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Review
Playing music to animals: an interdisciplinary approach to improving
our understanding of animals' responses to music
Buddhamas P. Kriengwatana
a
,
*
, Ruedi G. Nager
b
, Alex South
c
,
d
,
Martin Ullrich
e
, Emily L. Doolittle
c
a
Department of Biosystems, Division of Animal and Human Health Engineering, KU Leuven, Leuven, Belgium
b
School of Biodiversity, One Health and Veterinary Medicine, College of Biomedical, Veterinary and Life Sciences, University of Glasgow, Glasgow, U.K.
c
Royal Conservatoire of Scotland, Glasgow, U.K.
d
Centre for Social Learning and Cognitive Evolution and Sea Mammal Research Unit, University of St Andrews, Fife, U.K.
e
Nuremberg University of Music, Nuremberg, Germany
article info
Article history:
Received 22 February 2024
Initial acceptance 28 May 2024
Final acceptance 24 October 2024
Available online 4 February 2025
MS. number: 24-00103R
Keywords:
acoustic enrichment
aesthetics
animal welfare
anthropogenic noise
environmental enrichment
ethics
interdisciplinary
music
preference
Humans have profoundly changed the global soundscape. Studying how nonhuman animals respond to
music can contribute to a better understanding of the effect of sound on animals. Animals are frequently
exposed to human music, whether intentionally (for example, in laboratory settings), or unintentionally
(for example, when animals live in close proximity to humans). Although several papers examine ani-
mals' responses to music, these typically do so from a purely animal behavioural perspective, sometimes
missing relevant details about salient features of the music being played. An interdisciplinary approach
that places musical and scientific knowledge on equal footing can improve our understanding of how
animals respond to music and music-like sounds, in new and exciting ways. Here, we show with a
systematic review that crucial factors (intrinsic music properties, listener properties, playback context
and producer properties and contexts; ILPP) are not being adequately considered or reported in recently
published scientific articles on the effects of music on animals, which hinders scientific reproducibility
within this area of study. These problems are caused by improper referencing of music sources,
misunderstanding of music and unexamined assumptions about individual variation and preferences
between individuals of the same or different species. We then suggest that Berlyne’s psychobiological
theory might provide a useful framework for studying how animals respond to human-generated
sounds.
©2025 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.
As humans alter the landscape to meet our ever-increasing de-
mands for space and resources, we are profoundly changing the
global soundscape (Fritschi et al., 2011). This means a large and
growing number of free-living and managed nonhuman animals
(hereafter, animals) are surrounded by acoustic environments high
in anthrophony (i.e. human-generated sound, Krause, 2008;
Pijanowski et al., 2011), because they share the same habitat with
humans (Applebaum et al., 2023;Eurostat, 2023;Spotswood et al.,
2021). For most animals, sound is a vital source of information
about their environment and about conspecifics and/or hetero-
specifics. Anthropogenic sound clearly affects animals' behaviour
and physiology (Kunc &Schmidt, 2019). A general understanding of
why and what aspects of sounds are salient and meaningful for
animals could improve knowledge of how they respond to
anthrophony, leading to better management practices that reduce
the potentially detrimental effects of anthrophony on wild and
captive animals (Elmer et al., 2021).
Some animals actively avoid areas with high levels of anthro-
pogenic sound because anthropogenic noise itself can have nega-
tive and costly effects on their behaviour, cognition, immune and
other health status indicators (Kleist et al., 2018;Masud et al., 2020;
Slabbekoorn &Den Boer-Visser, 2006). In some cases, however,
animals may approach human-generated sounds which differ from
those of the natural soundscape. For instance, black-chinned
hummingbirds, Archilochus alexandri, prefer to nest near natural
gas wells with noise-producing compressors (Francis et al., 2009).
Utilizing animals' attraction to certain sound sources or sound-
scapes has real conservation potential, as research demonstrates
that animals settle into degraded habitats that are made to sound
like healthy, thriving habitats (Gordon et al., 2019) and are more
*Corresponding author.
E-mail address: pralle.kriengwatana@kuleuven.be (B. P. Kriengwatana).
Contents lists available at ScienceDirect
Animal Behaviour
journal homepage: www.elsevier.com/locate/anbehav
https://doi.org/10.1016/j.anbehav.2025.123074
0003-3472/©2025 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.
Animal Behaviour 221 (2025) 123074
likely to use passageways with conspecific calls present (Testud
et al., 2020). Outside of certain contexts, such as intraspecific
communication, prey and predator detection (e.g. Ryan et al., 1993;
Ryan &Tuttle, 1983), however, the difference between sounds that
animals are attracted to or avoid is not well understood.
MUSIC AND ITS RELEVANCE TO ECOLOGY AND CONSERVATION
Human music (hereafter, music), an instance of anthropogenic
sound, is only rarely considered as relevant for ecology or conser-
vation research (e.g. Hooker et al., 2023;Leduc, Nunes et al., 2021).
Yet music is present almost everywhere humans are, can inten-
tionally mimic biophony (Doolittle, 2008) and is currently used in
animal management practices (e.g. for increasing milk production,
Lemcke et al., 2021; reducing anxiety, Kinnaird &Wells, 2022).
Thus, music, whether live or recorded, pervades soundscapes in
areas where humans spend their time (i.e. live, work and partici-
pate in recreation) and could impact the animals that also spend
time in those areas.
Understanding how animals respond to music may contribute
valuable insights into how animals respond to sound in general.
This is because we can vary and test parameters in music, using
those known to be important for human perception and response
as a starting point. Music can be characterized at different levels:
physical acoustic features (e.g. Wiener entropy, amplitude, peak
frequency); psychoacoustic features (e.g. timbre, pitch, loudness),
which vary across species depending on auditory perceptual sys-
tems; and/or musical features (e.g. rhythm, melody, harmony),
which emerge due to interactions between perceptual and cogni-
tive systems. Many species of animals show remarkable music
perception abilities, which could have arisen homologously or
convergently to human faculties for music perception (Hoeschele
et al., 2018), and sometimes these abilities are similar to our own
(Panksepp &Bernatzky, 2002). This similarity generates predictions
about what aspects of sounds might be perceptually salient to an-
imals. For example, some species can recognize particular fre-
quencies (‘perfect pitch’;Hoeschele, 2017) and can discriminate
between consonant and dissonant intervals, a component of many
human musical systems (Izumi, 2000;Watanabe et al., 2005), and a
few species can coordinate their vocalizations or movements with
an external beat (Cook et al., 2013;Patel, 2021). A better under-
standing of the saliency of different sound aspects to animals will
help to predict their response to music in particular and to
anthropogenic sound in general.
Contributions of Musicology to Studies of Animals' Responses to
Music
Musicology is a broad, multidisciplinary field in which meth-
odologies from the humanities, arts, social and/or natural sciences
are applied to the study of music and its production, perception and
appreciation (Parncutt, 2007). Although musicology’s focus is on
human music, recent decades have seen increasing numbers of
cross-species studies. On the one hand, these have come from the
natural scientific perspective of evolutionary musicology, or bio-
musicology (Fitch, 2006;Honing, 2018;Wallin et al., 2000), which
aims to illuminate human capacities ‘to perceive, appreciate and
produce music’and its variation across other animals (Hoeschele
et al., 2018, p. 149). On the other hand, zo€
omusicology has turned
musicological expertise and methods from the humanities, arts and
social sciences, as well as from ethology and bioacoustics, towards
animal sounds (e.g. Doolittle, 2020;Doolittle &Gingras, 2015;
Martinelli, 2009;Rothenberg, 2008;Taylor, 2017). Further musi-
cological disciplines relevant to understanding how animals
respond to music include acoustics (the physical properties of
sound), psychoacoustics (the correlation of sound with psycho-
logical sensations and perceptions), music cognition (how music is
perceived and cognitively assessed, and the connection between
perception and affective and bodily responses) and music infor-
mation retrieval (the extraction of computational measures of
acoustic and musical features from audio).
Rather than review all the effects of music on animals (which
has been covered elsewhere: e.g. Alworth &Buerkle, 2013;
Kriengwatana et al., 2022;Kühlmann et al., 2018;Wells, 2009), this
review highlights important gaps in how animals' perception of
music is considered and how this limits our understanding of the
diverse effects of music that have been reported. This paper de-
scribes features important for music perception in humans and
argues that many of these also apply to animals. It subsequently
uses a systematic review of studies on the effects of music on an-
imals to evaluate how musical knowledge is critically needed to: (1)
improve stimulus choice through a better understanding and
characterization of music; (2) ensure sufficient reporting of music
sources and description of music stimuli to increase scientific
reproducibility; (3) elucidate how animals' perception and
response to music are affected by the interaction between music’s
acoustic features, listener characteristics, playback conditions and
producer characteristics and contexts; (4) explore how assump-
tions of individualism and agency influence the kind of research
being conducted on how animals respond to human-generated
sound. Finally, we describe how Berlyne’s(1971)psychobiological
theory may help to understand variable responses and generate
novel predictions about how animals might respond to human-
generated sound.
Intrinsic and Extrinsic Features in Music Perception
In this section, we describe the contributions of key factors that
affect human responses to music. In humans, the perception of
music depends not only on (1) its intrinsic properties (e.g. acous-
tics) but also on extrinsic features; that is, (2) listener properties, (3)
playback contexts and (4) producercontext and properties (we will
collectively refer to these as ILPP). Here, we use the term ‘producer’
to refer to human composers and performers. We use the term
‘listener’to refer to humans or animals that are close enough to the
source of music playback to hear it. Fig. 1 visually summarizes the
interaction of intrinsic and extrinsic features that influence re-
sponses to music in listeners and Table 1 lists examples of each ILPP
factor. First, we discuss features known to affect music perception
in humans.
Intrinsic features
Intrinsic features refer broadly to time-varying acoustic, psy-
choacoustic or musical features that can be reproduced from
recorded audio. No two recordings of the same piece of music will
have identical intrinsic features, due to variation in producer
properties and context (see below). The faithfulness of this repro-
duction will depend on the quality of the playback technology. At
the lowest level, the acoustic features of sound waves are typically
represented in time and frequency domains, or as spectrograms.
Independent of a listener, they include the duration, energy and
peak frequency of a sound. Psychoacoustic features, including pitch,
timbre, consonance and loudness (Schneider, 2018), arise from the
interaction of sound waves with the perceptual system of the
listener. Recognition of musical features, such as rhythm, melody
and harmony, requires higher-level perceptual and cognitive abil-
ities. The perception of both psychoacoustic and musical features
hence depends on the interaction between acoustic features,
listener properties and playback context. If music is altered to test
B. P. Kriengwatana et al. / Animal Behaviour 221 (2025) 1230742
the effect of certain parameters on perception and behaviour, de-
tails of any deliberate manipulation made by experimenters to alter
how the music sounds (e.g. shifting pitch) should be reported so
that music-intrinsic features can be accurately reproduced in future
studies.
Listener properties
Listener properties constrain what a listener can perceive, how
they will interpret what they perceive and consequently how they
will respond behaviourally, physiologically or affectively to music
(Kriengwatana et al., 2022). In humans, the effects of music can be
influenced by slow-changing (or permanent) and fast-changing
listener characteristics. Slow-changing characteristics may include
age (Nieminen et al., 2011), sex (Nieminen et al., 2012), perceptual
abilities (e.g. assessed through the frequency dependence of audi-
bility thresholds), learning (Krumhansl, 2000), musical training
(Broughton et al., 2021), history of musical exposure (Lahdelma
et al., 2021), familiarity (Lahdelma &Eerola, 2020;Senn et al.,
2021) and preferences (Bodner &Bensimon, 2016). Faster-
changing properties include fluctuating internal physical and psy-
chological states, such as mood (Xue et al., 2018) and attention
(Koehler &Broughton, 2017;Loui &Wessel, 2007), which are partly
determined by activities before or during music exposure. Some of
these characteristics can be either slow or fast changing, such as
when learning might affect how the person perceives the next
sound they hear, or when sudden hearing loss alters what was
previously a constant perceptual ability.
Playback context
Playback context refers to the physical and social environment
where music is perceived (e.g. concert hall, at home, during a
commute), as well as the playback equipment used to delivermusic
stimuli. The physical space and playback equipment control
acoustic and psychoacoustic features of the sounds (e.g. through
reverberation and the loudspeaker frequency bandwidth), and can
directly influence the emotional excitation of human listeners
(P€
atynen &Lokki, 2016). In humans, affective responses to music
are also modulated by social aspects, such as social feedback
(Koehler &Broughton, 2017).
Producer properties and context
Whether precomposed, or produced spontaneously (improv-
ization), the acoustic features of recorded musical performances are
only partly determined by properties of their composers and per-
formers (such as voice and/or instruments used). They also depend
on the context; that is, the physical, social and cultural-historical
setting in which the music is created, performed and/or recorded,
as well as on the recording technologies used.
Social
structure
Cognitive
abilities
Perceptual
abilities
Current
activities
Prior
experience
Mood
Attention Habituation Familiarity Life history Personality Age & sex
Rhythm Tempo
Pitch
loudness
Speakers
Ecology
Health &
welfare
Ethical
implications
Individual
response
Applied animal sciences
Conservation
Social
Other
stimuli
Internal
state
Energy
Skill Functions Culture Politics
Arousal
Piece or
song
Voice
instrument
Audience
venue
Temporal
structure
Musical
training
Melody
harmony
Texture
timbre
Duration
repetition
Acoustic
space
Spectral
structure
INTRINSIC
FEATURES
PRODUCER
PROPERTIES
& CONTEXT
LISTENER
PROPERTIES
(FAST-CHANGING)
LISTENER
PROPERTIES
(SLOW-CHANGING)
PLAYBACK
CONTEXT
Ambient
sounds
Biological
relevance
Figure 1. Individual responses to music playback (whether physiological, affective, behavioural or cognitive) may be influenced not only by intrinsic features of audio recordings
used as music stimuli, but also by the properties of producers and listeners, and by physical and social aspects of the playback context. In animal studies, experimental designs
(represented by the dark grey circle) should take into account all these factors when assessing the impact of music on animals. Ethical implications (represented by the light grey
background) should also be considered when designing experiments interpreting the results of animals' responses to music (e.g. related to agency and individualism).
B. P. Kriengwatana et al. / Animal Behaviour 221 (2025) 123074 3
Research in animal communication suggests that ILPP factors
also have potent effects on some species other than humans. Spe-
cifically, animals communicate with each other to inform about
their internal states, the presence of ecologically relevant events in
their environment (Seyfarth et al., 2010) or to manipulate or
manage the behaviour of recipients (Rendall et al., 2009). The
acoustic structure of a signal is related to the communicative pur-
pose of that signal (formefunction relationship) and is evolution-
arily conserved over many taxa (Briefer, 2012;Bryant, 2013). We
would therefore expect that intrinsic acoustic properties, the
listener and the playback context can play a role in influencing
animals' responses. In Table 2, we give examples and evidence of
why these factors should be considered and reported in animal
studies. If there is evidence that animals respond to a feature, we
suggest that this must be reported. If evidence is limited to humans,
we suggest that it could be reported.
DO ANIMAL STUDIES CONSIDER INTRINSIC AND EXTRINSIC
FEATURES OF MUSIC PERCEPTION?
We have made the case that animals not only respond to
intrinsic music features, but they may also have the capacity to be
influenced by extrinsic features. To determine whether animal
studies are considering the whole range of ILPP factors, we con-
ducted a systematic review of original, peer-reviewed research ar-
ticles in Web of Science published over the last 3 years (between
2021 and 2024) that tested animal responses to music. This is not
intended to be an exhaustive review of all studies but to provide a
representative insight into the current state of research on effects of
music on animals. The search terms we used are listed in full in the
Supplementary Material. We chose a range of animal terms to
ensure a broad survey of different species while also targeting
species that we know have been the focus of research on animal
responses to music (e.g. laboratory rodents, Kühlmann et al., 2018;
farm animals, Ciborowska et al., 2021; zoo-housed nonhuman
primates, Robbins &Margulis, 2014;Wallace et al., 2017; com-
panion animals, Lindig et al., 2020).
We included articles that met the following criteria: (1) animals
were passively or actively exposed to music, and their behavioural,
physiological, affective or neural responses were measured; (2) the
authors described the stimuli as ‘music’or ‘music-like’.We
excluded articles that were: (1) not original, peer-reviewed
research articles (e.g. review articles or commentaries); (2) arti-
cles that analysed song or music production by animals; (3) articles
where neither human experimenters nor animal subjects had
control over music stimulus exposure; (4) articles that examined
music perception and/or discrimination abilities. For each included
article (N¼76), we noted if 24 parameters pertaining to ILPP were
reported (see Table 1). We additionally noted whether music
stimuli were chosen based on genre labels and whether animals
had the choice of turning music on or off (i.e. control; total 26 pa-
rameters extracted). Full details of our search strategy, PRISMA
flowchart and list of included articles are available in the Supple-
mentary Material.
A visual summary of the results is given in Fig. 2. Most studies
(97.4%) were on captive animals (74 out of 76 studies). We discuss
findings related to each ILPP feature and genre.
Intrinsic Features
Duration of each music stimulus (e.g. song) was reported in
22.7% of studies, which did not include the study using a single
frequency (dos Santos et al., 2023). Range of sound pressure levels
(SPL) was reported in 46.8% and average SPL was reported in 33.8%
of studies.
Listener Properties
Although there is good evidence that both sex and age of an
animal can affect its response to sound (Table 1), sex of the study
animals was reported in only 77.3% and age was reported in 73.7%
of the studies. Some studies reported only the life history stage (e.g.
‘pregnant’,‘lactating’or ‘adult’) rather than the chronological age of
the animal (e.g. Kamar &Yusof, 2023;Kochewad et al., 2022).
Whether animals had prior music exposure was explicitly reported
in only 12.2% of studies performed on captive animals, even though
captive animals could have experienced music at some stage.
Playback Context
Playback equipment was specified in 36.4% of studies. Room
acoustics (room size, enclosure size, soundproofing or building
materials used) was reported in 29.7% of studies that were con-
ducted indoors. Ambient sound levels were reported in 20.8% of
studies. Social housing conditions (whether animals were housed
alone or with other conspecifics) was reported in 67.6% of studies
that used captive animals. Human presence during music playback
was rarely reported (14.5%).
Duration of total music exposure was reported in 86.5% of
studies (this did not include two studies where animals could
control how much music they heard; Hirskyj-Douglas &
Kankaanp€
a€
a, 2022, pp. 1497e1511; Truax &Vonk, 2023). Intervals
between music playback sessions (when no music was presented)
was reported in 78.6% of studies. Intervals were not reported in the
two studies where animals could control music presentation and in
four studies with only one playback session. Intervals between
playback of each music stimulus within a playback session (e.g.
transition between songs) was reported in 18.9% of studies not
including the two studies where animals controlled music pre-
sentation. Repetition (i.e. how many times a music stimulus was
Table 1
Examples of intrinsic, listener, playback and producer (ILPP) features
Intrinsic features Duration of the piece of music
Average sound pressure level (SPL)
SPL range
Frequency range
Listener properties Species
Music experience (fast changing)
Age (slow changing)
Sex
Playback context Ambient sound levels
Deliberate manipulations by experimenters
Room acoustics
Playback equipment
Presence of humans
Presence of other conspecifics
Pauses between pieces within a playback session
Pauses between playback sessions
Number of repetitions
Section of the piece of music
Total duration of playback
Producer properties
and context
Piece of music
Composer(s)
Performer(s)
Live or studio recording
Release date
Sampling rate
Producer properties and context refer to aspects of the composition, performance
and recording of the piece of music. The resulting recording (audio), when played
back, has intrinsic features (acoustic, psychoacoustic, musical) of which some can be
measured objectively and described with summary statistics. The responses of an-
imals are further determined by the playback context (including controlled aspects
of experimental design), and slow- and fast-changing listener properties.
B. P. Kriengwatana et al. / Animal Behaviour 221 (2025) 1230744
Table 2
Suggested features of stimuli and playback conditions to report (analogous to ARRIVE guidelines, Percie du Sert et al., 2020)
Feature Must-have (M)
or Could-have (C)
Rationale for inclusion with supporting evidence that animals respond to this feature
Species M Listener properties that are relevant for music perception and response at the level of species include the
species' cognitive ability, hearing ability and social structure/organization. Although listener properties have not
yet been systematically evaluated, some of their influences have been noticed. A few studies on music for animal
welfare paid attention to animals' perceptual capacities, such as hearing range (e.g. Hampton et al., 2020;
Snowdon et al., 2015;Zapata Cardona et al., 2023) or properties of conspecific vocalizations, in predicting how
animals will respond (Snowdon et al., 2015;Snowdon &Teie, 2010). Attention is needed to ensure that a
stimulus is within the hearing capacity of the species and is not close to sounds that are aversive to that species.
There is also an as yet underappreciated relationship between animal communication and social behaviour
(Briefer et al., 2024) and it is possible that responses to acoustic stimuli may vary with social organization of a
species.
There are also listener attributes that vary within species. The most important, and those that can readily be
assigned, are discussed in the following rows.
Sex M Sex can affect the response to music. For example, in budgerigars, Melopsittacus undulatus, males preferred
arrythmic stimuli whereas females preferred rhythmic stimuli (Hoeschele &Bowling, 2016). Sex differences
may be due to differences in sex-specific behaviours or due to differences in hormonal state. Within a sex,
variation in hormone levels associated with different reproductive stages may also affect the response to music;
e.g. the probability of response of female túngara frogs, Physalaemus pustulosus, to male calls increased with
increased gonadotropin levels (Lynch et al., 2006). If the sex of the animals used in a study was not known or not
possible to distinguish, then this should be explicitly stated.
Age M Sensory ability to perceive stimulus and cognitive capacity to discriminate can vary with age of the animal
(Frisina, 2009).
There are also other interindividual variation in traits that may influence how an animal responds to music that
are less readily identified. Individual responses to acoustic stimuli can vary according to ‘temperament’of
individuals (e.g. shyer eastern chipmunks, Tamias striatus, reacted more strongly to alarm calls; Couchoux et al.,
2018) or according to the ‘current state and perceived threat’(e.g. at higher ambient temperatures, great tits,
Parus major, responded in more risk-aversive ways to conspecific mobbing calls; Cordonnier et al., 2023).
Animals in different ‘affective welfare states’can also respond differently to the same sound in judgement bias
tests (Mendl et al., 2009).
Prior experience with
music
M An animal’s prior experience with the same or similar stimulus, and experience associated with that, can
influence subsequent responses. An animal’s response to species-specific sounds can depend on prior
experience; e.g. in zebra finches, Taeniopygia guttata, females showed a preference for a previously experienced
male song (Riebel et al., 2009). We have not found studies testing the effect of prior experience to artificial
sounds but, for instance, prior experience of consecutive, repeated exposure to a music stimulus caused
habituation in dogs (Bowman et al., 2015).
Piece of music
(composer,
performer, release
date, section)
M Providing information that pinpoints the music stimulus used is crucial for understanding what animals respond
to and for reproducibility. Citation of music sources according to academic conventions can solve this problem,
but if this is not possible then specifying performance and release date (in addition to composer) and exact
section of music used (if applicable) can help remedy this problem as well.
Animals can discriminate between different pieces of music (e.g. Porter &Neuringer, 1984). However, variability
among recorded performances, whether by the same or different performers (e.g. differences in expressive style,
tempo, vocal timbre and instrumentation) means that specifying the music stimuli by name of the piece or song
and the recording artist (or composer, for classical music) does not adequately specify the acoustic,
psychoacoustic or musical features of the music stimulus. The release date can specify the exact performance
used. Animals such as songbirds can discriminate between birdsong from different populations and individuals
(Nelson &Poesel, 2007) and rats, Rattus norvegicus, can discriminate between the same piece or song performed
by different artists (Okaichi &Okaichi, 2001).
Pieces of music usually contain contrasting sections (resulting in ‘intraperformance variability’). This means if
only part of a larger piece of music is used, the section that has been used should be specified (e.g. track number,
movement title, timing) as different sections within a piece of music can be very different.
Duration, pauses and
repetition
M Animals can habituate to a stimulus and stop responding to it, or respond to it negatively, if it is presented often
and/or repetitively (Bowman et al., 2017;Sierro et al., 2023). Pauses between music stimuli are also important to
report, as animals behave differently towards intermittent versus continuous sounds (Neo et al., 2014) and
sometimes prefer silence over sound (McDermott &Hauser, 2007;Truax &Vonk, 2023).
Frequency range and
sound pressure level
(SPL)
M Pieces of music will range both in amplitude and frequency. Animals vary in their sensitivity to different
frequencies, thus knowing what parts of a stimulus can be heard is important. Loud sounds can be stressful
(Kight &Swaddle, 2011) and sounds with higher or lower pitches can induce different behavioural responses
from listeners (Morton, 1977). Reporting whether sound meters used to measure SPL are set for human hearing
(i.e. ‘A’filter or not) may also enable knowledge of what animals are hearing.
Details of deliberate
manipulation of
stimuli by
experimenters
M During playback of a piece of music its exact attributes can be deliberately manipulated (e.g. sped up, slowed
down, played backwards and/or had the tuning altered). Songbirds, for example, have extraordinary temporal
discrimination abilities (Dooling &Prior, 2017) and may perceive manipulations not detected by a human
listener. Animals can distinguish between variation in sounds produced by humans or conspecifics (Nelson &
Poesel, 2007;Okaichi &Okaichi, 2001). Researchers should also report, to the best of their ability, if the music
has been compressed and/or equalized, as this will affect the frequencies heard and the variability between loud
and quiet sounds. This is required to understand the animal’s listening experience and for stimulus replicability.
Ambient sound levels M The stimulus of interest may be at least partially masked by background sounds (e.g. machinery in animal rooms,
vocalization of other animals present). For a listener to perceive a stimulus, it requires that the ratio of the signal
to the background noise (i.e. signal-to-noise ratio, SNR: The difference between signal and noise amplitudes
measured in decibels) within a frequency band exceeds a critical detection threshold (e.g. for birds this can range
from 18 dB to 37 dB depending on the frequency), and discrimination of the signal from other signals (e.g.
perceiving certain attributes of the music as threatening or reassuring) requires an even higher SNR than
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B. P. Kriengwatana et al. / Animal Behaviour 221 (2025) 123074 5
repeated) was reported in 9.6% of studies, and did not include the
two studies where animals controlled music presentation and the
study where a single frequency was used.
Most studies did not intentionally manipulate or alter music
stimuli (67.5% of studies). However, out of the 24 studies that did
manipulate music stimuli, only six studies (25%) reported sufficient
information to replicate the manipulation. In at least one case, the
researchers were unaware that the music they played had been
lowered in pitch from the original (‘“The Division Bell”(pitch at
432 Hz) by Pink Floyd was selected’;Russo et al., 2021), potentially
leading to problems should anyone try to replicate the experiment.
Producer Properties and Contexts
Relevant information was extracted from 75 studies (excluding
one study that used a single frequency; dos Santos et al., 2023).
Sampling rate was reported in 7.8% of studies. Composer informa-
tion was given in 58.7% of studies. Information about composers as
for example ‘French classical piano’and ‘flute music’(Kamar &
Yusof, 2023) refers to a style as determined by region and era,
and a broad instrumental category that could include music from
many contrasting genres; neither is sufficiently specific. In two
cases the music was described as using a particular Indian raga
(Kochewad et al., 2022;Rathod &Vaidya, 2024), which refers to a
set of melodic patterns and pitch relationships (Kaufmann, 1968).
Although naming the composer is not standard for a raga, naming
the performer or improvisor and recording would be essential for
replicability, as interpretation varies widely, not only between
performers but between different performances by the same
performer. Several studies used one or more pieces from the same
composer. Mozart’s compositions were the stimulus in 36.8% of the
studies: out of those, 71.4% used Mozart’s‘Sonata for Two Pianos in
D Major’, K.448 and 14.3% used Mozart’s string quartets K.428,
K.525, K.458. Different pieces by Bach were also used in four studies
(dos Santos Lemes Lechuga et al., 2023;Lippi et al., 2022,2023;
Palermo Mendes et al., 2023).
Performer information was reported in only 14.7% of studies, but
the same piece of music may differ between performers and
animals have been shown to distinguish between vocalizations
from different individuals (reviewed in Carlson et al., 2020). Release
date (needed to distinguish between multiple recordings made by
the same performer that may be different) was reported in only
5.3% of studies. Similarly, live or studio recordings were reported in
5.3% of studies, but live recordings may contain sounds from the
audience that animals may respond to. The musical work was
named in 58.7% of studies but the specific section(s) of the work the
animals were exposed to were named in only 13.7% of studies.
Hence, only a minority of studies provided sufficient information
on the music stimulus to allow replicability of the study.
Genre
Music stimuli were grouped according to genre in 24 out of 70
studies (34.3%). Six studies did not report sufficient information to
deduce whether genres were used (Dong et al., 2024;Epstein et al.,
2021;Liu et al., 2022;Mao et al., 2022;Wang et al., 2022;Xu et al.,
2022). For example, Mao et al. (2022) and Dong et al. (2024)
described their stimuli as ‘soothing music’and ‘background mu-
sic’, respectively, without further explanation or examples.
WHERE CAN MUSICOLOGY HELP TO IMPROVE STUDY
DESIGNS?
Our results show that recent studies on how animals respond to
music do not sufficiently consider, control for or report many ILPP
factors. As a result, music recordings were generally not described
in sufficient detail to pin down specifically which sounds were
experienced by an animal. This raises the issue of scientific repli-
cability and confuses any attempts to identify the salient features of
the sound environment that animals respond to. It potentially
marks an underlying assumption that certain differences that are
important to human listeners are not important to other animals
(e.g. the performers of a piece of music, even though this factor
leads to considerable differences in the audio). We further identi-
fied problems related to assumptions about music categories (ac-
cording to composer or genre), misinformation about music and
Table 2 (continued )
Feature Must-have (M)
or Could-have (C)
Rationale for inclusion with supporting evidence that animals respond to this feature
detection (de Kort et al., 2024;Lohr et al., 2003). Ambient sounds before and during music playback could be
provided as supplementary information.
Social environment C The disruption of customary social relationships by the music stimulus may cause stress and change an animal ’s
response to music (Hawkins et al., 2024). Presence of a familiar or unfamiliar human may also affect stress and
hence an animal’s response to music.
Room acoustics C The capacity of a listener to perceive a signal depends on the distance between source and listener and the
signal’s frequency (as sounds of different frequencies carryover different distances), and this also depends on the
environment in which the stimulus is broadcast. So, the distance between sound source and listener as well as
room acoustics (size of room, size of enclosure, hard vs soft surfaces) may play a role in how the listener
perceives a stimulus. Size of the enclosure will affect whether animals are able to vary their perception
(proximity, volume) of the sound stimulus. Audio recordings of the space where animals are exposed to music
before and after music playback may be needed to better understand what features of music become more or
less salient and/or distorted.
Recording conditions C Original recording conditions of a music stimulus should be reported (e.g. studio/live, close-up) to account for
the existence of sounds that may change the effect of the stimulus. Some recordings may also contain sounds
that are not part of the musical work but may potentially affect the listener (e.g. crowd sounds in live recordings,
or Glenn Gould’s audible singing in his piano recordings). Acoustic features of live recordings will vary from
performance to performance, as performers are influenced by audience presence and feedback, and
intentionally or incidentally alter their performance in subtle or obvious ways (LeBlanc et al., 1997;Moelants
et al., 2012).
Playback equipment C Loudspeaker make and model are important to know, because they affect the frequency distribution and
bandwidth that can be produced (for example, bass frequencies are not produced in small speakers). Equipment,
therefore, might affect animal responses. All the studies in our literature survey used digital recordings but if a
different recording medium or format is used (e.g. analogue tapes) then it should be mentioned.
Most of this information will be captured by appropriately referencing the music stimuli.
B. P. Kriengwatana et al. / Animal Behaviour 221 (2025) 1230746
views of individualism and animal agency that not only have ethical
implications but can implicitly/unconsciously shape experimental
designs and questions.
Referencing and Citation of Music
Music stimuli should be cited in research papers according to
standard scholarly practices used in human music research, giving
sufficient information to identify the precise recording(s) used: for
example, ‘Collins, J. (1970) “Farewell to Tarwathie”, Whales and
Nightingales [Vinyl]. New York: Elektra Records’. In the case of
hard-to-find recordings, citations could be usefully supplemented
by providing audio stimuli in digital repositories, or as journal
supplementary information. If copyright issues prevent this,
recording metadata such as the International Standard Recording
Code (https://isrc.ifpi.org/en/) could be provided for each digital
track. Table 2 lists and describes the most important aspects of
stimuli and playback conditions that animal studies should report
to allow understanding of what was played and achieve replica-
bility and reproducibility.
Assumption of Homogeneity for Music From the Same Composer and
Genre
Over 30% of studies in the past 3 years continue to simply use
genre to characterize the music stimulus. A smaller, but related
assumption seems to be that music from the same composer will
either sound the same or have similar effects on animals.
Duration total
Pauses between sessions
Sex
Age
Social conditions
Piece of music
Range SPL
Playback equipment
Genre
Average SPL
Room acoustics
Manipulations
Duration piece
Ambient sound levels
Pauses within sessions
Performer
Human presence
Section
Music experience
Repetitions
Sampling rate
Release date
Live or studio
0255075
% Studies re
p
orted
Attribute
Intrinsic
Listener
Playback
Producer
Figure 2. Results of systematic review showing percentage of studies within the last 3 years that report features listed in Table 1.
B. P. Kriengwatana et al. / Animal Behaviour 221 (2025) 123074 7
Musicological research into automated music genre recognition
shows that musical genre categories are not objectively defined nor
stable, and that there is considerable variability within genres and
overlap between them.
Subjectivity in construction of genre taxonomies and human
classification
Music genre recognition (MGR) is used by digital music services
to classify music and make recommendations to its users. Within
MGR it is widely acknowledged that genre taxonomies (usually
conceived hierarchically) are subjective because different taxon-
omies, whether used for commercial systems (e.g. Spotify) or for
research, rarely agree on the names of taxons (e.g. ‘classical’,‘hard
rock’) or their location within the hierarchy. Moreover, taxonomies
depend on both intrinsic (acoustic, psychoacoustic, musical) and
extrinsic (artist, album, country, period) features (Aucouturier &
Pachet, 2003;Ramírez &Flores, 2020). This is one reason why in-
dividual items may be located in multiple genres (McKay &Fujinaga,
2006;Ramírez &Flores, 2020). Even when categorizations are based
on intrinsic features only, these almost inevitably include some
features that are predicated on perceptual features of the human
auditory system (e.g. Mel Frequency Cepstral Coefficients; Fu et al.,
2011) or on human biological responses, and may be inapplicable
to other animals. For example, subgenres of electronic dance music
are defined in part by the tempo, which is determined in relation to
typical human heart rates and/or movement (Burger &Toiviainen,
2020;Chen et al., 2021;Leimeister et al., 2014). There is also
disagreement in allocation of musical items within taxonomies
between human classifiers, with each other and with commercial
systems (Ramírez &Flores, 2020;Schedl et al., 2014).
Change and instability of genre classifications
Rather than taxonomies describing a stable ‘order of things’,any
system of classification depends on who is doing the classifying,
what they are doing it for and the methods employed (Sturm,
2014). As tastes and technologies change, genres evolve, spawn
new genres and disappear (Aucouturier &Pachet, 2003;Nie, 2022).
We note that the number of named genres identified by Spotify
using a blend of machine learning and human curation, expanded
from 1742 in 2018 (Johnston, 2018) to almost 6000 in 2023 (Krogh,
2023). Furthermore, even the most standard genre labels may refer
to categorizations of vastly different bin sizes. For example the
word ‘classical’is commonly used to refer to enormously varied
styles of Western classical music composed throughout Europe and
countries influenced by European music since the 9th century, and
typically performed in more formal contexts (although it may also
be used more narrowly to describe music written in Europe and
European-influenced countries during the Classical era,
1750e1820), while ‘country’refers to a less diverse grouping of
music written and performed primarily (although not exclusively)
in the U.S.A. over the past 100 years. Therefore, rather than relying
on genres, it may be more fruitful to test how specific ILPP factors
and their interactions affect animals.
Misinformation About Music
Because so much musical knowledge is practice based (experi-
ential) rather than traditionally academic, it can be hard for non-
musicians to recognize which sources of musical information are
reliable, and which are popular oversimplifications, mis-
conceptions or even conspiracy theories. While most papers in our
systematic review avoided perpetuating outright musical false-
hoods, some showed the influence of popular music mis-
conceptions, such as belief in the now-largely-debunked ‘Mozart
effect’(previously discussed in Kriengwatana et al., 2022). While
scientists may choose to continue to use Mozart K.448 to maintain
consistency across studies, it is important to recognize that Mozart
K.448 is not in itself unique, nor does it have a specific‘function in
stimulating brain activity’(Luo et al., 2021), and that most of its
positive effects could probably be replicated with other pieces of
music with similar qualities (such as other piano pieces, other
Classical-era compositions, other pieces with a steady rhythm,
other pieces in a similar frequency range, etc.), or with nonmusical
forms of cognitive stimulation.
Even more troubling is when scientific researchers fall prey to
popular music conspiracy theories, including those about the note
A being tuned to 432 Hz, and ‘solfeggio frequencies’. Substantiable
statements, such as ‘Evidence supports that music can modulate
many physiological roles, exerting clear effects on the central ner-
vous system’(Bidari et al., 2023) become intertwined with con-
spiracy theories, e.g. ‘For this effect to be positive, music should be
tuned at a frequency of 432 Hz’;Bidari et al., 2023), and false his-
tories, e.g. ‘in the past, musical instruments were tuned to 432 Hz’
(Bidari et al., 2023;Russo et al., 2021). In fact, A¼432 Hz has never
been a widespread or universally agreed upon tuning (Haynes,
2002;Rosenberg, 2021), and according 432 Hz any special signifi-
cance has to do with numerology based on the scales used in
Western classical music and the (arbitrary) length of seconds,
rather than any intrinsic characteristics of the frequency itself or its
‘resonance’with human or nonhuman bodies or the universe. One
study also conflated a legitimate music pedagogical technique (the
use of solf
ege syllables, solfeggio in Italian, to sing the notes of a
scale) with a similar-sounding fabrication popularized by New Age
entrepreneur David Hulse (2009),‘solfeggio frequencies’, citing
papers about the pedagogical value of solf
ege in support of their
choice to play ‘solfeggio frequencies’to zebrafish (dos Santos et al.,
2023). These misconceptions about music hamper a thorough
exploration of the effect of music on animals but can be prevented
through interdisciplinary collaborations.
Individualism, Agency and Ethical Implications
Among the 76 studies that we reviewed, only two (2.6%), both
conducted on captive zoo-housed animals (Hirskyj-Douglas &
Kankaanp€
a€
a, 2022, pp. 1497e1511; Wu et al., 2021), give animals
control over their music listening experiences. In the majority of
studies we reviewed, animals exposed to music were confined to a
small space (cage or stall) and were unable to remove themselves
from the music. Human audiences in most cases deliberately
choose to listen to music (or are at least free to stop listening).
Donaldson and Kymlicka (2011) differentiate between domesti-
cated animals living in close proximity and mutual dependence
with humans (‘citizens’), liminal animals living in proximity to
humans but independent of them (‘denizens’) and animals who live
independently from humans in their own environment (‘sover-
eigns’). This threefold concept helps situate our results in a broader
context. Playing music to animals living in mutual dependence with
humans (‘citizens’) on the one hand means that these animals are
often habituated to human-generated sounds and might even have
been exposed regularly to music, with which they may have posi-
tive, negative or neutral associations. On the other hand, they may
live in a confined situation that makes it difficult or even impossible
for them to choose to avoid music exposure. Free-living, wild ani-
mals can have more control over their interaction with human
music than domesticated or liminal animals, by adjusting their
behaviour (Francis &Barber, 2013). So, the scientific need for a
controlled environment for the collection of musical and behav-
ioural data can exist in tension with the ethical need of providing
animal participants the choice between listening and not listening.
B. P. Kriengwatana et al. / Animal Behaviour 221 (2025) 1230748
We would argue that in cases where animals do have control
over their sound environment, many species will willingly exert it.
The ability to control sound exposure (e.g. by moving away or to-
wards sounds) may contribute to animals' sense of agency, defined
as ‘the capacity of animals to engage in voluntary, self-generated
and goal-directed behaviour that they are motivated to perform’
(Littlewood et al., 2023). Increasing sonic agency could lead to
immediate and/or delayed impacts on behaviour, physiology and
welfare beyond the effects of the sound stimulus itself. Being able
to control the sound environment may also affect animals' behav-
iours towards music and other human-generated sounds in the
future. Overall, the present and past choices that animals have had
to influence their sound environments, combined with their
proximity, experiences and relationships with humans, will prob-
ably affect their responses towards music and other human-
generated sounds. These ideas are partially supported by
Kleinberger et al. (2020), who created a system for a zoo-housed
parrot to turn music on or off. The animal subsequently used this
to control when they wanted to interact with visitors, as more
visitors assembled and interacted with them when they turned the
music on, and left when they turned the music off.
Moreover, patterns of sound preference (when and how much
sound) exhibited by animals may not match what animals are
typically exposed to in passive listening studies. Zoo-housed lemurs
that were able to enter or exit a device to trigger or terminate sound
playback showed a strong preference for very short durations of
sound exposure (order of seconds) and usage peaked during spe-
cific hours of the day (Hirskyj-Douglas &Kankaanp€
a€
a, 2022, pp.
1497e1511). This preference contrasts with the majority of the
passive exposure studies, where animals are exposed to music
(sometimes the same piece repeated, e.g. 19 out of 76 studies
played only Mozart K.448) over and over for hours at a time and at
times chosen by experimenters. Although preferences undoubtedly
vary between species and are influenced by feeding and husbandry
schedules, Hirskyj-Douglas and Kankaanp€
a€
a’s results suggest that
there is a good chance that the duration and timing of music
exposure used in passive exposure studies may not be what (most)
animals want, but the impact of self-chosen short exposure of
music on the animals' behaviour and physiology still needs to be
studied. In human contexts, the positive or negative effects of music
can vary greatly according to when and how the music is played,
even if the musical stimulus itself remains the same. Listening to a
song once, voluntarily, could be a source of pleasure, while being
forced to listen to that same song repeatedly for many hours a day
can be unpleasant, or even a form of torture (Grüny, 2020). Other
species of animals may well be similar.
While research on music for animals has often been species-
oriented, it might be fruitful to consider a broad spectrum of
musical interest and receptivity within a species. Animal individ-
ualism views that even within a species, each animal is a unique
individual, with specific needs, desires, habits, preferences and
experiences. It places critical importance on the presence of indi-
vidual variation: animals in a group or population are not equiva-
lent nor homogeneous. The study of animal personality aligns to a
degree with the concept of individualism, in that it recognizes that
there is between-individual variation in behavioural responses and
that this variation is adaptive (Dall et al., 2004;Dingemanse &Wolf,
2010). While the existence of individual variation in response and
preference is not contested and has been reported in music studies
(Hirskyj-Douglas &Kankaanp€
a€
a, 2022, pp. 1497e1511; Truax &
Vonk, 2023;Watanabe &Nemoto, 1998), we believe it needs to
be emphasized more strongly in studies of animal responses to
music (and perhaps to human-generated sounds in general).
Only 12% of studies in our literature search explicitly reported
whether animals had prior experience with music and several
studies only specify animal age as ‘adult’. Here, the assumption may
be that responses to sounds do not change with experience and will
remain stable and unchanging once animals reach adulthood
(perhaps with exceptions during life history stages such as repro-
duction), which fails to recognize the dynamicity and adaptability
of animals.
Lestel (2004) has philosophically conceptualized the ‘singular
animal’, thus hinting at the possibility that in some species of an-
imals, a high degree of musicality and interest in music may be
found only in certain individuals. For example, composer and
environmentalist Jim Nollman writes of his experience playing
music with orcas: ‘I next discovered it was not “the orcas”playing
with me, but two whales in particular that gravitated to the boat
whenever we transmitted.’(Nollman, 2008). Furthermore, whether
an individual animal demonstrates a particular response or
behaviour or not may be partially context specific. Evidence for
highly individualized drumming skills by a single chimpanzee
(Dufour et al., 2015,2017) speaks for the transferability of Lestel’s
concept to the musical domain. Thus, captive environments require
a high degree of awareness for individual and even context-
dependent variations in musical preferences and responses, and
future studies could pay more attention to individual variation in
the response of animals to music and possible factors that may
explain between-individual variation.
Assuming that animals must have similar musical interests and
preferences to humans would be anthropocentric, and it is neces-
sary to take precautions against unwarranted anthropocentric
biases as far as possible. At the same time, anthropomorphism is a
more ambiguous concept, neither inherently helpful nor inherently
harmful. A cautious, considered, ‘critical’anthropomorphism that
recognizes the possibility of similar responses to sounds based on
shared biological or psychological traits and/or environments may
lead to a more considerate use of music with animals. Prum’s
research on evolutionary aesthetics emphasizes that the subjective
experiences and aesthetic decisions of animals have contributed to
the evolution of biodiversity, and permits plausible conjectures
about the evolutionary shared ancestry of aesthetic preferences
and the aesthetic receptivity of different species (Prum, 2018). In
this sense, a refined and cautious, animal-centric anthropomor-
phism (de Waal, 1999) could help us to better understand in which
ways animals share human music preferences and interests.
THE PSYCHOBIOLOGICAL THEORY
Berlyne’s(1971)psychobiological theory, which aims to explain
human aesthetic preferences for art (not only music) by proposing
that intrinsic and extrinsic ILPP features interact to affect human
listener responses, may provide a suitable framework for under-
standing animals' responses to sounds in general; for example,
music and anthropogenic noise. Even without presuming to know if
or how other species of animals might distinguish between ‘music’
and ‘noise’(something that varies even among human cultures), we
can use Berlyne’s theory to help guide choices about what kind of
music exposure may be helpful or harmful for animals.
The theory assumes that there is an optimal level of arousal that
follows an inverted U-shape function, where arousal is the physi-
ological and psychological state of wakefulness. The hedonic value
of an auditory stimulus therefore depends on how close it brings
the listener to their optimal arousal level. Psychophysical proper-
ties (e.g. loudness and pitch), ecological properties associated with
harmful and distressing or gratifying and beneficial outcomes, and
collative properties of the stimulus (which require comparing or
collating information from two or more sources, e.g. stimulus
novelty, complexity, ambiguity) are presumed to increase or
decrease arousal, towards or away from the optimal level.
B. P. Kriengwatana et al. / Animal Behaviour 221 (2025) 123074 9
A key strength of the psychobiological theory is that it accounts
for ILPP factors by recognizing that a listener’s arousal levels are not
always constant, so the hedonic value of the same stimulus can
change-over time and context for a single listener. Whether the
stimulus is evaluated as having a positive or negative hedonic value
can therefore be predicted by assessing the listener’s current
arousal levels and the amount of increase or decrease in arousal a
stimulus is likely to induce.
As a hypothetical example, in quiet environments with minimal
stimulation, music and noise (depending on their psychophysical,
ecological and collative properties) could increase the animal’s
arousal to a moderate level, which the animal would find pleasur-
able. In noisy environments, however, the animal may already be in
a high state of arousal and the addition of music or noise could in-
crease it further and beyond the optimal level, which the animal
would then find distressing. Berlyne (1966) describes a study where
rats were housed in a noisy or quiet environment. Rats housed in the
noisy environment were more motivated to hear previously pre-
sented sounds rather than a novel buzzer sound (measured as
number of lever presses in a Skinner box), whereas rats housed in
quiet environments showed the opposite motivation. The psycho-
biological theory’s emphasis on arousal modification could also
explain why sound playback might decrease arousal if it completely
masks potentially arousing environmental sounds such as predator
cues, but increase arousal if it partially masks the predator cue (due
to increased ambiguity of the sound; Berlyne, 1971).
The psychobiological theory also has useful predictions for the
effects of anthropogenic noise on cognitive performance. Distrac-
tion by noise while performing essential activities that require
cognitive or attentional resources (e.g. foraging, assessing mates
and monitoring predators) is one of the explanations as to why
anthropogenic noise is harmful for animals (Chan et al., 2010;
Dominoni et al., 2020;Potvin, 2017). The psychobiological theory
predicts that (1) noise should affect cognition only if it substantially
alters arousal and (2) as with music, how much arousal noise in-
duces depends on the variability of sound characteristics within
and among anthropogenic noise recordings used as experimental
stimuli (intrinsic features determined by producer properties and
context), the individual’s perceptual and cognitive capacities and
current arousal state, and playback context (including difficulty of
the cognitive task).
Noise with very highly arousing properties would disturb per-
formance regardless of task difficulty and the individual’scognitive
capacity or current arousal state. If noise causes arousal to increase
dramatically, the animal could be less motivated to perform the task
and more motivated to escape the acoustic stimulation. Noise with
only moderately arousing properties would disturb performance in
individuals already in a moderate or higher arousal state, and be
more disruptive as task difficulty increases and as individuals have
fewer cognitive resources available to devote to the task.
Some of these predictions are indirectly supported by a few
empirical studies, but none measures arousal. For instance, pink
noise matching spectral and temporal properties of traffic noise
affected cognitive performance of laboratory-raised great tits, Parus
major, in a difficult task, but not in an easy one (Halfwerk &van
Oers, 2020). The ability of wild shore crabs, Carcinus maenas, and
the tide-pool damselfish Sergeant major, Abudefduf saxatilis,tofind
food in laboratory settings was also not affected by boat noise in
relatively simple tasks (i.e. in an open arena or in a maze with only
two options; Hubert et al., 2021;Leduc, Costa, et al., 2021;Wale
et al., 2013). However, performance on more difficult tasks was
not measured. Urban noise did not affect learning performance of
free-living Australian magpies, Gymnorhina tibicen, or black-capped
chickadees, Poecile atricapillus, that had daily exposure to urban
noise (Connelly et al., 2022,2024;Templeton et al., 2023), which
could suggest that familiarity with the noise prevented a significant
change in arousal and subsequent effects on cognition. In
laboratory-raised zebra finches, Taeniopygia guttata, traffic noise
was found to affect inhibitory control, motor learning and spatial
memory in one study but not in another (Daria et al., 2023;Osbrink
et al., 2021). Individual variation in noise-induced arousal might
possibly explain these contradicting results.
It would be worthwhile to learn more about the properties of
anthropogenic sounds, including noise and music, that increase
psychophysiological arousal and by how much, and subsequently
to test the relationships between sound presence and characteris-
tics, arousal and cognition, to understand whether cognitive per-
formance is significantly affected when the combination of task
difficulty and properties of sound substantially alters arousal.
CONCLUSIONS
Currently, music stimuli are poorly characterized or improperly
referenced in the literature, significantly impeding our under-
standing of how music affects animals' behaviour, physiology and
cognition. Genre labels are not a shorthand for intrinsic features of
music and do not reliably predict the acoustic, psychoacoustic or
musical features of works contained within them. We strongly
encourage scientists to collaborate with musicologists to prevent
these and other misunderstandings about music in the future. The
contributions of sound environments to animal agency remains a
relatively unexplored but exciting topic of research. Research into
the effect of music on animals can also benefit from the flexible and
diverse concept of aesthetic preferences. Thus, we encourage crit-
ical reflection on the roles of choice and control of music exposure,
and of individual and aesthetic preferences, in scientific efforts to
understand how animals respond to music. Finally, viewing music
and other anthropogenic sounds from the psychobiological theory’s
perspective generates novel ideas that can be used to help predict
instances where individuals would be more strongly affected and
allows for individuality in responses. Equipped with this interdis-
ciplinary approach, we believe future research might avoid the
pitfalls we have described, and significantly improve our under-
standing of how animals respond to music (and possibly other
sounds) in new and exciting ways.
Author Contributions
Alex South: Writing ereview &editing, Writing eoriginal draft,
Investigation, Conceptualization, Methodology, Visualization. Bud-
dhamas P. Kriengwatana: Writing ereview &editing, Writing e
original draft, Visualization, Supervision, Project administration,
Investigation, Formal analysis, Data curation, Conceptualization,
Methodology. Emily L. Doolittle: Writing ereview &editing,
Writing eoriginal draft, Methodology, Conceptualization, Investi-
gation, Funding. Martin Ullrich: Writing ereview &editing, Writing
eoriginal draft, Investigation, Conceptualization, Methodology,
Funding. Ruedi G. Nager: Writing ereview &editing, Writing e
original draft, Investigation, Conceptualization, Methodology.
Declaration of Interest
The authors have no relevant conflict of interest to disclose.
Acknowledgments
We thank two anonymous referees for their helpful feedback
and suggestions on the manuscript. We thank the RCS Athenaeum
Award Funding and the Open Access Fund of Nuremberg University
of Music for open-access funding.
B. P. Kriengwatana et al. / Animal Behaviour 221 (2025) 12307410
Supplementary Material
Supplementary material associated with this article is available at
https://doi.org/10.1016/j.anbehav.2025.123074.
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