Third Class: Neuroscience of Hearing and Speech: Neurolinguistics
Third Class: Neuroscience of Hearing and Speech: Neurolinguistics
third class:
neuroscience of hearing and speech
the ear
price!
THE HEARING BR
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air
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Part of sec
cortex, with
projections
auditory co
Parabelt r
Part of sec
cortex, rec
projections
adjacent b
Tonotopic
The princip
close to ea
frequency
by neurons
spatially cl
other in th
A membrane within the cochlea, termed the basilar membrane, contains tiny
hair cells linked to receptors. Sound induces mechanical movement of the
basilar membrane and the hair cells on it. These movements induce a flow of
ions through stretch-sensitive ion channels, that initiates neural activity (release
of neurotransmitters)
Au or
or e
Au or
or e
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100.000.000
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500.000
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brainstem
236 THE
STUDENTS GUIDE TO COGNITIVE NEUROSCIENCE
tonotopic
organization
and those responding to lower frequencies more centrally (Kiang et al., 1965).
central regions toTolower
frequencies
some extent,
this organization is carried upwards to the early cortical
Sparse scanning
stages. In both humans (Formisano et al., 2003) and other animals (Merzenich
In
fMRI,
a
short
break
in
outer
regions to ethigher
al., 1973) frequencies
there is evidence that the central region of the primary auditory cortex
scanning to enable
responds to lower frequencies and the outer regions, on both sides, to higher
sounds to be presented
frequencies.
in relative silence.
Core area
dierent pathways:
where vs. what
Parabelt area
900 Hz
300 Hz
900 Hz
3,000 Hz
Belt area
1,800 Hz
3,000 Hz
primary auditory
cortex (A1)
1,800 Hz
KEY TERM
cognitive
science
questions
VERY SPECIFIC!
4.3 Processes: the neuroanatomy of language
Classical Model
Brocas Aphasia
Conduction Aphasia
fluentness, prosody,
articulation, word finding, and
complex grammar disturbes
how independent is
speech processing
from acoustic processing?
Wernickes Aphasia
comprehension disturbed
prosody preserved, production undisturbed to
overshooting
03.02.2016
price!
SPEECH
SPEECH
AUDITORY
AUDITORY
AUDITORY
AUDITORY
PATTERN
PATTERN
PATTERN
PATTERN
RECOGNITION
RECOGNITION RECOGNITION
RECOGNITION
LOCALIZATION
LOCALIZATION
EARLYEARLY
HEARING
HEARING
(a)
(a)
SPEECH
SPEECH
EARLY
EARLY
HEARING
HEARING
(b)
(b)
LOCALIZATION
LOCALIZATION
double dissociation
between agnosia and
speech comprehension
Speech comprehension
Pure word
deafness
Auditory
agnosia
Cortical
deafness
impaired
impaired
impaired
impaired
impaired
impaired
Recognition of music
+/-
impaired
Hearing sensitivity
impaired
non-speech sounds
(audiometry)
Language I:
Spontaneous speech
Language II:
Reading comprehension
Language III:
Writing
+-sign indicates adequate performance in a given domain.
high functional
specialization
(encapsulation?
modules?
inpdenendency?)
of dierent
components
previous studies..
Dominance of le, hemisphere in Language clearly
observable from lesion studies
BUT prior func<onal imaging studies have shown bilateral
ac<va<on by speech in healthy individuals
These studies contrasted speech with rest, simple tones
or noise
Speech is a complex acous<c signal
BeAer baseline(s) might reveal le, lateraliza<on of speech
and subtle dis<nc<ons within
Intelligible Condi-ons:
Sp: Normal Speech
Vco: Noise Vocoded Speech (harsh whisper but
comprehensible a8er training)
Unintelligible Condi-ons:
RSp: Rotated Speech (preserve some phone-c features,
e.g. frica?ves, voiced-unvoiced, and intona?on)
RvCo: Rotated Noise-Vocoded Speech (completely
dis?nct from speech)
Phone&cs Contrast
Le# STG ac*vated
(Orange)
Intelligibility Contrast
Le# (more anterior)
STS ac*vated (yellow)
CONCLUSION:
with subtle exp. design
specificity of left hemisphere for
intelligible signals in passive listening
anterolateral stream of neural information
from the primary auditory cortex
(Superior Temporal Sulcus: only activated
with intelligible speech)
monosynaptic connections between
A1 and STS
confirmed by single-cell recording
in primates (specie-specific vocalizations)
linguistic analysis
manipulating speech-relevant
acoustic process
Scrambling
methodology
increasing activation
decreasing activation
suppressed
activation
for degraded
object pics
speech segments
length varied parametrically (30 to 960 ms)
quilts:
Frequency
1
1
2
30 ms
30-960 ms
3
X
[CnR (t, f )
...
L
Cn+1
(t, f )]2
t,f
Quilt
Frequency
3 9 5 2
3
...
Time
30ms Segments
960msS1Segments
CoModulation Control Ex 1
Frequency (Hz)
9413
2731
Speech
680
hypothesis:
50
2731
680
50
9413
9413
5113
5113
5113
10
-10 2731
10
2731
10
15
1411
15
1411
15
680
20
-20
680
20
680
20
274
25
274
25
274
25
2731
1411
dB
1
30
30
9413
5113
2731
1411
680
274
1
9413
-30
30
S1 CoModulation Control Ex 2
Frequency (Hz)
quency (Hz)
Modulation!
Control
30 ms Quilt
00
9413
960 ms Quilt
9413
960 ms Quilt
30 ms Quilt
9413
9413
5113
5113
10
2731
10
2731
10
15
1411
15
1411
15
20
680
20
680
20
25
274
25
274
25
30
30
30
defined anatomically
(primary Auditory cortex, Heschl
Gyrus, A1
Planum Temporale non-primary
auditory cortex, A2)
defined functionally
(Superior Temporal Sulcus,
associative auditory area)
contrast between response
to 960 ms quilts vs. 30 ms quilts
A2
ROI
ROI
segmented speech
plateau
at
480 ms
A2
ROI
ROI
segmented speech
CONTROLS!
similar response
in A1/A2
2,057
100
10
10
Audio Frequency (Hz)
8,844
0
-9
6
-3
0
od
30
Condition
10
919
Condition
M
od
90
0.2
S-
Speech 30
Speech 960
Mod 30
Mod 960
0.2
S-
80
0.4
Mod 960
0.4
-9
6
10
4
0.6
od
70
0.6
-3
0
60
0.8
od
1
4
16
128
Modulation channel (Hz)
0.8
4,325
50
1.0
S30
60
8,844
40
347
1.0
30
1.2
919
1.2
S96
Long term power spectra are very similar between source and quilt signal
2
Right hemisphere
Left hemisphere
96
b
Proportion of response
to localizer (960 ms)
Speech 960
synthesized
stimuli with103
4
8,844
algorithm decomposing
4,425 speech signal using an auditory
3
model
2,057
Level (dB)
Speech
8,844
2,856
820
Modulation control
Co-modulation control
8,844
no modulation2,856
nor response in STS
820
l sounds that
generated an
correlations
power distrime statistical
gain showed
eak effect of
of quilt type
ion between
, H2p = 0.95)
ontrol quilts
quilts in the
e to that for
animal vocalisations
A R T(dogs
I C Lbarking,
E S birds songs)
human actions (footsteps, sawing wood)
Left hemisphere
Proportion of response to localizer (960 ms)
esponses
k) to speech
egment
e participants
ition set.
Right hemisphere
1.2
1.2
1.0
1.0
0.8
0.8
0.6
0.6
0.4
0.4
0.2
S-30
S-960
Env-30 Env-960
Condition
0.2
S-30
S-960
Env-30 Env-960
Condition
similar response
in A1/A2
Quilts made from environmental sounds evoked an overall response To test the importance of p
cortical
analysis
of speech-specific
temporal
in the individual fROIs that was much the
lower
than that
for any
of the from noise-vocoded
speech
Overath et al. 2015 structure
revealed
responses
speech quilts, irrespective of segment length
(main
effect ofby
quilt
type: to
of sound
each ofquilts
our source speec
F(1,4) = 72.59, P = 0.001, H2p = 0.95; Fig. 5). The effect of segment length of ten frequency bands (co
on the fROI response was also much larger for speech quilts than cochlea, covering the audi
speech
fornoise-vocoded
environmental sound
quilts, producing an interaction between which relies on fine spectra
version
of speech
eliminate
quiltnoise-vocoded
type and segment
length
(F(1,4) =stimuli
9.45, Pthat
< 0.05,
H2p =pitch
0.7), but coarse spectral conten
but maintain
coarse
spectral between
content sufficient
for phonetic identification
remains present. When Eng
although
a pairwise
comparison
the two environmental
(cf.quilt
Scott
et al. conditions
2000)
sound
control
was significant (P < 0.05). In contrast, intelligibility remains high2
The fROI response to quil
Left hemisphere
Right hemisphere
lower than that to normal
similar response
effect
of segment length (m
1.2
1.2
in A1/A2
P < 0.01, H2p = 0.85, withou
Although the response to
1.0
1.0
thatbig
to modulation!
noise-vocoded spee
(F(1,4)
= 91.55,
P = 0.001, H2
(effect
of segment-size)
0.8
0.8
but smaller
response
stimuli
were nonetheless
m
than real speech
non-speech
quilts that we t
in STS ANOVAs for th
measures
0.6
0.6
0.4
0.4
0.2
S-30
S-960 NV-30
Condition
NV-960
0.2
S-30
S-960 NV-30
Condition
NV-960
this 6
effect
is not to noise-v
Figure
Responses
due to pitch
(os.e.m.)
in HGproperties
(red), PT (blue
(as noise-vocoded
quilts
(solid) and noise-vocode
30speech
and 960
ms.not
Data
are aver
does
have
scanned
with the noise-vocode
these properties)
30
ROI response
b 1.0
Proportion of response to localizer (960 ms)
Naturalness rating
naturaleness
rating
0.9
smaller response
N = 15
0.8
N = 11
0.7
0.6
30
60
120
240
480
960
conclusions
STS (superior temporal sulcus)
is bilaterally tuned to the acoustic structure of speech
particularly, temporal structure (use of quilts)
amplitude modulation
prosodic pitch variation
hierarchy of selectivity
STS responses are stimulus relevant
speech-specific analysis
critiques?
what is STS processing?
which speech related representations?
sillables
speech related
acoustic statistics
words
phonemes
for linguists:
these results are quite trivial
intermezzo
LINGUISTIC THEORY
NEUROSCIENCE
brain activation
phonology
phonemes
?
cortical networks & areas
functional specialization
syntax
constituents, phrase structures,
syntactic operations and dependencies
semantics
interpretation, meaning
logic, abstraction, concepts
pragmatics
intentions, communications,
goals, language use, context
intermezzo
NEUROSCIENCE
brain activation
how the brain reacts to
specific behaviour
LINGUISTIC THEORY
phonology
phonemes
syntax
cortical networks & areas
functional specialization
is this the best way
to understand how
language works (in the brain)?
semantics
interpretation, meaning
logic, abstraction, concepts
pragmatics
intentions, communications,
goals, language use, context
NEUROSCIENCE
and
LINGUISTIC THEORY
1) COMPUTATIONAL LEVEL
2) ALGORITHMIC LEVEL
3) IMPLEMENTATION LEVEL
LINGUISTIC THEORY
representations
LINGUISTIC THEORY
2) ALGORITHMIC LEVEL
how it performs its computation
operation/processes
3) IMPLEMENTATION LEVEL
how it is realized in the brain
NEUROSCIENCE
NEUROSCIENCE
compress information
allow for ABSTRACTION and GENERALIZATION
compress information
allow for ABSTRACTION and GENERALIZATION
trade-off: a representation makes certain information explicit
at the expenses of other information that is pushed in the background
it may be effortful to recover certain information
back to neurolinguistics
LINGUISTIC THEORY
NEUROSCIENCE
brain activation
cortical networks & areas
functional specialization
representations
phonemes
phonetic sounds
operation/processes
continuous
non-symbolic
more numerous than
phonological cat.
phonemes
(/t/, /d/, /a/)
phonetic
representations
phonological/phonetic processing/recognition
phonemes
phonological minimal pairs in English
phonemes are different for 1 feature
Conclusion: /p/, /b/, /m/, /s/, /n/ and /g/ are phonemes in
Fall2014
English.
36
phonology
phonetic category
phonological category
(phoneme)
discrete
Phonology
Continuous/Gradient
Discrete/Symbolic/
Categorical
Physical
Abstract
General
Language Specific
P
En pil
B
En bil
Fall2014
Release of
stop
Short VOT
38
oddball paradigm
X X X X X Y X X X X Y X X X X X X Y X X X Y X ...
MMF
Standards and deviants elicit dierent MEG responses
Fall2014
oddball paradigm
52
Fall2014
60 msec
40
real experiment
stimuli randomly sampled from one of the phonological categories
many-to-one ratio (oddball is the one)
acoustic
difference
withincategory
was greater
than
between
category
many-to-one ratio
with phono contrast
VOT shifted by
20 ms
same acoustic
dierence
many-to-one
phono contrast
was dramatically
modified
results
strong bipolar
MMF in
the phonological
experiment
experiment
-14
Phonological Experiment
4
experiment
x10
1
Acoustic Experiment
0
-100
100
200
Time(msec)
300
400
500
discussion
MMF in Phonological experiment
absence of MMF in
Acoustic experiment
abstract discrete linguistic categories are available in a part of the brain (A1)
that is known to be involved in relatively low-level auditory processing
David Poeppel