Musical imagery: sound of silence activates auditory cortex.
ABSTRACT Auditory imagery occurs when one mentally rehearses telephone numbers or has a song 'on the brain'--it is the subjective experience of hearing in the absence of auditory stimulation, and is useful for investigating aspects of human cognition. Here we use functional magnetic resonance imaging to identify and characterize the neural substrates that support unprompted auditory imagery and find that auditory and visual imagery seem to obey similar basic neural principles.
- [Show abstract] [Hide abstract]
ABSTRACT: Speech signals are often compromised by disruptions originating from external (e.g., masking noise) or internal (e.g., inaccurate articulation) sources. Speech comprehension thus entails detecting and replacing missing information based on predictive and restorative neural mechanisms. The present study targets predictive mechanisms by investigating the influence of a speech segment’s predictability on early, modality-specific electrophysiological responses to this segment’s omission. Predictability was manipulated in simple physical terms in a single-word framework (Experiment 1) or in more complex semantic terms in a sentence framework (Experiment 2). In both experiments, final consonants of the German words Lachs ([laks], salmon) or Latz ([lats], bib) were occasionally omitted, resulting in the syllable La ([la], no semantic meaning), while brain responses were measured with multi-channel electroencephalography (EEG). In both experiments, the occasional presentation of the fragment La elicited a larger omission response when the final speech segment had been predictable. The omission response occurred ∼125-165 msec after the expected onset of the final segment and showed characteristics of the omission mismatch negativity (MMN), with generators in auditory cortical areas. Suggestive of a general auditory predictive mechanism at work, this main observation was robust against varying source of predictive information or attentional allocation, differing between the two experiments. Source localization further suggested the omission response enhancement by predictability to emerge from left superior temporal gyrus and left angular gyrus in both experiments, with additional experiment-specific contributions. These results are consistent with the existence of predictive coding mechanisms in the central auditory system, and suggestive of the general predictive properties of the auditory system to support spoken word recognition.Cortex 01/2014; · 6.16 Impact Factor
- [Show abstract] [Hide abstract]
ABSTRACT: Neuroimaging studies have repeatedly reported findings of activation in frontoparietal regions that largely overlap across various cognitive functions. Part of this frontoparietal activation has been interpreted as reflecting attentional mechanisms that can adaptively be directed towards external stimulation as well as internal representations (internal attention), thereby generating the experience of distinct cognitive functions. Nevertheless, findings of material- and task-specific activation in frontal and parietal regions challenge this internal attention hypothesis and have been used to support more modular hypotheses of cognitive function. The aim of this review is twofold: First, it discusses evidence in support of the concept of internal attention and the so-called dorsal attention network (DAN) as its neural source with respect to three cognitive functions (working memory, episodic retrieval, and mental imagery). While DAN activation in all three functions has been separately linked to internal attention, a comprehensive and integrative review has so far been lacking. Second, the review examines findings of material- and process-specific activation within frontoparietal regions, arguing that these results are well compatible with the internal attention account of frontoparietal activation. A new model of cognition is presented, proposing that supposedly different cognitive concepts actually rely on similar attentional network dynamics to maintain, reactivate and newly create internal representations of stimuli in various modalities. Attentional as well as representational mechanisms are assigned to frontal and parietal regions, positing that some regions are implicated in the allocation of attentional resources to perceptual or internal representations, but others are involved in the representational processes themselvesProgress in Neurobiology 01/2014; · 9.04 Impact Factor
- [Show abstract] [Hide abstract]
ABSTRACT: The aim of this paper was to investigate the neurological underpinnings of auditory-to-motor translation during auditory repetition of unfamiliar pseudowords. We tested two different hypotheses. First we used functional magnetic resonance imaging in 25 healthy subjects to determine whether a functionally defined area in the left temporo-parietal junction (TPJ), referred to as Sylvian-parietal-temporal region (Spt), reflected the demands on auditory-to-motor integration during the repetition of pseudowords relative to a semantically mediated nonverbal sound-naming task. The experiment also allowed us to test alternative accounts of Spt function, namely that Spt is involved in subvocal articulation or auditory processing that can be driven either bottom-up or top-down. The results did not provide convincing evidence that activation increased in either Spt or any other cortical area when non-semantic auditory inputs were being translated into motor outputs. Instead, the results were most consistent with Spt responding to bottom up or top down auditory processing, independent of the demands on auditory-to-motor integration. Second, we investigated the lesion sites in eight patients who had selective difficulties repeating heard words but with preserved word comprehension, picture naming and verbal fluency (i.e., conduction aphasia). All eight patients had white-matter tract damage in the vicinity of the arcuate fasciculus and only one of the eight patients had additional damage to the Spt region, defined functionally in our fMRI data. Our results are therefore most consistent with the neurological tradition that emphasizes the importance of the arcuate fasciculus in the non-semantic integration of auditory and motor speech processing.Frontiers in Human Neuroscience 01/2014; 8:24. · 2.91 Impact Factor
Sound of silence
activates auditory cortex
subjective experience of hearing in the
absence of auditory stimulation, and is use-
ful for investigating aspects of human cog-
nition1. Here we use functional magnetic
resonance imaging to identify and charac-
terize the neural substrates that support
unprompted auditory imagery and find
that auditory and visual imagery seem to
obey similar basic neural principles.
The few studies that have examined the
topic ofauditory imagery2–5have focused on
the neural substrates ofdirected imagery (for
example, “imagine a tone”). What is not
known,however,is whether similar principles
guide the more pervasive and spontaneous
forms of imagery that punctuate everyday
life.We used functional magnetic resonance
imaging to investigate the recruitment of
auditory cortex during spontaneous auditory
imagery ofexcerpts ofpopular music.
During scanning, subjects passively lis-
tened to excerpts of songs with lyrics (for
example, Satisfaction by the Rolling Stones)
and to instrumentals that contained no lyrics
(for example,the theme from ThePink Pan-
ther). Each piece of music was pre-rated by
subjects as either familiar or unknown,and a
unique soundtrack was created for each indi-
vidual. Short sections of music (lasting for
uditory imagery occurs when one
mentally rehearses telephone numbers
or has a song ‘on the brain’ — it is the
Our findings offer a neural basis for the
spontaneous and sometimes vexing experi-
ence of hearing a familiar melody in one’s
head.Whereas previous investigations have
explicitly directed subjects to imagine a
specific auditory experience2–4,we provided
no instruction. Instead, simply muting
short gaps offamiliar music was sufficient to
trigger auditory imagery — a finding that
indicates the obligatory nature of this phe-
nomenon. Corroborating this observation,
all subjects reported subjectively hearing
a continuation ofthe familiar songs,but not
of the unfamiliar songs, during the gaps in
We note also that the extent of neural
activity in the primary auditory cortex was
determined by the linguistic features of
the imagined experience. When semantic
knowledge (that is, lyrics) could be used to
generate the missing information, recon-
struction terminated in auditory association
areas. When this meaning-based route to
reconstruction was unavailable (as in instru-
mentals), activity extended to lower-level
regions of the auditory cortex,most notably
the primary auditory cortex (Fig.1b,d).
These findings parallel those in the
domain ofvisual imagery.For example,visual
imagery elicited when considering names of
objects (known as figural imagery) does not
rely on the primary visual cortex6,7.As these
‘low-resolution’images do not demand fine-
grained perceptual processing, activity in
visual-association areas is sufficient to
reconstruct the relevant representation. By
contrast, when semantic information is
absent or irrelevant (known as depictive
imagery), a ‘high-resolution’ perceptual
image is needed to reconstruct a representa-
tion,hence activity extends into the primary
visual cortex8. Our results provide evidence
that auditory imagery obeys the same basic
David J.M.Kraemer*,C.Neil Macrae*†,
Adam E.Green*,William M.Kelley*
*Department of Psychological and Brain Sciences,
Dartmouth College, Hanover, New Hampshire
†School of Psychology, University of Aberdeen,
Aberdeen AB24 2UB, UK
1. Kosslyn,S.M.,Ganis,G.& Thompson,W.L. Nature Rev.
Neurosci. 2, 635–642 (2001).
2. Halpern, A. R. & Zatorre, R. J. Cereb. Cortex 9, 697–704
3. Wheeler, M. E., Petersen, S. E. & Buckner, R. L.Proc. Natl Acad.
Sci. USA 97, 11125–11129 (2000).
4. Yoo, S. S., Lee, C. U. & Choi, B. G.Neuroreport 12, 3045–3049
5. Hughes, H. C. et al. Neuroimage 13, 1073–1089 (2001).
6. Fletcher,P.C. et al. Neuroimage 2, 195–200 (1995).
7. Kosslyn, S. M. & Thompson,W. L.Psychol. Bull. 129, 723–746
8. Kosslyn,S.M.,Thompson,W.L.,Kim,I.J.& Alpert,N.M.
Nature 378, 496–498 (1995).
9. Van Essen,D.C. et al. J. Am. Med. Inform. Assoc. 41, 1359–1378
Supplementary information accompanies this communication on
Competing financial interests:declared none.
NATURE|VOL 434|10 MARCH 2005|www.nature.com/nature
2–5s) were extracted at different points dur-
ing the soundtrack and replaced with silent
gaps.We then monitored the neural activity
in subjects that occurred during these gaps.
(For details of methods, see supplementary
Brain activity in the primary auditory
cortex and in the auditory association cortex
(Brodmann’s area 22) (Fig. 1a) was com-
pared during gaps of silence in familiar and
unknown songs.The results revealed a func-
tional dissociation within the left auditory
cortex (region?music-type interaction:
F[1,14]?48.92, P?0.0001; Fig. 1b). Silent
gaps embedded in familiar songs induced
greater activation in auditory association
areas than did silent gaps embedded in
unknown songs (Fig.1b);this was true for gaps
in songs with lyrics (F[1,14]?5.46,P?0.05;
Fig.1c) and without lyrics (F[1,14]?11.56,
songs contained no lyrics, cortical activity
extended into the left primary auditory
We confirmed that these effects were
uniquely attributable to the gaps ofsilence in
the music, rather than simply the result of
differences in activation in response to hear-
ing different music categories. By contrast
with the gap responses,listening to unknown
songs produced greater activity in auditory
association areas than did familiar songs
(lyrics: F[1,14]?11.24, P?0.005; instru-
mentals: F[1,14]?31.74, P?0.0001), and
activity in the primary auditory cortex
did not differ as a function of familiarity
(see supplementary information).
FLFI UL UI
Figure 1 Auditory cortex activation during silent gaps in music. a, An inflated rendering of the left hemisphere9illustrates primary auditory
cortex (PAC; red) and auditory association cortex, also known as Brodmann’s area 22 (green). The superior temporal sulcus (STS) and
inferior temporal sulcus (ITS) are indicated for reference. b, Signal change (arbitrary units) in PAC (red) and Brodmann’s area 22 (green)
during gaps in familiar songs with lyrics (FL), familiar instrumentals (FI), unknown songs with lyrics (UL) and unknown instrumentals (UI).
Error bars denote s.e.m. c, d, Difference in activity, which is greater for familiar songs, during silent gaps embedded in songs with (c) and
without (d) lyrics,projected on to flattened views of the left temporal lobe.Dark-grey regions represent sulci; lighter grey regions denote gyri.