Massive diurnally modulated photoreceptor membrane turnover in crab light and dark adaptation

Abstract

1. Effects of light and dark adaptation on the photoreceptor membranes of the rock crabGrapsus have been quantitatively studied using light and electron microscopy to document changes in rhabdom size and their fine structural basis. Some comparative data were also obtained for the ghost crabOcypode. Animals in the laboratory were maintained on a daily light (L)-dark (D)cycle 12L∶12D. Periods of adaptation ranged from 3–28 h (Table 1) and light adaptation was effected at one moderate intensity equal to the light period in the daily sequence. 2. Massive rapid reversible changes in rhabdom size (Fig. 1) comprising mainly diameter modulation (Figs. 2, 3) and some alteration in length, normally resulted in maximum increases in receptor organelle volume of about 20× between a thinner, shorter fully light adapted condition (2×290 μm) to the fully dark adapted state (7.9× 336 μm). 3. Rhabdom growth during dark adaptation (Table 2) was due mainly (Fig. 8) to microvillus elongation (+154%) as well as substantial increase (+117%) in the number of microvilli present in cross sections (Figs. 2, 3, 10); microvillus diameter (Figs. 4, 5, 9) also increased (+14%) but accounts for only a minor part of the overall change commensurate with observed differences in rhabdom length (+16%). Calculation shows that from fully light adapted to fully dark adapted state the area of photoreceptor membrane increased nearly 20× in a few hours. 4. Although duration of adaptation affected its amplitude, six hours of either light or darkness evoked the maximum changes documented; shorter and longer periods had less effect (Figs. 6, 7). 5. Time of day had a strong periodic influence on the amplitude of membrane adaptation produced by a given exposure to dark or light. Thus maximum dark adaptation occurred with 6 h of darkness terminating at midnight and maximum light adaptation with 6 h of light ending at noon of the diurnal light cycle (Fig. 11). 6. Direct effects of light and dark on synthesis and degradation of photoreceptor membrane must therefore be superimposed on indirect or direct cyclic neuroendocrine control presumably coupled to an entrained circadian oscillator.