Psychophysiology, 37 ~2000!, 2942. Cambridge University Press. Printed in the USA.

Copyright 2000 Society for Psychophysiological Research

Early attention effects in human

auditory-evoked potentials

JRG HOORMANN, MICHAEL FALKENSTEIN, and JOACHIM HOHNSBEIN

Institut fr Arbeitsphysiologie an der Universitt Dortmund ~IfADo!, Dortmund, Germany

Abstract
A fundamental question in attention theory concerns the earliest processing stages that can be modulated by selective
attention. A series of experiments is reported in which very early attention effects are found under specific conditions
in the frequency-following potential ~FFP!, a brain stem response to low-frequency tone stimuli. In two experiments,
stimuli of two different modalities were applied, and attention directed to one of the modalities. In two further
experiments, only auditory stimuli were presented. In the first of these last two experiments, a dichotic paradigm with
sustained attention to one ear was used, in the second a monotic paired-stimuli paradigm was used, in which the first
stimulus served as reference for the second one. Only in the last experiment significant attention effects were found in
the latency, but not in the amplitude of the FFP. The results show that a very early attention effect on the latency of the
FFP can be demonstrated, but only under highly specific conditions. The size and preconditions of the attention effect
suggest that it reflects subtle intramodal tuning mechanisms in the cochlea or in the lower brain stem.
Descriptors: Brain stem, Evoked potentials, Frequency-following potential, FFP, Attention

Peripheral Sensory Gating
It has been long known from animal studies that efferent neurons
can modulate activity within afferent pathways, even at the level of
the sensory receptor ~Granit, 1955; Hernandez-Peon, Scherrer, &
Jouvet, 1956!. On the basis of these data, Hernandez-Peon ~1966!
proposed the peripheral filter model, which assumes that a de-
scending tonic inhibition can modulate sensory neurons. In fact,
the hardware for such a filter exists, at least in the auditory system
of mammals, namely the olivocochlear bundle ~OCB!. The OCB
consists of two subsystems, a medial and a lateral subsystem
~Guinan, 1988!. In the medial OCB, efferent fibers arise from cell
bodies in the superior olivary complex and terminate in the co-
chlea. This OCB appears to modulate the active micromechanical
properties of the outer hair cells ~Mountain, 1980; Siegel & Kim,
1982!. Scharf, Quigley, Aoki, Peachey, and Reeves ~1987!, who
conducted psychophysical studies, postulated that this modulation
is an active process to control specifically those frequencies that
are task relevant. Hence it is reasonable to assume that early at-
tention effects can be mediated by efferent pathways such as the
olivocochlear bundle. Indeed, classical animal studies have dem-
onstrated effects of auditory attention at the level of the acoustical
We thank Michael Smieja for his support with the hardware develop-
ment, Ludger Blanke for developing the software for EEG processing and
presentation, Brigitta Voss for her helpful assistance with the data evalu-
ation, Christiane Westedt for producing the figures, and Carl R. Cavonius
for checking the manuscript. We also thank Steven Hackley for most valu-
able comments and discussions during the preparation of this paper.
Address reprint requests to: J. Hoormann, Institut fr Arbeitsphysiolo-
gie an der Universitt Dortmund, Ardeystr. 67, D-44139 Dortmund, Ger-
many. E-mail: hoormann@arb-phys.uni-dortmund.de.
nerve ~Oatman, 1976; Oatman & Anderson, 1977! and the brain
stem ~Gabriel, Saltwick, & Miller, 1975; Olesen, Ashe, & Wein-
berger, 1975!. In humans, evoked otoacoustic emissions ~EOAEs!
have been used recently to demonstrate attention effects, because
these emissions are sensitive to mechanical alterations of the outer
hair cells. The evidence of the studies on otoacoustic emissions is
not uniform. Earlier studies ~Avan & Bonfils, 1991; Freedman,
Adler, Waldo, Patchman, & Franks, 1990; Froehlich, Collet, Cha-
nal, & Morgon, 1990; Meric & Collet, 1992, 1993; Puel, Bonfils,
& Pujol, 1988! suffered from methodological problems, that is,
comparison of passive with discrimination conditions. This criti-
cism does not apply to two more recent studies by Giard, Collet,
Bouchet, and Pernier ~1994! and Michie, LePage, Solowij, Haller,
and Terry ~1996!. However, whereas Giard et al. ~1994! did find a
small ~average of 0.5 dB! but consistent effect of attention on
otoacoustic emissions in a narrow frequency band around the evok-
ing frequency, Michie et al. ~1996! did not find such an effect.
In sum, from animal studies there is ample evidence for pe-
ripheral gating even at the cochlear level, although attempts to
demonstrate such gating for humans in recent psychoacoustical
studies have been doubtful. To clarify this issue, since the early
1970s electroencephalographic data were used to study early at-
tention effects, which led to the results summarized in the following.
Electrophysiology of Sensory Gating
The physiological and psychological mechanisms of selective at-
tention can be analyzed in humans by recordings of event-related
potentials ~ERPs! from the scalp. Attention effects have been ob-
served consistently for long-latency auditory ~and visual! ERPs.
The most prominent attention-related phenomenon is a negative

29
30
displacement for attended ~as opposed to unattended! auditory
stimuli, which has been called the N1 effect ~Hillyard, Hink,
Schwent, & Picton, 1973!, or the processing negativity ~Ntnen,
1982; Ntnen, Gaillard, & Mntysalo, 1978!. This phenomenon
is probably not a modulation of the N1, but an endogenous com-
ponent that under certain conditions overlaps the N1. However,
apart from the processing negativity, real amplitude modulations
not only of the N1, but of other exogenous components ~e.g., the
P2! can be observed under conditions that are optimized for early
selection ~e.g., Woldorff & Hillyard, 1991!. McCallum, Curry,
Cooper, Pocock, and Papakostopoulos ~1983! even observed a
negative shift in the ERP to attended versus unattended stimuli as
early as 15 ms after stimulus onset. In contrast, with a different
task, Woldorff and Hillyard ~1991! found the ERP to be more
positive in the time range of 2050 ms for attended versus un-
attended high-frequency tone pips ~P20-50!. By recording neuro-
magnetic fields, Woldorff et al. ~1993! could locate the sources of
the P20-50 as well as the later effect around 100 ms in the auditory
cortex. This result confirms the findings of Scherg and von Cra-
mon ~1985!, who located the sources of middle-latency compo-
nents in the 2030-ms range in the primary and secondary auditory
cortex with the help of dipole estimation.
From these studies it is clear that selective attention can influ-
ence sensory processing in the auditory cortex. However, the ques-
tion remains open whether any modulation of auditory processing
can operate at a more peripheral level, that is, in the brain stem or
even the cochlea.
The Brain Stem Evoked Response (BER)
Beginning in the early 1970s, considerable effort has been devoted
to investigate possible attention effects on very early auditory ERP
components in humans ~Picton & Hillyard, 1974; Picton, Hillyard,
Galambos, & Schiff, 1971; Woods & Hillyard, 1978!. Picton et al.
~1971! studied the cochlear nerve response, which has a latency of
about 2 ms after click stimuli, in which high frequencies predom-
inate. These authors failed to find any attention-related modulation
of this very early auditory potential. Only a short time later the
far-field electrical activity from the human brain stem was discov-
ered by Jewett and Williston ~1971!. These brain stem evoked
responses ~BERs!, which are evoked by short click stimuli, consist
of five to seven positive components ~I to VII! with peak latencies
between 1.5 and 7 ms. Picton and Hillyard ~1974! examined BERs
during click intensity discrimination and during a passive control
condition. They found no differences in any BER component be-
tween the attention ~discrimination! and control conditions, whereas
later ERP components showed a strong amplitude modulation.
Woods and Hillyard ~1978! considered the possibility that periph-
eral gating might operate mainly in the context of ecologically
relevant, complex stimuli, such as speech. They presented clicks
superimposed on different spoken prose passages presented to the
two ears, while one of the passages had to be attended to. Again the
click-evoked BERs were unaffected by attention. Together these
results were interpreted as clear evidence against a peripheral gat-
ing mechanism of auditory attention.
In the studies cited, attention was manipulated only within one
~the auditory! modality. However, peripheral gating or filtering
may be more effective if the difference between the attended and
ignored stimuli is greater. Hence the optimum strategy to demon-
strate early attention effects could be to use intermodal paradigms
in which stimuli of two ~or more! modalities are presented, one of
which has to be attended. In fact, in the 1970s, a series of animal
J. Hoormann et al.
studies conducted by Oatman and co-workers ~Oatman, 1971, 1976;
Oatman & Anderson, 1977, 1980; Glenn & Oatman, 1977! showed
clear attention effects on brain stem potentials in a bimodal para-
digm, when attention was directed to or away from the visual
modality. Lukas ~1980, 1981! adapted the bimodal methodology
for experiments with humans. In both studies by Lukas, visual and
auditory stimuli occurred at random. In the first experiment of the
1980 study, the subjects had either to count visual target letters or
to listen to tone pips of 1000 Hz. In the visual discrimination
condition, the amplitude of wave V was smaller, and its latency
longer, than in the listening condition. In a second experiment with
higher stimulus frequencies ~2000 and 8000 Hz!, similar attention
effects were observed, but only for 2000 Hz.
The problem of this study is that tasks of different complexity
were assigned to the visual ~discrimination! and the auditory ~lis-
tening! modality. Hence in a second study ~Lukas, 1981! subjects
had to detect targets in both modalities. The auditory stimuli were
8000-Hz tone pips ~targets had a longer duration than nontargets!.
No attention effect was found on the nontarget BER, but an effect
on wave I for the targets.
Lukas explained this targetnontarget difference by the differ-
ent frequency content of his targets and nontargets ~due to the
different duration!, which hence stimulate slightly different areas
of the basilar membrane. However, this explanation is unlikely,
because the power spectra of the two stimuli differ only slightly.
Furthermore, the specific delay of wave I ~and not of the later
waves IIV! seems improbable, because wave I reflects the summed
activity of the eighth nerve action potentials ~Achor & Starr, 1980;
Moller, Jannetta, Bennett, & Moller, 1981!. A final critical point in
that study is the small number of averaged trials for targets.
Picton, Stapells, and Campbell ~1981! tried to replicate the
results of Lukas ~1980! with an auditory listen0visual discrimina-
tion paradigm. They found no differences between the conditions
for waves I and V. In a preliminary report, Brix ~1984! measured
BERs to ~unspecified! clicks in 100 subjects who had either to
attend to the clicks ~trying to count them! or to read a newspaper.
The author found a shorter interpeak latency between waves I and
V in the attention condition. However, it remained unclear whether
this shortening was due solely to a shortening of wave V latency.
Also, the number of averaged epochs was rather small, and no
significance level for the effect was reported.
Sommer ~1985! conducted a cross-modal attention study with
individuals with schizophrenia. For a subgroup of the patients who
showed no electrodermal response to orienting stimuli ~nonrespond-
ers!, the author found longer wave V latencies when visual rather
than auditory stimuli were attended.
Connolly, Aubry, McGillivary, and Scott ~1989! used both intra-
and intermodal attention paradigms with high-frequency clicks as
auditory stimuli. These authors also found no significant change in
the BER waves across the attention conditions, which was again a
failure to replicate Lukas ~1980! earlier results. However, their
Experiment 1 was not likely to have manipulated attention effec-
tively, because the visual task was too easy and coupled with the
auditory task.
Gregory, Heath, and Rosenberg ~1989! compared BERs in a
passive condition versus an auditory versus a visual selective at-
tention condition. In the two latter conditions, subjects had to
count occasional high-frequency targets ~8-kHz tones! among lower-
frequency pips, or to count deviant-colored shapes among other
shapes. The authors found no attention-related effects on the BER.
Hackley, Woldorff, and Hillyard ~1990! investigated cross-
modal selective attention effects on retinal, myogenic, brain stem,
Early attention effects and evoked potentials
and cerebral potentials. The visual stimuli were flashes, the audi-
tory stimuli high-frequency ~4000 Hz! tone pips. The target stimuli
~about 5%! were less intense in both modalities. Whereas the longer-
latency components showed clear attention-related enhancements,
neither the BER nor the early visual components ~electroretino-
gram, occipital components between 40 and 70 ms! were signifi-
cantly affected by attention.
Bauer and Bayles ~1990! conducted an intermodal study in-
cluding the auditory and the somatosensory modalities. To focus
attention within certain time periods, trains ~10s long! of clicks
were presented to the left ear via headphones; the trains were
separated by pauses. subjects awaited either an auditory target ~a
500-Hz tone presented from an additional loudspeaker placed be-
hind the subjects! or a somatosensory target presented with near-
threshold intensity at a random time during a train. Waves II and
V of the BER were significantly larger and peaked earlier in the
auditory attention than in the somatosensory attention condition.
The authors suggested two possible factors for their positive re-
sults: ~i! the high task demands, that is, the use of low-intensity
targets, which led to a highly focused attention, and ~ii! the large
separation of targets and nontargets, that is, by frequency, time-
course, and location. A further reason might be the good signal-
to-noise ratio, which is usually obtained when the subjects could
close their eyes, which is possible when visual stimuli are not
involved.
Hirschhorn and Michie ~1990! attempted to compare inter-
modal with intramodal attention effects on the BER, using visual
and auditory stimuli. The auditory stimuli were loud clicks with
main frequencies around 1400 Hz ~nontargets! or 900 Hz ~targets;
4%!. The visual stimuli were nonsemantic shapes ~containing 25%
targets!. The stimuli were randomized. subjects had to attend to
one of the modalities either in a passive listen0look condition or in
an active discrimination condition ~which was varied in difficulty!.
Several comparisons were made for the amplitudes and latencies
of all BER waves. Among other significant findings, the latency of
wave I was prolonged during the difficult visual discrimination
condition compared with the listen condition. This result replicated
the early results of Lukas ~1980! and suggests an inhibition of all
auditory stimuli during highly focused visual attention. However,
this result was not reflected in the later BER waves. Also, the large
number of statistical tests ~80 altogether! greatly increases the
possibility of reaching significance by chance. On the other hand,
the only significant effects were found for waves I and II, which is
unlikely to be due to chance.
In summary, whereas the intramodal BER studies so far showed
no effects, the results of the intermodal BER studies are less con-
sistent. Although most of the intramodal BER studies likewise
failed to demonstrate any attention effects, a few intermodal stud-
ies did show effects on the BER waves I, II, and0or V. It cannot be
excluded that the positive results were chance findings. On the
other hand, it is possible that under certain conditions positive
effects may be real. It seems worthwhile to delimit such condi-
tions. One basic precondition seems to be a sufficiently complex
task. Also, positive effects were observed mainly for latencies
rather than for amplitudes of the BER waves. Finally, there seems
to be a tendency for positive results when lower stimulus frequen-
cies are used. This tendency suggests the possibility that there are
no peripheral gating mechanisms for high-frequency stimuli, but
that there are for low-frequency stimuli such as speech ~which are
ecologically more valid for humans!. This possibility has already
been mentioned by Woods and Hillyard ~1978!. Unfortunately, the
frequencies that contribute to the BER are usually higher than the
31
main speech frequencies. Hence high-frequency click stimuli, such
as also used by Woods and Hillyard, might not be optimally ap-
propriate. Consequently, brain stem responses that are sensitive to
lower frequencies might be more usefully examined.
The Frequency-Following Potential (FFP)
The FFP ~Marsh, Smith, & Worden, 1972; Moushegian, Rupert, &
Stillman, 1973! is a brain stem evoked response, which can be
elicited by relatively low-frequency stimuli. The FFP is best re-
corded from the vertex with reference at the mastoid contralateral
to the stimulated ear. However, because at least two sources or
pathways for the FFP are assumed ~Marsh, Brown, & Smith, 1974!,
an ipsilateral reference is also frequently used ~e.g., Stillman, Crow,
& Moushegian, 1978!. The FFP reproduces the frequency of the
waveform of the eliciting stimulus, and its onset is delayed by
about 6 ms relative to stimulus onset, which suggests a lower brain
stem origin ~Galbraith & Brown, 1990! ~cf. also Figure 1!.
Marsh et al. ~1972! suggested the inferior colliculus as the
origin of the FFP. However, some years later, evidence was found
that more peripheral brain stem sites, such as the superior olivary
complex or even the cochlear nucleus produce the FFP ~Gardi,
Merzenich, & McKean, 1979; Hoormann, Falkenstein, & Hohns-
bein, 1992a!. The FFP is thought to depend on phase-locking
neural elements ~Gardi et al., 1979; Marsh et al., 1972; Stillman
et al., 1978!, which code other stimulus properties than do neurons
responsive to transients. The amplitude of the FFP is estimated by
calculating the fast Fourier transform and measuring the peak at
the stimulus frequency. The FFP latency is estimated by calculat-
ing the cross-correlation function between stimulus and FFP and
determining the latency of the maximum of this function. Hence a
methodological advantage of the FFP over the BER is that ampli-
tude and latency measures depend on the entire waveshape and not
on a single peak, which enhances the accuracy of the measure-
ment. The maximum amplitude FFP is obtained with tones around
350 Hz and is diminished strongly for frequencies beyond 500 Hz.
For very low frequencies ~100 to about 200 Hz! the FFP shows in
addition a strong second harmonic of the eliciting frequency ~Hoor-
mann et al., 1992a!. The FFP can be also elicited by vowels and
natural speech stimuli ~Galbraith, Arbagey, Branski, Comerci, &
Rector, 1995; Galbraith, Jhaveri, & Kuo, 1997!. Hence the FFP
appears to be most sensitive to frequencies in the main speech
range. If attention affects mainly speech information ~Woods &
Hillyard, 1978!, the FFP should be the most appropriate tool for
assessing very early auditory attention effects.
Oatman and Anderson ~1977, 1980! used the FFP in their an-
imal studies. Cats were trained to perform a visual attention task.
The visual stimuli consisted of two concentric rings presented with
varying stimulus onset asynchrony ~SOA!. The cats had to press a
lever after but not before the occurrence of the second ring. Tone
bursts of different frequencies were presented simultaneously. The
authors demonstrated a clear suppression of the FFP amplitude for
all stimulus frequencies during visual attention compared with pre-
and posttest passive control conditions.
In a first attempt to study attention effects on the FFP in hu-
mans, Hillyard and Picton ~1979! used an intermodal paradigm in
which attention was directed to the auditory or to the visual mo-
dality. Hillyard and Picton could find no effects on the amplitude
of the FFP despite significant effects on later components. Possible
effects on FFP latency were not mentioned.
Galbraith and Kane ~1993! also investigated intermodal atten-
tion effects on the FFP. In a visual attention condition, subjects
32
Figure 1. Example for a pure tone burst ~320 Hz; upper panels! that elicits a frequency-following potential ~FFP; lower panels,
averaged response of a single S!. The left panels present the time course, the right panels the frequency contents of both stimulus and
response. The response is delayed by about 6 ms relative to the stimulus and reproduces its frequency.
performed a complex visual task, while ignoring simultaneously
presented 230-Hz tone pips. In an auditory attention condition with
closed eyes, the subjects had to silently count complex auditory
stimuli interspersed among the tones. Although the cortical-evoked
potential to the tones showed a clear attention-related modulation,
no attention effect on the FFP ~averaged across 750 sweeps! was
found. The authors attributed their negative results to the very high
discriminability of targets from nontargets in the auditory attention
condition, which may have rendered a strong focusing of attention
unnecessary. Moreover, both of these studies used low stimulus
rates ~which might be ineffective in terms of maximum focussing
of attention! because the investigators intended to record long-
latency ERPs in addition to FFPs. Finally, the number of sweeps
~about 750! for the FFP in the latter study may be too low for
detecting small differences.
In a further study, Galbraith and Arroyo ~1993! used the FFP in
an intramodal selective attention task. They presented different
tone bursts ~with frequencies of 200 and 400 Hz, respectively!
randomly and asynchronously to the two ears, the interstimulus
interval ~ISI! was randomized between 120 and 320 ms. In differ-
ent blocks, subjects had to attend to one particular ear and fre-
quency ~e.g., high tones in the left ear! and press a key to each
target, which was less intense than the other tones. Because the
FFP waveshape appeared to consist of two parts, the FFP ampli-
tude was determined for the first and second parts separately. A
complex result was found: for attended 400-Hz stimuli, the am-
plitude of the second part of the FFP was reduced compared with
the first part, whereas the opposite was found for nonattended
400-Hz stimuli. For 200-Hz stimuli, the result for the ignored
stimuli was in the opposite direction. The authors stated cautiously
that some form of attention-related modulation may be occurring
in the lower brain stem. The complex result was explained by the
J. Hoormann et al.
overlapping activity of two FFP generators, which had been claimed
by Stillman et al. ~1978!.
In a further study, Galbraith and Doan ~1995! presented stimuli
with a different quality, namely tones and missing fundamental
~MF! stimuli. MF stimuli have the same pitch as the pure tones of
the fundamental frequency, but do not physically contain the fun-
damental frequency. MF stimuli have been shown to evoke similar
FFPs as tone stimuli ~Galbraith, 1994; Greenberg, 1980!. Galbraith
and Doan ~1995! assumed that different anatomical and physio-
logical properties are involved in the coding of MF and pure tones
and could hence yield different FFP results with respect to atten-
tion. For both types of stimuli, the targets differed from nontargets
either by a lower intensity ~8-dB difference! or by a longer dura-
tion ~75 vs. 25 ms!. Attention was directed to one of the ears in a
block; the direction of attention was changed between blocks. The
FFP amplitude determination was based on the entire length of the
waveshape ~i.e., not divided into two parts as in Galbraith & Ar-
royo, 1993!. Again a complex result was found, namely that the
FFP was only larger for attended ~vs. unattended! MF stimuli,
whereas the FFP was even smaller for attended tones, particularly
when intensity was the discriminative cue. The latter discrimina-
tion was also the most difficult one, as shown in the d9 scores. This
pattern of results is hard to explain and may be partly due to
differences in task difficulty between the conditions, as was also
discussed by the authors. However, the fact that the task with the
most difficult discrimination yielded a negative attention effect is
puzzling. If the FFP effects for the MF and the easy duration
discrimination were due to attention, they should become even
larger for a harder discrimination, because a harder discrimination
requires even more attention. In addition, the overall discrimina-
bility of targets and nontargets proved to be high, as revealed in a
mean hit rate of 90%. Also, no effects on late ERP components in
Early attention effects and evoked potentials
the N1 range were reported. Hence it is doubtful whether attention
was effectively manipulated in that study, and a nonattention ex-
planation for the results should be considered. A further problem
may have been that only 500 sweeps were averaged to obtain the
FFP in each condition, a number that does not seem to be sufficient
considering the small size of the expected attention effects.
In sum, the results of attention effects on FFPs are contradic-
tory, and no latency results were reported. However, the two most
recent reports of the Galbraith group yielded at least partially
positive results.
The Present Study
The present study was aimed to continue the research on attention
effects with the FFP, because early attention effects are expected to
occur in the low frequencies in the speech band. Besides looking
for amplitude effects, we were particularly interested in latency
effects, because these effects have not been reported up to now, but
can be expected, because latency effects were reported several
times for the click-evoked BER. Moreover, the variance of FFP
latency is smaller than that of FFP amplitude ~Hoormann et al.,
1992a!. We studied FFPs with several paradigms; the bimodal
paradigm was thought to be the most promising, because the to-
be-attended channels ~the modalities! have maximum separation
~cf. Connolly et al., 1989!. To further enhance the focusing of
attention, the discriminability of targets and nontargets was made
difficult ~Woldorff & Hillyard, 1991!. In two experiments we used
a bimodal paradigm with one modality attended. Attention was
either sustained for longer time intervals ~Experiment 1! or the
relevant modality was cued in each trial ~Experiment 2!. To study
possibly different distraction mechanisms for different modalities,
the modality pairs were varied ~auditory-visual or auditory-
somatosensory! in Experiment 2. In two further experiments, only
auditory stimuli were used ~unimodal paradigm!. In the first of
these ~Experiment 3!, a dichotic listening paradigm with sustained
attention was used; during one block subjects had to attend exclu-
sively to one ear and press a key after the occurrence of an occa-
sional deviant. As in the study of Galbraith and Doan ~1995!, pure
tones and missing fundamental stimuli were used. To control for
the effectiveness of the attention manipulation, middle- and long-
latency ERPs ~up to 200 ms! were also measured. In the second of
these experiments ~Experiment 4! ~short report in Hoormann, Falk-
enstein, & Hohnsbein, 1994! monotic paired-stimuli paradigm with
short intervals between the pairs was used, in which the first stim-
ulus served as reference for the second, which could be the same
or ~occasionally! slightly different. subjects had to focus attention
very strongly after the first stimulus to be able to distinguish for
the subsequent stimulus a target from a nontarget and could then
reduce the focussing of attention until the next trial, since the first
stimulus was always the same. This paradigm was thought to lead
to a strong phasic focusing of attention to the critical time segment.
Methods
General Remarks
Because we expected only small ~if any! attention effects, careful
method refinements were carried out in advance to enhance the
quality of the FFP. First, we avoided the use of earphones because
they either lead to stimulus artifacts in the response or, if shielded
by mu-metal, they may distort the signal ~Batra, Kuwada, & Maher,
1986; Hoormann, Falkenstein, & Hohnsbein, 1992b!. Instead, for
each stimulated ear we used a separate tube device that delayed the
33
acoustic signal by about 30 ms relative to the electromagnetic
signal, hence excluding stimulus artifacts in the response window.
The device consists of a mu-metal shielded low-frequency speaker,
which was coupled via an exponential horn to a long PVC tube.
The tube was led into a sound-attenuated, electrically shielded
chamber. The proximal end of the tube was coupled to a circu-
maural ear cushion. The total distance between the speaker mem-
brane and the eardrum was 9.2 m, which causes the 30-ms delay of
the auditory stimulus. The electrical signal to the speaker was
prefiltered with an active filter to equalize the frequency charac-
teristic of the entire system. Distortions of the signal at the tube-
exit induced by the entire system were negligible, which was
monitored via a miniature microphone in the ear cushion. Here the
second harmonic of the signal was less than 55 dB ~rel. funda-
mental! in the range of 1281088 Hz. ~For further technical details
cf. Hoormann et al., 1992b.!
Second, several techniques were used to enhance the signal-to-
noise ratio: ~i! to reduce the total electromyogram ~EMG!, the
subjects lay on a vacuum mattress shaped exactly to their body,
which enabled them to relax very comfortably; ~ii! special low-
noise preamplifiers were used and the gain was adjusted individ-
ually; ~iii! the passband was filtered very sharply with 10-pole
Butterworth filters; and ~iv! a sufficient number of artifact-free
sweeps, that is, at least 2,500 per condition, were averaged to
obtain the FFP. To resolve very small latency differences, the
electroencephalogram ~EEG! was sampled with 250 kHz per
channel.
EXPERIMENT 1
This was a cross-modal attention study with sustained attention to
one modality.
Methods
Subjects
Twenty healthy subjects ~10 men, 10 women, mean age 5 25
years; naive as to the purpose of the experiment! participated. All
subjects had normal ~or corrected-to-normal! vision. Before the
EEG session their hearing threshold was measured at 10 test fre-
quencies by an audiometer and found to be within normal limits
~65 dB; American National Standards Institute, 1970!. The sub-
jects were paid 15 DM per hour for participating.
Stimuli
The auditory stimuli were low-frequency tone bursts presented to
the left ear via the auditory delay line. The stimulus frequency was
320 Hz ~nontargets! or 337 Hz ~targets!. The stimulus envelope
was symmetrically Gaussian-shaped enclosing a plateau ~for de-
tails cf. Hoormann, Voss, Falkenstein, & Hohnsbein, 1997!; the
total stimulus length was 10 cycles ~i.e., circa 30-ms duration as
were all stimulus durations in all four experiments!, including two
cycles rise and two cycles fall time. The pitch difference between
targets and nontargets was chosen in pilot studies to establish
roughly a 75% probability of correct target identification. The
intensity of the stimuli was adjusted individually to 80 dB HL.
The high intensity was chosen to obtain maximum amplitudes of
the FFPs ~Hoormann et al., 1992a!. To avoid the overlap of FFPs
from both lateral brain stem sites ~as elicited by binaural stimula-
tion!, the right ear was masked with white noise of 65 dB SPL. The
visual stimuli were LED flashes of 50 ms length. The LED was a
duo-LED ~capable of emitting green or red light or a mixture of
34
both!, which was controlled such that either pure green flashes ~non-
targets! or green flashes with a slight addition of red ~targets! were
emitted. The intensity was held constant for nontargets and targets.
The hue difference between targets and nontargets was chosen in
pilot studies to establish a roughly 75% correct target identification.
Procedure
Each session had three parts: audiometry, attention part, and pas-
sive part. In the attention part, auditory and visual stimuli were
presented in random sequence with an ISI randomized around
320 ms ~240 400 ms!. The stimuli were presented in blocks of
100 stimuli; before each block a visual or an auditory cue was
given that indicated the to-be-attended modality during the follow-
ing block. The visual cue was a pure red stimulus from the duo-
LED, the auditory cue was a 1-kHz tone. Visual and auditory cues
were given in alternation, that is, the subjects had to change the
attended modality every block. About 320 ms after the cue the
stimulus train began. In the attended modality, 5% of the nontar-
gets were replaced randomly by targets, whereas in the non-
attended modality only nontargets were presented to avoid the
possibility of distraction of attention by targets in the nonattended
modality. The task was to press a key after each target. The first
three sweeps were excluded from averaging in each block. After
each 10 min a short break was inserted. Altogether the attention
part had a length of about 80 min. In the passive part, only auditory
stimuli were presented, with the same ISI as in the attention part.
The subjects had simply to relax, and ignore the stimuli. Also in
the passive part 3,000 artifact-free sweeps were collected.
EEG Recording and Data Evaluation
The EEG was recorded between Cz and right ~contralateral! as
well as left ~ipsilateral! mastoid; the forehead served as ground.
Electrode resistance was kept below 2 kV. The subjects were asked
to relax and to avoid any tension in the head muscles. The pre-
amplifier gain was adjusted individually between 392,000 and
758,000 to make best use of the full range of the preamplifiers. The
EEG was analog-filtered by high-pass ~100 Hz! and low-pass
~600 Hz! 10-pole Butterworth filters, and sampled with 250 kHz
per channel ~i.e., sampling interval of 4 ms; further processing of
the data was conducted with the same resolution!. A total of 3,000
artifact-free sweeps were collected and averaged for the nontargets
in the two ~contra- and ipsilateral! channels; artifacts ~trials that
exceeded the preamplifier range! were discarded online. The re-
cording window had a length of 55 ms beginning 8 ms before
~acoustic! stimulus onset. The amplitude of the averaged FFP was
estimated by its spectral power, which was calculated using the
DADiSP software package. The latency of the FFP was estimated
in the time domain by the latency of the maximum of the cross-
correlation function of the response with the acoustic stimulus
~using BMDP2T; for further details see Hoormann et al., 1992a!.
Both were fed through the same set of analog filters, hence group
delay and phase shift were the same for both. An artificial ear
~Brel & Kjaer 4153, Naerum, Denmark! and an impulse sound
level meter ~Brel & Kjaer 2209! were used to measure the sound
pressure level and the reference signal for the cross-correlation
with the stimulus. Within a multivariate design, the dependent
variables amplitude and latency and the independent within-factors
recording channel ~contralateral, ipsilateral! and attention ~audito-
ry, visual, passive! were analyzed by a two-way multivariate analy-
sis of variance ~MANOVA!. The degrees of freedom for the
independent factor attention with three levels were adjusted after
Geisser and Greenhouse ~1958!.
J. Hoormann et al.
Results and Discussion
A total of 81% of the auditory targets and 78% of the visual targets
were identified correctly. Although the difference between these
levels was not significant, the difference between correct auditory
~415 ms! and visual ~462 ms! reaction times ~RTs! was significant,
F~1,19! 5 23.84, p , .0001. The mean FFP amplitude was 380 nV.
The FFP was larger with contralateral ~410 nV! than with ipsilat-
eral reference ~351 nV!, F~1,19! 5 20.35, p , .0005. No further
amplitude effects were found, F , 1. The mean FFP latency was
6.34 ms. No significant effects at all were found for the FFP
latency. An explanation for the absence of any attention effect on
the FFP may be the high intensity of the stimuli, because the
olivocochlear bundle exerts efferent control mainly for low-intensity
stimuli ~e.g., Siegel and Kim, 1982!. Hence softer stimuli were
used in the subsequent experiments. A further reason may be that
attention cannot be focused strongly enough for an entire block of
32 s ~in our case! to influence early auditory processing. Hence in
a subsequent experiment the focusing of attention was directed to
short time periods only. Furthermore, the somatosensory modality
was included as an alternative distractor modality, in addition to
the visual modality.
EXPERIMENT 2
In this experiment stimulus pairs were presented to the same or to
different modalities. The S1-S2-interval ~SSI! was short, whereas
the interpair0intertrial interval ~ITI! was long. The first stimulus
specified the to-be-attended modality: when the second stimulus
was presented to the same modality as the first stimulus, it was
attended, otherwise it was unattended. Hence, subjects had to focus
attention during the SSI, whereas they could relax during the ITI.
Methods
Subjects
Twelve healthy naive subjects ~6 men, 6 women, mean age 5 24
years! participated. All subjects had normal ~or corrected-to-
normal! vision and hearing thresholds ~see Experiment 1!. The
subjects were paid 15 DM per hour for participating.
Stimuli and Procedure
Several short sessions were administered to induce a high perfor-
mance level and to obtain a sufficient number of sweeps per con-
dition. All subjects participated in six sessions, each on a separate
day. In three of the sessions, auditory and visual stimuli were
presented, in the three remaining sessions auditory and somato-
sensory stimuli were presented. The order of sessions was balanced
across subjects. The auditory stimuli were presented monaurally
via the delay line, the visual stimuli via a duo-LED, as described
in Experiment 1. The nontargets and targets had the same charac-
teristics as in Experiment 1. The somatosensory stimuli were ap-
plied to the skin of the left lower leg ~about 20 cm above the outer
ankle! by a mechanical stimulation device, which avoided electro-
magnetic artifacts in the chamber. The stimulation device con-
sisted of a small plastic frame fastened to the leg in which two
movable pins are mounted at a distance of 4 cm. Both pins could
be moved ~about 2-mm extension! independently by two 5-m long
cables in a tube and were controlled from outside the chamber.
Two possible stimuli were applied by this device: a nontarget was
a short ~500 ms long! touch of one pin, a target was a synchronous
touch of both pins. The pitch difference ~auditory stimuli!, the hue
Early attention effects and evoked potentials
difference ~visual stimuli!, and the touch difference ~somatosen-
sory stimuli! between targets and nontargets were determined in
pilot studies to establish a roughly 75% probability of correct
target identification. The intensity of the auditory stimuli was set to
65 dB HL, which was much less than in Experiment 1. As in
Experiment 1, targets occurred with 5% probability in the attended
modality. The stimuli were presented as pairs ~S1 and S2! with an
ISI of 600 ms between S1 and S2, and an ITI of 2,400 ms between
the pairs. The first stimulus in a pair specified the to-be-attended
modality, the second stimulus was presented to the to-be-attended
or to the to-be-unattended modality with equal probability. As in
Experiment 1, the subjects had to press a key after a target.
FFPs were recorded to all auditory stimuli, which could be cue,
attended auditory stimulus ~with a preceding auditory cue!, or
unattended auditory stimulus ~with a preceding visual or somato-
sensory cue!. As in Experiment 1, both contra- and ipsilateral
references were used. In each condition 3,000 artifact-free sweeps
were averaged and evaluated as described above. The maximum of
the cross-correlation function was evaluated in addition as a mea-
sure for the quality of the averaged FFP. Long-latency auditory as
well as visual and somatosensory evoked potentials ~EPs! were
computed in addition. The late auditory EPs were not further eval-
uated, because the factor attention is confounded with the ISI,
which is known to influence the amplitude of the N1 ~e.g., Davis,
Mast, Yoshie, & Zerlin, 1966!: for attended tones ~auditory cue
preceding auditory stimulus! the time interval between successive
auditory stimuli is short ~600 ms!, which is likely to suppress N1
amplitude, whereas for nonattended tones ~nonauditory cue pre-
ceding auditory stimulus! it is long ~more than 3,000 ms!, which is
likely to enhance N1 amplitude. Hence the ISI effect on N1 am-
plitude would counteract and obscure a possible attention effect.
The dependent variables FFP amplitude and latency were analyzed
by a three-way repeated-measures MANOVA with the independent
within-factors recording channel ~contralateral, ipsilateral!, con-
current modality ~visual, somatosensory!, and condition ~cue, au-
ditory attention, auditory nonattention!.
Results and Discussion
The maximum value of the cross-correlation function was higher
for the contralateral ~0.80! than for the ipsilateral recordings ~0.73!,
which shows the better quality of the contralateral recordings. Also
the FFP appeared to be larger with contralateral ~275 nV! than
with ipsilateral reference ~239 nV!; however, this difference was
not significant. No significant result was obtained for attention,
though there was an FFP latency difference in the expected direc-
tion ~attended 6.54 ms, nonattended 6.58 ms!.
In sum, there was no significant attention effect on any FFP
parameter. Hence, even if attention is strongly focused to a par-
ticular modality over a split-second period, there are no significant
attention effects on FFP latency or amplitude. The possibility that
the different ISIs between subsequent attended and nonattended
stimuli ~see above! have influenced the FFP result is very unlikely,
because the FFP is invariant to different ISIs down to 100 ms ~e.g.,
Glaser, Suter, Dasheiff, & Goldberg, 1976!. A further reason may
be the switching of attention across modalities, which has to be
expected from trial to trial, and which has to be done every other
trial on average. After 600 ms, ISI attention can be expected to be
focused on the to-be-attended modality; however, a cost of switch-
ing between modalities cannot be excluded. This may have led to
a relative dilution of attention, which may have cancelled the
expected effect due to focusing of attention to a narrow time segment.
35
Altogether the results of the cross-modal Experiments 1 and 2
revealed no significant effects of focusing of attention on the am-
plitude or latency of the FFP. In the subsequent experiments we
investigated intramodal attention effects with auditory stimuli only.
EXPERIMENT 3
In this first intramodal experiment, we applied an equivalent of the
classical Hillyard paradigm ~Hillyard et al., 1973!, that is, auditory
stimuli were presented at a fast rate to the two ears with different
frequencies; one ear had to be attended to. A crucial point in the
Hillyard paradigm is a large separation of the attended and non-
attended channels, and a difficult targetnontarget discrimination.
Hence we used both ear position and frequency as channels, and
made the discriminability of targets from nontargets difficult. Be-
cause Galbraith and Doan ~1995! found positive results mainly
with missing fundamental stimuli, and we also intended to repli-
cate parts of that study, we did not only apply pure tones as stimuli,
but also missing fundamental stimuli. Finally, to control the effec-
tiveness of the attention procedure, middle- and long-latency
auditory-evoked potentials ~AEPs! were recorded in addition to
the FFP.
Methods
Subjects
Twelve healthy female subjects with a mean age of 24 years par-
ticipated in this experiment. One subject had to be excluded, be-
cause she did not meet the hearing threshold criteria ~see Experiment
1!; one further subject had no measurable FFP. The remaining 10
subjects had a mean age of 23 years.
Stimuli and Procedure
All subjects participated in three sessions, each on a separate day.
On the first day, the audiometry was conducted, and the subjects
practiced the main task. Particularly they were trained to direct
attention exclusively to one ear and switch attention to the other
ear after a cue. On the next 2 days, pure tone stimuli and missing
fundamental stimuli were presented. The order of the tone and
missing fundamental days was balanced across subjects. The au-
ditory stimuli ~circa 30 ms duration! were presented to both ears
via two independent auditory delay lines ~cf. Experiments 1 and 2!.
For the pure tone stimulation, 240-Hz tones ~nontargets! or 200-Hz
tones ~targets! were presented to the right ear, and 380-Hz tones
~nontargets! or 340-Hz tones ~targets! to the left ear. These fre-
quencies were chosen so as to yield maximum channel separation
~maximum frequency difference; Hillyard et al., 1973! within the
most sensitive frequency range of the FFP ~Hoormann et al., 1992a!.
The nontargets were presented in random sequence to the ears; in
the attended ear, about 8% of the nontargets were replaced ran-
domly by targets. In this experiment no targets were presented to
the nonattended ear to avoid ~automatic! distraction of attention to
the nonattended ear. Indeed, the optimal focusing of attention seemed
so important to us that we accepted the asymmetry of the exper-
imental design with respect to the distribution of targets between
the two ears. The ISI was randomized between 200 and 400 ms.
The intensity of the 380-Hz tone was 65 dB SL; the intensity of the
240-Hz tone was adjusted individually until it was perceived with
the same loudness as the 380-Hz tone. The stimulus trains were
divided into blocks of 400 stimuli. Each block was started by the
subjects with a key press, which was followed by a cue ~a 1-kHz,
50-dB SPL tone of 2-s length! that was given to the ear that had to
36
be attended in the following block. The order of right0left cues0
blocks was regular: Four right cues0blocks were followed by four
left cues0blocks; this sequence was repeated 300 ms after the cue
the stimulus train began. Again the subjects had to press a key after
each target in the attended ear. The first three sweeps of each block
were excluded from averaging.
For the missing fundamental stimulation, complex stimuli were
produced by superpositioning third, fourth, and fifth harmonics of
fundamentals, which correspond to the pure tones used in the pure
tone stimulation. Hence the right ear was stimulated with a com-
plex of 720, 960, and 1200 Hz ~nontargets! or 600, 800, and
1000 Hz ~targets!, the left ear with a complex of 1140, 1520, and
1900 Hz ~nontargets! or 1020, 1360, and 1700 Hz ~targets!. In all
other respects the stimulation procedure was the same as for the
pure tones.
Because we found no attention specific differences for FFPs
after ipsi- versus contralateral stimulation in Experiments 1 and 2,
we used a vertical and horizontal derivation as proposed by attended stimuli for two subjects.
Galbraith ~1994! for better distinction of the different sources of
tone and missing fundamental FFPs. The vertical derivation was
Cz versus left mastoid and the horizontal derivation was right
versus left mastoid. The FFPs were recorded to all nontarget stim-
uli, which could be attended or unattended, and evaluated as de-
scribed above; 2,500 artifact-free sweeps were averaged separately
for each condition.
In addition to the FFPs, middle-latency and late-evoked poten-
tials were recorded from the three midline electrodes, Fz, Cz, and
Pz, with the left mastoid as reference. The middle-latency poten-
tials were low-pass filtered with 250 Hz and sampled with 800 Hz,
the late potentials were low-pass filtered with 60 Hz and sampled
with 200 Hz. The group delays introduced by the low pass filters
were calculated and subtracted from the raw latency values. Again,
2,500 sweeps were collected per condition; electrooculogram ~EOG!
and slow artifacts were eliminated offline, which resulted in about
2,100 artifact-free sweeps per condition. The averages were
smoothed digitally ~Ruchkin & Glaser, 1978! below 117 Hz ~for
middle-latency potentials! and below 17.6 Hz ~for late potentials!,
and then corrected for EOG artifacts ~Verleger, Gasser, & Mcks,
1982!. The middle-latency potentials revealed three deflections in
the latency range between 10 and 50 ms after stimulus onset, called
P11, N22, and P32. In the late range, the N100 was detected as the
largest relative minimum in the window 80160 ms after stimulus
onset. All deflections were measured relative to electrical zero
~i.e., zero voltage at the differential amplifiers!; for the middle-
latency potentials the mean amplitude in the window 1550 ms
poststimulus was computed in addition ~see below!.
The FFP could not be measured reliably in the horizontal chan-
nel for the 380-Hz missing fundamental stimulus. Hence, it was mea-
sured in the vertical channel for both frequencies and its amplitude
and latency ~dependent variables! analysed ~separately for pure tones
and missing fundamental stimuli! by a two-way repeated-measures
MANOVA with the dependent within-factors frequency0ear ~2400
right, 3800left!, and attention ~attended, nonattended!. In addition,
the 240-Hz data were evaluated with the design channel ~vertical,
horizontal! and attention ~attended, nonattended!. For the middle-
latency potentials only the data for the tones were evaluated ~be-
cause the missing fundamental data showed no clear peaks! with the
design electrode ~Fz, Cz, Pz!, frequency, and attention. The ampli-
tude and latency of the N100 were evaluated separately for tones
and missing fundamental stimuli with the design electrode ~Fz, Cz,
Pz!, frequency, and attention. Where appropriate the degrees of
freedom were adjusted after Geisser and Greenhouse ~1958!.
J. Hoormann et al.
Results and Discussion
The RTs for pure tones were 459 ms ~right ear stimulation! and
420 ms ~left ear!, and for missing fundamental stimuli 407 ms ~right
ear! and 391 ms ~left ear!. The RT differences between right and left
were significant, F~1,9! 5 31.65, p , .0005, as were the differ-
ences between the RTs after tones and missing fundamental stimuli,
F~1,9! 5 18.58, p , .005; the interaction was not significant. A to-
tal of 8.4% ~right ear! and 5.0% ~left ear! of the tone targets were
missed, and 6.1% ~right ear! and 7.7% ~left ear! of the missing
fundamental targets. The main effects were not significant, F , 1,
although the interaction was significant, F~1,9! 5 5.74, p , .05!.
The FFP was clearly larger in the vertical than in the horizontal
channel for the 240-Hz stimuli; no other significant amplitude or
latency effects were found. For the vertical FFP ~including both
stimulus frequencies! no significant amplitude or latency effect
was found. Figure 2 shows the FFPs after attended and non-
The N100 following tone stimuli was larger at Cz than at the
two other electrodes, F~2,18! 5 6.29, p , .05, E 5 0.72, which is
the usual finding. A similar, though weaker, effect was found after
missing fundamental stimuli. More important, the N100 after tone
stimuli was larger for attended ~21.7 mV! than for unattended
stimuli ~21.0 mV!, F~1,9! 5 16.41, p , .005 ~Figure 3!. A similar
effect was found for missing fundamental stimuli: the N100 was
larger for attended ~22.4 mV! than for unattended stimuli
~21.3 mV!, F~1,9! 5 27.83, p , .001. As also seen in Figure 3, not
only the N100, but also the subsequent ERP segment appears to be
shifted in the negative direction for attended versus nonattended
stimuli.
For the middle-latency potentials ~Figure 4! no significant la-
tency effects were found. For the amplitudes the P11 and P32
showed no significant attention effects, though the P32 was nu-
merically less positive after attended stimuli, as also seen in Fig-
ure 4. The N22 was more negative after attended than after
nonattended stimuli, F~1,9! 5 5.64, p , .05. As also seen in
Figure 4, not only the N22, but a broader time window between 15
and 50 ms ~averaged across stimuli! appears to be shifted in the
negative direction for attended compared with nonattended stimuli.
However, this effect failed to reach significance, p 5 .14.
These results demonstrate the effectiveness of the attention
manipulation, as revealed in the amplitude effects on N100 and
N22. The latter effect shows a negative displacement of the N22
after attended versus unattended stimuli, which is similar to the
findings of McCallum et al. ~1983!. On the other hand, the present
data are different from the results of Woldorff et al. ~1987!, be-
cause we could not detect any positive shift such as the P20-50.
This result may be due to task differences, that is, much lower
stimulus frequencies and larger ISIs in the present task versus that
of Woldorff et al. . In any case, the present results show coherent
effects, namely negative displacements for attended versus un-
attended stimuli in middle and late AEPs.
In contrast, no effect at all was seen for the FFP, that is, we could
not find larger FFP amplitudes after attended than after unattended
tone or MF stimuli, a finding that contradicts the results of Gal-
braith and Doan ~1995!. The negative result is not likely due to the
fact that discriminability was lower in our experiment compared with
Galbraith and Doans; attention should be focused even more ef-
fectively when the discriminability of targets and nontargets is lower.
Also, we could show a strong modulation of the N100 with atten-
tion, which proves the effectiveness of the attention manipulation.
However, the discrepancy may be due to the use of a frequency dif-
Early attention effects and evoked potentials 37

Figure 2. Examples of two typical individual frequency-following potentials ~FFPs! after attended and nonattended stimuli ~upper part
subject EZ, lower part subject BV!. Left panels: FFPs after 380 Hz stimulus, right panels: FFPs after 240 Hz stimulus. att 5 attended
ear; non-att 5 nonattended ear; S 5 stimulus onset.

ference in the present study instead of duration0intensity differ-
ences in the study of Galbraith and Doan ~1995!, because early
attention mechanisms may depend on the nature of the task required
~frequency vs. intensity vs. duration discrimination!. Such possible
mechanisms are discussed below ~Experiment 4!.
Hence, it appears that a classical unimodal condition with sus-
tained attention to one ear, which does produce the well-known
attention effects on the N100, does not affect very early process-
ing, as reflected in the FFP. This finding may be due to the fact that
a sustained allocation of attention allows only for suboptimal fo-
cussing of attention. In our next experiment, a unimodal monaural
paradigm was administered, in which attention was highly focused
over very short intervals.
EXPERIMENT 4
To focus attention over short time intervals, a double stimulus
paradigm was applied. In contrast to the cross-modal study with
stimulus pairs ~Experiment 2!, the first stimulus was not a cue, but
served as the reference for the second stimulus. The interval be-
38 J. Hoormann et al.

Figure 3. Grand means ~n 5 10 subjects; Experiment 3! for the late auditory-evoked potentials ~up to 200 ms after stimulus onset!
after 240-Hz tones ~upper panel! and after 380-Hz tones ~lower panel! for attended ~ATT; heavy lines! and unattended stimuli ~NATT;
thin lines!. The later part of the event-related potential ~ERP! is more negative after attended than after nonattended stimuli.

tween first and second stimulus was chosen to be very short, which presented with a short time interval under passive conditions, a
should permit a very effective focusing of attention restricted to suppression of the amplitude of a middle-latency auditory poten-
this period ~Ntnen, 1990!. When auditory stimulus pairs are tial, the P50, is usually found for the second compared with the

Figure 4. Grand means ~n 5 10 subjects; Experiment 3! for the middle-latency auditory-evoked potentials ~up to 70 ms after stimulus
onset! after 240-Hz tones ~upper panel! and after 380-Hz tones ~lower panel! for attended ~heavy lines! and unattended stimuli ~thin
lines!. The event-related potential ~ERP! is more negative after attended than after nonattended stimuli in the time segment 1550 ms
~averaged across stimuli!.
Early attention effects and evoked potentials 39

first stimulus ~Freedman, Adler, Waldo, Patchman, & Franks, Results and Discussion
1983; Waldo & Freedman, 1986!. Guterman, Josiassen, and
Bashore ~1992! could show that this P50 suppression was not No significant effects at all were found for the FFP amplitude.
evident when subjects attended to the second stimulus to detect ~The amplitude of the FFP was numerically larger for the contra-
a target. Guterman et al. concluded that this cancellation of the lateral than for ipsilateral channel, however, the difference did not
P50 suppression reflects an active attention control process, which reach the significance level.! The maxima of the cross-correlation
already influences rather early auditory processing. Hence, we functions between stimuli and responses ~that were calculated to
expected to have a chance to find even earlier attention effects determine the latencies of the FFPs! were between 0.79 ~contra-
with this paradigm. lateral! and 0.74 ~ipsilateral!. Of course, these values are highly
significant and demonstrate the high quality of the latency estima-
tions. No significant main effects and interactions of the factor
Methods recording channel were found for the FFP latency. The results
for the other two factors on FFP latency are summarized in Fig-
Subjects ure 5. The factor condition revealed a strong tendency, F~1,10! 5
Twelve healthy young subjects ~5 men, 7 women! with normal 4.71, p 5 .055, for a shorter FFP latency in the attention than in the
auditory thresholds participated. One female subject had to be ignore condition. The factor stimulus was highly significant,
excluded because no FFP could be measured. The remaining 11 F~1,10! 5 17.68, p , .005. Most importantly, the condition 3
subjects had a mean age of 21 years ~range, 1829 years!. stimulus interaction was also significant, F~1,10! 5 5.34, p , .05.
The latter result allowed the computation of simple effects for both
Stimuli and Procedure attention conditions. This analysis revealed that the FFP latency
The stimuli were low-frequency tone bursts ~c. 30 ms duration, 85 was highly significantly shorter for the second stimulus ~6.32 6
dB SPL! presented to the left ear via the auditory delay line ~see 0.061 ms @mean 6 SEM #! than for the first stimulus ~6.37 6
above!; the right ear was masked with white noise of 65 dB SPL. 0.063 ms! in the attention condition, F~1,10! 5 13.54, p , .005,
The duration and envelope of the stimuli was as described for the whereas there was no latency difference at all between the two
earlier experiments. The stimuli were presented in pairs with a stimuli in the ignore condition ~S1: 6.37 6 0.063 ms, S2: 6.37 6
very short S1-S2 interval ~300 ms!, and an interpair interval of 0.066 ms!. The latency reduction is quantitatively small. However,
about 1,850 ms ~randomized between 1,300 and 2,400 ms!. The the reliability of this effect is not only revealed by the clear AN-
first stimulus of each pair had a frequency of 340 Hz and served as OVA results, but also is underlined by the fact that in 22 compar-
the reference for the second stimulus, which could be either iden- isons ~11 subjects, 2 sides! only a single latency showed a slight
tical with the first stimulus ~nontarget!, or a 358-Hz stimulus ~tar- increase.
get!. Targets occurred on between 0.5% and 5% of the presentations The fact that in the ignore condition no latency difference was
in different blocks ~see below!. The pitch differences between seen between the FFPs to the first and the second stimulus shows
targets and nontargets were chosen in pilot studies to establish a
roughly 75% probability of correct target identification. ~The crit-
ical pitch difference was relatively small because of the immediate
succession of the to-be-discriminated stimuli.! The task of the
subjects was to count the targets covertly and report the result at
the end of a block, which consisted of 150 stimulus pairs. After
each block, feedback was given as to the quality of the perfor-
mance by giving the actual number of targets in the past block. The
average block length was 5 min. These blocks were repeated until
a sufficient number of artifact-free sweeps were collected for each
of the 340-Hz stimuli ~see below!; for this task about 30 blocks
were necessary on average. Because the presentation of pairs may
have an automatic suppressive effect on the FFP to the second
stimulus ~as found for the P50!, we established a control condition,
in which the same stimulus pairs were presented, while the sub-
jects had to ignore the stimuli and relax. In the control condition no
targets were presented. Again, blocks of 150 pairs were given until
the same number of artifact-free sweeps was collected as in the
attention condition.
FFPs were recorded to all 340-Hz stimuli, that is, reference
stimuli and nontargets in both attention and ignore conditions. The
FFP was measured at Cz relative to the contralateral and the ipsi-
lateral mastoid. A total of 3,000 artifact-free sweeps were averaged
per condition. The FFP amplitude and latency were evaluated as
described above.
The amplitude and latency ~dependent variables! of the FFP Figure 5. Mean latencies ~n 5 11 subjects; Experiment 4! for the frequency-
was analyzed by a three-way repeated-measures MANOVA with following potentials ~FFPs! after the first and the second stimulus of the
the independent within-factors recording channel ~contralateral, paired stimuli in the attention and the ignore condition. The FFP latency is
ipsilateral!, stimulus ~first, second!, and condition ~attend, significantly shorter after the second stimulus in the attention, but not in the
ignore!. ignore condition.
40
that automatic facilitation cannot be the reason for the reduced
latency in the attention condition. Also the reduced arousal in the
ignore condition ~indeed, some of the subjects were actually asleep
in this condition! is unlikely to be the reason for the latency re-
duction in the attention condition, because the FFP latency was
exactly the same for the first stimulus in both the attention and the
ignore condition. Any effect due to a difference in arousal should
have influenced the responses after the first stimulus as well as
after the second stimulus, which was not the case. It seems also
unlikely that general arousal has changed between the first and the
second stimulus ~i.e., within 300 ms in the attention condition! for
several thousand times in a row. Hence, these results suggest that
in the unimodal monaural condition the processing of the second
auditory stimuli in pairs that were highly attended, were facilitated
on the level of the lower brain stem or even at a more peripheral
level.
The effect is, however, numerically very small, and clearly
irrelevant for reaction tasks. It may be speculated what kind of
mechanism could be affected by this small peripheral effect. One
possibility is that the effect supports the auditory localization of
low-frequency stimuli in the horizontal plane ~lateralization!, be-
cause latency differences in the range of 50 ms lead to detectable
lateralization changes. Moreover, Hall ~1979! assumed that FFPs
reflect neural information relevant for lateralization. Hence it may
be possible that our FFP latency result reflects the action of a
neural mechanism that underlies the sharpening of spatial atten-
tion. A further possibility is that the effect reflects a sharpening of
pitch discrimination, because highly accurate pitch discrimination
is in fact required to cope with the task. This idea is supported by
the fact that the auditory efferent fibers, particularly the olivoco-
chlear bundle, support a sharpening of the frequency tuning in the
cochlea by modulating the active micromechanical properties of
the outer hair cells ~Mountain, 1980; Scharf et al., 1987; Siegel &
Kim, 1982!.
GENERAL DISCUSSION AND CONCLUSION
In none of the four experiments reported we could find any effect
of attention on FFP amplitude. This finding is in line with the
negative findings of Hillyard and Picton ~1979! and Galbraith and
Kane ~1993!, but disagrees with some positive results of Galbraith
and Arroyo ~1993!1 and Galbraith and Doan ~1995!. In a quasi-
replication of the experiment by Galbraith and Doan ~1995!, we
could demonstrate the effectiveness of our attention manipulation
by effects on late ~N100! and middle-latency AEPs ~N22!, whereas
no attention effect on the FFP was found. Hence we conclude that
with cross-modal and with intramodal paradigms there is no effect
on FFP amplitude, even when attention is focused strongly to
narrow time segments.
As to FFP latency, there were no significant attention effects in
cross-modal sustained and cued attention paradigms ~Experiments
1 and 2!. Hence, it may be concluded tentatively that peripheral
gating as reflected in the FFP is not relevant or possible for inter-
modal attention mechanisms, which facilitate or attenuate an entire
modality.
As to the intramodal experiments ~Experiments 3 and 4!, no
early attention effect was seen for a sustained attention paradigm
1
Contrary to these authors we could not find any differential attention
effect for the first vs. second half of the FFP. The reason for this discrep-
ancy is not clear to us, but it is a fact that the signal-to-noise ratio of the
FFPs was much higher in our experiment.
J. Hoormann et al.
~Experiment 3!, although the task was difficult and the attention
manipulation effective, as seen in the N100 and N22 effects. Hence,
it may be concluded that mechanisms supporting sustained atten-
tion to one ear0frequency are likewise not influenced by peripheral
gating, as reflected in the FFP. Only when attention was directed
exclusively to one ear, and focused to short time segments to cope
with a difficult frequency discrimination task, could early attention
effects be demonstrated ~Experiment 4!.
We therefore conclude that early auditory attention effects are
evident mainly in unimodal situations with unilateral stimuli, when
attention is highly focused to a restricted time interval. Hence
peripheral gating does not support a general facilitation or inhibi-
tion of an entire stimulus modality, but serves subtle intramodal
mechanisms, which may be necessary for a better localization
and0or frequency discrimination of task-relevant stimuli. The pre-
cise origin of the observed FFP latency effect within the auditory
pathway is still unclear, and can be located, at present, only be-
tween the cochlea and the FFP source. Because we provided ev-
idence for the cochlear nucleus as the source of the FFP ~Hoormann
et al., 1992a!, the origin of the effect is probably the cochlea or the
cochlear nucleus. Recent studies of otoacoustic emissions suggest
control of outer hair cells via the olivocochlear bundle ~Giard
et al., 1994; Meric & Collet, 1992; Michie et al., 1996!. In partic-
ular, it has been demonstrated that activating the medial olivoco-
chlear system by contralateral acoustic stimulation contributes to
shortening otoacoustic emission latency ~Giraud, Perrin, Chry-
Croze, Chays, & Collet, 1996; Giraud et al., 1997!. This result may
be relevant to the FFP-latency shortening we observed. Hence, we
favor the view that the attention effect on FFPs has its origin in the
cochlea.
An issue that has to be discussed in more detail is whether
the double-stimulus paradigms ~Experiments 2 and 4! stimulate
an involuntary capture of attention ~exogenous orienting! rather
than a voluntary allocation of attention ~endogenous orienting!
~Spence & Driver, 1994! and whether the latency shortening of
the FFP in Experiment 4 is due to exogenous or endog-
enous orienting of attention or, alternatively, to general arousal
rather than to attention. For visual stimuli, Rafal, Henik, and
Smith ~1991! found that subcortical structures are only involved
in exogenous attention. Schrger and Eimer ~1996! showed
late negative modulations of the ERP ~Nd! after peripheral
stimuli presented at the same position as an immediately preced-
ing cue stimulus even when the cue was uninformative or mis-
leading ~invalid!. However, the Nd was seen only in an active
condition, but not in an ignore condition. This observation sug-
gests that exogenous attention is ~i! not fully automatic ~as
also claimed by Spence & Driver, 1994!, and ~ii! occurs in
paradigms with paired auditory stimuli ~cue and imperative stim-
ulus!, as applied in our study. Taken together, the results of the
two cited studies support our view that the early attention ef-
fects we found in Experiment 4 may be mainly due to exog-
enous attention. However, because of the difficulty of the task
in our paradigm, a substantial contribution of endogenous at-
tention, that is, a voluntary sustained attention to the left ear and
additional allocation of attention after S1, seems possible. Con-
cerning the possible contribution of arousal, we cannot defi-
nitely exclude some influence on the observed latency effect.
However, two findings seem to contradict this interpretation: ~i!
Only in the attention condition of Experiment 4 was the FFP-
latency after the second stimulus shorter in comparison with all
other conditions. The latency after the first stimulus was virtu-
ally the same in the control and the attention condition. If gen-
Early attention effects and evoked potentials
eral arousal indeed played a role, these results meant, as discussed
above, that, on the one hand, arousal was the same in both
conditions ~sleep and attention!, but that, on the other hand, To us this absence suggests that the latency reduction of our
arousal changed within the SOA of 300 ms for 3,000 times in a
row. ~ii! Possibly even more relevant may be the result of our
Experiment 2, which was a cross-modal task with stimulus pairs.
Indeed, in a similar cross-modal task, Low, Larson, Burke, and
Hackley ~1996! demonstrated an immediate arousal effect ~alert-
ing! for the R50 component ~50 ms after the evoking stimulus!
of the eyeblink reflex, when the evoking visual stimulus was
preceded by an accessory tone by only 40 ms. Therefore, one
should expect an immediate arousal effect in our Experiment 2
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~Received July 23, 1998; Accepted March 9, 1999!