Eye-Head Coordination for Visual Cognitive Processing

We investigated coordinated movements between the eyes and head (“eye-head coordination”) in relation to vision for action. Several studies have measured eye and head movements during a single gaze shift, focusing on the mechanisms of motor control during eye-head coordination. However, in everyday life, gaze shifts occur sequentially and are accompanied by movements of the head and body. Under such conditions, visual cognitive processing influences eye movements and might also influence eye-head coordination because sequential gaze shifts include cycles of visual processing (fixation) and data acquisition (gaze shifts). In the present study, we examined how the eyes and head move in coordination during visual search in a large visual field. Subjects moved their eyes, head, and body without restriction inside a 360° visual display system. We found patterns of eye-head coordination that differed those observed in single gaze-shift studies. First, we frequently observed multiple saccades during one continuous head movement, and the contribution of head movement to gaze shifts increased as the number of saccades increased. This relationship between head movements and sequential gaze shifts suggests eye-head coordination over several saccade-fixation sequences; this could be related to cognitive processing because saccade-fixation cycles are the result of visual cognitive processing. Second, distribution bias of eye position during gaze fixation was highly correlated with head orientation. The distribution peak of eye position was biased in the same direction as head orientation. This influence of head orientation suggests that eye-head coordination is involved in gaze fixation, when the visual system processes retinal information. This further supports the role of eye-head coordination in visual cognitive processing.     

Static Postural Stability in Women during and after Pregnancy: A Prospective Longitudinal Study

This longitudinal study aimed to compare static postural stability in women between early pregnancy, advanced pregnancy, and at 2 and 6 months postpartum. Forty-five pregnant women were enrolled and 31 completed the protocol. Data were collected at 7–16 and 34–39 weeks gestation, and at 6–10 and 26–30 weeks postpartum. For each subject, the center of foot pressure path length and mean velocity (with directional subcomponents) were computed from 30-s long quiet-standing trials on a stationary force plate with eyes open or closed. The body mass, stance width, and sleep duration within 24 h before testing were also recorded. Static postural stability was not different between pregnancy and postpartum, except for the anterior posterior sway tested in the eyes-closed condition, which was significantly increased in late pregnancy compared to that at 2 and 6 months postpartum. Pregnant/postpartum women’s body mass weakly positively correlated with anterior-posterior sway in the eyes-closed condition and their stance width weakly positively correlated with the anterior-posterior sway in the eyes-open condition. No effect of sleep duration on postural sway was found. Our findings indicate that under visual deprivation conditions women in advanced pregnancy may have decreased static stability compared to their non-pregnant state.     

Conduction of excitation in the cat visual system

Unit responses in area 17 of the visual cortex to stimulation of the lateral geniculate body and optic tract were studied in experiments on unanesthetized cats immobilized with D-tubocurarine. Of the neurons tested, 53.6% responded to stimulation of the lateral geniculate body. In 92% of these cells the responses were orthodromic with latent periods of between 2 and 12.5 msec. Most cells responded with latent periods of 2.0–2.5, 3.0–3.5, and 4.0–4.5 msec, corresponding to latent periods of the components of the electropositive wave of the primary response. Antidromic responses to stimulation of the lateral geniculate body were given by 8% of neurons. The difference between the latent periods of responses of the same visual cortical neurons to stimulation of the optic tract and lateral geniculate body was 0.1–1.8 msec, but for most neurons (55.8%) it was 0.5–1 msec. The histograms of response latencies of visual cortical neurons to stimulation of the above-mentioned formations were found to be similar. It is concluded that the optic radiation contains three principal groups of fibers with conduction velocities of 28.5–16.6, 11.7–8.9, and 7.4–6.0 m/sec, respectively.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 7, No. 6, pp. 589–596, November–December, 1975.     

Benefit of Bi-Ocular Visual Stimulation for Postural Control in Children with Strabismus

Vision is important for postural control as is shown by the Romberg quotient (RQ): with eyes closed, postural instability increases relative to eyes open (RQ = 2). Yet while fixating at far distance, postural stability is similar with eyes open and eyes closed (RQ = 1). Postural stability can be better with both eyes viewing than one eye, but such effect is not consistent among healthy subjects. The first goal of the study is to test the RQ as a function of distance for children with convergent versus divergent strabismus. The second goal is to test whether vision from two eyes relative to vision from one eye provides better postural stability. Thirteen children with divergent strabismus and eleven with convergent strabismus participated in this study. Posturtography was done with the Techno concept device. Experiment 1, four conditions: fixation at 40 cm and at 200 cm both with eyes open and eyes covered (evaluation of RQ). Experiment 2, six conditions: fixation at 40 cm and at 200 cm, with both eyes viewing or under monocular vision (dominant and non-dominant eye). For convergent strabismus, the groups mean value of RQ was 1.3 at near and 0.94 at far distance; for divergent, it was 1.06 at near and 1.68 at far. For all children, the surface of body sway was significantly smaller under both eyes viewing than monocular viewing (either eye). Increased RQ value at near for convergent and at far for divergent strabismus is attributed to the influence of the default strabismus angle and to better use of ocular motor signals. Vision with the two eyes improves postural control for both viewing distances and for both types of strabismus. Such benefit can be due to complementary mechanisms: larger visual field, better quality of fixation and vergence angle due to the use of visual inputs from both eyes.     

Reflection of temporal parameters of an optical stimulus in unitary responses of the sensomotor and visual cortex in cats

Unit activity in the visual (area 17) and sensomotor (areas 4 and 6) cortex in response to an optical stimulus up to 1000 msec in duration was investigated by extracellular recording in acute experiments on cats anesthetized with chloralose (70 mg/kg body weight). Comparative analysis of the types of unitary responses and the durations of the intervals of functional changes showed that: 1) The number of neurons generating on-off responses was about 25% in the visual cortex and 100% in the sensomotor cortex; 2) the intervals of functional changes of the neurons were equal in length to the time intervals of on-off discharges; 3) together with a single time range (200–500 msec), for each area studied specific ranges also exist: from 0 to 200 msec for the visual cortex and from 500 msec and more for the sensomotor cortex; 4) the latent period of after-discharge is equal to the duration of the intervals of functional changes. The results were analyzed from the standpoint of reflection of temporal parameters of optical stimuli by neurons of the sensomotor cortex.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 7, No. 4, pp. 365–371, July–August, 1975.     

Evoked potentials in the visual and association cortex of alert cats during paired uniform stimulation of the lateral geniculate body and pulvinar

Recovery curves of evoked potentials in the association and visual cortex during paired stimulation of the pulvinar in chronic experiments on alert cats were shown to be similar in character. Depression of the test response was observed only if the interval between stimuli was of the order of 10 msec, but if it was 40 msec considerable (2–4 times) facilitation of the second response was observed, mainly on account of an increase in the negative component N₁. Facilitation was less marked if the intervals were from 60 to 100 msec, and they decreased gradually to an interval of 200 msec. The recovery curve of cortical evoked potentials during paired stimulation of the lateral geniculate body differed considerably from the recovery curve during paired stimulation of the pulvinar and was characterized by a gradual increase in amplitude of the second response — from its almost total suppression with an interval of 10 msec to slight facilitation with an interval of 200 msec. If intervals of 10 to 80 msec were used, the test response was restored more slowly in the association cortex than in the visual cortex. The results are discussed from the standpoint of differences in the character of intracortical spread of excitation as a result of activation of geniculo-cortical and pulvinar-cortical pathways of conduction of information.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 16, No. 4, pp. 497–505, July–August, 1984.     

The Influence of Very Low Illumination on the Postural Sway of Young and Elderly Adults

The purpose of the present study was to evaluate the influence of very low ambient illumination and complete darkness on the postural sway of young and elderly adults. Eighteen healthy young participants aged 23.8±1.5 years and 26 community-dwelling elderly aged 69.8±5.6 years were studied. Each participant performed four tests while standing on a force platform in the following conditions: in normal light (215 lx) with open eyes and with closed eyes, in very low illumination (0.25 lx) with open eyes, and in complete darkness with open eyes. The sequences of the tests in the altered visual conditions were determined by random blocs. Postural sway was assessed by means of the force platform measurements. The centre of pressure variables: the medio-lateral and antero-posterior path lengths, mean velocities, sway areas, and fractal dimensions were analysed. Very low illumination resulted in a statistically significant increase in postural sway in both the young and elderly groups compared to normal light, although the increase was significantly smaller than those observed in the eyes closed and complete darkness condition, and no significant effects of illumination on fractal dimensions were detected. The gains of the sways in the very low or no illumination conditions relative to the normal light condition were significantly larger in the group of young participants than in the group of elderly participants (up to 50% and 25%, respectively). However, the response patterns to changes in illumination were similar in the young and elderly participants, with the exception of the short-range fractal dimension of the medio-lateral sway. In conclusion, very low illumination resulted in increased postural sway compared to normal illumination; however, in the closed eye and complete darkness conditions, postural sway was significantly higher than in the very low illumination condition regardless of the age of the participants.     

The Development of Vision in the Zebrafish (Danio rerio)

We studied the development and maturation of the visual system by determining when zebrafish begin to see and to move their eyes. This information was correlated with the time courses of the development of the retina, the retinofugal projection, the retinal image, and the extraocular muscles, to obtain an integrated picture of early visual development. Two visual behaviors were monitored over 48–96 hr postfertilization (hpf). The startle response (body twitch) was evoked by an abrupt decrease in light intensity. The optokinetic response (tracking eye movements) was evoked by rotation of a striped drum. Visually evoked startle developed over 68–79 hpf, more than 20 hr after the onset of a touch-evoked startle. It was not seen in eyeless fish, excluding a role for nonretinal light senses. Tracking eye movements developed over 73–80 hpf. They were always in the direction of drum rotation, even when the fish had been light deprived from blastula stage, ruling out a “trial and error” period of learning to track the drum. The image formed by the ocular lens was examined in intact fish made transparent by suppressing the formation of melanin. The eye was initially far sighted and gradually improved, so that by 72 hpf the image plane coincided with the photoreceptor layer. The extraocular muscles assumed their adult configuration between 66 and 72 hpf. Thus, the retinal image and functional extraocular muscles appeared nearly simultaneously with the onset of tracking eye movements and probably represent the last events in the construction of this behavior.     

Human upright posture control in a virtual visual environment

The sagittal and frontal components of the stabilogram were monitored in 14 healthy subjects standing on a rigid or pliant support under three different conditions of visual control: with the eyes opened (EO), with the eyes closed (EC), or in a virtual visual environment (VVE). Under the VVE conditions, the subjects looked at a three-dimensional image of elements of a room (a 3-D artificial room) that was generated by a computer and locked to the fluctuations of the body center of gravity (CG) so that the visual connection between body sway and shifts of the visual environment typical of normal visual conditions was reproduced. Frequency filtration of the fluctuations of the foot’s center of pressure (FCP) was used to isolate the movements of the vertical projection of the CG and determine the difference between these two variables. The changes in the variables (CG and FCP-CG) were estimated using spectral analysis followed by the calculation of the root mean square (RMS) amplitudes of their spectral fluctuations. In subjects standing on a rigid support, the RMS amplitudes of the spectra of both variables were the highest under the VVE and EC conditions and the lowest under the EO conditions. In subjects standing on a pliant support, body sway was considerably enhanced, which was accompanied by a different pattern of visual influences. The RMS values were the highest under the EC conditions and were lower by a factor of 2–2.5 under the EO and VVE conditions. Thus, it has been demonstrated that the cerebral structures controlling posture ignore the afferent input from the eyes under VVE conditions, if the subject is standing on a rigid support and the CG fluctuations are relatively small; however, this afferentation is efficiently used for maintaining the posture on a pliable support, when the body sway is substantially enhanced.     

Visual gaze control during peering flight manoeuvres in honeybees

As animals travel through the environment, powerful reflexes help stabilize their gaze by actively maintaining head and eyes in a level orientation. Gaze stabilization reduces motion blur and prevents image rotations. It also assists in depth perception based on translational optic flow. Here we describe side-to-side flight manoeuvres in honeybees and investigate how the bees’ gaze is stabilized against rotations during these movements. We used high-speed video equipment to record flight paths and head movements in honeybees visiting a feeder. We show that during their approach, bees generate lateral movements with a median amplitude of about 20 mm. These movements occur with a frequency of up to 7 Hz and are generated by periodic roll movements of the thorax with amplitudes of up to ±60°. During such thorax roll oscillations, the head is held close to horizontal, thereby minimizing rotational optic flow. By having bees fly through an oscillating, patterned drum, we show that head stabilization is based mainly on visual motion cues. Bees exposed to a continuously rotating drum, however, hold their head fixed at an oblique angle. This result shows that although gaze stabilization is driven by visual motion cues, it is limited by other mechanisms, such as the dorsal light response or gravity reception.     

Changes in the direction of vestibulomotor response in the course of adaptation to protracted static head turning in man

Vestibulomotor response during the course of adaptation to prolonged (10 min) static head turning to the furthest limit was investigated in healthy subjects standing upright with the eyes closed. The head was either actively or passively maintained in this position. The sensation of a decline in the angle of head turning was experienced during adaptation to the position by five of the 12 subjects tested. Error in appreciating this angle ranged up to 70–80°. Matching changes occurred in the direction of vestibulomotor response to electrical stimulation of the vestibular apparatus. When true and perceived head position conflict, direction of vestibulomotor response thus matches spatial perception rather than actual orientation of the head.Institute for Research into Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 21, No. 2, pp. 210–217, March–April, 1989.     

Unit responses of the sensomotor cortex to paired electrical stimuli

Unit responses of the sensomotor cortex to paired electrical stimulation and visual cortex, applied either simultaneously or after various delays (from 0 to 200 msec) depend on the order of application of the stimuli and on the interval between them. If stimulation of the sensomotor cortex was used in a conditioning role the response continued unchanged when the intervals between stimuli were increased to 200 msec. If, however, stimulation of the sensomotor cortex had a testing role interaction was observed between the stimuli so that responses to both first and second stimuli were blocked; this was exhibited most clearly for intervals of 40–80 msec between stimuli. The blocking effect persisted on some neurons with delays of up to 200 msec between stimuli, while the response of others to both the first and the second stimulus was restored.Institute of Higher Nervous Activity and Neurophysiology, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 5, No. 6, pp. 628–635, November–December, 1973.     

Coordinates of Human Visual and Inertial Heading Perception

Heading estimation involves both inertial and visual cues. Inertial motion is sensed by the labyrinth, somatic sensation by the body, and optic flow by the retina. Because the eye and head are mobile these stimuli are sensed relative to different reference frames and it remains unclear if a perception occurs in a common reference frame. Recent neurophysiologic evidence has suggested the reference frames remain separate even at higher levels of processing but has not addressed the resulting perception. Seven human subjects experienced a 2s, 16 cm/s translation and/or a visual stimulus corresponding with this translation. For each condition 72 stimuli (360° in 5° increments) were delivered in random order. After each stimulus the subject identified the perceived heading using a mechanical dial. Some trial blocks included interleaved conditions in which the influence of ±28° of gaze and/or head position were examined. The observations were fit using a two degree-of-freedom population vector decoder (PVD) model which considered the relative sensitivity to lateral motion and coordinate system offset. For visual stimuli gaze shifts caused shifts in perceived head estimates in the direction opposite the gaze shift in all subjects. These perceptual shifts averaged 13 ± 2° for eye only gaze shifts and 17 ± 2° for eye-head gaze shifts. This finding indicates visual headings are biased towards retina coordinates. Similar gaze and head direction shifts prior to inertial headings had no significant influence on heading direction. Thus inertial headings are perceived in body-centered coordinates. Combined visual and inertial stimuli yielded intermediate results.     

Analysis of spike discharge of a neuron population in the cat motor cortex during a conditioned change of posture reflex

Spike responses of area 4 neurons in the projection area of the contralateral forelimb to acoustic stimulation (1 sec), which became the conditioned stimulus after training, and to dropping of the platform beneath the test limb, which served as reinforcing stimulus, were studied in trained and untrained cats. Responses only of those neurons which were activated during a passive movement caused by dropping of the platform were studied. In trained animals the number of these neurons which responded to the conditioned stimulus if a reflex occurred was 100%, and in the absence of conditioned-reflex movements to the conditioned stimulus it was 70%, much greater than the number of neurons responding to the same acoustic stimulus in untrained animals (45%). On peristimulus histograms of responses of the test neuron population in untrained and trained animals to acoustic stimulation (in the absence of movements) only the initial spike response with a latent period of under 50 msec and a duration of up to 100 msec could be clearly distinguished. In the presence of reflex movement multicomponent spike responses were observed: an initial spike response and early and late after-responses linked with performance of conditioned-reflex limb flexion. Early after-responses 100–200 msec in duration, appearing after a latent period of 100–150 msec, were linked to the time of application of the conditioned stimulus, whereas the appearance and duration of late after-responses were determined by the time of onset of conditioned-reflex movement. The magnitude of the neuronal response to reinforcement in trained animals does not depend on the appearance of the conditioned movement.A. A. Bogomolets Institute of Physiology, Academy of Sciences of the Ukrainian SSR, Kiev. Translated from Neirofiziologiya, Vol. 17, No. 1, pp. 93–102, January–February, 1985.     

Characteristics of electrical responses by cochlear nuclei in Vespertilionidae and Rhinolophidae to ultrasonic stimuli with various fill frequencies

The total electrical response and action potentials of separate neurons in the cochlear nuclei in Vespertilionidae and Rhinolophidae were investigated. Maximum sensitivity to ultrasound was recorded in Vespertilionidae in the frequency ranges 10–30 and 70–80 kc/sec, and in Rhinolophidae in the frequency ranges 10–30 and 84–86 kc/sec. Mininum off-response thresholds were observed in Vespertilionidae in the range 50–60 kc/sec, and in Rhinolophidae in the range 78–80 kc/sec. The areas of responses by neurons in the cochlear nuclei in both species of bats were similar in shape to those recorded in the same structure in other animals. An exception was provided by Rhinolophidae, in which three peculiar types of neurons were observed: 1) neurons whose response area lay in the frequency ranges up to 78 kc/sec or from 80 to 90 kc/sec; 2) neurons responding in the range 40–90 kc/sec, but not sensitive to stimuli with a fill frequency of 78–80 kc/sec; and 3) neurons whose response area lay in the range 78–80 kc/sec, but in which the character of the response changed from tonic to phasic when there was a change in the fill frequency of the stimulus. Maximum selectivity with regard to fill frequency of stimulus was observed in the neurons of Rhinolophidae in the frequency range 70–90 kc/sec.The term "fill frequency" can be rendered as frequency — Consultants Bureau.A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 3, No. 4, pp. 379–385, July–August, 1971.     

Effect of intracortical chemical stimulation on spontaneous sensomotor cortical unit activity in cats

The effect of microelectrophoresis of glutamate on spontaneous activity of sensomotor cortical neurons located 80–250 µ from the point of application of glutamate was studied in cats anesthetized with pentobarbital. If glutamate was applied at distances of under 100 µ from the neurons the predominant response was one of excitation, evidently due to the direct action of the excitatory mediator. With more distant application inhibition of spontaneous activity predominated: at distances from 100 to 200 µ it was observed in 57%, and between 200 and 250 µ, in 70% of cases. Application of picrotoxin close to the neuron weakened inhibition induced by microelectrophoresis of glutamate through a neighboring microelectrode.A. A. Ukhtomskii Physiological Research Institute, A. A. Zhdanov Leningrad State University. Translated from Neirofiziologiya, Vol. 14, No. 4, pp. 347–352, July–August, 1982.     

Human Postural Responses to Vibratory Stimulation of Calf Muscles under Conditions of Visual Inversion

The authors studied postural responses to bilateral vibratory stimulation (70 Hz, 1 mm, 2 s) of the calf triceps proprioceptors or anterior tibial muscles. Anteroposterior body tilts evoked by vibration were recorded by stabilography. The authors compared the values of postural responses under various conditions of visual control, namely, with normal vision, eyes closed, right–left inversion of the visual space by prismatic spectacles, central vision, and diffuse light. Visual inversion influenced the subjects' proprioceptive postural responses. The amplitude of vibration-evoked shifts of the feet pressure center was minimal with eyes open and significantly increased with eyes closed and inverted vision. Postural responses with visual inversion were significantly stronger than with eyes closed. Since inversion spectacles enabled a subject to see only the central part of the visual field (20°), the reference point was the condition of central vision, i.e., spectacles with same visual angle and without prisms. Postural responses were significantly weaker under these conditions than with visual inversion and eyes closed. Visual field inversion by prismatic spectacles made it impossible to use visual information for stabilizing the human upright posture and, moreover, destabized it. True, this holds only for a randomized experimental protocol, which prevents adaptation to prisms.     

Figure-ground activity in V1 and guidance of saccadic eye movements.

Every day we shift our gaze about 150.000 times mostly without noticing it. The direction of these gaze shifts are not random but directed by sensory information and internal factors. After each movement the eyes hold still for a brief moment so that visual information at the center of our gaze can be processed in detail. This means that visual information at the saccade target location is sufficient to accurately guide the gaze shift but yet is not sufficiently processed to be fully perceived. In this paper I will discuss the possible role of activity in the primary visual cortex (V1), in particular figure-ground activity, in oculo-motor behavior. Figure-ground activity occurs during the late response period of V1 neurons and correlates with perception. The strength of figure-ground responses predicts the direction and moment of saccadic eye movements. The superior colliculus, a gaze control center that integrates visual and motor signals, receives direct anatomical connections from V1. These projections may convey the perceptual information that is required for appropriate gaze shifts. In conclusion, figure-ground activity in V1 may act as an intermediate component linking visual and motor signals.     

Motion and vision: why animals move their eyes

Nearly all animals with good vision have a repertoire of eye movements. The majority show a pattern of stable fixations with fast saccades that shift the direction of gaze. These movements may be made by the eyes themselves, or the head, or in some insects the whole body. The main reason for keeping gaze still during fixations is the need to avoid the blur that results from the long response time of the photoreceptors. Blur begins to degrade the image at a retinal velocity of about 1 receptor acceptance angle per response time. Some insects (e.g. hoverflies) stabilise their gaze much more rigidly than this rule implies, and it is suggested that the need to see the motion of small objects against a background imposes even more stringent conditions on image motion. A third reason for preventing rotational image motion is to prevent contamination of the translational flow-field, by which a moving animal can judge its heading and the distances of objects. Some animals do let their eyes rotate smoothly, and these include some heteropod molluscs, mantis shrimps and jumping spiders, all of which have narrow linear retinae which scan across the surroundings. Hymenopteran insects also rotate during orientation flights at speeds of 100–200° s⁻¹. This is just consistent with a blur-free image, as are the scanning speeds of the animals with linear retinae. Accepted: 29 April 1999     

Visual cortex contribution to the organization of eye movements produced by local electrical stimulation of the cat lateral geniculate body

Eye movements evoked by local electrical stimulation of the dorsal nucleus of the lateral geniculate body were analyzed after removal of the visual cortex and in intact animals during trials on awake cats. No significant difference was observed between the eye movement patterns of the two animal groups evoked by electrical stimulation. These movements could be classed into three main groups: those unassociated with the starting position of the eyes in orbit (or unidirectional movements), goal-directed, and centered movements, with direction depending on the initial position of the eyes in their orbits. Our findings indicate that the cortical visual areas are neither the principal nor an indispensable link in the chain for transmitting signals evoked by (electrically) stimulating the geniculate body from the cortical structures of the direct visual pathway towards the operative links of the oculomotor system. Potential pathways for conducting information from the dorsal nucleus of the lateral geniculate body to oculomotor system structures are discussed.I. P. Pavlov Institute of Physiology, Academy of Sciences of the USSR, Leningrad. Translated from Neirofiziologiya, Vol. 19, No. 2, pp. 164–170, March–April, 1987.