Attention, Perception, & Psychophysics (2022) 84:2360–2383

https://doi.org/10.3758/s13414-021-02364-4

The influence of auditory rhythms on the speed of inferred motion

Timothy B. Patrick 1,2 & Richard B. Anderson 1

Accepted: 26 July 2021 / Published online: 25 August 2021

# The Psychonomic Society, Inc. 2021

Abstract
The present research explored the influence of isochronous auditory rhythms on the timing of movement-related prediction in
two experiments. In both experiments, participants observed a moving disc that was visible for a predetermined period before
disappearing behind a small, medium, or large occluded area for the remainder of its movement. In Experiment 1, the disc was
visible for 1 s. During this period, participants were exposed to either a fast or slow auditory rhythm, or they heard nothing. They
were instructed to press a key to indicate when they believed the moving disc had reached a specified location on the other side of
the occluded area. The procedure measured the (signed) error in participants’ estimate of the time it would take for a moving
object to contact a stationary one. The principal results of Experiment 1 were main effects of the rate of the auditory rhythm and of
the size of the occlusion on participants’ judgments. In Experiment 2, the period of visibility was varied with size of the occlusion
area to keep the total movement time constant for all three levels of occlusion. The results replicated the main effect of rhythm
found in Experiment 1 and showed a small, significant interaction, but indicated no main effect of occlusion size. Overall, the
results indicate that exposure to fast isochronous auditory rhythms during an interval of inferred motion can influence the
imagined rate of such motion and suggest a possible role of an internal rhythmicity in the maintenance of temporally accurate
dynamic mental representations.

Keywords Attention . Timing . Auditory rhythm . Inferred motion

Introduction structure of external events, leading to future oriented attending

To accurately anticipate how events will unfold, people must
convert dynamic sensory input into internal representations of
time and space (Bar, 2009; Engel et al., 2001; Friston, 2005;
Graf et al., 2007; Grush, 2004; Mehta & Schaal, 2002;
Morillon et al., 2014; Morillon et al., 2015; Pezzulo, 2008;
Press et al., 2011; Schroeder et al., 2010; Springer & Prinz,
2010; Tian & Poeppel, 2010; Wolpert & Flanagan, 2001;
Wolpert & Kawato, 1998). The orienting of attention to dy-
namic sensory events requires an attentional system that is as
active and flexible as the events to which it attends (Balkenius
& Hulth, 1999).
Dynamic attentional models propose that rhythmic neural
oscillations adapt to and become entrained with the temporal
* Timothy B. Patrick
tbpatric@iu.edu
1
Department of Psychology, Bowling Green State University,
Bowling Green, OH, USA
2
Indiana University, 1101 E. 10th St., Bloomington, IN 47405, USA
and sensorimotor prediction (Barnes & Jones, 2000; Barnhart
et al., 2018; Engel et al., 2001; Jones, 1976; Jones & Boltz,
1989; Large, 2008; Large & Jones, 1999; McAuley & Jones,
2003; van Noorden & Moelants, 1999). Rhythmic activity is
indeed characteristic of neural populations (Buzsaki et al.,
2013; Schroeder & Lakatos, 2009). Intrinsic and event-related
oscillations are usually categorized into delta (0.5–3.5 Hz),
theta (4–7 Hz), alpha (8–12 Hz), beta (13–30 Hz) and gamma
(> 30 Hz) frequencies (Buzsáki & Draguhn, 2004; Engel &
Fries, 2010). Cortical oscillations appear to play a role in syn-
chronizing activity across brain regions. Evidence suggests that
higher frequency oscillations are utilized for localized interac-
tion whereas lower frequencies are characteristic of longer-
range interaction (Gupta & Chen, 2016; Sarnthein et al.,
1998; von Stein & Sarnthein, 2000).
Additionally, differing frequencies are also associated with
particular cognitive processes. Delta, theta, and alpha frequen-
cy waves have been associated with initial learning processes
as well as visual perception and imagery, working memory
retention and internal timekeeping (Arnal & Giraud, 2012;
Engel & Fries, 2010; Keitel et al., 2019; Lakatos et al.,
2008; Portnova et al., 2011; Saleh et al., 2010; Schroeder &
Atten Percept Psychophys (2022) 84:2360–2383
Lakatos, 2009; von Stein & Sarnthein, 2000). Whereas beta
oscillations are most commonly associated with motor func-
tions (Baker, 2007; Gu et al., 2015; Wiener & Kanai, 2016).
Beta band oscillations are also linked with temporal process-
ing (Merchant et al., 2013). They have been implicated in the
entrainment with both visual and auditory rhythms, as well as
the synchronization of activity between auditory and motor
regions (Arnal, 2012; Baker, 2007; Comstock et al., 2018a;
Engel & Fries, 2010; Fujioka et al., 2009, 2012; Fujioka et al.,
2015; Salmelin et al., 1995; Salmelin & Hari, 1994).
Gamma band activity is related to the selective orientation
of attention toward expected sensory events (Bhattacharya
et al., 2001; Bouyer et al., 1981; Buschman & Miller, 2007;
Fell et al., 2003; Fries et al., 2001; Fries et al., 2002; Fries
et al., 2008; Gruber et al., 1999; Jenson et al., 2007; Mitchell
et al., 2007; Müller et al., 2001; Munk et al., 1996; Snyder &
Large, 2005; Steriade & Amzica, 1996; Steriade et al., 1996;
Tallon-Baudry & Bertrand, 1999; Tallon-Baudry et al., 1997;
Vidal et al., 2006; Von Stein et al., 2000).
More generally, Arnal and Giraud (2012) propose that low-
frequency oscillations (delta-alpha) are more often involved in
predicting when an event will occur, whereas higher frequen-
cy oscillations are more often associated with the prediction of
what the event will be. Still, it is unlikely that a particular
frequency band is solely limited to a single cognitive function
(Arnal & Giraud, 2012; Engel & Fries, 2010). Almost all
frequency bands have been linked with temporal processing
at some level (Wiener & Kanai, 2016). This reflects the sig-
nificant contribution of these oscillations in processing the
passage of time within different contexts. This is most clearly
revealed in the dynamic response of these oscillations in the
presence of temporally consistent sensory input. Exposure to
auditory or visual rhythms produces neural responses at the
exact frequency of these sensory events (Barnes & Jones,
2000; Escoffier et al., 2010; Henao et al., 2020; Large &
Jones, 1999; McAuley & Jones, 2003; Nozaradan et al.,
2012; Tierney & Kraus, 2014; Van Noorden & Moelants,
1999). The entrainment of oscillatory activity with the rate
of an external stimulus produces steady-state evoked poten-
tials (SSEPs). SSEPs are stimulus driven and frequency
locked to the rate of sensory events, allowing for dynamic
attentional processes to be directed toward perceptually rele-
vant moments in time (Engel et al., 2001).
Evidence for the alignment of rhythmic attentional process-
es is reflected in the enhancement of perceptual abilities when
stimuli are presented simultaneously with a perceived beat
(Bolger, et al., Bolger, Coull, & Schon, 2013, b; Brochard
et al., 2013; Correa et al., 2005; Coull & Nobre, 1998; Jones
et al., 2011; Jones et al., 2002; Kösem & van Wassenhove,
2012; Mathewson et al., 2009; McDonald & Ward, 2000;
Miller et al., 2013; Nobre, 2001; Noesselt et al., 2008;
Rohenkohl et al., 2012; Schroeder et al., 2010; Ten Oever
et al., 2014; Zou et al., 2012).
2361
The entrainment of attentional pulses with the temporal
regularities of environmental stimuli serves to minimize the
processing of irrelevant sensory events, while focusing atten-
tional resources at the most relevant moments in time (Cravo
et al., 2013; Herrmann et al., 2016; McAuley & Fromboluti,
2014; Miller et al., 2013; Nobre, 2001; Rohenkohl et al.,
2012; Thut et al., 2011; Voss et al., 2006).
There is growing evidence supporting the notion that sen-
sorimotor attention is in fact a dynamic, rhythmic process
(Fiebelkorn et al., 2013; Fiebelkorn & Kastner, 2019;
Helfrich et al., 2018; Zalta et al., 2020). Brain rhythms have
been implicated in the dynamic perception of sensory events
in the environment (Buhusi & Meck, 2005; Busch et al., 2009;
Costa-Faidella et al., 2017; Engel & Fries, 2010; Laje &
Buonomano, 2013; Mathewson et al., 2009; Merchant &
Yarrow, 2016; Rohenkohl & Nobre, 2011). The identification
of these sensory events occurs as a result of environmental
sampling that is modulated by oscillating brain rhythms
(Dugue et al., 2015; Fiebelkorn et al., 2013; Fiebelkorn &
Kastner, 2019; Helfrich et al., 2018; Henry et al., 2014;
Henry & Obleser, 2012; Landau & Fries, 2012).
Environmental sampling involves the rhythmic scanning of
the external environment for relevant stimuli as well as the
subsequent enhancement of the processing of relevant stimu-
lus features. Rhythmic oscillations in cortical excitability are
believed to be the primary mechanism by which we select and
focus our attention toward external stimuli (Calderone et al.,
2014; Lakatos et al., 2008; Schroeder & Lakatos, 2009).
Interestingly, researchers have introduced the notion that the
seemingly continuous experience of the world is instead con-
structed from the rapid, sequential sampling of the environ-
ment (Fiebelkorn & Kastner, 2019; Helfrich et al., 2018).
Recent evidence suggests that the rate of environ-
mental sampling is constrained by the activity of the
motor system (Zalta et al., 2020). The motor system’s
interaction with perceptual processes provides a basis
for sensorimotor prediction, allowing us to respond ap-
propriately to future events (Merchant & Yarrow, 2016).
This interaction has been proposed to occur via the
coupling of gamma band oscillations across sensory
and motor regions (Gupta & Chen, 2016).
Auditory rhythms in particular initiate rhythmic activation
in both the auditory and motor systems (Arnal, 2012; Chen
et al., 2006; Chen et al., 2008; Janata et al., 2012; Large &
Snyder, 2009; Penhune et al., 1998; Ross et al., 2016; Thaut
et al., 2014). This occurs as a result of the high degree of
connectivity between the auditory and motor systems
(Comstock et al., 2018b; Grahn, 2009; Grahn et al., 2011;
Grahn & Brett, 2007; Kung et al., 2013; Nozaradan et al.,
2017; Patel & Iversen, 2014; Phillips-Silver & Trainor,
2005; Phillips-Silver & Trainor, 2007; Zatorre et al., 2007).
The modality-appropriateness hypothesis proposes that
sensory information is ultimately obtained from the modality
2362
that is best suited for processing that type of information
(Welch & Warren, 1980; Welch & Warren, 1986). For exam-
ple, visual input provides the most accurate spatial informa-
tion such as location, movement, and distance, and is therefore
the dominant source for this information (Alais & Burr, 2004;
Bertelson & Aschersleben, 1998; Hove et al., 2010; Hove
et al., 2012; Hove et al., 2013; Kitajima & Yamashita, 1999;
Soto-Faraco et al., 2003; Soto-Faraco et al., 2004). The audi-
tory system is apparently more effective at processing the
precise timing and frequency of sensory events (Burr et al.,
2009; Comstock et al., 2018a, 2018b; Conway &
Christiansen, 2005; Rencanzone, 2003; Repp & Penel, 2002;
Welch et al., 1986).
Conflicting information from the two senses does not seem
to interfere with the process of multisensory integration.
Rather, the dominant sense is known to capture the other
(Guttman et al., 2005). For example, when the directions of
auditory and visual movement are in conflict, the resulting
perception is that the auditory stimulus is moving in the direc-
tion of the visual stimulus (Soto-Faraco et al., 2004).
Similarly, temporal judgments relating to visual stimuli are
often biased by the rate of simultaneously presented sounds
(Fendrich & Corballis, 2001; Rencanzone, 2003).
Research has revealed that spatiotemporal visual rhythms can
promote the perception of a beat as well as the synchronization of
motor activation (Chen et al., 2006; Grahn, 2012; Hove et al.,
2012; Su & Jonikaitis, 2011). However, the encoding of a beat in
the visual modality is believed to involve the automatic transfor-
mation of visual signals into an auditory-motor representation,
thereby promoting this visuomotor connection (Grahn, 2012;
Guttman et al., 2005; Karabanov et al., 2009; Su & Jonikaitis,
2011). While the findings are mixed regarding the automatic
nature of the auditory encoding of visual rhythms, research has
revealed that prior exposure to auditory rhythms can promote
more accurate temporal judgments regarding visual rhythms
(McAuley & Henry, 2010). Prior exposure to visual rhythms
does not seem to have the same priming effect on auditory
rhythm perception (Grahn et al., 2011). So, while some degree
of motor activation can occur as a result of exposure to visual
rhythms, the motor response is noticeably greater for rhythms
presented in the auditory modality (Grahn et al., 2011;
Balasubramaniam, 2018a, 2018b’; Jancke et al., 2000).
The strength of the connection between the auditory and
motor systems promotes the spontaneous synchronization of
movement with the timing of a beat, regardless of our inten-
tions to do so (Bouvet et al., 2019; Varlet et al., 2018).
Because of the motor system’s role in event timing, this link
promotes greater temporal accuracy for and synchronization
with events perceived in the auditory modality (Comstock
et al., 2018a, 2018b; Grahn, 2012; Grahn et al., 2011; Hove
et al., 2013; McAuley & Henry, 2010; Pasinski et al., 2016).
Recent research has revealed that both time perception and
movement synchronization is further increased for lower
Atten Percept Psychophys (2022) 84:2360–2383
pitched tones (≤ 130 Hz), implying a more pronounced neural
response to deeper tones in associated motor centers (Hove
et al., 2014; Lenc et al., 2018; Varlet et al., 2018).
Motor activation during passive listening to auditory
rhythms also recruits the motor system’s predictive influence
over attentional processes (Schubotz, 2007). The motor sys-
tem’s influence over the frequency and timing of attentional
and sensorimotor oscillations may provide the neural basis for
temporally oriented sensory prediction (Arnal, 2012; Arnal &
Giraud, 2012; Morillon et al., 2014; Morillon et al., 2015). In
other words, rhythmic auditory stimuli are capable of
entraining sensorimotor attention (Arnal, 2012; Bolger,
Coull, & Schon Bolger, Coull, & Schon, 2013, b;
Nozaradan et al., 2012; Nozaradan et al., 2015). Prolonged
exposure to a rhythmic auditory stimulus leads to the more
precise entrainment of neural activity at the frequency of the
input stream (Henao et al., 2020; Henry et al., 2014; Henry &
Obleser, 2012; Jones et al., 2002; Schroeder et al., 2010;
Schroeder & Lakatos, 2009; Will & Berg, 2007).
Timely allocation of attentional resources in the form
of entrainment implies that attentional processes are of-
ten driven by temporal predictions (Engel et al., 2001;
Proksch et al., 2020). The alignment of endogenous at-
tentional rhythms with an ongoing sensory event is also
believed to provide the basis for the temporal experi-
ence of that event (Jones & Boltz, 1989; Kösem
et al., 2014; Large & Jones, 1999; Portnova et al.,
2011). It is well established that the amount of attention
directed toward an event has a significant impact on
that event’s perceived duration (Coull et al., 2004;
Flaherty, 1993). Specifically, the greater the amount of
attention devoted to the temporal aspects of an event,
the longer it is judged to last (Casini & Macar, 1997).
Similarly, increasing the temporal frequency of an ex-
ternal stimulus can also increase an event’s perceived
duration (Kanai et al., 2006; Portnova et al., 2011).
Duration judgments are strongly correlated with an in-
crease in the frequency of neural responses during a
given interval (Eagleman & Pariyadath, 2009; Horr
et al., 2016). In particular, research has shown that tem-
porally regular or isochronous sounds or flashes of light
can alter the subjective experience of time (Droit-Volet
& Wearden, 2002; Eagleman & Pariyadath, 2009; Horr
et al., 2016; Horr & Di Luca, 2015a, 2015b; Kanai
et al., 2006; Penton-Voak et al., 1996; Portnova et al.,
2011; Rammsayer & Lima, 1991; Thomas & Brown,
1974; Treisman et al., 1990; Wearden et al., 1999;
Wearden et al., 2007). As the pattern moves further
away from isochrony, its ability to influence the subjec-
tive experience of a duration and induce neural entrain-
ment is gradually reduced (Horr & Di Luca, 2015a;
Lenc et al., 2018). These findings strongly suggest that
exposure to isochronous rhythms of higher rates can
Atten Percept Psychophys (2022) 84:2360–2383
create fundamentally different temporal experiences of
the same dynamic event by exploiting the motor sys-
tem’s influence over attentional processes. It is currently
unclear if dynamic mental representations are also vul-
nerable to this temporal manipulation.
Inferred motion
Inferred motion (sometimes called “predicted motion”) exper-
iments have long been used to examine the nature of how we
are able to judge the current position of moving objects that
are not available to visual perception (Alderson & Whiting,
1974; Caird & Hancock, 1994; Gottsdanker, 1955; Lyon &
Waag, 1995; Peterken et al., 1991; Rosenbaum, 1975;
Schiff & Oldak, 1990). Most inferred motion tasks in-
volve requiring participants to judge the time point at
which a moving object will arrive at a specified location
(Makin, 2018; Rosenbaum, 1975).
In inferred motion tasks, an object is visible for a specific
duration, known as the viewing time or viewing interval. At a
certain point during its movement, it disappears by moving
behind an occlusion of some kind. The assumption in these
tasks is that the object continues to move, though out of sight
(Alderson & Whiting, 1974; Peterken et al., 1991). A speci-
fied location is provided as the endpoint or point of contact for
the moving object. Participants are required to respond when
they believe the moving object has arrived at or made contact
with the endpoint. This is often referred to as the object’s time
to contact (TTC). The difference observed between the time
point of the participants’ judged TTC and the actual TTC of
the moving object is the judgment's timing error (Battaglini
et al., 2013).
The cognitive mechanism being measured in these tasks is
referred to as motion extrapolation, which is the ability to form
internal representations of an invisible object’s motion
(Rosenbaum, 1975; Tresilian, 1995). Motion extrapolation
may depend to some degree on a cognitive model constructed
based on the object’s visible motion prior to its disappearance
(Tresilian, 1995). Researchers have found support for the in-
volvement of a perceptual memory system in the visual mo-
dality that may transfer prior experience into later representa-
tions during the period of occlusion (Battaglini et al., 2013).
Others have concluded that motion extrapolation involves
some degree of visuospatial attention in tracking moving,
but unseen objects (Lyon & Waag, 1995). Previous research
points to a possible overlap in the systems required for oculo-
motor control and those required for extrapolation judge-
ments, regardless of whether visual tracking or fixation is
involved (Makin & Poliakoff, 2011).
Factors such as travel time can influence the accuracy of
participants’ judgments regarding the TTC of a moving object
(Caird & Hancock, 1994; Schiff & Oldak, 1990). More spe-
cifically, increasing the period of invisibility relative to the
2363
period of visibility tends to increase the degree of absolute
error in judgments (Alderson & Whiting, 1974; Peterken
et al., 1991). Errors in estimating TTC are found even when
the period of invisibility exceeds 1 s, reflecting a rapid break-
down of the cognitive mechanisms used to maintain this in-
ternal representation (Bennett et al., 2010). The rate of move-
ment also influences the accuracy of temporal judgments.
Judgment timing error tends to increase as the speed of the
moving object decreases (Lyon & Waag, 1995; Schiff &
Oldak, 1990). In general, participants tend to overestimate
the TTC for slower velocities and underestimate the TTC for
faster velocities (Bennett et al., 2010). This provides some
evidence that temporal estimations are based on internal pro-
cesses that can be influenced by external sensory events.
Surprisingly, few studies have examined the spatiotempo-
ral predictions of occluded movement in the presence of both
visual and auditory cues. The few that have been conducted
reveal an advantage in accuracy of spatiotemporal predictions
of audiovisual stimuli relative to stimuli perceived strictly in
the auditory domain, but no greater advantage in accuracy
when stimuli are perceived only in the visual domain
(Hofbauer et al., 2004; Peterken et al., 1991; Schiff &
Oldak, 1990). Previous research has identified the potential
for auditory interference in inferred motion tasks requiring
motion extrapolation (Gordon & Rosenblum, 2005; Prime &
Harris, 2010). These results seem to demonstrate the primary
importance of visual input in motion extrapolation relative to
strictly auditory or even audiovisual input (DeLucia et al.,
2016; Hofbauer et al., 2004).
More recent research has revealed that auditory cues are
not completely ignored during inferred motion tasks and can
indeed influence the timing of spatiotemporal judgments
(Gordon & Rosenblum, 2005; Prime & Harris, 2010).
Ascending pitch of pure tones during vertical movement has
been shown to produce earlier judgments of TTC of an oc-
cluded object (King & Prime, 2019). Evidence seems to sug-
gest that the addition of supplementary auditory cues to visual
stimuli in inferred motion tasks does not ensure improved
accuracy (DeLucia et al., 2016; Gordon & Rosenblum,
2005). Rather, the addition of auditory information may in-
stead provide external temporal cues that can be incorporated
into participant’s judgments of inferred motion (Prime &
Harris, 2010).
Recent evidence has also pointed to the possibility that
auditory rhythms of different speeds can have a measurable
impact on the timing of motion extrapolation when they are
presented during a period of occlusion (Chotsrisuparat et al.,
2017). Similar to our current design, this prior research ex-
plored the impact of differing rhythms on participant’s judg-
ments of TTC. While the results of this prior research did
reveal a noticeable influence of rhythm speed on temporal
predictions, the researchers failed to explore the influence of
differing occlusion sizes and travel distances on the timing of
2364
these predictions. Currently it is unclear if and how these
factors may differentially impact the nature of these predic-
tions. Consistent with earlier findings (Chotsrisuparat et al.,
2017), we propose that concurrent exposure to regularly oc-
curring auditory events during an inferred motion task will
systematically impact participant’s judgments of TTC. We
theorize that by recruiting the motor system’s influence over
sensorimotor attentional processes through exposure to iso-
chronous rhythms, it is possible to alter the temporal experi-
ence of sensory events. This influence could be observed in
the form of consistent differences in perceived time of arrival
of an object that is assumed to be moving but cannot be seen.
The present research also aims to examine the impact of au-
ditory rhythms of varying rates on judgments of arrival time in
an inferred motion task that varies the travel time by utilizing
occlusions of different sizes. It is predicted that auditory
rhythms of varying rates, each having their own unique effects
on participants’ attentional processes, will produce distinct
patterns of temporal judgments. Additionally, it is predicted
that the differing periods of occlusion may potentially magni-
fy these effects.
Experiment 1
The present research was designed to examine whether judgments
of TTC can be influenced by exposure to regularly occurring
auditory stimuli that can affect the subjective experience of time
duration. Isochronous auditory rhythms are known to influence
temporal perception in a particular manner. Additionally, judg-
ments of duration are known to correlate with the frequency of
neural responses during a given interval (Eagleman & Pariyadath,
2009; Horr et al., 2016; Portnova et al., 2011). Therefore, it was
expected that exposure to more frequent auditory events in the
form of faster rhythms would result in the relatively premature
anticipation of the object’s TTC when compared to judgments
made in the presence of slower rhythms or silence. This was
predicted to occur across all levels of occlusion. Additionally, we
predicted that the anticipated temporal underestimation resulting
from exposure to faster rhythms would increase with the length of
the period of invisibility. Greater exposure to the temporal infor-
mation contained in these fast rhythms was expected to become
increasingly more influential as the amount of visual information is
made less available. Therefore, the effect of rhythm should be
greater when the period of invisibility is long than when it is short.
Method
Participants
One hundred and sixty-five participants (85 women and 80
men) were recruited from Bowling Green State University in
Atten Percept Psychophys (2022) 84:2360–2383
Ohio. Commonly used software for a priori power analysis
accommodates designs in which there is only one repeated-
measures factor. However, as described below, our experi-
mental design consisted of more than one repeated-measures
factor. Even for a one-factor repeated-measures design – with
three conditions, a power of 0.8, and a small, estimated
Cohen's f effect size of 0.1 – the required sample would be
N = 965. Given the resources available for the project, our data
collection strategy was simply to include as many participants
as we could within the span of an academic term.
The participants consisted of undergraduate students rang-
ing in age from 18 to 22 years participated in the experiment.
Participants received course credit in exchange for their par-
ticipation. Informed consent was obtained from each partici-
pant prior to the experiment being conducted.
Materials and design
The experiment was a 3 (size of occlusion: small (25%), me-
dium (50%), large (75%)) × 3 (sound condition: fast rhythm,
slow rhythm, silent) design, with both factors varying within
participants. Note that in this design, the length of the period
of invisibility necessarily varied with occlusion size, because
we used occlusion size as a way to manipulate the length of
the period of invisibility period. Participants completed this
experiment in a psychology laboratory on the campus of
Bowling Green State University. The visual stimuli were
displayed on a Dell Optiplex 780 desktop computer with a
Pentium 4 processor and 4GB of RAM. The monitors were
19-in., flat panel, wide format (16:9) Dell monitors and the
screen resolution was 1,440 × 900 pixels; refresh rate 75 Hz).
The operating system was Windows 7 Enterprise.
Stimuli
Two-dimensional stimuli were constructed using Synfig
Studio v. 1.3.10 open source 2-D animation software.
The auditory stimuli were created using the trial version
of FL Studio 20.1.2 music production software. The
auditory stimuli were then edited using WavePad
Sound Editor v. 6.44 audio editing software developed
by NCH Software. The auditory patterns created includ-
ed 12-s clips of fast and slow isochronous rhythms. The
slow rhythm consisted of 200-ms, 60-Hz tones alternat-
ing with 320-ms pauses (roughly 2 Hz; 120 bpm) and
the fast rhythm consisted of 200-ms, 60-Hz tones alter-
nating with 50-ms intervals (4 Hz; 240 bpm) (see Fig.
1). The rates of these patterns were selected because
they are known to fall within the frequency range typ-
ically heard in music (London, 2004). Additionally,
rates ranging from 1.25 to 5 Hz have also been utilized
in previous research demonstrating neural synchroniza-
tion (Lenc et al., 2018; Tierney & Kruas, 2014).
Atten Percept Psychophys (2022) 84:2360–2383 2365

Fig. 1 The tonal structure of the auditory patterns used for both
experiments

All auditory stimuli were presented at 70 dB. Each rhythm

was presented from both monitor speakers simultaneously and
contained no spatial information. The rhythmic auditory pat-
terns and the animated videos were synchronized using the
Adobe® Premiere Pro 2018 timeline-based video editing ap-
plication. The experiment itself was constructed in
OpenSesame v. 3.2.7, an open source, graphical experiment
program for the social sciences.
One of three levels of a blue colored occlusion, consisting
of small, medium, and large sizes were presented on a gray
background that was 600 × 900 pixels (px). The coordinates
of the display were oriented at 300 px to -300 px from top to
bottom and from -450 px to 450 px, left to right. Each animat-
ed video included a red disc that was 20 px in diameter (rough-
ly 2° visual angle) located at -440 px × 0 px (far left center) of
the display that served as the moving object. On the opposite
side of each occlusion was a white disc 20 px in diameter
(roughly 2° visual angle) that represented the point of collision
for the moving disc. This disc was located 15 px passed the far Fig. 2 The graphical layout of Experiment 1. The time displayed in the
edge of the occlusion in each condition. The small occlusion darker region (blue, in color versions of the figure) represents the period
of invisibility. Visible time remained constant at 1,000 ms for all
made up 25% of the animated display (h = 600 px, w = 225 occlusion sizes
px), the medium occlusion made up 50% of the display (h =
600 px, w = 450 px), and the large occlusion made up 75% of
the display (h = 600px, w = 675 px). The disc was designed to period) for the large occlusion. These periods of invisibility
move at a constant rate of approximately 80.5 pixels/s in order were significantly longer than those traditionally used (1–4 s)
to better isolate the nature of the relationship between the to create more measurable differences in performance
sound conditions and the different occlusion conditions. The (Alderson & Whiting, 1974; Chotsrisuparat et al., 2017; Lyon
disc began its movement 1,000 ms after the presentation of the & Waag, 1995; Prime & Harris, 2010) (see Fig. 2).
display. The visible time was held constant for all trials at
1,000 ms before the disc disappeared behind each occlusion.
The actual time to contact for the red disc to reach the white Procedure
disc in the small occlusion condition was 4,100 ms (3,100-ms
invisibility period), 6,900 ms (5,900-ms invisibility period) for The three levels of the sound condition (silent, slow, fast) and
the medium occlusion and 9,700 ms (8,700-ms invisibility three levels of occlusion size (25%, 50%, 75%) yielded nine
2366 Atten Percept Psychophys (2022) 84:2360–2383

conditions. Each participant experienced seven trials in each

condition, yielding a total of 63 trials.
Participants were run one at a time. They were seated in
front of the computer monitor and provided with identical
instructions prior to beginning. Two monitor speakers were
placed on both sides of the screen. Participants were also pre-
sented with written instructions on the screen prior to starting.
Participants were instructed to press the spacebar at the precise
moment they felt the red disc would reach the white disc on
the opposite side of the occlusion. The red disc would never
re-appear after it traveled behind the occluded area.
Participants had to anticipate when this moment would occur
and respond at that moment. All participants were required to
perform three practice trials before beginning the actual ex-
periment. Each of the three practice trials included an example
of one of the sound conditions and one of the occlusion con-
ditions. Each trial began by instructing the participants to click
a "continue" button at the bottom of the screen with the mouse
cursor in order to hear two tones that were separated by 1 s.
This was to be used as a reference for their temporal judgment.
Once the tones were presented, they would then be asked to
click "continue" again to start the actual trial.
Each experimental trial began with the layout of the screen Fig. 3 Experiment 1. Error in judged time-to-contact (TTC) as a function
presented. The red disc would be stationary, prior to its initial of occlusion size, sound condition, and experiment. The figure shows
movement. In all trials the disc would move from left to right. means surrounded by 95% confidence intervals that are adjusted
When the disc reached the area of occlusion, either no audi- (Cousineau, 2005) for within-subject comparisons. The spikes indicate
± 1 SD. Offset to the right of the means are dark dots indicating the
tory pattern was presented, or a fast or slow isochronous medians. In the background are point-clouds consisting of the
rhythm started to play. In the rhythm conditions, the sound individual scores (from which the summary statistics have been
continued until a response was given in the form of pressing computed). The shape of each point-cloud shows a frequency
the spacebar. No feedback was given after the responses were distribution: At each level on the vertical axis, the cloud's width
indicates the relative frequency of the binned scores
recorded. After that, participants were asked to click the "con-
tinue" button once again to hear the two tones. They then had
to click the "continue" button once more to start the next trial. The results were further examined using sets of planned com-
All 63 trials were presented in random order and took approx- parisons. (Note that we used planned comparisons with individ-
imately 30 min to complete. ual, un-pooled variances, rather than, say, linear contrasts be-
cause our ANOVA results violated the sphericity assumption,
and we did not want to risk computing and reporting contrasts
Results that might be based on faulty assumptions.) In Set A, three com-
parisons assessed differences in TTC judgment error across the
The actual TTC (for the red disc to reach the white disc) was levels of sound condition (averaged over the levels of occlusion
4,100 ms, 6,900 ms, and 9,700 ms for the small, medium, and size). The mean difference for the slow versus silent comparison
large occlusion size, respectively. The actual TTC was was small and non-significant, but there was a significant,
subtracted from the participant’s judged TTC to compute the medium-sized difference for the fast versus slow, and for the fast
error in judged TTC. versus silent comparisons (Table 1). Being planned comparisons,
Figure 3 illustrates the results. Using the Greenhouse- the tests within each set of comparisons were not corrected for
Geisser correction, a 3 × 3 analysis of variance on error in familywise error.
judged TTC, with both factors varying within participants, In Set B, another three comparisons assessed differences in
revealed a significant main effect of sound condition, TTC judgment error across the levels of occlusion size (aver-
F(1.83, 300.71) = 40.70, p < .001, η2 = .20, a significant main aged over the levels of sound condition). Each of those com-
effect of occlusion size, F(1.10, 180.93) = 134.37, p < .001, η2 parisons yielded a significant, medium-to-large-sized effect
= .45), and a significant but small interaction between the size (Table 1).
of the occlusion and sound condition (F(3.40, 557.99) = 8.13, In Set C, nine comparisons assessed differences in TTC
p <.001, η2 = .05). judgment error across the levels of sound condition, separately
Atten Percept Psychophys (2022) 84:2360–2383 2367

Table 1 Planned comparisons for mean differences in TTC judgment error (ms) in Experiment 1

Comparison Mean of differences SD of differences Cohen's d t* df p pHolm

Set A
Slow - Silent -35.53 561.34 -0.06 -0.81 164 .417 .417
Fast - Silent -391.99 703.92 -0.56 -7.15 164 <.001 <.001
Slow - Fast 356.46 576.75 0.62 7.94 164 <.001 <.001
Set B
Small - Medium 480.94 706.74 0.68 8.74 164 <.001 <.001
Large - Medium -752.44 737.19 -1.02 -13.11 164 <.001 <.001
Small - Large 1,233.39 1,343.23 0.92 11.79 164 <.001 <.001
Set C
Small
occlusion
Slow - Silent 48.94 429.93 0.11 1.46 164 .146 .291
Fast - Silent -215.13 515.22 -0.42 -5.36 164 <.001 <.001
Slow - Fast 264.07 483.62 0.55 7.01 164 <.001 <.001
Medium
occlusion
Slow - Silent -27.53 744.51 -0.04 -0.48 164 .635 .635
Fast - Silent -384.94 840.59 -0.46 -5.88 164 <.001 <.001
Slow - Fast 357.41 689.40 0.52 6.66 164 <.001 <.001
Large
occlusion
Slow - Silent -128.00 947.34 -0.14 -1.74 164 .085 .254
Fast - Silent -575.90 1071.16 -0.54 -6.91 164 <.001 <.001
Slow - Fast 447.90 974.57 0.46 5.90 164 <.001 <.001

t is Student's t-test. The value pHolm is the Holm-corrected p value (corrected for familywise error), computed separately for each set of comparisons

for each level of occlusion size. The results (Table 1) indicated
non-significant, small effects for the three slow-versus-silent
comparisons, but indicated significant, medium-sized effects
for all other comparisons.
Overall, the ANOVA, the planned comparisons, and Fig. 3
together indicate that the positivity of error in judged TTC (i.e.,
the overestimation of time-to-contact) was greater when the
accompanying auditory rhythm was slow than when it was fast
and was greater when the occlusion size was smaller rather than
larger. The significant interaction in the ANOVA appears to be
a result of the TTC judgment being especially high in the silent,
large-occlusion condition.
Discussion
Experiment 1 attempted to identify whether exposure to audi-
tory stimuli in the form of isochronous rhythms could impact
participants’ estimations of a hidden object’s actual TTC. If
differing rates of auditory rhythms were found to produce
consistent patterns of over or underestimations across condi-
tions, this would provide support for the proposed
manipulation of attentional processes that are believed to be
involved in the temporal perception of dynamic events. The
results of Experiment 1 are consistent with this conclusion.
Faster rhythms led to more premature judgments of TTC
across occlusion conditions relative to the other sound condi-
tions. These results demonstrate that increasing the rate of
regularly occurring auditory stimuli can alter the subjective
experience of a dynamic event in predictable ways, even when
the event is not visible. Contrary to predictions, the interaction
between rhythm and the period of invisibility was miniscule
and not clearly interpretable (though it was statistically
significant).
All of our predictions concerned the effects of rhythm and
the period of invisibility on the mean error in judged TTC. We
made no predictions regarding effects on the absolute devia-
tion of that mean judgment error from zero. However, previ-
ous studies have observed that increasing the period of invis-
ibility relative to the period of visibility tends to increase the
absolute error of temporal judgments in inferred motion tasks
(Alderson & Whiting, 1974; Peterken et al., 1991; Schiff &
Oldak, 1990). Progressively more premature estimations of
TTC across occlusion conditions reflects an apparent
2368
increased reliance on the temporal information contained
within the auditory rhythm. It appears that within the present
experiment, as visual information becomes increasingly un-
available, participants may rely to a greater extent on the tem-
poral information contained within the auditory rhythms as
alternative cues regarding the rate of movement. It should be
noted that the constant velocity of a moving object can be
misperceived if the object is only visible for short periods.
Previous research has discovered that when objects start mov-
ing at a constant speed, rather than slowly accelerating as in
natural movement, viewers perceive these objects as moving
faster at the start (Brouwer et al., 2002; Runeson, 1974).
However, these effects are found to be minimal as visibility
approaches 1,000 ms (Brouwer & Brenner, 2002).
Additionally, this effect would have applied to all conditions
and therefore would have been distributed evenly throughout
the experiment.
Attentional and sensory processes are considered to be
rhythmic (Buhusi & Meck, 2005; Calderone et al., 2014;
Fiebelkorn et al., 2013; Fiebelkorn & Kastner, 2019;
Helfrich et al., 2018; Laje & Buonomano, 2013; Lakatos
et al., 2008; Merchant & Yarrow, 2016; Schroeder &
Lakatos, 2009). It is important to consider the likelihood that
the nature of the attentional processes involved in perceiving
an ongoing sensory event are at least somewhat specific to
each particular event. These distinct internal states would be
preserved if we continue to attend to the spatiotemporal qual-
ities of the dynamic event they are constructed around. The
reliability of the attentional rhythms established in the pres-
ence of dynamic visual input in making accurate temporal
judgments would depend on their degree of alignment with
the rate of external events. While we did not directly examine
this, we propose that the increases in the mean error in judged
TTC from zero observed with greater periods of invisibility
occur as a result of the attentional processes involved in visu-
ally tracking a moving object becoming increasingly unreli-
able in the absence of confirmatory visual evidence. This
could effectively explain why the error in judged TTC tends
to increase along with the period of invisibility in inferred
motion tasks.
More specifically, this helps to explain the current
findings. If distinct attentional rhythms are established
or beginning to be established in the presence of brief
visual information, these rhythms would become in-
creasingly vulnerable to external influence in the ab-
sence of this sensory input. The replacement of visuo-
spatial information with auditory rhythms would neces-
sitate a shift in both the nature and the rate of the
attention required for processing the new stimulus.
The rate of these newly constructed attentional rhythms
would more closely align with the rate of the auditory
rhythm (Arnal, 2012; Eagleman & Pariyadath, 2009;
Henry & Obleser, 2012; Henry et al., 2014; Horr
Atten Percept Psychophys (2022) 84:2360–2383
et al., 2016; Jones et al., 2002; Nazoradan et al.,
2012; Schroeder & Lakatos, 2009; Schroeder et al.,
2010; Tierney & Kraus, 2014). Faster auditory rhythms are
known to create the impression that more time has passed within
a given duration, thereby leading to earlier responses in timing
tasks (Kanai et al., 2006; Portnova et al., 2011). The replacement
of visual information with incompatible auditory information
allowed us to predictably alter the participants’ experience of
time during this period of invisibility. Although we did not col-
lect electrophysiological measures, we propose that this resulted
from the manipulation of the rate of the attentional rhythms in-
volved in the representation of imagined dynamic events. The
results of Experiment 1 revealed response times that are consis-
tent with these proposed effects.
The present experiment examined how altering the
relationship between the rate of auditory rhythms and
the length of the period of invisibility would impact
participants’ judgments of TTC in an inferred motion
task. The results support our hypotheses. They also pro-
vide evidence for the proposal that the underlying
mechanism behind motion extrapolation involves the
formation of attentional rhythms required for preserving
the temporal nature of movement when it is visually
unavailable. When visual input is removed, the atten-
tional rhythms involved in the initial perception of a
dynamic event will likely become increasingly unreli-
able indicators of the temporal nature of that event.
The further absence of visual input consistent with lon-
ger predictive periods, will effectively produce greater
error in judged TTC. However, when an alternative
source of sensory information is presented, capable of
reestablishing an internal rhythmicity, the rate of this
attentional rhythm will shift toward the rate of these
regularly occurring sensory cues. When these cues pro-
vide temporal information that is inconsistent with the
unfolding of the actual event, the error in judged TTC
increases in a predictable direction. This is what we
observed. It is also what would be expected if the in-
ternal mechanism involved in accurate motion extrapo-
lation relied on both the establishment and maintenance
of an internal rhythmicity consistent with the rate of an
external event.
It remains unclear whether similar effects would be
observed if visibility is increased relative to invisibil-
ity. If participants are given more exposure to dynam-
ic visual information, will they still produce such
consistent patterns of responses? Or will prolonged
visual exposure protect against the effects of auditory
rhythms on promoting timing errors? Experiment 2
was designed to explore how the error in judged
TTC is impacted when participants are exposed to
fast isochronous rhythms following longer periods of
visibility.
Atten Percept Psychophys (2022) 84:2360–2383 2369

Experiment 2

Experiment 2 was designed to replicate the results of

the first experiment while more precisely examining
the influence of varying periods of visibility relative to
invisibility, on the degree of error in judged TTC. By
maintaining a constant travel time and travel distance
for all levels of occlusion it will be possible to explore
this relationship. Again, previous research has identified
the length of the invisibility period relative to viewing
time as a major contributor to the amount of judgment
error in inferred motion tasks requiring temporal judg-
ments (Alderson & Whiting, 1974; Peterken et al.,
1991; Schiff & Oldak, 1990). If the length of the invis-
ibility period relative to the visibility period is a critical
factor in producing greater error in judged TTC, it is
expected that longer invisibility periods should continue
to promote increasingly greater error in judged TTC.
Additionally, if the rate of an internal representation of
a dynamic event can be influenced by external factors,
as was observed in Experiment 1, the error in judged
TTC should again be biased towards a particular direc-
tion. Therefore, exposure to fast rhythms during invisi-
bility periods should result in consistently premature
estimations of TTC, and the degree of this prematurity
should increase along with increases in the invisibility
period.

Method

Participants

Ninety participants (53 women and 37 men) were recruited

from Bowling Green State University in Northwest Ohio.
These participants consisted of undergraduate students
ranging in age from 18 to 22 years. Participants received
research credit for psychology or communication classes in
exchange for their participation. Prior informed consent
was obtained from each participant. The number of partic-
ipants was determined in the same manner as was done for
Experiment 1.
Materials and design
The experiment was again a 3 (amount of occlusion: small
(25%), medium (50%), large (75%) × 3 (sound condition:
slow rhythm, fast rhythm, silent) within-participant design.
Participants completed this experiment in the same cognitive
psychology laboratory on campus at Bowling Green State
University on the same equipment used in Experiment 1.
Fig. 4 The graphic layouts of Experiment 2. The time displayed in the
darker region (blue, in color versions of the figure) represents the period
of invisibility. The time in the gray region represents the visible
time
Stimuli
The auditory stimuli were the same as those used in
Experiment 1. In the present experiment, the sizes of
the occluded areas remained the same. However, there
was additional, initial travel time that varied across the
levels of the occlusion-size so that the total travel time
was constant at 10.5 s, regardless of occlusion size. In
the small occlusion condition (25%), the moving disc
was visible for 7,300 ms (3,200-ms invisibility period),
in the medium occlusion condition (50%) it was visible
for 4,700 ms (5,800-ms invisibility period), and in the
2370
large occlusion condition (75%) the moving disc was
visible for 2,100 ms (8,400-ms invisibility period) (see
Fig. 4).
Movement velocity was approximately the same as it was
in the first experiment (82.7 pps). Similar to Experiment 1, the
velocity of the moving disc remained constant for all trials.
Procedure
The procedure was the same as that of Experiment 1.
However, the duration of the experiment was increased to
approximately 35 min as a result of increasing the length of
the individual trials.
Results
The actual time for the red disc to reach the white disc (the
actual TTC) was 10.5 s for all occlusion conditions. As in
Experiment 1, the actual TTC was subtracted from the partic-
ipant’s judged TTC to compute a judgment error score.
Figure 5 shows the pattern of the results. Using the
Greenhouse-Geisser correction, a 3 × 3 repeated-measures
analysis of variance revealed a small, significant main effect
of sound condition (F(1.61, 143.10) = 6.95, p = .003, η2 =
.07), and a small, significant interaction between sound con-
dition and occlusion size (F(3.39, 301.32) = 3.86, p = .007, η2
= .04), but no main effect of occlusion size, F(1.33,118.63) =
1.48, p = .23, η2 = 0.02.
Fig. 5 Error in judged time-to-contact (TTC) as a function of occlusion
size, sound condition, and experiment. The figure shows means
surrounded by 95% confidence intervals that are adjusted (Cousineau,
2005) for within-subject comparisons. The spikes indicate ± 1 SD.
Offset to the right of the means are dark dots indicating the medians. In
Atten Percept Psychophys (2022) 84:2360–2383
As in Experiment 1, The results were further examined
using sets of planned comparisons. In Set A, three compari-
sons assessed differences in TTC judgment error across the
levels of sound condition (averaged over the levels of occlu-
sion size). The mean difference for the slow versus silent
comparison was small and non-significant, but there were sig-
nificant, small-to-medium-sized differences for the fast versus
the slow, and for the fast versus the silent comparisons
(Table 2).
In Set B, three comparisons assessed differences in
TTC judgment error across the levels of occlusion size
(averaged over the levels of sound condition). The ef-
fects were small, and only the small-versus-medium
comparison was statistically significant (Table 1).
In Set C, nine comparisons assessed differences in TTC
judgment error across the levels of sound condition, separately
for each level of occlusion size. The results (Table 2) indicated
small-to-medium-sized, significant effects for the three slow-
versus-fast comparisons. The results were inconsistent for
comparisons involving the silent condition.
Overall, the results indicated that the positivity of the
error in judged TTC (i.e., the overestimation of time-to-
contact) was greater when the accompanying auditory
rhythm was slow than when it was fast. As in
Experiment 1, the significant interaction in the
ANOVA appears to be a result the TTC judgment being
especially high in the silent, large-occlusion condition.
the background are point-clouds consisting of the individual scores (from
which the summary statistics have been computed). The shape of each
point-cloud shows a frequency distribution: At each level on the vertical
axis, the cloud's width indicates the relative frequency of the binned
scores. The Experiment 1 data are included for purposes of comparison
Atten Percept Psychophys (2022) 84:2360–2383 2371

Table 2 Planned comparisons for mean differences in TTC judgment error (ms), in Experiment 2

Comparison Mean of differences SD of differences Cohen's d t df p pHolm

Set A
Slow - Silent 21.89 438.96 0.05 0.47 89 .637 .637
Fast - Silent -181.73 556.69 -0.33 -3.10 89 .003 .008
Slow - Fast 203.62 684.66 0.30 2.82 89 .006 .012
Set B
Small - Medium 155.72 605.45 0.26 2.44 89 .017 .050
Large - Medium 108.23 813.69 0.13 1.26 89 .210 .421
Small - Large 47.49 1,138.23 0.04 0.40 89 .693 .693
Set C
Small
occlusion
Slow - Silent 100.56 435.07 0.23 2.19 89 .031 .217
Fast - Silent -73.83 475.76 -0.16 -1.47 89 .145 .434
Slow - Fast 174.39 528.70 0.33 3.13 89 .002 .019
Medium
occlusion
Slow - Silent 96.71 680.02 0.14 1.35 89 .181 .361
Fast - Silent -103.32 763.31 -0.14 -1.28 89 .203 .203
Slow - Fast 200.03 878.67 0.23 2.16 89 .034 .201
Large
occlusion
Slow - Silent -131.60 802.21 -0.16 -1.56 89 .123 .493
Fast - Silent -368.00 844.61 -0.44 -4.13 89 <.001 <.001
Slow - Fast 236.40 1,068.22 0.22 2.10 89 .039 .193

t is Student's t-test. The value pHolm is the Holm-corrected p value (corrected for familywise error), computed separately for each set of comparisons

Comparison across experiments Discussion

To assess whether there were significant differences be-
tween Experiment 1 and Experiment 2, we conducted an
analysis of variance that included experiment as a fac-
tor. Sound condition and occlusion size varied within-
subjects, and the dependent variable was error in judged
TTC.
There was no main effect of experiment (p =.822).
However, there were two significant interactions in-
volving experiment as a factor. One was a small
experiment-by-sound-condition interaction, F(1.87,
474.28) = 3.81, p = .025, η2 = 0.01. Another was a
substantial experiment-by-occlusion-size interaction,
F(1.18, 298.03) = 49.26, p < .001, η2 = 0.16. There
was no significant three-way interaction (p = .550).
Fig. 5 shows the data pattern.
Consistent with the results of the experiments analyzed
separately, there was a main effect of sound condition,
F(1.87, 474.28) = 34.58, p < .001, η2 = 0.12. Consistent with
the results of Experiment 1 only, there was a main effect of
occlusion size, F(1.18, 298.03) = 53.81, p < .001, η2 = 0.18.
The present experiment provides a more precise look at the na-
ture of the relationship between visible time and the invisibility
period in inferred motion tasks. While Experiment 1 showed
significant and substantial main effects of the rate of auditory
patterns and of occlusion size on the error of judged TTC, the
duration of the visible time interval was kept constant at 1,000 ms
for all trials, while total travel time increased. As a result, the
duration of invisibility was always greater relative to the amount
of time that the moving disc was visible, leading to increasingly
more error in judged TTC judgments across conditions. In
Experiment 2, travel time and distance were kept constant, so
that the ratio varied as a product of the size of the occluded area.
While the consistent effects of faster rhythms in producing earlier
estimations of TTC were still observed, the effect of occlusion
size on the mean error in judged TTC was smaller in Experiment
2 than in Experiment 1. The results appear to reveal a protective
function of increased visibility on the manipulation of timing
estimation that results from exposure to fast auditory rhythms.
The principal finding was a replication of the main effect of
rhythm found in Experiment 1.
2372
Distance travelled (d) is a product of a moving object’s
velocity (r) multiplied by the amount of time (t) that has
passed since its departure (d = rt). In the absence of movement
related cues, like velocity (r), participants in the present study
were likely forced to rely on the remaining source of evidence
regarding the disc’s rate of movement, the passage of time (t).
Auditory input provides the most reliable and influential
source of temporal information (Conway & Christiansen,
2005; Fendrich & Corballis, 2001; Grondin, 1993; Grondin
& McAuley, 2009; Hove et al., 2010; Keetels & Vroomen,
2011; Spence & Driver, 2000; Vroomen & de Gelder, 2004;
Vroomen & Keetels, 2010; Welch et al., 1986). Without ac-
cess to visual information regarding the distance travelled, the
temporal cues present in the auditory rhythms would
have offered the next best source of information regard-
ing travel speed.
Because of the strong links between the auditory and motor
systems, the auditory system is more strongly suited for accu-
rate temporal perception than the visual system (Comstock
et al., 2018a, 2018b; Grahn, 2012; Grahn et al., 2011;
McAuley & Henry, 2010; Pasinski et al., 2016; Repp &
Penel, 2002). Unfortunately, this relationship also leaves the
motor system’s role in event timing vulnerable to exploitation
from input received in the auditory modality. Prolonged ex-
posure to visual stimuli seems to counteract these distorting
effects. So, while the auditory sense may be more directly
involved in the accurate temporal perception of sensory
events, the visual sense appears to aid in preserving the accu-
rate representation of these events in the face of contradictory
auditory information.
At all three levels of occlusion, the period of visibility was
greater in Experiment 2 relative to Experiment 1. Consistent
with previous findings, increased access to visual information
resulted in a reduced effect of occlusion size on the mean error
of participants’ judged TTC (Alderson & Whiting, 1974;
Peterken et al., 1991). Prolonged access to relevant sensory
information may provide the attentional system with more
durable and longer lasting impressions regarding the future
rate of an object’s movement, even if the object can no longer
be seen. An increase in the durability and resulting reliability
of these impressions would be reflected in the reduced effect
of occlusion size on the mean error of judged TTC, similar to
what was observed.
Contrary to our hypothesis, the results failed to reflect a
consistent change in the error of judged TTC along with in-
creases in the period of invisibility. Why would participants
show an increase in accuracy in predicting TTC in both the
silent and slow rhythm conditions, when visual information is
made more unavailable? This is an interesting and unexpected
finding that on the surface appears to contradict our claims
regarding the increasing vulnerability of attentional processes
to outside influence. Therefore, it deserves serious
examination.
Atten Percept Psychophys (2022) 84:2360–2383
Research suggests that prior experience with dynamic stim-
uli provides a foundation for later sensorimotor prediction
(Balkenius & Hulth, 1999; Bar, 2009; Friston, 2005; Huber
& Krist, 2004; Kording & Wolpert, 2006; Wilson &
Knoblich, 2005). Additionally, both event timing and rhyth-
mic attunement are found to improve with increased exposure
(Drake et al., 2000; McAuley et al., 2006; Thompson et al.,
2015). A possible explanation for the pattern of results re-
quires the consideration that participants became better able
to preserve the temporal qualities of dynamic mental imagery
due to repeated exposure to constant travel times. Therefore, it
is plausible that carryover effects were present in the current
design. If so, this most certainly would be a factor in minimiz-
ing error across all conditions. Nevertheless, the direction of
the differences observed were still mostly consistent with
those observed in Experiment 1. So, while carryover effects
may have been present, they likely impacted all conditions
equally.
At present it is unclear why the effect of occlusion size on
the error in judged TTC would be substantially smaller in
Experiment 2 than in Experiment 1 (thus bringing the
Experiment 2 errors closer to zero). Still, there was a clear
trend toward earlier responses when participants were ex-
posed to the faster rhythm. This supports the proposal that
the underlying mechanism by which temporal judgments are
believed to be made involve attentional rhythms that are pro-
duced in response to exposure to dynamic stimuli. These
rhythms provide a means of tracking time by aligning with
the course of events as they occur. The results of Experiment 2
support the prediction that exposure to external stimuli capa-
ble of altering the rate of these attentional rhythms will influ-
ence the rate at which the mental representations of dynamic
events unfold. These results also introduce the possibility that
the temporal qualities of internal representations may in some
ways be preserved as a result of prolonged exposure to con-
firmatory visual input.
It must be emphasized that the two experiments in the
present study were designed as separate, distinct experiments.
They differed from one another in multiple ways. Therefore,
any interpretations of the results of cross-experiment analyses
are tentative and would need to be confirmed (or
disconfirmed) by future research.
General discussion
Both experiments were designed to explore the effect of dif-
ferent speeds of an auditory rhythm on the rate of motion
extrapolation in an inferred motion task that measured TTC
judgments. As predicted, exposure to a faster rhythmic pattern
during the period of invisibility tended to produce more-
premature judgments of an invisible object’s TTC than did
exposure to a slower pattern. The degree of the overall effect
Atten Percept Psychophys (2022) 84:2360–2383
was at least partially moderated by the ratio of the visibility to
invisibility interval (Alderson & Whiting, 1974; Peterken
et al., 1991).
The present research attempted to elucidate the role of ex-
ternal sensory input on predictive representations of move-
ment under the assumption that the attentional system is both
dynamic and oscillating. In both experiments, participants
were required to make temporal judgments regarding TTC
when a moving disc was invisible. It is proposed that in the
absence of any visual input, these oscillating attentional
rhythms would lack reliable dynamic sensory information
with which to become aligned. Participants were instead ex-
posed to isochronous auditory rhythms of different rates dur-
ing this period of invisibility. These patterns would have pro-
vided supplementary temporal information that could be
attended to and incorporated with the dynamic representations
of the moving disc. Neurobehavioral evidence reveals an as-
sociation between the rate of sensorimotor oscillations and the
rate of sensory stimuli (Morillon et al., 2014; Morillon et al.,
2015; Portnova et al., 2011). Entrainment to these rhythms
during periods of invisibility would effectively provide a sub-
stitute sensory and consequent neural pattern from which to
make temporal judgments regarding the rate of movement
over time. Although this has not been directly explored here,
the results are consistent with this theoretical interpretation.
Our findings are consistent with those previously found by
Chotsrisuparat et al. (2017). Participants appear to underesti-
mate the TCC of a moving object when faster auditory
rhythms are played while the object is invisible. However,
ours is the first experiment to explore the relationship between
the rate of auditory rhythms and the ratio of visibility to
invisibility on judgments of TTC. An additional distinction
should be made between these earlier findings and the
current results. Chotsrisuparat et al. (2017) concluded that
the described effects occurred as a result of participants asso-
ciating faster tones with the more rapid movement of a trav-
elling object. This seems like a logical conclusion since faster
walking or running would produce the sound of more rapid
footsteps, and since the processing of temporal information
contained within auditory rhythms requires the contributions
of the motor system. However, the seemingly innate and rel-
atively automatic recruitment of motor regions in response to
auditory rhythms would suggest a more general, non-
associative link (Chen et al., 2008; Comstock et al., 2018a,
2018b; Grahn, 2009; Grahn et al., 2011; Grahn & Brett, 2007;
Kung et al., 2013; Phillips-Silver & Trainor, 2005, 2007;
Zatorre et al., 2007).
We offer instead that exposure to auditory rhythms leads to
the recruitment of the motor system, and this recruitment is
what promotes the entrainment of sensorimotor attentional
processes with the rate of the perceived rhythm (Arnal,
2012; Arnal & Giraud, 2012; Bolger, et al., Bolger, Coull, &
Schon, 2013, b; Henao et al., 2020; Henry & Obleser, 2012;
2373
Henry et al., 2014; Jones et al., 2002; Morillon et al., 2014;
Morillon et al., 2015; Nozaradan et al., 2012; Nozaradan et al.,
2017; Nozaradan et al., 2015; Schroeder & Lakatos, 2009;
Schroeder et al., 2010; Schubotz, 2007; Will & Berg, 2007).
Prior research has determined that exposure to faster rhythms
produces higher rates of SSEPs, which are correlated with
differences in the subjective experience of a temporal interval
(Eagleman & Pariyadath, 2009; Horr et al., 2016; Portnova
et al., 2011).
Shepard and Metzler (1971) originally demonstrated a tem-
poral connection between the manipulation of both real and
imagined objects. Longer periods of mental imagery are found
to be correlated with the imaginary walking of longer dis-
tances (Decety et al., 1989). Mental imagery experiments also
provide support for the existence of an overlapping system
used for motor imagery and execution (Decety, 1996; Hari
et al., 1998; Munzert et al., 2009; Naito et al., 2002;
Schnitzler et al., 1997). Neurophysiological research has re-
vealed similar levels of sensory and motor activation during
mental imagery of dynamic events (Press et al., 2011; Stadler
et al., 2012; Tian & Poeppel, 2010). A sensorimotor system
that processes observed and imagined events using very sim-
ilar neurocognitive mechanisms likely applies similar dynam-
ic expectations to both types of experiences (Decety &
Jeannerod, 1996). While the movement utilized in the current
experiments was in no way naturalistic, evidence suggests a
high degree of flexibility in terms of the motor system’s ability
to formulate accurate representations of motion. Regardless of
the degree of reproducibility of a movement, the motor system
seems to impose its own expectations onto dynamic events
(Jeannerod, 2001; Kunde et al., 2004; Schubotz, 2007;
Wilson & Knoblich, 2005; Wolpert & Flanagan, 2001).
Therefore, the effects observed in the present experiments
would most certainly be involved in the perception of more
naturalistic movement.
Early behavioral evidence for the predictive capacity of our
representational systems was identified by Freyd and Finke
(1984, 1985) who examined participants’ memories of a mov-
ing target’s final position prior to its disappearance. When
given three possible target orientations from which to choose,
participants incorrectly recalled the target in a position of for-
ward displacement, relative to its true end position. They re-
ferred to this phenomenon as representational momentum.
Representational momentum describes the forward displace-
ment of a mental representation of a dynamic sensory experi-
ence. This displacement is consistent with real-world expec-
tations in that it can be influenced by the speed of movement,
whereby greater displacement is reported for visual objects
displaying faster rotational speed (Freyd & Finke, 1985; for
a review, see Hubbard, 1995). Additionally, movement-
related factors such as final position, stimulus velocity and
motion direction also produce variations in the degree of for-
ward displacement (Hecht & Bertamini, 2000; Hubbard &
2374
Bharucham, 1988; Jordan et al., 2002; Schmiedchen et al.,
2013).
Dynamic stimuli require dynamic expectations. Dynamic
movements as well as rhythmic auditory stimuli can both lead
to forward displacements in memory (Feinkohl et al., 2014;
Schmiedchen et al., 2013; Wolpert et al., 1995). These expec-
tations must incorporate the passage of time into the formation
of accurate predictive representations. It is worth considering
that this forward displacement may be a product of ongoing
attentional rhythms that serve to preserve these representa-
tions into the future, beyond what is witnessed in the external
environment. Importantly, the extent of forward displacement
can be reduced by delaying the time at which the moving
object vanishes relative to our expectations (Jordan et al.,
2002).
Sound condition was the principal independent variable in
the present studies, and its effect was similar in both experi-
ments. Occlusion size, on the other hand, played a secondary
role, providing an opportunity for sound condition to have its
effects. The effect of occlusion size was substantially smaller
in Experiment 2 than in Experiment 1, thus raising the ques-
tion of why this occurred. A complicating factor is that the two
procedures were intentionally different (the Experiment 2 pro-
cedure was designed to address potential temporal confounds
in Experiment 1). We therefore believe further research would
be required to address this across-experiment effect. Because
the effect of occlusion size was inconsistent across experi-
ments, we focus our theoretical interpretation on the effects
of sound condition.
We introduce the possibility that internal rhythmic states
are intimately involved in the construction of dynamic mental
representations as well as the resulting expectations regarding
their outcome. More specifically, the accurate representation
of movement may rely to a large degree on the corresponding
rate of endogenous attentional rhythms relative to the rate of
visible movement.
While there does appear to be strong evidence for the spe-
cialization of processing between the auditory and visual
sense, there are times when we must depend on auditory input
for spatial information. When visual input is severely degrad-
ed or absent, there is an increased reliance on auditory input
during the process of spatial localization (Alais & Burr, 2004).
The capture of the timing of a visual event by an auditory
stimulus is generally referred to as temporal ventriloquism.
This effect is proposed to be a product of intersensory integra-
tion, whereby auditory stimuli are able to influence the per-
ceived timing of co-occurring visual events or intervals.
Research has revealed that this effect is seemingly indepen-
dent of the spatial distance between the visual and auditory
stimuli but is highly dependent on the temporal distance be-
tween the two sensory events (for a review, see Chen &
Vroomen, 2013). While most, if not all, previous research
has demonstrated this effect through the close temporal
Atten Percept Psychophys (2022) 84:2360–2383
pairing of perceivable auditory and visual events, the present
research reveals that this effect may also result from co-
occurring visual imagery as well.
The results from the present experiments reveal that if a
context rhythm is provided in the absence of conflicting visual
information, it can be incorporated into the timing of the rep-
resented event(s). The speed of the imagined event is pro-
posed to be a product of the rate of the sensorimotor attention-
al rhythm which can be influenced by the rate of context
rhythm. In the current experiments, 60-Hz pitch tones were
utilized in both the slow and fast sound conditions. Lower
pitched tones (≤ 130 Hz) have been found to be more effective
in producing stronger sensorimotor synchronization (Hove
et al., 2014; Lenc et al., 2018; Varlet et al., 2018). It is possible
that the particular tones used, may have enhanced the tempo-
ral effects of the sound conditions in both experiments.
Nevertheless, it is evident that the incorporation of the sensory
information provided by the different sound conditions pro-
duced unique temporal experiences of the same sensory
events. It seems apparent that exposure to different rates of
rhythmic auditory patterns can influence the temporal quali-
ties of dynamic mental representations.
The current research introduces the proposal that the inter-
nal attentional states required for the active representation of
movement are vulnerable to external manipulation. Exposure
to auditory rhythms that exceed the rate of external events may
produce dynamic representations that exceed the actual move-
ment speed of the object, leading to premature estimations of
TTC. Moreover, the potential for the manipulative effects of
auditory rhythms is greatly reduced when visual input is made
more available.
Evidence from previous studies point to a specific network
of brain regions that is likely involved in the processing of the
stimuli used as well as the completion of the task involved in
the present experiments. These regions include: (i) the visual
area V5 or the middle temporal area (MT), commonly associ-
ated with visual motion processing (Soto-Faraco et al., 2004),
as well as (ii) the medial superior temporal area (MST), which
is known to remain active during visual pursuit of objects that
are temporarily out of sight, pointing to a possible role in dy-
namic visual imagery (Berthoz, 2000). Evidence suggests that
an area located within the Sylvian fissure, (iii) where the pari-
etal and temporal lobes intersect (Area Spt), is associated with
auditory-motor integration (Chen et al., 2006; Hickok et al.,
2003; Warren et al., 2005). This region shares direct connec-
tions with (iv) the premotor cortex (Deacon, 1992; Petrides &
Pandya, 2002). The premotor cortex experiences increased ac-
tivation during exposure to rhythmic auditory stimuli, similar to
Area Spt (Bengtsson et al., 2009; Chen et al., 2006; Chen et al.,
2008; Karabanov et al., 2009; Kornysheva et al., 2010;
Schubotz et al., 2003; Schubotz & von Cramon, 2001, 2002,
2003; Wolfensteller et al., 2007). Additionally, a portion of the
premotor cortex known as (v) the supplementary motor area
Atten Percept Psychophys (2022) 84:2360–2383
(SMA), which has been strongly linked with motor timing
(Coull et al., 2016; Merchant & Yarrow, 2016; Sakai et al.,
2004; Schwartze et al., 2012; Tanji, 2001), and (vi) the basal
ganglia, which share reciprocal connections with the SMA
(Hove et al., 2013). The basal ganglia are also found to be
active during tasks requiring temporal perception and are di-
rectly involved in sensorimotor synchronization (Grahn &
Brett, 2007; Grahn & Rowe, 2009; Hove et al., 2013; Van
Wassenhove et al., 2011). Beta oscillations are believed to be
the mechanism by which the SMA coordinates its activity with
these subcortical structures (Wiener & Kanai, 2016).
These oscillations are likely to entrain with the rate of the
auditory rhythm presented during the present experiment,
thereby providing temporal information that could be applied
to the rate of visual imagery maintained during the periods of
occlusion. This entrainment would provide the motor system
with inaccurate temporal information, leading to a consistent-
ly skewed perception of the passage of time. Consistent with
the current findings, greater visual exposure would essentially
minimize the extent to which the motor system would need to
rely on auditory cues for this temporal estimation. Therefore,
smaller degrees of error in judged TTC would be expected
when the rate of movement is visually perceivable for longer
periods of time.
This theoretical framework must also consider the possibility
that auditory rhythms may have the potential to be beneficial
during times in which visual information is unavailable. If partic-
ipants are exposed to an auditory rhythm that closely aligns with
the rate of a dynamic event, this should aid in preserving the
temporal qualities of dynamic representations when movement
is inferred rather than directly observed. This implies that if we
know an object’s end point and rate of movement, our accuracy in
predicting when it will reach this point may benefit from exposure
to supplemental stimuli while the object is unseen. This is an
intriguing possibility that is worth exploring. Regardless of the
potential implications of the current research, it is evident from
our results that motion extrapolation is indeed a temporally ori-
ented process that is increasingly vulnerable to external manipu-
lation when visual information is made more unavailable.
Conclusion
Researchers have proposed several different cognitive factors
to explain how motion extrapolation (the ability to internally
represent an invisible object’s motion) is possible (Battaglini
et al., 2013; DeLucia et al., 2016; DeLucia & Liddel, 1998;
Huber & Krist, 2004; Lyon & Waag, 1995; Makin &
Poliakoff, 2011; Schiff & Oldak, 1990; Tresilian, 1995). The
results of the present research suggest the possibility that the
internal representation of dynamic motion is based upon the
preservation of corresponding attentional states produced from
the initial sensory experience. Accurately representing temporal
2375
information in dynamic representations is likely dependent on
preserving or reproducing these internal states constructed dur-
ing the actual sensory experience (Schaefer et al., 2011). Neural
patterns consistent with sensory experiences can in fact be pro-
duced from imagined events (Decety & Jeannerod, 1996;
Jeannerod, 2001; Okawa et al., 2017; Wolpert & Flanagan,
2001). Maintaining these patterns would provide continuous,
albeit potentially unreliable information about the perceived
relationship between movement and the passage of time. In
the absence of verifying sensory input, the gradual deterioration
of the internal representation of a dynamic sensory event is a
potential source for error in temporal and spatial judgments.
Our results indicate that when the process of tracking the
passage of time is manipulated by exposure to external stim-
uli, the accuracy of such tracking is affected in predictable
ways. Specifically, the findings reveal that as the apparent
duration of the period of invisibility increases, so does the
apparent speed of the moving object. This suggests that the
mechanisms involved in mentally tracking both movement
and the passage of time are not acting independently of one
another in this type of task. How these processes are linked in
the brain becomes a question of primary importance. It is
proposed that information related to time is communicated
in the form of oscillatory attentional activity that is then syn-
chronized with the rate of external events. The merging of
these two sources of information may provide a basis for the
temporal perception of dynamic mental imagery. In the ab-
sence of confirmatory sensory (visual) information, appropri-
ate adjustments are unable to be made, leading to the eventual
breakdown of this representation and consequently, greater
error in temporal predictions observed over time. In the pres-
ence of temporally consistent sensory information, our atten-
tional system would effectively incorporate this input into
these representations, thereby preserving their accuracy.
In the current experiments, the presence of isochro-
nous auditory pulses of different rates led to predictable
shifts in the perceived timing of dynamic events. Our
results have demonstrated that exposure to faster rhyth-
mic patterns influences participants’ estimations of the
velocity of invisible objects. They have also revealed
that this influence is drastically reduced when visual
input is made more available, likely preserving a more
accurate mental representation during periods of invisi-
bility. Taken together, this reveals a possible mechanism
by which the temporal qualities of mental imagery may
be understood. While these experiments did not examine
the neurophysiology associated with motion extrapola-
tion, previous research as well as the current results
point to a possible role of endogenous rhythms in
representing the temporal qualities of dynamic imagery.
Merging both the representation of the moving object as
well as the representation of passing time could be the
mechanism by which motion extrapolation occurs. Future
2376
neurophysiological research could certainly help to identify the
role of neural rhythms in both constructing and maintaining ac-
curate mental representations of dynamic events.
Open Practices Statement
Data and materials are available, and are located in a secure
laboratory in the Psychology building on the Bowling Green
State University campus. None of the experiments were
preregistered.
Author Note Timothy B. Patrick is now at the Department of
Psychological and Brain Sciences, Indiana University
Bloomington. The authors have no known conflicts of interest
to disclose.
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