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

“The intraparietal and premotor cortices are known to use visual, somatosensory, and motor information to guide the moving arm (Mountcastle et al., 1975; Bioulac and Lamarre, 1979; Hyvarinen, 1982; Kalaska et al., 1983; Georgopoulos and Massey, 1985; Colby and Duhamel, 1991; Caminiti et al., 1996; Fogassi et al., 1996;…”

“…evidence has shown that body representation is strictly linked with that of the space immediately surrounding it, the so-called peripersonal space (Fogassi et al., 1996), through the strong integration of multisensory input, in particular tactile stimuli on the skin and visual stimuli in the…”

“The region of space around the body that lies within reach (often known as peripersonal space) is prioritized in many areas of dorsal (Duhamel, Bremmer, Ben Hamed, & Graf, 1997; Galletti, Battaglini, & Fattori, 1993) and frontal (Fogassi et al., 1996) cortex.”

“Indeed, VIP neurons are highly selective to the direction and velocity of stimuli (Colby et al. 1993) and receive dense proprioceptive inputs (Lewis and Van Essen 2000), while the receptive field of many F4 neurons expands in depth when the stimulus speed increases (Fogassi et al. 1996).”

“…regions of the premotor (F2, F3, F4 and F5), cingulate and lateral parietal/opercular cortices, which host peripersonal ((Graziano et al., 1994; Fogassi et al., 1996); see (Graziano and Cooke, 2006)), mirror ((Gallese et al., 1996; Rozzi et al., 2008; Hihara et al., 2015); see (Bonini, 2017))…”

“…Graziano, 1999; Graziano and Cooke, 2006) and are located in cortical regions that support a multimodal system for upcoming action: parietal cortex, premotor cortex, and the putamen (Graziano and Gross, 1994; Fogassi et al., 1996; Duhamel et al., 1998; Graziano, 1999; Graziano and Cooke, 2006).”

“…in space perception. in particular, their rFs are anchored to a particular body part and when the body part is moved, the rF moves along with it (Fogassi et al. 1996; graziano and gross 1998). hence, it has been proposed that area F4 could be involved in the integration of multisensory…”

“These cells, found in the ventral intraparietal cortex, the putamen and in a small region of the premotor area of the monkey's brain (Fogassi et al., 1996) respond to tactile, visual and auditory stimulation that takes place in so called, peripersonal space or near space.”

“…reported to reflect spatial target location (Godschalk et al., 1985; Gentilucci et al., 1988; Mushiake et al., 1997; Hoshi and Tanji, 2002), the three-dimensional shape of motor targets (Murata et al., 1997) and visual objects in the peripersonal space (Graziano et al., 1994; Fogassi et al., 1996).”

“Several investigators (Fogassi et al. 1992, 1996; Graziano et al. 1994, 1997) have suggested that the location of these visual RFs is independent of eye position and that the PMv neurons encode space in a coordinate system that is not centered on the retina but rather on other body parts such as…”

“…for arm reaching (Borghi and Riggio 2015: 9).24 Thus, canonical neurons space dependency is due to the connections between area F5 and area F4 (Fogassi et al. 1996; Matelli et al. 1996; Borghi and Riggio 2015, p. 7; Costantini et al. 2011a, b).25 Accordingly, 23 See also Vainio et al. (2011),…”

“These neurons feature visual receptive fields (RFs) that extend up to 40 cm from the skin surface (Fogassi et al., 1996), that are spatially anchored to the limb (and follow changes in limb position) (Graziano, Hu, & Gross, 1997) and that are independent of the direction of gaze (Graziano & Gross,…”

“…sidewalls next to the participants either in extrapersonal space, i.e., beyond reach (large room-size condition), or in peripersonal space, i.e., the space surrounding the body (narrow room-size condition; Fogassi et al., 1996; Rizzolatti et al., 1997; Ladavas et al., 1998; Teneggi et al., 2013).”

“The current work is centered around these neural dynamics and designed to address whether the recruitment of PPS neurons in IPS and vPM results in notable alteration of the EEG signal.”

“Moreover, the firing rates of some of these neurons change dynamically with stimulus velocity [14].”

“In F4, the space is encoded in body-parts-centred coordinates and the visual receptive fields are also located in the peripersonal space (Grazziano et al. 1994; Fogassi et al. 1996).”

“PMv neurons respond to tactile and proprioceptive somatosensory stimuli (Fogassi et al., 1996; Graziano et al., 1997; Graziano, 1999).”

“These results 642 comport well with the known multisensory convergence in vPM [58, 59], but also represent the first evidence 643 that these neurons can integrate this information.”

“…revealed that activity in the premotor cortex (Ehrsson et al., 2004, 2005; Convento et al., 2018) is associated with feelings of owning the rubber hand, and that neurons in the ventral premotor cortex react to multisensory stimuli that guide action (Graziano et al., 1994; Fogassi et al., 1996).”

“These multisensory populations establish a somatosensory connection to visual stimuli in the environment and are regarded as the main encoders of peripersonal space (Graziano and Gross 1993; Fogassi et al. 1996).”

“…Radboud University Nijmegen, Nijmegen, The Netherlands and neurally dissociateable from the EPS in both primates (Rizzolatti et al. 1981a, b; Fogassi et al. 1996, 1999; Graziano et al. 1994, 1997; Murata et al. 1997; Duhamel et al. 1998; Caggiano et al. 2009) and humans (di Pellegrino et…”

“We speculate that this finding relates to movement close to a hand being a strong cue for stimulation in peripersonal space, a stimulus that activates visuotactile neuronal populations in the fronto-parietal association cortices (Brozzoli et al., 2011; Fogassi et al., 1996; Graziano et al., 1997; Makin et al., 2007),…”

“…stream involved in the vision for action has been confirmed by various neuropsychological, neurophysiological, and human neuroimaging studies (Fogassi et al., 1992, 1996; Clower et al., 1996; Kawashima et al., 1996; Rizzolatti et al., 1996; Robertson et al., 1997; Karnath and Perenin, 2005;…”

“The PMv receives abundant corticocortical projections from the parietal cortex [19,26,39], including the anterior intraparietal area (AIP) [34], through which visual and somatosensory information is provided.”

“mimic the visual body-part-centered receptive fields (RFs) of neurons that were observed in monkeys [9].”

“…the metrics of space representation are contingent upon the equilibrium emerging in a widespread neuronal network of space-coding neurons (such as those investigated by, for example, Graziano and Gross, 1994; Galletti et al., 1995; Fogassi et al., 1996) from a system of functional counterforts.”

“…receptive fields of F4 and VIP neurons: they are primarily centered around the head and the face, although F4 also shows more receptive fields around the hands and upper limbs than VIP (Gentilucci et al., 1983; Colby et al., 1993; Graziano et al., 1994; Fogassi et al., 1996; Duhamel et al., 1998).”

“In particular, multimodal neurons coding peripersonal space are activated by changes in the distance of objects from the body and their velocity (Fogassi et al., 1996; Holmes & Spence, 2004).”

“tation of space can be used to construct multiple frames of reference, each one related to a spatial location or a specific class of action, and these can be used by the motor system to code appropriate movements (Arbib 1991; Colby and Duhamel 1996; Fogassi et al. 1996; Ghafouri et al. 2002; Graziano et al. 1994; Rizz…”

“About 20 years ago, neurophysiologists observed neurons in the premotor cortex of the macaque monkey that respond to both visual and tactile stimulation (Fogassi et al., 1996; Graziano & Gross, 1994; Graziano, Hu, & Gross, 1997).”

“…& Schubö, 2014) to events localized in PPS. Seminal studies in non-human primate have shown that the ventral premotor cortex (vPMc), (e.g., Fogassi et al., 1996), and intraparietal sulcus, specifically in the ventral intraparietal area (VIP; e.g., Avillac et al., 2005), may be responsible…”

“…fronto-parietal network of the macaque brain, integrate tactile information on the body with visual (or auditory) stimuli occurring close to the body, i.e. within the PPS (Graziano and Cooke, 2006; Rizzolatti et al., 1981a, 1981b; Duhamel et al., 1998; Fogassi et al., 1996; Graziano et al., 1994).”

“…to both tactile and visual stimuli applied to the contralateral side of the body have been identified in the putamen, the parietal cortex and the inferior premotor cortex (Avillac et al., 2007; Duhamel et al., 1998; Fogassi et al., 1996; Graziano and Gross, 1993; Graziano and Gross, 1995).”

“Unlike studies in monkeys (Fogassi et al., 1996; Graziano et al., 1997), studies investigating the mapping of nociceptive stimuli in a peripersonal frame of reference in humans have mainly focused on a representation of the peripersonal space of the whole body using the body’s main axes, such as the midsagittal plane,…”

“The advantage of seeing the tactile stimulation on tactile feeling might be due to feedback projections from high-level multimodal areas (e.g., visual–tactile areas in the parietal cortex [3], [11], [12]) to more specialized unimodal areas (somatosensory areas in this case) [13], [14].”

“The visual receptive field would therefore represent a three-dimensional expansion of the cutaneous field (Fogassi et al., 1996).”

“The multimodal neurons may correspond to cells in the parietal and frontal cortex (observed via electrophysiological measures in animals; Fogassi et al., 1996; Graziano et al., 1997, 1999, and via neuroimaging studies in humans; Bremmer et al., 2001; Makin et al., 2007) having visual, auditory, and…”

“A large proportion of them also respond to all possible sensory stimuli (tactile, visual, acoustic) present on the skin or close to the skin, in correspondence of a specific tactile receptive field (Fogassi et al., 1996; Graziano et al., 1997, 1999; Rizzolatti et al., 1988).”

“The visual and tactile receptive fields of these neurons are in the spatial register; that is, the visual receptive fields match the location of the tactile receptive fields on the body surface, and extend in depth to the space immediately surrounding the body [Fogassi et al., 1996].”

“…close, as opposed to far from the location of tactile stimulation (Spence et al. 2004b; Shore et al. 2006), perhaps due to the response properties of visuotactile neurons, which respond more strongly for visual stimuli positioned close to the body (e.g. Fogassi et al. 1996; Rizzolatti et al. 2002).”

“In order to establish a comparison with the neurophysiological literature, we then studied the subset of neurons in the multisensory layer that positively respond to touch, to compare our artificial neural population to the one typically studied in primates (Fogassi et al., 1996; Graziano et al., 1994, 1997).”

“Our finding that F4 processes visual stimuli, regardless of execution of reaching movements towards them, is in agreement with previous studies reporting visual responses in F4 (Boussaoud et al., 1993; Fogassi et al., 1996a; Graziano et al., 1997).”

“…McCollum, Klam, & Graf, 2012; Sereno & Huang, 2006); (4) precentral polysensory zone (PZ) and ventral premotor cortex (PMv) (Bremmer et al., 2001; Fogassi et al., 1996; Graziano & Gandhi, 2000; Graziano, Yap, & Gross, 1994; Graziano, Hu, & Gross, 1997; Huang & Sereno, 2007, 2018); and (5) area…”

“Second, PMv hosts bimodal neurons that respond to both tactile and visual (Rizzolatti et al. 1981b; Graziano et al. 1994; Fogassi et al. 1996; Guipponi et al. 2015) or tactile and auditory (Graziano et al. 1999) stimuli delivered within peripersonal space sectors linked with specific body parts…”

“In fact, cognitive neuroscience has provided evidence that the brain uniquely represents this space by its potential for action and interaction with the physical world (Fogassi et al., 1996; Graziano & Gross, 1994; Rizzolatti, Fadiga, Fogassi, & Gallese, 1997).”

“Previous neurophysiological studies in monkeys (Fogassi et al. 1996; Gentilucci et al. 1983, 1988; Graziano 2006) and humans (Jeannerod 1994; Johnson-Frey 2004) have attributed to inferior parietal lobule and premotor cortex the role of affordance instantiation on the basis of intrinsic (grasp) and extrinsic (reach) o…”