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

“This finding is reinforced by the neuronal response obtained during active movements [31,45].”

“[4] on the other hand, although also arguing for a representational failure, link it instead to changes in neurones in areas F4 [8], area 6[11] and especially area V6 (PO) [10] in the visual dorsal stream.”

“In contrast, PMv neurons respond to somatosensory stimuli applied to either the face or the arm and to visual stimuli corresponding to peripersonal stimuli (Fogassi et al. 1996; Graziano et al. 1997).”

“Tactile responses in the ventral premotor cortex (PMv; areas F4 and F5) have already been described, with receptive fields preferentially located on the face, near the mouth, and on the hand (Fogassi et al. 1996; Gentilucci et al. 1988; Rizzolatti et al. 1981).”

“This, added to the fact that the firing rate of the multimodal neurons in both areas (Rizzolatti et al., 1981a,b; Graziano and Gross, 1993; Fogassi et al., 1996; Duhamel et al., 1998; Graziano, 1999; Graziano et al., 2000) increases as more visuo-tactile stimuli approach the face, leads to propose…”

“In this context, it has been found that the core part of the cortex involved in the encoding process of the peripersonal space is premotor area F4 [78].”

“For instance, stimuli looming toward the body enhance tactile sensitivity at the spatial and temporal location where observers expect impact to occur [9], and PPS enlarges as the speed of incoming stimuli grows [10].”

“The neural substrate supporting this direct perceptual mechanism is called the “mirror neuron” system that has recently been discovered in monkeys and humans (Buccino et al., 2001; Fadiga, Fogassi, Pavesi, & Rizzolatti, 1995; Fogassi et al., 1996; Iacoboni & Dapretto, 2006; Rizzolatti, Fadiga, Gallese et al., 1996; Ri…”

“…putamen, and the intraparietal sulcus, have overlapping visual and tactile receptive fields (vRFs and tRFs, respectively), typically on the face or hand (Rizzolatti et al., 1981a,b; Graziano and Gross, 1993; Graziano et al., 1994; Fogassi et al., 1996; Graziano, 1999; Graziano and Gandhi, 2000).”

“Accordingly, neurophysiological research in animals has identified a network of cortical and subcortical brain areas, including the ventral premotor cortex, the posterior parietal cortex, the putamen, and the superior colliculus, that subserves the visuotactile representation of peripersonal space [1, 4, 5, 7].”

“…movement [Boussaoud, 1985]/guiding reaching area [Cisek and Kalaska, 2005]) and in F4/F5 (sensory guidance of movement [Graziano et al., 1994] and peripersonal space [Fogassi et al., 1996]/hand shaping during grasping, vocalization [Coudè et al., 2011] and mirror neurons [Kohler et al., 2002]).”

“Similar bimodal neurons have been reported in the premotor cortex and the putamen (Graziano et al. 1994; Fogassi et al. 1996; Duhamel et al. 1998).”

“Fogassi et al. (1996) reported that the visual receptive fields of visuotactile neurons in area F4 expanded with an approaching visual object’s increasing velocity.”

“…has been hidden; Obayashi, Tanaka, & Iriki, 2000), and that their visual receptive fields are anchored to the hand and extend into the space adjacent to the skin surface (i.e., perihand space) up to 40 cm (Fogassi et al., 1996; Graziano & Gross, 1996; Graziano et al., 1997; Guterstam et al., 2016).”

“Some neurons in supplementary motor area and premotor area fire when stimuli of each of the three modalities are presented; however, they may fire preferentially for one or more of them (Fogassi et al. 1996; Gentilucci et al. 1988; Graziano and Gandhi 2000; Graziano et al. 1997, 1999; Kurata and Tanji 1986; Rizzolatti…”

“…neurons with small visual RFs (~30 cm), a differential rate of firing occurs for stimuli within the animal's reach (Graziano and Gross, 1995; Fogassi et al., 1996; Duhamel et al., 1998), thus coding for the limited portion of space immediately surrounding the body (i.e. near peripersonal…”

“Two are the main circuits involved in this type of coding: the LIP-FEF, devoted to the movement programming of the eyes (Barash, Bracewell, Fogassi, Gnadt, & Andersen, 1991); and the VIP-F4 dedicated to the encoding of movements of head, arm and hand (Fogassi et al., 1996).”

“…regions are activated (Weiss et al. 2000), and different coordinate systems are involved in the processing of near space compared to far space (Fogassi et al. 1996) These findings lead to the question of whether the perception/action dissociation found in nearspace tasks generalizes to action…”

“PPNs are mostly found in brain area F4 and are multimodal and somatocentered (Fogassi et al., 1996).”

“Individual neurons were shown to encode peripersonal spaces, responding to stimuli touching or appearing close by a particular part of the hand or face (Fogassi et al., 1996; Maravita et al., 2003).”

“In macaques, bimodal visuotactile neurons are distinguished by their characteristic response properties in peripersonal space [34] [35] [36] [37] [30] [38] [39] [40] [41] [42].”

“For example, bimodal neurons, which code for peripersonal and extrapersonal space, have been described in inferior parietal areas and the ventral premotor cortex (Duhamel et al., 1998; Fogassi et al., 1992, 1996; Graziano and Cooke, 2006; Graziano et al., 1994; Graziano and Gross, 1998).”

“VIP shares connections with area F4 of premotor cortex in the macaque,(39) which contains neurons that represent face, neck, and proximal arm movements and most likely codes peripersonal space in body- or headcentered reference frames.(48,49) In humans, resting-state functional connectivity and dMRI studies suggest a…”

“These bimodal neurons show increasing response to visual stimuli as they get closer to the hand (Foggassi et al., 1996; Graziano, Hu, & Gross, 1997).”

“The known input– output connectivity of PMv supports the likelihood of converging somatosensory information from SII (directly to PMv and indirectly via AIP and ventral intraparietal area (VIP)) and visual target information from AIP (Dancause et al., 2006) and VIP (Duhamel et al., 1997).”

“…a specific body part, and by visual stimuli close to the same body part, thus promoting a body part-centered representation of the near space (e.g., Rizzolatti et al., 1981, 1988, 1997; Fogassi et al., 1992, 1996; Graziano and Gross, 1993, 1995, 1998; Graziano et al., 1994, 1997; Graziano, 1999).”

“The visual and tactile receptive fields (RFs) of these bimodal neurons are in spatial register, that is the visual RF matches the location of the tactile RF on the body surface, and extends in depth to the space immediately surrounding the body [3], [4].”

“For example, Fogassi et al. (1996) described bimodal neurons in macaque premotor cortex with both tactile and visual receptive fields (RFs).”

“Most visual receptive fields encode nearby space in a body-part-centered reference frame (Fogassi et al, 1992; Fogassi et al, 1996; Gentilucci et al, 1983; Graziano et al, 1994; Graziano et al, 1997).”

“…distributed representation of space can be used to elaborate a synthetic ‘‘construct” that is the representation of the body’s spatial orientation that encodes the appropriate movements (Arbib, 1991; Gurfinkel et al., 1993; Fogassi et al., 1996; Ghafouri et al., 2002). ed by Elsevier Ireland Ltd.”

“…in premotor and parietal cortical areas are responsive to nearby stimuli relative to a particular body surface, anticipate approaching stimuli, and partially relocate their receptive fields during tool usage (Canzoneri et al. 2013; Fogassi et al. 1996; Holmes and Spence 2004; Iriki et al. 1996).”

“Furthermore, the visual and tactile receptive fields remain in correspondence when the arm moves, but almost all macaque bimodal units centered on the arm do not move with either the eye or the head (Fogassi et al., 1996; Graziano & Gross, 1994, 1995; Graziano et al., 1997).”

“(Fogassi et al., 1996; Graziano and Cooke, 2006; Rizzolatti et al., 1997).”

“Electrophysiological studies at a single neuron level indicate that an interaction between vision and proprioception may change cellular discharge (Duhamel et al. 1998; Fogassi et al. 1996; for review, Graziano and Botvinick 2002).”

“Damage to the posterior ventral pre-motor cortex is responsible for peripersonal neglect, while damage to the frontal field of vision is responsible for extra-personal neglect [9][10].”

“Separate networks are devoted to the movement programming of the eyes [17–19] and to the encoding of reaching movements of the head, arm, and hand [20].”

“In these (and human-analogue) sensorimotor regions, electrophysiological studies in non-human primates [5,6,14,15,37,38] and neuroimaging work in healthy humans [27,39,40] have identified populations of neurons exhibiting multisensory processing.”

“All participants gave informed consent prior to participation and were remunerated with 20 Swiss Francs for their time.”

“Surprisingly, under specific circumstances, the visual properties of these neurons can be extended to a portion of space beyond that covered by the tactile receptive field [38].”

“, [3]; [1] for a review) firing when some part of the skin is stimulated or a visual stimulus is presented nearby.”

“In addition to this, the premotor cortex is recognized for its involvement with responses to auditory, visual, and somatosensory inputs [52].”

“Cortical areas integrating multiple sensory modalities such as area VIP and sectors of area PF in the posterior parietal cortex, and area F4 in the ventral premotor cortex, code the presence of objects within a limited space sector, peripersonal space (Gentilucci et al., 1988; Fogassi et al., 1996; Rizzolatti et al.,…”

“Neurophysiological studies in monkeys have shown that faster-looming stimuli elicit responses in PPS neurons at farther distances than slower stimuli (Fogassi et al. 1996).”

“Furthermore, Fogassi et al. [12] demonstrated that the magnitude of visual receptive fields in the F4 region (premotor cortex), which is involved in defensive PPS, depends on the speed of approaching stimuli.”

“This phenomenon can be easily explained by neurophysiological recordings in monkeys, which stress the bimodal characteristic of neurons coding the peripersonal space surrounding each part of the body and especially the hand (Fogassi et al., 1996; Grazziano & Gross, 1995).”

“Effects of Animation The role of the motor system in shaping and maintaining the bodily self and body ownership in particular has been welldocumented by neuroimaging studies showing the emergence of premotor cortex activity lying at the root of our body schema (Graziano et al., 1994; Fogassi et al., 1996; Ehrsson et a…”

“These results are in accordance with the existence of a network devoted specifically to the representation of peripersonal space [13, 16, 17, 43].”

“When the animal moved its body part, the auditory or visual fields accompanied the tactile receptive field, representing the visual or auditory space centered on that body part [21–27].”

“…the visual stimulus is closer to the body rather than far away from it, as revealed by electrophysiological studies in monkey (Graziano, Reiss, & Gross, 1999; Duhamel, Bremmer, Hamed, & Graf, 1997; Fogassi et al., 1996) as well as humans (Noel, Serino, & Wallace, 2019; Bernasconi et al., 2018).”

“It was conceived as an interface between the body and the environment contributing to the orientation of attention toward objects that represent potential targets for motor actions, and would thereby serve two essential functions: selecting potential actions toward incentive objects and protecting the body from threat…”

“However, here we speculate that a careful comparison between motor processes (Fogassi et al., 1992, 1996) in DP versus psychosis may reveal important differences, as demonstrated by previous studies on schizophrenia patients (see Ardizzi et al., 2020).”