Auditory aversion in absolute pitch possessors

Abstract

Absolute pitch (AP) refers to the ability of identifying the pitch of a given tone without reliance on any reference pitch. The downside of possessing AP may be the experience of disturbance when exposed to out-of-tune tones. Here, we investigated this so-far unexplored phenomenon in AP, which we refer to as auditory aversion. Electroencephalography (EEG) was recorded in a sample of AP possessors and matched control musicians without AP while letting them perform a task underlying a so-called affective priming paradigm: Participants judged valenced pictures preceded by musical primes as quickly and accurately as possible. The primes were bimodal, presented as tones in combination with visual notations that either matched or mismatched the actually presented tone. Both samples performed better in judging unpleasant pictures over pleasant ones. In comparison with the control musicians, the AP possessors revealed a more profound discrepancy between the two valence conditions, and their EEG revealed later peaks at around 200 ms (P200) after prime onset. Their performance dropped when responding to pleasant pictures preceded by incongruent primes, especially when mistuned by one semitone. This interference was also reflected in an EEG deflection at around 400 ms (N400) after picture onset, preceding the behavior responses. These findings suggest that AP possessors process mistuned musical stimuli and pleasant pictures as affectively unrelated with each other, supporting an aversion towards out-of-tune tones in AP possessors. The longer prime-related P200 latencies exhibited by AP possessors suggest a delay in integrating musical stimuli, underlying an altered affinity towards pitch–label associations.

Full text

Research Report
Auditory aversion in absolute pitch possessors
Lars Rogenmoser
a,b,*
, H.Charles Li
a
, Lutz J€
ancke
c,d
and
Gottfried Schlaug
a
a
Department of Neurology, Beth Israel Deaconess Medical Center and Harvard Medical School, Boston, MA, USA
b
Department of Medicine, University of Fribourg, Fribourg, Switzerland
c
Department of Psychology, University of Zurich, Zurich, Switzerland
d
University Research Priority Program (URPP), Dynamics of Healthy Aging, University of Zurich, Zurich,
Switzerland
article info
Article history:
Received 30 March 2020
Reviewed 9 June 2020
Revised 7 August 2020
Accepted 9 November 2020
Action editor Gus Buchtel
Published online 5 December 2020
Keywords:
Affective priming
Fluency processing
Negativity bias
N400
N100/P200
abstract
Absolute pitch (AP) refers to the ability of identifying the pitch of a given tone without
reliance on any reference pitch. The downside of possessing AP may be the experience of
disturbance when exposed to out-of-tune tones. Here, we investigated this so-far unex-
plored phenomenon in AP, which we refer to as auditory aversion. Electroencephalography
(EEG) was recorded in a sample of AP possessors and matched control musicians without
AP while letting them perform a task underlying a so-called affective priming paradigm:
Participants judged valenced pictures preceded by musical primes as quickly and accu-
rately as possible. The primes were bimodal, presented as tones in combination with visual
notations that either matched or mismatched the actually presented tone. Both samples
performed better in judging unpleasant pictures over pleasant ones. In comparison with
the control musicians, the AP possessors revealed a more profound discrepancy between
the two valence conditions, and their EEG revealed later peaks at around 200 ms (P200) after
prime onset. Their performance dropped when responding to pleasant pictures preceded
by incongruent primes, especially when mistuned by one semitone. This interference was
also reflected in an EEG deflection at around 400 ms (N400) after picture onset, preceding
the behavior responses. These findings suggest that AP possessors process mistuned
musical stimuli and pleasant pictures as affectively unrelated with each other, supporting
an aversion towards out-of-tune tones in AP possessors. The longer prime-related P200
latencies exhibited by AP possessors suggest a delay in integrating musical stimuli, un-
derlying an altered affinity towards pitchelabel associations.
©2020 The Author(s). Published by Elsevier Ltd. This is an open access article under the CC
BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
*Corresponding author. Department of Medicine, University of Fribourg, Chemin du Mus
ee 8, 1700, Fribourg, Switzerland.
E-mail address: lars.rogenmoser@unifr.ch (L. Rogenmoser).
Available online at www.sciencedirect.com
ScienceDirect
Journal homepage: www.elsevier.com/locate/cortex
cortex 135 (2021) 285e297
https://doi.org/10.1016/j.cortex.2020.11.020
0010-9452/©2020 The Author(s). Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://
creativecommons.org/licenses/by-nc-nd/4.0/).

1. Introduction
Absolute pitch (AP), the ability to identify the chroma of a
tone without the aid of any reference (Takeuchi &Hulse,
1993), is sparsely distributed in the population (approximate
<1%) but yet bears phylogenetic and ontogenetic significance:
The phylogenetic argument relies on the fact that hearing
with a preference for absolute over relational cues appears to
be conserved in humans (Levitin &Rogers, 2005) given that
songbirds fail to demonstrate octave generalization while
experiments confirm this particular skill in rhesus monkeys
(Cynx, 1993;Wright, Rivera, Hulse, Shyan, &Neiworth, 2000).
Some ontogenetic evidence suggest that humans prefer ab-
solute auditory cues during early infancy, shifting towards a
preference for relational ones as they mature (Saffran &
Griepentrog, 2001; for a contradictory finding see Plantinga
&Trainor, 2005). A genetic component in combination with
specific learning factors such as early music engagement and
exposure to language, and possibly the presence of a partic-
ular brain anatomy prevent the maturation-related decline in
the ability of processing pitches absolutely and thus account
for AP acquisition (Deutsch, Henthron, Marvin, &Xu, 2006;
Gregersen, Kowalsky, Kohn, &Marvin, 1999,2001). Similar to
the successful language acquisition that requires early lan-
guage exposure during a child’s development (Newport,
1990), AP may only emerge on condition that music engage-
ment takes place during a critical period, a maturational
stage at which cognitive processing still functions in a rather
unidimensional mode and the brain is still highly malleable
and responsive to environmental inputs (Chin, 2003;Russo,
Windel, &Cuddy, 2003). As a result, adult AP possessors
show an increased left-sided asymmetry of the planum
temporale (Keenan, Thangaraj, Halpern, &Schlaug, 2001;
Schlaug, J€
ancke, Huang, &Steinmetz, 1995;Wilson, Lusher,
Wan, Dudgeon, &Reutens, 2009), which is likely due to the
reduced size on the right side (Keenan et al., 2001;
Wengenroth et al., 2013). This structural asymmetry might
create a functional dominance directing auditory stimuli to
categorization centers within the left temporal lobe such as
the superior temporal sulcus (J€
ancke, Langer, &H€
anggi, 2012;
Loui, Li, Hohmann, &Schlaug, 2010). A further brain structure
related to AP is the posterior dorsal frontal cortex. This brain
region might potentially drive conditional associative mem-
ory and, in the context of AP, be responsible for the process
underlying the association between categorized pitches and
verbal labels or other sensorimotor codes (Zatorre &Beckett,
1989;Zatorre, Perry, Beckett, Westbury, &Evans, 1998). In AP
possessors, within this left-sided frontotemporal network an
interplay is detectable even at rest in electroencephalography
(EEG) recordings (Elmer, Rogenmoser, Ku
¨hnis, &J€
ancke,
2015), suggesting functional optimization that may enable
the automatic nature of AP. Furthermore, right-sided net-
works comprising auditory and non-auditory structures have
been reported (Kim &Kn€
osche, 2016,2017;Wengenroth et al.,
2013), possibly mediating AP perception (Wengenroth et al.,
2013). Automaticity is, however, accompanied by the diffi-
culty in suppression, as many studies using interference
tasks have demonstrated, revealing a drop in identification
performance under incongruent trial conditions among AP
possessors (Akiva-Kabiri &Henik, 2012;Itoh, Suwazono,
Arao, Miyazaki, &Nakada, 2005;Rogenmoser, Arnicane,
J€
ancke, &Elmer, 2020;Schulze, Mueller, &Koelsch, 2013;Ziv
&Radin, 2014). Further studies revealed that in comparison
to non-AP (NAP) participants, AP possessors are disadvan-
taged in recognizing transposed melodies or in identifying
musical intervals within out-of-tune contexts (Miyazaki,
1993;Miyazaki &Rakowski, 2002). AP possessors incur more
cognitive load for processing out-of-tune tones, this certain
sensitivity towards mistuned tones increasing the later the
onset of musical training takes place during childhood
(Rogenmoser, Elmer, &J€
ancke, 2015). From an affective
perspective, this sensitivity is corroborated by anecdotal ev-
idence suggesting that AP possessors become agitated when
exposed to out-of-tune tones (Levitin &Rogers, 2005). Here,
we investigated this so-far unexplored phenomenon in AP,
which we refer to as auditory aversion. Understanding the
mechanisms underlying auditory aversion might advance
research on cognitive-affective development. It is further
crucial given that some AP possessors report a distortion in
frequency perception as a consequence of aging (Athos et al.,
2007;Vernon, 1977). Thus, aversion would no longer exclu-
sively be triggered externally by certain out-of-tune tones but
additionally internally by perceiving in-tune tones as out-of-
tune. In the present study, we measured the affective re-
actions in response to musical stimuli in a sample of AP
possessors and in one of matched control participants by
recording EEG during the performance of a task underlying an
affective priming paradigm (Fazio, 2001). This particular
paradigm has turned out to be promising in capturing the
automatic processing of emotional experiences (Fazio, 2001;
Klauer, 1997), even though semantic network effects (i.e.,
semantic priming) may contribute to the measurement (Eder,
Leuthold, Rothermund, &Schweinberger, 2012;Steinbeis &
Koelsch, 2008,2011). The affective priming procedure in-
volves the sequential presentation of two valenced stimuli,
namely the preceding so-called primes and the following so-
called targets. The task is to handle the targets whereby the
preceding primes are either affectively related or affectively
unrelated to them. It is well documented that affectively
unrelated prime-target pairs induce interference that varies
as a function of the extent of involved emotional experience
(Fazio, 2001). In EEG, this interference is reflected by a
negative-going deflection around 400 ms after target-onset,
which is referred to as the N400 (Kutas &Hillyard, 1980;
Steinbeis &Koelsch, 2008,2011;Zhang, Li, Gold, &Jiang, 2010).
EEG has an excellent time resolution and has been promising
in capturing the early perceptual and cognitive processes
underlying AP (Burkhard, Elmer, &J€
ancke, 2019;Elmer,
Sollberger, Meyer, &J€
ancke, 2013;Greber, Rogenmoser,
Elmer, &J€
ancke, 2018;Rogenmoser et al., 2015). In the pre-
sent study, the participants were instructed to judge as
quickly and accurately as possible whether pictures (targets)
were either pleasant or unpleasant by pressing one of two
response buttons. In the experimental condition, these tar-
gets were preceded by musical stimuli (primes) which were
bimodal, comprising visually presented notations in combi-
nation with auditorily presented tones. The auditory
cortex 135 (2021) 285e297286

counterpart either matched the notations (congruent) or
deviated from them (incongruent) in a higher semitone (i.e.,
sharp) or quarter-tone. Based on the assumption that out-of-
tune tones trigger aversion in AP possessors, we predicted an
interference in the AP possessors’ response behavior
accompanied by a greater N400 deflection in conditions in
which the incongruent primes preceded pleasant targets.
2. Materials and methods
2.1. Participants
We report how we determined our sample size, all data ex-
clusions (if any), all inclusion/exclusion criteria, whether in-
clusion/exclusion criteria were established prior to data
analysis, all manipulations, and all measures in the study.
Inclusion (i.e., possession or non-possession of AP) and
exclusion criteria (i.e., history of neurological, psychiatric and
audiological disorders) were established a priori. The open
source software G*Power (Faul, Erdfelder, Lang, &Buchner,
2007) was used to estimate the sample sizes required for the
detection of within-between interaction effects.
Twenty-one AP possessors (14 males) and 21 matched NAP
control musicians (13 males) participated in this study. The
two samples were comparable regarding age (t
40
¼.80,
P¼.428, d¼.25), general cognitive capability (t
39.91
¼.677,
P¼.502, d¼.21) as measured by the ShipleyeHartford Retreat
Scale (Shipley, 1940), and comparable regarding the distribu-
tion of handedness (one left-hander in each sample) and of
the sexes (Х
21
¼.10, P¼.747). Both AP and NAP participants
commenced their musical training at a comparable age range
(t
40
¼1.64, P¼.106, d¼.51) and trained for a comparable
number of hours per day (t
40
¼.45, P¼.656, d¼.14) and years
in total (t
40
¼.11, P¼.910, d¼.04). Across both samples, the
participants were comparably skilled in music performance,
as confirmed using pitch and rhythm discrimination tasks
underlying psychophysical adaptive staircase procedures
(Fujii &Schlaug, 2013;Loui, Alsop, &Schlaug, 2009). Regarding
pitch discrimination, the minimum detectable frequency dif-
ference around the center frequency of 500 Hz was measured.
The two samples performed at comparable pitch discrimina-
tion thresholds (t
40
¼1.18, P¼.244, d¼.36). Regarding rhythm
discrimination (i.e., Beat Interval Test (Fujii &Schlaug, 2013)),
sequences of 21 woodblock tones with varying interval
lengths were presented. The sensory threshold for discrimi-
nating temporal changes as well as the performance in tap-
ping in synchrony while adapting to temporal changes were
measured. The two groups performed at comparable rhythm
discrimination thresholds (t
31.83
¼.011, P¼.992, d¼.00) and
with a comparable tapping synchronization ability (t
40
¼.67,
P¼.507, d¼.21). The values on characteristics and musical
background are reported in Table 1.
All participants gave written informed consent, and the
study was approved by the institutional review board of Beth
Israel Deaconess Medical Center. No part of the study pro-
cedures or analyses was pre-registered prior to the research
being conducted.
2.2. Absolute pitch (AP) verification
AP was confirmed using an established pitch-labeling test
(Keenan et al., 2001) in which 52 trials were presented. Each trial
contained one computer-generated sine wave tone (500-ms
duration with a 50-ms rise and decay time) with a funda-
mental frequency covering one octave from 370 Hz (F#3) to
739.97 Hz (F#4) in the equal-tempered Western scale. The par-
ticipant’s task was to label each pitch by writing down the letter
name on an answer sheet upon hearing each tone. The inter-
tone interval was 2 sec. Semitone errors were counted as
incorrect responses. The AP possessors performed (% correct
responses) considerably better on this test (mean correct: 94.15,
SD ¼10.05) than the NAP participants (mean correct 5.77,
SD ¼6.26; t
40
¼34.21, P¼3.08 10
31
,d¼10.56). NAP partici-
pants did not perform better than chance level (8.33%;
t
20
¼1.89, P¼.075). The individual scores are depicted in Fig. 1.
2.3. Experimental task and stimulus material
To measure the participants’ affective reaction in response to
musical stimuli, EEG was recorded during performance of a
task underlying an affective priming paradigm. In this task,
the participants were instructed to judge as quickly and
accurately as possible whether pictures (targets) preceded by
musical stimuli (primes) were either pleasant or unpleasant
by pressing one of two response buttons. The targets were 144
pictures (see Appendix) taken from the International Affective
Picture System (Lang, Bradley, &Cuthbert, 2008) on the basis
of normative ratings (9-point scales) regarding the two affec-
tive dimensions of valence (1 ¼most unpleasant; 9 ¼most
pleasant) and arousal (1 ¼most calm; 9 ¼most aroused). Half
of the picture collection was pleasant (M¼7.60, SD ¼.41) while
the other was unpleasant (M¼2.66, SD ¼.79; t
107.21
¼46.91,
P¼2.76 10
73
,d¼7.82), but across both valences the pic-
tures were controlled for arousal (unpleasant pictures:
M¼5.59, SD ¼.71; pleasant pictures: M¼5.41, SD ¼.95;
t
131.31
¼1.33, P¼.185, d¼.21). The preceding primes were
Table 1 eCharacteristics and data on the musical
background of both samples.
AP NAP
Age (years) 22.38 (4.86) 23.47 (3.94)
Cognitive capability (IQ scores) 115.24 (8.62) 113.47 (8.22)
Age at commencement of musical
practice (years)
6.14 (2.44) 7.43 (2.60)
Practice intensity (hours per day) 2.92 (1.82) 2.67 (1.79)
Duration of musical training (years) 16.24 (5.70) 16.05 (5.08)
Pitch discrimination threshold (Hz) at
500 Hz
4.89 (5.43) 7.55 (8.77)
Rhythm; perceptual discrimination
threshold (ms)
1.53 (1.72) 1.54 (3.01)
Rhythm; tapping synchronization
threshold (ms)
1.61 (3.56) 2.39 (4.03)
Listed are the means with the standard deviations in brackets. All
independent-samples t-test calculated for each variable revealed
values of P>.1. AP: Absolute pitch, NAP: non-absolute pitch.
cortex 135 (2021) 285e297 287

bimodal, comprising visually presented notations in combi-
nation with auditorily presented pure tones. The set of pre-
sented notations comprised the scale of 12 subsequent notes
ranging from F#4 to F5. The auditory counterpart either
matched the notations or deviated from them in a higher
semitone (sharp) or quarter-tone. The 144 targets were pre-
sented 4 times in total; once per each matching condition and
additionally once as control condition without any prime.
Since the tuning of the musical stimulus is expected to impact
the affective reaction exceptionally in AP participants, the
particular prime-target pairs in this experimental set-up
reflect either affectively unrelated or affectively related con-
ditions. Predicated on this constellation, 6 particular prime-
target pair conditions result (see Table 2), within which each
of the 12 scaled primes were paired 4 times with different
targets (72 times per valence). The prime-target pairs as well
as the order of trials were pseudorandomized. Each trial began
with a fixation cross for gaze stabilization that was presented
for a duration of 500 ms. The following primes were presented
with a duration of 500 ms and the targets with one of 800 ms
while the prime-target interval was of 200 ms. The inter-trial
interval was jittered between 1000 and 1500 ms. The proced-
ure of one trial is illustrated in Fig. 2. The auditory stimuli
included 10 ms of fade-in/out phases and were delivered via
Sennheiser HD 205 headphones at the sound pressure level of
80 dB. The visual material was shown in the center of a PC
laptop monitor. Stimuli presentation and behavior recordings
were controlled by the software PsyTask. Before starting the
actual experiment, the participants were given 8 practice tri-
als to get familiar with the task.
2.4. EEG recording
EEG was recorded via 19 channels (Ag/AgCl electrodes, BEE
Medic GmbH) placed according to the “10e20”system with the
following electrode locations: Fp1, Fp2, F7, F3, Fz, F4, F8, T3, C3,
Cz, C4, T4, T5, P3, Pz, P4, T6, O1, O2. A further Ag/AgCl elec-
trode embedded in the cap horizontally centered between Cz
and Fz served as ground and two Ag/AgCl electrodes clipped
on both earlobes as reference. EEG recording was done with an
amplifier (EEG NeuroAmp x23) and the ERPrec software. The
signal was recorded with a sampling rate of 500 Hz and a
bandpass filter of .1e100 Hz. Impedances were kept below 5 kU
using conductive gel (Elecro-Gel).
2.5. Data analyses
The conditions of our ethics approval do not permit public
archiving of anonymised study data. Readers seeking access to
the data should contact the first or last author. Access will be
granted following completion of a formal data sharing agree-
ment. Legal copyright restrictions prevent public archiving of
the IAPS stimuli used in the current study which can be ob-
tained from (Lang et al. 2008), and see Appendix for stimulus
catalogue numbers. Presentation code, absent the IAPS stim-
uli, may be freely downloaded from https://osf.io/a6qfd.
2.5.1. Preprocessing
Raw EEG data were imported into EEGLAB v.13.2.1 (Delorme &
Makeig, 2004), an open source toolbox running under MAT-
LAB. The data were down-sampled to 250 Hz, band-pass
filtered at 1e20 Hz and re-referenced to the mean of the T3
and T4 electrodes. Epochs were created ranging from 200 to
500 ms after prime-onset and from 200 to 1000 ms after
target-onset, respectively. A baseline correction relative to the
200 to 0 ms pre-stimulus time period was applied. Epochs
contaminated by unsystematic artifacts were rejected. Sys-
tematic artifacts corresponding to non-cortical sources were
removed by using independent component analysis (Jung et
al., 2000). Condition-wise, event-related potentials (ERP)
were calculated by averaging the epochs belonging to the
respective condition. The central channels (F3, Fz, F4, C3, Cz,
C4, P3, Pz, P4) were pooled together. The maxima/minima
peak values and their timepoints (latencies) were extracted
Table 2 ePrime-target pair conditions.
Affectively related
1. Congruent prime þpleasant target
2. Incongruent prime (semitone) þunpleasant target
3. Incongruent prime (quarter-tone) þunpleasant target
Affectively unrelated
4. Congruent prime þunpleasant target
5. Incongruent prime (semitone) þpleasant target
6. Incongruent prime (quarter-tone) þpleasant target
Fig. 1 ePitch-labeling performance. Depicted are the
individual scores (%) achieved by the participants with
absolute pitch (AP, blue circles, N¼21) and participants
without absolute pitch (NAP, red circles, N¼21) from the
pitch-labeling test. The dotted line represents the baseline
at 8.3%.
cortex 135 (2021) 285e297288

from the N100eP200 complex of the prime-induced ERPs. The
minima peak values and their latencies of the negative-going
deflection centered around 400 ms after target-onset (N400)
were extracted from the difference waves, calculated between
the two ERPs (affectively unrelatedeaffectively related)
belonging to the same matching condition. In case of the ERPs
induced by targets without preceding primes, minima values
and their latencies of the negative-going deflection centered
Fig. 2 eSchematic representation of the task. Each trial began with a fixation cross (A) that was presented on the monitor for
500 ms, followed by a prime (B) for a duration of 500 ms. The prime was bimodal, comprising visually presented notations in
combination with auditorily presented pure tones via earphones. The auditory counterpart either matched the notations
(congruent) or deviated from them (incongruent) in a higher semitone or quarter-tone. Afterwards, the monitor turned blank
for 200 ms (C). Then, the target (D) followed, presented for 800 ms. The target was either a pleasant or an unpleasant picture
taken from the International Affective Picture System. The participants were instructed to indicate as quickly and accurately
as possible the valence of the target by pressing on either the right (pleasant) or left (unpleasant) mouse button. The next
trial followed after the monitor turned blank for a duration jittered between 1000 and 1500 ms (E).
cortex 135 (2021) 285e297 289

around 400 ms after target-onset were extracted from the
difference waves, subtracted from each other in both di-
rections (pleasanteunpleasant and vice versa).
2.5.2. Statistical analyses
The amplitudes and the latencies as well as the mean reaction
times (RT) and accuracy scores obtained from each condition
by each participant were imported into the SPSS software for
statistical analyses. The mean RTs as well as the accuracy
scores were subjected to three-way mixed analyses of vari-
ance (ANOVA) with group (AP, NAP) as between-factor and
two within-factors, namely “matching condition”with four
levels (no prime, congruent, incongruent semitone, incon-
gruent quarter-tone) and “valence”with two levels (pleasant,
unpleasant). The amplitudes and latencies of the ERPs (N100,
P200, N400) were subjected to two-way mixed ANOVAs with
group (AP, NAP) as between-factor and “matching condition”
as within-factor with three levels (congruent, incongruent
semitone, incongruent quarter-tone). The amplitudes and la-
tencies of the negative-going waves of the control difference
waves were subjected to independent-samples t-tests. Sta-
tistical analyses were adjusted for non-sphericity using
GreenhouseeGeiser Epsilon when equal variances could not
be assumed. Group-interaction effects were further inspected
with post-hoc analyses. The relation between group-specific
ERP findings and the AP scores achieved from the pitch-
labeling test were investigated using Pearson’s
productemoment correlations. Correlation analyses as well
as t-tests were run two-tailed. Effect size measures were
calculated; namely the Cohen’s dfor t-tests and Generalized
Eta Squared (h
2G
) for ANOVAs.
3. Results
3.1. Behavioral findings
The mean reaction times (RT) and accuracy scores achieved at
each condition by both samples are depicted in Fig. 3.Both
samples responded not only faster (F
1, 40
¼63.19,
P¼9.26 10
10
,h
2G
¼.071) but also with higher accuracy (F
1,
40
¼12.64, P¼9.87 10
4
,h
2G
¼.074) to unpleasant targets than
to pleasant ones. The matching condition of the primes had a
group-independent influence on the RT (F
1.18, 47.07
¼22.06,
P¼910
6
,h
2G
¼.027). The two samples responded with
comparable RT, as no group effect (F
1, 40
¼2.90, P¼.096,
h
2G
¼.06) or any group-interaction effects (matching x group:
F
1.18, 47.07
¼1.31, P¼.265, h
2G
¼.002; valence x group: F
1,
40
¼2.42, P¼.128, h
2G
¼.003) were revealed. Regarding the
accuracy scores, the two samples differed in general (group
effect: F
1, 40
¼4.10, P¼.050, h
2G
¼.055) but differed also as
interaction with the matching condition (F
1.53, 61.44
¼3.83,
P¼.037, h
2G
¼.012) and the valence factor (F
1, 40
¼4.28, P¼.045,
h
2G
¼.026). Post-hoc pairwise comparisons (Bonferroni-
adjusted at the level of a¼.05for8paired-samplest-tests) of
the accuracy scores between the valences at all matching
conditions across both samples revealed a significant differ-
ence only in the AP sample at the incongruent semitone con-
dition (t
20
¼3.16, Bonferron i-adjusted P¼.039, d¼.69).
3.2. EEG findings
3.2.1. N100eP200 complex
The ERP induced by the primes are depicted in Fig. 4 and their
values are plotted in Fig. 5. Regarding the amplitude and the
latency of the N100, no group (amplitude: F
1, 40
¼.59, P¼.446,
h
2G
¼.014; latency: F
1, 40
¼.41, P¼.526, h
2G
¼.008), nor condition
differences (amplitude: F
2, 80
¼2.89, P¼.061, h
2G
¼.004; latency:
F
2, 80
¼2.35, P¼.102, h
2G
¼.012) nor groupecondition in-
teractions (amplitude: F
2, 80
¼2.748, P¼.070, h
2G
¼.004; latency:
F
2, 80
¼.769, P¼.467,h
2G
¼.004) were found. Within the P200, AP
possessors peaked later than NAP participants (F
1, 40
¼5.41,
P¼.025, h
2G
¼.093) but with comparable height (F
1, 40
¼.35,
P¼.559, h
2G
¼.008). TheP200 latencies correlated positively with
the participants’ AP scores achieved from the pitch-labeling test
(congruent: r
40
¼.37, P¼.017; incongruent semitone: r
40
¼.31,
P¼.042; incongruent quarter-tone: r
40
¼.32, P¼.041). The
matching condition, however, had no impact on the P200
Fig. 3 ePerformances achieved from the affective priming task. The mean RTs (left) as well as the accuracy scores (right) are
depicted for each condition for the absolute pitch (AP, blue, N¼21) and the non-absolute pitch (NAP, red, N¼21) samples.
The solid lines present pleasant targets and the dashed lines present unpleasant targets. The bars depict standard errors.
Two-tailed Bonferroni-adjusted *P<.05.
cortex 135 (2021) 285e297290

amplitudes (F
2, 80
¼.32, P¼.649, h
2G
<.001) nor on the P200
latencies(F
2, 80
¼1.02, P¼.366, h
2G
¼.006), and nor did it interact
with the samples(amplitude: F
1.42, 56.63
¼.59, P¼.531, h
2G
¼.002;
latency: F
1.68, 67.35
¼.11, P¼.862, h
2G
¼.001) in this regard.
3.2.2. N400
The ERPs induced by the targets including the difference
waves calculated between the two valence factors referring to
each prime matching condition are depicted in Fig. 6. The
values of the difference waves are plotted in Fig. 7. The
negative-going deflection of the difference waves calculated
between the affectively related and affectively unrelated pair
conditions varied as a function of the matching condition
(amplitude: F
1.63, 65.36
¼12.13, P¼1.03 10
4
,h
2G
¼.148; la-
tency: F
1.66, 66.43
¼12.73, P¼6.3 10
5
,h
2G
¼.178). No group
differences were found in the amplitudes (F
1, 40
¼2.30,
P¼.137, h
2G
¼.024) nor in the latencies (F
1, 40
¼1.68, P¼.203,
h
2G
¼.013). The latencies did not reveal a group-interaction
effect (F
1.66, 66.43
¼1.68, P¼.068, h
2G
¼.048), whereas the
amplitudes did (F
1.63, 65.36
¼5.10, P¼.013, h
2G
¼.068). This
effect was not impacted by the participants’ response
behavior, as their RTs did not correlate with the amplitudes;
neither in the AP (“un-/pleasant”x“congruent”:r
21
<.2, P>.5;
“un-/pleasant”x“incongruent semitone”:r
21
>.3, P>.2;
“un-/pleasant”x“incongruent quarter-tone”:.1 >r
21
<.1, P>
.6) nor in the NAP group (“un-/pleasant”x“congruent”:r
21
<.2,
P>.4; “un-/pleasant”x“incongruent semitone”:r
21
>.03,
P>.9; “un-/pleasant”x“incongruent quarter-tone”:r
21
>.4,
P>.07). The AP scores, however, correlated negatively with
the “incongruent semitone”amplitudes (r
40
¼e.35, P¼.025)
and trended negatively with the “incongruent quarter-tone”
amplitudes (r
40
¼e.29, P¼.060) but correlated positively with
the “congruent”amplitudes (r
40
¼.31, P¼.044). Post-hoc
comparisons (Bonferroni-adjusted at the level of a¼.05 for 6
paired-samples t-tests) of amplitudes between all matching
conditions across both samples revealed significant differ-
ences only in the AP possessors between the congruent and
incongruent semitone conditions (t
20
¼4.85, Bonferroni-
adjusted P¼5.88 10
4
,d¼1.64) and between the
congruent and incongruent quarter-tone conditions
(t
20
¼3.20, Bonferroni-adjusted P¼.028, d¼1.14).
Independent-samples t-tests across the two samples revealed
a difference at the incongruent semitone condition (t
40
¼2.37,
uncorrected P¼.023, d¼.73).
3.2.3. Control difference waves
Comparing the negative-going deflections of the difference
wave calculated by subtraction of the ERPs induced by the
unpleasant from the ERPs induced by the pleasant targets
revealed no group difference in amplitude (t
40
¼.09, P¼.930,
d¼.03) or latency (t
40
¼.14, P¼.891, d¼.04). Vice versa, the
negative-going deflections of the difference wave calculated
by subtraction of the ERPs induced by the pleasant from the
ERPs induced by the unpleasant targets revealed no group
difference in amplitude (t
40
¼1.18, P¼.244, d¼.37) or latency
(t
39.37
¼.36, P¼.720, d¼.11). The amplitudes also did not
correlate with the AP scores (pleasanteunpleasant: r
40
¼e.06,
P¼.714; unpleasantepleasant: r
40
¼.17, P¼.294). The ERPs
induced by the targets including the difference waves calcu-
lated between the two valence factors without preceding
primes are also depicted in Fig. 6. The values of the difference
waves are plotted in Fig. 7.
Fig. 4 eGroup-averaged ERPs in response to the primes. On top, the ERPs averaged across the absolute pitch (N¼21) and the
non-absolute pitch (N¼21) samples are displayed. Below, the respective current distribution of the scalps are displayed
derived from the peaks of the N100 (first) and P200 (second).
cortex 135 (2021) 285e297 291

4. Discussion
The participants processed the unpleasant pictures more
efficiently than the pleasant ones, as evident from the faster
and more accurate responses in the unpleasant target condi-
tions. These results mirror the well documented finding that
unpleasant stimuli induce stronger affective effects than
pleasant ones, supporting the idea that the processing system
is sensitized to aversion ( €
Ohman &Mineka, 2001). From an
evolutionary perspective, this system has adaptive value
given that it favors avoidance/defense over approach
response behavior. This discrepancy, referred to as negativity
bias in the literature, underlies the recruitment of the
subcortical route that involves the thalamus and amygdala,
leading to a more rapid orienting of attention when exposed
with aversive, potentially harmful events (Carreti
e, Mercado,
Tapia, &Hinojosa, 2001;Ito, Larsen, Smith, &Cacioppo, 1998;
LeDoux, 1995).
Compared to NAP participants, the AP possessors judged
the targets with accuracy scores varying as function of the
prime-target pair condition. They underperformed strongest
in the condition in which the pleasant targets were preceded
by incongruent primes that were one semitone off. In line with
the behavioral results, the N400 amplitudes differed between
the two samples in dependence of the matching condition.
This interaction effect was due to the conditions in which the
primes were incongruent, particularly in terms of one semi-
tone. The interference in the response behavior together with
the N400 effects occurring in the conditions in which the
pleasant targets were preceded by incongruent primes
confirm that the AP possessors processed these prime-target
pairs as affectively not in line with each other, supporting
an aversion towards out-of-tune tones in AP possessors.
Furthermore, these findings provide evidence that aversion is
experienced more intensely when detuning reaches one
semitone, possibly underlying the categorical nature of AP
(Harris &Siegel, 1975). Instead of relying on a sensory coding
strategy, AP has shown to rather be driven by a categorical
perception mechanism (Siegel, 1974). Within the critical in-
fantile period during which AP acquisition takes place,
Western pitchelabel associations are formed in units of
Fig. 5 eValues of the prime-induced ERPs. Plotted are the averaged amplitudes (top) and latencies (bottom) from the
absolute pitch (N¼21) and non-absolute pitch (N¼21) sample across all prime matching conditions. The bars depict
standard errors. A: N100. B: P200.
cortex 135 (2021) 285e297292

semitones, leading to a susceptibility specified for these highly
familiarized units. Thus, in comparison with within-
categorical violations (e.g., quarter-tone), between-categori-
cal violations (i.e., semitone) intensify the interference. In AP
possessors, detuning especially across semitone categories
not only impairs music performance (Miyazaki 1993,1995;
Miyazaki &Rakowski, 2002;Siegel, 1974) but also, as the pre-
sent findings suggest, affect the affective responses. These
findings are in line with the more general relation between
categorization difficulty (i.e., low fluency processing) and
negative affect (Reber, Schwarz, &Winkielman, 2004). How-
ever, it shall be noted that here we only explored the affective
effects in response to a limited range of detuning degrees in
only one (i.e., sharp) direction by using an indirect measuring
approach. The advantage of the affective priming paradigm
lies in that it bypasses introspection and captures automatic
emotional responses outside of awareness (Bargh &Morsella,
2008;Fazio, 2001;Greenwald, Draine, &Abrams, 1996),
avoiding potential top-down biases. Nevertheless, the partic-
ipants were not asked to directly rate the primes. Thus,
further research is required to validate and generalize our
findings on auditory aversion covering also detuning degrees
in the “flat”direction.
Regarding the maturational aspect of AP, these findings are
revealing given that they expand the interlinkage between AP
and other developmental conditions in which AP sometimes
co-exists; such as congenital synesthesia (Bouvet et al., 2014;
Gregersen et al., 2013;H€
anggi, Beeli, Oechslin, &J€
ancke, 2008;
J€
ancke, Rogenmoser, Meyer, &Elmer, 2012) and blindness
(Hamilton, Pascual-Leone, &Schlaug, 2004), autism spectrum
disorder (Bonnel et al., 2003;Brenton, Devries, Barton,
Minnich, &Sokol, 2008;Heaton, 2003;Heaton, Davis, &
Happ
e, 2008), Williams syndrome (Lenhoff, Perales, &
Hickok, 2001). A range of auditory abnormalities are reported
in Williams syndrome such as hyper- and odynacusis, audi-
tory allodynia or attraction to certain everyday life sounds
(Levitin, 2005;Levitin, Cole, Lincoln, &Bellugi, 2005). Likewise,
hyperacusis and hypersensitivity to certain, especially com-
plex, sounds have been described in autism spectrum disorder
(O’Connor, 2012;Rosenhall, Nordin, Sandstr€
om, Ahls
en, &
Fig. 6 eGroup-averaged ERPs induced by the targets including the difference waves. The difference waves are calculated
between the two ERPs (affectively unrelatedeaffectively related) belonging to the same matching condition. In case of the
control condition (no primes), differences waves were calculated by subtracting the ERPs induced by the two targets from
each other in both directions (pleasant-unpleasant and vice versa). The current distributions derived from deflection-peak
of each difference curve are plotted as scalp maps. Absolute pitch (left; N¼21), non-absolute pitch (right; N¼21).
cortex 135 (2021) 285e297 293

Gillberg, 1999). Furthermore regarding common characteris-
tics, AP possessors have shown to exhibit more autistic traits
(Brown et al., 2003;Dohn, Garza-Villarreal, Heaton, &Vuust,
2012;Wenhart, Bethlehem, Baron-Cohen, &Altenmu
¨ller,
2019) and rather an “autistic-typical”cognitive style, namely
the processing of local information in expense of global ones
(Foxton et al., 2003;Mottron, Peretz, &Menard, 2000), which is
observable not only in the auditory but also in the visual
domain (Chin, 2003;Costa-Giomi, Gilmour, Siddell, &
Lefebvre, 2001;Wenhart &Altenmu
¨ller, 2019;Wenhart,
Hwang, &Altenmu
¨ller, 2019;Ziv &Radin, 2014). Imaging
studies pointed out that specific hyperfunctioning underlie
local hyperconnectivity between the involved areas in the
brain; a further overlapping feature not only documented in
AP but also synesthesia or other savant skills (Brauchli,
Leipold, &J€
ancke, 2019;Elmer et al., 2015;H€
anggi et al., 2008;
J€
ancke, Langer et al., 2012;Loui et al., 2010;Mottron et al.,
2013). A similar phenomenon to the AP-specific aversion
confirmed in this study has been described in congenital
synesthesia, a condition that overlaps phenotypically and
genotypically with AP (Gregersen et al., 2013;Loui, Zamm, &
Schlaug, 2012a) and also co-occurs with autism (Bouvet et
al., 2014;Mottron et al., 2013;Neufeld et al., 2013). In
particular, synesthetes react aversively when exposed to
inducereconcurrent pairs mismatching their familiarized
idiosyncratic own (Callejas, Acosta, &Lupi
a~
nez, 2007).
Beyond the interference reflected in the present data
pattern, the AP possessors showed overall poorer accuracy
scores specifically when judging pleasant pictures. A group
difference in mere affective visual processing, however, was
not supported by the EEG results, as the control difference
curves did not differ between the two samples. Thus, the
question raises to what extent AP possessors differ in affective
processing not limited to out-of-tune responses or beyond the
auditory dimension in general. As already mentioned, non-
auditory differences between AP and NAP participants have
been reported in visual cognition, such as in the ability for
spatial and binocular rivalry resolution (Costa-Giomi,
Gilmour, Siddell, &Lefebvre, 2001;Kim, Blake, Lee, &Kim,
2017). Worth mentioning, a further linkage between the vi-
sual system and AP is reflected in a higher incidence rate of AP
among congenitally blind (Hamilton et al., 2004). In the pre-
sent study, compared to NAP participants the AP possessors
exhibited longer P200 latencies in response to the auditory-
visual primes regardless of the matching condition. An ERP
modulation within this relatively early component has been
Fig. 7 eValues of the difference curves. Plotted are the group-averaged amplitudes (top) and latencies (bottom) extracted
from deflection at the N400 across all prime matching (A) and control (B) conditions. Absolute pitch (N¼21), non-absolute
pitch (N¼21). The bars depict standard errors. Two-tailed Bonferroni-adjusted ***P<.001, *P<.05, and unadjusted
#P<.05.
cortex 135 (2021) 285e297294

ascribed to the intrinsic relevance of the stimulus and to af-
fective integration of multiple connotations (Carreti
e et al.,
2001;Spreckelmeyer, Kutas, Urbach, Altenmu
¨ller, &Mu
¨nte,
2006). Thus, the findings provide evidence that AP posses-
sors exhibit a delay in affectively integrating musical stimuli,
possible due to altered affinity to them. In earlier studies, AP
possessors differed from NAP participants in brain activation
during music listening (Loui et al., 2012a) and in response to
the performance of an emotional rating task during music
listening (Loui et al., 2012a;Loui, Zamm, &Schlaug, 2012b). In
particular, AP possessors exhibited increased functional acti-
vation as well as greater functional interconnections with
strongest effects around the left superior temporal gyrus
including brain areas involved in multisensory integration as
well as in emotion and reward processing (Loui et al., 2012b).
Altogether, some previous studies (Loui et al., 2012a,2012b)
and the present findings suggest that the differences under-
lying the affective processing system in AP possessors may go
beyond auditory aversion.
Acknowledgements
This work was supported by the Swiss National Science
Foundation (P1ZHP1_158642, P2ZHP1_168587, P300P1_177744
to LR and 138668, 163149 to LJ) and the National Institute on
Deafness and Other Communication Disorders (RO1-
DC009823 to GS).
Appendix
The catalog numbers for the pictures used in this study are as
follows.
Pleasant: 1440, 1460, 1463, 1510, 1600, 1710, 1750, 1920,
1999, 2040, 2045, 2050, 2058, 2070, 2071, 2080, 2091, 2150, 2151,
2160, 2165, 2170, 2208, 2209, 2216, 2260, 2274, 2311, 2332, 2340,
2341, 2345, 2347, 2395, 2550, 2660, 4622, 4660, 5470, 5480, 5600,
5621, 5629, 5829, 5830, 5833, 7200, 7230, 7330, 7502, 7580, 8030,
8080, 8158, 8170, 8178, 8179, 8180, 8185, 8186, 8190, 8200, 8206,
8210, 8370, 8400, 8420, 8470, 8490, 8496, 8499, 8501.
Unpleasant: 1033, 1110, 1111, 1120, 1202, 1220, 1271, 1274,
1280, 1930, 1931, 1932, 2053, 2095, 2120, 2205, 2490, 2590, 2661,
2717, 2750, 2800, 2811, 2900, 2981, 3005.1, 3015, 3030, 3064,
3071, 3100, 3140, 3160, 3181, 3220, 3225, 3230, 3300, 3301, 3350,
6020, 6021, 6520, 6555, 6563, 7380, 9006, 9007, 9008, 9040, 9042,
9043, 9075, 9102, 9140, 9182, 9183, 9265, 9301, 9320, 9325, 9373,
9405, 9490, 9530, 9561, 9570, 9571, 9582, 9584, 9592, 9594.
CRediT authorship contribution statement
Lars Rogenmoser: Conceptualization, Methodology, Software,
Formal analysis, Investigation, Data Curation,
WritingeOriginal Draft, WritingeReview &Editing, Visualiza-
tion, Funding acquisition. H. Charles Li: Software. Lutz J€
ancke:
Resources, WritingeReview &Editing, Supervision, Funding
acquisition. Gottfried Schlaug: Resources, Data Curation,
WritingeReview &Editing, Supervision, Project administra-
tion, Funding acquisition.
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