Visually elicited head rotation in pigeons (Columba livia).

Horizontal rotational head movements were video-taped from pigeons standing freely in a rotating cylinder. The cylinder carried vertically striped patterns approximating a sinusoidally modulated horizontal intensity distribution. We altered systematically various stimulus parameters: spatial wavelength and contrast of the pattern, angular velocity of the pattern motion and mode of motion onset. We found: (1) both gradual acceleration of the patterned cylinder as well as immediate onset of pattern motion elicit the sequence of smooth following and saccadic resetting movement typical of the rotational "stare" head nystagmus; (2) in experiments with rapid onset of pattern motion, velocity of the smooth following response gradually increases to its steady-state level over a period of about 10 sec; (3) the saccadic head rotations are not stereotyped: larger and shorter saccades follow in an irregular sequence, saccadic velocity and average size varies with stimulus conditions; (4) in the range of 0.9-95 deg/sec, the velocity of the following phase increases in parallel with stimulus speed; (5) in the range of spatial wavelengths of the striped patterns from 6 to 45 deg, at a given drum velocity, patterns of short wavelengths elicit optokinetic head rotations with higher gain (head velocity/drum velocity) than patterns of long wavelengths; (6) response velocity increases with pattern contrast (Michaelson contrast 5, 32 and 75%), following approximately a logarithmic relation; (7) our results on rotational optokinetic head movements support the notion that the neural mechanism underlying motion detection operates like a correlation mechanism.

Binocular interaction in the optokinetic system of the crab Carcinus maenas (L.): optokinetic gain modified by bilateral image flow.

We recorded optokinetic eye movements of the crab, Carcinus maenas, in split-drum experiments. The patterns were either oscillated in antiphase on both sides mimicking translational image flow or they were oscillated in phase producing rotational image flow. Eye movements elicited by the rotational stimulus were larger than those produced by the pseudotranslational pattern movements. The smaller response to the latter is mainly a consequence of binocular interaction, the strength of which depends on both the phase-shift and amplitude of pattern oscillation. We develop two hypotheses to explain our results: either (1) signals from each eye modify the gain of the linkage signals coming from the other eye, or (2) the signals coming from the other eye modify the gain of the control loop itself. Quantitative evaluation of the data favors the second of these two hypotheses, which comprises the models of Barnes and Horridge (1969) and Nalbach et al. (1985). In addition, we found that it is the signals from the two slow channels of the crab's movement-detecting system that are transferred from one eye to the other, while signals of the fastest channel act almost exclusively ipsilaterally. We discuss our results as an adaptation by which an animal with panoramic vision compensates exclusively the rotational component of image flow during locomotion. The fact that freely walking crabs distinguish the two components of image flow better than restrained crabs indicates that further visual and nonvisual signals help to disentangle image flow.

Interactions of local movement detectors enhance the detection of rotation. Optokinetic experiments with the rock crab, Pachygrapsus marmoratus.

Walking crabs move their eyes to compensate for retinal image motion only during rotation and not during translation, even when both components are superimposed. We tested in the rock crab, Pachygrapsus marmoratus, whether this ability to decompose optic flow may arise from topographical interactions of local movement detectors. We recorded the optokinetic eye movements of the rock crab in a sinusoidally oscillating drum which carried two 10-deg wide black vertical stripes. Their azimuthal separation varied from 20 to 180 deg, and each two-stripe configuration was presented at different azimuthal positions around the crab. In general, the responses are the stronger the more widely the stripes are separated. Furthermore, the response amplitude depends also strongly on the azimuthal positions of the stripes. We propose a model with excitatory interactions between pairs of movement detectors that quantitatively accounts for the enhanced optokinetic responses to widely separated textured patches in the visual field that move in phase. The interactions take place both within one eye and, predominantly, between both eyes. We conclude that these interactions aid in the detection of rotation.

Translational head movements of pigeons in response to a rotating pattern: characteristics and tool to analyse mechanisms underlying detection of rotational and translational optical flow.

Pigeons freely standing in the centre of a two-dimensionally textured cylinder not only rotate but also laterally translate their head in response to the pattern sinusoidally oscillating or unidirectionally rotating around their vertical axis. The translational head movement dominates the response at high oscillation frequencies, whereas in a unidirectionally rotating drum head translation declines at about the same rate as the rotational response increases. It is suggested that this is a consequence of charging the 'velocity storage' in the vestibulo-ocular system. Similar to the rotational head movement (opto-collic reflex), the translational head movement is elicited via a wide-field motion sensitive system. The underlying mechanism can be described as vector integration of movement vectors tangential to the pattern rotation. Stimulation of the frontal visual field elicits largest translational responses while rotational responses can be elicited equally well from any azimuthal position of a moving pattern. Experiments where most of the pattern is occluded by a screen and the pigeon is allowed to view the stimulus through one or two windows demonstrate a short-range inhibition and long-range excitation between movement detectors that feed into the rotational system. Furthermore, the results obtained from such types of experiments suggest that the rotational system inhibits the translational system. These mechanisms may help the pigeon to decompose image flow into its translational and rotational components. Because of their translational response to a rotational stimulus, it is concluded, however, that pigeons either generally cannot perfectly perform the task or they need further visual information, like differential image motion, that was not available to them in the paradigms.

The pigeon's eye viewed through an ophthalmoscopic microscope: orientation of retinal landmarks and significance of eye movements.

The retina of live, anaesthetized pigeons was inspected with an ophthalmoscopic microscope mounted on a goniometer. Retinal landmarks (optic axis, pecten, fovea, border between the yellow and red field) and the ora terminalis were projected into the visual field of the eye and related to existing data. The resting position of the eye is determined by an orientation of the pecten 45 degrees to the horizontal plane and the optic axis pointing to the horizon with an azimuth angle of 70 degrees relative to the bill. The binocular overlap is maximal (approximately 30 degrees) some 15 degrees above the eye-bill axis. In the resting position of the eye the red field is directed to the lower frontal visual field with only marginal binocular overlap. Binocular overlap of the area dorsalis with the red field, however, during frontal fixation is brought about by eye movements in the range we have demonstrated. The fixation point is 10 degrees below the eye-bill axis.