A role for hydrolysis of inositol 1,4,5-trisphosphate in terminating the response to inositol 1,4,5-trisphosphate and to a flash of light in Limulus ventral photoreceptors.

Injection of inositol 1,4,5-trisphosphate (Ins 1,4,5-P3) into Limulus ventral photoreceptors produces excitation similar to that produced by light. One process which might contribute to rapid termination of the responses to Ins 1,4,5-P3 and to light is the hydrolysis of Ins 1,4,5-P3 by an InsP3-5-phosphatase to form inositol 1,4-bisphosphate. Inositol 2,4,5-trisphosphate (Ins 2,4,5-P3) is known to be less hydrolysable by the InsP3-5-phosphatase than is Ins 1,4,5-P3. Whereas ventral photoreceptors respond to an injection of Ins 1,4,5-P3 with a single wave of depolarization, the response to Ins 2,4,5-P3 is a burst of waves of depolarization. Our hypothesis is that it is the resistance to hydrolysis by the InsP3-5-phosphatase which accounts for the burst of waves produced by Ins 2,4,5-P3. To test this idea we injected ventral photoreceptors with Ins 1,4,5-P3 in the presence of the non-specific phosphatase inhibitors, vanadate and fluoride, which prolong the response to a flash of light in ventral photoreceptors (D.W. Corson, A. Fein, W.W. Walthall, J. Gen. Physiol. 82 (1983) 659-677). In the presence of fluoride or vanadate the response to Ins 1,4,5-P3 was composed of a burst of waves rather than a single wave of depolarization. We conclude that hydrolysis of Ins 1,4,5-P3 by the InsP3-5-phosphatase plays a role in terminating the ventral photoreceptors response to Ins 1,4,5-P3 and also to light.

Membrane conductances involved in amplification of small signals by sodium channels in photoreceptors of drone honey bee.

1. Voltage signals of about 1 mV evoked in photoreceptors of the drone honey bee by shallow modulation of a background illumination of an intensity useful for behaviour are thought to be amplified by voltage-dependent Na+ channels. To elucidate the roles of the various membrane conductances in this amplification we have studied the effects of the Na+ channel blocker tetrodotoxin (TTX) and various putative K+ channel blockers on the membrane potential, Vm. 2. Superfusion of a slice of retina with 0.5-10 mM-4-aminopyridine (4-AP) depolarized the membrane and, in fifty of sixty-three cells induced repetitive action potentials. Ionophoretic injection of tetraethylammonium produced similar effects. 3. In order to measure the depolarization caused by 4-AP, action potentials were prevented by application of TTX: 4-AP was applied when the membrane was depolarized to different levels by light. 4-AP induced an additional depolarization at all membrane potentials tested (-64 to -27 mV). We conclude that there are 4-AP-sensitive K+ channels that are open at constant voltage over this range. 4. 4-AP slowed down the recovery phase of the action potential induced by a light flash by a factor that ranged from 0.51 to 0.16. This reduction could be accounted for by the reduction in a voltage-independent K+ conductance estimated from the steady-state depolarization. 5. After the voltage-gated Na+ channels had been blocked by TTX, exposure to 4-AP further changed the amplitude of the response to a small (approximately 10%) decremental light stimulus. The change was an increase when the background illumination brought Vm to potentials more negative than about -40 mV; it was a decrease when Vm > -40 mV. The data could be fitted by a circuit representation of the membrane with a light-activated conductance and a K+ conductance (EK = -66 mV) that was partly blocked by 4-AP. The voltage range studied was from -52 to -27 mV; neither conductance in the model was voltage dependent. 6. The responses to small changes in light intensity in the absence of TTX were mimicked by a model. We conclude that a voltage-dependent Na+ conductance described by the Hodgkin-Huxley equations can amplify small voltage changes in a cell membrane that is also capable of generating action potentials; the magnitude of the K+ conductance is critical for optimization of signals while avoiding membrane instability.

A method for estimating the minimum visual stimulus that evokes a behavioural response in the drone, Apis mellifera male.

Drones, attracted to an observation site by a scented lure, left the lure to fly directly towards a small object suspended nearby. An object the size of a queen bee was detected at distances up to 2 m. It is estimated that at this maximum distance the object will have decreased the light incident on a single rhabdom by 6%.