Research paper Stimulus-dependent activations and attention-related modulations in the auditory cortex: A meta-analysis of fMRI studies Kimmo Alho a, b, * , Teemu Rinne b , Timothy J. Herron c , David L. Woods c, d a Helsinki Collegium for Advanced Studies, University of Helsinki, PO Box 4, FI 00014 Helsinki, Finland b Institute of Behavioural Sciences, University of Helsinki, PO Box 9, FI 00014 Helsinki, Finland c Veterans Affairs Northern California Health Care System (VANCHSC), Research Building R4 VAMC,150 Muir Road, Martinez, CA 94553-4612, USA d Department of Neurology, Center for Neuroscience, and Center for Mind and Brain, University of California at Davis, 267 Cousteau Place, Davis, CA 95618, USA article info Article history: Received 17 May 2013 Received in revised form 22 July 2013 Accepted 1 August 2013 Available online 11 August 2013 abstract We meta-analyzed 115 functional magnetic resonance imaging (fMRI) studies reporting auditory-cortex (AC) coordinates for activations related to active and passive processing of pitch and spatial location of non-speech sounds, as well as to the active and passive speech and voice processing. We aimed at revealing any systematic differences between AC surface locations of these activations by statistically analyzing the activation loci using the open-source Matlab toolbox VAMCA (Visualization and Meta-analysis on Cortical Anatomy). AC activations associated with pitch processing (e.g., active or passive listening to tones with a varying vs. fixed pitch) had median loci in the middle superior temporal gyrus (STG), lateral to Heschl’s gyrus. However, median loci of activations due to the processing of infrequent pitch changes in a tone stream were centered in the STG or planum temporale (PT), significantly posterior to the median loci for other types of pitch processing. Median loci of attention-related modulations due to focused attention to pitch (e.g., attending selectively to low or high tones delivered in concurrent sequences) were, in turn, centered in the STG or superior temporal sulcus (STS), posterior to median loci for passive pitch processing. Activations due to spatial processing were centered in the posterior STG or PT, significantly posterior to pitch processing loci (processing of infrequent pitch changes excluded). In the right-hemisphere AC, the median locus of spatial attention-related modulations was in the STS, significantly inferior to the median locus for passive spatial processing. Activations associated with speech processing and those associated with voice processing had indistinguishable median loci at the border of mid-STG and mid-STS. Median loci of attention-related modulations due to attention to speech were in the same mid-STG/STS region. Thus, while attention to the pitch or location of non-speech sounds seems to recruit AC areas less involved in passive pitch or location processing, focused attention to speech predominantly enhances activations in regions that already respond to human vocalizations during passive listening. This suggests that distinct attention mechanisms might be engaged by attention to speech and attention to more elemental auditory features such as tone pitch or location. This article is part of a Special Issue entitled <Human Auditory Neuroimaging>. Ó 2013 Elsevier B.V. All rights reserved. 1. Introduction Measurement of blood oxygenation level dependent (BOLD) brain responses with functional magnetic resonance imaging (fMRI) allows millimeter-scale localization of human brain activity. Hundreds of fMRI studies have investigated different aspects of auditory processing and meta-analyses have been performed comparing spatial and nonspatial auditory processing (Arnott et al., 2004) and comparing the processing of speech sounds, human vocalizations and other complex sounds (Frühholz and Granjean, 2013; Samson et al., 2011; Schirmer et al, 2012; Turkeltaub and Abbreviations: 2D, two-dimensional; 3D, three-dimensional; AC, auditory cor- tex; AG, angular gyrus; ALE, activation likelihood estimate; BOLD, blood oxygena- tion level dependent; ERP, event-related brain potential; FM, frequency-modulated; fMRI, functional magnetic resonance imaging; HG, Heschl’s gyrus; MEG, magne- toencephalography; MMN, mismatch negativity; MNI, Montreal Neurological Institute; MTG, middle temporal gyrus; PET, positron emission tomography; PP, planum polare; PT, planum temporale; SMG, supramarginal gyrus; STG, superior temporal gyrus; STS, superior temporal sulcus; VAMCA, Visualization and Meta- analysis on Cortical Anatomy * Corresponding author. Helsinki Collegium for Advanced Studies, University of Helsinki, PO Box 4, FI 00014 Helsinki, Finland. Tel.: þ358 405577578 (mobile). E-mail addresses: kimmo.alho@helsinki.fi (K. Alho), teemu.rinne@helsinki.fi (T. Rinne), tjherron@ebire.org (T.J. Herron), dlwoods@ucdavis.edu (D.L. Woods). Contents lists available at ScienceDirect Hearing Research journal homepage: www.elsevier.com/locate/heares 0378-5955/$ e see front matter Ó 2013 Elsevier B.V. All rights reserved. http://dx.doi.org/10.1016/j.heares.2013.08.001 Hearing Research 307 (2014) 29e41