BIOELECTRIC POTENTIALS IN HALICYSTIS : VIII. THE EFFECTS OF LIGHT.
The effects of light upon the potential difference across the protoplasm of impaled Halicystis cells are described. These effects are very slight upon the normal P.D., increasing it 3 or 4 per cent, or at most 10 per cent, with a characteristic cusped time course, and a corresponding decrease on darkening. Light effects become much greater when the P.D. has been decreased by low O(2) content of the sea water; light restores the P.D. in much the same time course as aeration, and doubtless acts by the photosynthetic production of O(2). There are in both cases anomalous cusps which decrease the P.D. before it rises. Short light exposures may give only this anomaly. Over part of the potential range the light effects are dependent upon intensity. Increased CO(2) content of the sea water likewise depresses the P.D. in the dark, and light overcomes this depression if it is not carried too far. Recovery is probably due to photosynthetic consumption of CO(2), unless there is too much present. Again there are anomalous cusps during the first moments of illumination, and these may be the only effect if the P.D. is too low. The presence of ammonium salts in the sea water markedly sensitizes the cells to light. Subthreshold NH(4) concentrations in the dark become effective in the light, and the P.D. reverses to a negative sign on illumination, recovering again in the dark. This is due to increase of pH outside the cell as CO(2) is photosynthetically reduced, with increase of undissociated NH(3) which penetrates the cell. Anomalous cusps which first carry the P.D. in the opposite direction to the later drift are very marked in the presence of ammonia, and may represent an increased acidity which precedes the alkaline drift of photosynthesis. This acid gush seems to be primarily within the protoplasm, persisting when the sea water is buffered. Glass electrode measurements also indicate anomalies in the pH drift. There are contrary cusps on darkening which suggest temporarily increased alkalinity. Even more complex time courses are given by combining low O(2) and NH(4) exposures with light; these may have three or more cusps, with reversal, recovery, and new reversal. The ultimate cause of the light effects is to be found in an alteration of the surface properties by the treatments, which is overcome (low O(2), high CO(2)), or aided (NH(4)) by light. This alteration causes the surface to lose much of its ionic discrimination, and increases its electrical resistance. Tests with various anion substitutions indicate this, with recovery of normal response in the light. A theory of the P.D. in Halicystis is proposed, based on low mobility of the organic anions of the protoplasm, with differences in the two surfaces with respect to these, and the more mobile Na and K. ions.
Blinks
L R
LR
Jacques Loeb Laboratory, Hopkins Marine Station, Pacific Grove, and the School of Biological Sciences, Stanford University.