Auditory attention — focusing the searchlight on sound

doi:10.1016/j.conb.2007.07.011

Current Opinion in Neurobiology

Volume 17, Issue 4, August 2007, Pages 437-455

https://doi.org/10.1016/j.conb.2007.07.011 Get rights and content

Some fifty years after the first physiological studies of auditory attention, the field is now ripening, with exciting recent insights into the psychophysics, psychology, and neural basis of auditory attention. Current research seeks to unravel the complex interactions of pre-attentive and attentive processing of the acoustic scene, the role of auditory attention in mediating receptive-field plasticity in both auditory spatial and auditory feature processing, the contrasts and parallels between auditory and visual attention pathways and mechanisms, the interplay of bottom-up and top-down attentional mechanisms, the influential role of attention, goals, and expectations in shaping auditory processing, and the orchestration of diverse attentional effects at multiple levels from the cochlea to the cortex.

Section snippets

Introduction and overview

Auditory attention allows us to rapidly and precisely direct our acoustic searchlight toward sounds of interest in our acoustic environment. Attention can be top-down (voluntary or task-dependent) or bottom-up (sound-based salience). At the interface of perception and action, top-down attention leads to enhanced information processing, behavioral sensitivity, and shortened response latencies. Top-down attention is a selection process that focuses cortical processing resources on the most

Relationship between pre-attentive and attentive processes in auditory scene analysis

In order to focus auditory attention on specific acoustic objects of interest in the real world, we typically make use of a combination of auditory spatial cues and auditory feature cues to solve the pattern recognition problem of foreground–background decomposition (FBD). This is illustrated by one of the best known examples of auditory attention, the ‘cocktail party effect’, in which we can attend and selectively eavesdrop on different speakers in a crowded room brimming with multiple

Auditory spatial attention

Depending upon whether an auditory task requires attending to a spatial location, or to an auditory feature or object, there may be differential activation of the auditory ‘what’ and ‘where’ pathways [73, 74, 75]. Attentional mechanisms can modulate neural activity encoding the spatial location and/or the acoustic attributes of the selected targets and the early sensory representation of attended stimuli [2^•]. For simplicity, we shall distinguish between auditory spatial and non-spatial

Auditory feature and object attention — extracting signals from background

A variety of complex, shifting acoustic soundscapes present enormous challenges for acoustic scene analysis and for attentional focus on auditory features or objects such as environmental soundscapes (such as a morning chorus of birds), polyphonic music and speech. Top-down attention can selectively focus on a limited range of an acoustic feature dimensions [7], or can even focus on the expected (or recalled) features of an auditory target [15]. Although bottom-up salience certainly plays a

Auditory attention in time

Precisely focused temporal expectancies, such as musical expectancies, are likely to be very important in auditory processing since many auditory patterns unfold in time. A recent ERP study has shown that auditory attention can be temporally directed to focus on events that are projected to occur at a particular future point in time [106]. In another study, subjects engaged in an auditory task could avoid involuntary attentional capture by distracting acoustic stimuli, by foreknowledge of when

Attentive imagery in silence and hallucinations

In auditory induction (such as the familiar psychoacoustic phenomena of FM completion or phonemic restoration) the auditory system fills in occluded information, as when missing foreground sound segments are perceptually restored in the presence of background sound [23, 117, 118]. Although it might seem at first glance like a reasonable candidate for top-down effects, there is considerable evidence for pre-attentive mechanisms in auditory induction [118], though this may be influenced by

Effects of auditory attention on receptive-field plasticity

The adaptive functions of the cerebral cortex rely upon flexibility and plasticity of information processing networks. Since the topic of plasticity of auditory cortical processing has been recently reviewed [135], we will focus in this section on the evidence that attention may play a decisive role in triggering auditory plasticity, particularly in the adult brain [12, 136]. Parallel studies have shown that attention can initiate plasticity in other sensory cortices, as well as in motor cortex

Intermodal and crossmodal interactions between auditory and visual attention

There are many similarities between attention in the auditory and visual modalities, where a two-component framework for attentional selection (top-down and bottom-up) has also emerged from psychophysical, behavioral, and neurobiological studies. Two sets of mechanisms are thought to operate in parallel in both modalities: using either bottom-up, automatic, image-based saliency cues or top-down, attentional, task-dependent cues. Another fundamental similarity is that attention can modulate both

Neural networks of auditory attention

Auditory attention can be selectively directed to a rich variety of acoustic features including spatial location, auditory pitch, frequency or intensity, tone duration, timbre, FM direction or slope, speech versus nonspeech streams, and characteristics of individual voices. Given the multiplicity of acoustic dimensions to which we can attend and the richly interconnected auditory processing networks, there are likely to be multiple neural loci for auditory attention. In fact, the locations of

Summary

Auditory attention involves a distributed network of auditory cortical and subcortical structures that are activated selectively in a task-specific manner during auditory processing, which also integrate with a generalized multisensory attentional network that includes parietal, frontal, and anterior cingulate cortical regions [74, 207, 208, 209, 210]. Recent research has revealed a richly interconnected network for auditory attention that assists in the computation of early auditory features

References and recommended reading

Some articles have been marked as worthy of special interest. All papers in this subjective category are recent (publication within the last five years) and relevant (results making an important contribution to the field of auditory attention).

• Of special interest

Acknowledgement

We gratefully acknowledge funding from NIH R01 DC005779.

Glossary

A1: Primary auditory cortex.
ACC: Anterior cingulate cortex — medial prefrontal structure likely to be important in control of attention.
ASA: Auditory scene analysis — decomposition of complex mixture of incoming sounds into individual sound sources and sound streams.
ERP: Event-related brain potentials (averaged EEG segments time-locked to stimulus onset).
FBD: Foreground–background decomposition — separation of foreground sound stream of interest from background acoustic scene.
MMN: Mismatch negativity — a negative

References (213)

S. Kastner et al.
Visual attention as a multilevel selection process (review)
Cogn Affect Behav Neurosci
(2004)
Nobre AC, Correa A, Coull JT: Temporal effects of attention (review). Curr Opin Neurobiol 2007, (this...
J.B. Fritz et al.
Differential dynamic plasticity of A1 receptive fields during multiple spectral tasks
J Neurosci
(2005)
D.H. Hubel et al.
Attention units in the auditory cortex
Science
(1959)
H. Spitzer et al.
Increased attention enhances both behavioral and neuronal performance
Science
(1988)
C.E. Boudreau et al.
Effects of task difficulty and target likelihood in area V4 of macaque monkeys
J Neurophysiol
(2006)
A.S. Bregman
Auditory Scene Analysis: The Perceptual Organization of Sounds
(1990)
U. Groger et al.
Classification of human breathing sounds by the common vampire bat, Desmodus rotundus
BMC Biol
(2006)
E.S. Sussman et al.
The role of attention in the formation of auditory streams
Percept Psychophys
(2007)
E. Sussman et al.
Dynamic sensory updating in the auditory system
Brain Res Cogn Brain Res
(2001)

E. Brattico et al.

Musical scale properties are automatically processed in the human auditory cortex

Brain Res

(2006)

B. Opitz et al.

Sensory and cognitive mechanisms for preattentive change detection in auditory cortex

Eur J Neurosci

(2005)

M.G. Woldorff et al.

Magnetoencephalographic recordings demonstrate attentional modulation of mismatch-related neural activity in human auditory cortex

Psychophysiology

(1998)

C. Alain et al.

Attention modulates auditory pattern memory as indexed by event-related brain potentials

Psychophysiology

(1997)

E. Sussman et al.

An investigation of the auditory streaming effect using event-related brain potentials

Psychophysiology

(1999)

I.P. Jaaskelainen et al.

Human posterior auditory cortex gates novel sounds to consciousness

Proc Natl Acad Sci USA

(2004)

N. Ulanovsky et al.

Multiple time scales of adaptation in auditory cortex neurons

J Neurosci

(2004)

S. Watkins et al.

Brain mechanisms mediating auditory attentional capture in humans

Cereb Cortex

(2007)

J.S. Snyder et al.

Effects of attention on neuroelectric correlates of auditory stream segregation

J Cogn Neurosci

(2006)

S.R. Arnott et al.

Assessing the auditory dual-pathway model in humans

Neuroimage

(2004)

A. Bellmann et al.

Two types of auditory neglect

Brain

(2001)

S. Clarke et al.

Auditory neglect: what and where in auditory space

Cortex

(2004)

L. Spierer et al.

Extinction of auditory stimuli in hemineglect: space versus ear

Neuropsychologia

(2007)

T. Moore et al.

Selective gating of visual signals by microstimulation of frontal cortex

Nature

(2003)

A. Garg et al.

Orienting auditory spatial attention engages frontal eye fields and medial occipital cortex in congenitally blind humans

Neuropsychologia

(2007)

G.S. Russo et al.

Frontal eye field activity preceding aurally guided saccades

J Neurophysiol

(1994)

D.E. Winkowski et al.

Top-down gain control of the auditory space map by gaze control circuitry in the barn owl

Nature

(2006)

D.E. Winkowski et al.

Modulation of visual responses in optic tectum by microstimulation of forebrain gaze fields in barn owls

C. Alain et al.

Selectively attending to auditory objects

Front Biosci

(2000)

Fritz JB, Elhilali M, Shamma SA: Adaptive changes in cortical receptive fields induced by attention to complex sounds....

K. Alho et al.

Brain activity during selective listening to natural speech (review)

Front Biosci

(2007)

J. Fan et al.

Human attentional networks

Psychiatr Prax

(2004)

J. Ahveninen et al.

Task-modulated ‘what’ and ‘where’ pathways in human auditory cortex

Proc Natl Acad Sci USA

(2006)

J.A. Johnson et al.

Neural substrates for dividing and focusing attention between simultaneous auditory and visual events

Neuroimage

(2006)

E.I. Knudsen

Fundamental components of attention

Annu Rev Neurosci

(2007)

C. Kayser et al.

Mechanisms for allocating auditory attention: an auditory saliency map

Curr Biol

(2005)

E.R. Hafter et al.

Auditory attention and filters (review)

C.D. Gilbert et al.

Brain states: top-down influences in sensory processing (review)

Neuron

(2007)

J. Fritz et al.

Rapid task-related plasticity of spectrotemporal receptive fields in primary auditory cortex

Nat Neurosci

(2003)

J.B. Fritz et al.

Does attention play a role in dynamic receptive field adaptation to changing acoustic salience in A1? (Review)

Hear Res

(2007)

A. Brechmann et al.

Hemispheric shifts of sound representation in auditory cortex with conceptual listening

Cereb Cortex

(2005)

H. Scheich et al.

The cognitive auditory cortex: task-specificity of stimulus representations

Hear Res

(2007)

R.J. Zatorre

There's more to auditory cortex than meets the ear (review)

Hear Res

(2007)

S.A. Hillyard et al.

Electrical signs of selective attention in the human brain

Science

(1973)

M.G. Woldorff et al.

Modulation of early auditory processing during selective listening to rapidly presented tones

Electroencephalogr Clin Neurophysiol

(1991)

E.C. Cherry

Some experiments on the recognition of speech, with one and two ears

J Acoust Soc Am

(1953)

H. Asari et al.

Sparse representations for the cocktail party problem

J Neurosci

(2006)

R.P. Carlyon

How the brain separates sounds

Trends Cogn Sci

(2004)

C. Alain

Breaking the wave: effects of attention and learning on concurrent sound perception (review)

Hear Res

(2007)

B.G. Shinn-Cunningham et al.

A sound element gets lost in perceptual competition

Proc Natl Acad Sci USA

(2007)

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Binaural reproduction aims at recreating a realistic sound scene at the ears of the listener using headphones. Unfortunately, externalization for frontal and rear sources is often poor (virtual sources are perceived inside the head, instead of outside the head). Nevertheless, previous studies have shown that large head-tracked movements could substantially improve externalization and that this improvement persisted once the subject had stopped moving his/her head. The present study investigates the relation between externalization and evoked response potentials (ERPs) by performing behavioral and EEG measurements in the same experimental conditions. Different degrees of externalization were achieved by preceding measurements with 1) head-tracked movements, 2) untracked head movements, and 3) no head movement. Results showed that performing a head movement, whether the head tracking was active or not, increased the amplitude of ERP components after 100 ms, which suggests that preceding head movements alters the auditory processing. Moreover, untracked head movements gave a stronger amplitude on the N1 component, which might be a marker of a consistency break in regards to the real world. While externalization scores were higher after head-tracked movements in the behavioral experiment, no marker of externalization could be found in the EEG results.
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Auditory cortical neurons modify their response profiles in response to numerous external factors. During task performance, changes in primary auditory cortex (A1) responses are thought to be driven by top-down inputs from the orbitofrontal cortex (OFC), which may lead to response modification on a trial-by-trial basis. While OFC neurons respond to auditory stimuli and project to A1, the function of OFC projections to A1 during auditory tasks is unknown. Here, we observed the activity of putative OFC terminals in A1 in mice by using in vivo two-photon calcium imaging of OFC terminals under passive conditions and during a tone detection task. We found that behavioral activity modulates but is not necessary to evoke OFC terminal responses in A1. OFC terminals in A1 form distinct populations that exclusively respond to either the tone, reward, or error. Using tones against a background of white noise, we found that OFC terminal activity was modulated by the signal-to-noise ratio (SNR) in both the passive and active conditions and that OFC terminal activity varied with SNR, and thus task difficulty in the active condition. Therefore, OFC projections in A1 are heterogeneous in their modulation of auditory encoding and likely contribute to auditory processing under various auditory conditions.
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Auditory attention — focusing the searchlight on sound

Section snippets

Introduction and overview

Relationship between pre-attentive and attentive processes in auditory scene analysis

Auditory spatial attention

Auditory feature and object attention — extracting signals from background

Auditory attention in time

Attentive imagery in silence and hallucinations

Effects of auditory attention on receptive-field plasticity

Intermodal and crossmodal interactions between auditory and visual attention

Neural networks of auditory attention

Summary

References and recommended reading

Acknowledgement

Glossary

Cogn Affect Behav Neurosci

J Neurosci

Science

Science

J Neurophysiol

BMC Biol

Percept Psychophys

Brain Res Cogn Brain Res

Brain Res

Eur J Neurosci

Psychophysiology

Psychophysiology

Psychophysiology

Proc Natl Acad Sci USA

J Neurosci

Cereb Cortex

J Cogn Neurosci

Neuroimage

Brain

Cortex

Neuropsychologia

Nature

Neuropsychologia

J Neurophysiol

Nature

Front Biosci

Front Biosci

Human attentional networks

Psychiatr Prax

Task-modulated ‘what’ and ‘where’ pathways in human auditory cortex

Proc Natl Acad Sci USA

Neural substrates for dividing and focusing attention between simultaneous auditory and visual events

Neuroimage

Fundamental components of attention

Annu Rev Neurosci

Mechanisms for allocating auditory attention: an auditory saliency map

Curr Biol

Auditory attention and filters (review)

Brain states: top-down influences in sensory processing (review)

Neuron

Rapid task-related plasticity of spectrotemporal receptive fields in primary auditory cortex

Nat Neurosci

Does attention play a role in dynamic receptive field adaptation to changing acoustic salience in A1? (Review)

Hear Res

Hemispheric shifts of sound representation in auditory cortex with conceptual listening

Cereb Cortex

The cognitive auditory cortex: task-specificity of stimulus representations

Hear Res

There's more to auditory cortex than meets the ear (review)

Hear Res

Electrical signs of selective attention in the human brain

Science

Modulation of early auditory processing during selective listening to rapidly presented tones

Electroencephalogr Clin Neurophysiol

Some experiments on the recognition of speech, with one and two ears

J Acoust Soc Am

Sparse representations for the cocktail party problem

J Neurosci

How the brain separates sounds

Trends Cogn Sci

Breaking the wave: effects of attention and learning on concurrent sound perception (review)

Hear Res

A sound element gets lost in perceptual competition

Proc Natl Acad Sci USA