Neural Underpinnings of Learning in Dementia Populations: A Review of Motor Learning Studies Combined with Neuroimaging.

The intent of this review article is to serve as an overview of current research regarding the neural characteristics of motor learning in Alzheimer disease (AD) as well as prodromal phases of AD: at-risk populations, and mild cognitive impairment. This review seeks to provide a cognitive framework to compare various motor tasks. We will highlight the neural characteristics related to cognitive domains that, through imaging, display functional or structural changes because of AD progression. In turn, this motivates the use of motor learning paradigms as possible screening techniques for AD and will build upon our current understanding of learning abilities in AD populations.

Motion state-dependent motor learning based on explicit visual feedback has limited spatiotemporal properties compared with adaptation to physical perturbations.

We recently showed that subjects can learn motion state-dependent changes to motor output (temporal force patterns) based on explicit visual feedback of the equivalent force field (i.e., without the physical perturbation). Here, we examined the spatiotemporal properties of this learning compared with learning based on physical perturbations. There were two human subject groups and two experimental paradigms. One group (n = 40) experienced physical perturbations (i.e., a velocity-dependent force field, vFF), whereas the second (n = 40) was given explicit visual feedback (EVF) of the force-velocity relationship. In the latter, subjects moved in force channels and we provided visual feedback of the lateral force exerted during the movement, as well as the required force pattern based on movement velocity. In the first paradigm (spatial generalization), following vFF or EVF training, generalization of learning was tested by requiring subjects to move to 14 untrained target locations (0° to ±135° around the trained location). In the second paradigm (temporal stability), following training, we examined the decay of learning over eight delay periods (0 to 90 s). Results showed that learning based on EVF did not generalize to untrained directions, whereas the generalization for the vFF was significant for targets ≤ 45° away. In addition, the decay of learning for the EVF group was significantly faster than the FF group (a time constant of 2.72 ± 1.74 s vs. 12.53 ± 11.83 s). Collectively, our results suggest that recalibrating motor output based on explicit motion state information, in contrast to physical disturbances, uses learning mechanisms with limited spatiotemporal properties.NEW & NOTEWORTHY Adjustment of motor output based on limb motion state information can be achieved based on explicit information or from physical perturbations. Here, we investigated the spatiotemporal characteristics of short-term motor learning to determine the properties of the respective learning mechanisms. Our results suggest that adjustments based on physical perturbations are more temporally stable and applied over a greater spatial range than the learning based on explicit visual feedback, suggesting largely separate learning mechanisms.

Moving a missing hand: children born with below elbow deficiency can enact hand grasp patterns with their residual muscles.

Children with a unilateral congenital below elbow deficiency (UCBED) have one typical upper limb and one that lacks a hand, ending below the elbow at the proximal/mid forearm. UCBED is an isolated condition, and affected children otherwise develop normal sensorimotor control. Unlike adults with upper limb absence, the majority of whom have an acquired loss, children with UCBED never developed a hand, so their residual muscles have never actuated an intact limb. Their ability to purposefully modulate affected muscle activity is often assumed to be limited, and this assumption has influenced prosthetic design and prescription practices for this population as many modern devices derive control signals from affected muscle activity. To better understand the motor capabilities of the affected muscles, we used ultrasound imaging to study 6 children with UCBED. We examined the extent to which subjects activate their affected muscles when performing mirrored movements with their typical and missing hands. We demonstrate that all subjects could intentionally and consistently enact at least five distinct muscle patterns when attempting different missing hand movements (e.g., power grasp) and found similar performance across affected and typically developed limbs. These results suggest that although participants had never actuated the missing hand they could distinctively and consistently activate the residual muscle patterns associated with actions on the unaffected side. These findings indicate that motor control still develops in the absence of the normal effector, and can serve as a guide for developing prostheses that leverage the full extent of these children's motor control capabilities.

Motion state-dependent motor learning based on explicit visual feedback is quickly recalled, but is less stable than adaptation to physical perturbations.

Recent studies have shown that adaptation to visual feedback perturbations during arm reaching movements involves implicit and explicit learning components. Evidence also suggests that explicit, intentional learning mechanisms are largely responsible for savings-a faster recalibration compared with initial training. However, the extent explicit learning mechanisms facilitate learning and early savings (i.e., the rapid recall of previous performance) for motion state-dependent learning is generally unknown. To address this question, we compared the early savings/recall achieved by two groups of human subjects. One experienced physical perturbations (a velocity-dependent force-field, vFF) to promote adaptation that is thought to be a largely implicit process. The second was only given visual feedback of the required force-velocity relationship; subjects moved in force channels and we provided visual feedback of the lateral force exerted during the movement, as well as the required force pattern based on the movement velocity. Thus, subjects were shown explicit information on the extent the applied temporal pattern of force matched the required velocity-dependent force profile if the force-field perturbation had been applied. After training, both groups experienced a decay and washout period, which was followed by a reexposure block to assess early savings/recall. Although decay was faster for the explicit visual feedback group, the single-trial recall was similar to the physical perturbation group. Thus, compared with visual feedback perturbations, conscious modification of motor output based on motion state-dependent feedback demonstrates rapid recall, but this adjustment is less stable than adaptation based on experiencing the multisensory errors that accompany physical perturbations.NEW & NOTEWORTHY The extent explicit feedback facilitates motion state-dependent changes to motor output is largely unknown. Here, we examined motor adaptation for subjects that experienced physical perturbations and another that made adjustments based on explicit visual feedback information of the required force-velocity relationship. Our results suggest that adjustment of motor output can be based on explicit motion state-dependent information and demonstrates rapid recall, but this learning is less stable than adaptation based on physical perturbations to movement.

Transsaccadic visual perception of foveal compared to peripheral environmental changes.

The maintenance of stable visual perception across eye movements is hypothesized to be aided by extraretinal information (e.g., corollary discharge [CD]). Previous studies have focused on the benefits of this information for perception at the fovea. However, there is little information on the extent that CD benefits peripheral visual perception. Here we systematically examined the extent that CD supports the ability to perceive transsaccadic changes at the fovea compared to peripheral changes. Human subjects made saccades to targets positioned at different amplitudes (4° or 8°) and directions (rightward or upward). On each trial there was a reference point located either at (fovea) or 4° away (periphery) from the target. During the saccade the target and reference disappeared and, after a blank period, the reference reappeared at a shifted location. Subjects reported the perceived shift direction, and we determined the perceptual threshold for detection and estimate of the reference location. We also simulated the detection and location if subjects solely relied on the visual error of the shifted reference experienced after the saccade. The comparison of the reference location under these two conditions showed that overall the perceptual estimate was approximately 53% more accurate and 30% less variable than estimates based solely on visual information at the fovea. These values for peripheral shifts were consistently lower than that at the fovea: 34% more accurate and 9% less variable. Overall, the results suggest that CD information does support stable visual perception in the periphery, but is consistently less beneficial compared to the fovea.

Dissociating effects of error size, training duration, and amount of adaptation on the ability to retain motor memories.

Extensive computational and neurobiological work has focused on how the training schedule, i.e., the duration and rate at which an environmental disturbance is presented, shapes the formation of motor memories. If long-lasting benefits are to be derived from motor training, however, retention of the performance improvements gained during practice is essential. Thus a better understanding of mechanisms that promote retention could lead to the design of more effective training procedures. The few studies that have investigated how retention depends on the training schedule have suggested that the gradual exposure of a perturbation leads to improved retention of motor memory compared with an abrupt exposure. However, several of these previous studies showed small effects, and although some controlled the training duration and others the level of learning, none have controlled both. In the present study we disambiguated both of these effects from exposure rate by systematically varying the duration of training, type of trained dynamics, and exposure rate for these dynamics in human force-field adaptation. After controlling for both training duration and the amount of learning, we found essentially identical retention when comparing gradual and abrupt training for two different types of force-field dynamics. By contrast, we found that retention was markedly higher for long-duration compared with short-duration training for both types of dynamics. These results demonstrate that the duration of training has a far greater effect on the retention of motor memory than the exposure rate during training. We show that a multirate learning model provides a computational mechanism for these findings.NEW & NOTEWORTHY Previous studies have suggested that a gradual, incremental introduction of a novel environment is helpful for improving retention. However, we used experimental and computational approaches to demonstrate that previously reported improvements in retention associated with gradual introductions fail to persist when other factors, including the duration of training and the degree of initial learning, are accounted for.

The 24-h savings of adaptation to novel movement dynamics initially reflects the recall of previous performance.

Humans rapidly adapt reaching movements in response to perturbations (e.g., manipulations of movement dynamics or visual feedback). Following a break, when reexposed to the same perturbation, subjects demonstrate savings, a faster learning rate compared with the time course of initial training. Although this has been well studied, there are open questions on the extent early savings reflects the rapid recall of previous performance. To address this question, we examined how the properties of initial training (duration and final adaptive state) influence initial single-trial adaptation to force-field perturbations when training sessions were separated by 24 h. There were two main groups that were distinct based on the presence or absence of a washout period at the end of day 1 (with washout vs. without washout). We also varied the training duration on day 1 (15, 30, 90, or 160 training trials), resulting in 8 subgroups of subjects. We show that single-trial adaptation on day 2 scaled with training duration, even for similar asymptotic levels of learning on day 1 of training. Interestingly, the temporal force profile following the first perturbation on day 2 matched that at the end of day 1 for the longest training duration group that did not complete the washout. This correspondence persisted but was significantly lower for shorter training durations and the washout subject groups. Collectively, the results suggest that the adaptation observed very early in reexposure results from the rapid recall of the previously learned motor recalibration but is highly dependent on the initial training duration and final adaptive state.NEW & NOTEWORTHY The extent initial readaptation reflects the recall of previous motor performance is largely unknown. We examined early single-trial force-field adaptation on the second day of training and distinguished initial retention from recall. We found that the single-trial adaptation following the 24-h break matched that at the end of the first day, but this recall was modified by the training duration and final level of learning on the first day of training.

Visual Responses in FEF, Unlike V1, Primarily Reflect When the Visual Context Renders a Receptive Field Salient.

When light falls within a neuronal visual receptive field (RF) the resulting activity is referred to as the visual response. Recent work suggests this activity is in response to both the visual stimulation and the abrupt appearance, or salience, of the presentation. Here we present a novel method for distinguishing the two, based on the timing of random and nonrandom presentations. We examined these contributions in frontal eye field (FEF; N = 51) and as a comparison, an early stage in the primary visual cortex (V1; N = 15) of male monkeys (Macaca mulatta). An array of identical stimuli was presented within and outside the neuronal RF while we manipulated salience by varying the time between stimulus presentations. We hypothesized that the rapid presentation would reduce salience (the sudden appearance within the visual field) of a stimulus at any one location, and thus decrease responses driven by salience in the RF. We found that when the interstimulus interval decreased from 500 to 16 ms there was an approximate 79% reduction in the FEF response compared with an estimated 17% decrease in V1. This reduction in FEF response for rapid presentation was evident even when the random sequence preceding a stimulus did not stimulate the RF for 500 ms. The time course of these response changes in FEF suggest that salience is represented much earlier (<100 ms following stimulus onset) than previously estimated. Our results suggest that the contribution of salience dominates at higher levels of the visual system.SIGNIFICANCE STATEMENT The neuronal responses in early visual processing [e.g., primary visual cortex (V1)] reflect primarily the retinal stimulus. Processing in higher visual areas is modulated by a combination of the visual stimulation and contextual factors, such as salience, but identifying these components separately has been difficult. Here we quantified these contributions at a late stage of visual processing [frontal eye field (FEF)] and as a comparison, an early stage in V1. Our results suggest that as visual information continues through higher levels of processing the neural responses are no longer driven primarily by the visual stimulus in the receptive field, but by the broader context that stimulus defines-very different from current views about visual signals in FEF.

The decay of motor adaptation to novel movement dynamics reveals an asymmetry in the stability of motion state-dependent learning.

Motor adaptation paradigms provide a quantitative method to study short-term modification of motor commands. Despite the growing understanding of the role motion states (e.g., velocity) play in this form of motor learning, there is little information on the relative stability of memories based on these movement characteristics, especially in comparison to the initial adaptation. Here, we trained subjects to make reaching movements perturbed by force patterns dependent upon either limb position or velocity. Following training, subjects were exposed to a series of error-clamp trials to measure the temporal characteristics of the feedforward motor output during the decay of learning. The compensatory force patterns were largely based on the perturbation kinematic (e.g., velocity), but also showed a small contribution from the other motion kinematic (e.g., position). However, the velocity contribution in response to the position-based perturbation decayed at a slower rate than the position contribution to velocity-based training, suggesting a difference in stability. Next, we modified a previous model of motor adaptation to reflect this difference and simulated the behavior for different learning goals. We were interested in the stability of learning when the perturbations were based on different combinations of limb position or velocity that subsequently resulted in biased amounts of motion-based learning. We trained additional subjects on these combined motion-state perturbations and confirmed the predictions of the model. Specifically, we show that (1) there is a significant separation between the observed gain-space trajectories for the learning and decay of adaptation and (2) for combined motion-state perturbations, the gain associated to changes in limb position decayed at a faster rate than the velocity-dependent gain, even when the position-dependent gain at the end of training was significantly greater. Collectively, these results suggest that the state-dependent adaptation associated with movement velocity is relatively more stable than that based on position.

Temporal specificity of the initial adaptive response in motor adaptation.

Repeated exposure to a novel physical environment eventually leads to a mature adaptive response whereby feedforward changes in motor output mirror both the amplitude and temporal structure of the environmental perturbations. However, adaptive responses at the earliest stages of learning have been found to be not only smaller, but systematically less specific in their temporal structure compared to later stages of learning. This observation has spawned a lively debate as to whether the temporal structure of the initial adaptive response is, in fact, stereotyped and non-specific. To settle this debate, we directly measured the adaptive responses to velocity-dependent and position-dependent force-field perturbations (vFFs and pFFs) at the earliest possible stage of motor learning in humans-after just a single-movement exposure. In line with previous work, we found these earliest stage adaptive responses to be more similar than the perturbations that induced them. However, the single-trial adaptive responses for vFF and pFF perturbations were clearly distinct, and the disparity between them reflected the difference between the temporal structure of the perturbations that drove them. Critically, we observed these differences between single-trial adaptive responses when vFF and pFF perturbations were randomly intermingled from one trial to the next within the same block, indicating perturbation response specificity at the single trial level. These findings demonstrate that the initial adaptive responses to physical perturbations are not stereotyped. Instead, the neural plasticity in sensorimotor areas is sensitive to the temporal structure of a movement perturbation even at the earliest stage in learning. This insight has direct implications for the development of computational models of early-stage motor adaptation and the evolution of this adaptive response with continued training.

Saccadic Corollary Discharge Underlies Stable Visual Perception.

Saccadic eye movements direct the high-resolution foveae of our retinas toward objects of interest. With each saccade, the image jumps on the retina, causing a discontinuity in visual input. Our visual perception, however, remains stable. Philosophers and scientists over centuries have proposed that visual stability depends upon an internal neuronal signal that is a copy of the neuronal signal driving the eye movement, now referred to as a corollary discharge (CD) or efference copy. In the old world monkey, such a CD circuit for saccades has been identified extending from superior colliculus through MD thalamus to frontal cortex, but there is little evidence that this circuit actually contributes to visual perception. We tested the influence of this CD circuit on visual perception by first training macaque monkeys to report their perceived eye direction, and then reversibly inactivating the CD as it passes through the thalamus. We found that the monkey's perception changed; during CD inactivation, there was a difference between where the monkey perceived its eyes to be directed and where they were actually directed. Perception and saccade were decoupled. We established that the perceived eye direction at the end of the saccade was not derived from proprioceptive input from eye muscles, and was not altered by contextual visual information. We conclude that the CD provides internal information contributing to the brain's creation of perceived visual stability. More specifically, the CD might provide the internal saccade vector used to unite separate retinal images into a stable visual scene. SIGNIFICANCE STATEMENT: Visual stability is one of the most remarkable aspects of human vision. The eyes move rapidly several times per second, displacing the retinal image each time. The brain compensates for this disruption, keeping our visual perception stable. A major hypothesis explaining this stability invokes a signal within the brain, a corollary discharge, that informs visual regions of the brain when and where the eyes are about to move. Such a corollary discharge circuit for eye movements has been identified in macaque monkey. We now show that selectively inactivating this brain circuit alters the monkey's visual perception. We conclude that this corollary discharge provides a critical signal that can be used to unite jumping retinal images into a consistent visual scene.

The absence or temporal offset of visual feedback does not influence adaptation to novel movement dynamics.

Delays in transmitting and processing sensory information require correctly associating delayed feedback to issued motor commands for accurate error compensation. The flexibility of this alignment between motor signals and feedback has been demonstrated for movement recalibration to visual manipulations, but the alignment dependence for adapting movement dynamics is largely unknown. Here we examined the effect of visual feedback manipulations on force-field adaptation. Three subject groups used a manipulandum while experiencing a lag in the corresponding cursor motion (0, 75, or 150 ms). When the offset was applied at the start of the session (continuous condition), adaptation was not significantly different between groups. However, these similarities may be due to acclimation to the offset before motor adaptation. We tested additional subjects who experienced the same delays concurrent with the introduction of the perturbation (abrupt condition). In this case adaptation was statistically indistinguishable from the continuous condition, indicating that acclimation to feedback delay was not a factor. In addition, end-point errors were not significantly different across the delay or onset conditions, but end-point correction (e.g., deceleration duration) was influenced by the temporal offset. As an additional control, we tested a group of subjects who performed without visual feedback and found comparable movement adaptation results. These results suggest that visual feedback manipulation (absence or temporal misalignment) does not affect adaptation to novel dynamics, independent of both acclimation and perceptual awareness. These findings could have implications for modeling how the motor system adjusts to errors despite concurrent delays in sensory feedback information.NEW & NOTEWORTHY A temporal offset between movement and distorted visual feedback (e.g., visuomotor rotation) influences the subsequent motor recalibration, but the effects of this offset for altered movement dynamics are largely unknown. Here we examined the influence of 1) delayed and 2) removed visual feedback on the adaptation to novel movement dynamics. These results contribute to understanding of the control strategies that compensate for movement errors when there is a temporal separation between motion state and sensory information.

The temporal stability of visuomotor adaptation generalization.

Movement adaptation in response to systematic motor perturbations exhibits distinct spatial and temporal properties. These characteristics are typically studied in isolation, leaving the interaction largely unknown. Here we examined how the temporal decay of visuomotor adaptation influences the spatial generalization of the motor recalibration. First, we quantified the extent to which adaptation decayed over time. Subjects reached to a peripheral target, and a rotation was applied to the visual feedback of the unseen motion. The retention of this adaptation over different delays (0-120 s) 1) decreased by 29.0 ± 6.8% at the longest delay and 2) was represented by a simple exponential, with a time constant of 22.5 ± 5.6 s. On the basis of this relationship we simulated how the spatial generalization of adaptation would change with delay. To test this directly, we trained additional subjects with the same perturbation and assessed transfer to 19 different locations (spaced 15° apart, symmetric around the trained location) and examined three delays (~4, 12, and 25 s). Consistent with the simulation, we found that generalization around the trained direction (±15°) significantly decreased with delay and distance, while locations >60° displayed near-constant spatiotemporal transfer. Intermediate distances (30° and 45°) showed a difference in transfer across space, but this amount was approximately constant across time. Interestingly, the decay at the trained direction was faster than that based purely on time, suggesting that the spatial transfer of adaptation is modified by concurrent passive (time dependent) and active (movement dependent) processes.NEW & NOTEWORTHY Short-term motor adaptation exhibits distinct spatial and temporal characteristics. Here we investigated the interaction of these features, utilizing a simple motor adaptation paradigm (recalibration of reaching arm movements in response to rotated visual feedback). We examined the changes in the spatial generalization of motor adaptation for different temporal manipulations and report that the spatiotemporal generalization of motor adaptation is generally local and is influenced by both passive (time dependent) and active (movement dependent) learning processes.

Quantifying the spatial extent of the corollary discharge benefit to transsaccadic visual perception.

Extraretinal information, such as corollary discharge (CD), is hypothesized to help compensate for saccade-induced visual input disruptions. However, support for this hypothesis is largely for one-dimensional transsaccadic visual changes, with little comprehensive information on the spatial characteristics. Here we systematically mapped the two-dimensional extent of this compensation by quantifying the insensitivity to different displacement metrics. Human subjects made saccades to targets positioned at different amplitudes (4° or 8°) and directions (rightward, oblique, or upward). After the saccade the initial target disappeared and, after a blank period, reappeared at a shifted location-a collinear, diagonal, or orthogonal displacement. Subjects reported the perceived shift direction, and we determined the displacement detection based on the perceptual judgments. The two-dimensional insensitivity fields resulting from the perceptual thresholds had spatial features similar to the saccadic eye movement variability: 1) scaled with movement amplitude, 2) oriented (less sensitive to the change) along the saccade vector, and 3) approximately constant in shape when normalized by movement amplitude. In addition, comparing the postsaccadic perceptual estimate of the presaccadic target location to that based solely on the postsaccade visual error showed that overall the perceptual estimate was approximately 50% more accurate and 35% less variable than estimates based solely on this visual information. However, this relationship was not uniform: The benefit of extraretinal information was observed largely for displacements with a component parallel to the saccade vector. These results suggest a graded use of extraretinal information when forming the postsaccadic perceptual evaluation of transsaccadic environmental changes.

The effect of saccade metrics on the corollary discharge contribution to perceived eye location.

Corollary discharge (CD) is hypothesized to provide the movement information (direction and amplitude) required to compensate for the saccade-induced disruptions to visual input. Here, we investigated to what extent these conveyed metrics influence perceptual stability in human subjects with a target-displacement detection task. Subjects made saccades to targets located at different amplitudes (4°, 6°, or 8°) and directions (horizontal or vertical). During the saccade, the target disappeared and then reappeared at a shifted location either in the same direction or opposite to the movement vector. Subjects reported the target displacement direction, and from these reports we determined the perceptual threshold for shift detection and estimate of target location. Our results indicate that the thresholds for all amplitudes and directions generally scaled with saccade amplitude. Additionally, subjects on average produced hypometric saccades with an estimated CD gain <1. Finally, we examined the contribution of different error signals to perceptual performance, the saccade error (movement-to-movement variability in saccade amplitude) and visual error (distance between the fovea and the shifted target location). Perceptual judgment was not influenced by the fluctuations in movement amplitude, and performance was largely the same across movement directions for different magnitudes of visual error. Importantly, subjects reported the correct direction of target displacement above chance level for very small visual errors (<0.75°), even when these errors were opposite the target-shift direction. Collectively, these results suggest that the CD-based compensatory mechanisms for visual disruptions are highly accurate and comparable for saccades with different metrics.

Compression and suppression of shifting receptive field activity in frontal eye field neurons.

Before each saccade, neurons in frontal eye field anticipate the impending eye movement by showing sensitivity to stimuli appearing where the neuron's receptive field will be at the end of the saccade, referred to as the future field (FF) of the neuron. We explored the time course of this anticipatory activity in monkeys by briefly flashing stimuli in the FF at different times before saccades. Different neurons showed substantial variation in FF time course, but two salient observations emerged. First, when we compared the time span of stimulus probes before the saccade to the time span of FF activity, we found a striking temporal compression of FF activity, similar to compression seen for perisaccadic stimuli in human psychophysics. Second, neurons with distinct FF activity also showed suppression at the time of the saccade. The increase in FF activity and the decrease with suppression were temporally independent, making the patterns of activity difficult to separate. We resolved this by constructing a simple model with values for the start, peak, and duration of FF activity and suppression for each neuron. The model revealed the different time courses of FF sensitivity and suppression, suggesting that information about the impending saccade triggering suppression reaches the frontal eye field through a different pathway, or a different mechanism, than that triggering FF activity. Recognition of the variations in the time course of anticipatory FF activity provides critical information on its function and its relation to human visual perception at the time of the saccade.

Dissociating the Influence of Limb Posture and Visual Feedback Shifts on the Adaptation to Novel Movement Dynamics.

Accurate movements of the upper limb require the integration of various forms of sensory feedback (e.g., visual and postural information). The influence of these different sensory modalities on reaching movements has been largely studied by assessing endpoint errors after selectively perturbing sensory estimates of hand location. These studies have demonstrated that both vision and proprioception make key contributions in determining the reach endpoint. However, their influence on motor output throughout movement remains unclear. Here we used separate perturbations of posture and visual information to dissociate their effects on reaching dynamics and temporal force profiles during point-to-point reaching movements. We tested human subjects (N = 32) and found that vision and posture modulate select aspects of reaching dynamics. Specifically, altering arm posture influences the relationship between temporal force patterns and the motion-state variables of hand position and acceleration, whereas dissociating visual feedback influences the relationship between force patterns and the motion-state variables of velocity and acceleration. Next, we examined the extent these baseline motion-state relationships influence motor adaptation based on perturbations of movement dynamics. We trained subjects using a velocity-dependent force-field to probe the extent arm posture-dependent influences persisted after exposure to a motion-state dependent perturbation. Changes in the temporal force profiles due to variations in arm posture were not reduced by adaptation to novel movement dynamics, but persisted throughout learning. These results suggest that vision and posture differentially influence the internal estimation of limb state throughout movement and play distinct roles in forming the response to external perturbations during movement.

Multimodal processing of noisy cues in bumblebees.

Multimodal cues can improve behavioral responses by enhancing the detection and localization of sensory cues and reducing response times. Across species, studies have shown that multisensory integration of visual and olfactory cues can improve response accuracy. However, in real-world settings, sensory cues are often noisy; visual and olfactory cues can be deteriorated, masked, or mixed, making the target cue less clear to the receiver. In this study, we use an associative learning paradigm (Free Moving Proboscis Extension Reflex, FMPER) to show that having multimodal cues may improve the accuracy of bees' responses to noisy cues. Adding a noisy visual cue improves the accuracy of response to a noisy olfactory cue, despite neither the clear nor noisy visual cue being sufficient when paired with a novel olfactory cue. This may provide insight into the neural mechanisms underlying multimodal processing and the effects of environmental change on pollination services.

Understanding the capacity of children with congenital unilateral below-elbow deficiency to actuate their affected muscles.

In recent years, commercially available dexterous upper limb prostheses for children have begun to emerge. These devices derive control signals from surface electromyography (measure of affected muscle electrical activity, sEMG) to drive a variety of grasping motions. However, the ability for children with congenital upper limb deficiency to actuate their affected muscles to achieve naturalistic prosthetic control is not well understood, as compared to adults or children with acquired hand loss. To address this gap, we collected sEMG data from 9 congenital one-handed participants ages 8-20 years as they envisioned and attempted to perform 10 different movements with their missing hands. Seven sEMG electrodes were adhered circumferentially around the participant's affected and unaffected limbs and participants mirrored the attempted missing hand motions with their intact side. To analyze the collected sEMG data, we used time and frequency domain analyses. We found that for the majority of participants, attempted hand movements produced detectable and consistent muscle activity, and the capacity to achieve this was not dissimilar across the affected and unaffected sides. These data suggest that children with congenital hand absence retain a degree of control over their affected muscles, which has important implications for translating and refining advanced prosthetic control technologies for children.

Suppressive surrounds of receptive fields in monkey frontal eye field.

A critical step in determining how a neuron contributes to visual processing is determining its visual receptive field (RF). While recording from neurons in frontal eye field (FEF) of awake monkeys (Macaca mulatta), we probed the visual field with small spots of light and found excitatory RFs that decreased in strength from RF center to periphery. However, presenting stimuli with different diameters centered on the RF revealed suppressive surrounds that overlapped the previously determined excitatory RF and reduced responses by 84%, on average. Consequently, in that overlap area, stimulation produced excitation or suppression, depending on the stimulus. Strong stimulation of the RF periphery with annular stimuli allowed us to quantify this effect. A modified difference of gaussians model that independently varied center and surround activation accounted for the nonlinear activity in the overlap area. Our results suggest that (1) the suppressive surrounds found in FEF are fundamentally the same as those in V1 except for the size and strength of excitatory and suppressive mechanisms, (2) methodically assaying suppressive surrounds in FEF is essential for correctly interpreting responses to large and/or peripheral stimuli and therefore understanding the effects of stimulus context, and (3) regulating the relative strength of the surround clearly changes neuronal responses and may therefore play a significant part in the neuronal changes resulting from visual attention and stimulus salience.