Visual and Nonvisual Contributions to Three-Dimensional Heading Selectivity in the Medial Superior Temporal Area

“It can be shown that such a network is capable of estimating an arbitrary function, given a sufficient number of hidden units (Bishop, 1995).”

“The overall heading estimate of the network, resulting from the activity of the output units, was defined by a population vector (Georgopoulos et al., 1986) and was computed as the vector sum of the responses of the 26 output units.”

“Both monkeys were also implanted with scleral coils for measuring eye movements in a magnetic field (Robinson, 1963).”

“As demonstrated previously by Zipser and Andersen (1988), the transformation of optic flow signals from an eye-centered to head-centered reference frame was implemented in the hidden layer of our model through modulation of hidden unit responses by eye position (i.”

“The changes in receptive field location of MT neurons across guide tube locations were as expected from the known topography of MT (Zeki, 1974; Gattass and Gross, 1981; Van Essen et al., 1981; Desimone and Ungerleider, 1986; Albright and Desimone, 1987; Maunsell and Van Essen, 1987).”

“…area, area 7a, and superior temporal polysensory area] are involved in coding patterns of optic flow that typically result from self-motion (Tanaka et al., 1986, 1989; Duffy and Wurtz, 1991; Schaafsma and Duysens, 1996; Siegel and Read, 1997; Anderson and Siegel, 1999; Bremmer et al., 2002a,b).”

“…area, area 7a, and superior temporal polysensory area] are involved in coding patterns of optic flow that typically result from self-motion (Tanaka et al., 1986, 1989; Duffy and Wurtz, 1991; Schaafsma and Duysens, 1996; Siegel and Read, 1997; Anderson and Siegel, 1999; Bremmer et al.,…”

“, 1972 ; Fernandez and Goldberg , 1976a , b ) .”

“The changes in receptive field location of MT neurons across guide tube locations were as expected from the known topography of MT (Zeki, 1974; Gattass and Gross, 1981; Van Essen et al., 1981; Desimone and Ungerleider, 1986; Albright and Desimone, 1987; Maunsell and Van Essen, 1987).”

“The average RF size was 44 Ϯ 8 ϫ 58 Ϯ 13° SE, which is similar to RF sizes reported previously for MSTd (Van Essen et al., 1981; Desimone and Ungerleider, 1986; Komatsu and Wurtz, 1988a).”

“MT neurons were identified according to several properties, including smaller receptive fields (diameter eccentricity), sensitivity to small visual stimuli as well as large stimuli, and similar direction preferences within penetrations approximately normal to the cortical layers (Albright et al., 1984).”

“The changes in receptive field location of MT neurons across guide tube locations were as expected from the known topography of MT (Zeki, 1974; Gattass and Gross, 1981; Van Essen et al., 1981; Desimone and Ungerleider, 1986; Albright and Desimone, 1987; Maunsell and Van Essen, 1987).”

“These findings extend previous work done in MSTd using 1D or 2D stimulation (Duffy, 1998; Bremmer et al., 1999). In an experiment involving 1D (fore-aft) motion on a parallel swing, Bremmer et al. (1999) reported that approximately half of MSTd neurons showed opposite heading preferences for visual and vestibular stim…”

“The changes in receptive field location of MT neurons across guide tube locations were as expected from the known topography of MT (Zeki, 1974; Gattass and Gross, 1981; Van Essen et al., 1981; Desimone and Ungerleider, 1986; Albright and Desimone, 1987; Maunsell and Van Essen, 1987).”

“The MRI data were converted to a flat map using CARET software (Van Essen et al., 2001), and the flat map was morphed to match a standard macaque atlas.”

“…area, area 7a, and superior temporal polysensory area] are involved in coding patterns of optic flow that typically result from self-motion (Tanaka et al., 1986, 1989; Duffy and Wurtz, 1991; Schaafsma and Duysens, 1996; Siegel and Read, 1997; Anderson and Siegel, 1999; Bremmer et al.,…”

“The dorsal subdivision of the medial superior temporal area (MSTd) has been a main focus of investigation, because single neurons in MSTd appear well suited to signal heading based on optic flow (Duffy and Wurtz, 1995).”

“Although our data do not allow us to make strong conclusions regarding hemispheric differences, they are consistent with findings of a right-hemispheric dominance of vestibular responses in functional MRI (fMRI) experiments involving caloric stimulation (Fasold et al., 2002).”

“…object motion all confound the optic flow that results from head translation, such that visual information alone is not always sufficient to judge heading accurately (Warren and Hannon, 1990; Royden et al., 1992; Royden, 1994; Banks et al., 1996; Royden and Hildreth, 1996; Crowell et al., 1998).”

“However, eye movements, head movements, and object motion all confound the optic flow that results from head translation, such that visual information alone is not always sufficient to judge heading accurately (Warren and Hannon, 1990; Royden et al., 1992; Royden, 1994; Banks et al., 1996; Royden and Hildreth, 1996; C…”

“In contrast , visual motion is thought to be encoded in terms of velocity signals ( Rodman and Albright , 1987 ; Lisberger and Movshon , 1999 ) ( but see Cao et al .”

“In addition, electrical microstimulation of MSTd can bias monkeys' heading percepts based on optic flow (Britten and van Wezel, 1998, 2002), and lesions to the human homolog of MST can seriously impair one's ability to navigate using optic flow (Vaina, 1998).”

“Recent studies have also shown that MSTd neurons respond to translation of the body in darkness, suggesting that they might integrate visual and vestibular signals to code heading more robustly (Duffy, 1998; Bremmer et al., 1999; Page and Duffy, 2003).”

“The changes in receptive field location of MT neurons across guide tube locations were as expected from the known topography of MT (Zeki, 1974; Gattass and Gross, 1981; Van Essen et al., 1981; Desimone and Ungerleider, 1986; Albright and Desimone, 1987; Maunsell and Van Essen, 1987).”

“Second , MSTd responses during iner - tial motion in the absence of optic flow might also arise from skin receptors and body proprioceptors ( for review , see Lackner and DiZio , 2005 ) .”

“, 1972 ; Fernandez and Goldberg , 1976a , b ) .”

“In contrast , visual motion is thought to be encoded in terms of velocity signals ( Rodman and Albright , 1987 ; Lisberger and Movshon , 1999 ) ( but see Cao et al .”

“…area, area 7a, and superior temporal polysensory area] are involved in coding patterns of optic flow that typically result from self-motion (Tanaka et al., 1986, 1989; Duffy and Wurtz, 1991; Schaafsma and Duysens, 1996; Siegel and Read, 1997; Anderson and Siegel, 1999; Bremmer et al., 2002a,b).”

“It is often assumed that optic flow signals must first be converted into a headcentered reference frame, before being useful for heading perception (Royden, 1994; Royden et al., 1994; Banks et al., 1996).”

“eye and head movements (Bradley et al., 1996; Page and Duffy, 1999; Shenoy et al., 1999, 2002; Ilg et al., 2004), this does not necessarily indicate that MST codes motion in a head- or worldcentered reference frame (i.”

“Vestibular contributions to heading perception have not been studied extensively , but there is evidence that humans integrate visual and vestibular signals to estimate heading more robustly (Telford et al., 1995; Ohmi, 1996; Harris et al., 2000; Bertin and Berthoz, 2004).”

“The changes in receptive field location of MT neurons across guide tube locations were as expected from the known topography of MT (Zeki, 1974; Gattass and Gross, 1981; Van Essen et al., 1981; Desimone and Ungerleider, 1986; Albright and Desimone, 1987; Maunsell and Van Essen, 1987).”

“The reference frame for heading selectivity based on optic flow in MSTd has not yet been examined during static fixation at different eye positions, although MSTd receptive fields have been reported to be primarily eye centered with gain modulations (Squatrito and Maioli, 1996, 1997; Bremmer et al., 1997).”

“…area, area 7a, and superior temporal polysensory area] are involved in coding patterns of optic flow that typically result from self-motion (Tanaka et al., 1986, 1989; Duffy and Wurtz, 1991; Schaafsma and Duysens, 1996; Siegel and Read, 1997; Anderson and Siegel, 1999; Bremmer et al., 2002a,b).”

“However, eye movements, head movements, and object motion all confound the optic flow that results from head translation, such that visual information alone is not always sufficient to judge heading accurately (Warren and Hannon, 1990; Royden et al., 1992; Royden, 1994; Banks et al., 1996; Royden and Hildreth, 1996; C…”

“However, eye movements, head movements, and object motion all confound the optic flow that results from head translation, such that visual information alone is not always sufficient to judge heading accurately (Warren and Hannon, 1990; Royden et al., 1992; Royden, 1994; Banks et al., 1996; Royden and Hildreth, 1996; C…”

“…area, area 7a, and superior temporal polysensory area] are involved in coding patterns of optic flow that typically result from self-motion (Tanaka et al., 1986, 1989; Duffy and Wurtz, 1991; Schaafsma and Duysens, 1996; Siegel and Read, 1997; Anderson and Siegel, 1999; Bremmer et al., 2002a,b).”

“Recent studies have also shown that MSTd neurons respond to translation of the body in darkness, suggesting that they might integrate visual and vestibular signals to code heading more robustly (Duffy, 1998; Bremmer et al., 1999; Page and Duffy, 2003).”

“8 mm apart) that allowed insertion of microelectrodes vertically into the brain via transdural guide tubes that were passed through a small burr hole in the skull (Dickman and Angelaki, 2002).”

“These properties are typical of neurons in area MSTd and distinct from the lateral subdivision of area MST (Komatsu and Wurtz, 1988a,b; Tanaka et al., 1993).”

“These findings extend previous work done in MSTd using 1D or 2D stimulation (Duffy, 1998; Bremmer et al., 1999).”

“For example , the observed responses could be driven by a pursuit - like signal related to suppression of the vestibulo - ocular reflex ( VOR ) ( Thier and Erickson , 1992a , b ; Fukushima et al .”

“However, eye movements, head movements, and object motion all confound the optic flow that results from head translation, such that visual information alone is not always sufficient to judge heading accurately (Warren and Hannon, 1990; Royden et al., 1992; Royden, 1994; Banks et al., 1996; Royden and Hildreth, 1996; C…”

“eye and head movements (Bradley et al., 1996; Page and Duffy, 1999; Shenoy et al., 1999, 2002; Ilg et al., 2004), this does not necessarily indicate that MST codes motion in a head- or worldcentered reference frame (i.”

“Vestibular contributions to heading perception have not been studied extensively , but there is evidence that humans integrate visual and vestibular signals to estimate heading more robustly (Telford et al., 1995; Ohmi, 1996; Harris et al., 2000; Bertin and Berthoz, 2004).”

“These results suggest that heading tuning in the vestibular condition reflects sensory signals that arise from vestibular and / or proprioceptive inputs rather than a VOR suppression or pursuit - like signal ( see Discussion ) .”

“, 1997), whereas vestibular signals are head/body centered (Shaikh et al., 2004).”

“Because changes in vergence angle can be elicited by radial optic flow under some circumstances (Busettini et al., 1997), we examined how vergence angle depended on heading direction within the horizontal place (eight azimuth angles, 45° apart).”

“…area, area 7a, and superior temporal polysensory area] are involved in coding patterns of optic flow that typically result from self-motion (Tanaka et al., 1986, 1989; Duffy and Wurtz, 1991; Schaafsma and Duysens, 1996; Siegel and Read, 1997; Anderson and Siegel, 1999; Bremmer et al., 2002a,b).”

“eye and head movements (Bradley et al., 1996; Page and Duffy, 1999; Shenoy et al., 1999, 2002; Ilg et al., 2004), this does not necessarily indicate that MST codes motion in a head- or worldcentered reference frame (i.”

“It is often as - sumed that optic flow signals must first be converted into a head - centered reference frame , before being useful for heading perception ( Royden , 1994 ; Royden et al .”

“Vestibular contributions to heading perception have not been studied extensively , but there is evidence that humans integrate visual and vestibular signals to estimate heading more robustly (Telford et al., 1995; Ohmi, 1996; Harris et al., 2000; Bertin and Berthoz, 2004).”

“For example , the observed responses could be driven by a pursuit - like signal related to suppression of the vestibulo - ocular reflex ( VOR ) ( Thier and Erickson , 1992a , b ; Fukushima et al .”

“Vestibular contributions to heading perception have not been studied extensively , but there is evidence that humans integrate visual and vestibular signals to estimate heading more robustly (Telford et al., 1995; Ohmi, 1996; Harris et al., 2000; Bertin and Berthoz, 2004).”

“Specifically , visual motion signals originate in retinal coordinates ( Squatrito and Maioli , 1996 , 1997 ; Bremmer et al .”

“In addition, electrical microstimulation of MSTd can bias monkeys' heading percepts based on optic flow (Britten and van Wezel, 1998, 2002), and lesions to the human homolog of MST can seriously impair one's ability to navigate using optic flow (Vaina, 1998).”

“Specifically , visual motion signals originate in retinal coordinates ( Squatrito and Maioli , 1996 , 1997 ; Bremmer et al .”

“For inertial motion, these accelerations far exceed vestibular thresholds (for review, see Gundry, 1978) (Benson et al., 1986; Kingma, 2005).”

“First, reference frames for visual and vestibular heading signals in MSTd remain distinct, although evidence clearly shows that these neurons might mediate multisensory cue integration (Gu et al. 2006b).”

“Details of the apparatus and stimuli (Gu et al., 2006), as well as the basic task design and training (Gu et al.”