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

“…objects and the body (Bufacchi and Iannetti, 2018), thus contributing to shape defensive behavior (Graziano and Cooke, 2006; Graziano, 2016; Kaas et al., 2018) and other ethologically relevant functions, such as reaching to bring food to the mouth (Gentilucci et al., 1988; Fogassi et al., 1996).”

“Subsequently, several neurophysiological studies focused on these areas, mostly investigating the dorsal portion of F4 (Gentilucci et al., 1988; Fogassi et al., 1996) and the whole area F5 (Rizzolatti et al.”

“Multimodal neurons in the ventral premotor cortex map visual and auditory stimuli in spatial relation to the space immediately surrounding the body, or peri-personal space (Rizzolatti et al., 1981; Graziano et al., 1994; Fogassi et al., 1996).”

“The physiological properties of neurons in the PMV have been studied in macaques, and many neurons respond to tactile stimuli (Rizzolatti et al., 1981; Gentilucci et al., 1988; Fogassi et al., 1996; Graziano et al., 1997).”

“The final circuit involves reciprocal parieto-frontal links between the PPC and PMv (Cavada & Goldman-Rakic, 1989; Clower, West, Lynch, & Strick, 2001; Fogassi et al., 1996; Graziano, Hu, & Gross, 1997; Kurata, 1991; Neal, Pearson, & Powell, 1990a,b).”

“The startle is not spatially directed;auditory stimuli (Fogassi et al., 1996; Gentilucci et al., 1988; Graziano and Gandhi, 2000; Graziano et al., 1997, it is typically bilaterally symmetric, including, for exam- ple, a bilateral blink.”

“…of cross-modal neurones whose visual response is typically anchored to a particular point of the skin corresponding to their tactile receptive field (Gentilucci et al., 1983; Graziano et al., 1994; Graziano and Gross, 1995; Fogassi et al., 1996), even when such a body part is moved in visual space.”

“…“peripersonal” neurons fire either when a tactile stimulus is delivered to the animal skin and when a visual stimulus is presented in the space close to the body part where the tactile field is located (Fogassi et al., 1996; Gentilucci et al., 1988; Graziano and Gross, 1995; Graziano et al., 1994).”

“…title or abstract (see also Murray et al. (2013) and Stein et al. (2010)). extend in depth as the speed of an approaching visual stimulus increased (Fogassi et al., 1996), one could argue that an extension of RFs in depth may also depend on the speed of movement with which humans move through…”

“1 3 representation of a spatial location (that is signalled by the encroaching object) allowing a potential action directed towards that location (Fadiga et al. 2000; Fogassi et al. 1996).”

“…" neurons fire both when a tactile stimulus is delivered to the animal's skin and when a visual stimulus is presented in the space near the part of the body where the tactile field is located (Fogassi et al., 1996; Gentilucci et al., 1988; Graziano and Gross, 1995; Graziano et al., 1994).”

“Furthermore, the spatial resolution we have selected (with taxels of around 2 cm in diameter on the skin surface) is likely unnecessarily high—the body-part-centeredreceptive fields of parietal cortex neurons are typically much larger (e.g. spanning a whole upper arm [5]).”

“Most of these cells show a response gradient: typically, their firing rate increases when visual stimuli are presented closer to the somatosensory body part (hand) and decreases when it is more distant (Duhamel et al., 1991, 1998; Fogassi et al., 1996; Fogassi et al., 1999; Graziano & Gross, 1995; Rizzolatti et al., 1…”

“Owing to these functional properties, it has been suggested that the premotor cortex, parietal areas and putamen form an interconnected system for multisensory coding of near peripersonal space centered on body parts (Colby et al. 1993; Fogassi et al. 1996; Graziano et al. 1997; Duhamel et al. 1998; Fogassi et al. 199…”

“This effect can be explained by the anatomical connections of this sector of area F5 with the adjacent area F4 (Matelli et al., 1986), whose neurons encode monkey’s peripersonal space in a pragmatic format (Fogassi et al., 1996).”

“The firing rate of some of these neurons increases as function of the velocity of the looming stimulus, suggesting that these neurons might be computing the time to impact on the body (Fogassi et al., 1996).”

“…that the fronto-parietal brain areas implied in multisensory integration during the RHI contain neurons with body part-centered visuo-tactile receptive fields, which may help code the space around the body (Fogassi et al., 1996; Graziano, 1999; Graziano et al., 2000; Graziano and Cooke, 2006).”

“…et al., 2004; Tsakiris et al., 2007; Evans and Blanke, 2013) and analogous regions in the monkey brain have been found to contain trimodal neurons that integrate tactile, visual and proprioceptive signals (Fogassi et al., 1996; Iriki et al., 1996; Graziano et al., 1997; Graziano and Gandhi, 2000).”

“…of studies with non-human primates revealing that both premotor and intraparietal areas contain neurons that integrate visual, tactile, and proprioceptive information in body part-centered reference frames (Avillac, Deneve, Olivier, Pouget, & Duhamel, 2005; Fogassi et al., 1996; Graziano, 1999).”

“This is in line with evidence in monkeys showing that neurons integrating multisensory information within the PPS around a certain body part are also involved in movements of the same body part (Duhamel et al., 1998; Fogassi et al., 1996; Graziano et al., 1994; Rizzolatti et al., 1997).”

“VIP also projects to an area in the ventral premotor cortex (Luppino et al., 1999), which is involved in head movements (Fogassi et al., 1992, 1996).”

“…Colby, & Goldberg, 1998; Graziano & Gross, 1994) and the ventral premotor cortex, F4 THE QUARTERLY JOURNAL OF EXPERIMENTAL PSYCHOLOGY, 2009, 62 (12) 2457 (Fogassi et al., 1992, 1996; Graziano, Yap, & Gross, 1994) with the VIP having projections to F4 (Luppino, Murata, Govoni, & Matelli, 1999).”

“1 mg/kg), followed by postsurgical antibiotic and pain medications (Fogassi et al., 1996; Rozzi et al., 2006).”

“…namely parietal area VIP (Duhamel et al., 1997, 1998; Avillac et al., 2005; Schlack et al., 2005; Bremmer et al., 2013; Guipponi et al., 2013) and premotor area F4 (Cléry et al., 2015b; Fogassi et al., 1996; Graziano et al., 1994, 1997, 1999; Matelli and Luppino, 2001; Rizzolatti et al., 1981).”

“Two types of visual stimuli were used: the same stimuli that served as targets were presented for 100 ms and an experimenter’s hand moving toward the monkey’s eyes (Fogassi et al. 1992, 1996).”

“The visual recepive fields of many F4 neurons are anchored on the tactile receptive elds, and are independent of eye position (Fogassi et al., 1996), lthough for some F4 neurons the visual receptive field is in retinoopic coordinates (Mushiake et al.”

“In line with such a defensive role, PHS boundaries may expand as a function of the speed of approaching objects (Fogassi et al., 1996), thus allowing individuals to respond to objects approaching the body at higher speeds at even farther distances.”

“Remarkably, many neurons display dual visual/tactile receptive fields organized in a bodyanchored reference frame; this kind of neurons respond to visual stimuli approaching a specific body part (Rizzolatti et al., 1981; Graziano et al., 1994; Fogassi et al., 1996).”

“This scheme is consistent with evidence that the shoulder-centered position of the arm is represented in premotor and motor cortices (Fogassi et al., 1996; Graziano, Hu, & Gross, 1997).”

“The existence of the PPS was firstly documented in monkeys (Fogassi et al., 1996; Iriki et al., 1996) through neural responses recording to stimuli belonging to different sensory domains.”

“Numerous findings have been accumulated by using different methodologies: single-cell recordings in animals (Hyvarinen, 1981; Rizzolatti, Scandolara, Matelli, & Gentilucci, 1981; Graziano & Gross, 1995; Fogassi et al., 1996; Graziano, Hu, & Gross, 1997; Duhamel, Colby, & Goldberg, 1998; Avillac, Ben, & Duhamel, 2007),…”

“These areas receive projections from earlier motion processing regions such as MSTd, and as a consequence it is no surprise that they are velocity (Fogassi et al. 1996, Noel et al. 2018c) and motion-direction (Duhamel et al. 1998) selective (particularly sensitive to looming stimuli).”

“ously to dynamic stimuli (Fogassi et al., 1996) and that proprioceptive inputs contribute to remapping of the peri-hand space upon limb movements (Brozzoli et al.”

“…(areas F4 and F5, illustrated in Figure 1, see Matelli et al. 1985) showed that these areas code several types of motor act executed with axial (trunk), proximal (arm) and distal (hand and mouth) effectors (Gentilucci et al. 1988; Rizzolatti et al. 1988; Fogassi et al. 1996; Ferrari et al. 2003).”

“Findings from primate and human studies suggest that the brain region is involved in coding locations of somatosensory (visual, auditory, and tactile) stimuli in extrapersonal space with respect to a body-centred reference frame (Fogassi et al., 1996; Galati et al., 2001).”

“The notion of PPS was initially proposed by neurophysiological studies describing the properties of a particular ensemble of neurons with bimodal (tactile, auditory, visual) receptive fields anchored to specific body parts and extending over a limited sector of space surrounding the same body part (Duhamel et al., 199…”

“That is, making use a latent variable of the network (i.e., the activity of the multisensory neuron), we replicate the exact electrophysiological analysis as in Fogassi et al. (1996) (except that we used auditory probing stimuli, instead of visual ones).”

“Such questions led Rizzolatti and colleagues to claim, ''The fundamental problem now is to understand how these different space maps interact and give an introspectively unitary space percept'' (Fogassi et al., 1996).”

“The premotor cortex receives visual and somatosensory information (Fogassi et al., 1996; Graziano et al., 1997) and is involved in visually guided movements and limb positioning in space (Godschalk et al.”

“…fields covering the space around the hand, face and trunk, within a network of fronto-parietal areas spanning from the ventral premotor cortex (Fogassi et al., 1996, Graziano et al., 1997), the ventral intraparietal area (Avillac et al., 2005; Duhamel et al., 1997,) and parietal area 7…”

“If the body part (e.g., the arm) to which the tactile receptive field is anchored moves, the spatial location of the visual or auditory receptive field shifts congruently from the initial spatial position to its new location (Fogassi et al., 1996; Graziano et al., 1997).”

“Other neurons, especially in inferior parietal lobule are active during both action execution and observation, suggesting similar mirror-like properties in the parietal lobe (Fogassi et al., 1996; Fogassi and Gallese, 2000; Fogassi et al., 2005; Nelissen et al., 2005).”

“Area F4 contains neurons that respond to objects and are related to the description of the peripersonal space with respect to reaching (Graziano et al., 1997; Fogassi et al., 1996).”

“, 1988) or F4 neurons to the visual stimuli with their visual receptive field encoded in the body-parts-centered coordinates (Fogassi et al., 1996; Graziano et al., 1994).”

“…in area 7b (Graziano and Gross 1994), the intraparietal sulcus (Colby et al. 1993; Duhamel et al. 1998; Iriki et al. 1996), the ventral premotor (PMv) area (Rizzolatti et al. 1981; Graziano and Gross 1994; Fogassi et al. 1996), and the superior temporal polysensory (STP) area (Bruce et al. 1981).”

“, 2005) and frontal and prefrontal cortices (Fogassi et al., 1996; Graziano et al., 1999; Romanski, 2007), the cat anterior ectosylvian (Clemo et al.”

“Electrophysiological studies in non-human primates showed that neurons in the anterior superior parietal lobule and the premotor cortex integrate visual and somatosensory stimuli involving the arm and the trunk (Fogassi et al., 1996; Graziano et al., 1997, 2000; Maravita and Iriki, 2004).”