Physiol. Res. 60 (Suppl. 1): S93-S99, 2011
Auditory Modulation of the Inferior Temporal Cortex Neurons in Rhesus Monkey P. KAPOSVÁRI1, P. CSIBRI1, G. CSETE1, T. TOMPA1, GY. SÁRY1 1
Department of Physiology, University of Szeged, Szeged, Hungary
Received January 3, 2011 Accepted February 16, 2011 On-line July 19, 2011
Summary We performed a systematic study to check whether neurons in the area TE (the anterior part of inferotemporal cortex) in rhesus monkey, regarded as the last stage of the ventral visual pathway, could be modulated by auditory stimuli. Two fixating rhesus monkeys were presented with visual, auditory or combined audiovisual stimuli while neuronal responses were recorded. We have found that the visually sensitive neurons are also modulated by audiovisual stimuli. This modulation is manifested as the change of response rate. Our results have shown also that the visual neurons were responsive to the sole auditory stimuli. Therefore, the concept of inferotemporal cortex unimodality in information processing should be re-evaluated. Key words Monkey • Inferotemporal cortex • Auditory modulation Corresponding author Gy. Sáry, Dept. of Physiology, University of Szeged, 10. Dóm tér, Szeged,
Hungary.
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part of the inferotemporal cortex (TE). Inferotemporal cortex ITC is considered to be the last unimodal visual stage in the ventral visual stream (Tanaka 1996, Ungerleider and Mishkin 1982), consisting of several subregions, like anterior, middle and posterior part (Tamura and Tanaka 2001). In one study (Iwai et al. 1987), one third of the recorded neuronal population in the ITC responded to pure tones. Another study (Watanabe and Iwai 1991) reported neurons of the posterior part of inferotemporal region (TEO) reacting to auditory signals, while others obtained similar results in ITC in split-brain monkeys (Ringo and O'Neill 1993). Baylis and his co-workers reported neurons receiving auditory signals in TS and TAa areas of the temporal cortex (Baylis et al. 1987). However, these studies might have found attention or action dependent modulated responses. In this study, we tested whether the neurons of the TE area of inferotemporal cortex, responsive to visual stimulation, could react to sole auditory stimulation.
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Material and Methods
Introduction
Subjects
It is more and more evident that we cannot regard the brain as made up of independent unimodal streams of information joining only at the association areas (Budinger et al. 2006, Schiller 1996). We also know from everyday experience and from psychophysical studies how intricately visual and auditory information acts together in shaping our knowledge of our surroundings and our acting in them. Thus, we found it advisable to study the effects of auditory stimulation in that area regarded par excellence unimodal, the anterior
Two adult Rhesus monkeys (monkey Ch, a male and monkey Z, a female; 6.8 kg and 6 kg, respectively) participated in the study. The monkeys worked under a controlled water access paradigm in daily recording sessions, 5 days a week. During weekend, they were allowed to drink water ad libitum and fruits and vegetables were added to their diet. All procedures conformed to the guidelines of the National Institutes of Health Guide for the Care and Use of Laboratory Animals and the guidelines of the University of Szeged Animal Care and Use Committee.
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Surgery Prior to the recordings, the monkeys underwent two aseptic surgeries (Sáry et al. 2008) performed under general anesthesia to receive a headpost and recording chamber over the ITC on the skull, and a scleral search coil in the eye (Judge et al. 1980). Anesthesia was induced with Calypsol (Ketamine, 20 mg/kg i.m.) and maintained with 1.5 % Halothane in a 2:1 mixture of N2O and O2. Atropine (0.05 mg/kg i.m.), was injected to reduce bronchial secretion and salivation. Body temperature, heart rate, respiratory rate, end-tidal CO2 and peripheral O2 saturation were monitored throughout the surgery. Analgesic was given after surgery (Nalbuphine, 0.15 mg/kg) for 5 days. Stimuli and recordings Colorful, complex images (Sary et al. 2004) were shown on a display (Philips Brilliance 17A having 17’ diameter; refresh rate, resolution and viewing distance were 74 Hz, 800x600 pixels and 57 cm, respectively). Visual stimuli had a viewing angle of 5x6 degs, and mean luminance was 7.9 cd/m2. Auditory stimuli were presented from a computer loudspeaker positioned on top of the stimulation display (app. 60 dB, 440, 466, 493, 523, 554, 587, 622, 659, 698, 739, 783, 830, 880, 932, 987, 1046, 1108, 1174, 1244 and 1318 Hz, respectively). Three stimulation conditions were used: one unimodal condition contained only the visual images (VIS); the other unimodal condition had only the auditory stimulus (AUDIO). For bimodal stimulation (condition: AUDIOVIS) the visual images were paired with one of the auditory stimuli. This pairing was fixed, i.e., an image was always presented with the same sound. A screen with a grey background, without the sound served as control (see stimulus sequence). Standard electrophysiological equipment was used (FHC inc., Bowdoin, ME, USA) with electrodes having impedance values of 1-3 MΩ (FHC). During the recording sessions, the monkeys sat with their heads fixed. A custom-made software running on a PC recorded the eye movements (200 Hz sampling rate), delivered the reward, controlled the animals' behavior, and presented the stimuli. Another PC collected the electrophysiological data. The background luminance in the sound-proof experimental room was kept constant at a level
