Electron Microscope Observations on Form Changes in Photoreceptor Outer Segments and Their Saccules in Response to Osmotic Stress

Isolated retinas of rats or frogs were incubated in various salt or sucrose solutions over a wide range of osmolarities and then fixed in 1% osmium tetroxide solutions of matching osmolarities. Light and electron microscope observations were concentrated on the outer segments of the intact photoreceptors. These became globular and of increasing size with increasing hypoosmolarity and irregularly linear and condensed in hyperosmotic solutions. Isoosmotic incubations of rat retinas in solutions containing potassium as the only cation also produced swelling of the outer segments when chloride or acetate was present; but swelling was less when the cation was sodium and it was not seen with either cation when the anion was methylsulfate. The effects of various metabolic and membrane poisons are also reported. The behavior of the saccules within the outer segments was equivocal. While there was a tendency toward more saccules with wider lumina with hypoosmolarity, most of the saccules were not swollen. Surprisingly, intrasaccular space was consistently enlarged in rat retinas exposed to hyperosmotic sucrose but not to salt. The saccules or their derivatives within swollen outer segments tend to maintain their intersaccule spacing and approximation to the cell membrane. It was also noted that the ciliary connectives resist swelling and that retinal Müller cells swell readily.