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

“In these cases, a position in peri-personal space can be specified in a number of ways: sound, sight, and touch (Duhamel, Colby, & Goldberg, 1998; Fogassi et al., 1996a; Gentilucci et al., 1988; Graziano & Gross, 1995; Graziano et al., 1994; Rizzolatti et al., 1997).”

“(Fogassi et al., 1996; Graziano et al., 1997a).”

“As in area VIP, F4 neurons can be subdivided into two categories according to their responses to sensory stimuli: bimodal neurons (56%) and unimodal neurons (44%; Fogassi et al., 1996).”

“As such, the ventral premotor cortex has been postulated to play an important role in spatial attentional processing and in encoding the locations of objects in peripersonal space (Fogassi et al. 1996; Graziano et al. 1994).”

“Fogassi et al. (1996) studied the responses of F4 neurons to 3D objects moved toward the monkey at different velocities.”

“These areas are part of a system that controls actions and they contain many neurons that integrate visual, tactile and proprioceptive information in head and body-part centred reference frames [7,41,42,43,47].”

“Studies of the visual properties of F4 neurons showed that in most F4 neurons the receptive fields do not change position with respect to the observer's body when the eyes move (Gentilucci et al. 1983; Fogassi et al. 1992, 1996; Graziano et al. 1994).”

“One class of precentral neuron has a distinc1993; Sanes et al., 1995). tive type of response to tactile and visual stimuli (Rizzo- latti et al., 1981; Fogassi et al., 1996; Graziano et al., 1997).”

“…to the location of the tactilePopulations of neurons that respond to both visual and somatic stimuli have been found in several brain regions, RF (Fogassi et al. 1996; Gentilucci et al. 1983; Graziano and Gross 1994a; Leinonen and Nyman 1979; Leinonen 1980).with similarities and differences with…”

“Surgical and recording procedures were described in detail response to the latter of ¢20%. in our previous articles on premotor cortex (Fogassi et al. 1996; Selectivity for a single object (ring, plate, cylinder) was Gentilucci et al. 1988; Rizzolatti et al. 1990).”

“…for the position of a tactile stimulus delivered to the animal skin and for a visual stimulus presented in the space near the part of the body where the tactile field is located (bimodal neurons, Fogassi et al., 1996; Graziano, Yap, & Gross, 1994; Graziano & Gross, 1995; Gentilucci et al., 1988).”

“, 2011), which are well-known cortical nodes for integrating visual, tactile, and proprioceptive information from the body and the space immediately surrounding the body in body-part-centered coordinates (Graziano and Gross, 1993; Fogassi et al., 1996; Graziano et al., 1997; Duhamel et al., 1998; Graziano, 1999).”

“Premotor regions are responsive to auditory, visual, and somato- 392 Neuron 69, 387–396, January 27, 2011 ª2011 Elsevier Inc. sensory stimuli (Fogassi et al., 1996; Graziano et al., 1994, 1999; Lemus et al., 2009), and temporal regions are involved in the cross-modal integration of audiovisual…”

“In the studies of Fogassi et al. (1996) and Graziano et al.”

“The popula- It has been suggested (Fogassi et al. 1992, 1996) that tion of visual receptive fields moved 157, maintaining precise such modulation of activity in PMv neurons is not caused register with the head.”

“RF, and declining as it is moved away [13,15,18].”

“…with the real counterpart that remains out of view. neurons were proposed to build an enlarged and flexible representation of the space surrounding the body—the peripersonal space—that mainly serves to guide our movements (Graziano and Gross, 1994; Fogassi et al., 1996; Graziano and Cooke, 2006).”

“…actions in space and perceiving peri-personal space (the area of space reachable by body parts) (Fogassi, Gallese, Fadiga, & Rizzolatti , 1996a; Fogassi et al., 1996b; Fogassi et al., 1992; Gentilucci et al., 1988; Gentilucci, Scandolara, Pigarev, & Rizzolatti, 1983; Rizzolatti & Gentilucci,…”

“…described above and the consistency of the neuronal data with those of our previous experiments (see Rizzolatti et al., 1981a, b; Gentilucci et al., 1988; Fogassi et al., 1996; Gallese et al., 1996; Murata et al., 1997) leave little doubt about the areas that were inactivated in the latter monkey.”

“Although there are findings to the contrary (Fogassi et al. 1996; Godschalk et al. 1985; Graziano et al. 1997; Wise and Mauritz 1985), recent studies have shown that arm movement– related premotor activity can be modulated by the direction of gaze of the eyes in tasks in which the direction of…”

“…contributes to the defense of the body surface, neurons in this region of cortex typically respond to tactile stimuli on the face and arms and to visual stimuli looming toward the tactile receptive fields (Fogassi et al. 1996, Gentilucci et al. 1988, Graziano et al. 1997, Rizzolatti et al. 1981).”

“At least some neurons in premotor cortex, however, respond to visual stimuliresponses are most common lie outside the boundaries of the single map of postures found in our stimulation in untrained monkeys and even in anesthetized ones (Rizzolatti et al., 1981; Fogassi et al., 1996; Graziano etstudy.”

“A convergence of different sensory modalities on neurons coding motor acts has been previously described also in the premotor cortex (Gentilucci et al., 1983; Fogassi et al., 1996a, b; Graziano et al., 1997).”

“The ventral premotor cortex (area F4), the parietal areas (area 7b), and the putamen form an integrated (visual–tactile) network for the coding of peripersonal space centered on body parts (Colby, Duhamel, & Goldberg, 1993; Duhamel et al., 1991; Fogassi et al., 1996; Graziano, Hu, & Gross, 1997).”

“…receptive fields appear to be centred on the particular body part on which the tactile field lies (e.g. the hand), such that the visual and tactile receptive fields remain in correspondence even when that body part is moved separately from the eyes (Fogassi et al., 1996; Graziano & Gross, 1995).”

“, 2015a) are modulated by the direction of incoming sensory cues, with stronger response for looming, as compared to receding stimulation, and larger receptive fields when probed with faster as compared to slower stimuli (Fogassi et al., 1996; Noel et al., 2018a).”

“The spatial alignment of visual and somatosensory receptive fields allows the construction of a body-part-centered representation of the peripersonal space (Graziano and Gross, 1993; Fogassi et al., 1996).”

“Among these functions, were the coding of the goal of motor acts rather than movements that form them [7,8], the responsiveness to objects in area F5 ([9] and see [10]) and the coding of the peripersonal space in area F4 [11,12].”

“To this end, we employed a two-bytwo factorial design in which we systematically varied the type of object observed (humanoid body versus wooden object) and the timing of the visual and tactile stimuli (synchronous versus asynchronous) and computed the interaction term, which identified areas showing a greater effect…”

“These surgical procedures were the same as previously described (Fogassi et al. 1996; Rozzi et al. 2006).”

“Several studies consistently showed that premotor and parietal areas contain neurons that perceptually respond to visual, auditory and somatosensory inputs [7,8,12–19].”

“1), and abundant functional and anatomical studies support crossmodal interactions in these regions (Benevento et al. 1977; Hyvarinen and Shelepin 1979; Bruce et al. 1981; Rizzolatti et al. 1981; Hikosaka et al. 1988; Graziano et al. 1994, 1999; Cusick et al. 1995; Fogassi et al. 1996; Seltzer et al. 1996; Duhamel et…”

“Fogassi et al. (1996) showed that an increase in the velocity of approaching stimuli produced an online expansion of visual RF depth in most F4 neurons, such that fast-moving stimuli were signaled earlier than slow-moving ones.”

“, 1997) and premotor (Graziano and Gross, 1994, 1998; Fogassi et al., 1996) cortices.”

“The surgical procedures for single-neuron recordings were the same as previously described (Fogassi et al., 1996).”

“While recording from F4 neurons, Fogassi et al. (1996) and Graziano, Hu, and Gross (1997a) found that this region is involved in coding space relevant for action.”

“In either case, however, environmental features are coded as a reference frame for a particular set of effectors (Fogassi et al., 1996; Gentilucci et al., 1988; Rizzolatti et al., 1988).”

“In addition, mainly in the ventral part of the premotor cortex, there are neurons that discharge also to visual sti- Ž mulation Rizzolatti et al., 1981b, 1988; Gentilucci et al., 1983, 1988; Fogassi et al., 1996a,b; Gallese et al., 1996; Murata et al., .”

“Fogassi et al. (1996), for example, found visual RFs ranging in extent from 5 to 35 cm from tactile RFs.”

“cortex, both in areas F4 and F5, in rostral area 6 (Gentilucci et al., 1988; Rizzolatti et al., 1988; Graziano et al., 1994; Gross and Graziano, 1995; Fogassi et al., 1996).”

“In a similar vein, Fogassi et al. (1996) also found that the visual RF of F4’s visuo–tactile neurons expand when the visual stimulus velocity increases while approaching the cutaneous RF, a property that could be crucial for preparing and/or executing actions towards nearby objects.”

“effect shown for the IN sound is coherentwith previousfindings showing higher relevance of looming stimuli for PPS neurons [12–15].”

“PPS representation is particularly sensitive to stimuli approaching towards or receding from the body (Colby et al. 1993; Graziano et al. 1997; Fogassi et al. 1996).”

“Third, overlapping activation was found in the left precentral gyrus (dorsal PMv/BA 6), when all the touch observation conditions were pooled together.”

“Second, this paradigm directly resembles the stimulations used in monkey neurophysiology to study PPS bimodal or trimodal neurons, where a visual or an auditory stimulus was presented, as approaching to or receding from the animal’s body part where the neuron’s tactile receptive field was located [6,8,9,11].”

“…PM, appear to direct specific aspects of complex motor activities, including bimanual, reaching, grasping, and sequenced movements in relation to internal states or external stimuli (Kurata and Wise, 1988; Mushiake et al., 1991; Tanji and Shima, 1994; Fogassi et al., 1996; Graziano et al., 1997).”

“These visual receptive fields extend from the tactile receptive fields into adjacent space and move as a unit with it (Fogassi et al., 1996; Graziano et al., 1997).”

“All rights reserved. oi:10.1016/j.neuropsychologia.2004.11.010 of the distance of the visual stimulus from the hand somatosensory RF, increasing when the stimulus comes closer, and decreasing at farther distances (Duhamel, Colby, & Goldberg, 1998; Fogassi et al., 1996, 1999).”

“Thus, among other brain areas which contain neurons representing similar bimodal properties for the body parts in the direct environments (Leinonen et al., 1979; Leinonen and Nyman, 1979; Colby and Duhamel, 1991; Colby et al., 1993; Graziano and Gross, 1993, 1994; Fogassi et al., 1996; Graziano et al., 1997; Rizzolatt…”

“This may relate to multimodal neurons found in several areas of monkey cortex, responding to both vision and touch with spatially corresponding receptive fields in the two modalities [2,8–11].”

“, some respond more strongly to snakes than to apples) [21,25,28,46].”

“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., 1997b; Fogassi et al., 1996).”

“…is compatible with the functional and anatomical characteristics of the macaque’s premotor neurons clustered in front of the spur of the arcuate sulcus, namely, arm-centered, motionsensitive visuotactile receptive fields (Graziano & Gandhi, 2000; Graziano, Hu, & Gross, 1997; Fogassi et al., 1996).”

“Area PZ receives sensory input—its neurons respond with short latency to specific tactile, visual, and sometimes auditory stimuli (Fogassi et al. 1996; Gentilucci et al. 1988; Graziano and Gandhi 2000; Graziano et al. 1997, 1999; Rizzolatti et al. 1981).”

“Neurons coding for sectors of the peripersonal space around the face and arm show considerable overlap, though it is not clear whether at the single neurons level a shared representation of hand and mouth is present (Fogassi et al., 1996; Gentilucci et al., 1988; Graziano et al., 1994).”

“According to the effective sensory stimulus, F4 neurons were subdivided into two categories: pure somatosensory neurons and bimodal, somatosensory and visual neurons (Gentilucci et al., 1988; Graziano et al., 1994; ogassi et al., 1996).”

“Accordingly, higher-level cortical areas involved in representing the peripersonal space (Fogassi et al., 1996; Macaluso and Maravita, 2010; Legrain, 2011) and in detecting potentially dangerous stimuli near the body (Cooke and Graziano, 2003; Lloyd et al.”

“Subsequent studies revealed that premotor neurons represent visuospatial information related to a cue (Boussaoud et al. 1993; Fogassi et al. 1996; Graziano et al. 1997), an object’s three-dimensional features (Fogassi et al. 1999; Murata et al. 1997), and attention to a spatial location (Boussaoud…”