..L. S. J. Cortese... and visual capture through the mirror box paradigm). Phantom pain. Armel. NeuroReport 8. 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