A. Ylinen et al., SHARP WAVE-ASSOCIATED HIGH-FREQUENCY OSCILLATION (200-HZ) IN THE INTACT HIPPOCAMPUS - NETWORK AND INTRACELLULAR MECHANISMS, The Journal of neuroscience, 15(1), 1995, pp. 30-46
Sharp wave bursts, induced by a cooperative discharge of CA3 pyramidal
cells, are the most synchronous physiological pattern in the hippocam
pus. In conjunction with sharp wave bursts, CA1 pyramidal cells displa
y a high-frequency (200 Hz) network oscillation (ripple), In the prese
nt study extracellular field and unit activity was recorded simultaneo
usly from 16 closely spaces sites in the awake rat and the intracellul
ar activity of CA1 pyramidal cells during the network oscillation was
studied under anesthesia, Current source density analysis of the high-
frequency oscillation revealed circumscribed sinks and sources in the
vicinity of the pyramidal layer, Single pyramidal cells discharged at
a low frequency but were phase locked to the negative peak of the loca
lly derived field oscillation, Approximately 10% of the simultaneously
recorded pyramidal cells fired during a given oscillatory event. Puta
tive interneurons increased their discharge rates during the field rip
ples severalfold and often maintained a 200 Hz frequency during the os
cillatory event, Under urethane and ketamine anesthesia the frequency
of ripples was slower(100-120 Hz) than in the awake rat(180-200 Hz), H
alothane anesthesia prevented the occurrence of high-frequency field o
scillations in the CA1 region. Both the amplitude(1-4 mV) and phase of
the intracellular ripple, but not its frequency, were voltage depende
nt, The amplitude of intracellular ripple was smallest between -70 and
-80 mV, The phase of intracellular oscillation relative to the extrac
ellular ripple reversed when the membrane was hyperpolarized more than
-80 mV, A histologically verified CA1 basket cell increased its firin
g rate during the network oscillation and discharged at the frequency
of the extracellular ripple. These findings indicate that the intracel
lularly recorded fast oscillatory rhythm is not solely dependent on me
mbrane currents intrinsic to the CA1 pyramidal cells but it is a netwo
rk driven phenomenon dependent upon the participation of inhibitory in
terneurons. We hypothesize that fast field oscillation (200 Hz) in the
CA1 region reflects summed IPSPs in pyramidal cells as a result of hi
gh-frequency barrage of interneurons. The sharp wave associated synchr
onous discharge of pyramidal cells in the millisecond range can exert
a powerful influence on retrohippocampal targets and may facilitate th
e transfer of transiently stored memory traces from the hippocampus to
the entorhinal cortex.