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Attention powerfully influences auditory perception, but little is understood about the mechanisms whereby attention sharpens responses to unattended sounds. We used high-resolution surface mapping techniques (using functional magnetic resonance imaging, fMRI) to examine activity in human auditory cortex during an intermodal selective attention task. Stimulus-dependent activations (SDAs), evoked by unattended sounds during demanding visual tasks, were maximal over mesial auditory cortex. They were tuned to sound frequency and location, and showed rapid adaptation to repeated sounds. Attention-related modulations (ARMs) were isolated as response enhancements that occurred when subjects performed pitch-discrimination tasks. In contrast to SDAs, ARMs were localized to lateral auditory cortex, showed broad frequency and location tuning, and increased in amplitude with sound repetition. The results suggest a functional dichotomy of auditory cortical fields: stimulus-determined mesial fields that faithfully transmit acoustic information, and attentionally labile lateral fields that analyze acoustic features of behaviorally relevant sounds.

Simple reaction time (SRT), the minimal time needed to respond to a stimulus, is a basic measure of processing speed. SRTs were first measured by Francis Galton in the 19th century, who reported visual SRT latencies below 190 ms in young subjects. However, recent large-scale studies have reported substantially increased SRT latencies that differ markedly in different laboratories, in part due to timing delays introduced by the computer hardware and software used for SRT measurement. We developed a calibrated and temporally precise SRT test to analyze the factors that influence SRT latencies in a paradigm where visual stimuli were presented to the left or right hemifield at varying stimulus onset asynchronies (SOAs). Experiment 1 examined a community sample of 1469 subjects ranging in age from 18 to 65. Mean SRT latencies were short (231, 213 ms when corrected for hardware delays) and increased significantly with age (0.55 ms/year), but were unaffected by sex or education. As in previous studies, SRTs were prolonged at shorter SOAs and were slightly faster for stimuli presented in the visual field contralateral to the responding hand. Stimulus detection time (SDT) was estimated by subtracting movement initiation time, measured in a speeded finger tapping test, from SRTs. SDT latencies averaged 131 ms and were unaffected by age. Experiment 2 tested 189 subjects ranging in age from 18 to 82 years in a different laboratory using a larger range of SOAs. Both SRTs and SDTs were slightly prolonged (by 7 ms). SRT latencies increased with age while SDT latencies remained stable. Precise computer-based measurements of SRT latencies show that processing speed is as fast in contemporary populations as in the Victorian era, and that age-related increases in SRT latencies are due primarily to slowed motor output.

BackgroundWhile human auditory cortex is known to contain tonotopically organized auditory cortical fields (ACFs), little is known about how processing in these fields is modulated by other acoustic features or by attention.Methodology/Principal FindingsWe used functional magnetic resonance imaging (fMRI) and population-based cortical surface analysis to characterize the tonotopic organization of human auditory cortex and analyze the influence of tone intensity, ear of delivery, scanner background noise, and intermodal selective attention on auditory cortex activations. Medial auditory cortex surrounding Heschl's gyrus showed large sensory (unattended) activations with two mirror-symmetric tonotopic fields similar to those observed in non-human primates. Sensory responses in medial regions had symmetrical distributions with respect to the left and right hemispheres, were enlarged for tones of increased intensity, and were enhanced when sparse image acquisition reduced scanner acoustic noise. Spatial distribution analysis suggested that changes in tone intensity shifted activation within isofrequency bands. Activations to monaural tones were enhanced over the hemisphere contralateral to stimulation, where they produced activations similar to those produced by binaural sounds. Lateral regions of auditory cortex showed small sensory responses that were larger in the right than left hemisphere, lacked tonotopic organization, and were uninfluenced by acoustic parameters. Sensory responses in both medial and lateral auditory cortex decreased in magnitude throughout stimulus blocks. Attention-related modulations (ARMs) were larger in lateral than medial regions of auditory cortex and appeared to arise primarily in belt and parabelt auditory fields. ARMs lacked tonotopic organization, were unaffected by acoustic parameters, and had distributions that were distinct from those of sensory responses. Unlike the gradual adaptation seen for sensory responses, ARMs increased in amplitude throughout stimulus blocks.Conclusions/SignificanceThe results are consistent with the view that medial regions of human auditory cortex contain tonotopically organized core and belt fields that map the basic acoustic features of sounds while surrounding higher-order parabelt regions are tuned to more abstract stimulus attributes. Intermodal selective attention enhances processing in neuronal populations that are partially distinct from those activated by unattended stimuli.

We measured digit span (DS) in two experiments that used computerized presentation of randomized auditory digits with performance-adapted list length adjustment. A new mean span (MS) metric of DS was developed that showed reduced variance, improved test-retest reliability, and higher correlations with the results of other neuropsychological test results when compared to traditional DS measures. The MS metric also enhanced the sensitivity of forward versus backward span comparisons, enabled the development of normative performance criteria with sub-digit precision, and elucidated changes in DS performance with age- and education level. Computerized stimulus delivery and improved scoring metrics significantly enhance the precision of DS assessments of short-term verbal memory.

While auditory cortex in non-human primates has been subdivided into multiple functionally specialized auditory cortical fields (ACFs), the boundaries and functional specialization of human ACFs have not been defined. In the current study, we evaluated whether a widely accepted primate model of auditory cortex could explain regional tuning properties of fMRI activations on the cortical surface to attended and non-attended tones of different frequency, location, and intensity. The limits of auditory cortex were defined by voxels that showed significant activations to non-attended sounds. Three centrally located fields with mirror-symmetric tonotopic organization were identified and assigned to the three core fields of the primate model while surrounding activations were assigned to belt fields following procedures similar to those used in macaque fMRI studies. The functional properties of core, medial belt, and lateral belt field groups were then analyzed. Field groups were distinguished by tonotopic organization, frequency selectivity, intensity sensitivity, contralaterality, binaural enhancement, attentional modulation, and hemispheric asymmetry. In general, core fields showed greater sensitivity to sound properties than did belt fields, while belt fields showed greater attentional modulation than core fields. Significant distinctions in intensity sensitivity and contralaterality were seen between adjacent core fields A1 and R, while multiple differences in tuning properties were evident at boundaries between adjacent core and belt fields. The reliable differences in functional properties between fields and field groups suggest that the basic primate pattern of auditory cortex organization is preserved in humans. A comparison of the sizes of functionally defined ACFs in humans and macaques reveals a significant relative expansion in human lateral belt fields implicated in the processing of speech.

Is attentional selection between local and global forms based on spatial frequency? This question was examined by having subjects identify local or global forms of stimuli that had been "contrast balanced," a technique that eliminates low spatial frequencies. Response times (RTs) to global (but not local) forms were slowed for contrast-balanced stimuli, suggesting that low spatial frequencies mediate the global RT advantage typically reported. In contrast, the beneficial effect of having targets appear at the same, as opposed to a different, level as that on the immediately preceding trial was unaffected by contrast balancing. This suggests that attentional selection between different levels of structure is not based on spatial frequency. The data favor an explanation in terms of "priming," rather than in terms of adjustments in the diameter of an attentional "spotlight."Numerous investigators have suggested that the analysis of hierarchically organized stimuli might depend on the differences in spatial frequency between local and global forms (Badcock, Whitworth, Badcock, & Lovegrove, 1990;Hughes, Fendrich, & Reuter-Lorenz, 1990;LaGasse, 1993;Lovegrove & Pepper, 1994;Navon, 1991;Sergent, 1982Sergent, , 1987Shulman, Sullivan, Gish, & Sakoda, 1986). Local information (e.g., the Ss in Figure Ia) is carried in relatively high spatial frequencies, whereas global information (e.g., the A in Figure 1a) is carried in relatively low spatial frequencies. Several studies have provided evidence that the speed or efficiency with which local and global levels of structure are analyzed depends on these differences in spatial frequency (LaGasse, 1993; Lamb & Yund, 1993, in press; Badcock et aI., 1990; Shulman et aI., 1986 was lower than the adapting frequency that most affected performance on the local task.The role that spatial frequency might play in the analysis of hierarchical structure is complicated by recent neuropsychological evidence suggesting that several interacting but separate mechanisms are involved in the analysis of hierarchically organized stimuli (Lamb, Robertson, & Knight, 1989, 1990Robertson & Lamb, 1991;Robertson, Lamb, & Knight, 1988). These studies have demonstrated lesion-site-specific selective impairment of (I) the speed or efficiency with which targets at different levels of structure are identified (referred to as reaction time, or RT,advantage), (2) the degree to which distractors at one level affect responding to targets at another (referred to as interference), and (3) changes in RTs occurring in response to changing task demands (referred to as attention). Similar dissociations have been demonstrated in neurologically intact subjects as well (LaGasse, 1993;Lamb & Yund, 1993;Lovegrove & Pepper, 1994;Navon & Norman, 1983;Paquet, 1992). To the extent that several different mechanisms are involved in the analysis ofhierarchically organized stimuli, it becomes necessary to determine the role that spatial frequency plays for each mechanism, because it is quite possible that spatial frequency is impo...

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