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Dynamics of single neuron activity in monkey primary motor cortex related to sensorimotor transformation.

We investigated the dynamics of neuronal activity related to sensorimotor transformation during single experimental trials of a given stimulus-response (S-R) association task. A monkey was trained to perform wrist extension/flexion movements in the horizontal plane to align a pointer with a visual target while single unit activity in the primary motor cortex (MI) was being recorded. The stimulus was a colored light-emitting diode (LED) presented to either the left or right of a central reference point. The monkey had to point directly at the target ("compatible" S-R mapping) or point to the opposite side of the target position ("incompatible" S-R mapping), with the mapping rule specified by the color of the LED. Single neuron activities on the four correct trials (left/right stimulus x compatible/incompatible S-R mapping) were compared to determine whether such activities were more related to stimulus encoding and representation, to response preparation and execution, or to the "decision" processes translating the stimulus representation into a response representation. A novel mathematical technique, called LOCUS ANALYSIS, has been developed to quantitatively analyze and visualize the contribution of neuronal activity toward the sensory, motor, or sensorimotor (i.e., decisional) aspects of the task. Our data show that as a trial evolves, neuronal activity in MI, at a population level, is first correlated with the representation of the specific stimulus (the side of LED), then with the representation of the S-R mapping rule (the color of LED) as well as trial-specific S-R association (the conjunction of stimulus side and stimulus color), and finally with the representation of the behavioral response (extension or flexion wrist movement). Immediately after the issuance of the movement command, the populational activity in MI remains correlated with the trial-specific stimulus-response conjunctions, i.e., the context of the motor decision that the monkey has just made. Cells recorded successively in a single penetration tend to resemble each other in their pattern of firing on the four correct trials, suggesting a modular organization of neurons based on their functional role in the processing of the S-R association task. Our results indicate that MI belongs to a distributed network such that its neuronal activity reflects the underlying network dynamics that translate a stimulus representation into a response representation via the activation and application of appropriate S-R mapping rule.

Animals↗

Spatial orientation and balance control changes induced by altered gravitoinertial force vectors.

To better understand the mechanisms of human adaptation to rotating environments, we exposed 19 healthy subjects and 8 vestibular-deficient subjects ("abnormal"; four bilateral and four unilateral lesions) to an interaural centripetal acceleration of 1 g (resultant 45 degrees roll-tilt of 1.4 g) on a 0.8-m-radius centrifuge for periods of 90 min. The subjects sat upright (body z-axis parallel to centrifuge rotation axis) in the dark with head stationary, except during 4 min of every 10 min, when they performed head saccades toward visual targets switched on at 3- to 5-s intervals at random locations (within +/- 30 degrees) in the earth-horizontal plane. Eight of the normal subjects also performed the head saccade protocol in a stationary chair adjusted to a static roll-tilt angle of 45 degrees for 90 min (reproducing the change in orientation but not the magnitude of the gravitoinertial force on the centrifuge). Eye movements, including voluntary saccades directed along perceived earth- and head-referenced planes, were recorded before, during, and immediately after centrifugation. Postural center of pressure (COP) and multisegment body kinematics were also gathered before and within 10 min after centrifugation. Normal subjects overestimated roll-tilt during centrifugation and revealed errors in perception of head-vertical provided by directed saccades. Errors in this perceptual response tended to increase with time and became significant after approximately 30 min. Motion-sickness symptoms caused approximately 25% of normal subjects to limit their head movements during centrifugation and led three normal subjects to stop the test early. Immediately after centrifugation, subjects reported feeling tilted 10 degrees in the opposite direction, which was in agreement with the direction of their earth-referenced directed saccades. Postural COP, segmental body motion amplitude, and hip-sway frequency increased significantly after centrifugation. These postural effects were short-lived, however, with a recovery time of several postural test trials (minutes). There were also asymmetries in the direction of postcentrifugation COP and head tilt which depended on the subject's orientation during the centrifugation adaptation period (left ear or right ear out). The amount of total head movements during centrifugation correlated poorly or inversely with postcentrifugation postural stability, and the most unstable subject made no head movements. There was no decrease in postural stability after static tilt, although these subjects also reported a perceived tilt briefly after return to upright, and they also had COP asymmetries. Abnormal subjects underestimated roll-tilt during centrifugation, and their directed saccades revealed permanent spatial distortions. Bilateral abnormal subjects started out with poor postural control, but showed no postural decrements after centrifugation, while unilateral abnormal subjects had varying degrees of postural decrement, both in their everyday function and as a result of experiencing the centrifugation. In addition, three unilateral, abnormal subjects, who rode twice in opposite orientations, revealed a consistent orthogonal pattern of COP offsets after centrifugation. These results suggest that both orientation and magnitude of the gravitoinertial vector are used by the central nervous system for calibration of multiple orientation systems. A change in the background gravitoinertial force (otolith input) can rapidly initiate postural and perceptual adaptation in several sensorimotor systems, independent of a structured visual surround.

Acceleration↗

Eye movements of monkey observers viewing vocalizing conspecifics.

Primates, including humans, communicate using facial expressions, vocalizations and often a combination of the two modalities. For humans, such bimodal integration is best exemplified by speech-reading - humans readily use facial cues to enhance speech comprehension, particularly in noisy environments. Studies of the eye movement patterns of human speech-readers have revealed, unexpectedly, that they predominantly fixate on the eye region of the face as opposed to the mouth. Here, we tested the evolutionary basis for such a behavioral strategy by examining the eye movements of rhesus monkeys observers as they viewed vocalizing conspecifics. Under a variety of listening conditions, we found that rhesus monkeys predominantly focused on the eye region versus the mouth and that fixations on the mouth were tightly correlated with the onset of mouth movements. These eye movement patterns of rhesus monkeys are strikingly similar to those reported for humans observing the visual components of speech. The data therefore suggest that the sensorimotor strategies underlying bimodal speech perception may have a homologous counterpart in a closely related primate ancestor.

Animals↗

Altered inhibition of motor responses in Tourette Syndrome and Obsessive-Compulsive Disorder.

OBJECTIVES: The Gilles de la Tourette Syndrome (TS) and Obsessive Compulsive Disorder (OCD) have been shown to display impaired cognitive and motor inhibition. This study investigated inhibitory mechanisms of motor responses in order to expand the understanding of sensorimotor integration processes in both disorders. We hypothesized that both patient groups would display altered frontal inhibitory activity. MATERIAL AND METHODS: To this end event-related brain potentials (ERPs) were recorded in a STOP-paradigm in groups of TS and OCD patients and in a control group. The paradigm required the execution of a motor response after a "go" signal was given and the occasional suppression of this response after a second "stop" signal occurred. RESULTS: Behavioral parameters and Lateralized Readiness Potential (LRP) confirmed that both patient groups were well able to initiate motor responses. "Go" and "stop" stimuli elicited an enhanced frontal negative activity in both patient groups. In addition, "stop" stimuli were associated with a frontal shift of the NoGo-Anteriorization (NGA) in the TS group but not in the OCD group. CONCLUSIONS: The data are interpreted to indicate altered frontal inhibitory functions. Similarities and dissimilarities between the findings for TS and OCD are discussed with respect to other pathophysiologic aspects of the disorders.

Adolescent↗

Facilitated beam-walking recovery during acute phase by kynurenic acid treatment in a rat model of photochemically induced thrombosis causing focal cerebral ischemia.

We previously demonstrated the presence of activated areas in the non-injured contralateral sensorimotor cortex in addition to the ipsilateral sensorimotor cortex of the area surrounding a brain infarction, using a rat model of focal photochemically induced thrombosis (PIT) and functional magnetic resonance imaging. Using this model, we next applied gene expression profiling to screen key molecules upregulated in the activated area. RNA was extracted from the ipsilateral and contralateral sensorimotor cortex to the focal brain infarction and from the sham controlled cortex, and hybridized to gene-expression profiling arrays containing 1,322 neurology-related genes. Results showed that glycine receptors were upregulated in both the ipsilateral and contralateral cortex to the focal ischemic lesion. To prove the preclinical significance of upregulated glycine receptors, kynurenic acid, an endogenous antagonist to glycine receptors on neuronal cells, was administered intrathecally. As a result, the kynurenic acid significantly improved behavioral recovery within 10 days from paralysis induced by the focal PIT (p < 0.0001), as evaluated with beam walking. These results suggest that intrathecal administration of a glycine receptor antagonist may facilitate behavioral recovery during the acute phase after brain infarction.

Animals↗

The role of nitric oxide in the pathophysiology of thromboembolic stroke in the rat.

Although nitric oxide (NO) has been shown to play an important role in the pathophysiology of cerebral ischemia, its contribution to the pathogenesis of experimentally induced thromboembolic stroke is unknown. In this study, we pharmacologically manipulated NO levels in the acute post-thrombotic stage and determined the effects on behavior and histopathology. The following drugs were used: nitro-L-arginine-methyl ester (L-NAME), a non-specific endothelial and neuronal nitric oxide synthase (eNOS and nNOS) inhibitor, 3-bromo-7-nitroindazole (7-NI), a specific inhibitor for nNOS, the NO precursor, exogenous L-arginine and the NO-donor, 3-morpholino-sydnonimine (SIN-1). Male Wistar rats (n = 76) were randomly assigned to receive vehicle or drug immediately after common carotid artery thrombosis (CCAT). Regional measurements of cortical NOS activity using the [3H]L-arginine to [3H]L-citrulline conversion assay were decreased 1 h after treatment with L-NAME and 7-NI by 50 and 65%, respectively; hippocampal NOS activity was reduced with L-NAME by 35% and with 7-NI by 65%. L-NAME significantly worsened forelimb placing as compared to other groups. 7-NI accelerated sensorimotor recovery. Water maze retention deficits were noted 48 h after CCAT and these were exacerbated by L-NAME treatment. Histopathological protection was conferred in the hippocampus by 7-NI and SIN-1; conversely, L-NAME increased neuronal injury in the contralateral cortex. L-arginine had no effect on these outcomes. In conclusion, both structural and functional consequences of CCAT can be aggravated by limiting endothelial NO production in the acutely post-thrombotic brain. In contrast, inhibition of nNOS and infusion of an NO donor has a beneficial effect on pathology.

Animals↗

Short-latency excitatory postsynaptic potentials are evoked in primate spinothalamic tract neurons by corticospinal tract volleys.

The responses of 8 wide-dynamic-range (WDR) STT neurons to electrical stimulation of the sensorimotor cortex or the cerebral peduncle were recorded intracellularly in anesthetized monkeys. EPSPs and spikes were evoked in STT neurons with short latencies ranging from 0.4 to 5.3 msec after the corticospinal volley, indicating a monosynaptic connection in at least some STT neurons. Spatial and temporal summation seem necessary for the cortex to maintain an effective excitatory influence on STT neurons. It is postulated that the cortical facilitation of STT neurons helps them to acquire more nociceptive signals during behavioral tasks such as exploring a new object so that the brain can make decisions about whether to continue or to abandon a certain task. Possible implications for central pain states are discussed.

Animals↗

Forelimb use after focal cerebral ischemia in rats treated with an alpha 2-adrenoceptor antagonist.

Atipamezole, a selective alpha(2)-adrenoceptor antagonist, enhances recovery of sensorimotor function after focal cerebral ischemia in rats. The aim of the present study was to further characterize the effects of atipamezole treatment combined with enriched-environment housing in ischemic rats by evaluating spontaneous exploratory activity in the cylinder test. The right middle cerebral artery (MCA) of rats was occluded for 120 min using the intraluminal filament method. Atipamezole (1.0 mg/kg) or 0.9% NaCl was administered on postoperative days 2 through 11 and 15, 19, and 23. Spontaneous behavior of rats in a transparent cylinder was videotaped before, and 6 and 23 days after surgery 20 min after drug administration. Constant asymmetry in forelimb use was observed in the cylinder test on postoperative days 6 and 23. Ischemic rats used the impaired forelimbs (contralateral to lesion) during lateral exploration less than did sham-operated rats (P<.001). Ischemic rats also preferred to turn contralateral to the lesion (P<.05). Atipamezole increased the simultaneous, but not independent, use of the forelimbs during lateral exploration (P<.05). The data suggest that noradrenergic manipulation does not significantly enhance recovery in a test that does not depend on practice following focal cerebral ischemia.

Adrenergic alpha-2 Receptor Antagonists↗

Lack of task specificity and absence of posttraining effects of atropine on learning.

Three experiments are reported whose purpose was to examine the effect of the cholinergic antagonist atropine on the acquisition of different learning tasks known to be sensitive or insensitive to impairment by hippocampal lesions; on the retention of performance acquired in the absence of the drug; and on memory consolidation immediately after daily training trials. In Experiment 1, atropine sulfate (10 or 50 mg/kg, ip), injected 30 min prior to training, severely impaired learning of both spatial and nonspatial discrimination tasks when compared with saline or atropine methylnitrate (50 mg/kg). In Experiment 2, atropine sulfate (50 mg/kg) also impaired spatial discrimination accuracy in rats previously trained to asymptote under drug-free conditions. These deficits were not due to either peripheral drug effects or gross sensorimotor impairments. In Experiment 3, daily posttraining injections of atropine sulfate (50 mg/kg) failed to influence either learning or subsequent retention of place navigation in rats that were trained to find a single hidden escape platform. The data confirm that profound learning deficits occur when training is conducted under atropine but offer no support to the hypothesis that cholinergic neurons play an important role in memory consolidation or other posttraining processes. Furthermore, these results point to dissimilarities between the behavioral impairments induced by cholinergic blockade and hippocampal lesions under appropriate test regimes.

Animals↗

Combined exposure to DEET (N,N-diethyl-m-toluamide) and permethrin: pharmacokinetics and toxicological effects.

Permethrin and DEET are concurrently used for pests control inside homes, in public places, and in military shelters. Combined exposure to these compounds produced greater biochemical, behavioral, and metabolic alterations in animals compared to each individual compound. Concurrent application of DEET and permethrin induced urinary excretion of 3-nitrotyrosine and 8-hydroxy-2'-deoxyguanosine, markers of DNA damage and oxidative stress in rats, increased the release of rat brain mitochondrial cytochrome c, disrupted the blood-brain barrier (BBB) in rats, decreased m2 muscarinic acetylcholine receptor ligand binding density in rat brain, increased urinary excretion of 6 beta-hydroxycortisol, a marker CYP3A4 induction, altered sensorimotor and locomotor activities in rats, and changed in vivo and in vitro metabolism and pharmacokinetic profiles of the individual compound. These findings show that more research is needed to examine adverse effects of the combined use of DEET and permethrin on other biochemical/physiological system(s) and to predict mechanistic pathways for these effects, particularly mechanism of action at cellular and molecular levels and alterations of genes transcription.

Animals↗

Contextual modulation of central thalamic delay-period activity: representation of visual and saccadic goals.

This study examines the influence of behavioral context on the activity of visuomotor neurons in primate central thalamus. Neurons that combine information about sensory stimuli and their behavioral relevance are thought to contribute to the decision mechanisms that link specific stimuli to specific responses. We reported in a previous study that neurons in central thalamus carry spatial information throughout the instructed delay period of a visually guided delayed saccade task. The goal of the current study was to determine whether the delay-period activity of thalamic neurons is modulated by behavioral context. Single neurons were evaluated during performance of visually guided and memory-guided variants of a saccadic choice task in which a cue designated the response field stimulus as the target of a rewarded saccade or as an irrelevant distracter. The relative influence of the physical stimulus and context on delay-period activity suggested a minimum of 3 neural groups. Some neurons signaled the locations of visible stimuli regardless of behavioral relevance. Other neurons preferentially signaled the locations of current saccadic goals and did so even in the absence of the physical stimulus. A third group signaled only the locations of currently visible saccadic goals. For the latter 2 groups, activity was the product of both stimulus and context, suggesting that central thalamic neurons play a role in the context-dependent linkage of sensory signals and saccadic commands. More generally, these data suggest that the anatomical substrate of sensorimotor decision making may include the cortico-subcortical loops for which central thalamus serves as the penultimate synapse.

Action Potentials↗

The "staircase test": a measure of independent forelimb reaching and grasping abilities in rats.

A novel reaching test for the rat has been developed to assess the independent use of forelimbs in skilled reaching and grasping tasks. The apparatus is a plexiglas box with a removable baited double staircase. Food pellets are placed on the staircase and presented bilaterally at 7 graded stages of reaching difficulty to provide objective measures of side bias, maximum forelimb extension and grasping skill. In the present experiment, the apparatus was used to assess the reaching performance of rats following unilateral lesions of the sensorimotor cortex, unilateral lesions of the posterior cortex or bilateral lesions of the olfactory bulbs. The task has the advantage of objective over rating measurement, and the simplicity of the apparatus permits many animals to be tested concurrently.

Animals↗

Unilateral AMPA lesions of nucleus basalis magnocellularis induce a sensorimotor deficit which is differentially altered by arecoline and nicotine.

One week after unilateral alpha-amino-3-hydroxy-5-methyl-4-isoxazole propionic acid (AMPA) lesions of nucleus basalis magnocellularis, rats showed significant lateralised bias in spontaneous turning and in turning induced by tail pinch or by placing the rat on a 45 degrees grid. Turning was biased to the lesioned side and this side also showed increased responsiveness to pin-prick stimulation of the skin (somaesthesia), snout and whisker stimulation and ammonia olfaction. Arecoline (0.5 mg/kg), at a dose which did not affect responses to sensorimotor stimulation in sham-operated rats, corrected the lesion-induced biased turning to tail pinch and the 45 degrees grid test and reduced the bias in the open field. In contrast, nicotine (0.05 mg/kg), at a dose which also did not substantially affect responses to sensorimotor stimulation in sham-operated rats, switched the lesion-induced turning bias towards the contralateral side. Neither cholinoceptor agonist reduced the lesion-induced increased sensory responsiveness. The effects of nicotine were blocked by the centrally acting nicotinic antagonist, mecamylamine (1.0 mg/kg), but not by hexamethonium (1.0 mg/kg), or ondansetron (0.01 mg/kg). Amphetamine (up to 1.0 mg/kg) did not affect the lesion-induced motor asymmetry. The results confirm that the basal forebrain cholinergic system plays a role in sensorimotor cortical functions, but suggest different functional roles for muscarinic and nicotinic receptors.

Afferent Pathways↗

Biofeedback-produced hemispheric asymmetry of slow cortical potentials and its behavioural effects.

Two studies served to examine behavioural effects of slow cortical potentials (SPs). SPs were manipulated by means of a biofeedback procedure. The ability of human subjects to alter SPs differentially between the two hemispheres--specifically over the lateral aspects of the central sulcus--was tested by providing feedback of the SP difference between C3 and C4. In Expt. I, 21 of the 45 subjects produced hemispheric asymmetries of more than 2 microV between C3 and C4 on an average after 80 trials of analogue, continuous and immediate feedback. In Expt. II, SP changes were fed back digitally at the end of each trial. Within 120 trials, 20 of the 48 subjects reached the criterion of a minimum 2-microV difference in SPs between C3 and C4 on the average. Average differentiation remained significantly below the SP differentiations achieved for continuous feedback. Trials with feedback were followed by 'task' trials without feedback, during which subjects were still requested to produce SP changes but also had to complete a task: Either sensorimotor tasks (Expt. I) or forced choice handedness tasks (Expt. II) were presented to evaluate behavioural consequences of hemispheric SP differences. In subjects achieving the required SP differentiation it affected the behavioural output in agreement with the known functions of the respective cortical area.

Adult↗

Sensorimotor behaviour following incomplete cervical spinal cord injury in the rat.

Rats are one of the most commonly used species for spinal cord injury research. Since the advent of the Basso, Beattie, Bresnahan (BBB) locomotor rating scale, the majority of spinal cord injury research relies upon evaluating locomotor behaviour in thoracic spinal cord injury rat models. Slightly more than 50% of all traumatic spinal cord injuries in humans, however, occur at the level of the cervical spinal cord. Further, therapies aimed at thoracic spinal cord injuries may not be directly transferable to cervical spinal cord injuries. This could be due to (1) differences in distance between the cell bodies of injured axons and the injury site and (2) because some behaviours (e.g. stepping movements) used to evaluate the therapeutic potential of a given treatment are governed primarily by intraspinal neuronal circuitry while other behaviours (e.g. skilled reaching) require more sophisticated conscious integration of the sensorimotor system. Consequently, there is a need to develop and use experimental cervical spinal cord injury models and understand the behavioural characteristics of such models. The present review highlights the sensorimotor abilities of cervical spinal cord-injured rats, including both forelimb, hind limb, and whole body behaviours. We also provide insight into the neuroanatomic substrates important for performing a given behaviour, information which may prove essential in the development of site-directed therapeutic strategies.

Animals↗

Behavioural analysis of unilateral monoamine depletion in the marmoset.

Unilateral stereotaxic injections of 6-hydroxydopamine (6-OHDA) into the nigrostriatal bundle of marmosets (Callithrix jacchus) produced substantial losses of tyrosine hydroxylase immunoreactive neurons from the substantia nigra, and mean dopamine (DA) depletions of 98-99% in the caudate nucleus, putamen and nucleus accumbens, and of 91-97% in frontal cortex, on the side of the lesion. Noradrenaline (NA) and 5-hydroxytryptamine (5-HT) levels were also affected. Behavioural tests conducted pre-operatively and at regular intervals during the 6 mths following surgery revealed persistent deficits in the lesioned marmosets as a group compared with sham-lesioned controls, although individual marmosets sometimes recovered or showed no initial deficit on some tests. The main behavioural effects of the lesion were as follows: (i) an increase in the time spent with the head positioned ipsilaterally with respect to the rest of the body; (ii) ipsilateral spontaneous and amphetamine-induced rotation, although occasional intermittent periods of contralateral rotation and head biases were also recorded; (iii) contralateral apomorphine-induced rotation; (iv) reduced spontaneous activity; (v) ipsilateral hand preference on a conveyor belt task, although hand skill (measured as percentage errors when the speed of the belt was increased) was not affected; (vi) neglect of contralateral stimuli, both at the conveyor belt where lesioned monkeys often failed to respond on trials on which apple pieces arrived from the contralateral side, and on a test of sensorimotor neglect in which adhesive labels were placed around both feet. Comparisons of biochemical measures of the lesion with behavioural scores in individual monkeys suggest that DA depletions in excess of 95% are essential for long-term behavioural deficits.

Animals↗

[The effect of stimulation of the lateral hypothalamus on the spike trains of the rabbit neocortical neurons].

Cross- and autocorrelation histograms of the impulse activity of the visual and sensorimotor cortical neurons were plotted before and after stimulation of the right (RH) and left lateral hypothalamus (LH) which provoked food motivation reactions. Reorganization in correlated activity of neocortical neurons was more expressed after the LH than the RH stimulation. Only after the LH stimulation the number of neuronal pairs with co-related activity in the left hemisphere increased by 41% and some order of priority appeared in the neuronal discharges. The visual neurons discharged after the sensorimotor ones with delays up to 120 ms. The conclusion was drawn that cortical interhemispheric asymmetry of electrical activity during hunger was associated with unequal operation of the right and left lateral hypothalamus.

Animals↗

Visuomotor processing as reflected in the directional discharge of premotor and primary motor cortex neurons.

Premotor and primary motor cortical neuronal firing was studied in two monkeys during an instructed delay, pursuit tracking task. The task included a premovement "cue period," during which the target was presented at the periphery of the workspace and moved to the center of the workspace along one of eight directions at one of four constant speeds. The "track period" consisted of a visually guided, error-constrained arm movement during which the animal tracked the target as it moved from the central start box along a line to the opposite periphery of the workspace. Behaviorally, the animals tracked the required directions and speeds with highly constrained trajectories. The eye movements consisted of saccades to the target at the onset of the cue period, followed by smooth pursuit intermingled with saccades throughout the cue and track periods. Initially, an analysis of variance (ANOVA) was used to test for direction and period effects in the firing. Subsequently, a linear regression analysis was used to fit the average firing from the cue and track periods to a cosine model. Directional tuning as determined by a significant fit to the cosine model was a prominent feature of the discharge during both the cue and track periods. However, the directional tuning of the firing of a single cell was not always constant across the cue and track periods. Approximately one-half of the neurons had differences in their preferred directions (PDs) of >45 degrees between cue and track periods. The PD in the cue or track period was not dependent on the target speed. A second linear regression analysis based on calculation of the preferred direction in 20-ms bins (i.e., the PD trajectory) was used to examine on a finer time scale the temporal evolution of this change in directional tuning. The PD trajectories in the cue period were not straight but instead rotated over the workspace to align with the track period PD. Both clockwise and counterclockwise rotations occurred. The PD trajectories were relatively straight during most of the track period. The rotation and eventual convergence of the PD trajectories in the cue period to the preferred direction of the track period may reflect the transformation of visual information into motor commands. The widely dispersed PD trajectories in the cue period would allow targets to be detected over a wide spatial aperture. The convergence of the PD trajectories occurring at the cue-track transition may serve as a "Go" signal to move that was not explicitly supplied by the paradigm. Furthermore, the rotation and convergence of the PD trajectories may provide a mechanism for nonstandard mapping. Standard mapping refers to a sensorimotor transformation in which the stimulus is the object of the reach. Nonstandard mapping is the mapping of an arbitrary stimulus into an arbitrary movement. The shifts in the PD may allow relevant visual information from any direction to be transformed into an appropriate movement direction, providing a neural substrate for nonstandard stimulus-response mappings.

Animals↗