Planning of reach-and-grasp movements: effects of validity and type of object information.

Individuals are assumed to plan reach-and-grasp movements by using two separate processes. In 1 of the processes, extrinsic (direction, distance) object information is used in planning the movement of the arm that transports the hand to the target location (transport planning); whereas in the other, intrinsic (shape) object information is used in planning the preshaping of the hand and the grasping of the target object (manipulation planning). In 2 experiments, the authors used primes to provide information to participants (N = 5, Experiment 1; N = 6, Experiment 2) about extrinsic and intrinsic object properties. The validity of the prime information was systematically varied. The primes were succeeded by a cue, which always correctly identified the location and shape of the target object. Reaction times were recorded. Four models of transport and manipulation planning were tested. The only model that was consistent with the data was 1 in which arm transport and object manipulation planning were postulated to be independent processes that operate partially in parallel. The authors suggest that the processes involved in motor planning before execution are primarily concerned with the geometric aspects of the upcoming movement but not with the temporal details of its execution.

Where's the ball? Two- and three-year-olds reason about unseen events.

Children 2, 2 1/2, and 3 years of age engaged in a search task in which they opened 1 of 4 doors in an occluder to retrieve a ball that had been rolled behind the occluder. The correct door was determined by a partially visible wall placed behind the occluder that stopped the motion of the unseen ball. Only the oldest group of children was able to reliably choose the correct door. All children were able to retrieve a toy that had been hidden in the same apparatus if the toy was hidden from the front by opening a door. Analysis of the younger children's errors indicated that they did not search randomly but instead used a variety of strategies. The results are consistent with the Piagetian view that the ability to use representations to guide action develops slowly over the first years of life.

Proximodistal structure of early reaching in human infants.

Nine infants were tested, at the age of onset of reaching, seated on their parent's lap and reaching for a small plastic toy. Kinematic analysis revealed that infants largely used shoulder and torso rotation to move their hands to the toy. Many changes in hand direction were observed during reaching, with later direction changes correcting for earlier directional errors. Approximately half of the infants started many reaches by bringing their hands backward or upward to a starting location that was similar across reaches. Individual infants often achieved highly similar peak speeds across their reaches. These results support the hypothesis that infants reduce the complexity of movement by using a limited number of degrees-of-freedom, which could simplify and accelerate the learning process. The proximodistal direction of maturation of the neural and muscular systems appears to restrict arm and hand movement in a way that simplifies learning to reach.

Midreach correction in 7-month-olds.

In recent work, it has been suggested that infant reaching is composed of a sequence of perception-action cycles in which the positions of the hand and the target are assessed and actions are generated that bring the hand to the target. In the present work, the author examined the ability of 7-month-old infants (N = 12) to correct reaches in midflight when a target location was shifted. The results showed that on the majority of shift trials, the infants corrected hand direction in midreach and that the latency of correction was 200-400 ms. Although the present results are limited to the case of the infant's response to a target shift, they are consistent with the hypothesis that infants monitor the positions of the hand and the target during the reach and are able to adjust for any errors of movement.

Visual Information and Object Size in the Control of Reaching.

The role of vision in the control of reaching and grasping was investigated by varying the available visual information. Adults (N = 7) reached in conditions that had full visual information, visual information about the target object but not the hand or surrounding environment, and no visual information. Four different object diameters were used. The results indicated that as visual information and object size decreased, subjects used longer movement times, had slower speeds, and more asymmetrical hand-speed profiles. Subjects matched grasp aperture to object diameter, but overcompensated with larger grasp apertures when visual information was reduced. Subjects also qualitatively differed in reach kinematics when challenged with reduced visual information or smaller object size. These results emphasize the importance of vision of the target in reaching and show that subjects do not simply scale a command template with task difficulty.

Multimodal perception in the control of infant reaching.

Six-month-old infants were presented with sounding objects under 3 conditions of illumination: in full vision, in the dark with target location specified by a glowing and sounding object, and in the dark with location specified by sound alone. Reaching behavior was videotaped with an infrared camera, and hand movement was measured by infrared-emitting diodes on the hand that were tracked by a motion analysis system. No differences were found in reaching behavior for objects in the light and glowing objects in the dark. Reaches for sounding objects in the dark had higher speeds, shorter durations, and more errors compared to the other 2 conditions. These findings indicate that vision of the hand did not appear to affect infants' reaching in this situation, whereas vision of the target did.

Distributed representation of limb motor programs in arrays of adjustable pattern generators.

Abstract This paper describes the current state of our exploration of how motor program concepts may be related to neural mechanisms. We have proposed a model of sensorimotor networks with architectures inspired by the anatomy and physiology of the cerebellum and its interconnections with the red nucleus and the motor cortex. We proposed the concept of rubrocerebellar and corticocerebellar information processing modules that function as adjustable pattern generators (APGs) capable of the storage, recall, and execution of motor programs. The APG array model described in this paper extends the single APG model of Houk et al. (1990) to an array of APGs whose collective activity controls movement of a simple two degree-of-freedom simulated limb. Our objective was to examine the APG array theory in a simple computational framework with a plausible relationship to anatomy and physiology. Results of simulation experiments show that the APG array model is capable of learning how to control movement of the simulated limb by adjusting distributed motor programs. Although the model is based on many simplifying assumptions, and the simulated motor control task is much simpler than an actual reaching task, these results suggest that the APG array model may provide a useful step toward a more comprehensive understanding of how neural mechanisms may generate motor programs.

Muscle activity during unconditioned and conditioned eye blinks in the rabbit.

EMGs were recorded from the orbicularis oculi, retractor bulbi and superior rectus muscles in rabbits to investigate the time course of muscle activation during unconditioned and conditioned eye blinks. EMGs from the three muscles showed two responses, with the responses of the orbicularis oculi and retractor bulbi showing the same latency, and the responses of the superior rectus lagging. The latency of responses to periorbital electrostimulation was about 5 ms, and to air puff stimulation about 10 ms. Results showed a tight coupling of activity between muscles, with cross-correlograms peaking at 0.65 to 0.85 and showing little time shift. Stimulus-response curves showed clear non-linearities in the response of the muscle to changes in stimulus strength. Local anesthesia of the cornea had little effect on unconditionally evoked responses. The form of unconditionally evoked responses was similar with periorbital electrostimulation and air puff stimuli but differed in latency. These results show the form of the eye blink reflex response and will be of importance in interpreting electrophysiological studies of the classically conditioned eye blink of rabbits.

Linear systems analysis of the relationship between firing of deep cerebellar neurons and the classically conditioned nictitating membrane response in rabbits.

The correlation of the activity of neurons in the interposed and dentate nuclei of the cerebellum with conditioned movements of the nictitating membrane was investigated using linear systems analysis. The activity of single deep cerebellar nuclear cells was assumed to be the input to a linear system that produced nictitating membrane movement. Data were initially analyzed with a causal model to assess the degree to which past neural activity predicted the conditioned response. 55 of 165 cells had correlation coefficients of 0.50 or greater between the model's moment-to-moment output and the actual output, with two interpositus cells having correlation coefficients of greater than 0.90. Double-sided impulse responses indicated that afference from the face and efference copy probably affect deep cerebellar neural activity. Nonlinearities were also found in the relationship between neuronal activity and conditioned movement. It was concluded that cerebellar deep nuclear firing is highly correlated with future nictitating membrane movements but that the firing-movement relationship contains noncausal and nonlinear components.

Activity of deep cerebellar nuclear cells during classical conditioning of nictitating membrane extension in rabbits.

The activity of neurons in the interposed and dentate nuclei of the cerebellum was investigated during differential classical conditioning of the rabbit eye blink nictitating membrane response. Forty-seven percent of the 165 cells in the study responded to the orbital stimulation used as the unconditioned stimulus (US). The latency distribution of US-elicited responses was bimodal with peaks at 7 and 19 ms. Twenty-one percent of the cells responded with short latencies to the tones used as conditioned stimuli (CSs). These cells typically responded to both the reinforced and nonreinforced CSs. Forty-one percent of the cells responded on conditioned response (CR) trials but not on trials without CRs. The average lead of the neural response to the CR was 71.4 ms. Cells that responded on CR trials were more likely to respond to the CSs, or to the CSs and the US, than cells that did not respond on CR trials. For about half of the cells that responded on CR trials the latency of response followed trial-by-trial variations of CR latency. For the remainder, the response was time-locked to CS-onset. Cells whose responses paralleled the CR may be involved in the initiation or modulation of the CR, while those whose responses were time-locked to the CS may be involved in sensory processing underlying the initiation of the movement. The pathways that may underlie the US- and CS-elicited responses are also discussed.

Adaptively timed conditioned responses and the cerebellum: a neural network approach.

Conditioned responses often reflect knowledge about the timing of a US. This knowledge is manifested in the dependence of response topography on the CS-US interval employed in training. A neural network model and set of learning rules capable of simulating temporally adaptive features of conditioned responses is reviewed, and simulations are presented. In addition, we present a neural network implementation of the model which is designed to reconcile empirical studies of long-term synaptic depression in the cerebellum with neurobiological evidence from studies of the classically conditioned nictitating membrane response of the rabbit.

Simulation of the classically conditioned nictitating membrane response by a neuron-like adaptive element: response topography, neuronal firing, and interstimulus intervals.

A neuron-like adaptive element with computational features suitable for classical conditioning, the Sutton-Barto (S-B) model, was extended to simulate real-time aspects of the conditioned nictitating membrane (NM) response. The aspects of concern were response topography, CR-related neuronal firing, and interstimulus interval (ISI) effects for forward-delay and trace conditioning paradigms. The topography of the NM CR has the following features: response latency after CS onset decreases over trials; response amplitude increases gradually within the ISI and attains its maximum coincidentally with the UR. A similar pattern characterizes the firing of some (but not all) neurons in brain regions demonstrated experimentally to be important for NM conditioning. The variant of the S-B model described in this paper consists of a set of parameters and implementation rules based on 10-ms computational time steps. It differs from the original S-B model in a number of ways. The main difference is the assumption that CS inputs to the adaptive element are not instantaneous but are instead shaped by unspecified coding processes so as to produce outputs that conform with the real-time properties of NM conditioning. The model successfully simulates the aforementioned features of NM response topography. It is also capable of simulating appropriate ISI functions, i.e. with maximum conditioning strength with ISIs of 250 ms, for forward-delay and trace paradigms. The original model's successful treatment of multiple-CS phenomena, such as blocking, conditioned inhibition, and higher-order conditioning, are retained by the present model.

Cerebellar Purkinje cell activity related to the classically conditioned nictitating membrane response.

Because of the purported critical role of cerebellar lobule HVI in classical conditioning of the nictitating membrane response of the rabbit, we recorded extracellularly from HVI Purkinje cells (PCs) during differential conditioning. Rabbits were trained using tonal conditioned stimuli (CSs) and stimulation of the periocular region as the unconditioned stimulus (US). Many PCs responded to the US, the most frequently observed response being a burst of simple spikes. PCs in HVI showed a variety of responses to CSs that were related to conditioned responses (CRs). The most frequently observed response was an increase in simple spikes correlated with CRs. The activity of many of these cells antedated CRs by 20-200 ms. A smaller proportion of cells exhibited inhibition of simple spike activity that antedated CRs. The existence of PCs that alter their firing before CRs suggests that they may be causally involved in this behavior, and in this respect they reinforce reports that lesions of HVI or its connections disrupt nictitating membrane CRs. Although complex spike activity was not generally related to the US or to CRs, a few PCs responded in relation to CRs with only complex spikes. In demonstrating CR-related activity in cerebellar PCs, this study supports theories of cerebellar learning such as those of Marr and Albus.

The role of the extraocular muscles in the rabbit nictitating membrane response: a re-examination.

Berthier and Moore showed that the rabbit nictitating membrane (NM) response principally results from contracting the retractor bulbi muscle which pulls the globe into the socket thereby passively effecting NM extension. They concluded that the remaining extraocular muscles can effect NM extension if the retractor bulbi is denervated. A re-examination of the role of the recti and oblique extraocular muscles in nictitating membrane extension was undertaken in the light of recent results of Marek et al., suggesting that the facial nerve, and not the extraocular muscles, participates in extension of the NM. In contrast to Marek et al., the present results indicated that section of the extraocular muscles was necessary to abolish eyeshock or tactilly elicited NM extension when the abducens and facial nerves were severed. It is therefore likely that extraocular (recti and oblique) muscles participate in globe retraction and NM extension, as originally noted by Lorente de No.

Effects of hypothalamic stimulation on unit responses recorded from neurons of sensorimotor cortex of awake cats during conditioning.

1. The activity of single units of the coronal pericruciate (CPC) cortex was studied in 11 awake cats during sessions in which a click conditioned stimulus (CS) was repeatedly paired with glabella tap unconditioned stimulus (US) and hypothalamic stimulation (HS). Effects of HS on the activity of cortical units were also studied during sessions in which HS alone was delivered repeatedly every 10 s. 2. HS evoked an increase in spike activity of less than 60 ms latency in 89 of 116 units tested. 3. Repeated presentation of HS that was effective in producing rapid behavioral conditioning resulted in a characteristic reduction in the latency of discharge evoked by HS in cortical units. 4. Short-latency activation (less than 20 ms) of units of the sensorimotor cortex appeared to be characteristic of HS that led to enhanced rates of conditioned response (CR) acquisition. One of the cells responding in this way was identified as a pyramidal cell of layer V by intracellular injection of horseradish peroxidase (HRP). 5. Further analyses of activity were performed on 16 units of the CPC cortex that were followed through conditioning (or reconditioning) and extinction of the CR. After less than 10 CS-US-HS pairings, there was a selective augmentation of unit response to the CS but not of response to an explicitly unpaired discriminative stimulus (DS). Responses to the CS were not similarly augmented when presentations of HS preceded rather than followed the presentations of the CS and US. The rapid development of CS-evoked unit activity coincided with the rapid acquisition of discriminative CRs behaviorally. 6. During conditioning, the most conspicuous increases in CS-evoked unit response occurred at latencies 100 ms or more after onset of the click CS. This corresponded with the behavioral observation that the majority of eye blink CRs occurred with onset latencies longer than 100 ms.

The nictitating membrane response: an electrophysiological study of the abducens nerve and nucleus and the accessory abducens nucleus in rabbit.

The nictitating membrane response to periocular electrostimulation was investigated in anesthetized and paralyzed rabbits. Recordings from the abducens nerve, which carries the fibers innervating retractor bulbi muscles that are primarily involved in this reflex, showed two distinct volleys to effective stimulation: a short duration volley with a minimum latency of approximately 4 ms and a longer duration volley beginning approximately 10 ms after stimulus onset and lasting up to 25 ms. Recordings of antidromically evoked field potentials via microelectrodes indicated large responses to abducens nerve stimulation in the vicinity of the accessory abducens nucleus. Single unit recordings from the accessory abducens nucleus produced spike trains with minimal latencies of 3.7-5 ms to eyeshock. The latency of spike discharge was inversely related to stimulus current. Units in the abducens nucleus did not show stimulus-elicited spiking, suggesting that the accessory abducens, but not the abducens, is primarily involved in the reflex pathway. Transverse knife cuts which separated caudal areas of the sensory trigeminal complex from the accessory abducens nucleus did not attenuate the efferent volley to suprathreshold stimulation, suggesting that more rostral components of the trigeminal complex are primarily involved in the reflex pathway.