A milestone on which relies the voluntary control of behavior is the ability to shape our motor o... more A milestone on which relies the voluntary control of behavior is the ability to shape our motor output to meet the needs of the context which we are continuously facing. Even though it is solidly established that contextual information inXuence movement generation few studies have so far explored their eVects on inhibitory processes. We compared the inhibitory control of arm movements of ten healthy right-handed volunteers in a countermanding reaching paradigm with and without the presence of a temporal gap between the oVset of the central target and the peripheral target appearance. We found that this perceptual gap reduces the reaction times of hand movements and, at the same time, increases the duration of the stop process, the stop signal reaction time. The two eVects are not correlated implying that inhibition and execution of reaching movement are two independent processes inXuenced by a common factor: the disengagement of selective attention from the central target. Therefore our results support the idea of the existence of a link between spatial selective attention and inhibitory processes.
The capacity to rapidly suppress a behavioral act in response to sudden instruction to stop is a ... more The capacity to rapidly suppress a behavioral act in response to sudden instruction to stop is a key cognitive function. This function, called reactive control, is tested in experimental settings using the stop signal task, which requires subjects to generate a movement in response to a go signal or suppress it when a stop signal appears. The ability to inhibit this movement fluctuates over time: sometimes, subjects can stop their response, and at other times, they can not. To determine the neural basis of this fluctuation, we recorded local field potentials (LFPs) in the alpha (6-12 Hz) and beta (13-35 Hz) bands from the dorsal premotor cortex of two nonhuman primates that were performing the task. The ability to countermand a movement after a stop signal was predicted by the activity of both bands, each purportedly representing a distinct neural process. The beta band represents the level of movement preparation; higher beta power corresponds to a lower level of movement preparation, whereas the alpha band supports a proper phasic, reactive inhibitory response: movements are inhibited when alpha band power increases immediately after a stop signal. Our findings support the function of LFP bands in generating the signatures of various neural computations that are multiplexed in the brain.
The capacity to rapidly suppress a behavioral act in response to sudden instruction to stop is a ... more The capacity to rapidly suppress a behavioral act in response to sudden instruction to stop is a key cognitive function. This function, called reactive control, is tested in experimental settings using the stop signal task, which requires subjects to generate a movement in response to a go signal or suppress it when a stop signal appears. The ability to inhibit this movement fluctuates over time: sometimes, subjects can stop their response, and at other times, they can not. To determine the neural basis of this fluctuation, we recorded local field potentials (LFPs) in the alpha (6-12 Hz) and beta (13-35 Hz) bands from the dorsal premotor cortex of two nonhuman primates that were performing the task. The ability to countermand a movement after a stop signal was predicted by the activity of both bands, each purportedly representing a distinct neural process. The beta band represents the level of movement preparation; higher beta power corresponds to a lower level of movement preparation, whereas the alpha band supports a proper phasic, reactive inhibitory response: movements are inhibited when alpha band power increases immediately after a stop signal. Our findings support the function of LFP bands in generating the signatures of various neural computations that are multiplexed in the brain.
The Journal of neuroscience : the official journal of the Society for Neuroscience, Jan 12, 2014
The exquisite ability of primates to grasp and manipulate objects relies on the transformation of... more The exquisite ability of primates to grasp and manipulate objects relies on the transformation of visual information into motor commands. To this end, the visual system extracts object affordances that can be used to program and execute the appropriate grip. The macaque anterior intraparietal (AIP) area has been implicated in the extraction of affordances for the purpose of grasping. Neurons in the AIP area respond during visually guided grasping and to the visual presentation of objects. A subset of AIP neurons is also activated by two-dimensional images of objects and even by outline contours defining the object shape, but it is unknown how AIP neurons actually represent object shape. In this study, we used a stimulus reduction approach to determine the minimum effective shape feature evoking AIP responses. AIP neurons responding to outline shapes also responded selectively to very small fragment stimuli measuring only 1-2°. This fragment selectivity could not be explained by differences in eye movements or simple orientation selectivity, but proved to be highly dependent on the relative position of the stimulus in the receptive field. Our findings challenge the current understanding of the AIP area as a critical stage in the dorsal stream for the extraction of object affordances.
In the study of decision making, emphasis is placed on different forms of perceptual integration,... more In the study of decision making, emphasis is placed on different forms of perceptual integration, while the influence of other factors, such as memory, is ignored. In addition, it is believed that the information underlying decision making is carried in the rate of the neuronal response, while its variability is considered unspecific. Here we studied the influence of recent experience on motor decision making by analyzing the activity of neurons in the dorsal premotor area of two monkeys performing a countermanding arm task. We observe that the across-trial variability of the neural response strongly correlates with trial history-dependent changes in reaction time. Using a theoretical model of decision making, we show that a trial history-monitoring signal can explain the observed behavioral and neural modulation. Our study reveals that, in the neural processes that culminate in motor plan maturation, the evidence provided by perception and memory is reflected in mean rate and variance respectively.
Del Giudice P, Ferraina S, Wurtz RH. The control of the initiation and the inhibition of voluntar... more Del Giudice P, Ferraina S, Wurtz RH. The control of the initiation and the inhibition of voluntary movements: experimental analysis and theoretical modeling of the neural correlates at the population level. In: Garaci E, D'Addazio C, Giuliano F, ed. Programma di collaborazione ISS-NIH: ...
Cognitive functions like motor planning rely on the concerted activity of multiple neuronal assem... more Cognitive functions like motor planning rely on the concerted activity of multiple neuronal assemblies underlying still elusive computational strategies. During reaching tasks, we observed stereotyped sudden transitions (STs) between low and high multiunit activity of monkey dorsal premotor cortex (PMd) predicting forthcoming actions on a single-trial basis. Occurrence of STs was observed even when movement was delayed or successfully canceled after a stop signal, excluding a mere substrate of the motor execution. An attractor model accounts for upward STs and high-frequency modulations of field potentials, indicative of local synaptic reverberation. We found in vivo compelling evidence that motor plans in PMd emerge from the coactivation of such attractor modules, heterogeneous in the strength of local synaptic self-excitation. Modules with strong coupling early reacted with variable times to weak inputs, priming a chain reaction of both upward and downward STs in other modules. Such web of "flip-flops" rapidly converged to a stereotyped distributed representation of the motor program, as prescribed by the long-standing theory of associative networks.
Depth information is necessary for adjusting the hand to the three-dimensional (3-D) shape of an ... more Depth information is necessary for adjusting the hand to the three-dimensional (3-D) shape of an object to grasp it. The transformation of visual information into appropriate distal motor commands is critically dependent on the anterior intraparietal area (AIP) and the ventral premotor cortex (area F5), particularly the F5p sector. Recent studies have demonstrated that both AIP and the F5a sector of the ventral premotor cortex contain neurons that respond selectively to disparity-defined 3-D shape. To investigate the neural coding of 3-D shape and the behavioral role of 3-D shape-selective neurons in these two areas, we recorded single-cell activity in AIP and F5a during passive fixation of curved surfaces and during grasping of real-world objects. Similar to those in AIP, F5a neurons were either first- or second-order disparity selective, frequently showed selectivity for discrete approximations of smoothly curved surfaces that contained disparity discontinuities, and exhibited mostly monotonic tuning for the degree of disparity variation. Furthermore, in both areas, 3-D shape-selective neurons were colocalized with neurons that were active during grasping of real-world objects. Thus, area AIP and F5a contain highly similar representations of 3-D shape, which is consistent with the proposed transfer of object information from AIP to the motor system through the ventral premotor cortex.
Anatomical studies indicate that area F5 in the macaque ventral premotor cortex consists of three... more Anatomical studies indicate that area F5 in the macaque ventral premotor cortex consists of three different sectors. One of these is F5a in the posterior bank of the inferior arcuate sulcus, but no functional characterization of F5a at the single-cell level exists. We investigated the neuronal selectivity for three-dimensional (3D) shape and grasping activity in F5a. In contrast to neighboring regions F5p and 45B, the great majority of F5a neurons showed selectivity for disparity-defined curved surfaces, and most neurons preserved this selectivity across positions in depth, indicating higher-order disparity selectivity. Thus, as predicted by monkey fMRI data, F5a neurons showed robust 3D-shape selectivity in the absence of a motor response. To investigate the relationship between disparity selectivity and grasping activity, we recorded from 3D-shape-selective F5a neurons during a visually guided grasping task and during grasping in the dark. F5a neurons encoding the depth profile of curved surfaces frequently responded during grasping of real-world objects in the light, but not in the dark, whereas nearby neurons were also active in the dark. The presence of 3D-shape-selective and "visual-dominant" neurons demonstrates that the F5a sector is distinct from neighboring regions of ventral premotor cortex, in line with recent anatomical connectivity studies.
A milestone on which relies the voluntary control of behavior is the ability to shape our motor o... more A milestone on which relies the voluntary control of behavior is the ability to shape our motor output to meet the needs of the context which we are continuously facing. Even though it is solidly established that contextual information inXuence movement generation few studies have so far explored their eVects on inhibitory processes. We compared the inhibitory control of arm movements of ten healthy right-handed volunteers in a countermanding reaching paradigm with and without the presence of a temporal gap between the oVset of the central target and the peripheral target appearance. We found that this perceptual gap reduces the reaction times of hand movements and, at the same time, increases the duration of the stop process, the stop signal reaction time. The two eVects are not correlated implying that inhibition and execution of reaching movement are two independent processes inXuenced by a common factor: the disengagement of selective attention from the central target. Therefore our results support the idea of the existence of a link between spatial selective attention and inhibitory processes.
The ability of rapidly adapting our motor behaviour in order to face the unpredictable changes in... more The ability of rapidly adapting our motor behaviour in order to face the unpredictable changes in the surrounding environment is fundamental for survival. To achieve such a high level of efficiency our motor system has to assess continuously the context in which it acts, gathering all available information that can be relevant for planning goal-oriented movements. One still-debated aspect of movement organization is the nature and timing of motor planning. While motor plans are often taken to be concerned with the setting of kinematic parameters as a function of perceptual and motor factors, it has been suggested that higher level, cognitive factors may also affect planning. To explore this issue further, we asked 18 right-handed human participants to perform speeded hand-reaching movement toward a visual target in two different experimental settings, a reaction time (RT) paradigm (go-only task) and a countermanding paradigm. In both tasks participants executed the same movements, but in the countermanding task no-stop trials were randomly intermixed with stop trials. In stop trials participants were required to withhold the ongoing movement whenever a stop signal was shown. It is known that the presence of stop trials induces a consistent increase of the RTs of no-stop trials with respect to the RTs of go-only trials. However, nothing is known about a similar effect for movement times (MTs). We found that RTs and MTs exhibit opposing tendencies, so that a decrease in the RT correspond to an increase in the MT and vice versa. This tendency was present in all our participants and significant in 90% of them. Furthermore we found a moderate, but again very consistent, anticorrelation between RTs and MTs on a trial-by-trial base. These findings are consistent with strategic changes in movement programmes for the very same movements under different cognitive contexts, requiring different degrees of feedback-driven control during movement.
Canceling a pending movement is a hallmark of voluntary behavioral control because it allows us t... more Canceling a pending movement is a hallmark of voluntary behavioral control because it allows us to quickly adapt to unattended changes either in the external environment or in our thoughts. The countermanding paradigm allows the study of inhibitory processes of motor acts by requiring the subject
to withhold planned movements in response to an infrequent stopsignal. At present the neural processes underlying the inhibitory control of arm movements are mostly unknown. We recorded the activity of single units in the rostral and caudal portion of the dorsal premotor cortex (PMd) of monkeys trained in a countermanding reaching task. We found that among neurons with a movement preparatory activity, about one-third exhibit a modulation before the behavioral estimate of the time it takes to cancel a planned movement. Hence these neurons exhibit a pattern of activity suggesting that PMd plays a critical role in the brain networks involved in the control of
arm movement initiation and suppression.
A milestone on which relies the voluntary control of behavior is the ability to shape our motor o... more A milestone on which relies the voluntary control of behavior is the ability to shape our motor output to meet the needs of the context which we are continuously facing. Even though it is solidly established that contextual information inXuence movement generation few studies have so far explored their eVects on inhibitory processes. We compared the inhibitory control of arm movements of ten healthy right-handed volunteers in a countermanding reaching paradigm with and without the presence of a temporal gap between the oVset of the central target and the peripheral target appearance. We found that this perceptual gap reduces the reaction times of hand movements and, at the same time, increases the duration of the stop process, the stop signal reaction time. The two eVects are not correlated implying that inhibition and execution of reaching movement are two independent processes inXuenced by a common factor: the disengagement of selective attention from the central target. Therefore our results support the idea of the existence of a link between spatial selective attention and inhibitory processes.
The capacity to rapidly suppress a behavioral act in response to sudden instruction to stop is a ... more The capacity to rapidly suppress a behavioral act in response to sudden instruction to stop is a key cognitive function. This function, called reactive control, is tested in experimental settings using the stop signal task, which requires subjects to generate a movement in response to a go signal or suppress it when a stop signal appears. The ability to inhibit this movement fluctuates over time: sometimes, subjects can stop their response, and at other times, they can not. To determine the neural basis of this fluctuation, we recorded local field potentials (LFPs) in the alpha (6-12 Hz) and beta (13-35 Hz) bands from the dorsal premotor cortex of two nonhuman primates that were performing the task. The ability to countermand a movement after a stop signal was predicted by the activity of both bands, each purportedly representing a distinct neural process. The beta band represents the level of movement preparation; higher beta power corresponds to a lower level of movement preparation, whereas the alpha band supports a proper phasic, reactive inhibitory response: movements are inhibited when alpha band power increases immediately after a stop signal. Our findings support the function of LFP bands in generating the signatures of various neural computations that are multiplexed in the brain.
The capacity to rapidly suppress a behavioral act in response to sudden instruction to stop is a ... more The capacity to rapidly suppress a behavioral act in response to sudden instruction to stop is a key cognitive function. This function, called reactive control, is tested in experimental settings using the stop signal task, which requires subjects to generate a movement in response to a go signal or suppress it when a stop signal appears. The ability to inhibit this movement fluctuates over time: sometimes, subjects can stop their response, and at other times, they can not. To determine the neural basis of this fluctuation, we recorded local field potentials (LFPs) in the alpha (6-12 Hz) and beta (13-35 Hz) bands from the dorsal premotor cortex of two nonhuman primates that were performing the task. The ability to countermand a movement after a stop signal was predicted by the activity of both bands, each purportedly representing a distinct neural process. The beta band represents the level of movement preparation; higher beta power corresponds to a lower level of movement preparation, whereas the alpha band supports a proper phasic, reactive inhibitory response: movements are inhibited when alpha band power increases immediately after a stop signal. Our findings support the function of LFP bands in generating the signatures of various neural computations that are multiplexed in the brain.
The Journal of neuroscience : the official journal of the Society for Neuroscience, Jan 12, 2014
The exquisite ability of primates to grasp and manipulate objects relies on the transformation of... more The exquisite ability of primates to grasp and manipulate objects relies on the transformation of visual information into motor commands. To this end, the visual system extracts object affordances that can be used to program and execute the appropriate grip. The macaque anterior intraparietal (AIP) area has been implicated in the extraction of affordances for the purpose of grasping. Neurons in the AIP area respond during visually guided grasping and to the visual presentation of objects. A subset of AIP neurons is also activated by two-dimensional images of objects and even by outline contours defining the object shape, but it is unknown how AIP neurons actually represent object shape. In this study, we used a stimulus reduction approach to determine the minimum effective shape feature evoking AIP responses. AIP neurons responding to outline shapes also responded selectively to very small fragment stimuli measuring only 1-2°. This fragment selectivity could not be explained by differences in eye movements or simple orientation selectivity, but proved to be highly dependent on the relative position of the stimulus in the receptive field. Our findings challenge the current understanding of the AIP area as a critical stage in the dorsal stream for the extraction of object affordances.
In the study of decision making, emphasis is placed on different forms of perceptual integration,... more In the study of decision making, emphasis is placed on different forms of perceptual integration, while the influence of other factors, such as memory, is ignored. In addition, it is believed that the information underlying decision making is carried in the rate of the neuronal response, while its variability is considered unspecific. Here we studied the influence of recent experience on motor decision making by analyzing the activity of neurons in the dorsal premotor area of two monkeys performing a countermanding arm task. We observe that the across-trial variability of the neural response strongly correlates with trial history-dependent changes in reaction time. Using a theoretical model of decision making, we show that a trial history-monitoring signal can explain the observed behavioral and neural modulation. Our study reveals that, in the neural processes that culminate in motor plan maturation, the evidence provided by perception and memory is reflected in mean rate and variance respectively.
Del Giudice P, Ferraina S, Wurtz RH. The control of the initiation and the inhibition of voluntar... more Del Giudice P, Ferraina S, Wurtz RH. The control of the initiation and the inhibition of voluntary movements: experimental analysis and theoretical modeling of the neural correlates at the population level. In: Garaci E, D'Addazio C, Giuliano F, ed. Programma di collaborazione ISS-NIH: ...
Cognitive functions like motor planning rely on the concerted activity of multiple neuronal assem... more Cognitive functions like motor planning rely on the concerted activity of multiple neuronal assemblies underlying still elusive computational strategies. During reaching tasks, we observed stereotyped sudden transitions (STs) between low and high multiunit activity of monkey dorsal premotor cortex (PMd) predicting forthcoming actions on a single-trial basis. Occurrence of STs was observed even when movement was delayed or successfully canceled after a stop signal, excluding a mere substrate of the motor execution. An attractor model accounts for upward STs and high-frequency modulations of field potentials, indicative of local synaptic reverberation. We found in vivo compelling evidence that motor plans in PMd emerge from the coactivation of such attractor modules, heterogeneous in the strength of local synaptic self-excitation. Modules with strong coupling early reacted with variable times to weak inputs, priming a chain reaction of both upward and downward STs in other modules. Such web of "flip-flops" rapidly converged to a stereotyped distributed representation of the motor program, as prescribed by the long-standing theory of associative networks.
Depth information is necessary for adjusting the hand to the three-dimensional (3-D) shape of an ... more Depth information is necessary for adjusting the hand to the three-dimensional (3-D) shape of an object to grasp it. The transformation of visual information into appropriate distal motor commands is critically dependent on the anterior intraparietal area (AIP) and the ventral premotor cortex (area F5), particularly the F5p sector. Recent studies have demonstrated that both AIP and the F5a sector of the ventral premotor cortex contain neurons that respond selectively to disparity-defined 3-D shape. To investigate the neural coding of 3-D shape and the behavioral role of 3-D shape-selective neurons in these two areas, we recorded single-cell activity in AIP and F5a during passive fixation of curved surfaces and during grasping of real-world objects. Similar to those in AIP, F5a neurons were either first- or second-order disparity selective, frequently showed selectivity for discrete approximations of smoothly curved surfaces that contained disparity discontinuities, and exhibited mostly monotonic tuning for the degree of disparity variation. Furthermore, in both areas, 3-D shape-selective neurons were colocalized with neurons that were active during grasping of real-world objects. Thus, area AIP and F5a contain highly similar representations of 3-D shape, which is consistent with the proposed transfer of object information from AIP to the motor system through the ventral premotor cortex.
Anatomical studies indicate that area F5 in the macaque ventral premotor cortex consists of three... more Anatomical studies indicate that area F5 in the macaque ventral premotor cortex consists of three different sectors. One of these is F5a in the posterior bank of the inferior arcuate sulcus, but no functional characterization of F5a at the single-cell level exists. We investigated the neuronal selectivity for three-dimensional (3D) shape and grasping activity in F5a. In contrast to neighboring regions F5p and 45B, the great majority of F5a neurons showed selectivity for disparity-defined curved surfaces, and most neurons preserved this selectivity across positions in depth, indicating higher-order disparity selectivity. Thus, as predicted by monkey fMRI data, F5a neurons showed robust 3D-shape selectivity in the absence of a motor response. To investigate the relationship between disparity selectivity and grasping activity, we recorded from 3D-shape-selective F5a neurons during a visually guided grasping task and during grasping in the dark. F5a neurons encoding the depth profile of curved surfaces frequently responded during grasping of real-world objects in the light, but not in the dark, whereas nearby neurons were also active in the dark. The presence of 3D-shape-selective and "visual-dominant" neurons demonstrates that the F5a sector is distinct from neighboring regions of ventral premotor cortex, in line with recent anatomical connectivity studies.
A milestone on which relies the voluntary control of behavior is the ability to shape our motor o... more A milestone on which relies the voluntary control of behavior is the ability to shape our motor output to meet the needs of the context which we are continuously facing. Even though it is solidly established that contextual information inXuence movement generation few studies have so far explored their eVects on inhibitory processes. We compared the inhibitory control of arm movements of ten healthy right-handed volunteers in a countermanding reaching paradigm with and without the presence of a temporal gap between the oVset of the central target and the peripheral target appearance. We found that this perceptual gap reduces the reaction times of hand movements and, at the same time, increases the duration of the stop process, the stop signal reaction time. The two eVects are not correlated implying that inhibition and execution of reaching movement are two independent processes inXuenced by a common factor: the disengagement of selective attention from the central target. Therefore our results support the idea of the existence of a link between spatial selective attention and inhibitory processes.
The ability of rapidly adapting our motor behaviour in order to face the unpredictable changes in... more The ability of rapidly adapting our motor behaviour in order to face the unpredictable changes in the surrounding environment is fundamental for survival. To achieve such a high level of efficiency our motor system has to assess continuously the context in which it acts, gathering all available information that can be relevant for planning goal-oriented movements. One still-debated aspect of movement organization is the nature and timing of motor planning. While motor plans are often taken to be concerned with the setting of kinematic parameters as a function of perceptual and motor factors, it has been suggested that higher level, cognitive factors may also affect planning. To explore this issue further, we asked 18 right-handed human participants to perform speeded hand-reaching movement toward a visual target in two different experimental settings, a reaction time (RT) paradigm (go-only task) and a countermanding paradigm. In both tasks participants executed the same movements, but in the countermanding task no-stop trials were randomly intermixed with stop trials. In stop trials participants were required to withhold the ongoing movement whenever a stop signal was shown. It is known that the presence of stop trials induces a consistent increase of the RTs of no-stop trials with respect to the RTs of go-only trials. However, nothing is known about a similar effect for movement times (MTs). We found that RTs and MTs exhibit opposing tendencies, so that a decrease in the RT correspond to an increase in the MT and vice versa. This tendency was present in all our participants and significant in 90% of them. Furthermore we found a moderate, but again very consistent, anticorrelation between RTs and MTs on a trial-by-trial base. These findings are consistent with strategic changes in movement programmes for the very same movements under different cognitive contexts, requiring different degrees of feedback-driven control during movement.
Canceling a pending movement is a hallmark of voluntary behavioral control because it allows us t... more Canceling a pending movement is a hallmark of voluntary behavioral control because it allows us to quickly adapt to unattended changes either in the external environment or in our thoughts. The countermanding paradigm allows the study of inhibitory processes of motor acts by requiring the subject
to withhold planned movements in response to an infrequent stopsignal. At present the neural processes underlying the inhibitory control of arm movements are mostly unknown. We recorded the activity of single units in the rostral and caudal portion of the dorsal premotor cortex (PMd) of monkeys trained in a countermanding reaching task. We found that among neurons with a movement preparatory activity, about one-third exhibit a modulation before the behavioral estimate of the time it takes to cancel a planned movement. Hence these neurons exhibit a pattern of activity suggesting that PMd plays a critical role in the brain networks involved in the control of
arm movement initiation and suppression.
Uploads
Papers by Pierpaolo Pani
to withhold planned movements in response to an infrequent stopsignal. At present the neural processes underlying the inhibitory control of arm movements are mostly unknown. We recorded the activity of single units in the rostral and caudal portion of the dorsal premotor cortex (PMd) of monkeys trained in a countermanding reaching task. We found that among neurons with a movement preparatory activity, about one-third exhibit a modulation before the behavioral estimate of the time it takes to cancel a planned movement. Hence these neurons exhibit a pattern of activity suggesting that PMd plays a critical role in the brain networks involved in the control of
arm movement initiation and suppression.
to withhold planned movements in response to an infrequent stopsignal. At present the neural processes underlying the inhibitory control of arm movements are mostly unknown. We recorded the activity of single units in the rostral and caudal portion of the dorsal premotor cortex (PMd) of monkeys trained in a countermanding reaching task. We found that among neurons with a movement preparatory activity, about one-third exhibit a modulation before the behavioral estimate of the time it takes to cancel a planned movement. Hence these neurons exhibit a pattern of activity suggesting that PMd plays a critical role in the brain networks involved in the control of
arm movement initiation and suppression.