A. Arieli et al., COHERENT SPATIOTEMPORAL PATTERNS OF ONGOING ACTIVITY REVEALED BY REAL-TIME OPTICAL IMAGING COUPLED WITH SINGLE-UNIT RECORDING IN THE CAT VISUAL-CORTEX, Journal of neurophysiology, 73(5), 1995, pp. 2072-2093
1. We examined the spatiotemporal organization of ongoing activity in
cat visual areas 17 and 18, in relation to the spontaneous activity of
individual neurons. To search for coherent activity, voltage-sensitiv
e dye signals were correlated with the activity of single neurons by t
he use of spike-triggered averaging. In each recording session an area
of at least 2 X 2 mm of cortex was imaged, with 124 diodes. In additi
on, electrical recordings from two isolated units, the local field pot
ential (LFP) from the same microelectrodes, and the surface electroenc
ephalogram (EEG) were recorded simultaneously. 2. The optical signals
recorded from the dye were similar to the LFP recorded from the same s
ite. Optical signals recorded from different cortical sites exhibited
a different time course. Therefore real-time optical imaging provides
information that is equivalent in many ways to multiple-site LFP recor
dings. 3. The spontaneous firing of single neurons was highly correlat
ed with the optical signals and with the LFP. In 88% of the neurons re
corded during spontaneous activity, a significant correlation was foun
d between the occurrence of a spike and the optical signal recorded in
a large cortical region surrounding the recording site. This result i
ndicates that spontaneous activity of single neurons is not an indepen
dent process but is time locked to the firing or to the synaptic input
s from numerous neurons, all activated in a coherent fashion even with
out a sensory input. 4. For the cases showing correlation with the opt
ical signal, 27-36% of the optical signal during spike occurrence was
directly related to the occurrence of spontaneous spikes in a single n
euron, over an area of 2 X 2 mm. In the same cortical area, 43-55% of
the activity was directly related to the visual stimulus. 5. Surprisin
gly, we found that the amplitude of this coherent ongoing activity, re
corded optically, was often almost as large as the activity evoked by
optimal visual stimulation. The amplitude of the ongoing activity that
was directly and reproducibly related to the spontaneous spikes of a
single neuron was, on average, as high as 54% of the amplitude of the
visually evoked response that was directly related to optimal sensory
stimulation, recorded optically. 6. Coherent activity was detected eve
n at distant cortical sites up to 6 mm apart. 7. The spontaneous activ
ity of adjacent neurons, isolated by the same electrode and sharing th
e same orientation preference, was often correlated with different spa
tiotemporal patterns of coherent activity, suggesting that adjacent ne
urons in the same orientation column can belong to different neuronal
assemblies. 8. The averaged optical signal consisted of several tempor
al components, mostly in the frequency range of 0-14 Hz, as expected f
rom the anesthetized state. It contained only a small amount of fast s
ignals in the 15- to 60-Hz range normally found in alert states. 9. Th
e spatiotemporal patterns of coherent activity in areas 17 and 18 were
explored along the representation of the vertical meridian. The coher
ent activity in both areas was time locked to the spontaneous firing o
f single neurons over distances exceeding 6 mm. This activity changed
in an organized manner as a function of time, but the two cortical reg
ions tended to reach peaks or troughs of coherent activity at differen
t times. 10. Temporal Fourier analysis of the spatiotemporal patterns
imaged in these two visual areas revealed that activity at different c
ortical loci consisted of different frequencies. Within a given functi
onal area, however, the activity tended to contain the same pattern of
frequencies.