1. To study the encoding of input currents into output spike trains by
regular-spiking cells, we recorded intracellularly from slices of the
guinea pig visual cortex while injecting step, sinusoidal, and broadb
and noise currents. 2. When measured with sinusoidal currents, the fre
quency tuning of the spike responses was markedly band-pass. The prefe
rred frequency was between 8 and 30 Hz, and grew with stimulus amplitu
de and mean intensity. 3. Stimulation with broadband noise currents dr
amatically enhanced the gain of the spike responses at low and high fr
equencies, yielding an essentially flat frequency tuning between 0.1 a
nd 130 Hz. 4. The averaged spike responses to sinusoidal currents exhi
bited two nonlinearities: rectification and spike synchronization. By
contrast, no nonlinearity was evident in the averaged responses to bro
adband noise stimuli. 5. These properties of the spike responses were
not present in the membrane potential responses. The latter were rough
ly linear, and their frequency tuning was low-pass and well fit by a s
ingle-compartment passive model of the cell membrane composed of a res
istance and a capacitance in parallel (RC circuit). 6. To account for
the spike responses, we used a ''sandwich model'' consisting of a low-
pass linear filter (the RC circuit), a rectification nonlinearity, and
a high-pass linear filter. The model is described by six parameters a
nd predicts analog firing rates rather than discrete spikes. It provid
ed satisfactory fits to the firing rate responses to steps, sinusoids,
and broadband noise currents. 7. The properties of spike encoding are
consistent with temporal nonlinearities of the visual responses in V1
, such as the dependence of response frequency tuning and latency on s
timulus contrast and bandwidth. We speculate that one of the roles of
the high-frequency membrane potential fluctuations observed in vivo co
uld be to amplify and linearize the responses to lower, stimulus-relat
ed frequencies.