Eye structure and optics in the pelagic shrimp Acetes sibogae (Decapoda, Natantia, Sergestidae) in relation to light-dark adaptation and natural history

Abstract

The structure and optics of the compound eyes of the neritic sergestid shrimp, Acetes sibogae , are described. The eyes are nearly spherical and heavily pigmented. The facets are square, indicating that the eye operates by the recently recognized mechanism of reflecting superposition. The most distal portion of each ommatidium is the corneal lens, which is secreted by two underlying corneagenous cells. These two cells surround the crystalline cone and cone stalk and the four cells of which they are composed and extend proximally at least as far as the distal rhabdom. Near the base of the cone stalk the extensions of the corneagenous cells swell and enclose spheres which bear on their surfaces small particles similar to ribosomes in appearance. Beneath the corneagenous cells lie four crystalline cone cells, parts of which differentiate to form the crystalline cone and cone stalk. The latter structures are compound, one quarter of each being contributed by each crystalline cone cell. Distally the crystalline cone cells send a small projection, which is surrounded by the corneagenous cells, to the cornea. Proximal extensions of each of the four parts of the cone stalk extend between the retinula cells and meet within the basement membrane. Between the base of the cone stalk and the regularly layered rhabdom lies the distal rhabdom. It is surrounded by a cell that we have termed retinula cell eight (R8), by analogy with other crustacean systems, and consists of unordered microvilli projecting from the cell membrane into the extracellular space above the layered rhabdom. In addition to R 8, which contributes only to the distal rhabdom, seven other retinula cells contribute to the proximal rhabdom, which consists of alternating ordered layers of orthogonally arranged microvilli. Four of these retinula cells are arranged orthogonally and extend far distally along the crystalline tract. The other three do not extend as far distally and alternate with the first four in their position around the axis of the ommatidium. R8 is located still further proximally at the level of the distal rhabdom. All seven of the retinula cells which contribute to the proximal rhabdom contain proximal pigment and extend through the basement membrane. The basement membrane consists of a meshwork grid with each intersection supporting a rhabdom so at this point the retinula cell axons project into different squares of the meshwork. Tapetal pigment cells are present in the vicinity of the basement membrane and extend downward to the lamina. The granules of tapetal pigment are covered or exposed by movements of the proximal pigment and also change their intracellular distribution depending on illumination. In addition to the proximal (retinula cell) pigment and the tapetal pigment the eye contains four types of distal pigment. Moving inward from the cornea these are the distal yellow pigment (DYP) which surrounds the entire eye; the distal reflecting pigment (DRP), which forms a thin layer and is continuous with the tapetal pigment at the edge of the eye; and the black distal pigment and the mirror pigment (MP) both contained within distal pigment cells (DPC). In the light-adapted state the proximal pigment moves distally, surrounding the rhabdoms, and the tapetal pigment granules move proximally so that they are mainly found beneath the basement membrane. Movements of the distal pigments are less clearcut, but they all appear to move somewhat proximally in the light-adapted state. Multivesicular bodies are more abundant in the retinula cells shortly after dawn, and are possibly related to membrane turnover. Interommatidial angle, as measured on both fixed and fresh material, varied from 2.8 to 3.8° in different parts of the eye. The crystalline cones were found to have a uniform refractive index radially, which, combined with their square shape, indicates that they function by reflecting superposition. Total internal reflection from the sides of the cones is adequate to explain the maximum diameter of the eyeshine from the dark-adapted eye at night without the need for additional mirrors. Nevertheless, from its organization and appearance the mirror pigment could act as a reflector in the dark-adapted eye. Also, the size of the glow patch indicates that there would be a gain of nearly two log units in image brightness in going from the light-adapted to the dark-adapted state. Each corneal facet was found to act as a weak converging lens, with a focal length of approximately 300 μm. The eye structure of Acetes is discussed in relation to that of other shrimp and to the natural history of Acetes .