Heterogenic components of a fast electrical potential in Drosophila compound eye and their relation to visual pigment photoconversion.

The electroretinogram of the dipteran compound eye in response to an intense flash contains an early, diphasic potential that has been termed the M potential. Both phases of the M potential arise from the photostimulation of metarhodopsin. The early, corneal-negative component, the M1, can be recorded intracellularly in the photoreceptors and has properties similar to the classical early receptor potential (ERP). The M1 is resistant to cold, anaesthesia, and anoxia and has no detectable latency. It depends on flash intensity and metarhodopsin fraction in the manner predicted for a closed, two-state pigment system, and its saturation is shown to correspond to the establishment of a photoequilibrium in the visual pigment. On the other hand, the dominant, corneal-positive component, the M2, does not behave like an ERP. It arises, not in the photoreceptors, but deeper in the retina at the level of the lamina, and resembles the on-transient of the electroretinogram in its reversal depth and sensitivity to cooling or CO2. The on-transient, which is present over a much wider range of stimulus intensity than the M potential, has been shown to arise from neurons in the lamina ganglionaris. Visual mutants in which the on-transient is absent or late are also defective in the M2. It is proposed that the M2 and the on-transient arise from the same or similar groups of second-order neurons, and that the M2 is a fast laminar response to the depolarizing M1 in the photoreceptors, just as the on-transient is a fast laminar response to the depolarizing late receptor potential. Unlike the M1, the M2 is not generally proportional to the amount of metarhodopsin photoconverted, and the M2 amplitude is influenced by factors, such as a steady depolarization of the photoreceptor, which do not affect the M1.

Responses of Barnacle Photoreceptors to High Energy Flashes of Short Duration

In chromatic adapted barnacle median and lateral photo­receptors the two stable states of the photopigment (rhodopsin R and metarhodopsin M) were interconverted with intense, colored light flashes of 1 ms duration. Only after conversion of the red adapted photoreceptor in K+-Ringer solution with an intense flash the negative early receptor potential, ERP (of R) gradually appeared detected with an indicator flash. For the opposite conversion (blue adapted, R→M) the gradual appearence of the positive ERP (M) was not measurable in the same time span. In artificial seawater all flash stimuli yielded - irrespective of color - the transient component of the late receptor potential (LRP). ERP results for the lateral photoreceptor are dis­cussed in view of an existing kinetic model and an attempt is made to give an explanation which covers the new LRP transient and ERP results for both types of photoreceptor (appendix).

Alteration of sensitivity and time scale in invertebrate photoreceptors exposed to anoxia, dinitrophenol, and carbon dioxide.

The effects of anoxia, 2,4-dinitrophenol (DNP), and carbon dioxide (CO2) on the late receptor potential of Balanus lateral ocelli, Limulus ventral eyes, and the retinular cells of Linulus lateral eyes have been studied. Either anoxia, DNP, or exposure to 100% CO2 causes a depolarization of 5-30 mV and a gradual reduction and eventually abolition of the late receptor potential and an increase in the latency and time to peak of the response. This lengthening of the time scale is in contrast to the response obtained in photoreceptors that have been light-adapted or injected with calcium. In that case a loss in sensitivity is associated with a decrease in latency and time to peak. Because of these observed differences, the effects of metabolic inhibition cannot be attributed merely to a loss in regulation of intracellular free calcium. Rather, because alteration of intracellular pH (pHi) by using either (NH4)2SO4 or CO2 produced changes in the photoresponse similar to those caused by metabolic inhibition, it is suggested that changes in pHi during metabolic inhibition can account in part for the lengthening of the time scale. In addition to the changes in pHi and internal Ca++ concentration due to metabolic inhibition, the possible role of other consequences of metabolism in the transduction mechanism is also discussed.

Lability of the prolonged depolarizing afterpotential in Balanus photoreceptors.

A large cell to cell variability of the prolonged depolarizing afterpotential (PDA) decay time constant (tau) has been measured in Balanus eberneus lateral ocelli. While 25% of the cells had PDA's of long duration, tau greater than 10 min. 45% of the cells tested showed either weak (tau less than 60 s) PDA or none at all. The variability was not reflected in the late receptor potential. All the cells showed normal light-coincident responses. The variability was not due to some alteration of the thermal stability of the pigment states, since after monochromatic adaptation the amplitude of the early receptor potential remained unchanged for at least 30 min. In addition, in some cells that initially showed PDAs of long duration, the decay time was either shortened or abolished after exposure to anoxia. Again, the late receptor potential and the stability of the pigments remained unaffected. These results indicate that the mechanisms which give rise to the PDA are not always tightly coupled to the direct chain of events that lead to the light-coincident response.