Coding of peripersonal space in inferior premotor cortex (area F4).

“These neurons integrate multisensory information at the single-cell level, presenting both tactile and visual receptive fields (RFs), the latter limited to the region of space close to the body surface (Fogassi et al., 1996).”

“The stereotactic parameters were chosen on the basis of our previous single-neuron recording experience in the ventral premotor cortex see (Fogassi et al. 1996; Gallese et al. 1996; Gentilucci et al. 1988; Rizzolatti et al. 1988).”

“…visual stimuli are presented far (at more than about 30 cm) from the body part where the tactile receptive field is located (Colby, Duhamel, & Goldberg, 1993; Duhamel, Colby, & Goldberg, 1998; Fogassi et al., 1996; Graziano, Yap, & Gross, 1994; Rizzolatti, Scandolara, Matelli, & Gentilucci, 1981).”

“The design of object manipulation and communication capabilities should be inspired by interdisciplinary empirical and theoretical investigations of linguistic and cognitive development in children and adults, as well as of experiments with humanoid robots.”

“Neurons in the PMv receive visual and somatosensory inputs (Kubota and Hamada, 1978; Rizzolatti et al., 1981a,b; Fogassi et al., 1996; Graziano et al., 1997) and are active during the preparation for and execution of visually guided movements (Kubota and Hamada, 1978; Godschalk et al.”

“…position converge on bimodal visual–tactile neurones, some of which also respond to peripersonal visual stimuli centred on body parts (Colby et al., 1993; Fogassi et al., 1996; Graziano et al., 1997; Duhamel et al., 1998). when the monkey's vision of the arm is prevented (Graziano et al., 1994).”

“Ventral premotor area F4 is in fact involved in coding the peripersonal space for reaching [132], [133] and area F5 contains neurons coding hand/mouth grasping.”

“Crucial to this concept is the output of the so-called ‘real-position’ cells, found in the parietal areas V6A andVIP (Duhamel, Bremmer, Ben Hamed, & Graf, 1997; Galletti, Battaglini, & Fattori, 1993) and in premotor cortex (Fogassi et al., 1996, 1992; Graziano & Gross, 1998).”

“After chamber implantation, the ventral part of the agranular frontal cortex was functionally explored (single-unit recordings and intracortical microstimulation) in order to assess the location of areas F1 (primary motor cortex), F4 and F5 (PMv) and to find out the sector of F5 where neurons related to mouth actions…”

“Fogassi et al. (1996) recorded from single neurons in inferior premotor cortex and showed that the majority of cells responded to both visual and tactile stimulation.”

“Most neurons in both areas are bimodal, responding to visual and tactile stimuli (Bremmer et al. 2002; Colby et al. 1993; Duhamel et al. 1998; Fogassi et al. 1996; Graziano and Gandhi 2000; Graziano et al. 1997; Rizzolatti et al. 1991; Schaafsma and Duysens 1996).”

“Furthermore, canonical neurons fire only when objects are located in the peripersonal space, likely due to the connections between area F5 and area F4 (Fogassi et al., 1996; Matelli et al., 1996).”

“Procedures for the implantation of the head restraint and recording chamber were as described in previous studies (Fogassi et al. 1996).”

“Furthermore, this frontoparietal network is thought to subserve the multisensory representation of peripersonal space (Fogassi et al. 1996; Làdavas et al. 1998; Macaluso and Maravita 2010), as well as to respond to aversive visual stimuli presented within the peripersonal space, including stimuli representing potentia…”

“In monkeys, functional properties of bimodal neurons in premotor (Graziano et al., 1994, 1997; Fogassi et al., 1996; Rizzolatti et al., 1981a, 1981b) and parietal cortex (Avillac et al.”

“stimuli is spatially tuned, being stronger at shorter distances between the hand and the visual source, and vice versa (Duhamel et al., 1998; Fogassi et al., 1996; Fogassi et al., 1999; Graziano, Yap, & Gross, 1994).”

“…demonstrated in a number of cortical areas in both primate and infraprimate species (Bruce et al., 1981; Rizzolatti et al., 1981; Wallace et al., 1992; Ramachandran et al., 1993; Stein et al., 1993; Fogassi et al., 1996; Graziano et al., 1997, 1999; Duhamel et al., 1998; Schroeder et al., 2001).”

“In moneys, bimodal neurons have been described in inferior parietal reas (Hyvärinen & Poranen, 1974) and premotor cortex (Duhamel, olby, & Goldberg, 1998; Fogassi et al., 1996; Graziano & Cooke, 006; Graziano & Gross, 1995).”

“Neurons in premotor cortex of the macaque brain respond to tactile, visual and auditory stimuli (Rizzolatti et al. 1981; Gentilucci et al. 1988; Fogassi et al. 1996; Graziano et al. 1997, 1999).”

“For these characteristic properties bimodal VT populations have been suggested to encode the peripersonal space (Fogassi et al., 1996): through their vRFs these populations define in fact the region of space surrounding the body in which visual stimuli may trigger somatic sensations.”

“Similarly, movement in space necessarily implies a velocity; yet Fogassi et al. (1996) demonstrated that PPS neurons in the inferior premotor cortex are sensitive to such feature of movement kinematics and showed that receptive fields extend proportionally as the velocity of the incoming stimuli…”

“For example, certain neurons in the ventral and dorsal premotor cortex represent objects in hand-centered coordinates (Graziano et al., 1994, 1997; Fogassi et al., 1996, 1999; Graziano, 1999) and may be involved in the preparation and guidance of movements both toward and away from objects (as…”

“‘extrapersonal space’ [7,27,28]; for a review see[25]).”

“…to a stimulus at 57 cm. Bimodal visuotactile neurons have also been found in inferior area 6 (Rizzolatti et al., 1981; Graziano and Gross, 1995; Fogassi et al., 1996); they have somatosensory receptive fields on the face (24%), the arm (34%), both the face and the arm (29%), the chest (2%),…”

“Premotor area F4 ( a sector of area 6 in the macaque monkey brain ) contains neurons that integrate motor , visual , and somato - sensory modalities for the purpose of controlling actions in space and perceiving peripersonal space , the area of space reachable by head and limbs ( Fogassi et al .”

“However, evidence for sensory responses within monkey premotor cortex in the absence of any movement preparation (Fogassi et al., 1996; Graziano et al., 1999; Graziano and Gandhi, 2000) has begun to challenge this functional dichotomy in favor of a more distributed sensorimotor representation in…”

“Drawing more from the neural science literature, the results of Fogassi et al. (Fogassi et al., 1996) and Graziano et al. (Graziano et al., 1997) have shown the existence of neurons that respond to objects and are related to the description of the peripersonal space with respect to reaching (area F4 and VIP).”

“Similar responses have also been reported for a sub-set of single units in a multisensory region within the ventral premotor cortex (PMv; Fogassi et al., 1996; Graziano et al., 1997), an area that receives afferents from VIP but not from adjacent parietal regions (Luppino et al., 1999).”

“Furthermore, most bimodal neurons identified in primates had a boundary extending beyond 30 cm and many didn't have a clear border at all (Fogassi et al., 1996).”

“Some ventral PMA neurons respond to visual, auditory and tactile stimuli (Graziano et al., 1999), coding stimulus position in nonretinal, body-centred coordinates that are anchored to speci®c body parts (Fogassi et al., 1996; Graziano et al., 1997).”

“…in F2 (Caminiti et al. 1991; Crammond and Kalaska 1994, 1996; Hoshi and Tanji 2000; Johnson et al. 1996; Weinrich and Wise 1982) and in F4 (Fogassi et al. 1996a; Gentilucci et al. 1988) and distal movements in F2 (Kurata 1993, Raos et al. 1999) and in F5 (Kurata and Tanji 1986; Murata et…”

“Cell responses are highest when visual stimuli approach the tactile receptive field [5], and electrical stimulation of neurons in this circuit evokes defensive behaviors [32].”

“…the contribution of a well characterized visuotactile convergence network of post-arcuate premotor (premotor area F4; Gross and Graziano, 1995; Fogassi et al., 1996), intraparietal [ventral intraparietal area (VIP); Duhamel et al., 1997, 1998; Bremmer et al., 1997, 2000, 2002a,b; Guipponi et…”

“Expansion of near space Neurophysiological and neuropsychological studies have shown that near space representations can be dynamically expanded (e.g., Fogassi et al., 1996; Iriki et al. 1996).”

“In monkeys, bimodal neurons representing PPS were first described in the ventral premotor cortex, specifically in area F4 [1,2,28,29], which contains neurons representing specific body parts movements [30–32].”

“In particular, the visual receptive fields of F4 neurons around the hand extend from 5 to 35 cm from the tactile receptive fields [17].”

“These observations have prompted the hypothesis that the VIP–F4 circuit represents peripersonal space in a frame of reference centered on the body part involved in a given visually guided action (Graziano et al., 1994, 1997; Fogassi et al., 1996).”

“Thus, premotor neurons discharge not only during movement, but also during sensory (Rizzolatti et al., 1981b, 1988; Gentilucci et al., 1983, 1988) and somatosensory (Rizzolatti et al., 1981a) stimulation or both (Gentilucci et al., 1988; Fogassi et al., 1996a,b; Graziano and Gandhi, 2000).”

“The neurons of both these regions have bimodal visuotactile receptive fields representing close peripersonal space and the corresponding skin surface (Gentilucci et al., 1988; Rizzolatti et al., 1988; Colby et al., 1993; Graziano et al., 1994; Gross and Graziano, 1995; Fogassi et al., 1996; Duhamel et al., 1998).”

“…PMd: dorsal premotor cortex. muscle related coordinates between PM and the motor cortex (Crammond & Kalaska, 1996; Graziano, Hu, & Gross, 1997; Kalaska, Scott, Cisek, & Sergio, 1997), and an independence of many visual receptive fields with respect to eye position in PM (Fogassi et al., 1996).”

“In monkeys, a network of fronto-parietal regions, involving area F4 in the premotor cortex (PMc; Fogassi et al., 1996; Graziano et al., 1994, 1997, 1999; Rizzolatti et al., 1981) and the ventral intraparietal area (VIP; Avillac et al.”

“Bimodal neurons in the monkey parietal cortex and related structures perform this integration by responding to both somatosensory and visual information from a given body area and surrounding space (Fogassi et al. 1996 ; Duhamel et al. 1998; Graziano & Cooke, 2006).”

“Those neurons have been extensively investigated in non-human primates (Graziano et al., 1994; Fogassi et al., 1996) and their presence has been also documented in humans (Bremmer et al., 2001).”

“Whereas considerable work has focused on the coordinate frames in which premotor cortex neurons represent spatial information (Fogassi et al., 1992, 1996; Caminiti et al., 1996; Graziano et al., 1997; Kalaska et al., 1997; Graziano & Gross, 1998), their functional correlates remain uncertain.”

“1997), and ventral premotor cortex (vPM) (Fogassi et al. 1996; Graziano et al. 1997) have been reported to respond to tactile stimuli applied to different body parts.”

“Target location and movement direction can be defined in eye-centered, armcentered or object-centered coordinates depending on the cerebral area and the task (Batista et al. 1999; Caminiti et al. 1990; Fogassi et al. 1996; Lacquaniti et al. 1995; Tremblay et al. 2002).”

“F4 cells provide a representation of the space around the head and body (Fogassi et al., 1996); F2 cells focus on body rather than face regions (Fogassi et al.”

“, handcentered) (Bremmer, Schlack, Duhamel, Graf, & Fink, 2001; Duhamel, Golby, & Goldberg, 1991, 1998; Graziano, Tian Hu, & Gross, 1997; Fogassi et al., 1996; Graziano & Gross, 1993, 1995; Rizzolatti, Scandolara, Matelli, & Gentilucci, 1981).”

“The visual receptive fields are independent of eye position, likely reflecting coding of perip-ersonal space based on a body part–centered frame of reference (Graziano et al., 1994; Fogassi et al., 1996).”

“Additionally, these neurons are most responsive to moving, as opposed to static, stimuli (Fogassi et al., 1996).”

“In particular, the visual receptive fields of F4 neurons around the hand extend from 5 to 35 cm from the tactile receptive fields [23].”

“…of this finding with respect to the existence of neuronal populations containing multisensory receptive fields that encompass a large portion of the body is compatible with neurophysiological data (Rizzolatti et al., 1981a,b; Fogassi et al., 1996; Iwamura, 1998; Graziano and Gandhi, 2000).”

“, the somatosensory Receptive Field; sRF), and to visual information from the space (visual Receptive Field; vRF) adjacent to it [65,66].”

“…been found includes classic regions of higher-order cortex, including: posterior parietal cortex (Xing and Andersen 2000; Andersen and Buneo 2002), intraparietal cortex (Duhamel et al. 1998), premotor cortex (Fogassi et al. 1996; Graziano et al. 1997), and prefrontal cortex (Fuster et al. 2000).”

“where the tactile field is located (Fogassi et al., 1996; Gentilucci et al., 1988; Graziano & Gross, 1995; Graziano, Yap, & Gross, 1994).”

“Neurophysiological and neuroimaging investigations (Rizzolatti et al., 1981; Fogassi et al., 1996; Bremmer et al., 2001; Brozzoli et al., 2011, 2014) have shown that the premotor cortex (PM) plays a crucial role in motor mapping of sensory events occurring near the body, especially in humans…”