Abstract
In the compound eyes of the fruitflyDrosophila, the dioptric system of each ommatidium is able to form virtual images of the receptor terminals (rhabdomere tips) throughout the whole depth of the eye. It is shown (§ 3) that 3 characteristic superposition phenomena occur for images formed by distinct ommatidia (Figs. 3b and 5). The most remarkable superposition appears at the point where the optical axes of all ommatidia converge (center of curvature of the eye). At this level, highly magnified virtual and erect images of corresponding rhabdomeres are superimposed, giving rise to adeep pseudopupil (Fig. 9). Since in the ommatidia ofDrosophila the rhabdome shows a pattern of 7 distal endings (Fig. 8a), the resultingdeep pseudopupil consists of 7 light spots with a similar pattern (Figs. 8b, 7, 11). Conversely thedeep pseudopupil of compound eyes which have fused rhabdomes consists of a single light spot (Fig. 19). Such pseudopupils can be best observed either with antidromic or with orthodromic illumination of the eye, according to the specific transmission or reflection properties of the rhabdomes.
Thedeep pseudopupil of Dipterans is not to be confused with thecorneal pseudopupil (Fig. 13 a) and especially not with thereduced corneal pseudopupil observed with a reduced aperture of the microscope (Fig. 13 b), in spite of the remarkable similarity of these phenomena regarding the asymmetry and the dimension of their pattern (comp. Figs. 7 and 13b). Thereduced corneal pseudopupil consists of 7 facets whereas thedeep pseudopupil consists of 7 virtual images of the receptor endings.
From the results of Kirschfeld (1967), the appearance of areduced corneal pseudopupil like Fig. 13 b on the eye ofDrosophila proves that 7 receptors located in 7 neighbouring ommatidia look in the same direction in space (Fig. 14). The existence of such an optical arrangement favors the view that the eye ofDrosophila, like that ofMusca, belongs to the “neural superposition type”.
A comparative study between thedeep pseudopupil and thereduced corneal pseudopupil leads to the following geometric relation, which is specific of theDrosophila eye and probably of all compound eyes of the “neural superposition type”:
De=Rf′,
, whereD is the diameter of a facet,e the distance between the centers of two neighbouring rhabdomere endings,R the radius of curvature of the eye, andf′ the focal length (in air) of a corneal lens.
Other types of pseudopupils, commonly appearing as dark spots in compound eyes, are explained on a basis similar to thedeep pseudopupil of Drosophila (§5). In fact, the dioptric system of an ommatidium can give virtual images not only of its distal receptor endings but of the whole intensity distribution (i.e. the whole “luminous structure”) which is present in its internal focal plane. If this structure is simple, thedeep pseudopupil, resulting from superpositions of virtual images, is likewise simple (Figs. 16 and 17). If the “luminous structure” is complex, as for example in the eye of the butterflyVanessa (Fig. 18a schematized in Fig. 18c), then thedeep pseudopupil shows the same complexity (Fig. 18 b and d).
In compound eyes which lack screening pigment between their crystalline cones, one can seesecondary pupils of the 1st and 2nd order as described by Exner. Again they may be explained by superpositions of virtual images in the depth of the eye, according to Fig. 20. Moreover, thedeep pseudopupil of the “optical superposition eye” may be due to the fact that the more distal converging system of an ommatidium forms virtual images not of the rhabdome endings themselves but of real images of these endings (Fig. 21).
Although the phenomenon of thedeep pseudopupil is not perceived by the animal, it is of interest for the experimenter who can use it: 1) to study the light receptors easily in the eye of live and intact animals, 2) to measure the physiological divergence angle between adjoining ommatidia, 3) to study the movement of the visual axis and the retinomotor adaptation of the receptors, and 4) to stimulate simultaneously manycorresponding receptors belonging to different ommatidia. The advantages of thisin vivo technique are discussed in § 6.3.
