Visual working memory models of delayed estimation do not generalize to whole-report tasks.

Whole-report working memory tasks provide a measure of recall for all stimuli in a trial and afford single-trial analyses that are not possible with single-report delayed estimation tasks. However, most whole-report studies assume that trial stimuli are encoded and reported independently, and they do not consider the relationships between stimuli presented and reported within the same trial. Here, we present the results of two independently conducted whole-report experiments. The first dataset was recorded by Adam, Vogel, and Awh (2017) and required participants to report color and orientation stimuli using a continuous response wheel. We recorded the second dataset, which required participants to report color stimuli using a set of discrete buttons. We found that participants often group their reports by color similarity, contradicting the assumption of independence implicit in most encoding models of working memory. Next, we showed that this behavior was consistent across participants and experiments when reporting color but not orientation, two circular variables often assumed to be equivalent.Finally, we implemented an alternative to independent encoding where stimuli are encoded as a hierarchical Bayesian ensemble and found that this model predicts biases that are not present in either dataset. Our results suggest that assumptions made by both independent and hierarchical ensemble encoding models-which were developed in the context of single-report delayed estimation tasks-do not hold for the whole-report task. This failure to generalize highlights the need to consider variations in task structure when inferring fundamental principles of visual working memory.

Parietofrontal oscillations show hand-specific interactions with top-down movement plans.

To generate a hand-specific reach plan, the brain must integrate hand-specific signals with the desired movement strategy. Although various neurophysiology/imaging studies have investigated hand-target interactions in simple reach-to-target tasks, the whole brain timing and distribution of this process remain unclear, especially for more complex, instruction-dependent motor strategies. Previously, we showed that a pro/anti pointing instruction influences magnetoencephalographic (MEG) signals in frontal cortex that then propagate recurrently through parietal cortex (Blohm G, Alikhanian H, Gaetz W, Goltz HC, DeSouza JF, Cheyne DO, Crawford JD. NeuroImage 197: 306-319, 2019). Here, we contrasted left versus right hand pointing in the same task to investigate 1) which cortical regions of interest show hand specificity and 2) which of those areas interact with the instructed motor plan. Eight bilateral areas, the parietooccipital junction (POJ), superior parietooccipital cortex (SPOC), supramarginal gyrus (SMG), medial/anterior interparietal sulcus (mIPS/aIPS), primary somatosensory/motor cortex (S1/M1), and dorsal premotor cortex (PMd), showed hand-specific changes in beta band power, with four of these (M1, S1, SMG, aIPS) showing robust activation before movement onset. M1, SMG, SPOC, and aIPS showed significant interactions between contralateral hand specificity and the instructed motor plan but not with bottom-up target signals. Separate hand/motor signals emerged relatively early and lasted through execution, whereas hand-motor interactions only occurred close to movement onset. Taken together with our previous results, these findings show that instruction-dependent motor plans emerge in frontal cortex and interact recurrently with hand-specific parietofrontal signals before movement onset to produce hand-specific motor behaviors.NEW & NOTEWORTHY The brain must generate different motor signals depending on which hand is used. The distribution and timing of hand use/instructed motor plan integration are not understood at the whole brain level. Using MEG we show that different action planning subnetworks code for hand usage and integrating hand use into a hand-specific motor plan. The timing indicates that frontal cortex first creates a general motor plan and then integrates hand specificity to produce a hand-specific motor plan.

Effector-dependent stochastic reference frame transformations alter decision-making.

Psychophysical, motor control, and modeling studies have revealed that sensorimotor reference frame transformations (RFTs) add variability to transformed signals. For perceptual decision-making, this phenomenon could decrease the fidelity of a decision signal's representation or alternatively improve its processing through stochastic facilitation. We investigated these two hypotheses under various sensorimotor RFT constraints. Participants performed a time-limited, forced-choice motion discrimination task under eight combinations of head roll and/or stimulus rotation while responding either with a saccade or button press. This paradigm, together with the use of a decision model, allowed us to parameterize and correlate perceptual decision behavior with eye-, head-, and shoulder-centered sensory and motor reference frames. Misalignments between sensory and motor reference frames produced systematic changes in reaction time and response accuracy. For some conditions, these changes were consistent with a degradation of motion evidence commensurate with a decrease in stimulus strength in our model framework. Differences in participant performance were explained by a continuum of eye-head-shoulder representations of accumulated motion evidence, with an eye-centered bias during saccades and a shoulder-centered bias during button presses. In addition, we observed evidence for stochastic facilitation during head-rolled conditions (i.e., head roll resulted in faster, more accurate decisions in oblique motion for a given stimulus-response misalignment). We show that perceptual decision-making and stochastic RFTs are inseparable within the present context. We show that by simply rolling one's head, perceptual decision-making is altered in a way that is predicted by stochastic RFTs.

Confidence in predicted position error explains saccadic decisions during pursuit.

A fundamental problem in motor control is the coordination of complementary movement types to achieve a common goal. As a common example, humans view moving objects through coordinated pursuit and saccadic eye movements. Pursuit is initiated and continuously controlled by retinal image velocity. During pursuit, eye position may lag behind the target. This can be compensated by the discrete execution of a catch-up saccade. The decision to trigger a saccade is influenced by both position and velocity errors, and the timing of saccades can be highly variable. The observed distributions of saccade frequency and trigger time remain poorly understood, and this decision process remains imprecisely quantified. Here, we propose a predictive, probabilistic model explaining the decision to trigger saccades during pursuit to foveate moving targets. In this model, expected position error and its associated uncertainty are predicted through Bayesian inference across noisy, delayed sensory observations (Kalman filtering). This probabilistic prediction is used to estimate the confidence that a saccade is needed (quantified through log-probability ratio), triggering a saccade upon accumulating to a fixed threshold. The model qualitatively explains behavioral observations on the frequency and trigger time distributions of saccades during pursuit over a range of target motion trajectories. Furthermore, this model makes novel predictions that saccade decisions are highly sensitive to uncertainty for small predicted position errors, but this influence diminishes as the magnitude of predicted position error increases. We suggest that this predictive, confidence-based decision-making strategy represents a fundamental principle for the probabilistic neural control of coordinated movements.NEW & NOTEWORTHY This is the first stochastic dynamical systems model of pursuit-saccade coordination accounting for noise and delays in the sensorimotor system. The model uses Bayesian inference to predictively estimate visual motion, triggering saccades when confidence in predicted position error accumulates to a threshold. This model explains saccade frequency and trigger time distributions across target trajectories and makes novel predictions about the influence of sensory uncertainty in saccade decisions during pursuit.

Reaching around obstacles accounts for uncertainty in coordinate transformations.

When reaching to a visual target, humans need to transform the spatial target representation into the coordinate system of their moving arm. It has been shown that increased uncertainty in such coordinate transformations, for instance, when the head is rolled toward one shoulder, leads to higher movement variability and influence movement decisions. However, it is unknown whether the brain incorporates such added variability in planning and executing movements. We designed an obstacle avoidance task in which participants had to reach with or without visual feedback of the hand to a visual target while avoiding collisions with an obstacle. We varied coordinate transformation uncertainty by varying head roll (straight, 30° clockwise, and 30° counterclockwise). In agreement with previous studies, we observed that the reaching variability increased when the head was tilted. Indeed, head roll did not influence the number of collisions during reaching compared with the head-straight condition, but it did systematically change the obstacle avoidance behavior. Participants changed the preferred direction of passing the obstacle and increased the safety margins indicated by stronger movement curvature. These results suggest that the brain takes the added movement variability during head roll into account and compensates for it by adjusting the reaching trajectories.NEW & NOTEWORTHY We show that changing body geometry such as head roll results in compensatory reaching behaviors around obstacles. Specifically, we observed head roll causes changed preferred movement direction and increased trajectory curvature. As has been shown before, head roll increases movement variability due to stochastic coordinate transformations. Thus these results provide evidence that the brain must consider the added movement variability caused by coordinate transformations for accurate reach movements.

Hierarchical recruitment of competition alleviates working memory overload in a frontoparietal model.

The storage limitations of visual working memory have been the subject of intense research interest for several decades, but few studies have systematically investigated the dependence of these limitations on memory load that exceeds our retention abilities. Under this real-world scenario, performance typically declines beyond a critical load among low-performing subjects, a phenomenon known as working memory overload. We used a frontoparietal cortical model to test the hypothesis that high-performing subjects select a manageable number of items for storage, thereby avoiding overload. The model accounts for behavioral and electrophysiological data from high-performing subjects in a parameter regime where competitive encoding in its prefrontal network selects items for storage, interareal projections sustain their representations after stimulus offset, and weak dynamics in its parietal network limit their mutual interference. Violation of these principles accounts for these data among low-performing subjects, implying that poor visual working memory performance reflects poor control over frontoparietal circuitry, making testable predictions for experiments.

Saccade-induced changes in ocular torsion reveal predictive orientation perception.

Natural orienting of gaze often results in a retinal image that is rotated relative to space due to ocular torsion. However, we perceive neither this rotation nor a moving world despite visual rotational motion on the retina. This perceptual stability is often attributed to the phenomenon known as predictive remapping, but the current remapping literature ignores this torsional component. In addition, studies often simply measure remapping across either space or features (e.g., orientation) but in natural circumstances, both components are bound together for stable perception. One natural circumstance in which the perceptual system must account for the current and future eye orientation to correctly interpret the orientation of external stimuli occurs during movements to or from oblique eye orientations (i.e., eye orientations with both a horizontal and vertical angular component relative to the primary position). Here we took advantage of oblique eye orientation-induced ocular torsion to examine perisaccadic orientation perception. First, we found that orientation perception was largely predicted by the rotated retinal image. Second, we observed a presaccadic remapping of orientation perception consistent with maintaining a stable (but spatially inaccurate) retinocentric perception throughout the saccade. These findings strongly suggest that our seamless perceptual stability relies on retinocentric signals that are predictively remapped in all three ocular dimensions with each saccade.

Misperception of motion in depth originates from an incomplete transformation of retinal signals.

Depth perception requires the use of an internal model of the eye-head geometry to infer distance from binocular retinal images and extraretinal 3D eye-head information, particularly ocular vergence. Similarly, for motion in depth perception, gaze angle is required to correctly interpret the spatial direction of motion from retinal images; however, it is unknown whether the brain can make adequate use of extraretinal version and vergence information to correctly transform binocular retinal motion into 3D spatial coordinates. Here we tested this hypothesis by asking participants to reconstruct the spatial trajectory of an isolated disparity stimulus moving in depth either peri-foveally or peripherally while participants' gaze was oriented at different vergence and version angles. We found large systematic errors in the perceived motion trajectory that reflected an intermediate reference frame between a purely retinal interpretation of binocular retinal motion (not accounting for veridical vergence and version) and the spatially correct motion. We quantify these errors with a 3D reference frame model accounting for target, eye, and head position upon motion percept encoding. This model could capture the behavior well, revealing that participants tended to underestimate their version by up to 17%, overestimate their vergence by up to 22%, and underestimate the overall change in retinal disparity by up to 64%, and that the use of extraretinal information depended on retinal eccentricity. Since such large perceptual errors are not observed in everyday viewing, we suggest that both monocular retinal cues and binocular extraretinal signals are required for accurate real-world motion in depth perception.

Neck muscle spindle noise biases reaches in a multisensory integration task.

Reference frame transformations (RFTs) are crucial components of sensorimotor transformations in the brain. Stochasticity in RFTs has been suggested to add noise to the transformed signal due to variability in transformation parameter estimates (e.g., angle) as well as the stochastic nature of computations in spiking networks of neurons. Here, we varied the RFT angle together with the associated variability and evaluated the behavioral impact in a reaching task that required variability-dependent visual-proprioceptive multisensory integration. Crucially, reaches were performed with the head either straight or rolled 30° to either shoulder, and we also applied neck loads of 0 or 1.8 kg (left or right) in a 3 × 3 design, resulting in different combinations of estimated head roll angle magnitude and variance required in RFTs. A novel three-dimensional stochastic model of multisensory integration across reference frames was fitted to the data and captured our main behavioral findings: 1) neck load biased head angle estimation across all head roll orientations, resulting in systematic shifts in reach errors; 2) increased neck muscle tone led to increased reach variability due to signal-dependent noise; and 3) both head roll and neck load created larger angular errors in reaches to visual targets away from the body compared with reaches toward the body. These results show that noise in muscle spindles and stochasticity in general have a tangible effect on RFTs underlying reach planning. Since RFTs are omnipresent in the brain, our results could have implications for processes as diverse as motor control, decision making, posture/balance control, and perception. NEW & NOTEWORTHY We show that increasing neck muscle tone systematically biases reach movements. A novel three-dimensional multisensory integration across reference frames model captures the data well and provides evidence that the brain must have online knowledge of full-body geometry together with the associated variability to plan reach movements accurately.

Corrective response times in a coordinated eye-head-arm countermanding task.

Inhibition of motor responses has been described as a race between two competing decision processes of motor initiation and inhibition, which manifest as the reaction time (RT) and the stop signal reaction time (SSRT); in the case where motor initiation wins out over inhibition, an erroneous movement occurs that usually needs to be corrected, leading to corrective response times (CRTs). Here we used a combined eye-head-arm movement countermanding task to investigate the mechanisms governing multiple effector coordination and the timing of corrective responses. We found a high degree of correlation between effector response times for RT, SSRT, and CRT, suggesting that decision processes are strongly dependent across effectors. To gain further insight into the mechanisms underlying CRTs, we tested multiple models to describe the distribution of RTs, SSRTs, and CRTs. The best-ranked model (according to 3 information criteria) extends the LATER race model governing RTs and SSRTs, whereby a second motor initiation process triggers the corrective response (CRT) only after the inhibition process completes in an expedited fashion. Our model suggests that the neural processing underpinning a failed decision has a residual effect on subsequent actions. NEW & NOTEWORTHY Failure to inhibit erroneous movements typically results in corrective movements. For coordinated eye-head-hand movements we show that corrective movements are only initiated after the erroneous movement cancellation signal has reached a decision threshold in an accelerated fashion.

Learned rather than online relative weighting of visual-proprioceptive sensory cues.

When reaching to an object, information about the target location as well as the initial hand position is required to program the motor plan for the arm. The initial hand position can be determined by proprioceptive information as well as visual information, if available. Bayes-optimal integration posits that we utilize all information available, with greater weighting on the sense that is more reliable, thus generally weighting visual information more than the usually less reliable proprioceptive information. The criterion by which information is weighted has not been explicitly investigated; it has been assumed that the weights are based on task- and effector-dependent sensory reliability requiring an explicit neuronal representation of variability. However, the weights could also be determined implicitly through learned modality-specific integration weights and not on effector-dependent reliability. While the former hypothesis predicts different proprioceptive weights for left and right hands, e.g., due to different reliabilities of dominant vs. nondominant hand proprioception, we would expect the same integration weights if the latter hypothesis was true. We found that the proprioceptive weights for the left and right hands were extremely consistent regardless of differences in sensory variability for the two hands as measured in two separate complementary tasks. Thus we propose that proprioceptive weights during reaching are learned across both hands, with high interindividual range but independent of each hand's specific proprioceptive variability. NEW & NOTEWORTHY How visual and proprioceptive information about the hand are integrated to plan a reaching movement is still debated. The goal of this study was to clarify how the weights assigned to vision and proprioception during multisensory integration are determined. We found evidence that the integration weights are modality specific rather than based on the sensory reliabilities of the effectors.

Temporal Evolution of Target Representation, Movement Direction Planning, and Reach Execution in Occipital-Parietal-Frontal Cortex: An fMRI Study.

The cortical mechanisms for reach have been studied extensively, but directionally selective mechanisms for visuospatial target memory, movement planning, and movement execution have not been clearly differentiated in the human. We used an event-related fMRI design with a visuospatial memory delay, followed by a pro-/anti-reach instruction, a planning delay, and finally a "go" instruction for movement. This sequence yielded temporally separable preparatory responses that expanded from modest parieto-frontal activation for visual target memory to broad occipital-parietal-frontal activation during planning and execution. Using the pro/anti instruction to differentiate visual and motor directional selectivity during planning, we found that one occipital area showed contralateral "visual" selectivity, whereas a broad constellation of left hemisphere occipital, parietal, and frontal areas showed contralateral "movement" selectivity. Temporal analysis of these areas through the entire memory-planning sequence revealed early visual selectivity in most areas, followed by movement selectivity in most areas, with all areas showing a stereotypical visuo-movement transition. Cross-correlation of these spatial parameters through time revealed separate spatiotemporally correlated modules for visual input, motor output, and visuo-movement transformations that spanned occipital, parietal, and frontal cortex. These results demonstrate a highly distributed occipital-parietal-frontal reach network involved in the transformation of retrospective sensory information into prospective movement plans.

Transsaccadic memory of multiple spatially variant and invariant object features.

Transsaccadic memory is a process by which remembered object information is updated across a saccade. To date, studies on transsaccadic memory have used simple stimuli-that is, a single dot or feature of an object. It remains unknown how transsaccadic memory occurs for more realistic, complex objects with multiple features. An object's location is a central feature for transsaccadic updating, as it is spatially variant, but other features such as size are spatially invariant. How these spatially variant and invariant features of an object are remembered and updated across saccades is not well understood. Here we tested how well 14 participants remembered either three different features together (location, orientation, and size) or a single feature at a time of a bar either while fixating either with or without an intervening saccade. We found that the intervening saccade influenced memory of all three features, with consistent biases of the remembered location, orientation, and size that were dependent on the direction of the saccade. These biases were similar whether participants remembered either a single feature or multiple features and were not observed with increased memory load (single vs. multiple features during fixation trials), confirming that these effects were specific to the saccade updating mechanisms. We conclude that multiple features of an object are updated together across eye movements, supporting the notion that spatially invariant features of an object are bound to their location in memory.

Weighted integration of short-term memory and sensory signals in the oculomotor system.

Oculomotor behaviors integrate sensory and prior information to overcome sensory-motor delays and noise. After much debate about this process, reliability-based integration has recently been proposed and several models of smooth pursuit now include recurrent Bayesian integration or Kalman filtering. However, there is a lack of behavioral evidence in humans supporting these theoretical predictions. Here, we independently manipulated the reliability of visual and prior information in a smooth pursuit task. Our results show that both smooth pursuit eye velocity and catch-up saccade amplitude were modulated by visual and prior information reliability. We interpret these findings as the continuous reliability-based integration of a short-term memory of target motion with visual information, which support modeling work. Furthermore, we suggest that saccadic and pursuit systems share this short-term memory. We propose that this short-term memory of target motion is quickly built and continuously updated, and constitutes a general building block present in all sensorimotor systems.

Ca2+ removal by the plasma membrane Ca2+-ATPase influences the contribution of mitochondria to activity-dependent Ca2+ dynamics in Aplysia neuroendocrine cells.

After Ca(2+) influx, mitochondria can sequester Ca(2+) and subsequently release it back into the cytosol. This form of Ca(2+)-induced Ca(2+) release (CICR) prolongs Ca(2+) signaling and can potentially mediate activity-dependent plasticity. As Ca(2+) is required for its subsequent release, Ca(2+) removal systems, like the plasma membrane Ca(2+)-ATPase (PMCA), could impact CICR. Here we examine such a role for the PMCA in the bag cell neurons of Aplysia californica CICR is triggered in these neurons during an afterdischarge and is implicated in sustaining membrane excitability and peptide secretion. Somatic Ca(2+) was measured from fura-PE3-loaded cultured bag cell neurons recorded under whole cell voltage clamp. Voltage-gated Ca(2+) influx was elicited with a 5-Hz, 1-min train, which mimics the fast phase of the afterdischarge. PMCA inhibition with carboxyeosin or extracellular alkalization augmented the effectiveness of Ca(2+) influx in eliciting mitochondrial CICR. A Ca(2+) compartment model recapitulated these findings and indicated that disrupting PMCA-dependent Ca(2+) removal increases CICR by enhancing mitochondrial Ca(2+) loading. Indeed, carboxyeosin augmented train-evoked mitochondrial Ca(2+) uptake. Consistent with their role on Ca(2+) dynamics, cell labeling revealed that the PMCA and mitochondria overlap with Ca(2+) entry sites. Finally, PMCA-dependent Ca(2+) extrusion did not impact endoplasmic reticulum-dependent Ca(2+) removal or release, despite the organelle residing near Ca(2+) entry sites. Our results demonstrate that Ca(2+) removal by the PMCA influences the propensity for stimulus-evoked CICR by adjusting the amount of Ca(2+) available for mitochondrial Ca(2+) uptake. This study highlights a mechanism by which the PMCA could impact activity-dependent plasticity in the bag cell neurons.

Neural correlate of spatial (mis-)localization during smooth eye movements.

The dependence of neuronal discharge on the position of the eyes in the orbit is a functional characteristic of many visual cortical areas of the macaque. It has been suggested that these eye-position signals provide relevant information for a coordinate transformation of visual signals into a non-eye-centered frame of reference. This transformation could be an integral part for achieving visual perceptual stability across eye movements. Previous studies demonstrated close to veridical eye-position decoding during stable fixation as well as characteristic erroneous decoding across saccadic eye-movements. Here we aimed to decode eye position during smooth pursuit. We recorded neural activity in macaque area VIP during steady fixation, saccades and smooth-pursuit and investigated the temporal and spatial accuracy of eye position as decoded from the neuronal discharges. Confirming previous results, the activity of the majority of neurons depended linearly on horizontal and vertical eye position. The application of a previously introduced computational approach (isofrequency decoding) allowed eye position decoding with considerable accuracy during steady fixation. We applied the same decoder on the activity of the same neurons during smooth-pursuit. On average, the decoded signal was leading the current eye position. A model combining this constant lead of the decoded eye position with a previously described attentional bias ahead of the pursuit target describes the asymmetric mislocalization pattern for briefly flashed stimuli during smooth pursuit eye movements as found in human behavioral studies.

Effects of a pretarget distractor on saccade reaction times across space and time in monkeys and humans.

Previous studies have shown that the influence of a behaviorally irrelevant distractor on saccade reaction times (SRTs) varies depending on the temporal and spatial relationship between the distractor and the saccade target. We measured distractor influence on SRTs to a subsequently presented target, varying the spatial location and the timing between the distractor and the target. The distractor appeared at one of four equally eccentric locations, followed by a target (either 50 ms or 200 ms after) at one of 136 different locations encompassing an area of 20° square. We extensively tested two humans and two monkeys on this task to determine interspecies similarities and differences, since monkey neurophysiology is often used to interpret human behavioral findings. Results were similar across species; for the short interval (50 ms), SRTs were shortest to a target presented close to or at the distractor location and increased primarily as a function of the distance from the distractor. There was also an effect of distractor-target direction and visual field. For the long interval (200 ms) the results were inverted; SRTs were longest for short distances between the distractor and target and decreased as a function of distance from distractor. Both SRT patterns were well captured by a two-dimensional dynamic field model with short-distance excitation and long-distance inhibition, based upon known functional connectivity found in the superior colliculus that includes wide-spread excitation and inhibition. Based on these findings, we posit that the different time-dependent patterns of distractor-related SRTs can emerge from the same underlying neuronal mechanisms common to both species.

Computations underlying the visuomotor transformation for smooth pursuit eye movements.

Smooth pursuit eye movements are driven by retinal motion and enable us to view moving targets with high acuity. Complicating the generation of these movements is the fact that different eye and head rotations can produce different retinal stimuli but giving rise to identical smooth pursuit trajectories. However, because our eyes accurately pursue targets regardless of eye and head orientation (Blohm G, Lefèvre P. J Neurophysiol 104: 2103-2115, 2010), the brain must somehow take these signals into account. To learn about the neural mechanisms potentially underlying this visual-to-motor transformation, we trained a physiologically inspired neural network model to combine two-dimensional (2D) retinal motion signals with three-dimensional (3D) eye and head orientation and velocity signals to generate a spatially correct 3D pursuit command. We then simulated conditions of 1) head roll-induced ocular counterroll, 2) oblique gaze-induced retinal rotations, 3) eccentric gazes (invoking the half-angle rule), and 4) optokinetic nystagmus to investigate how units in the intermediate layers of the network accounted for different 3D constraints. Simultaneously, we simulated electrophysiological recordings (visual and motor tunings) and microstimulation experiments to quantify the reference frames of signals at each processing stage. We found a gradual retinal-to-intermediate-to-spatial feedforward transformation through the hidden layers. Our model is the first to describe the general 3D transformation for smooth pursuit mediated by eye- and head-dependent gain modulation. Based on several testable experimental predictions, our model provides a mechanism by which the brain could perform the 3D visuomotor transformation for smooth pursuit.

Ca2+-induced uncoupling of Aplysia bag cell neurons.

Electrical transmission is a dynamically regulated form of communication and key to synchronizing neuronal activity. The bag cell neurons of Aplysia are a group of electrically coupled neuroendocrine cells that initiate ovulation by secreting egg-laying hormone during a prolonged period of synchronous firing called the afterdischarge. Accompanying the afterdischarge is an increase in intracellular Ca2+ and the activation of protein kinase C (PKC). We used whole cell recording from paired cultured bag cell neurons to demonstrate that electrical coupling is regulated by both Ca2+ and PKC. Elevating Ca2+ with a train of voltage steps, mimicking the onset of the afterdischarge, decreased junctional current for up to 30 min. Inhibition was most effective when Ca2+ entry occurred in both neurons. Depletion of Ca2+ from the mitochondria, but not the endoplasmic reticulum, also attenuated the electrical synapse. Buffering Ca2+ with high intracellular EGTA or inhibiting calmodulin kinase prevented uncoupling. Furthermore, activating PKC produced a small but clear decrease in junctional current, while triggering both Ca2+ influx and PKC inhibited the electrical synapse to a greater extent than Ca2+ alone. Finally, the amplitude and time course of the postsynaptic electrotonic response were attenuated after Ca2+ influx. A mathematical model of electrically connected neurons showed that excessive coupling reduced recruitment of the cells to fire, whereas less coupling led to spiking of essentially all neurons. Thus a decrease in electrical synapses could promote the afterdischarge by ensuring prompt recovery of electrotonic potentials or making the neurons more responsive to current spreading through the network.

Saccade execution suppresses discrimination at distractor locations rather than enhancing the saccade goal location.

As we have limited processing abilities with respect to the plethora of visual information entering our brain, spatial selection mechanisms are crucial. These mechanisms result in both enhancing processing at a location of interest and in suppressing processing at other locations; together, they enable successful further processing of locations of interest. It has been suggested that saccade planning modulates these spatial selection mechanisms; however, the precise influence of saccades on the distribution of spatial resources underlying selection remains unclear. To this end, we compared discrimination performance at different locations (six) within a work space during different saccade tasks. We used visual discrimination performance as a behavioral measure of enhancement and suppression at the different locations. A total of 14 participants performed a dual discrimination/saccade countermanding task, which allowed us to specifically isolate the consequences of saccade execution. When a saccade was executed, discrimination performance at the cued location was never better than when fixation was maintained, suggesting that saccade execution did not enhance processing at a location more than knowing the likelihood of its appearance. However, discrimination was consistently lower at distractor (uncued) locations in all cases where a saccade was executed compared with when fixation was maintained. Based on these results, we suggest that saccade execution specifically suppresses distractor locations, whereas attention shifts (with or without an accompanying saccade) are involved in enhancing perceptual processing at the goal location.