Eye movements in Daphnia pulex (De Geer).

1. The various types of eye movement exhibited by the cyclopean eye of Daphnia pulex were studied using high speed motion photography. 2. This rudimentary eye, which consists of only 22 ommatidia, can move through approximately 150 degrees in the sagittal plane and 60 degrees in the horizontal plane. 3. Four classes of eye movement were found: (1) a high speed tremor at 16 Hz with an amplitude of 3-4 degrees, which resembles physiological nystagmus, (2) a slow rhythmic scanning movement at 4 Hz, and 5-6 degrees amplitude, (3) large fast eye movements similar to saccadic eye movements and (4) optokinetic nystagmus produced by moving striped patterns. 4. Where the fast tremor occurred concurrently with the slow rhythmic scan, a Fourier analysis revealed that the former was the fourth harmonic of the latter.     

Torsion movements of the human eye. IV. Passive body movements, effect of skeletal muscle reflexes, opticokinetic torsion tracking

A comparative study of torsional movement of the eye in passive and active tilting of the head and body of the object was carried out. Similarity of torsional movement of the eyes in passive and active movements was shown. It was found by the method of exclusion and selective stimulation of vestibular, cervikal, lumbar optokinetic reflexes, that neither the cervikal, nor lumbar reflexes elicited spontaneous torsional movements of the eyes and had no influence on them. A direct study (coinciding with rotation direction of the stimulus of head rotation) and the reverse (noncoinciding) torsional tracing of a rotating disc and tracing without head movements was investigated. During direct tracing depression of saccades and extention of the slow phase of torsion was found; during the reverse one--a decrease of the eye drist and increase of the amplitude and number of saccades. Phenomena of a seeming acceleration and deceleration of disc rotation etc. have been observed. It was found that with torsional saccades vision was retained. The presence of optokinetic control of phases of torsional eye movements formation has been recorded. Tracing without rotation of the head was accompanied by torsional nistagmus. Possible causes of incomplete stabilisation and optokinetic torsional tracing are discussed.     

Positioning of the Human Forearm in Tracking Movements and Their Reproduction under Conditions of Limited Visual Control

In 14 healthy persons, we studied movements of the forearm with its positioning on a target level. A double trapezium was used as the command trajectory (flexion in the elbow joint from the state of full extension, 0°, with positioning on the level of 50 or 60° and further flexion to the 100° angle, and a similar reverse movement). We compared (i) tracking movements, when the subject tried to adequately reproduce the movement of the target along the command trajectory visualized on the monitor screen and obtained visual information about the performed movement (shifts of the second light point in time/joint angle coordinates), and (ii) reproduction of these movements under conditions of limitation of the visual feedback (when there was no information about the performed movement). Parameters of the tracking movements and of their reproductions (delays of initiation of the movement phases as compared with the command signal, durations of these phases, and angle velocities of the forearm movement), as well as the quality of positioning after oppositely directed movements, were compared. Positioning on the target level performed under proprioceptive control (when visual control was limited) was accompanied by systematic errors, whose sign in most test series performed by most subjects coincided with the direction of the preceding movement phase. The pattern of signs of systematic positioning errors after movements of opposite directions was quite individual (typical of a given subject) and demonstrated no dependence on the value of the extensor loading. Averaged intragroup systematic errors of positioning after movement phase 1 (flexion to the target level) and phase 3 (extension to the same level) under conditions of a minimum extensor loading (0.5-1.0 N · m) were 2.57° and 2.52°, respectively. When the loading was substantial (3.6-6.0 N · m), the respective errors were 3.85° and 3.48°. The nonlinear properties of muscle stretch receptors in the elbow flexors and extensors (responsible for the significant dependence of the parameters of afferent signals produced in these receptors on the movement prehistory) are considered the primary reason for systematic errors when positioning is performed exclusively under proprioceptive control. The influence of alpha-gamma co-activation in active muscles on the characteristics of the above signals is discussed.     

The role of background movement in goldfish vision

In experiments described in the literature objects presented to restrained goldfish failed to induce eye movements like fixation and/or tracking. We show here that eye movements can be induced only if the background (visual surround) is not stationary relative to the fish but moving. We investigated the influence of background motion on eye movements in the range of angular velocities of 5–20° s⁻¹. The response to presentation of an object is a transient shift in mean horizontal eye position which lasts for some 10 s. If an object is presented in front of the fish the eyes move in a direction such that it is seen more or less symmetrically by both eyes. If it is presented at ±70° from the fish's long axis the eye on the side of the object moves in the direction that the object falls more centrally on its retina. During these object induced eye responses the typical optokinetic nystagmus of amplitude of some 5° with alternating fast and slow phases is maintained, and the eye velocity during the slow phase is not modified by presentation of the object. Presenting an object in front of stationary or moving backgrounds leads to transient suppression of respiration which shows habituation to repeated object presentations. Accepted: 14 April 2000     

Sensitivity of the Goldfish Motion Detection System Revealed by Incoherent Random Dot Stimuli: Comparison of Behavioural and Neuronal Data

Background

Global motion detection is one of the most important abilities in the animal kingdom to navigate through a 3-dimensional environment. In the visual system of teleost fish direction-selective neurons in the pretectal area (APT) are most important for global motion detection. As in all other vertebrates these neurons are involved in the control of slow phase eye movements during gaze stabilization. In contrast to mammals cortical pathways that might influence motion detection abilities of the optokinetic system are missing in teleost fish.

Results

To test global motion detection in goldfish we first measured the coherence threshold of random dot patterns to elicit horizontal slow phase eye movements. In addition, the coherence threshold of the optomotor response was determined by the same random dot patterns. In a second approach the coherence threshold to elicit a direction selective response in neurons of the APT was assessed from a neurometric function. Behavioural thresholds and neuronal thresholds to elicit slow phase eye movements were very similar, and ranged between 10% and 20% coherence. In contrast to these low thresholds for the optokinetic reaction and APT neurons the optomotor response could only be elicited by random dot patterns with coherences above 40%.

Conclusion

Our findings suggest a high sensitivity for global motion in the goldfish optokinetic system. Comparison of neuronal and behavioural thresholds implies a nearly one-to-one transformation of visual neuron performance to the visuo-motor output. In addition, we assume that the optomotor response is not mediated by the optokinetic system, but instead by other motion detection systems with higher coherence thresholds.     

Reproduction of tracking movements and target positioning of the forearm in humans in the absence of visual control

In 17 healthy volunteers, we studied movements of the forearm, which included episodes of positioning on the target level. The trajectory of the non-ballistic (relatively slow) movement looked like a double trapezium (flexion of the elbow joint from the state of full extension, 0 deg, positioning on the 50 deg level, further flexion to the limit angle of 100 deg, and a similar reverse sequence). The command trajectory and the trajectory of the realized movement were visualized with movements of cursors on a monitor in time/joint angle coordinates. We compared parameters of the tracking movements (in the presence of visual feedback) and their blindfold reproduction (with the complete absence of visual control). It was found that blindfold reproduction movements differ from sample tracking movements and their reproduction with partial limitation of visual control [16] in higher peak velocities and shorter durations, i.e., a trend toward conversion of such movements into ballistic ones was observed. Under conditions of elimination of visual control, movements that led to positioning were mostly hypermetric, i.e., positioning was usually accompanied by positive systematic errors (whose sign coincided with the direction of the preceding movement phase). The mean intragroup value of the systematic error of the first positioning (after flexion to the target level) was +6.73 ± 1.15 deg, while the respective mean for the second positioning (after extension to the same level) was +4.00 ± 1.31 deg. The nonlinear properties of stretch receptors of muscles whose activity provides the formation of a proprioceptive estimate of the joint angle are considered the crucial reason for systematic errors of blindfold positioning.Neirofiziologiya/Neurophysiology, Vol. 36, Nos. 5/6, pp. 393–404, September–December, 2004.This revised version was published online in April 2005 with a corrected cover date and copyright year.     

Movement sensitivities of cells in the fly's medulla

Summary Intracellular recordings were made in the medullae of intact, restrained females ofCalliphora vicina that faced a hemispherical, minimum-distortion surface upon which moving patterns and spots were projected from the rear (Fig. 2). In the distal medulla, noisy hyperpolarizations to light, most likely recorded in terminals of laminar (L) cells, had flicker-like oscillations to moving gratings of 15° spatial wavelength but not of 2.5° spatial wavelength (Fig. 3). Medullary (M) cells penetrated distally responded to grating movements with similar but depolarizing oscillations, in one cell 180° out of phase with a nearby laminar response (Figs. 4–6).A characteristic movement response recorded from most medullary cells consisted of abrupt, maintained nondirectional depolarizations in response to movements of gratings, often with directional ripple or spikes superimposed. When directions of movement reversed, there were brief repolarizations, but when movements stopped, depolarizations decayed away more slowly (Figs. 7 and 8). Magnitude of responses increased with increasing speeds of both 15° and 2.5° gratings (Figs. 9–11). In some cells, there were delayed decays of responses after stopping (Fig. 12). Still other cells seemed to receive inhibition from other, characteristically responding cells (Fig. 13).Receptive fields tested were simple and usually large, with only a suggestion of surround inhibition (Fig. 14). In general, intensity and position were interchangeable over a cell's receptive field (Figs. 15 and 16). Moving edges and dark spots elicited responses primarily within receptive field centers (Figs. 18–20).It is argued that waveforms of characteristic movement responses can be explained by multiplicative inputs from L- and M-cells to movement detectors (Figs. 21–26).Abbreviations L cells laminar (monopolar) cells - M cells medullary cells     

Eye-scanning during walking in the crabLeptograpsus variegatus

Summary The eyes of the crabLeptograpsus variegatus scan continually when the animal walks. The scanning movements are in the horizontal plane, have an amplitude of between 0.1° and 0.3° and a frequency of about 6 Hz if the animal is surrounded by a bright, contrasting visual field. The scanning movements are abolished if the animal is placed in the dark, or blinded. During scanning the two eyes are predominantly in phase with each other. It is proposed that the scanning is the result of a general increase of activity in the oculomotor neurons during walking, which causes the eyes to oscillate at a frequency which is set by the properties of the optokinetic feedback system. It is suggested that the main function of scanning is to prevent visual adaptation.I thank the members of the Department of Neurobiology for many helpful discussions relating to this study, and Roger Hardie for making the recordings from crab retinula cells, the results of which are referred to in the discussion.     

Static cervical-ocular reflex and optokinetic reflex interaction in rabbits

Summary The effect that tonic eye deviations, induced by angular deviation of the torso, have on the characteristics of optokinetic (OK) nystagmus was studied in rabbits. When the slow component of the OK nystagmus moved in the direction of the tonic eye deviation, the amplitude of the slow and fast components of the nystagmus was decreased and their frequency was increased, whereas when the slow component moved in the opposite direction, the amplitude and the frequency of the nystagmus were not different from those when the head and torso were aligned.Under the influence of neck reflexes, the total range of eye movements was double that when the torso was aligned with the head. The place in the orbit where the fast-component is initiated — the so-called fast-component threshold — was deviated in the direction of the neck-reflex-induced tonic eye deviation. The characteristics of the fast component, however, except for its amplitude, were not affected by the change of location of the fast-component threshold.These data indicate that the OK reflex function, as judged by measurement of the slow component velocity, is not affected by neck-vestibular reflexes. They also show that the fast-component threshold is dependent on parameters other than the actual orbital position and that there must be an internal representation of the range of possible eye movements within the brain to regulate the production of fast components.Abbreviations OK optokinetic - CW clockwise - CCW counterclockwise - CNS central nervous system This work was supported by grants NS07059, NS09823, and NS08335 from the National Institutes of Health     

Intracellular responses from cells of the medulla of the fly,Calliphora erythrocephala

1. Intracellular recordings were made from cells in medullae of immobilized, intact flies Calliphora erythrocephala. Stimuli were moving gratings or small spots projected onto translucent hemispheres before the fly.—2. Responses to stationary flashes included tonic and phasic slow potentials only. Sustaining and On/Off discharges were recorded from cells silent in the dark. Sustaining, dimming, On/Off, +On-Off, and-On/-Off discharges were recorded from cells spontaneous in the dark (Fig. 1, 2, and 3).—3. Some cells were relatively sensitive to 3 log unit changes in flash intensities; others were insensitive (Fig. 4).—4. Receptive fields of a few cells tested were small-field ipsilateral monocular, large-field ipsilateral monocular, or large-field binocular.—5. A number of types of nondirectional cells were found. Some gave stronger discharges to movement than to stationary flashes (Fig. 5).—6. Directionally-selective cells were generally spontaneous. Some simply fired faster in the preferred direction. Others (Fig. 6) had inhibition in the null diriction with or without hyperpolarizations.—7. Possibly-new nondirectional cells were found that were inhibited by changes of direction of movement (Fig. 7)—8. A number of cells were stained with Procion yellow, using high voltage pulses. Double stainings sometimes occurred (Fig. 8). Present Address: Psychological Laboratory, Downing Street, Cambridge CB2 3EB, U.K.     

The interstitial nucleus of Cajal of the cat. II. Effects of kainic acid lesion on vertical optokinetic nystagmus and after-nystagmus

The effects of bilateral lesions of the interstitial nucleus of Cajal (INC) by ka?nic acid on vertical optokinetic nystagmus (OKN) and after-nystagmus (OKAN) were studied in four cats: in three cats, in the acute stage from 1 to 60 days after the lesions; in the fourth cat, they were studied 3 years after the lesions were made. Histological control of lesions showed that the whole INC was bilaterally destroyed in two cats of the acute group and only the upper part of INC in the third cat. In the chronic cat, the density of cell bodies in both INC was lower than normal. In the acute group, the cats exhibited a spontaneous downward eye drift in light and in darkness. During an upward optokinetic stimulation, the effect of INC lesions was dramatic: upward slow phases and downward quick phases of OKN were abolished. Sixty days post lesions, small upward slow eye movements were again observed. During a downward optokinetic stimulation, the defect was much less; in particular, after a slight impairment of downward slow phases, during the first days post lesions, they recovered quickly. The secondary optokinetic after nystagmus (OKAN II) ensuing a downward OKN was cancelled and did not reappear 60 days post lesions. In the chronic stage, three years after the lesions, during an upward optokinetic stimulation, the cat showed upward slow phases with velocities close to normal. However, upward slow phases were curved: the velocity at the end of the slow phases was lower than at the beginning. After an upward OKN (the direction of slow phases gives the direction of the OKN and OKAN), the ensuing OKAN was present but abnormal.     

Body movements and retinal pattern displacements while approaching a stationary object in the walking fly,Calliphora erythrocephala

When a walking fly approaches a stationary object two types of body movements are distinguishable. Type I body movements are characterized by low frequencies (0.4–1.3 Hz) and large amplitudes (28–65°). Superimposed on these movements are type II body movements which are characterized by high frequencies (7.3–10.6 Hz) and small amplitudes (5.9–8.2°) (Figs. 3–6; Table 1). Type II movements occur no matter whether the fly is fixating a pattern or orientating itself in homogeneous surroundings without any pattern. In contrast, only 72% of the flies with immobilized heads and 62% of the flies with movable heads make type I body movements. The amplitude of type I and type II body movements increases slightly after immobilization of the head. Binocular as well as monocular pattern projection occurs for the whole walking trajectory (Fig. 7–9). Monocular pattern projection seems to be more frequent in flies with immobilized heads than in those with movable heads. The degree of pattern fluctuations in the visual field of the flies increases slightly along the walking trajectory. Near the starting point in the centre of the arena it amounts to 5–7°, while at the end of the walking trajectory it amounts to 8–10° (Table 2). The following conclusions and hypothesis can be drawn from these experiments. 1. The graph BT for the direction of the fly's logitudinal axis can be approximated by the first derivative of the walking trajectory WT, that means, dWT(x)/dxBT(x) (Fig. 11). 2. The amplitudes of type II body movements are caused by the alternating movements of the legs during forward motion, while type I body movements are classified as exploring movements. During evolution of visually guided behaviour it is possible that blowflies have adapted their elementary movement detector system to type II body movements. 3. The types of pattern projection into the visual field of the fly while approaching an object can be explained by a simple neuronal network characterized by either inhibitory and/or excitatory influences of the visually activated neurones on the motor neurones generating the propulsive forces, that means the forward motion. In addition it is postulated that the large frontal and antero-lateral receptive fields of these neurones are not coupled with the motor centres on the same side of the body (Fig. 12).     

Organization of a complex movement: fixed and variable components of the cockroach escape behavior

The escape behavior of the cockroach Periplaneta americana was studied by means of high speed filming (250 frames/s) and a computer-graphical analysis of the body and leg movements. The results are as follows: 1. The behavior begins with pure rotation of the body about the posteriorly located cerci, followed by rotation plus forward translation, and finally pure translation (Figs. 1, 2). 2. A consistent inter-leg coordination is used for the entire duration of the turn (Fig. 3A). At the start of the movement, five or all six legs execute their first stance phase (i.e. leg on the ground during locomotion) simultaneously. By the end of the turn the pattern has changed to the alternate 'tripod' coordination characteristic of insect walking. The change-over from all legs working together, to working alternately, occurs by means of a consistent pattern of delays in the stepping of certain legs. 3. The movements made by each leg during its initial stance phase are carried out using consistent movement components in the anterior-posterior (A-P) and the medial-lateral (M-L) axes (Fig. 4A). The movement at a particular joint in each middle leg is found to be diagnostic for the direction of turn. 4. The size and direction of a given leg's M-L movement in its initial stance phase depends on the same leg's prior A-P position (Fig. 5). No such feedback effects were seen among different legs. 5. Animals that are fixed to a slick surface on which they make slipping leg movements show the same inter-leg coordination (Fig. 3B), direction of initial stance movement (Fig. 4B) and dependence of the leg's initial M-L movement on its prior A-P position (Fig. 6), as did free-ranging animals. 6. Cockroaches that are walking at the moment they begin their escape reverse those ongoing leg movements that are contrary to escape movements. 7. These results are discussed in terms of the overall coordination of the complex movements, and in terms of the known properties of the neural circuitry for escape. Possibilities for neurobiological follow-up of certain of the findings presented here are also addressed.     

The wind-orientation of walking carrion beetles

The wind-orientation of carrion beetles (Necrophorus humator F.) was studied by use of a locomotion-compensator.

Movement learning of free flying honeybees

Summary Free flying honeybees were conditioned to moving black and white stripe patterns. Bees learn rapidly to distinguish the direction of movement in the vertical and horizontal plane.After being trained to a moving pattern bees do not discriminate the moving alternative from a stationary one. There is no significant velocity discrimination for patterns moving in the same direction.For vertical movements there are clear asymmetries in the spontaneous choice preference and in the learning curves for patterns moving upward or downward.After bees are trained to a stationary pattern they can discriminate it from an upward moving alternative. Learning curves involving movement are generally biphasic, suggesting different adaptive systems depending on the number of rewards.The flight pattern of bees which are trained to movement changes during the process of learning. At the beginning of the learning procedure bees reveal an optokinetic response to the moving patterns, this response is strongly reduced after a number of rewards on a moving pattern.     

fNIRS Exhibits Weak Tuning to Hand Movement Direction

Functional near-infrared spectroscopy (fNIRS) has become an established tool to investigate brain function and is, due to its portability and resistance to electromagnetic noise, an interesting modality for brain-machine interfaces (BMIs). BMIs have been successfully realized using the decoding of movement kinematics from intra-cortical recordings in monkey and human. Recently, it has been shown that hemodynamic brain responses as measured by fMRI are modulated by the direction of hand movements. However, quantitative data on the decoding of movement direction from hemodynamic responses is still lacking and it remains unclear whether this can be achieved with fNIRS, which records signals at a lower spatial resolution but with the advantage of being portable. Here, we recorded brain activity with fNIRS above different cortical areas while subjects performed hand movements in two different directions. We found that hemodynamic signals in contralateral sensorimotor areas vary with the direction of movements, though only weakly. Using these signals, movement direction could be inferred on a single-trial basis with an accuracy of ∼65% on average across subjects. The temporal evolution of decoding accuracy resembled that of typical hemodynamic responses observed in motor experiments. Simultaneous recordings with a head tracking system showed that head movements, at least up to some extent, do not influence the decoding of fNIRS signals. Due to the low accuracy, fNIRS is not a viable alternative for BMIs utilizing decoding of movement direction. However, due to its relative resistance to head movements, it is promising for studies investigating brain activity during motor experiments.     

Trained Eyes: Experience Promotes Adaptive Gaze Control in Dynamic and Uncertain Visual Environments

Current eye-tracking research suggests that our eyes make anticipatory movements to a location that is relevant for a forthcoming task. Moreover, there is evidence to suggest that with more practice anticipatory gaze control can improve. However, these findings are largely limited to situations where participants are actively engaged in a task. We ask: does experience modulate anticipative gaze control while passively observing a visual scene? To tackle this we tested people with varying degrees of experience of tennis, in order to uncover potential associations between experience and eye movement behaviour while they watched tennis videos. The number, size, and accuracy of saccades (rapid eye-movements) made around ‘events,’ which is critical for the scene context (i.e. hit and bounce) were analysed. Overall, we found that experience improved anticipatory eye-movements while watching tennis clips. In general, those with extensive experience showed greater accuracy of saccades to upcoming event locations; this was particularly prevalent for events in the scene that carried high uncertainty (i.e. ball bounces). The results indicate that, even when passively observing, our gaze control system utilizes prior relevant knowledge in order to anticipate upcoming uncertain event locations.     

Elementary detectors for vertical movement in the visual system of Drosophila

Optomotor thrust responses of the fruitfly Drosophila melanogaster to moving gratings have been analysed in order to determine the arrangement of elementary movement detectors in the hexagonal array of the compound eye. These detectors enable the fly to perceive vertical movement. The results indicate that, under photopic stimulation of a lateral equatorial eye region, the movement specific response originates predominantly from two types of elementary movement detectors which connect neighbouring visual elements in the compound eye. One of the detectors is oriented vertically, the other detector deviates 60° towards the anterior-superior direction (Fig. 5b). The maximum of the thrust differences to antagonistic movement is obtained if the pattern is moving vertically or along a superior/anterior — inferior/posterior direction 30° displaced from the vertical (Fig. 3d,e, Fig. 6). Only one of the detectors coincides with one of the two detectors responsible for horizontal movement detection. This indicates that a third movement specific interaction in the compound eye of Drosophila has to be postulated. — The contrast dependence of the thrust response (Fig. 2) yields the acceptance angle of the receptors mediating the response. The result coincides with the acceptance angle found by analysis of the turning response of Drosophila (Heisenberg and Buchner, 1977). This value corresponds to the acceptance angle expected, on the basis of optical considerations, for the receptor system R 1–6. — The movement-specific neuronal network responsible for thrust control is not homogeneous throughout the visual field of Drosophila. Magnitude and preferred direction of the thrust response in the upper frontal part of the visual field seem to vary considerably in different flies (Fig. 6).     

The course control system of beetles walking in an air-current field

The wind-orientated walk of carrion beetles Necrophorus humator F. was analysed under closed-loop conditions with a walking compensator and under openloop conditions with a paired tread wheel (Fig. 1).