PubMed Health⌕ Search

Biomedical subjects

John Porrill

Publications and source records attributed to John Porrill.

7 recordsLinked to original sources

Evidence for wide range of time scales in oculomotor plant dynamics: implications for models of eye-movement control.

Oculomotor-plant dynamics are not well characterised, despite their importance for modelling eye-movement control. We analysed the time course of the globe's return after horizontal displacements in three rhesus monkeys lightly anaesthetised with ketamine. The eye-position traces were well fitted by a sum of four exponentials (time constants 0.012, 0.099, 0.46, 7.8 s). The two long time-constant terms accounted for 25% of plant compliance, and led to a model that accounted for hitherto unexplained features of ocular motoneuron firing such as (i) hysteresis, and (ii) the inability of a 2 time-constant model to fit data for both fast and slow eye-movements.

Adaptation, Physiological↗

Report on a workshop concerning the cerebellum and motor learning, held in St Louis October 2004.

A one-day meeting on the cerebellum and motor learning was held in St Louis (October 2004), to address issues arising from a previous larger meeting (Tuebingen, June 2004). The learning tasks considered were VOR adaptation, saccadic adaptation and eyeblink conditioning. A theoretical development was reported that indicated how the cerebellum could use sensory error signals for adaptive control, by decorrelating them from an efferent copy of motor commands. The main topics for discussion were the nature of the error signals actually used by the cerebellum, and the evidence for multiple sites of synaptic plasticity. Reports of studies on VOR adaptation confirmed the presence of error signals in addition to retinal slip, in particular the eye-movement related simple-spike firing of floccular PCs. This firing appears to drive synaptic plasticity in the vestibular nuclei. From a theoretical perspective, a second site of plasticity in the brainstem has two advantages: it improves the high-frequency performance of the VOR given a delayed slip signal, and it allows VOR adaptation when smooth pursuit effectively removes the retinal slip signal. In contrast, some of the physiological data reported on saccadic adaptation seemed incompatible with current theoretical ideas about error signals. However, since other reported data were broadly consistent with those ideas, an important area of experimental disagreement was identified. Furthermore, behavioural studies indicated the presence of multiple sites of plasticity, consistent with earlier lesion studies that suggested one such site within cerebellar cortex and another outside it. Data from eyeblink conditioning suggested that the predictability of the error signal was important. Related ideas have previously emerged from studies of skeletal movement, but their theoretical implications for the cerebellar algorithm have yet to be fully explored. Finally, the long-standing controversy concerning sites of plasticity in eyeblink conditioning illustrated the technical difficulties involved in tracking down such sites.

Adaptation, Physiological↗

Recurrent cerebellar architecture solves the motor-error problem.

Current views of cerebellar function have been heavily influenced by the models of Marr and Albus, who suggested that the climbing fibre input to the cerebellum acts as a teaching signal for motor learning. It is commonly assumed that this teaching signal must be motor error (the difference between actual and correct motor command), but this approach requires complex neural structures to estimate unobservable motor error from its observed sensory consequences. We have proposed elsewhere a recurrent decorrelation control architecture in which Marr-Albus models learn without requiring motor error. Here, we prove convergence for this architecture and demonstrate important advantages for the modular control of systems with multiple degrees of freedom. These results are illustrated by modelling adaptive plant compensation for the three-dimensional vestibular ocular reflex. This provides a functional role for recurrent cerebellar connectivity, which may be a generic anatomical feature of projections between regions of cerebral and cerebellar cortex.

Cerebellum↗

Consistency of Listing's law and reciprocal innervation with pseudo-inverse control of eye position in 3-D.

Pseudo-inverse kinematics, under which small movements are produced by the least possible sum square changes in motor command, has been proposed as a unifying principle for the elimination of redundancy in general biological motor control systems (Pellionsz 1984) and in particular in the oculomotor system (Daunicht 1988, 1991). We have noted elsewhere (Dean et al, 1999) that this principle is incomplete without first specifying a parameterisation of motor command space and we proposed that the relevant motor-command parameter is summed motor unit firing rate. Under this assumption we were able to show that pseudo-inverse control of the horizontal extraocular muscles is consistent with available motor pool firing rate data. In this paper we extend this result to three dimensions and six extraocular muscles, showing that pseudo-inverse control is consistent with published firing rate date for a realistic model of oculomotor kinematics. We suggest that pseudo-inverse control may represent a common currency for modular control of many degree of freedom systems while its implementation may be a consequences of the minimisation of a more ecologically relevant parameter such as post-saccadic retinal slip.

Biomechanical Phenomena↗

Visual awareness and the cerebellum: possible role of decorrelation control.

The two roles in awareness most often suggested for the cerebellum are (i) keeping the details of motor skills away from forebrain computation, and (ii) signaling to the forebrain when a sensory event is not predictable from prior motor commands. However, it is unclear how current models of the cerebellum could carry out these roles. Their architecture, based on the seminal ideas of Marr and Albus, appears to need 'motor error' to learn correct motor commands. However, since motor error is the difference between the actual motor command and what the command should have been, it is a signal unavailable to the organism in principle. We propose a possible solution to this problem, termed decorrelation control, in which the cerebellum learns to decorrelate the motor command sent to the muscles from the sensory consequences of motor error. This method was tested in a linear model of oculomotor plant compensation in the vestibulo-ocular reflex. A copy of the eye-movement command was sent as mossy-fiber input to the flocculus, represented as a simple adaptive filter version of the Marr-Albus architecture. The sensory consequences of motor error were retinal slip, delivered as climbing fiber input to the flocculus. A standard anti-Hebbian learning rule was used to decorrelate the two. Simulations of the linearized problem showed the method to be effective and robust for plant compensation. Decorrelation control is thus a candidate algorithm for the basic cerebellar microcircuit, indicating how it could achieve motor learning using only signals available to the system. Such learning might then enable the cerebellum to free up visual awareness, and also, by providing a sensory signal decorrelated from motor command, supply awareness with crucial information about the external world.

Algorithms↗

Decorrelation control by the cerebellum achieves oculomotor plant compensation in simulated vestibulo-ocular reflex.

We introduce decorrelation control as a candidate algorithm for the cerebellar microcircuit and demonstrate its utility for oculomotor plant compensation in a linear model of the vestibulo-ocular reflex (VOR). Using an adaptive-filter representation of cerebellar cortex and an anti-Hebbian learning rule, the algorithm learnt to compensate for the oculomotor plant by minimizing correlations between a predictor variable (eye-movement command) and a target variable (retinal slip), without requiring a motor-error signal. Because it also provides an estimate of the unpredicted component of the target variable, decorrelation control can simplify both motor coordination and sensory acquisition. It thus unifies motor and sensory cerebellar functions.

Algorithms↗

Pseudo-inverse control in biological systems: a learning mechanism for fixation stability.

The problem of redundancy in motor control is common to both robotics and biology. Pseudo-inverse control has been proposed as a solution in robotics and appears to be used by the oculomotor system for eye position. Learning mechanisms for implementing pseudo-inverse control using a distributed system of ocular motor units were investigated by modelling integrator calibration for horizontal eye movements. Ocular motoneuron (OMN) input weights were adjusted with a gradient-descent learning rule, using a retinal-slip estimate as an error signal. Firing-rate threshold only became related to motor-unit strength when a noise term was added to OMN firing rates. The learning rule suppressed those units making the largest contribution to the noise-related error, causing the strongest units to have the highest thresholds (size principle). Because the size principle and pseudo-inverse control are related, the trained system approximated pseudo-inverse control over the central +/-35 degrees of the oculomotor range.

Journal Article↗