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Short Term Sleep Deprivation, Language Comprehension

and Auditory Temporal Resolution
Harvey Babkoff1, Leah Fostick2, Gil Zukerman2 and Elisheva Ben-Artzi3

1Department of Psychology, Bar-Ilan University, Ramat-Gan, Israel and

Department of Psychology, Ashkelon Academic College, Ashkelon,
Israel
2Department of Communication Disorders, Ariel University Center,
Samaria, Israel
3Kinneret College, Tzemach, Israel
ABSTRACT
Over the past several years there has been a shift in the theoretical
construct or models used by sleep researchers to explain the effects of
sleep deprivation on performance. The focus has changed from an
emphasis on arousal mechanisms to that of prefrontal cortical activation
and its associated cognitive functions. According to the latter theoretical
construct, cognitive functions that are considered most vulnerable to short
term sleep loss are language skills in communication, divergent thinking,
creativity and cognitive flexibility. Although the theoretical framework
for the impact of sleep deprivation on performance may have changed
over time, nevertheless there has not been any increased interest in
investigating the effects of sleep deprivation on sensory or perceptual
functions. In the present chapter we develop the arguments for studying
the impact of sleep loss on perceptual systems that are strongly associated
with a cognitive function, such as language. Language comprehension
and production have been strongly associated with auditory temporal
resolution. Consequently we believe that performance on tasks based on
auditory temporal resolution should be of paramount interest to
researchers of the impact of short term sleep deprivation on cognitive
functions.
We present a reanalysis of auditory temporal resolution data
published earlier to show how sleep loss impacts the ability to
discriminate dichotic temporal order. The meaning of deficit in auditory
temporal resolution and its possible relation to impairment in performing
language-based tasks is not only theoretically interesting but may also be
of practical use, e.g., when testing the effects of pharmacological or other
putative treatments to ameliorate the effects of sleep loss on perceptual or
cognitive performance.
INTRODUCTION
Studies of sleep deprivation, to date, have generally concentrated
on the effects of sleep loss on psychological, cognitive and motor
performance (Babkoff, et al 1991a,and b; Babkoff, et al , 2005; Dinges &
Kribbs, 1991). Very few studies have examined the effects of sleep loss
on more basic sensory and perceptual functions (Babkoff et al., 2005).
There was apparently a very good reason for the lack of interest in
studying the effects of sleep loss on basic sensory and perceptual
functions. In most of the earlier studies of the effects of sleep loss, the
main interest was in designing performance tasks that were prima facie
concordant with the "real life" complaints of individuals required to
perform effectively under conditions of sleep deprivation. In general,
these tasks tested: 1) for changes in mood, consistent with complaints of
changes in emotional responses; 2) either had face validity with the tasks
usually performed in the workplace; or 3) tested some very specific
cognitive or psychomotor capability that has been shown to be a basic
and necessary component of successful task performance, e.g., working
memory, choice reaction time. Individuals suffering from insomnia or
other sleep disturbances usually complain of fatigue and difficulty in
performing their normal daily activities, rather than complaining of
altered thresholds or other basic perceptual changes. Thus, superficially at
least, there seems to be no meaningful negative impact of sleep loss on
any of the basic sensory or perceptual functions that was worth
investigating.
Arousal Theory of Sleep Deprivation
However, there is a second, perhaps more important, reason for the lack
of interest in the impact of sleep loss on sensory and perceptual functions,
i.e., the theoretical framework within which researchers viewed and
studied the impact of sleep deprivation on task performance. The
prevailing thought regarding the brain mechanisms affected by sleep
deprivation during the latter part of the twentieth century centered
primarily on the arousal system. Up until the 1990s, many researchers
designed their studies and interpreted their results in terms of the Arousal
theory of sleep deprivation. They argued that a reduction in sleep length
affects performance by causing decreased arousal and slower cognitive
processing (Babkoff, et al, 1988, 1991a; Harrison & Horne, 2000a;
Kjellberg, 1977; Wilkinson, 1965; Wilkinson, 1992; Williams, et el,
1959). According to this view, sleep deprivation serves to reduce the non-
specific arousal level, but has no specific effect on any given sensory,
perceptual, or cognitive system (Wilkinson, 1992). The most extreme
version of this hypothesis (Lapse hypothesis) proposed that the only real
deficit due to sleep loss was an increase in errors of omission or lapses in
performance (Williams et al., 1959). The Lapse hypothesis considered the
lapse or an error of omission (defined behaviorally as a period of not
responding that is longer than several standard deviations beyond the
expected or mean response time) as the major dependent variable in
studies of sleep loss and posited the presence of microsleeps (defined as
an episode of sleep lasting from 0.5 to 10 seconds) during long hours of
sleep deprivation as the major factor in performance decline during long
term sleep deprivation.
In its radical form, the Lapse hypothesis implies that sleep loss
itself does not impact negatively on sensory, perceptual or cognitive
functions. The only deficit, which is systematically found, lapses, is due
to the subjects really having fallen asleep, physiologically, for short
periods of time (i.e., subjects experience microsleeps). According to this
view, sleep loss affects all cortical regions in a fairly similar manner.
Wilkinson (1992) even argued that sleep deprivation reduces the non-
specific arousal level, but has no specific effect on any sensory,
perceptual or cognitive function differently from any other. Even in its
milder form, Arousal theory posited a fairly uniform general slowing of
responses in all neurobehavioral systems, rather than deficit in any
specific system.
Dinges and Kribbs (1991) noted that the majority of studies of
sleep-deprivation and performance reported global changes in
performance ability, changes that may have been mediated as much by
microsleeps and lapses in attention as by more subtle or profound
alterations in information processing. Moreover, in the early 1990's
Koslowsky and Babkoff (1992) reviewed the results of 27 studies (from
1969-1990) with a total of 429 subjects that met the criteria for a
quantitative analysis, and applied meta-analytic techniques to test the
lapse hypothesis. They reported that speed rather than accuracy of
performance, and work-paced rather than self-paced tasks were most
affected by sleep loss. The authors concluded that the findings were
consistent with the Lapse hypothesis. In summary, according to the
Arousal hypothesis, there were no intrinsic differences in performance
decrement among different tasks and certainly not on any given sensory
or perceptual system, since the major source of variance caused by sleep
deprivation was basically the same for performance on all tasks, i.e.,
reduced arousal. The nature of the task itself and the sensory or
perceptual systems involved were apparently of lesser importance.
Although the major theoretical framework for studying and
understanding the effects of sleep loss on task performance has shifted
radically over the past decade, nevertheless the prevailing view seems to
continue to view the study of the effects of sleep loss on sensory and
perceptual functions as theoretically unimportant and practically
inapplicable to the workplace. In fact the emphasis of the more recent
view in positing that sleep loss has specific as well as general effects on
performance, has been on the higher cognitive level functions, rather than
on the more basis sensory or perceptual systems. The major alternative
hypothesis to that of Arousal, was proposed by Harrison and Horne
(1998, 2000a, 2000b, 2000c) and has been termed the ‘Sleep-Based
Neuropsychological Perspective’.
Sleep Based Neuropsychological Perspective
The 'sleep-based neuropsychological perspective' differs from the
Arousal theory in that; 1) it does assume that sleep loss might affect
different cognitive functions differently; and 2) it posits that sleep loss
may affect different parts of the brain differently (Harrison & Horne,
1998, 2000a, 2000b). Harrison and Horne (2000a, 2000b) have argued
that sleep reduction primarily impacts the functions associated with pre-
frontal cortex (PFC) activity. The prefrontal cortex is among the first
brain regions to be affected by sleep deprivation (e.g., Drummond et al.,
2000, 2001; Muzur, et al, 2002; Thomas et al., 2000, 2003). Thomas et al.
(2000, 2003) reported progressive decreases in relative cerebral metabolic
rate for glucose (CMRglu) in prefrontal and thalamic areas over the
course of 72 hours of sleep deprivation. These changes were positively
correlated with impairments in cognitive performance and saccadic
velocity. Relative thalamic activity was also correlated with alterations in
alertness. Altena et al. (2008) reported that insomnia interferes with
activation of prefrontal cortical systems during daytime task performance.
Some other recent work even indicates that prefrontal activation may not
be fully restored by recovery sleep. Wu et al. (2006) studied sleep
deprivation and recovery sleep in normal subjects and reported that sleep
deprivation resulted in a significant decrease in the relative metabolism of
the frontal cortex, thalamus, and striatum. They reported that recovery
sleep had only a partial restorative effect on frontal lobe function and
suggest that sleep may be especially important for maintenance of frontal
lobe activity.
According to the Sleep Based Neuropsychological hypothesis,
even with relatively mild to moderate sleep loss (24 hrs), tasks that are
highly dependent on intact prefrontal cortical activity will show deficit.
These researchers predict that deficit due to sleep deprivation mainly
affects complex tasks that measure higher cortical related skills such as:
language skills in communication, divergent thinking, creativity and
cognitive flexibility. Cognitive and motor tasks demanding only basic
skills, although requiring attention and arousal, should be relatively less
affected after mild to moderate sleep loss.
Does the shift in theoretical approach to a concentration on
cognitive functions associated with intact prefrontal activation mean that
studies of the effects of sleep loss on sensory and perceptual functions are
of no real theoretical interest? In the present chapter, we argue that, even
within the framework of the newer theoretical approach, there can be
merit to studies of sensory and perceptual functions during sleep
deprivation if the functions studied meet certain criteria: 1) the sensory or
perceptual system studied has been shown to be strongly related to a
cognitive function that is impacted by sleep deprivation; 2) that cognitive
function is, itself, of theoretical and/or practical interest because of its
necessity for optimal cognitive performance.
In the following section, we present the argument that auditory
temporal resolution meets the above enumerated criteria because of its
very strong association with language and language-based skills, and is,
therefore, a prime candidate for the study of the impact of sleep
deprivation.
LANGUAGE and SLEEP DEPRIVATION
The arguments and data analyses presented in the following section
are formulated in a four-step logical structure. We: 1) review evidence
that even short term sleep deprivation impacts negatively on language and
language-based tasks; 2) review the evidence that relates auditory
temporal resolution to intact language comprehension and/or production;
3) argue that consequently a psychophysical task that measures auditory
temporal resolution, e.g., dichotic temporal order judgment (TOJ), should
be negatively impacted by sleep deprivation; and finally 4) present
evidence to support the argument.
Language-based and language-dependent skills are the basic tools
of our civilization and almost ubiquitous for successful work
performance. The maintenance of a modern industrialized society
depends upon the ability to communicate clearly and comprehensively.
Today's work place is to a large extent, circadian-independent, operating
around the clock. Furthermore, many of the indispensable services
necessary for the maintenance of health and security must be available 24
hours a day, seven days a week. Very often this scheduling involves
either around-the-clock shift work or extended work time at the expense
of normal nocturnal sleep. Because of the major importance of intact
language performance in an information-based modern industrialized
society and given the requirement of continuous high level operations
around the clock even under the most stressful conditions, it becomes
important to study the effects of sleep deprivation on language
comprehension and the neuropsychological and neurophysiological
mechanisms involved in maintaining intact language skills. Indeed,
earlier studies had reported that short term sleep deprivation may impair
normal language generation and articulation (see e.g., Harrison & Horne,
1998). Sleep deprivation had also been reported to increase the number of
inaccuracies in sending and receiving messages (Neville, Bison, French,
& Boll, 1994; Schein, 1957).These findings may be taken to imply the
expectation of difficulties in language comprehension as well as in
language generation even after mild to moderate sleep loss.
The inclusion of language-based tasks in studies of sleep
deprivation is, by no means, a new phenomenon. Language-based tasks
have been a part of many of the protocols of long term sleep deprivation
studies for many years. For example, the effects of long term sleep
deprivation on Braddely's grammatical transformation (logical reasoning)
and working memory tasks were included in the test batteries of studies
by a number of authors over 25 years ago (see, e.g., Angus & Heslegrave,
1985; Babkoff, et al, 1988; 1991a).
More recently, however, as a result of the shift in the theoretical
approach that emphasizes the impact of sleep loss on executive control
processes and other cognitive skills that are highly related to prefrontal
and frontal cortex, as noted above, the interest in the impact of even short
term sleep loss or of reduced sleep on language and language-based tasks
has increased, and become a topic of interest by itself. In a recent study,
Pilcher et al. (2007) reported a complex impact of sleep loss on language-
based tasks. Specifically, Pilcher used a full and diverse series of
language tests and language-based tasks of a wide range of complexity to
study the effects of a combined 28-hour sleep deprivation and sustained
operations schedule on non-native English speaking subjects. These tasks
included, among others: sentence completion, word analogies, antonyms,
reading comprehension and logical reasoning (taken from the Law School
Admission Test). In addition, the authors tested their subjects on verbally
presented stories from four non fiction audio books to which subjects
listened and were required to identify a "keyword" and then after 25
minutes of listening summarize the information they had heard. Verbal
working memory and the PVT (a 10-minute vigilance and reaction time)
together with several additional tasks were also included as probes in the
test battery. The authors reported that some, but not all, of the language-
based tasks were negatively impacted. Language tasks that required
sustained attention or higher level processing, such as reading
comprehension showed significant deficit whereas other tasks that were
dependent on more basic language processing, e.g., antonym
identification were not negatively impacted by the experimental
conditions. Among the tasks significantly impacted by the sustained
operations-sleep deprivation protocol were logical reasoning and a verbal
working memory task. However, a number of the relatively short
duration probe tasks, e.g., the verbal working memory, followed by the
PVT tasks were good predictors of performance on eight of the ten
language tasks used in the study. The authors concluded…" that sustained
work conditions and sleep deprivation negatively affect some types of
language performance…"
Although there is fairly clear evidence from the Pilcher et al.
(2007) study of the negative impact of short term sleep loss on language
comprehension, there may, nevertheless be somewhat of a problem with
interpreting their results, since their subjects were not only sleep deprived
but also tested in a sustained operations protocol, with a relatively high
work-to-rest ratio. Beginning with Test Session I at 1830 until the end of
Test Session IV at 1200 the next day, their subjects were continuously
occupied in the testing program except for three one-half hours of rest,
for a total work time of 17.5 hours versus a total rest time of 1.5 hours.
Thus the sustained operations component of the protocol (i.e., the very
high work-to-rest ratio encompassing the testing period of over 10:1) may
be at least partially responsible for the deficits in performance rather than
just the sleep deprivation of 28 hours (see e.g., Angus and Hestlegrave,
Other studies, however, support the claim that short term sleep
deprivation negatively impacts language-based tasks even when the
work-to-rest ratio is low (Harrison & Horne, 1998; Linde & Berstrom,
1992; Monk & Carrier, 1997). For example, Monk and Carrier (1997)
tested the performance of eighteen young adults during 36 hours of
constant wakeful bed rest on a logical reasoning task. The authors
reported that even after microsleeps, lapses in attention, and general
slowing of motor responses were eliminated from the data, there was a
significant slowing of cognitive processing. It took subjects longer to
respond to items phrased in the negative voice than to items phrased in
the positive voice. The extra cognitive processing took longer at night and
on the day following sleep loss than it did during the day before the sleep
loss. The authors interpret these results to mean that human mental
processing slows down during the night under conditions of sleep loss
even when lapses have been removed from the data. Harrison and Horne
(1998) also tested subjects who were sleep deprived for 34 hours in a
single session and reported impairment on the word fluency task and the
Haylings test. In the latter test, the authors required their subjects to
complete a sentence with a single word which was either (a) congruous,
or (b) totally incongruous with the overall meaning of the sentence.
Harrison and Horne (1998) suggested that the novelty component in
certain language tasks may make those tasks more likely to be negatively
affected by sleep loss. Taken together, it seems that overall there is
sufficient evidence that even short term sleep deprivation can negatively
impact language and language-based tasks, although it may not yet be
clear what makes certain language tasks more sensitive than others to
sleep loss, nor are the possible underlying mechanisms known. Certainly,
the mechanisms by which sleep loss affects language-based performance
are, as yet, unclear.
Auditory Temporal Resolution as a Marker of Intact Language
Performance
Over the past several decades there have been a large number of
studies that have reported a strong relationship between auditory temporal
resolution and intact speech comprehension in a variety of different
populations (Farmer & Klein, 1995; Kiss, et al, 2008; Poeppel, 2003).
These studies have compared individuals with normal language abilities
to others with language difficulties that range from developmental
difficulties to pathological neurological conditions to the difficulties
reported by normally aging older individuals in comprehending speech.
Poppel (1997) has argued that intact auditory temporal resolution is
crucial to understand speech. Normal speech is produced at
approximately 150 to 250 words per minute, i.e., 4 to 7 syllables per
second (Goldman-Eisler, 1968; Huggins, 1964). Accordingly, syllable
duration is 67 to 177 msec (Klatt, 1976), vowel duration is 70 to 130
msec (Klatt, 1976), and consonant duration is 5 to 40 msec (Kent & Reed,
1975). Rapid spectral changes such as formant transitions with
information on place of articulation may occur within a temporal range of
20-40 msec (Poeppel (2003) and syllabicity and prosodic phenomena are
reported to occur within a temporal range of 100-200 msec (Rosen,
1992). Furthermore,the temporal separations necessary to accurately
identify two auditory stimuli (gap detection) and to correctly identify the
temporal order of two stimuli in the spectral and dichotic TOJ tasks range
between 5-120 msec (Fostick, 2006; He, et al, 1999; Phillips, et al, 2000;
Schneider & Hamstra, 1999; Snell, 1997; Snell & Frisina, 2000; Snell, et
al, 2002; Strouse, et al, 1998).
Auditory Temporal Resolution and Language Development in Children

Many developmental studies of children with language and reading

difficulties as compared with their normal reading cohorts have included
tests of auditory temporal resolution. The results, however, appear to be
inconclusive: Some researchers believe that intact auditory temporal
resolution is necessary for the development of normal reading ability
(e.g., Tallal, et al, 1997). For example, Visto, et al (1996) compared
children with specific language impairment (SLI) and children with
normal language learning in their ability to process binaural temporal
information. They reported that children with specific language
impairment appear to be impaired in their ability to use non linguistic
binaural acoustic information in a dynamic ongoing fashion. Visto et al.
(1996) suggested that requirements for processing such nonlinguistic
acoustic information in a "dynamic and ongoing" fashion may be similar
to those involved in the ongoing processing of rapid changes in the
temporal and spectral components of speech. Other researchers have
concluded that intact auditory temporal resolution does not necessarily
predict the development of good reading ability in children. McAnally, et
al, (2004) compared children who were good readers and those that were
delayed readers on the discrimination of differences in the temporal
properties of sound sequences and concluded that there were no
significant differences between the two groups. However, although
researchers may disagree concerning the issue of auditory temporal
resolution as a factor of causality in language disorders in children, all
researchers seem to agree that intact temporal resolution is an important
marker of intact language function (Marshall, et al, 2001).
Complaints of Language Difficulties and Auditory Temporal Resolution
in the Healthy Normal Elderly
Normal healthy aging often includes vulnerability to difficulties in
language comprehension and auditory temporal resolution. A common
complaint among the elderly is a difficulty in understanding speech,
especially attempting to do so under less favorable conditions, such as
when speech is accompanied by noise or when speech is rapid. Several
theories have been presented to explain this age-related difficulty in
comprehending speech. Since aging of otherwise healthy individuals is
accompanied by changes in a variety of peripheral and central
physiological, behavioral and cognitive functions, some of the theories
have suggested that these changes are associated with the reported
difficulty in understanding speech. The hearing-sensitivity theory argues
that presbycusis and other age-related changes in peripheral hearing are
the main causes for the difficulties in speech comprehension (Dubno, et
al, 1984, 1997; Halling & Humes, 2000; Humes et al., 1994; Humes,
1996; Schneider, et al, 2002; Wingfield, et al, 2000). However, a number
of reviews and studies have shown this explanation to be insufficient
since correction for audiometric hearing loss does not fully explain the
difficulties experienced by the elderly in speech comprehension (Drager
& Reichle, 2001; Fitzgibbons & Gordon-Salant, 1996; Gordon-Salant &
Fitzgibbons, 1993; He, et al, 1998; Snell, 1997; Tun, 1998).
Over the past two decades, some researchers have focused on age-
related degradation in auditory temporal resolution as an explanation for
speech comprehension difficulties observed in the elderly. The rationale
underlying this hypothesis is that the appropriate use of speech cues relies
on several types of auditory temporal resolution (Gordon-Salant, 2005;
Pichora-Fuller & Souza, 2003; Schneider & Pichora-Fuller, 2001;
Schneider et al., 2002), which have been shown to decline with aging.
For example, older adults perform poorer than younger adults in gap
detection tasks and need longer silent intervals to identify the presence of
a gap when the marker signal is 250 msec or shorter (Fink, et al, 2005;
Fitzgibbons & Gordon-Salant, 2001; Grose, et al, 2006; Lister & Roberts,
2005; Lister & Tarver, 2004; Roberts & Lister, 2004; Schneider &
Hamstra, 1999; Schneider, et al, 1998; Snell, 1997; Snell & Frisina, 2000;
Snell et al., 2002; Strouse et al., 1998). Older subjects have difficulty in
correctly identifying temporal order in a tonal sequence (Fitzgibbons &
Gordon-Salant, 1998; Gordon-Salant & Fitzgibons, 1999). A number of
studies have reported that older individuals require larger differences in
duration between two tones in order to detect a difference in duration
(Fitzgibbons & Gordon-Salant, 1994, 1996, 1998). Similar results,
indicating poorer discrimination, were found when comparing older and
younger adults on binaural temporal processing tasks such as locating a
tone in the front-back plane (Abel & Hay, 1996) , tone localization (Abel,
et al, 2000), and click lateralization (Babkoff et al., 2002; Strouse et al.,
1998).
Auditory Temporal Resolution and Pathological Language Conditions in
Adults
Deficits in the processing of rapidly changing signals, as, for
example, increased thresholds for the discrimination of the temporal order
of auditory stimuli, are often correlated with phoneme- identification and
phoneme-discrimination impairments in patients with left-hemispheric
lesions to the brain and aphasia, as well as children and adults with
dyslexia (e.g., Farmer & Klein, 1995; von Steinbuchel, et al, 1999;
Swisher & Hirsh, 1972; Tallal & Piercy, 1973; Wittmann, et al, 2004).
Based on these relationships, some authors have suggested that the
processing of rapid temporal changes in speech signals is required for
correct phonemic processing (Efron, 1963; Fink et al., 2005; Poepel,
1997; Tallal, 1980; Tallal, 1984; Wittmann, 1999).
Von Steinbuchel et al. (1999) and Fink et al. (2005) compared
aphasia patients with normal controls on two measures of auditory
temporal resolution, one of which was the dichotic temporal order
judgment of two clicks (termed by the authors: "alternating monaural
stimulation mode") and reported that patients with aphasia had
significantly higher temporal order thresholds than the normal controls.
In summary, there is increasing evidence of the strong relationship
between deficits in language comprehension found in a variety of
conditions, developmental, pathological and normal age-associated, and
reduced auditory temporal resolution.
A Study of Auditory Temporal Resolution and Short Term Sleep
Deprivation
Given the strong relationship between auditory temporal resolution
and intact speech comprehension and the evidence that sleep deprivation
impacts negatively on tasks requiring intact language comprehension as
reviewed above, we attempted to address the question whether short term
sleep deprivation impacts auditory temporal resolution negatively. We
chose an experimental protocol that included a low ratio work-to-rest
schedule environment (1.5 work: 1 rest) and tested auditory temporal
resolution by measuring dichotic temporal order judgment (TOJ). In the
paradigm used in our laboratory, dichotic temporal order judgments are
measured by presenting a tone of a given frequency to one ear and the
same frequency tone to the other ear separated by times ranging between
approximately 5-240 msec. Subjects are required to judge which of the
two ears received the first stimulus. The reasons for choosing this
paradigm to measure auditory temporal resolution are discussed below.
 For the purpose of the arguments presented in this paper, we
reanalyzed the data from our earlier study (Babkoff et al., 2005)
combining the data of the first four sessions only, beginning 0830 and
ending 1610. Data over that period of time represent 24-31 hours of sleep
deprivation.
The Choice of Measure of Auditory Temporal Resolution in the Present
Study
As discussed above and expanded upon below, the hypothesis
driving the present study was that mild to moderate sleep deprivation will
affect dichotic TOJ negatively (Babkoff et al., 2005). The hypothesis is
based on the following rationale. Intact auditory temporal resolution is
strongly related to and may be a necessary component of intact language
comprehension. Intact auditory temporal resolution requires normal
activation of the prefrontal cortex. As discussed earlier, even short term
sleep deprivation has been reported to impact negatively on functions
requiring normal activation of prefrontal cortex and on some language-
based tasks. Dichotic TOJ is a valid measure of auditory temporal
resolution. Therefore, we hypothesized that 24 hours of sleep loss should
impact negatively on dichotic TOJ.
But why choose dichotic temporal judgments (TOJ) to represent
auditory temporal resolution when studying the effects of sleep loss since
there are a number of other procedures that have been used by researchers
over the past several decades to measure auditory temporal resolution? In
fact, auditory temporal resolution has been measured by: 1) the ability to
discriminate compressed speech (Versfield and Dreschler, 2002); 2) gap
detection (Schneider and Hamstra, 1999); 3) the discrimination of
stimulus duration (Fitzgibbons and Gordon-Salant, 1994); 4) localization
and lateralization (Babkoff, French, Whitmore, & Sutherlin 2002), as
well as temporal order judgment (Ben-Artzi, Fostick, & Babkoff, 2005:
Farmer & Klein, 1995). There are several reasons why we preferred the
dichotic TOJ procedure over the aforementioned procedures especially
for measuring the effects of short term sleep deprivation on auditory
temporal resolution.
First, the study was designed to use a protocol for which there is
evidence that the procedure tests an auditory temporal resolution
phenomenon in the tens to hundred msec range. Resolution in this
temporal range is crucial for decoding and understanding language (see
discussion above). Since dichotic temporal order judgment thresholds are
most often reported in the tens to hundred msec range (see e.g., Ben-Artzi
et al., 2005), such data would represent auditory temporal resolution in
the same time range as that necessary for intact language comprehension
and perhaps provide a prime facie relationship between resolution in this
temporal range and intact language comprehension.
Second, our study was designed to use a protocol for which there is
evidence that the experimental procedure clearly tests central auditory
temporal processing, rather than one that might be strongly influenced by
peripheral auditory processing. For example, one of the more popular and
often used protocols to measure temporal order judgment uses the
following procedure. Two tones of different frequencies (one high and
one low) are presented either to one ear or to both ears separated in time,
and the subjects are required to report whether the order of presentation
was high-low or low-high (Ben-Artzi et al., 2005; Hirsh, 1959; Hirsh &
Sherrick, 1961; Reed, 1989; Tallal, 1980, 1984). The identification of the
tones depends upon the difference in frequency between the two.
However, two stimuli that differ in frequency, are presented to the same
ear and are separated by very short inter-stimulus intervals often give rise
to spectrally discriminable sequences when the order is reversed. The
result is that discrimination by the subjects of the temporal order of the
two tones, high and low, may be based not only on the temporal
resolution, but also on the spectral resolution of the auditory nervous
system that begins with the interaction of the two stimuli at the peripheral
level, i.e., at the basilar membrane of the same ears(s) (see Ben-Artzi et
al, 2005 for a full discussion). Warren (1974, 1976, 1993) had noted the
difficulty in using spectral temporal order to assess temporal resolution in
a completely independent manner because of the confounding presence of
the “temporal envelope” cue. In contrast, dichotic TOJ is one of the only
procedures for which the anatomy of the auditory system guarantees that
the two stimuli used (one to each ear) are processed together only in the
central nervous system (brain stem) where encoded information from the
two ears interact for the first time (see Pastore & Farrington, 1996, for
review). Consequently, to study the effects of short term sleep deprivation
on auditory temporal resolution, we used a variation of the temporal
order judgment task, in which pairs of equal-frequency tones are
presented dichotically with varying inter-stimulus intervals separating
their arrival time at the two ears (left–right or right–left). In the dichotic
form of the temporal order judgment (TOJ) paradigm, the location of the
two stimuli (i.e. left or right) distinguishes them so that their order may
be judged, although the two stimuli in each pair are identical in
frequency. Some authors who have also used a similar temporal order
judgment paradigm have termed the procedure: "alternating monaural
stimulation mode" (Fink, et al, 2006).
Third, the study was designed to use a procedure for which there is
evidence of prefrontal cortex involvement. There is accumulating
evidence of the involvement of areas in the prefrontal cortex (PFC) in the
comprehension of rapidly changing speech and in the resolution of non-
speech auditory stimuli (Temple et al., 2000). Several researchers have
argued that speech signals that are stable or that change only slowly in
time (over hundreds of milliseconds) appear to be processed bilaterally.
However, speech signals that change rapidly (over tens of milliseconds)
appear to be preferentially processed by left hemisphere mechanisms
(Johnsrude, et al, 1997). Johnsrude et al. (1997) reported that when
normal adults discriminated auditory stimuli with frequency glides of
either long or short durations (30 ms versus 100 ms) there was significant
cortical activation foci in the left orbito-frontal cortex, the left fusiform
gyrus and the right cerebral hemisphere. Temple and Posner (1998)
reported an increase in fMRI responses in left inferior frontal cortex with
increasing speech compression. Temple et al. (2000) also reported
increased left dorso-lateral prefrontal activation in response to rapidly
changing non-linguistic acoustic stimuli. They concluded that although
left superior temporal gyrus seems to be preferentially involved in
processing meaningful stimuli, the left prefrontal region is involved with
rapidly changing non-verbal acoustic stimuli as well. Their findings and
those of Belin et al. (1998) emphasize the role of the left prefrontal region
in processing linguistic as well as non-linguistic rapidly changing
acoustic stimuli. Joanisse and Gati (2003) found regions in the posterior
portion of the left and right superior temporal gyrus and the left inferior
frontal gyrus that show increased activation to tasks requiring rapid
temporal discrimination, whether the stimuli were speech or non-speech
auditory signals. They concluded that there is significant overlap in the
brain regions involved in processing the rapid temporal characteristics of
both speech and non-speech stimuli.
A Summary of the Study: Procedure and Results
See Babkoff et al (2005) for a full description of the study,
including choice of subjects, procedure and method. In summary, seven
men and thirteen women (mean age= 23.8 yrs), who reported no
psychiatric or sleep disorders served as subjects. None of the subjects
reported consuming more than three cups of coffee or any other caffeine-
containing beverage or smoking more than one cigarette per day.
The Study
One week separated each of the two testing days. One test day
followed normal nocturnal sleep, the other test day followed 24 hours of
sleep deprivation. Subjects spent the night prior to testing at home.
Subjects were told that during the night they were sleep deprived they
could read, watch TV or videos, but must refrain from physical exercises,
smoking or ingesting any foods or beverages that contained caffeine or
alcohol. In addition, they were to remain at home. Subjects were also
instructed that on the sleep deprivation night, they were to telephone the
lab every 20 minutes beginning from 2000 until 0600 the next morning
and to leave a short message verifying they were awake. Compliance with
the stay-at-home instruction could be monitored by noting the phone
number used to relay the message. Subjects were also told that if they
failed to call even once during the sleep deprivation night, they would not
be tested the following day. The 20 subjects who were tested complied
fully with the instructions. Subjects were paid the equivalent of 950 NIS
(approximately $210) for complete participation in the experiment.
The average TOJ performance data re-analyzed and reported in this
chapter are from the first four sessions of each of the testing days
(Babkoff et al, 2005). The four sessions took place at approximately
0830, 1030, 1245 and 1530 on each of the two test days. In addition to the
formal testing sessions, all subjects were familiarized with all of the tasks
in a full training session that took place prior to the first testing day. All
testing took place in a sound isolated testing room. A cognitive testing
session took approximately 70 minutes, of which the TOJ testing
procedure lasted approximately 25-30 minutes. Because of the difference
in the duration of the testing sessions, due to the addition of
electrophysiological recordings at three of the sessions, the inter-session
intervals were not equal in length. The work-to-rest ratio over the four
sessions was approximately 1.5:1 (1.5 hours of testing for every hour of
rest).
TOJ Testing Procedure and Stimuli.
The dichotic TOJ testing procedure in the present study was
identical to the procedure used in an earlier study (Ben Artzi et al, 2005),
except that the duration of the stimuli in the present study was 10 msec.
Each session consisted of three phases: 1) the preparation and
familiarization phase; 2) the training phase; and 3) the formal testing
phase. Subjects were initially presented with six examples of each one of
the two 10 msec duration tones, one high frequency (1.5 kHz), and one
low frequency (1 kHz) to identify. Then on the next 24 trials, the two
tones were presented to one of the two ears (right or left) and the subject
indicated whether the tone was presented to the right or left ear (while
disregarding the frequency of the tone) by pressing a key on the computer
keyboard. Visual feedback (“correct/incorrect”) was provided after each
response.
At the beginning of the training phase, the subjects were presented
with the stimuli at one of the two ears in random order and required to
identify the ear receiving the stimulus. No feedback was provided.
Training continued until a criterion of 8 correct responses out of 10 trials
were met (Binomial test, p<0.01). Any individual unable to meet this
criterion was dismissed from further participation. The training then
continued with the presentation of 16 dichotic tone pairs each consisting
of 10 msec duration tones of the same frequency (e.g., 1 kHz or 1.5 kHz)
and presented with an inter-stimulus interval (ISI) of 240 msec. Subjects
were required to indicate the order in which the tones were presented
(either right before left or left before right) by first pressing a key
indicating the first ear receiving the tone and then pressing a second key
indicating the second ear receiving the tone. Feedback was provided for
all of the 16 trials.
The formal testing phase followed and consisted of 576 trials, on
each of which a dichotic tone pair of the same frequency (either 1 kHz or
1.5 kHz) was presented with either the first member of the tone pair to the
right ear followed by the second tone of the pair to the left ear or the
reverse. On each trial the two tones of each pair were separated by one of
the following inter-stimulus intervals (ISI): 5, 10, 15, 30, 60, 90,120, 240
or 400 msec. The ear receiving the first tone and the ISI were randomized
by trial. Thus there were 64 trials at each ISI, i.e., 32 trials for each of the
two frequency pairs (1kHz and 1.5 kHz) on each session for each subject.
Accuracy and RT were recorded on each trial. No feedback was provided
during the formal testing phase. Since there was no significant difference
between dichotic temporal order judgments to tone pairs of 1-kHz or 1.5-
kHz, the data were combined for all further analyses.
Dichotic Temporal Order Judgments (TOJ Thresholds)
TOJ was assessed during each of six diurnal sessions during a
waking day following: 1) normal nocturnal sleep and 2) after 24 hours of
sleep deprivation. Eighteen of the 20 subjects successfully completed the
first four sessions (0830, 1030, 1245 and 1530) under both the non-sleep
deprived and the sleep-deprived conditions. The data analyses therefore
are based on the eighteen subjects and include only the first four sessions
of each day (after normal sleep and after 24 hours of sleep deprivation.
The Greenhouse-Geisser correction was applied to all tests of
significance.
A polynomial equation was fitted to each of the accuracy-ISI
curves for each subject under each of the eight conditions (four sessions
when non sleep deprived and four sessions when sleep deprived) to assess
the dichotic TOJ threshold. Dichotic TOJ threshold was defined as the ISI
at which the accuracy-ISI best fitting polynomial curve crossed 75%
correct. The threshold values were only assessed with interpolated, but
not extrapolated values, so that no thresholds were either shorter than 5
msec or longer than 400 msec. All of the eighteen subjects who
completed four full sessions under the non sleep deprivation and the sleep
deprivation conditions, yielded psychophysical accuracy-ISI curves after
averaging, with levels greater than 75% correct at the longer ISI values so
that a TOJ threshold could be evaluated as defined above. The dichotic
TOJ thresholds of the first four sessions were averaged for each subject
and the thresholds generated under the non sleep deprived condition were
compared with those generated under the sleep deprived condition.
Figure 1 is a histogram depiction of the distribution of dichotic TOJ
thresholds for the non sleep deprived and the sleep deprived conditions
averaged over the four sessions. The dichotic TOJ threshold was longer
under the sleep deprivation condition (proportion greater than 1) than
under the non-sleep deprivation condition for fourteen of the eighteen
subjects (Binomial distribution, p<0.03).

-----------------------
Fig. 1 about here
-----------------------
The variance of the dichotic TOJ threshold distribution under sleep
deprivation, however, was significantly larger (see Fig. 1) than that of the
non-sleep deprivation condition (F (1, 17) = 3.93p < 0.004). Two types of
data analyses were performed. First, the non-sleep deprived dichotic TOJ
thresholds were compared with the sleep deprived dichotic TOJ
thresholds by the non-parametric Wilcoxon Signed Ranks test for related
data. Dichotic TOJ thresholds under the sleep-deprived condition were
significantly longer than under the non-sleep deprived condition (Z=-2.
591; p<0.01). Second, the mean dichotic TOJ thresholds from the four
diurnal sessions on which a threshold could be assessed were log
transformed and were compared by a one-way repeated ANOVA
(NSD/SD). The results confirmed the significant difference between the
assessed thresholds under sleep deprivation and non sleep deprivation
(log threshold NSD= 1.8021, sd= .29157; log threshold SD= 1.9248 sd=
.3297; F(1,17)=9.193; p< 0.008; eta2= .35). The geometric means of the
non sleep derived and sleep deprived dichotic TOJ threshold conditions
were 63.4 msec and 84.2 msec respectively.
If, as proposed and developed above, dichotic TOJ is a measure of
the resolving power of the auditory temporal domain, then these data
indicate that 24-31 hours of sleep deprivation decrease that resolving
power by approximately 32.8%.
Auditory Temporal Resolution and 24-31 Hours of Sleep Deprivation
The meaning of 32.8% reduction in auditory temporal resolution
and the possible relationship to potential difficulties in language
comprehension may be understood better by comparing a reduction in
resolution of this magnitude with similar reductions in temporal
resolution, also measured by dichotic TOJ, reported in a variety of
populations whose symptoms include a variety of language difficulties.
The purpose is for comparison only, with no pretension of implying any
similarity in mechanism.
The data presented in Table 1 allow one to compare the extent of
the reduction in auditory resolution measured by dichotic TOJ in
pathological conditions to the extent of reduction experienced by an
individual who was sleep deprived for 24-31 hours. For example,
Szymaszek,et al (2006) compared a group of young subjects with a group
of older subjects on the dichotic (their alternating monaural condition)
TOJ thresholds of click stimuli. They reported average dichotic TOJ
thresholds of 66 msec for the young subjects and 88 msec thresholds for
the elderly, a reduction in resolution of 33%. In a recent dissertation,
Fostick (2006) also compared a group of young subjects with a group of
older subjects on the dichotic TOJ thresholds of 10 msec duration tones
and found an average of 65.44 msec thresholds for the young and 85.24
msec thresholds for the elderly subjects, an estimated reduction in
auditory temporal resolution of 30.3%.
Kinsbourne et al. (1991) compared TOJ in normal and dyslexic
adult readers, using 1 msec click stimuli presented dichotically. They
used an ascending method of limits to determine a 90% threshold, which
they reported to be 46.8 msec for the normal readers and 67.4 msec for
the dyslexic readers, a 44% difference in temporal resolution for the adult
dyslexics relative to the normal readers. Ben-Artzi et al. (2005) compared
the dichotic TOJ thresholds of normal adult readers with those of adult
dyslexics using 15 msec, 1000 Hz tones as stimuli. The average dichotic
TOJ threshold of the normal adult readers was 48.89 msec, while that of
the adult dyslexics was 80.66 msec, a reduction of ~65% in auditory
temporal resolution in adult dyslexics as compared with the normal
readers. In a recent dissertation, Bar-El (2009) compared the dichotic TOJ
of adult normal and dyslexic readers and found an average threshold of
65.44 msec for the normal readers and 108.28 msec for the dyslexic
readers, a reduction of 65 % in auditory temporal resolution.
Von Steinbuchel et al. (1999) compared dichotic TOJ in normal
adults with patients suffering from unilateral focal brain lesions, localized
in anterior or posterior regions of the left hemisphere (LH) (with
symptoms of non-fluent or fluent aphasia, respectively), or in
predominantly sub-cortical regions of the LH (without aphasic
symptoms) and in the anterior or posterior regions of the RH. Dichotic
TOJ threshold was found to be 57.7 msec for the normal population and
117.5 msec for the patients with fluent aphasia, a reduction of ~ 104% in
auditory temporal resolution.
In summary, it appears that the estimated average reduction in
auditory temporal resolution as measured by dichotic temporal judgment
thresholds due to 24-31 hours of sleep deprivation is numerically quite
similar to the reported reduction in auditory temporal resolution of the
elderly as compared to young subjects, around 30%. If we rank the
amount of deficit in dichotic TOJ in the various populations and
conditions reviewed, it appears that deficit in dichotic TOJ in the elderly
and light to moderately sleep deprived young individuals ranks lowest,
around 30%, followed by deficit in the dyslexic that ranges between 44-
65%, followed by individuals with brain lesions and aphasia, who show
deficits of 100% and over, i.e., the need for more than twice the dichotic
inter-stimulus interval before the correct order of stimuli can be judged.
Auditory Temporal Resolution and Language
Comprehension: A Suggestion for Future Research
Without assuming any identity or even relation of mechanisms, but
just arguing by analogy, the similarity between the 30% deficit in
auditory temporal resolution of the elderly and of younger individuals
who experienced mild to moderate sleep loss may suggest a further
direction of research. The major difficulty of the elderly in understanding
speech seems to occur when speech is somewhat degraded or the
conditions are less than favorable, e.g., when speech is accompanied by
noise or when speech is rapid. As discussed above, the rationale
underlying the hypothesis that relates auditory temporal resolution to
language comprehension deficit in the elderly is based on the theory that
the appropriate use of speech cues relies on several types of auditory
temporal resolution. Schneider and Pichora-Fuller (2001, 2002) have
described the relationship between auditory temporal resolution and
speech comprehension, by distinguishing between three levels of speech:
(1) The supra-segmental or prosodic level, which represents lexical and
syntactic processing, and relates to perceiving the rate and rhythm of
speech; (2) The segmental level, which represents phonemic
identification, and relates to the ability to perceive envelope properties,
including the processing of duration of tones and silent intervals; and (3)
The sub-segmental level, which represents the voice pitch, quality, and
clarity of speech and relates to periodicity or synchrony perception. They
have argued that although older adults are poorer than younger adults in
speech comprehension when speech is accompanied by background noise
or when speech is rapid or compressed, the supra-segmental level seems
to be unaffected by age (Wingfield et al., 2000; Wingfield and Tun,
2001). The deficiency of older adults seem to be mainly in auditory
temporal resolution abilities related to the segmental and sub-segmental
levels of speech comprehension, even when differences in hearing
sensitivity with younger individuals has been compensated. The evidence
for the deficit in auditory resolution by the elderly was discussed above.
By analogy, perhaps we should now attempt to concentrate research
efforts in studies of mild or moderate sleep loss on discrimination and
comprehension of the segmental and sub-segmental levels of speech.
Furthermore, since the elderly seem to have difficulty mainly when
speech is somewhat degraded or the conditions are less than favorable,
e.g., when speech is accompanied by noise or when speech is rapid,
perhaps those are the speech and language conditions that we should
concentrate on in our future research efforts in studying the effects of
sleep loss on language and language-based tasks?
 The estimate of 30% deficit in auditory temporal resolution during
24-31 hours of sleep deprivation in a low level work environment of
approximately 1.5: 1 work-to-rest ratio is of theoretical interest for
another reason as well. Almost 25 years ago, Angus and Heslegrave
(1985) discussed the differences between decrements in performance
during sleep deprivation in a fairly low level work –demand environment
as compared to sleep deprivation combined with a high work-demand
environment, such as sustained or continuous operations. Earlier studies
had indicated little or no performance decrement (approximately 10%)
during the day following a night of sleep loss (Naitoh, 1981 Opstad,
Ekanger, et al, 1978); whereas Angus and Heslegrave (1985) reported
fairly large decrements in performance during the day following one
night of sleep loss. Angus and Heslegrave (1985) pointed out that the
protocol of their study was one of intensive and sustained cognitive
performance in addition to sleep loss and consequently the decrements
they found represent the combined effects of sleep loss and the high
work-to-rest ratio. A number of language and language-based tasks were
included among the cognitive performance tasks tested by Angus and
Heslegrave (1985), e.g., logical reasoning, encoding-decoding, message
processing. They reported an average decrement of approximately 30% in
performance during the day following a night of sleep loss. Given the
fairly low work-to-rest ratio in the testing protocol of the study we
reported herein, the estimated deficit in auditory temporal resolution of
approximately 30%, seems quite high and perhaps reflects the sensitivity
of the neural language-processing mechanisms to even relatively short
term sleep deprivation as posited by Harrison and Horne (1998, 2000a,
2000b) in their 'sleep-based neuropsychological perspective' hypothesis
of the effects of sleep loss.
Language, Auditory Temporal Resolution and Sleep Deprivation: A
Summary and Caveat
We should not lose sight of the argument made earlier that interest
in the impact of sleep loss on very basic sensory/perceptual functions that
by themselves are not subjects of concern to the sleep deprivation
literature, have importance only to the extent that they are closely
associated with cognitive functions that are reported to be negatively
impacted by sleep loss. Auditory temporal resolution is, thus, of interest
because of its close association with language comprehension. Given the
present state of knowledge regarding sleep loss and language
comprehension, it is certainly not yet clear whether all aspects and
components of language are equally sensitive to short term sleep loss or
not. To date the most comprehensive study of the impact of sleep
deprivation on language-based tasks is that of Pilcher et al. (2007).
However, from the Pilcher et al. (2007) study reviewed above, it appears
that"… sustained work conditions and sleep deprivation negatively affect
some (but not all) types of language performance". We do not know
what class of language-based tasks is sensitive to short term sleep loss
and which class or classes are less sensitive. Consequently we do not
know whether the language-based tasks that are sensitive to sleep loss are
the same tasks that are closely associated with auditory temporal
resolution or not. What seems to be fairly clear from the data reviewed
and analyzed herein, is that auditory temporal resolution is sensitive to
short term sleep loss. Furthermore, since the association between auditory
temporal resolution and complaints of language difficulties, as reviewed
and discussed above, seems to be that of fairly basic comprehension of
spoken and written language under somewhat difficult conditions, we
predict that similarly, it is that aspect of language that would be most
sensitive to sleep deprivation, namely the basic understanding of written
and spoken language under fairly difficult conditions. So far, no
systematic studies have been undertaken to examine the impact of sleep
deprivation on a wide variety of basic and complex language and
language-based tasks and provide an answer to these questions.
Figure Legends
Figure 1. The distribution of Dichotic TOJ thresholds when subjects were
non sleep deprived and when they were sleep deprived are plotted
separately.
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 Table 1. This Table compares the reduction in dichotic TOJ resolution after 24-31
hours of sleep deprivation as compared with dichotic TOJ resolution after normal nocturnal
sleep with reductions in dichotic TOJ resolution found among various groups of individuals
with a variety of language disorders.
Author Stimuli Method Control Control Expt’l Expt’l Estimated
Group TOJ Group TOJ Reductio
Threshol Threshol n in
d d Temporal
Resolutio
n
Present 10 Constant Normal 63.35 Same 84.25 29.65 %
Study Msec stimuli. adults Msec subjects, Msec
dichoti 75% non tested
c tones threshold sleep- after 24-
s deprive 31Hr
d Sleep
(N=18) deprivatio
n
Ben-Artzi 15 Constant Adult 48.89 Adult 80.66 65%
et al. Msec stimuli. normal Msec dyslexic Msec
(2005) dichoti 75% readers readers
c tones threshold
s
Kinsbourne 1 Msec Ascendin Adult 46.8 Adult 67.4 44%
et al. dichoti g method normal Msec dyslexic Msec
(1991) c of limits. readers readers
clicks Threshol
d, 90%
Correct
Von Dichoti Adaptive Normal 57.7 Patients 117.5 104%
Steinbuchel c clicks staircase. adults Msec with Msec
et al. Criterion lesions in
(1999) set for posterior
75% region of
threshold LH
s (Fluent
Aphasia)
Szymaszek Clicks Yet Young 66 Msec Elderly 88 Msec 33%
et al. Another Subject Subjects
(2006) Adaptive s
Procedur
e
Bar-El, S. 10 Constant Adult 65.44 Adult 108.28 65%
Unpublishe msec Stimuli normal Msec dyslexic Msec
d dichoti 75% readers readers
dissertation c tones threshold
(2007) s

Fostick, L. 10 Constant Young 65.44 Elderly 85.24 30.3%

Unpublishe Msec Stimuli Adult Msec normal Msec
d dichoti 75% normal readers
dissertation c tones threshold readers
(2006) s