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Spectrotemporal modulation provides a unifying framework for auditory cortical asymmetries



The principles underlying functional asymmetries in cortex remain debated. For example, it is accepted that speech is processed bilaterally in auditory cortex, but a left hemisphere dominance emerges when the input is interpreted linguistically. The mechanisms, however, are contested, such as what sound features or processing principles underlie laterality. Recent findings across species (humans, canines and bats) provide converging evidence that spectrotemporal sound features drive asymmetrical responses. Typically, accounts invoke models wherein the hemispheres differ in time–frequency resolution or integration window size. We develop a framework that builds on and unifies prevailing models, using spectrotemporal modulation space. Using signal processing techniques motivated by neural responses, we test this approach, employing behavioural and neurophysiological measures. We show how psychophysical judgements align with spectrotemporal modulations and then characterize the neural sensitivities to temporal and spectral modulations. We demonstrate differential contributions from both hemispheres, with a left lateralization for temporal modulations and a weaker right lateralization for spectral modulations. We argue that representations in the modulation domain provide a more mechanistic basis to account for lateralization in auditory cortex.

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Fig. 1: Time, time–frequency and modulation domain representations for sound waveforms.
Fig. 2: The modulation asymmetry hypothesis.
Fig. 3: Overview of the filtering technique used to produce modulation domain filtered stimuli.
Fig. 4: Psychophysical performance as a function of temporal and spectral modulations in two separate experiments (diotic and dichotic).
Fig. 5: MEG results showing significant correlations between neural power and degree of stimulus modulation.
Fig. 6: Intracranial ECoG recordings in a patient with bilateral stereotactic depth electrodes, sampling superior temporal cortices.

Code availability

All stimuli construction code is available on a public repository Experiment and analysis code is available from the corresponding author upon reasonable request.

Data availability

The data that support the findings of this study are available from the corresponding author upon reasonable request.


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This work was supported by NIH F32 DC011985 and Charles H. Revson Senior Fellowships in Biomedical Science 15–28 to A.F., by NIH 2R01DC05660 to D.P. and by NIMH R21 MH114166-01 to A.D.M. The funders had no role in study design, data collection and analysis, decision to publish or preparation of the manuscript. We would like to thank I. T. Kim and N. Mei, who assisted in the setup and acquisition of psychophysical dichotic data, B. Mahmood and M. Hofstradter, who assisted in NYU ECoG data acquisition and setup, D. Groppe, who assisted in North Shore ECoG data acquisition and electrode reconstruction, and H. Wang, who provided electrode reconstruction at NYU.

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A.F. and D.P. designed the study and hypotheses, and wrote the manuscript. A.F. constructed the stimuli and filtering techniques and collected and analysed the data. A.D.M., O.D. and W.K.D. recruited clinical patients and performed clinical care.

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Correspondence to Adeen Flinker.

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Flinker, A., Doyle, W.K., Mehta, A.D. et al. Spectrotemporal modulation provides a unifying framework for auditory cortical asymmetries. Nat Hum Behav 3, 393–405 (2019).

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