Biomedical subjects

J W Morley

Publications and source records attributed to J W Morley.

At least 19 recordsLinked to original sources

PubMed2005

Synaptic plasticity in the dy2J mouse model of laminin alpha2-deficient congenital muscular dystrophy.

Laminin alpha2-deficient congenital muscular dystrophy is a debilitating disease affecting both muscle and neural tissue as a result of mutations in the LAMA2 gene. It presents at or soon after birth with muscle weakness and is further characterised by clinical central nervous system involvement. Laminin alpha2 is part of the extracellular matrix, linked to the cellular cystoskeleton via dystroglycan which is an integral part of the dystrophin-glycoprotein complex (DGC). We examined both short- and long-term synaptic plasticity in the C57BL6J/dy(2J) mouse, an animal model of laminin alpha2 deficient congenital muscular dystrophy. Using a cerebellar slice preparation, we show that the pre-synaptically mediated paired-pulse facilitation (PPF) was no different between dy(2J) and littermate controls. Approximately half (7/12) the dy(2J) Purkinje cells displayed a blunted LTD compared to littermate controls, and one third (4/12) of dy(2J) Purkinje cells displayed LTP. This study demonstrates that a defective laminin alpha2 causes a disruption in long-term synaptic plasticity at the Purkinje cell-parallel fibre synapse.

Animals↗

PubMed2002

Coding of disparity information in extrastriate cortex of the cat.

We have used information theory to analyse the responses of neurons in area 21a of the cat to disparity stimuli. Visual stimuli consisted of drifting sinusoidal gratings presented simultaneously to each eye. The relative spatial phase of the gratings varied between stimulus periods in a pseudo-random sequence of 45 degrees increments that covered the full 360 degrees. The mean information content of the responses of all neurons across all phases was 0.72 bits (+/-0.10, SE, n=29). The information conveyed by each neuron was well correlated with the extent to which the interocular phase difference modulated the response of the cell. However, information content was not simply related to firing rate, as there was usually significant information content in the neuronal responses to phase differences that elicited the minimum firing rate. In general, burst responses (impulse intervals <4 ms) did not convey more information than that conveyed by the total response. The contribution to the cumulative information of the response in successive 100-ms segments decreased over the course of the 1-s stimulus. The ratio of information transmitted at 200 ms to that transmitted over the full second had a median of 0.30 while the ratio of 500 ms to 1 s was 0.68.

Action Potentials↗

PubMed2002

Brain function in Duchenne muscular dystrophy.

Duchenne muscular dystrophy (DMD) is the second most commonly occurring genetically inherited disease in humans. It is an X-linked condition that affects approximately one in 3300 live male births. It is caused by the absence or disruption of the protein dystrophin, which is found in a variety of tissues, most notably skeletal muscle and neurones in particular regions of the CNS. Clinically DMD is characterized by a severe pathology of the skeletal musculature that results in the premature death of the individual. An important aspect of DMD that has received less attention is the role played by the absence or disruption of dystrophin on CNS function. In this review we concentrate on insights into this role gained from investigation of boys with DMD and the genetically most relevant animal model of DMD, the dystrophin-deficient mdx mouse. Behavioural studies have shown that DMD boys have a cognitive impairment and a lower IQ (average 85), whilst the mdx mice display an impairment in passive avoidance reflex and in short-term memory. In DMD boys, there is evidence of disordered CNS architecture, abnormalities in dendrites and loss of neurones, all associated with neurones that normally express dystrophin. In the mdx mouse, there have been reports of a 50% decrease in neurone number and neural shrinkage in regions of the cerebral cortex and brainstem. Histological evidence shows that the density of GABA(A) channel clusters is reduced in mdx Purkinje cells and hippocampal CA1 neurones. At the biochemical level, in DMD boys the bioenergetics of the CNS is abnormal and there is an increase in the levels of choline-containing compounds, indicative of CNS pathology. The mdx mice also display abnormal bioenergetics, with an increased level of inorganic phosphate and increased levels of choline-containing compounds. Functionally, DMD boys have EEG abnormalities and there is some preliminary evidence that synaptic function is affected adversely by the absence of dystrophin. Electrophysiological studies of mdx mice have shown that hippocampal neurones have an increased susceptibility to hypoxia. These recent findings on the role of dystrophin in the CNS have implications for the clinical management of boys with DMD.

Adult↗

PubMed1999

Signalling of static and dynamic features of muscle spindle input by external cuneate neurones in the cat.

1. The present experiments examined the capacity of external cuneate nucleus (ECN) neurones in the anaesthetized cat to respond to static and vibrotactile stretch of forearm extensor muscles. The aim was to compare their signalling capacities with the known properties of main cuneate neurones in order to determine whether there is differential processing of muscle spindle inputs at these parallel relay sites. 2. Static stretch (<= 2 mm in amplitude) and sinusoidal vibration were applied longitudinally to individual muscle tendons and responses recorded from single ECN neurones. The muscle-related ECN neurones that were sampled displayed a high sensitivity to both static and dynamic components of stretch, including muscle vibration at frequencies of 50-800 Hz, consistent with their dominant input being derived from primary spindle afferent fibres. 3. In response to ramp-and-hold muscle stretch, ECN neurones resembled their main cuneate counterparts in the pattern of their responses and in quantitative response measures. Their coefficients of variation in interspike intervals during steady stretch ranged from approximately 0.3 to 0.7, as they do in main cuneate responses, and their stimulus-response relations were graded as a function of stretch magnitude with low variability in responses at a fixed stretch amplitude. 4. In response to muscle vibration, ECN activity was tightly phase locked to the vibration waveform, in particular at frequencies of <= 150 Hz, where vector strength measures (R) were high (R >= 0.8) before declining as a function of frequency, with R values of approximately 0.6 at 300 Hz and <= 0.4 at 800 Hz. Both the qualitative and quantitative aspects of ECN responsiveness to the vibro-stretch disturbances were indistinguishable from those of the main cuneate neurones. 5. The results demonstrate a high transmission fidelity for muscle signals across the ECN and no evidence for differential synaptic transmission across the parallel main and external cuneate nuclei. Earlier limitations observed in the capacity of cerebellar Purkinje cells to respond to primary spindle inputs must therefore be imposed at synapses within the cerebellum.

Algorithms↗

PubMed1999

Binocular interactions in area 21a of the cat.

We investigated binocular suppression in area 21a cells of the anaesthetized cat using drifting sinusoidal gratings presented simultaneously to each eye. The grating presented to the dominant eye was always oriented optimally and the grating presented to the non-dominant eye was either at the same orientation, but at the least effective relative spatial phase, or orthogonal. The binocular response of approximately 80% of cells was less than the monocular dominant eye response when there was a mismatch in orientation or spatial location between stimuli. Response suppression in the two binocular stimulus conditions had a correlation coefficient (r) of 0.55. We propose a parsimonious model to account for the response facilitation and suppression by binocular stimulation of area 21a neurons.

Animals↗

PubMed1999

Binocular phase interactions in area 21a of the cat.

1. Binocular interactions related to retinal disparity were investigated in single neurons in area 21a of extrastriate cortex in the anaesthetized cat using sinusoidal luminance gratings. 2. The responses of approximately two-thirds of neurons were profoundly modulated by a relative phase difference between identical drifting gratings presented to each eye. This modulation included both facilitatory and inhibitory interocular interactions. The selectivity for binocular disparity was about twice as sharp as the selectivity for monocular spatial position. 3. Significant phase modulation was retained in many neurons at interocular orientation differences exceeding 45 deg. The response suppression associated with stimulation at a phase shift 180 deg from the optimum was stronger than the response suppression to an interocular orientation difference of 90 deg. 4. The proportion of phase modulated neurons and the potency of modulation in area 21a neurons exceed that reported for phase-selective complex cells in area 17. Neurons in area 21a show sharp disparity tuning that is relatively insensitive to changes in orientation and monocular position, which suggests that this extrastriate region has a role in stereoscopic depth perception.

Analysis of Variance↗

PubMed1998

Signalling of static and dynamic features of muscle spindle input by cuneate neurones in the cat.

1. The capacity of cuneate neurones to signal information derived from muscle spindle afferent fibres about static stretch or vibration of forearm extensor muscles was examined electrophysiologically in anaesthetized cats. 2. Static stretch (>= 2 mm in amplitude) and sinusoidal vibration (at frequencies of 50-800 Hz) were applied longitudinally to individual muscle tendons by means of a feedback controlled mechanical stimulator, and responses were recorded from individual cuneate neurones and from individual spindle afferent fibres. 3. Cuneate neurones sampled were located caudal to the obex and displayed a sensitivity to both vibration and static stretch of forearm muscles that was consistent with their input arising from primary spindle endings. In response to static muscle stretch, they displayed graded and approximately linear stimulus-response relations, and a stability of response level at fixed lengths that was consistent with these neurones contributing discriminative information about static muscle stretch. 4. In response to sinusoidal muscle vibration the cuneate neurones also showed graded stimulus-response relations (in contrast to spindle afferents which at low vibration amplitudes attain a plateau response level corresponding to a discharge of 1 impulse on each vibration cycle). Lowest thresholds were at 100-300 Hz and bandwidths of vibration sensitivity extended up to approximately 800 Hz. 5. Temporal precision in cuneate responses to muscle vibration was assessed by constructing phase scatter and cycle histograms from which measures of vector strength could be calculated. Cuneate responses displayed somewhat poorer phase locking (and lower vector strengths) than spindle afferent responses to vibration (a reflection of uncertainties associated with synaptic transmission). Nevertheless, the remarkable feature of cuneate responses to muscle vibration is the preservation of tight phase locking at frequencies up to 400-500 Hz, which presumably enables these central neurones to contribute accurate temporal information for the kinaesthetic sense in a variety of circumstances involving dynamic perturbations to skeletal muscle.

Animals↗

PubMed1997

Orientation-dependent binocular interactions in area 21a of the cat.

We investigated binocular interactions in area 21a cells of the anaesthetized cat. Visual stimuli were drifting sinusoidal gratings presented at the same optimal orientation in each eye (iso-oriented condition) or at the optimal orientation in the dominant eye and the orthogonal orientation in the other eye (orthogonal condition). In 68% of cells the response in the binocular iso-oriented condition was greater than the dominant eye monocular response, while in 88% of cells the response in the binocular orthogonal condition was less than the dominant eye monocular response. Our results suggest a possible role for this extrastriate region of cortex in binocular contour rivalry.

Animals↗

PubMed1997

Spatial and temporal frequency selectivity of cells in area 21a of the cat.

1. The spatial and temporal response properties of single cells in area 21a of the anaesthetized cat were assessed using drifting sinusoidal gratings presented at the optimum orientation for each cell. 2. Responses to sinusoidal gratings were dominated by an elevation of the mean discharge, with a relatively small modulated component at the temporal frequency of grating drift. The relative modulation ratio for the majority of cells was less than 1, similar to complex cells in the striate cortex. 3. Of those cells responsive to stimulation with sinusoidal gratings, 94% displayed spatial bandpass characteristics. Values derived from spatial frequency tuning curves were: mean optimum spatial frequency, 0.26 cycles deg-1; mean spatial resolution, 0.86 cycles deg-1; mean spatial bandwidth, 1.8 octaves; and mean normalized bandwidth, 1.3. Two cells (6%) displayed spatial low-pass characteristics. 4. Approximately half our sample of cells (44%) displayed temporal low-pass tuning, while 35% displayed temporal bandpass characteristics. The mean optimum temporal frequency of bandpass cells was 3.3 Hz and the mean temporal bandwidth 1.9 octaves. The remaining cells were classified as temporal broadband (17%) and temporal high-pass (4%). 5. We conclude that the dominant functional input to cells with relatively high spatial frequency selectivity and/or temporal low-pass response properties most probably arises from area 17. The responses of the remaining cells may be explained by input from area 17 or 18.

Animals↗

PubMed1997

Corticocortical connections between area 21a and primary visual cortex in the cat.

We investigated the corticocortical connections between area 21a and ipsilateral areas 17 and 18 in the cat. The anterograde/retrograde fluorescent tracer tetramethyl-rhodamine conjugated to dextran (Fluoro-Ruby) was injected into area 21a of the anaesthetized cat. Cell bodies labelled retrogradely from area 21a were consistently observed in both areas 17 and 18, primarily located in the supragranular layers of cortex where they formed discrete patches of cells. Similar numbers of cell bodies were labelled retrogradely in areas 17 and 18 of each animal. Our data are also consistent with previous reports of a reciprocal projection from area 21a back to areas 17 and 18 terminating principally in infragranular cortical layers.

Animals↗

PubMed1997

Development of contrast sensitivity and temporal-frequency selectivity in primate lateral geniculate nucleus.

We studied the development of spatial contrast-sensitivity and temporal-frequency selectivity for neurons in the monkey lateral geniculate nucleus. During postnatal week 1, the spatial properties of P-cells and M-cells are hardly distinguishable, with low contrast-sensitivity, sluggish responses, and poor spatial resolution. The acuity of P-cells improves progressively until at least 8 months, but there is no obvious increase in their maximum contrast-sensitivity with age. The contrast sensitivity of M-cells is already clearly higher than that of P-cells by 2 months, and at 8 months of age this characteristic difference between M- and P-cells approaches the adult pattern. There is a major increase in responsiveness during the first 2 postnatal months, especially for M-cells, the peak firing rate of which rises fivefold, on average, between birth and 2 months. Many P-cells in the neonatal and 2-month-old animals did not give statistically reliable responses to achromatic gratings, even at the highest contrasts: this unresponsiveness of P-cells might result from low gain and/or chromatic opponency. The upper limit of temporal resolution in the neonate is low--about one-third of that in the adult. Among M-cells, the improvement in temporal resolution, like that in contrast sensitivity, is rapid over the first 2 months, followed by a slower change approaching the adult value by 8 months of age. The development of contrast sensitivity, responsiveness and temporal tuning are little affected, if at all, by binocular deprivation of pattern vision from birth for even a prolonged period.

Aging↗

PubMed1997

Accommodation in binocular contour rivalry.

Although contour rivalry is known to suppress the contribution of the non-dominant eye to some visuomotor mechanisms such as the pupillary light reflex, there have been no reports of the impact of rivalry on accommodation control. In the situation where the accommodation demands in the two eyes are in dynamic conflict, it has been reported that the accommodation response can be modelled in terms of a vector average of the appropriate response in the two eyes. This study compared the binocular interactions in the accommodation system with rivalrous and non-rivalrous stimuli. Accommodation was continuously monitored with an infrared optometer, while the accommodation demand in the two eyes was dynamically modulated independently in the two eyes. When the visual target was perceptually rivalrous the previously described binocular interactions were abolished and the accommodation response closely followed the accommodation demand presented to the dominant eye.

Accommodation, Ocular↗

PubMed1995

Parallel processing of tactile information in cat cerebral cortex: effect of reversible inactivation of SII on SI responses.

1. Responsiveness of neurons in the distal forelimb region of primary somatosensory cortex (SI) was examined in cat in association with the cooling-induced, reversible inactivation of the corresponding region of the second somatosensory area (SII). The aim was to test whether a component of the stimulus-generated tactile input to SI came via an indirect, intracortical path from the thalamus through SII, or whether, when SI responsiveness fell in association with SII inactivation, the effect could be explained by a disfacilitation of the SI neuron; that is, a removal of a tonic facilitatory influence on the SI neuron that arises from within SII. 2. The responses of 33 SI neurons to controlled tactile stimuli, usually 1-s long trains of vibration or rectangular pulses delivered to the skin of the distal forelimb, were examined quantitatively before, during, and after the rapid, reversible inactivation of the SII area. 3. Nineteen of the 33 neurons (approximately 60%) were unaffected in their response level by SII inactivation. These included neurons of several functional classes whose input came from different classes of tactile afferent fibers, including the Pacinian corpuscle (PC) associated fibers, other rapidly adapting (RA) afferents from glabrous skin, and presumed hair follicle afferent (HFA) fibers. The remaining 14 neurons (approximately 40%), which also included different functional classes, displayed a reduction in response level with SII inactivation. Because this was not accompanied by significant prolongation of the SI spike waveforms, it is not attributable to direct spread of cooling from SII to SI. Construction of stimulus-response relations demonstrated that any effect of SII inactivation on individual SI neurons was consistent over the whole response range. 4. The fall in responsiveness for some SI neurons in association with SII inactivation may be attributable to disfacilitation, that is, a loss of tonic facilitation arising in SII, rather than to a block of peripherally generated inputs that traverse an indirect path from the skin to SI, via SII. There are three reasons for suggesting this. First, in the course of SII cooling, the latency and time course of SI evoked potentials were not delayed in a way that might be expected if part of the SI response had come via SII. Second, the SII inactivation could reduce the SI spontaneous activity (as well as the stimulus-related responsiveness). The facilitation from SII is therefore not necessarily dependent on overt tactile stimulation, and its source may therefore be endogenous to SII.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

PubMed1993

The role of single touch domes in tactile perception.

Detection thresholds to step indentations applied to touch domes were measured and compared with threshold values obtained on adjacent areas of hairy skin. The mean of the detection thresholds determined on touch domes for five subjects was 504 +/- 27 microns (SEM). The detection thresholds found off the touch domes (2-3 mm away) were somewhat higher in all ten cases (two studies per subject), with a mean value of 587 +/- 28 microns, which was significant at the 0.01 level. These thresholds are an order of magnitude greater than those that elicit responses in touch-dome-associated slowly adapting type I (SAI) fibres in cats, and, together with the small difference between the detection thresholds determined on and off touch domes, indicate that the activity of single touch dome SAI fibres does not mediate the detection of steady indentation in the hairy skin.

Adult↗

PubMed1992

Texture discrimination in carpal tunnel syndrome.

The ability to discriminate textured surfaces was measured in patients with carpal tunnel syndrome (CTS). Patients were initially diagnosed using clinical and electrophysiological criteria. The textured surfaces were gratings of alternating ridges and grooves. The gratings differed in their spatial period only, with the ratio of the ridge width to the groove width remaining constant (1:5). A two-alternative forced-choice paradigm was employed in which subjects, both patients and age-matched controls, rubbed their index finger (D2) or little finger (D5) back and forth across the surfaces. The grating spatial period at which subjects could discriminate a difference between the standard grating (spatial period = 2000 microns) and the comparison grating (spatial period in the range 2000-2900 microns) with a probability of 0.75 was taken as the measure of discriminative ability. Statistical comparison of the mean 75% values showed that: (i) when patients used D2 their discriminative ability was significantly impaired in comparison with the discriminative ability of controls using either D2 or D5; (ii) there was no significant difference in discriminative ability between patients and controls when patients used D5 to discriminate the textures; (iii) the 75% values for patients using D2 or D5 did not differ significantly. The degree of abnormality of each patient's sensory evoked potential did not allow us to predict their subsequent performance on the discrimination task.

Adult↗

PubMed1992

Parallel processing of tactile information in the cerebral cortex of the cat: effect of reversible inactivation of SI on responsiveness of SII neurons.

1. Localized cortical cooling was employed in anesthetized cats for the rapid reversible inactivation of the distal forelimb region within the primary somatosensory cortex (SI). The aim was to examine the responsiveness of individual neurons in the second somatosensory area (SII) in association with SI inactivation to evaluate the relative importance for tactile processing of the direct thalamocortical projection to SII and the indirect projection from the thalamus to SII via an intracortical path through SI. 2. Response features were examined quantitatively before, during, and after SI inactivation for 29 SII neurons, the tactile receptive fields of which were on the glabrous or hairy skin of the distal forelimb. Controlled mechanical stimuli that consisted of l-s trains of either sinusoidal vibration or rectangular pulses were delivered to the skin by means of small circular probes (4- to 8-mm diam). 3. Twenty-three of the 29 SII neurons (80%) showed no change in response level (in impulses per second) as a result of SI inactivation. These included seven neurons activated exclusively or predominantly by Pacinian corpuscle (PC) receptors, six that received hair follicle input, four activated by convergent input from hairy and glabrous skin, and six driven by dynamically sensitive but non-PC inputs from the glabrous skin. 4. Six SII neurons (20%), also made up of different functional classes, displayed a reduction in response to cutaneous stimuli when SI was inactivated. 5. Stimulus-response relations, constructed by plotting response level in impulses per second against the amplitude of the mechanical stimulus, showed that the effect of SI inactivation on individual neurons was consistent over the whole response range. 6. The reduced response level seen in 20% of SII neurons in association with SI inactivation cannot be attributed to direct spread of cooling from SI to the forelimb area of SII, as there was no evidence for a cooling-induced prolongation in SII spike waveforms, an effect that is known to precede any cooling-induced reduction in responsiveness. 7. As SI inactivation produced a fall in spontaneous activity in the affected SII neurons, we suggest that the inactivation removes a source of background facilitatory influence that arises in SI and affects a small proportion of SII neurons. 8. Phase-locking and therefore the precision of impulse patterning were unchanged in the responses of SII neurons to vibration during SI inactivation. This was the case whether response levels of neurons were reduced or unchanged by SI inactivation.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

PubMed1992

Role of monkey midbrain near-response neurons in phoria adaptation.

1. One striking characteristic of the way in which accommodation and convergence of the eyes are organized is that although the two responses are usually tightly coupled, a brief period of binocular viewing through prisms that require extra convergence brings about a persistent, adaptive alteration in the relationship between the two responses: the vergence during monocular viewing of a target at a given distance is biased in a convergent direction. We sought to discover the role of the near-response neurons we have previously described in the monkey midbrain in such phoria adaptation. 2. Phoria adaptation was produced in two monkeys by having them view binocularly stereoscopic targets under conditions that mimicked prism viewing, i.e., the mirrors of the stereoscope were set so as to require more convergence than that associated with a real target at the same distance as the images seen in the stereoscope. The activity of 57 near-response neurons located dorsally and dorsolaterally to the oculomotor nucleus was recorded before and after adaptation while the monkeys monocularly viewed targets at a range of distances. 3. Comparison of a neuron's response in normal binocular viewing with the response when the accommodation and vergence stimuli were in conflict allowed us to distinguish 24 neurons that behaved as though they were related exclusively to the vergence response. 5 neurons that behaved as though they were exclusively related to the accommodation response, and 12 neurons whose firing was not so simply related to either response. We were unable to classify the remaining 16 near-response cells by this method. 4. In accommodation-related neurons, the relationship between firing rate and accommodation did not alter, or only altered slightly, when the animal's phoria was adapted. 5. The relationship between firing rate and vergence was unaltered by phoria adaptation in only a small proportion of vergence-related neurons, showing that most neurons do not carry the entire signal responsible for phoria adaptation. On the other hand, in the majority of vergence-related neurons the relationship between firing rate and accommodation was altered by phoria adaptation, showing that most neurons do carry part of the phoria adaptation signal. 6. The implication is that the increase in vergence observed after adaptation is mediated at more than one site. A proportion of the phoria adaptation signal is present at the level of the midbrain vergence-related neurons, with the remainder of the signal being added later, presumably at the motoneurons.

Accommodation, Ocular↗

PubMed1992

Binocular interactions in accommodation control: effects of anisometropic stimuli.

In binocular viewing of real targets, the accommodative demand in the two eyes is not in general identical, yet the accommodation response in the two eyes is equal. In order to investigate how the accommodative signals from the two eyes are combined, this study has examined the effects of several forms of dynamic anisometropic stimulation on the accommodation response in both man and the rhesus monkey (Macaca mulatta). All experiments were performed in a computer-controlled haploscopic apparatus to allow independent control of the accommodative stimuli to the two eyes and of the vergence stimulus. The vergence stimulus was held constant while the accommodation demand was modulated independently in each eye. Accommodation was monitored continuously with a dynamic infrared optometer. Four anisometropic conditions were used. In two of these conditions, accommodation demand was varied sinusoidally with time in both eyes, but with phases differing by 90 degrees or 180 degrees between the two eyes. In the two remaining conditions, accommodation demand in one eye varied sinusoidally, while the accommodation demand was constant in the other. In all cases, the form of the target pattern was identified in the two eyes. The accommodation responses observed with these stimulus conditions were similar in both man and the monkey. When presented with conflicting stimuli in the two eyes, the accommodation response appeared to be best described as a compromise between the inputs to the two eyes; there were no indications of a purely random alternation of eye dominance of the form seen in binocular contour rivalry. When the accommodation demand was modulated in only one eye, there was a modulated accommodation response of similar phase to the control condition (i.e., both eyes modulated in phase) but with a much smaller gain (mean, 39% of control gain). When the accommodation demand was modulated in both eyes with a phase difference of 180 degrees, no significant modulation was observed in the accommodation response at the stimulation frequency. When the interocular phase difference was 90 degrees, a modulated response was observed that showed a mean phase lag 41 degrees more than that observed in the control condition (both eyes modulated in phase) and an appreciably smaller gain (mean, 55% of control gain). The extent to which the results can be described by a linear vector average of the uniocular inputs is considered.

Accommodation, Ocular↗