Chapter 11

Time Perception and Discrimination

in Individuals Suffering from Hallucinations
Flavie Waters

Abstract Hallucinations are perceptual experiences arising in the absence of external

stimulation, and that are perceived to be real. Auditory hallucinations have been intensively studied in the last 30 years or so, and different explanations have been proposed.
This chapter reviews evidence of timing abnormalities in people with schizophrenia
and auditory hallucinations. The first section of this chapter shows that abnormalities
in time perception and time discrimination are common in schizophrenia, with
difficulties occurring across all timing periods, stimulus modalities, and at all stages of
information processing. The second section synthesises different theoretical models of
auditory hallucinations, and demonstrates that abnormal timing mechanisms might
contribute in a significant way to cognitive difficulties and perceptual distortions that
underlie hallucinatory experiences. Timing dysfunctions arise from disruptions to the
integrity of neural circuits and neurobiological mechanisms in schizophrenia. A plausible explanation suggests that these might obstruct the normal coordination of internal
systems, the integration of bottomup and topdown processes, and self-monitoring
mechanisms linked to hallucinations. Timing problems might also induce distortions in
action causation, motivational significance, and higher-order cognitive functions.
Altogether, timing abnormalities provide a parsimonious explanation for neural, cognitive, and phenomenological findings in auditory hallucinations.

Abbreviations
DA
ERP

Dopamine
Event-related potential

F. Waters (*)
Centre for Clinical Research in Neuropsychiatry, Graylands Hospital, and The University
of Western Australia, Private Mail Bag No 1, Claremont, Perth, WA 6910, Australia
e-mail: flavie.waters@health.wa.gov.au
185
R. Jardri et al. (eds.), The Neuroscience of Hallucinations,
DOI 10.1007/978-1-4614-4121-2_11, Springer Science+Business Media New York 2013

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F. Waters

fMRI
PFC
SZ

Functional magnetic resonance imaging

Prefrontal cortex
Schizophrenia

11.1

Introduction

Time-keeping refers to an internal experience of the flow of time. The ability to

estimate time allows organisms to initiate behaviours at chosen moments, and
respond to situational demands. Time-keeping processes are believed to require the
use of an internal clock that codes temporal information. One role of this internal
clock is to orchestrate all aspects of human behaviour, from purely sensory information to higher- order cognitive processes (such as speech and thoughts), to
ensure fluid information processing. The precise synchronisation of motor, sensory
and cognitive information is also critical for generating self-awareness, given that
the modulation of voluntary efforts and incoming perceptual information enables
the distinction of actions that are self-generated from those that have an external
origin.
Timing processes are therefore of great interest for our understanding of disorders such as schizophrenia (SZ), since individuals with the disorder suffer from
fundamental difficulties differentiating between the self and others, and from distorted perceptions in which thoughts and actions are perceived to arise from external agents (see also Chap. 10, this volume). Of specific interest are auditory
hallucinations, where internal auditory events fail to be recognised and are misattributed to an external source. A well established finding is of timing abnormalities
in SZ irrespective of symptom presentation, but fewer studies have examined
whether this has any relevance to auditory hallucinations. Auditory hallucinations
occur frequently in many clinical disorders and in the general population, so unravelling the processes that underlie these symptoms has relevance beyond the immediate theoretical implications for SZ by increasing our understanding of a common
human experience.
The aim of this chapter is to evaluate the mechanistic processes by which
timing abnormalities might contribute to auditory hallucinations. The first section reviews the evidence linking timing deficits and SZ. This step is necessary
given that brain abnormalities in SZ provide the context in which hallucinations
occur. The term time representation will be used to refer to a broad range of
events that includes time perception (involving an estimation of the timing, or
duration, of events), and time discrimination (entailing a comparison of the
duration between different events). The second section of this chapter addresses
the proposal that timing abnormalities are a core abnormality in auditory hallucinations, which can explain distortions in mental and perceptual awareness,
and external attributions. The findings show that timing dysfunctions contribute
in a significant way to the processes underlying auditory hallucinations, and that

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a proposal of auditory hallucinations based on abnormalities in internal timing

provides a parsimonious explanation for many of the neural, cognitive, and phenomenological characteristics associated with these phenomena.

11.2

Timing Processes in Schizophrenia

Numerous studies have reported abnormal time representation in SZ. Altogether,

findings are generally consistent in showing broad difficulties across all timing periods, stimulus modalities, and task design. The following methods have been used to
assess timing mechanisms in SZ.
In time-estimation methods, participants are required to estimate the duration of
an event, or to estimate when a fixed duration has elapsed. In a prospective task
design, instructions are provided at the beginning of the task to alert the investigator every time a fixed duration has elapsed (ranging from 30 s to 5 min) (production task). Alternatively, a shorter duration is used (1 to 5 s), and participants are
required to pace (or reproduce) this time measure repeatedly by tapping on a button or key (reproduction tasks). In a retrospective task design, participants are
given instructions at the end of a task to think back and estimate how long has
passed, often using a verbal estimate of time (in minutes or seconds). Studies
using time-estimation methods have demonstrated that people with SZ are consistently less precise at estimating the duration of events than healthy comparison
controls, regardless of whether the task comprises a short or long duration, or
whether the passing interval includes a distracting activity or not (Densen 1977;
Johnson and Petzel 1971; Penney et al. 2005; Shum et al. 2004; Tracy et al. 1998;
Tysk 1990; Wahl and Sieg 1980).
Comparison-type tasks assess the sense of time by asking participants to compare
different time intervals. First, a fixed-duration auditory or visual (standard) stimulus is
presented. This is followed by a comparison stimulus which is of a longer or shorter
duration than the standard stimulus. Participants are then required to compare the durations of the standard and comparison stimuli. One version of the task requires participants to judge whether the comparison stimulus has same/different length than the
standard stimulus (temporal generalisation task). In another version, the comparison
stimulus needs to be classified as short or long, based on its perceived similarity
with the duration of the standard stimulus (temporal bisection task). Comparison-type
tasks traditionally require a pre-test training (or learning) phase, and this information
needs to be held in memory for subsequent comparison with the comparison stimulus. Using these tasks, studies show that people with SZ have greater difficulties discriminating time intervals compared to healthy controls. Deficits have been demonstrated
on tasks using time intervals ranging from milliseconds to several seconds or longer
(Carroll et al. 2008, 2009; Davalos et al. 2002, 2003; Elvevg et al. 2003), and in both
the auditory and visual modalities (Davalos et al. 2002).

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Early studies had reported conflicting results regarding the direction of error,
showing that individuals with SZ either overestimate time (Densen 1977; Johnson
and Petzel 1971; Wahl and Sieg 1980) or underestimate time (Johnson and Petzel
1971; Tysk 1990; Wahl and Sieg 1980). However, more recent studies involving
systematic comparisons of responding are largely consistent in showing that time
representation in SZ is better described as highly variable, without systematic
biases in either direction (Carroll et al. 2008, 2009; Elvevg et al. 2003, 2004; Lee
et al. 2009; Tracy et al. 1998). This is aligned with findings in general community
individuals, which show that time perception is subjective and highly variable, especially as the timing interval increases.
Altogether, these findings suggest that people with SZ show distortions and greater
timing imprecision when compared to healthy non-clinical control individuals.
Timing processes across long durations have typically been assessed using memory tasks where the temporal memory for events that occurred in the past is tested.
Often, memory of when actions were performed, or when items were presented, is
assessed. Tasks may require participants to make a recency judgment, indicate the
temporal position of a memory item, or classify items as a function of the test session in which they were presented. Studies of temporal memory show that people
with SZ perform less accurately than control participants (Dreher et al. 2001;
Elvevg et al. 2004; Rizzo et al. 1996; Schwartz et al. 1991; Waters et al. 2004),
even when the groups are matched on general memory abilities (Waters et al.
2004), providing evidence for the notion that long-term associative (or context)
memory involving a sense of time is impaired in this group.
A frequent criticism of the above tasks is the involvement of cognition other than
purely timing mechanisms. Difficulties in objectively measuring the internal clock
have led to a reliance on cognitive measures that involve a temporal component.
Thus, performance on such measures typically involves attention, working memory,
and memory processes (Lee et al. 2009; Ortuo et al. 2005; Shum et al. 2004; Tracy
et al. 1998), which are used to attend to the task, keep task instructions in mind, and
make response decisions. The concern is that performance deficits might be linked
to broader cognitive dysfunctions in SZ, rather than the internal clock. Thus, deficits
on timing tasks may be linked to difficulties performing the task rather on its timing
component. Many attempts have been made to control for cognitive involvement in
time-representation tasks (Elvevg et al. 2004; Waters et al. 2004), and findings
appear to suggest that timing processes are impaired irrespective of the level of
cognitive abilities. Nonetheless, other approaches have been used to circumvent
cognitive involvement when assessing time representation in SZ.
Time representation across short durations and/or with the use of preconscious measures. One method to avoid intervention from higher-order cognitive processes is to
use the fast and repetitive reproduction of simple movements (like finger tapping).
Some tasks require participants to tap in time to computer-generated tones, and then
to continue tapping at the same pace after the tones are discontinued. Studies show
that people with SZ demonstrate much variability on this type of task, and fail to keep
an even pace when compared to healthy control subjects (Carroll et al. 2009).

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Another method is to use preconscious measures of mental activity. Timing

studies often use physiological recordings to assess early stages of information
processing. These show that people with SZ are impaired on measures such as the
eyeblink conditioning method, thought to measure neural timing circuitry (Brown
et al. 2005), and a modified version of the mismatch negativity (MMN) task, in
which the neural detection of deviation in sound duration of tone intervals is measured using event-related potential (ERP) methodologies (Davalos et al. 2005;
Todd 2006). Electrophysiological studies have also demonstrated abnormal synchrony of neural timing processes, as demonstrated with EEG and magnetoencephalography. It has been proposed that the synchronisation of oscillatory
neural responses during task performance suggests a role for neural synchrony in
cortical networks and performance. Evidence has been presented that cognitive
performance in SZ was related to impaired neural synchrony of oscillatory activity
in the high frequency band (b and g) (Phillips and Silverstein 2002; Uhlhaas and
Singer 2006; Uhlhaas and Silverstein 2005). This was interpreted as showing that
interruptions to the fluid and coordinated timing of neural activity may explain
many of the cognitive dysfunctions in SZ, including deficits in attention, working
memory, and perceptual organisation.

11.2.1

Neuroimaging Findings

The above deficits demonstrated on behavioural tasks are supported by neuroimaging studies, which show that the brain systems normally supporting performance on
tasks of temporal representations (prefrontal and parietal cortices, thalamus, basal
ganglia, and cerebellum; Alexander et al. 2005; Harrington et al. 1998; Ivry and
Spencer 2004; Picton et al. 2006), are abnormally activated in SZ.
For instance, Volz et al. (2001) conducted a functional magnetic resonance
imaging (fMRI) study while volunteers underwent two tasks that required the discrimination of time and pitch (sound). Results showed that individuals with SZ
showed hypoactivity in the posterior putamen, anterior thalamus, and right medial
prefrontal cortex (PFC) that were related to timing performance. Ortuo et al.
(2005) also reported abnormal modulation of a corticalsubcortical network involving the supplementary motor area (SMA) in people with SZ during a timeestimation task. Finally, Ojeda et al. (2002) conducted a PET-015 water activation
study measuring relative cerebral blood flow during two counting tasks (counting
clicks, and then counting forward at the same rate after the clicks had stopped).
Results showed a differential pattern of performance depending on the task.
Counting clicks engaged more widespread brain regions in SZ than healthy controls, particularly in the inferior frontal regions. Counting forward unassisted,
however, yielded decreased activation in the frontal regions and inferior parietal
gyrus, suggesting that individuals with SZ failed to activate these regions at a similar level to controls during this task. Interestingly, behavioural performance was
not different from that of controls, suggesting that people with SZ may be using
different strategies in order to perform the task.

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11.2.2

F. Waters

Summary of Findings on Time Representation

Tasks in SZ

Altogether, the above studies show that timing deficits are wide-ranging in SZ.
Deficits have been demonstrated using time-estimation methods and comparisontype tasks, and on tasks assessing short and long timing durations. Deficits were
also demonstrated in the auditory, visual and motor modalities, and at all stages of
information processing, ranging from preconscious to conscious processing. Using
Lewis and Mialls (2003) terminology, this points to impairments on both the automatic aspects of temporal processing (referring to repetitive or unconscious timing
tasks), and the resource-dependent controlled processes, referring to cognitively
mediated neural systems associated with attention and memory.
Altogether, evidence points to extensive timing abnormalities in SZ. But how
does this link to auditory hallucinations?

11.3

Timing Processes in People with Auditory Hallucinations

Few studies have directly investigated timing processes in individuals with auditory
hallucinations. However, timing is central to many neurobiological, cognitive, and
psychological explanations of these experiences.
Neurobiological models posit that dopamine (DA) abnormalities give rise to positive psychotic symptoms such as auditory hallucinations. It is posited that such
symptoms arise by increasing DA release in the mesolimbic pathway, as shown
from evidence that the administration of DA pharmacological agents in healthy
people produce symptoms of psychosis, and from findings of abnormal DA transmission in people with psychosis. This DA theory is relevant to timing processes,
given that DA manipulations can modulate timing perception by altering the speed
of the internal clock (Cheng et al. 2006; Macdonald and Meck 2005; Meck 1996).
For example, increasing brain DA levels with the administration of a DA agonist is
linked to an increase in clock speed, whereby DA antagonists decrease clock speed
(Meck 2005). Given that hallucinations are associated with DA abnormalities, and
that DA is linked to timing perception, one plausible suggestion is that hallucinations are mediated by timing deficits.
Cognitive and psychological explanations provide a different level of description by
detailing the mechanistic processes and functional significance of brain abnormalities for hallucinations. The importance of such models is that they are more closely
tied to conscious experiences than neural models. They can therefore provide explanations for complex subjective symptoms, and generate predictions that are testable
using scientific methods of enquiry. Theoretical models have identified several characteristic features associated with auditory hallucinations, including the failure to
self-recognition (perhaps linked to abnormal connectivity, self-monitoring deficits,

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or a failure of integration of bottomup and topdown processes), and misattribution

of self-generated events to another agent (possibly linked to abnormal sense of causation,
salience or reasoning processes) (see also Chaps. 9 and 10, this volume). The following
describes how disturbances in timing processes might be linked to these features.

11.3.1

Abnormal Connectivity

A disconnection, or failure of integration, at the level of brain systems has been a

popular model of SZ and hallucinations for over a hundred years (Bleuler 1911;
Andreasen 1999; Friston and Frith 1995; Meehl 1989). It essentially suggests that
abnormal integration between the frontal and posterior brain regions in SZ leads to
a disruption in the temporal organisation (timing) of information sequences that
underlie normal behaviour and perceptions, giving rise to positive symptoms such
as hallucinations, thought disorder, and disorganisation. Evidence supports this proposal of abnormal temporal coordination of distributed neural processes. For
instance, neuroimaging and electrophysiological studies of people with SZ and hallucinations show abnormal connectivity both within, and between, brain regions
(Spencer 2008; Spencer et al. 2008; Hubl et al. 2004; Uhlhaas and Singer 2006;
Jardri et al. 2011).
Consequently, abnormal timing might underpin a failure of integration between
brain functions. Reduced connections between brain structures can explain other
cognitive abnormalities linked to auditory hallucinations. Within the cognitive
approach of psychopathology, this lack of integration between neural systems points
to a lack of synchrony between important cognitive functions such as perceptual
analysis, memory, and executive functioning. Excessive perceptual output, combined with a lack of monitoring and supervisory activity by the frontal lobes or
memory processes, might contribute to some of the key phenomenological features
of auditory hallucinations including recurrent perceptual intrusions, perception of
non-self origin, and abnormal reasoning processes linked to a sense of external
causation.

11.3.2

Forward Model (Self-Monitoring)

One motor-cognitive theory about auditory hallucinations suggests that a dysfunction of the forward-model system leads to difficulties in predicting the sensory consequences of ones intended actions and inner speech, and consequently a failure of
monitoring the origins of behaviours (self-monitoring) (Frith 1987; Frith 2005).
According to this model, incorrect sensorimotor predictions result in a failure to
recognise internally generated mental experiences so that they are perceived to have
a non-self origin. The importance of accurate timing for the forward-model system
and self-monitoring has been demonstrated in studies using self-produced tactile

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stimulation (tickling) (see review Blakemore and Frith 2003). In normal

circumstances, sensory predictions made by the forward model diminish the actual
sensory experience during self-stimulation, explaining the near impossibility of
tickling oneself. Blakemore and colleagues (1998) showed that the introduction of
longer delays between the tickling gesture and resultant body sensation resulted in
fundamental changes in subjective perceptual experience. Participants reported progressively decreased tickle sensations as the delay increased as the stimulation no
longer corresponded to the motor prediction. These results were interpreted as demonstrating that timing precision contributes to sensorimotor awareness by way of
forward-model processes. The application of this model to hallucinations was tested
by the same authors in a separate study. Blakemore et al. (2000) showed that people
with auditory hallucinations and other passivity symptoms lacked the normal
decrease in perceptual intensity for self-produced tactile stimuli, and were less able
to distinguish between self-produced and externally produced tickles. In other
words, they rated the self-tactile stimulations as more intense and tickly than
healthy people. Since timing processes contribute to tactile sensations, one possibility is that timing impairments in forward-model mechanisms contributed to increased
perceptual awareness in people with hallucinations.

11.3.3

Binding Effects Across Actions and Cause

(Perception of Action Causation)

An emerging literature suggests that the temporal relationship between events

(and particularly voluntary actions and their consequences in the external world)
critically affects our awareness of self, and to whom our actions are attributed.
Critically, this literature suggests that temporal precision is at the heart of this selfawareness. Studies show that, once an action has been performed, temporal contiguity between this action and its effect produces a distortion of the sense of time,
whereby the temporal interval between these events is perceived to be shorter than
it actually was (Haggard and Eimer 1999; Haggard et al. 2002; Haggard 2005).
In other words, the perceived timing of voluntary actions and their effects are shifted
across time towards each other. This processing does not reach awareness, as the
duration occurs in the range of milliseconds, but it is sufficient to induce a sense of
causation between voluntary actions and their consequences, so that the events are
bound together in the mind.
Proponents of this theory suggest that such processes are used to identify the
source of actions as coming from the self or others. One prediction, therefore, is that
an abnormality in time processing can lead to a misunderstanding regarding the
order of, and relationship between, events, and an incorrect attribution to external
agents. This is consistent with Spences proposal (1996), for instance, that a dysfunction in the timing of sensorimotor processes might result in the awareness of the
actual movement preceding awareness of the intention to act in SZ, which is contrary to the normal experience associated with self-generated actions.

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In support of such binding-across-time abnormalities in SZ, Franck and

colleagues (2005) showed that people with the disorder demonstrated unusually
strong binding effects (hyperbinding), so that temporal intervals between actions
and their sensory consequences appeared even shorter in this population than in
healthy controls. In addition, such hyperbinding effects were demonstrated for
voluntary actions, passive movements, and unrelated events. This general tendency
to perceive temporal intervals between events as being shortened was proposed to
disrupt processes involving the sense of causation between events, so that abnormalities might lead to deficits in action attribution. Using a temporal bisection task,
Waters and Jablensky (2009) demonstrated that time-estimation difficulties were
particularly more severe in individuals with auditory hallucinations and other passivity symptoms. The results showed that people with these symptoms experienced
time differently, and tended to underestimate the duration of time intervals. In addition, these deficits were independent of intentional activities. This is important,
because it suggests that motor intentions do not have privileged access to representations about the sense of self-causation, and therefore that the impact of timing
abnormalities on the sense of causation may occur across a range of events and may
be linked to psychotic symptoms such as hallucinations.
One functional explanation for the findings of timing abnormalities in individuals with auditory hallucinations is that such deficits may lead to a misunderstanding
regarding the causes of mental events such as inner speech or memory fragments, so
that they are not perceived as self-generated. Essentially, the subjective experience
would be akin to a perception without a clear understanding regarding its cause or
origin. In turn, this might lead to incorrect labelling and attribution regarding the
source of this event.

11.3.4

Hierarchical Systems Informed by Sensory Input

and TopDown Predictions (BottomUp
and TopDown Integration)

Other theoretical models of auditory hallucination propose that neural activity, and
therefore mental experiences, are shaped by both sensory input and topdown attentional mechanisms (Frith and Dolan 1997; Krishnan et al. 2011; see also Chap. 6,
this volume). In this model, self-awareness is articulated as the product of sensory
input that is modulated by prior knowledge and attention. Topdown processes
modify sensory information in a way that is dependent upon ones expectancy and
prior knowledge.
This notion arises from the proposal that internal events are based upon hierarchically organised cortical systems, so that each level is influenced both by the input
provided by the level below, and expectations generated by the level above. This
system allows a distinction between ones own actions and intentions, and those
of agents in the outside world. The model posits a failure of this mechanism in
auditory hallucinations, so that the relative weighting between these two forces is

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out-of-balance leading to excessive influences of topdown processes over sensory

input (Stephan et al. 2006). When the parameters and relative weight of bottomup
and topdown information are incorrectly estimated, topdown prediction errors
occur with regards to the nature and cause of experiences. Repeated errors lead to
enhanced learning of incorrect associations, which ultimately progresses to increased
prior aberrant expectations (Corlett et al. 2007). While deficits in timing are not
explicitly addressed in this model, it is easy to see how a timing disruption in the
fluid coordination of these events may lead to prediction errors and a general disruption at all levels of the hierarchical cortical system.

11.3.5

Aberrant Salience

A model that is somewhat linked to the previous explanation suggests a prominent

role of DA for mediating attention and reinforcements from the environment.
According to Kapur (2003), DA produces a heightened awareness towards external
objects and internal mental events, thereby producing an attribution of motivational
significance towards these stimuli. DA plays a key role towards the selection of
stimuli in the environment, and reinforcement of these new associations. One consequence of this abnormal salience is that it contributes to the formation of abnormal associations between unrelated events, so that a neutral stimulus can become
imbued with significance, placing excessive influence on thinking and behaviour.
The processes by which this occurs are less precisely described, although one may
speculate that aberrant salience is also linked to timing processes. Recent work ties
excessive modulation of motivational factors to temporal processes, in keeping with
DAs role in modulating the speed of the internal clock (Cools 2008; Galtress and
Kirkpatrick 2009). The exact role of timing in the attribution of salience is far from
clear, although one possibility is that DA may function to assign significance by
producing distortions in time perception through binding of associations between
unrelated events.

11.3.6

Cognitive Models of Context-Memory and Reasoning

(Self-Recognition, Misattribution)

Another class of theories proposes dysfunctions in memory and executive reasoning

processes that are involved in identifying the source of actions and mental events.
It is proposed that auditory hallucinations are associated with fundamental abnormalities in the higher- order processing of contextual information and cognitive
operations which are tied to internal mental events and memories (Bentall and Slade
1985; Brebion et al. 2000; Harvey 1985; Hemsley 1993; Waters et al.2006a, b).
Given that context of memories (the who and the when) provides cues that allow

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the differentiation of one memory from another, a loss of such qualitative information
would make it difficult to infer the origins of mental events. These dysfunctions are
posited to lead to difficulties with self-recognition and decision-making about the
origins of mental events (Waters et al.2006a, b).
Theoretically, timing processes are pivotal for the functioning of cognitive functions. Brain functions (including thoughts, perceptions, and other cognitive events)
rely upon the coordinated interaction of a large number of neurons that are distributed across specialist areas of the brain. This integration of responding across different cortical regions is needed for the organisation of sensory, perceptual, and cognitive
systems. Such integration may occur either in parallel, or in sequential order, but
essentially requires fluid and dynamic adjustments depending on the nature of the
task. Abnormal timing processes would therefore have an impact on the orchestration of such sequencing ordering, and cause problems for the systems that support
cognitive functions and perceptual awareness. Specifically, timing problems might
result in a failure to adjust neural activity to the nature and demands of the task.
So, what would the functional consequence of timing problems on memory and
prefrontal functions for auditory hallucinations? Several consequences might occur.
First, the ability to represent the past, present, and future, and to integrate information across time, is a key function of the PFC (Fuster 1999). Thus timing impairments would impact on the ability to fully represent temporal contextual information
(as proposed above), as well as to plan future events, sequence behaviours across
time, and predict later events (still to be empirically tested). Second, the binding of
context cues in memory is reliant upon the frontal lobes and its connections with the
hippocampus and medial temporal system. Hence, a failure of integration would
lead to faulty integration of memory and executive functions into a composite memory trace, and difficulties forming an intact representation regarding the origins of
mental events.
Unsuccessful or less effective cognitive executive functions might also be
expected to increase auditory hallucinations through the lack of, or use of
non-effective, strategies for controlling unwanted intrusive thoughts, and by faulty
reasoning processes.

11.4

Conclusion

In this review, the mechanistic processes by which timing problems make a contribution to auditory hallucinations have been reviewed. Evidence suggests that these
problems might lead to difficulties in coordinating and synchronising internal systems that are responsible for perceptual analysis and cognitive functioning. Timing
problems are also linked to difficulties with forward-model processes and actionbinding. Finally, an abnormal sense of time may cause aberrant significance towards
neutral events. Altogether, time-keeping processes are fundamentally involved in
key phenomenological features associated with auditory hallucinations, notably
self-monitoring and misattribution.

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Thus, a plausible key mechanism underlying auditory hallucinations, and

common denominator of all the theoretical models described above, involves abnormal timing processes. Specific predictions from this novel model however need to
be empirically tested. In particular, the specificity of timing deficits to auditory hallucinations needs to be tested. Are difficulties with time representation uniquely
associated with hallucinations, or do they also play a role in other positive psychotic
symptoms? Few studies have systematically investigated this notion, although it has
direct and important bearings for issues of specificity. Other questions involve the
contribution of medication, and the persistence of deficits across the course of the
illness. In addition, we must examine whether timing abnormalities become more
severe with increased severity of hallucinations.
In conclusion, time representation is a fascinating area of study, which has direct
relevance to all aspects of human behaviour. Its role in abnormal experiences such
as hallucinations and other psychotic symptoms is also of considerable interest, and
it provides a plausible explanation for a whole range of theoretical models at different levels of explanation.
Acknowledgment FW is supported by National Health and Medical Research Grant ID
634328.

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