Journal of Neurophysiology current issue

Preprint Servers and the Journal of Neurophysiology

Linden, D. J. — Wed, 04 Nov 2009 11:06:50 PST

Top-Down Control of Saccades as Part of a Generalized Model of Proactive Inhibitory Control

Ballanger, B. — Wed, 04 Nov 2009 11:06:50 PST

Lo and colleagues have recently described a recurrent network model of inhibitory control of saccadic eye movements based on neurophysiological observations in the frontal eye field (FEF) and superior colliculus (SC) of rhesus monkeys. This model emphasizes the proactive, inhibition-based, tonic neuronal activity that prevents the eye from moving in a countermanding paradigm. In this review I discuss the model with respect to existing literature that the authors did not mention, suggesting that proactive inhibitory control extends far beyond saccadic control and provides an interesting framework to interpret several attentional and movement disorders in humans.

Exploring the Superior Colliculus In Vitro

Isa, T., Hall, W. C. — Wed, 04 Nov 2009 11:06:50 PST

The superior colliculus plays an important role in the translation of sensory signals that encode the location of objects in space into motor signals that encode vectors of the shifts in gaze direction called saccades. Since the late 1990s, our two laboratories have been applying whole cell patch-clamp techniques to in vitro slice preparations of rodent superior colliculus to analyze the structure and function of its circuitry at the cellular level. This review describes the results of these experiments and discusses their contributions to our understanding of the mechanisms responsible for sensorimotor integration in the superior colliculus. The experiments analyze vertical interactions between its superficial visuosensory and intermediate premotor layers and propose how they might contribute to express saccades and to saccadic suppression. They also compare and contrast the circuitry within each of these layers and propose how this circuitry might contribute to the selection of the targets for saccades and to the build-up of the premotor commands that precede saccades. Experiments also explore in vitro the roles of extrinsic inputs to the superior colliculus, including cholinergic inputs from the parabigeminal and parabrachial nuclei and GABAergic inputs from the substantia nigra pars reticulata, in modulating the activity of the collicular circuitry. The results extend and clarify our understanding of the multiple roles the superior colliculus plays in sensorimotor integration.

Surround Motion Silences Signals From Same-Direction Motion

Neri, P., Levi, D. — Wed, 04 Nov 2009 11:06:50 PST

The response of motion-sensitive neurons to stimuli presented within their receptive field is often affected by stimulation in the surrounding region. These effects have perceptually relevant consequences that can be measured using behavioral techniques. We used psychophysical reverse correlation to characterize directional selectivity in human observers while they processed a local motion stimulus and studied the effect of adding an additional motion signal in the surrounding region. The surround had no effect on response gain for signals of opposite direction but selectively reduced gain for those of same direction. Surprisingly this reduction was close to 100%, effectively amounting to a gating process whereby signals of same direction were completely silenced. Our data indicate that by far the most prominent perceptual manifestation of center-surround antagonism is gain suppression by motion in the same direction without any appreciable change in directional tuning.

Organization of Hue Selectivity in Macaque V2 Thin Stripes

Lim, H., Wang, Y., Xiao, Y., Hu, M., Felleman, D. J. — Wed, 04 Nov 2009 11:06:50 PST

V2 has long been recognized to contain functionally distinguishable compartments that are correlated with the stripelike pattern of cytochrome oxidase activity. Early electrophysiological studies suggested that color, direction/disparity, and orientation selectivity were largely segregated in the thin, thick, and interstripes, respectively. Subsequent studies revealed a greater degree of homogeneity in the distribution of response properties across stripes, yet color-selective cells were still found to be most prevalent in the thin stripes. Optical recording studies have demonstrated that thin stripes contain both color-preferring and luminance-preferring modules. These thin stripe color-preferring modules contain spatially organized hue maps, whereas the luminance-preferring modules contain spatially organized luminance-change maps. In this study, the neuronal basis of these hue maps was determined by characterizing the selectivity of neurons for isoluminant hues in multiple penetrations within previously characterized V2 thin stripe hue maps. The results indicate that neurons within the superficial layers of V2 thin stripe hue maps are organized into columns whose aggregated hue selectivity is closely related to the hue selectivity of the optically defined hue maps. These data suggest that thin stripes contain hue maps not simply because of their moderate percentage of hue-selective neurons, but because of the columnar and tangential organization of hue selectivity.

Dependence of the Roll Angular Vestibuloocular Reflex (aVOR) on Gravity

Yakushin, S. B., Xiang, Y., Cohen, B., Raphan, T. — Wed, 04 Nov 2009 11:06:50 PST

Little is known about the dependence of the roll angular vestibuloocular reflex (aVOR) on gravity or its gravity-dependent adaptive properties. To study gravity-dependent characteristics of the roll aVOR, monkeys were oscillated about a naso-occipital axis in darkness while upright or tilted. Roll aVOR gains were largest in the upright position and decreased by 7–15% as animals were tilted from the upright. Thus the unadapted roll aVOR gain has substantial gravitational dependence. Roll gains were also decreased or increased by 0.25 Hz, in- or out-of-phase rotation of the head and the visual surround while animals were prone, supine, upright, or in side-down positions. Gain changes, determined as a function of head tilt, were fit with a sinusoid; the amplitudes represented the amount of the gravity-dependent gain change, and the bias, the gravity-independent gain change. Gravity-dependent gain changes were absent or substantially smaller in roll (5%) than in yaw (25%) or pitch (17%), whereas gravity-independent gain changes were similar for roll, pitch, and yaw (20%). Thus the high-frequency roll aVOR gain has an inherent dependence on head orientation re gravity in the unadapted state, which is different from the yaw/pitch aVORs. This inherent gravitational dependence may explain why the adaptive circuits are not active when the head is tilted re gravity during roll aVOR adaptation. These behavioral differences support the idea that there is a fundamental difference in the central organization of canal-otolith convergence of the roll and yaw/pitch aVORs.

Context-Dependent Effects of NMDA Receptors on Precise Timing Information at the Endbulb of Held in the Cochlear Nucleus

Pliss, L., Yang, H., Xu-Friedman, M. A. — Wed, 04 Nov 2009 11:06:50 PST

Many synapses contain both AMPA receptors (AMPAR) and N-methyl-d-aspartate receptors (NMDAR), but their different roles in synaptic computation are not clear. We address this issue at the auditory nerve fiber synapse (called the endbulb of Held), which is formed on bushy cells of the cochlear nucleus. The endbulb refines and relays precise temporal information to nuclei responsible for sound localization. The endbulb has a number of specializations that aid precise timing, including AMPAR-mediated excitatory postsynaptic currents (EPSCs) with fast kinetics. Voltage-clamp experiments in mouse brain slices revealed that slow NMDAR EPSCs are maintained at mature endbulbs, contributing a peak conductance of around 10% of the AMPAR-mediated EPSC. During repetitive synaptic activity, AMPAR EPSCs depressed and NMDAR EPSCs summated, thereby increasing the relative importance of NMDARs. This could impact temporal precision of bushy cells because of the slow kinetics of NMDARs. We tested this by blocking NMDARs and quantifying bushy cell spike timing in current clamp when single endbulbs were activated. These experiments showed that NMDARs contribute to an increased probability of firing, shorter latency, and reduced jitter. Dynamic-clamp experiments confirmed this effect and showed it was dose-dependent. Bushy cells can receive inputs from multiple endbulbs. When we applied multiple synaptic inputs in dynamic clamp, NMDARs had less impact on spike timing. NMDAR conductances much higher than mature levels could disrupt spiking, which may explain its downregulation during development. Thus mature NMDAR expression can support the conveying of precise temporal information at the endbulb, depending on the stimulus conditions.

Long-Lasting Context Dependence Constrains Neural Encoding Models in Rodent Auditory Cortex

Asari, H., Zador, A. M. — Wed, 04 Nov 2009 11:06:50 PST

Acoustic processing requires integration over time. We have used in vivo intracellular recording to measure neuronal integration times in anesthetized rats. Using natural sounds and other stimuli, we found that synaptic inputs to auditory cortical neurons showed a rather long context dependence, up to ≥4 s ( ~ 1 s), even though sound-evoked excitatory and inhibitory conductances per se rarely lasted 100 ms. Thalamic neurons showed only a much faster form of adaptation with a decay constant <100 ms, indicating that the long-lasting form originated from presynaptic mechanisms in the cortex, such as synaptic depression. Restricting knowledge of the stimulus history to only a few hundred milliseconds reduced the predictable response component to about half that of the optimal infinite-history model. Our results demonstrate the importance of long-range temporal effects in auditory cortex and suggest a potential neural substrate for auditory processing that requires integration over timescales of seconds or longer, such as stream segregation.

Time Course of Activity in Itch-Related Brain Regions: A Combined MEG-fMRI Study

Wed, 04 Nov 2009 11:06:51 PST

Functional neuroimaging studies have identified itch-related brain regions. However, no study has investigated the temporal aspect of itch-related brain processing. Here this issue was investigated using electrically evoked itch in ten healthy adults. Itch stimuli were applied to the left wrist and brain activity was measured using magnetoencephalography (MEG) and functional magnetic resonance imaging (fMRI). In the MEG experiment, the magnetic responses evoked by the itch stimuli were observed in the contralateral and ipsilateral frontotemporal regions. The dipoles associated with the magnetic responses were mainly located in the contralateral (nine subjects) and ipsilateral (eight subjects) secondary somatosensory cortex (SII)/insula, which were also activated by the itch stimuli in the fMRI experiment. We also observed an itch-related magnetic response in the posterior part of the centroparietal region in six subjects. MEG and fMRI data showed that the magnetic response in this region was mainly associated with itch-related activation of the precuneus. The latency was significantly longer in the ipsilateral than that in the contralateral SII/insula, suggesting the difference to be associated with transmission in the callosal fibers. The timing of activation of the precuneus was between those of the contralateral and ipsilateral SII/insula. Other sources were located in the premotor, primary motor, and anterior cingulate cortices (one subject each). This study is the first to demonstrate part of the time course of itch-related brain processing. Combining methods with high temporal and spatial resolution (e.g., MEG and fMRI) would be useful to investigate the temporal aspect of the brain mechanism of itch.

Asymmetric Changes in Cutaneous Reflexes After a Partial Spinal Lesion and Retention Following Spinalization During Locomotion in the Cat

Frigon, A., Barriere, G., Leblond, H., Rossignol, S. — Wed, 04 Nov 2009 11:06:51 PST

Locomotion involves dynamic interactions between the spinal cord, supraspinal signals, and peripheral sensory inputs. After incomplete spinal cord injury (SCI), interactions are disrupted, and remnant structures must optimize function to maximize locomotion. We investigated if cutaneous reflexes are altered following a unilateral partial spinal lesion and whether changes are retained within spinal circuits after complete spinal transection (i.e., spinalization). Four cats were chronically implanted with recording and stimulating electrodes. Cutaneous reflexes were evoked with cuff electrodes placed around left and right superficial peroneal nerves. Control data, consisting of hindlimb kinematics and electromyography (bursts of muscular activity and cutaneous reflexes), were recorded during treadmill locomotion. After stable control data were achieved (53–67 days), a partial spinal lesion was made at the 10th or 11th thoracic segment (T₁₀–T₁₁) on the left side. Cats were trained to walk after the partial lesion, and following a recovery period (64–80 days), a spinalization was made at T₁₃. After the partial lesion, changes in short-latency excitatory (P1) homologous responses between hindlimbs, evoked during swing, were largely asymmetric in direction relative to control values, whereas changes in longer-latency excitatory (P2) and crossed responses were largely symmetric in direction. After spinalization, cats could display hindlimb locomotion within 1 day. Early after spinalization, reflex changes persisted a few days, but over time homologous P1 responses increased symmetrically toward or above control levels. Therefore changes in cutaneous reflexes after the partial lesion and retention following spinalization indicate an important spinal plasticity after incomplete SCI.

Eye-Hand Coordination During Target Selection in a Pop-Out Visual Search

Song, J.-H., McPeek, R. M. — Wed, 04 Nov 2009 11:06:51 PST

We examined the coordination of saccades and reaches in a visual search task in which monkeys were rewarded for reaching to an odd-colored target among distractors. Eye movements were unconstrained, and monkeys typically made one or more saccades before initiating a reach. Target selection for reaching and saccades was highly correlated with the hand and eyes landing near the same final stimulus both for correct reaches to the target and for incorrect reaches to a distractor. Incorrect reaches showed a bias in target selection: they were directed to the distractor in the same hemifield as the target more often than to other distractors. A similar bias was seen in target selection for the initial saccade in correct reaching trials with multiple saccades. We also examined the temporal coupling of saccades and reaches. In trials with a single saccade, a reaching movement was made after a fairly stereotyped delay. In multiple-saccade trials, a reach to the target could be initiated near or even before the onset of the final target-directed saccade. In these trials, the initial trajectory of the reach was often directed toward the fixated distractor before veering toward the target around the time of the final saccade. In virtually all cases, the eyes arrived at the target before the hand, and remained fixated until reach completion. Overall, these results are consistent with flexible temporal coupling of saccade and reach initiation, but fairly tight coupling of target selection for the two types of action.

Effects of Canal Plugging on the Vestibuloocular Reflex and Vestibular Nerve Discharge During Passive and Active Head Rotations

Sadeghi, S. G., Goldberg, J. M., Minor, L. B., Cullen, K. E. — Wed, 04 Nov 2009 11:06:51 PST

Mechanical occlusion (plugging) of the slender ducts of semicircular canals has been used in the clinic as well as in basic vestibular research. Here, we investigated the effect of canal plugging in two macaque monkeys on the horizontal vestibuloocular reflex (VOR) and the responses of vestibular-nerve afferents during passive head rotations. Afferent responses to active head movements were also studied. The horizontal VOR gain decreased after plugging to <0.1 for frequencies <2 Hz but rose to about 0.6 as frequency was increased to 15 Hz. Afferents innervating plugged horizontal canals had response sensitivities that increased with the frequency of passive rotations from <0.01 (spikes/s)/(°/s) at 0.5 Hz to values of about 0.2 and 0.5 (spikes/s)/(°/s) at 8 Hz for regular and irregular afferents, respectively (<50% of responses in controls). An increase in phase lead was also noted following plugging in afferent discharge, but not in the VOR. Because the phase discrepancy between the VOR and afferent discharge is much larger than that seen in control animals, this suggests that central adaptation shapes VOR dynamics following plugging. The effect of canal plugging on afferent responses can be modeled as an increase in stiffness and a reduction in the dominant time constant and gain in the transfer function describing canal dynamics. Responses were also evident during active head rotations, consistent with the frequency content of these movements. We conclude that canal plugging in macaques is effective only at frequencies <2 Hz. At higher frequencies, afferents show significant responses, with a nearly 90° phase lead, such that they encode near-rotational acceleration. Our results demonstrate that afferents innervating plugged canals respond robustly during voluntary movements, a finding that has implications for understanding the effects of canal plugging in clinical practice.

Visual Field Maps, Population Receptive Field Sizes, and Visual Field Coverage in the Human MT+ Complex

Amano, K., Wandell, B. A., Dumoulin, S. O. — Wed, 04 Nov 2009 11:06:51 PST

Human neuroimaging experiments typically localize motion-selective cortex (MT+) by contrasting responses to stationary and moving stimuli. It has long been suspected that MT+, located on the lateral surface at the temporal–occipital (TO) boundary, contains several distinct visual field maps, although only one coarse map has been measured. Using a novel functional MRI model–based method we identified two maps—TO-1 and TO-2—and measured population receptive field (pRF) sizes within these maps. The angular representation of the first map, TO-1, has a lower vertical meridian on its posterior side at the boundary with the lateral–occipital cortex (i.e., the LO-2 portion). The angular representation continues through horizontal to the upper vertical meridian at the boundary with the second map, TO-2. The TO-2 angle map reverses from upper to lower visual field at increasingly anterior positions. The TO maps share a parallel eccentricity map in which center-to-periphery is represented in the ventral-to-dorsal direction; both maps have an expanded foveal representation. There is a progressive increase in the pRF size from V1/2/3 to LO-1/2 and TO-1/2, with the largest pRF sizes in TO-2. Further, within each map the pRF size increases as a function of eccentricity. The visual field coverage of both maps extends into the ipsilateral visual field, with larger sensitivity to peripheral ipsilateral stimuli in TO-2 than that in TO-1. The TO maps provide a functional segmentation of human motion-sensitive cortex that enables a more complete characterization of processing in human motion-selective cortex.

Brain Switch for Reflex Micturition Control Detected by fMRI in Rats

Wed, 04 Nov 2009 11:06:51 PST

The functions of the lower urinary tract are controlled by complex pathways in the brain that act like switching circuits to voluntarily or reflexly shift the activity of various pelvic organs (bladder, urethra, urethral sphincter, and pelvic floor muscles) from urine storage to micturition. In this study, functional magnetic resonance imaging (fMRI) was used to visualize the brain switching circuits controlling reflex micturition in anesthetized rats. The fMRI images confirmed the hypothesis based on previous neuroanatomical and neurophysiological studies that the brain stem switch for reflex micturition control involves both the periaqueductal gray (PAG) and the pontine micturition center (PMC). During storage, the PAG was activated by afferent input from the urinary bladder while the PMC was inactive. When bladder volume increased to the micturition threshold, the switch from storage to micturition was associated with PMC activation and enhanced PAG activity. A complex brain network that may regulate the brain stem micturition switch and control storage and voiding was also identified. Storage was accompanied by activation of the motor cortex, somatosensory cortex, cingulate cortex, retrosplenial cortex, thalamus, putamen, insula, and septal nucleus. On the other hand, micturition was associated with: 1) increased activity of the motor cortex, thalamus, and putamen; 2) a shift in the locus of activity in the cingulate and insula; and 3) the emergence of activity in the hypothalamus, substantia nigra, globus pallidus, hippocampus, and inferior colliculus. Understanding brain control of reflex micturition is important for elucidating the mechanisms underlying neurogenic bladder dysfunctions including frequency, urgency, and incontinence.

Neural Representations of Complex Temporal Modulations in the Human Auditory Cortex

Ding, N., Simon, J. Z. — Wed, 04 Nov 2009 11:06:51 PST

Natural sounds such as speech contain multiple levels and multiple types of temporal modulations. Because of nonlinearities of the auditory system, however, the neural response to multiple, simultaneous temporal modulations cannot be predicted from the neural responses to single modulations. Here we show the cortical neural representation of an auditory stimulus simultaneously frequency modulated (FM) at a high rate, f_FM 40 Hz, and amplitude modulation (AM) at a slow rate, f_AM <15 Hz. Magnetoencephalography recordings show fast FM and slow AM stimulus features evoke two separate but not independent auditory steady-state responses (aSSR) at f_FM and f_AM, respectively. The power, rather than phase locking, of the aSSR of both decreases with increasing stimulus f_AM. The aSSR at f_FM is itself simultaneously amplitude modulated and phase modulated with fundamental frequency f_AM, showing that the slow stimulus AM is not only encoded in the neural response at f_AM but also encoded in the instantaneous amplitude and phase of the neural response at f_FM. Both the amplitude modulation and phase modulation of the aSSR at f_FM are most salient for low stimulus f_AM but remain observable at the highest tested f_AM (13.8 Hz). The instantaneous amplitude of the aSSR at f_FM is successfully predicted by a model containing temporal integration on two time scales, ~25 and ~200 ms, followed by a static compression nonlinearity.

Age-Related Declines in Visuospatial Working Memory Correlate With Deficits in Explicit Motor Sequence Learning

Bo, J., Borza, V., Seidler, R. D. — Wed, 04 Nov 2009 11:06:51 PST

Numerous studies have shown that older adults exhibit deficits in motor sequence learning, but the mechanisms underlying this effect remain unclear. Our recent work has shown that visuospatial working-memory capacity predicts the rate of motor sequence learning and the length of motor chunks formed during explicit sequence learning in young adults. In the current study, we evaluate whether age-related deficits in working memory explain the reduced rate of motor sequence learning in older adults. We found that older adults exhibited a correlation between visuospatial working-memory capacity and motor sequence chunk length, as we observed previously in young adults. In addition, older adults exhibited an overall reduction in both working-memory capacity and motor chunk length compared with that of young adults. However, individual variations in visuospatial working-memory capacity did not correlate with the rate of learning in older adults. These results indicate that working memory declines with age at least partially explain age-related differences in explicit motor sequence learning.

Changing the "When" and "What" of Intended Actions

Obhi, S. S., Matkovich, S., Chen, R. — Wed, 04 Nov 2009 11:06:51 PST

Humans often have to modify the timing and/or type of their planned actions on the basis of new sensory information. In the present experiments, participants planned to make a right index finger keypress 3 s after a warning stimulus but on some trials were interrupted by a temporally unpredictable auditory tone prompting the same action (experiment 1) or a different action (experiment 2). In experiment 1, by comparing the reaction time (RT) to tones presented at different stages of the preparatory period to RT in a simple reaction time condition, we determined the cost of switching from an internally generated mode of response production to an externally triggered mode in situations requiring only a change in when an action is made (i.e., when the tone prompts the action at a different time from the intended time of action). Results showed that the cost occurred for interruption tones delivered 200 ms after a warning stimulus and remained relatively stable throughout most of the preparatory period with a reduction in the magnitude of the cost during the last 200 ms prior to the intended time of movement. In experiment 2, which included conditions requiring a change in both when and what action is produced on the tone, results show a larger cost when the switched to action is different from the action being prepared. We discuss our results in the light of neurophysiological experiments on motor preparation and suggest that intending to act is accompanied by a general inhibitory mechanism preventing premature motor output and a specific excitatory process pertaining to the intended movement. Interactions between these two mechanisms could account for our behavioral results.

Intrinsic Neuronal Excitability Is Reversibly Altered by a Single Experience in Fear Conditioning

McKay, B. M., Matthews, E. A., Oliveira, F. A., Disterhoft, J. F. — Wed, 04 Nov 2009 11:06:51 PST

Learning is known to cause alterations in intrinsic cellular excitability but, to date, these changes have been seen only after multiple training trials. A powerful learning task that can be quickly acquired and extinguished with a single trial is fear conditioning. Rats were trained and extinguished on a hippocampus-dependent form of fear conditioning to determine whether learning-related changes in intrinsic excitability could be observed after a few training trials and a single extinction trial. Following fear training, hippocampal slices were made and intrinsic excitability was assayed via whole cell recordings from CA1 neurons. Alterations in intrinsic excitability, assayed by the postburst afterhyperpolarization and firing frequency accommodation, were observed after only three trials of contextual or trace-cued fear conditioning. Animals that had been trained in contextual and trace-cued fear were then extinguished. Context fear-conditioned animals extinguished in a single trial and the changes in intrinsic excitability were reversed. Trace-cue conditioned animals only partially extinguished in a single trial and reductions in excitability remained. Thus a single learning experience is sufficient to alter intrinsic excitability. This dramatically extends observations of learning-specific changes in intrinsic neuronal excitability previously observed in paradigms requiring many training trials, suggesting the excitability changes have a basic role in acquiring new information.

Transformation in the Neural Code for Whisker Deflection Direction Along the Lemniscal Pathway

Bale, M. R., Petersen, R. S. — Wed, 04 Nov 2009 11:06:51 PST

A prominent characteristic of neurons in the whisker system is their selectivity to the direction in which a whisker is deflected. The aim of this study was to determine how information about whisker direction is encoded at successive levels of the lemniscal pathway. We made extracellular recordings under identical conditions from the trigeminal ganglion, ventro-posterior medial thalamus (VPM), and barrel cortex while varying the direction of whisker deflection. We found a marked increase in the variability of single unit responses along the pathway. To study the consequences of this for information processing, we quantified the responses using mutual information. VPM units conveyed 48% of the mutual information conveyed by ganglion units, and cortical units conveyed 12%. The fraction of neuronal bandwidth used for transmitting direction information decreased from 40% in the ganglion to 24% in VPM and 5% in barrel cortex. To test whether, in cortex, population coding might compensate for this information loss, we made simultaneous recordings. We found that cortical neuron pairs conveyed 2.1 times the mutual information conveyed by single neurons. Overall, these findings indicate a marked transformation from a subcortical neural code based on small numbers of reliable neurons to a cortical code based on populations of unreliable neurons. However, the basic form of the neural code in ganglion, thalamus, and cortex was similar—at each stage, the first poststimulus spike carried the majority of the information.

Question of Reference Frames: Visual Direction-Selective Neurons in the Accessory Optic System of Goldfish

Masseck, O. A., Hoffmann, K.-P. — Wed, 04 Nov 2009 11:06:51 PST

We investigated if visual direction-selective neurons in the pretectal area (APT) of goldfish (Carassius auratus auratus) preferred visual stimuli resulting from rotations around axes corresponding to the best responsive axes of the semicircular canals [optic flow that is consistent to a maximal activation of the horizontal canal pair (yaw), to a maximal activation of the right anterior/left posterior semicircular canal pair (RALP), and to a maximal activation of the left anterior/right posterior semicircular canal pair (LARP)]. Our sample of neurons recorded in the left pretectum had two preferred axes of rotation: first, rotation around the yaw axis and second, rotation around the RALP axis. Both axes of rotation correspond to best responsive axes of the semicircular canals. For this reason, coding in a reference frame defined by the vestibular system or the pulling direction of the eye muscles is suggested. In our population of recorded APT neurons, we did not find segregation of different preferred axes of rotation into different anatomical structures. Furthermore in all axes no bias for clockwise or counterclockwise rotations was obvious. This is particularly noteworthy for the yaw axis because preference for temporo-nasal and naso-temporal rotations was found at the same recording side. Hence we conclude that in fish the accessory optic system may consist of one nucleus on each side of the midbrain only, the APT. Segregation into different nuclei coding for different axes and different senses of rotation probably first developed in amphibians.

Generalization of Visuomotor Learning Between Bilateral and Unilateral Conditions

Wang, J., Sainburg, R. L. — Wed, 04 Nov 2009 11:06:51 PST

A long history of behavioral and physiological research has suggested that bilateral coordination invokes unique neural processes that are not involved in unilateral movements. This hypothesis predicts that motor learning should show limited transfer between unilateral and bilateral conditions, which is consistent with a recent finding that indicated partial, but not complete, transfer of learning between the two conditions. However, during learning of new motor skills, transformations must also be made between visual and proprioceptive coordinate systems, a process that may occur upstream to the processes that differentiate bilateral from unilateral movements. We now investigate whether visuomotor adaptations are shared between unilateral and bilateral movement conditions. Our results indicate substantial transfer from bilateral to subsequent unilateral conditions for both arms. Interestingly, whereas the nondominant arm never showed complete adaptation to visual rotation under bilateral conditions, this interference, or lack of improvement, in bilateral performance did not disturb the visuomotor adaptation process or transfer, as reflected by superb unilateral performances immediately following the bilateral conditions. These findings unambiguously indicate that visuomotor adaptation can extensively generalize between bilateral and unilateral conditions, thus suggesting a substantial overlap in the neural processes underlying visuomotor transformations between the two movement conditions. Our findings provide support for a two-stage model of motor planning, in which the visuomotor transformation process precedes the processes that convert the visuomotor plan into effector-specific commands that incorporate bilateral synergies and that result in the forces that determine motion.

On the Open-Loop and Feedback Processes That Underlie the Formation of Trajectories During Visual and Nonvisual Locomotion in Humans

Pham, Q.-C., Hicheur, H. — Wed, 04 Nov 2009 11:06:51 PST

We investigated the nature of the control mechanisms at work during goal-oriented locomotion. In particular, we tested the effects of vision, locomotor speed, and the presence of via points on the geometric and kinematic properties of locomotor trajectories. We first observed that the average trajectories recorded in visual and nonvisual locomotion were highly comparable, suggesting the existence of vision-independent processes underlying the formation of locomotor trajectories. Then by analyzing and comparing the variability around the average trajectories across different experimental conditions, we were able to demonstrate the existence of on-line feedback control in both visual and nonvisual locomotion and to clarify the relations between visual and nonvisual control strategies. Based on these insights, we designed a model in which maximum-smoothness and optimal feedback control principles account, respectively, for the open-loop and feedback processes. Taken together, the experimental and modeling findings provide a novel understanding of the nature of the motor, sensory, and "navigational" processes underlying goal-oriented locomotion.

Transfer of Dynamic Learning Across Postures

Ahmed, A. A., Wolpert, D. M. — Wed, 04 Nov 2009 11:06:51 PST

When learning a difficult motor task, we often decompose the task so that the control of individual body segments is practiced in isolation. But on re-composition, the combined movements can result in novel and possibly complex internal forces between the body segments that were not experienced (or did not need to be compensated for) during isolated practice. Here we investigate whether dynamics learned in isolation by one part of the body can be used by other parts of the body to immediately predict and compensate for novel forces between body segments. Subjects reached to targets while holding the handle of a robotic, force-generating manipulandum. One group of subjects was initially exposed to the novel robot dynamics while seated and was then tested in a standing position. A second group was tested in the reverse order: standing then sitting. Both groups adapted their arm dynamics to the novel environment, and this movement learning transferred between seated and standing postures and vice versa. Both groups also generated anticipatory postural adjustments when standing and exposed to the force field for several trials. In the group that had learned the dynamics while seated, the appropriate postural adjustments were observed on the very first reach on standing. These results suggest that the CNS can immediately anticipate the effect of learned movement dynamics on a novel whole-body posture. The results support the existence of separate mappings for posture and movement, which encode similar dynamics but can be adapted independently.

5-HT and GABA Modulate Intrinsic Excitability of Type I Interneurons in Hermissenda

Jin, N. G., Tian, L.-M., Crow, T. — Wed, 04 Nov 2009 11:06:51 PST

The sensory neurons (photoreceptors) in the visual system of Hermissenda are one site of plasticity produced by Pavlovian conditioning. A second site of plasticity produced by conditioning is the type I interneurons in the cerebropleural ganglia. Both photoreceptors and statocyst hair cells of the graviceptive system form monosynaptic connections with identified type I interneurons. Two proposed neurotransmitters in the graviceptive system, serotonin (5-HT) and -aminobutyric acid (GABA), have been shown to modify synaptic strength and intrinsic neuronal excitability in identified photoreceptors. However, the potential role of 5-HT and GABA in plasticity of type I interneurons has not been investigated. Here we show that 5-HT increased the peak amplitude of light-evoked complex excitatory postsynaptic potentials (EPSPs), enhanced intrinsic excitability, and increased spike activity of identified type I_e(A) interneurons. In contrast, 5-HT decreased spike activity and intrinsic excitability of type I_e(B) interneurons. The classification of two categories of type I_e interneurons was also supported by the observation that 5-HT produced opposite effects on whole cell steady-state outward currents in type I_e interneurons. Serotonin produced a reduction in the amplitude of light-evoked complex inhibitory PSPs (IPSPs), increased spontaneous spike activity, decreased intrinsic excitability, and depolarized the resting membrane potential of identified type I_i interneurons. In contrast to the effects of 5-HT, GABA produced inhibition in both types of I_e interneurons and type I_i interneurons. These results show that 5-HT and GABA can modulate the intrinsic excitability of type I interneurons independent of the presynaptic effects of the same transmitters on excitability and synaptic efficacy of photoreceptors.

Distinct Electrophysiological Properties in Subtypes of Nonspiking Olfactory Local Interneurons Correlate With Their Cell Type-Specific Ca2+ Current Profiles

Husch, A., Paehler, M., Fusca, D., Paeger, L., Kloppenburg, P. — Wed, 04 Nov 2009 11:06:51 PST

A diverse population of local interneurons (LNs) helps to process, structure, and spatially represent olfactory information in the insect antennal lobe. In Periplaneta americana, we identified two subtypes of nonspiking local interneurons (type II LNs) by their distinct morphological and intrinsic electrophysiological properties. As an important step toward a better understanding of the cellular mechanisms that mediate odor information processing, we present a detailed analysis of their distinct voltage-activated Ca²⁺ currents, which clearly correlated with their distinct intrinsic electrophysiological properties. Both type II LNs did not posses voltage-activated Na⁺ currents and apparently innervated all glomeruli including the macroglomerulus. Type IIa LNs had significant longer and thicker low-order neurites and innervated each glomerulus entirely and homogeneously, whereas type IIb LNs innervated only parts of each glomerulus. All type II LNs were broadly tuned and responded to odorants of many chemical classes with graded changes in the membrane potential. Type IIa LNs responded with odor-specific elaborate patterns of excitation that could also include "spikelets" riding on the depolarizations and periods of inhibition. In contrast, type IIb LNs responded mostly with sustained, relatively smooth depolarizations. Consistent with the strong active membrane properties of type IIa LNs versus type IIb LNs, the voltage-activated Ca²⁺ current of type IIa LNs activated at more hyperpolarized membrane potentials and had a larger transient component.

Influence of Vagotomy on Monosynaptic Transmission at Second-Order Nucleus Tractus Solitarius Synapses

Swartz, J. B., Weinreich, D. — Wed, 04 Nov 2009 11:06:51 PST

Manipulations of vagal activity are used to treat medical pathologies, but the underlying CNS changes caused by these treatments are not well understood. Furthermore, heart and lung transplant as well as treatments for many gastrointestinal disorders result in section of the vagus nerve (vagotomy). Following unilateral vagotomy under isoflurane anesthesia of Sprague-Dawley rats, electrophysiological properties were recorded with whole cell patch techniques in horizontal brain stem slices. Vagotomy significantly reduced the median amplitude of evoked excitatory postsynaptic currents (evEPSCs; –121; n = 43) in the nucleus tractus solitarius (NTS) when compared with controls (–157 pA; n = 66; P < 0.05) but had no significant effect on the passive properties or on the average amplitude or frequency of miniature EPSCs. The degree of synaptic failure exhibited during a 50-Hz train of stimuli was used to define two separate classes of synapses: "low failure" and "high failure" (HF); failure rates <5 and ≥5%, respectively. HF synapses had significantly smaller median evEPSCs (–88 vs. –184 pA; P < 0.05). After vagotomy, the percentage of HF synapses nearly doubled to 56% (n = 24/43) when compared with controls (30%; n = 20/66). Additionally, the overall percentage of failures after the second to fifth stimuli significantly increased by at least twofold. These results suggest that vagotomy causes a decrease in synaptic efficacy by both increasing the overall percentage of synaptic failures and shifting the population of NTS synapses toward more HF transmission. In addition, the alterations due to vagotomy are likely to be presynaptic in nature.

Bilateral Limb Phase Relationship and Its Potential to Alter Muscle Activity Phasing During Locomotion

Alibiglou, L., Lopez-Ortiz, C., Walter, C. B., Brown, D. A. — Wed, 04 Nov 2009 11:06:51 PST

It is well established that the sensorimotor state of one limb can influence another limb and therefore bilateral somatosensory inputs make an important contribution to interlimb coordination patterns. However, the relative contribution of interlimb pathways for modifying muscle activation patterns in terms of phasing is less clear. Here we studied adaptation of muscle activity phasing to the relative angular positions of limbs using a split-crank ergometer, where the cranks could be decoupled to allow different spatial angular position relationships. Twenty neurologically healthy individuals performed the specified pedaling tasks at different relative angular positions while surface electromyographic (EMG) signals were recorded bilaterally from eight lower extremity muscles. During each experiment, the relative angular crank positions were altered by increasing or decreasing their difference by randomly ordered increments of 30° over the complete cycle [0° (in phase pedaling); 30, 60, 90, 120, 150, and 180° (standard pedaling); and 210, 240, 270, 300, and 330° out of phase pedaling]. We found that manipulating the relative angular positions of limbs in a pedaling task caused muscle activity phasing changes that were either delayed or advanced, dependent on the relative spatial position of the two cranks and this relationship is well-explained by a sine curve. Further, we observed that the magnitude of phasing changes in biarticular muscles (like rectus femoris) was significantly greater than those of uniarticular muscles (like vastus medialis). These results are important because they provide new evidence that muscle phasing can be systematically influenced by interlimb pathways.

Differential Activation of Projection Neurons by Two Sensory Pathways Contributes to Motor Pattern Selection

Hedrich, U. B. S., Smarandache, C. R., Stein, W. — Wed, 04 Nov 2009 11:06:51 PST

Sensorimotor integration is known to occur at the level of motor circuits as well as in upstream interneurons that regulate motor activity. Here we show, using the crab stomatogastric nervous system (STNS) as a model, that different sensory systems affect the same set of projection neurons. However, they have qualitatively different effects on their activities (excitation vs. inhibition), and these differences contribute to the selection of motor patterns from multifunctional circuits. We compare the actions of the proprioceptive anterior gastric receptor (AGR) and the inferior ventricular (IV) neurons, which relay chemosensory information from the brain to the STNS, on modulatory commissural neurons 1 and 5 (MCN1 and MCN5) and commissural projection neuron 2 (CPN2) and their resulting actions on the gastric mill central pattern generating circuit in the stomatogastric ganglion. When stimulated, AGR and the IV neurons affect all three projection neurons but elicit distinct gastric mill rhythms. The effects of both sensory pathways on the projection neurons differ in the type of excitation provided to CPN2 and MCN5 (electrical vs. chemical) and the effect on MCN1 (direct inhibition by AGR vs. polysynaptic excitation by the IV neurons). The latter is functionally important because a restoration of MCN1 activity during the AGR rhythm made it more similar to that elicited by IV neuron stimulation. Our results thus support the hypothesis that sensory pathways activate different combinations of projection neurons to select distinct outputs from the same neuronal circuit.

Maintenance of Thalamic Epileptiform Activity Depends on the Astrocytic Glutamate-Glutamine Cycle

Bryant, A. S., Li, B., Beenhakker, M. P., Huguenard, J. R. — Wed, 04 Nov 2009 11:06:52 PST

The generation of prolonged neuronal activity depends on the maintenance of synaptic neurotransmitter pools. The astrocytic glutamate-glutamine cycle is a major mechanism for recycling the neurotransmitters GABA and glutamate. Here we tested the effect of disrupting the glutamate-glutamine cycle on two types of neuronal activity patterns in the thalamus: sleep-related spindles and epileptiform oscillations. In recording conditions believed to induce glutamine scarcity, epileptiform oscillations showed a progressive reduction in duration that was partially reversible by the application of exogenous glutamine (300 µM). Blocking uptake of glutamine into neurons with -(methylamino) isobutyric acid (5 mM) caused a similar reduction in oscillation duration, as did blocking neuronal GABA synthesis with 3-mercaptoproprionic acid (10 µM). However, comparable manipulations did not affect sleep spindles. Together, these results support a crucial role for the glutamate-glutamine cycle in providing the neurotransmitters necessary for the generation of epileptiform activity and suggest potential therapeutic approaches that selectively reduce seizure activity but maintain normal neuronal activity.

Multicomponent Control Strategy Underlying Production of Maximal Hand Velocity During Horizontal Arm Swing

Kim, Y.-K., Hinrichs, R. N., Dounskaia, N. — Wed, 04 Nov 2009 11:06:52 PST

Movement control responsible for generation of maximal hand velocity was studied on the example of horizontal arm swing that is a component of various sports activities. The movement was performed with the nondominant arm in similarity with the baseball bat swing. The task was to generate maximum hand velocity at a target. The movement included trunk long-axis rotation and horizontal shoulder and elbow extension. Kinematics and torque analyses were performed to study the organization of fastest movements and to compare trials representing the best and worst performance in each subject. Results revealed complex control strategy, with the trunk, shoulder, and elbow playing unique roles in generation of maximal hand velocity. The trunk provided a crucial contribution, directly, rotating the entire arm, and indirectly, exerting interaction torque that caused swift elbow extension. The major role of the shoulder was to transfer the mechanical effect of trunk motion to the elbow. However, the shoulder became the primary motion generator when the trunk reached its limits of rotation, revealing sequential organization of control. The role of the elbow was to maximally comply with passive influence of proximal joints. The findings are discussed in light of the leading joint hypothesis that offers a straightforward interpretation of control of horizontal arm swing as well as practically efficient recommendations for increases in movement speed. The revealed role of intersegmental dynamics in production of high movement speed suggests that movement slowness characteristic for some motor disorders may be partially a compensatory strategy that facilitates regulation of interaction torque.

Area Summation in Human Visual System: Psychophysics, fMRI, and Modeling

Nurminen, L., Kilpelainen, M., Laurinen, P., Vanni, S. — Wed, 04 Nov 2009 11:06:52 PST

Contextual modulation is a fundamental feature of sensory processing, both on perceptual and on single-neuron level. When the diameter of a visual stimulus is increased, the firing rate of a cell typically first increases (summation field) and then decreases (surround field). Such an area summation function draws a comprehensive profile of the receptive field structure of a neuron, including areas outside the classical receptive field. We investigated area summation in human vision with psychophysics and functional magnetic resonance imaging (fMRI). The stimuli were similar to those used drifting sine wave gratings in previous macaque single-cell area summation studies. A model was developed to facilitate comparison of area summation in fMRI to area summation in psychophysics and single cells. The model consisted of units with an antagonistic receptive field structure found in single cells in the primary visual cortex. The receptive field centers of the model neurons were distributed in the region of the visual field covered by a single voxel. The measured area summation functions were qualitatively similar to earlier single-cell data. The model with parameters derived from psychophysics captured the spatial structure of the summation field in the primary visual cortex as measured with fMRI. The model also generalized to a novel situation in which the neural population was displaced from the stimulus center. The current study shows that contextual modulation arises from similar spatially antagonistic and overlapping excitatory and inhibitory mechanisms, both in single cells and in human vision.

Postural Feedback Scaling Deficits in Parkinson's Disease

Kim, S., Horak, F. B., Carlson-Kuhta, P., Park, S. — Wed, 04 Nov 2009 11:06:52 PST

Many differences in postural responses have been associated with age and Parkinson's disease (PD), but until now there has been no quantitative model to explain these differences. We developed a feedback control model of body dynamics that could reproduce the postural responses of young subjects, elderly subjects, and subjects with PD, and we investigated whether the postural impairments of subjects with PD can be described as an abnormal scaling of postural feedback gain. Feedback gains quantify how the nervous system generates compensatory joint torques based on kinematic responses. Seven subjects in each group experienced forward postural perturbations to seven different backward support surface translations ranging from 3- to 15-cm amplitudes and with a constant duration of 275 ms. Ground reaction forces and joint kinematics were measured to obtain joint torques from inverse dynamics. A full-state feedback controller with a two-segment body dynamic model was used to simulate joint kinematics and kinetics in response to perturbations. Results showed that all three subject groups gradually scaled postural feedback gains as a function of perturbation amplitudes, and the scaling started even before the maximum allowable ankle torque was reached. This result implies that the nervous system takes body dynamics into account and adjusts postural feedback gains to accommodate biomechanical constraints. PD subjects showed significantly smaller than normal ankle feedback gain with low scaling and larger hip feedback gain, which led to an early violation of the flat-foot constraint and unusually small (bradykinetic) postural responses. Our postural feedback control model quantitatively described the postural abnormality of the patients with PD as abnormal feedback gains and reduced ability to modify postural feedback gain with changes in postural challenge.

Adaptation to Visuomotor Rotation Through Interaction Between Posterior Parietal and Motor Cortical Areas

Tanaka, H., Sejnowski, T. J., Krakauer, J. W. — Wed, 04 Nov 2009 11:06:52 PST

Studying how motor adaptation to visuomotor rotation for one reach direction generalizes to other reach directions can provide insight into how visuomotor maps are represented and learned in the brain. Previous psychophysical studies have concluded that postadaptation generalization is restricted to a narrow range of directions around the training direction. A population-coding model that updates the weights between narrow Gaussian-tuned visual units and motor units on each trial reproduced experimental trial-by-trial learning curves for rotation adaptation and the generalization function measured postadaptation. These results suggest that the neurons involved in rotation adaptation have a relatively narrow directional tuning width (~23°). Population coding models with units having broader tuning functions (such as cosine tuning in motor cortex and Gaussian sum in the cerebellum) could not reproduce the narrow single-peaked generalization pattern. Visually selective neurons with narrow Gaussian tuning curves have been identified in posterior parietal cortex, making it a possible site of adaptation to visuomotor rotation. We propose that rotation adaptation proceeds through changes in synaptic weights between neurons in posterior parietal cortex and motor cortex driven by a prediction error computed by the cerebellum.

Substantia Nigra Output to Prefrontal Cortex Via Thalamus in Monkeys. I. Electrophysiological Identification of Thalamic Relay Neurons

Tanibuchi, I., Kitano, H., Jinnai, K. — Wed, 04 Nov 2009 11:06:52 PST

A few studies have been performed in primate basal ganglia–thalamo–prefrontal pathways. Nevertheless, their electrophysiological properties and anatomical arrangements remain obscure. This study examined them in nigro-thalamo-cortical pathways from the substantia nigra pars reticulata (SNr) to the frontal cortex (FRC) via the mediodorsal (MD) and ventral anterior (VA) thalamus in monkeys. First, single thalamocortical neurons with SNr input were identified by antidromic responses to FRC stimulation and by inhibitory orthodromic responses with short latencies (<5 ms) to SNr stimulation. Second, single nigrothalamic neurons were found by antidromic responses to stimulation of the portions of the MD and VA where the thalamocortical neurons were recorded. The inhibitory orthodromic responses in the thalamocortical neurons were considered to be monosynaptically induced by nigral stimulation because the latency distribution of the orthodromic responses in the thalamocortical neurons was similar to that of the antidromic responses in the nigrothalamic neurons. Almost all relay neurons in the rostrolateral MD received inhibitory afferents from the caudolateral SNr and projected to the prefrontal area ventral to the principal sulcus, which constituted the densest nigro-thalamo-cortical projections. Meanwhile, neurons in the VA received inhibitory signals from the whole rostrocaudal extent of the SNr and projected to wide regions of the FRC; neurons in its pars magnocellularis mostly projected to different prefrontal areas, while those in its pars parvocellularis projected to motor areas. This report substantiated the topography of thalamocortical neurons monosynaptically receiving inhibitory SNr input and projecting to the FRC in the primate MD and VA at the single-neuron level.

Substantia Nigra Output to Prefrontal Cortex Via Thalamus in Monkeys. II. Activity of Thalamic Relay Neurons in Delayed Conditional Go/No-Go Discrimination Task

Tanibuchi, I., Kitano, H., Jinnai, K. — Wed, 04 Nov 2009 11:06:52 PST

The present report investigated the involvement of primate nigro-thalamo-cortical projections in discrimination of visual signals with behavioral meaning. We tested the extracellular unit activity of mediodorsal (MD) and ventral anterior (VA) thalamic neurons monosynaptically receiving inhibitory input from the substantia nigra pars reticulata (SNr) and projecting to the frontal cortex in Japanese monkeys performing a delayed conditional go/no-go discrimination task. In the task two colored stimuli (S1, S2) intervened by delay period required the monkeys lifting a lever (go) or not (no-go); the same and different colored pairs of S1 and S2 meant go and no-go signals, respectively. Prominent task-relevant responses were sustained activity with color preference to S1 during delay period and S2-related activity with different firing rates between go and no-go trials. In particular, a high proportion of such go/no-go differential S2-related activity was found in thalamic relay neurons, receiving input from the caudolateral SNr and projecting to the prefrontal area (PSv) ventral to the principal sulcus, in the rostrolateral MD. The findings suggest that the caudolateral SNr–rostrolateral MD–PSv pathways may be possible conduits of signals coding the behavioral meaning of the visual stimuli and thus may be responsible for generating similar neuronal activity in the PSv.

Experience-Dependent Intrinsic Plasticity in Interneurons of Barrel Cortex Layer IV

Sun, Q.-Q. — Wed, 04 Nov 2009 11:06:52 PST

It is unclear whether intrinsic excitabilities of specific interneurons are modulated by sensory experiences. Here, I examined the intrinsic excitabilities of interneurons in "sensory-spared" and "sensory-deprived" cortices of GAD67-GFP mice. The results showed that whisker trimming, begun at postnatal day 7 for 3 wk, induced significant changes in intrinsic and firing properties of fast-spiking (FS) but not regular spiking nonpyramidal (RSNP) cells. Firing threshold, spike frequency, spike adaptation index, and input resistance of FS cells were significantly altered by sensory deprivation such that FS cells became less excitable. An up-regulation of IA currents in FS cells appeared to be responsible. Along with changes in the intrinsic properties of FS cells, whisker trimming also induced a robust reduction in the number of vesicular glutamate transporter 2 positive varicosities and parvalbumin expression and the strength of thalamocortical (TC) excitatory postsynaptic currents in FS cells in the "sensory-deprived barrels." The probability of spike induction by TC stimulus was reduced by 30% and the spike jitter was increased in sensory-deprived FS cells. These results suggest that the FS networks are selectively inhibited by sensory deprivation. The concurrent changes of intrinsic properties and expression of parvalbumin in FS but not RSNP cells with TC synapses support a contribution from the TC pathway and glutamate to sensory-induced activity-dependent intrinsic plasticity of inhibitory networks in barrel cortex.

Radial Biases in the Processing of Motion and Motion-Defined Contours by Human Visual Cortex

Clifford, C. W. G., Mannion, D. J., McDonald, J. S. — Wed, 04 Nov 2009 11:06:52 PST

Luminance gratings reportedly produce a stronger functional magnetic resonance imaging (fMRI) blood oxygen level–dependent (BOLD) signal in those parts of the retinotopic cortical maps where they are oriented radially to the point of fixation. We sought to extend this finding by examining anisotropies in the response of cortical areas V1–V3 to motion-defined contour stimuli. fMRI at 3 Tesla was used to measure the BOLD signal in the visual cortex of six human subjects. Stimuli were composed of strips of spatial white noise texture presented in an annular window. The texture in alternate strips moved in opposite directions (left–right or up–down). The strips themselves were static and tilted 45° left or right from vertical. Comparison with maps of the visual field obtained from phase-encoded retinotopic analysis revealed systematic patterns of radial bias. For motion, a stronger response to horizontal was evident within V1 and along the borders between V2 and V3. For orientation, the response to leftward tilted contours was greater in left dorsal and right ventral V1–V3. Radial bias for the orientation of motion-defined contours analogous to that reported previously for luminance gratings could reflect cue-invariant processing or the operation of distinct mechanisms subject to similar anisotropies in orientation tuning. Radial bias for motion might be related to the phenomenon of "motion streaks," whereby temporal integration by the visual system introduces oriented blur along the axis of motion. We speculate that the observed forms of radial bias reflect a common underlying anisotropy in the representation of spatiotemporal image structure across the visual field.

Direct Activation and Temporal Response Properties of Rabbit Retinal Ganglion Cells Following Subretinal Stimulation

Tsai, D., Morley, J. W., Suaning, G. J., Lovell, N. H. — Wed, 04 Nov 2009 11:06:52 PST

In the last decade several groups have been developing vision prostheses to restore visual perception to the profoundly blind. Despite some promising results from human trials, further understanding of the neural mechanisms involved is crucial for improving the efficacy of these devices. One of the techniques involves placing stimulating electrodes in the subretinal space between the photoreceptor layer and the pigment epithelium to evoke neural responses in the degenerative retina. This study used cell-attached and whole cell current-clamp recordings to investigate the responses of rabbit retinal ganglion cells (RGCs) following subretinal stimulation with 25-µm-diameter electrodes. We found that direct RGC responses with short latency (≤2 ms using 0.1-ms pulses) could be reliably elicited. The thresholds for these responses were reported for on, off, and on–off RGCs over pulse widths 0.1–5.0 ms. During repetitive stimulation these direct activation responses were more readily elicited than responses arising from stimulation of the retinal network. The temporal spiking characteristics of RGCs were characterized as a function of stimulus configurations. We found that the response profiles could be generalized into four classes with distinctive properties. Our results suggest that for subretinal vision prostheses short pulses are preferable for efficacy and safety considerations, and that direct activation of RGCs will be necessary for reliable activation during high-frequency stimulation.

Functional Asymmetries Revealed in Visually Guided Saccades: An fMRI Study

Wed, 04 Nov 2009 11:06:52 PST

Because eye movements are a fundamental tool for spatial exploration, we hypothesized that the neural bases of these movements in humans should be under right cerebral dominance, as already described for spatial attention. We used functional magnetic resonance imaging in 27 right-handed participants who alternated central fixation with either large or small visually guided saccades (VGS), equally performed in both directions. Hemispheric functional asymmetry was analyzed to identify whether brain regions showing VGS activation elicited hemispheric asymmetries. Hemispheric anatomical asymmetry was also estimated to assess its influence on the VGS functional lateralization. Right asymmetrical activations of a saccadic/attentional system were observed in the lateral frontal eye fields (FEF), the anterior part of the intraparietal sulcus (aIPS), the posterior third of the superior temporal sulcus (STS), the occipitotemporal junction (MT/V5 area), the middle occipital gyrus, and medially along the calcarine fissure (V1). The present rightward functional asymmetries were not related to differences in gray matter (GM) density/sulci positions between right and left hemispheres in the precentral, intraparietal, superior temporal, and extrastriate regions. Only V1 asymmetries were explained for almost 20% of the variance by a difference in the position of the right and left calcarine fissures. Left asymmetrical activations of a saccadic motor system were observed in the medial FEF and in the motor strip eye field along the Rolando sulcus. They were not explained by GM asymmetries. We suggest that the leftward saccadic motor asymmetry is part of a general dominance of the left motor cortex in right-handers, which must include an effect of sighting dominance. Our results demonstrate that, although bilateral by nature, the brain network involved in the execution of VGSs, irrespective of their direction, presented specific right and left asymmetries that were not related to anatomical differences in sulci positions.

Long-Latency Responses During Reaching Account for the Mechanical Interaction Between the Shoulder and Elbow Joints

Kurtzer, I., Pruszynski, J. A., Scott, S. H. — Wed, 04 Nov 2009 11:06:52 PST

Although considerable research indicates that reaching movements rely on knowledge of the arm's mechanical properties and environment to anticipate and counter predictable loads, far less research has examined whether this degree of sophistication is present for on-line corrections during reaching. Here we examine the R2/3 response to mechanical perturbations (45–100 ms, also called the long-latency reflex), which is highly flexible and includes the fastest possible contribution from primary motor cortex, a key neural substrate for self-initiated action. Torque perturbations were occasionally and unexpectedly applied to the subject's shoulder and/or elbow in the course of performing reaching movements. Critically, these perturbations would evoke different patterns of feedback corrections from a shoulder extensor muscle if it countered only the local shoulder displacement relative to unperturbed motion or accounted for the mechanical interactions between the shoulder and elbow joints and countered the underlying shoulder torque. Our results show that the earliest response (R1: 20–45 ms) reflected local shoulder displacement, whereas the R2/3 response (45–100 ms) reflected knowledge of multijoint dynamics. Moreover, the same pattern of feedback occurred whether the shoulder muscle helped initiate the movement (during its agonist phase) or helped terminate the movement (during its antagonist phase). These results contribute to the accumulating evidence that highly sophisticated feedback control underlies motor behavior and are consistent with a shared neural substrate, such as primary motor cortex, for feedforward and feedback control.

Type of Featural Attention Differentially Modulates hMT+ Responses to Illusory Motion Aftereffects

Wed, 04 Nov 2009 11:06:52 PST

Activity in the human motion complex (hMT⁺/V5) is related to the perception of motion, be it either real surface motion or an illusion of motion such as apparent motion (AM) or motion aftereffect (MAE). It is a long-lasting debate whether illusory motion-related activations in hMT⁺ represent the motion itself or attention to it. We have asked whether hMT⁺ responses to MAEs are present when shifts in arousal are suppressed and attention is focused on concurrent motion versus nonmotion features. Significant enhancement of hMT⁺ activity was observed during MAEs when attention was focused either on concurrent spatial angle or color features. This observation was confirmed by direct comparison of adapting (MAE inducing) versus nonadapting conditions. In contrast, this effect was diminished when subjects had to report on concomitant speed changes of superimposed AM. The same finding was observed for concomitant orthogonal real motion (RM), suggesting that selective attention to concurrent illusory or real motion was interfering with the saliency of MAE signals in hMT⁺. We conclude that MAE-related changes in the global activity of hMT⁺ are present provided selective attention is not focused on an interfering feature such as concurrent motion. Accordingly, there is a genuine MAE-related motion signal in hMT⁺ that is neither explained by shifts in arousal nor by selective attention.

Transient Firing of Dorsal Raphe Neurons Encodes Diverse and Specific Sensory, Motor, and Reward Events

Ranade, S. P., Mainen, Z. F. — Wed, 04 Nov 2009 11:06:52 PST

Serotonin (5-hydroxytryptamine [5-HT]) is known to influence a wide range of behaviors and physiological processes, but relatively little is known about events that trigger 5-HT release. To address this issue, we recorded from neurons in the dorsal raphe nucleus (DRN) in rats performing an odor-guided spatial decision task. A large fraction of DRN neurons showed transient firing time locked to behavioral events on timescales as little as 20 ms. DRN transients were sometimes correlated with reward parameters, but also encoded specific sensorimotor events, including stimulus identity and response direction. These behavioral correlates were diverse but showed no apparent relationship with waveform or other firing properties indicative of neurochemical identity. These results suggest that the 5-HT system does not encode a unitary signal and that it will broadcast specific information to the forebrain with speed and precision sufficient not only to modulate but also to dynamically sculpt ongoing information processing.

Excitatory and Inhibitory Synapses in Neuropeptide Y-Expressing Striatal Interneurons

Wed, 04 Nov 2009 11:06:52 PST

Although rare, interneurons are pivotal in governing striatal output by extensive axonal arborizations synapsing on medium spiny neurons. Using a genetically modified mouse strain in which a green fluorescent protein (GFP) is driven to be expressed under control of the neuropeptide Y (NPY) promoter, we identified NPY interneurons and compared them with striatal principal neurons. We found that the bacteria artificial chromosome (BAC)-npy mouse expresses GFP with high fidelity in the striatum to the endogenous expression of NPY. Patch-clamp analysis from NPY neurons showed a heterogeneous population of striatal interneurons. In the majority of cells, we observed spontaneous firing of action potentials in extracellular recordings. On membrane rupture, most NPY interneurons could be classified as low-threshold spiking interneurons and had high-input resistance. Voltage-clamp recordings showed that both GABA and glutamate gated ion channels mediate synaptic inputs onto these striatal interneurons. AMPA receptor–mediated spontaneous excitatory postsynaptic currents (sEPSCs) were small in amplitude and infrequent in NPY neurons. Evoked EPSCs did not show short-term plasticity but some rectification. Evoked N-methyl-d-aspartate (NMDA) EPSCs had fast decay kinetics and were poorly sensitive to an NR2B subunit containing NMDA receptor blocker. Spontaneous inhibitory postsynaptic currents (sIPSCs) were mediated by GABA_A receptors and were quite similar among all striatal neurons studied. On the contrary, evoked IPSCs decayed faster in NPY neurons than in other striatal neurons. These data report for the first time specific properties of synaptic transmission to NPY striatal interneurons.

EEG Generator--A Model of Potentials in a Volume Conductor

Avitan, L., Teicher, M., Abeles, M. — Wed, 04 Nov 2009 11:06:52 PST

EEG generator—a model of potentials in a volume conductor. The potential recorded over the cortex electro-corticogram (ECoG) or over the scalp [electroencephalograph (EEG)] derives from the activity of many sources known as "EEG generators." The recorded amplitude is basically a function of the unitary potential of a generator and the statistical relationship between different EEG generators in the recorded population. In this study, we first suggest a new definition of the EEG generator. We use the theory of potentials in a volume conductor and model the contribution of a single synapse activated to the surface potential. We then model the contribution of the generator to the surface potential. Once the generator and its contribution are well defined, we can quantitatively assess the degree of synchronization among generators. The measures obtained by the model for a real life scenario of a group of generators organized in a specific statistical way were consistent with the expected values that were reported experimentally. The study sheds new light on macroscopic modeling approaches which make use of mean soma membrane potential. We showed major contribution of activity of superficial apical synapses to the ECoG signal recorded relative to lower somatic or basal synapses activity.

Characterizing Learning by Simultaneous Analysis of Continuous and Binary Measures of Performance

Wed, 04 Nov 2009 11:06:52 PST

Continuous observations, such as reaction and run times, and binary observations, such as correct/incorrect responses, are recorded routinely in behavioral learning experiments. Although both types of performance measures are often recorded simultaneously, the two have not been used in combination to evaluate learning. We present a state-space model of learning in which the observation process has simultaneously recorded continuous and binary measures of performance. We use these performance measures simultaneously to estimate the model parameters and the unobserved cognitive state process by maximum likelihood using an approximate expectation maximization (EM) algorithm. We introduce the concept of a reaction-time curve and reformulate our previous definitions of the learning curve, the ideal observer curve, the learning trial and between-trial comparisons of performance in terms of the new model. We illustrate the properties of the new model in an analysis of a simulated learning experiment. In the simulated data analysis, simultaneous use of the two measures of performance provided more credible and accurate estimates of the learning than either measure analyzed separately. We also analyze two actual learning experiments in which the performance of rats and of monkeys was tracked across trials by simultaneously recorded reaction and run times and the correct and incorrect responses. In the analysis of the actual experiments, our algorithm gave a straightforward, efficient way to characterize learning by combining continuous and binary measures of performance. This analysis paradigm has implications for characterizing learning and for the more general problem of combining different data types to characterize the properties of a neural system.

Are There Nociceptive-Specific Brain Potentials?

Baumgartner, U., Treede, R.-D. — Wed, 04 Nov 2009 11:06:52 PST

Are There Nociceptive-Specific Brain Potentials? Reply to Baumgartner and Treede

Mouraux, A., Plaghki, L., Iannetti, G. D. — Wed, 04 Nov 2009 11:06:52 PST

Corrigendum

Wed, 04 Nov 2009 11:06:52 PST