Noradrenergic modulation of functional selectivity in the cat visual cortex: an in vivo extracellular and intracellular study

In vitro intracellular studies have shown that norepinephrine modulates cellular excitability and synaptic transmission in the cortex. Based on these effects, norepinephrine has been proposed to enhance the signal-to-noise ratio and to improve functional selectivity by potentiating strong synaptic responses and reducing weak ones. Here we have studied the functional effects of iontophoretic applications of norepinephrine during in vivo extracellular and intracellular recordings from neurons of the primary visual cortex of kittens and adult cats. Analysis of extracellular data concentrated on norepinephrine-induced changes in spontaneous and evoked activities, in signal-to-noise ratio, and in orientation and direction selectivity. Analysis of the intracellular data concentrated on actions of norepinephrine on spike firing accommodation, which has been shown to be reduced by norepinephrine in vitro, and on synaptic responses. Application of norepinephrine resulted in a depression of both spontaneous and evoked spiking activity. However, no systematic change in signal-to-noise ratio was observed. The suppressive effect of norepinephrine was exerted with no significant sharpening of direction or orientation selectivity tuning. The overall reduction in visual activity by norepinephrine affected the orientation tuning curves in a way compatible with a divisive effect, that is a normalization or gain control with no change in tuning width. Norepinephrine applied during intracellular recordings reduced the visually evoked depolarizing potentials whereas no change in the responsiveness of the cell to current-induced depolarizations was observed. In conditions of optimal visual stimulation which produced large depolarizations of several hundreds of milliseconds and sustained repetitive firing comparable to that obtained by direct current injection, we were unable to observe a facilitation of the evoked responses by norepinephrine as it would be expected from the well-documented increase in excitability induced by norepinephrine in vitro. In conclusion, from these results we suggest that norepinephrine released in the primary visual cortex primarily reduces the level of cortical activation by afferent signals, without affecting the cortical functional selectivity nor increasing the signal-to-noise ratio.