The night flying scarabaeid beetle
Anoplognathus
provides an example of a dark-adapted clear-zone compound eye in which rays from a distant point source, entering by a large patch of facets, are imperfectly focused upon the receptor layer. The optical system of the eye was investigated by six methods, all of which give similar results: (1) ray tracing through structures of known refractive index, (2) measurement of visual fields of single receptors, (3) measurement of the divergence of eyeshine, and (4) of the optomotor response to stripes of decreasing width, and (5) by direct observation of distribution of light within the eye. Finally (6) anatomically there is no single plane upon which an image could be focused. In each ommatidium, beneath the thick cornea, with its short corneal cone, lies a non-homogeneous crystalline cone (range of r. i. 1.442-1.365) that is significant in partially focusing rays across the wide clear zone (340 μm) in the dark-adapted eye. On the proximal side of the clear zone the rhabdoms form 7-lobed columns, isolated from each other over half their length by a tracheal tapetum. In the light-adapted eye the cone cells extend to form a crystalline tract (70-90 μm long) which is surrounded by dense pigment, and the optical path across the clear zone is completed by retinula cell columns that are of higher density than the surrounding cells. Pigment movement upon adaptation takes about 10 min to complete. Dark adaptation can be induced only at night on account of a strong diurnal rhythm. Eyeshine can be seen in the dark-adapted eye so long as the distal pigment leaves free the tips of the crystalline cones. Eyeshine falls to 50% at an angle of 12° from the direction of a parallel beam shining on the eye, as is consistent with a partial focus in which the distribution of light on the receptor layer is 18°-24° wide at the 50% contour. This distribution was confirmed by direct examination of the inside of the eye and by measurement of receptor fields as follows. The mean acceptance angle for 13 light-adapted units was 12.57° ± 1.97° s. d. and that of 10 dark-adapted ones 20.3° ± 3.36° s. d. The sensitivity to a point source on axis is increased at least 1000 fold by dark adaptation. Rays traced through a scale drawing of the eye, with refractive index measured for each component, show how the eye as a whole comes to be partially focused, and predicts an acceptance angle of 12° in the light-adapted and 20°-24° in the dark-adapted eye. In optomotor experiments dark-adapted
Anoplognathus
does not respond to stripes narrower than 18° repeat period, but light-adapted beetles respond down to 10°. The optomotor experiments also show a 1000 fold increase in sensitivity when dark-adapted at night. The eye has poor acuity that goes with wide visual fields of its receptors, and this is surprising when other excellently focused clear zone eyes are known. A possible compensation for the poor acuity is that the aperture of the eye can be larger, so that sensitivity although only to large objects, is that much increased.
The Eye Muscle of Calliphora Vomitoria L