BRAIN
RESEARCH
ELSEVIER Brain Research 678 (1995) 177-190
Research report
The spatiotemporal organization of auditory, visual, and auditory-visual
evoked potentials in rat cortex
Daniel S. Barth *, Neil Goldberg, Barbara Brett, Shi Di
Department of Psychology, Universityof Colorado, Boulder, CO 80309-0345, USA
Accepted 24 January 1995
Abstract
Four placements of an 8 x 8 channel microelectrode array were used to map auditory, visual, and combined auditory-visual
evoked potentials (AEP, VEP, AVEP) from a total of 256 electrode sites over a 7 x 7 mm 2 area including most of somatosensory,
auditory, and visual cortex in the right hemisphere of the rat. The unimodal AEP and VEP consisted of an archetypal response
sequence representing a systematic spatial and temporal activation of primary and secondary sensory cortex. Spatiotemporal
analysis of these waveforms indicated that they could be decomposed into a small number of spatial and temporal components;
components that are related to patterns of specific and non-specific thalamocortical projections connecting the auditory and
visual nuclei of the thalamus with primary and secondary auditory and visual cortex. These data suggest that the AEP and VEP
complex are the cortical reflection of asynchronous activation of parallel thalamocortical projection systems. The areal
distribution of the AEP and VEP also overlapped, primarily in secondary auditory and visual cortex, indicating that these regions
contain populations of cells responding to either modality. Polymodal auditory-visual stimulation resulted in unique activation of
two isolated populations of neurons positioned in secondary auditory and secondary visual cortex which were revealed by
difference waveforms, computed by subtracting the sum of the AEP and VEP from the AVEP complex. Retrograde labeling of
the polymodal zones indicated that they receive parallel thalamocortical projections primarily from non-specific auditory and
visual thalamic nuclei including the medial and dorsal divisions of the medial geniculate nucleus (MGm and MGd), the
suprageniculate nucleus (SGN), and the lateral posterior nucleus (LP). The polymodal zone in visual cortex also receives specific
projections from the dorsal division of the lateral geniculate nucleus (LGd). These data conform to a general model of
thalamocortical organization in which specific thalamic nuclei with a high degree of modality specificity make restricted
projections to primary sensory cortex and parts of secondary sensory cortex, and association thalamic nuclei with a high degree of
sensory convergence make more divergent cortical projections. Primary and secondary sensory cortex, as well as distinct zones of
polysensory cortex appear to be activated in tandem via parallel thalamocortical projections. Thus, the cerebral cortex must have
simultaneous access to both unimodal and polymodal sensory information.
Keywords: Auditory; Visual; Polysensory; Evoked potential
1. Introduction
In the anesthetized rat, simple sensory stimuli evoke
consistent spatial and temporal patterns of field poten-
tial waveforms that can be recorded extracranially and
on the surface of the cortex. When recently developed
methods of numerical analysis are applied to epicorti-
cal waveform distributions, the sensory evoked poten-
* Corresponding author. Department of Psychology,Campus Box
345, University of Colorado at Boulder, Boulder, CO 80309-0345,
USA. Fax: (1) (303) 492-2967.
0006-8993/95/$09.50 © 1995 Elsevier Science B.V. All rights reserved
SSDI 0006-8993(95)00182-4
tial complex may be decomposed into a small number
of spatial and temporal components that are directly
related to the functional anatomy of thalamocortical
sensory projection systems [2-4,14,15].
Of particular interest in this recent work is the
observation that epicortical distributions of the evoked
potential complex extend well beyond cytoarchitectoni-
cally defined primary sensory cortex [4] into interposed
secondary sensory cortex with possible associative or
polysensory information processing functions [16].
These results are consistent with earlier studies in cat
and monkey [6-8,32,50,51], and more recent evoked
potential [53] and microelectrode [40] studies in the