Neuropsychologia 50 (2012) 11781189

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Neuropsychologia
journal homepage: www.elsevier.com/locate/neuropsychologia

How does phasic alerting improve performance in patients with unilateral

neglect? A systematic analysis of attentional processing capacity
and spatial weighting mechanisms
Kathrin Finke a, , Ellen Matthias a , Ingo Keller b , Hermann J. Mller a ,
Werner X. Schneider c , Peter Bublak d
a
General and Experimental Psychology/Neuro-Cognitive Psychology, Ludwig Maximilian University, Munich, Germany
b
Schoen Clinic Bad Aibling, Bad Aibling, Germany
c
Neuro-Cognitive Psychology, University of Bielefeld, Bielefeld, Germany
d
Neuropsychology Unit, Hans Berger Department of Neurology, University Clinic, Jena, Germany

a r t i c l e i n f o a b s t r a c t

Article history: In visual hemi-neglect, non-spatial decits such as reduced intrinsic alertness can signicantly modulate
Received 10 January 2012 the degree of left visual eld inattention. However, to date, the precise mechanisms mediating this effect
Received in revised form 6 February 2012 are hardly understood. In the present study, we assessed the inuence of increased alertness on both gen-
Accepted 15 February 2012
eral attentional capacity (perceptual processing speed) and spatial attentional selection processes (spatial
Available online 23 February 2012
distribution of attentional weighting). For this purpose, a whole-report paradigm based on Bundesens
theory of visual attention (TVA) was combined with a non-spatial, visual alerting cue. Three different
Keywords:
cue-target stimulus onset asynchronies (SOAs; of 80, 200, and 650 ms), allowed us to observe the time
Attention
Psychophysics course of the alerting-cue effects. A group of six patients with visual hemi-neglect was examined and
Intrinsic alertness their performance compared with six healthy control subjects matched for age, gender, and education.
Phasic alertness In neglect patients, the alerting cue evoked a phasic increase of perceptual processing speed. However,
Thalamus this effect was mainly found in the ipsilateral, i.e. in the preserved hemield. Importantly, however,
patients displayed a fast-evolving and short-lasting, phasic modulation of spatial attentional weighting,
with a re-distribution of attentional weights from the pathological rightward bias to a normal, more bal-
anced distribution of visual attention. In control participants, the cueing effects on perceptual processing
speed and spatial weighting were generally less pronounced than in neglect patients. Replicating results
of a prior study, cueing induced a stable, slightly leftward, distribution of attentional weights, whilst in
the no-cue condition, a temporary rightward shift of attentional weights was found.
This pattern of effects suggests a close interaction between alertness and spatial-attentional weighting
in the syndrome of visual hemi-neglect. It supports the hypothesis that the manifestation of spatial neglect
involves at least in part intrinsic alertness decits. It also provides clues to a more detailed account of
the mechanisms responsible for alleviating neglect in patients following manipulations of the alertness
level, both in the short (cueing) and in the long term (alertness training).
2012 Elsevier Ltd. All rights reserved.

1. Introduction
It is now well established that both spatially lateralized and
non-lateralized decits of attention contribute to the clinical pic-
ture of left-sided visual hemi-neglect (Husain & Rorden, 2003;
Samuelsson, Hjelmquist, Jensen, Ekholm, & Blomstrand, 1998; van
Kessel, van Nes, Brouwer, Geurts, & Fasotti, 2010). Although, at
the behavioural level, a spatially lateralized preference for stim-
uli and objects on the right side of space combined with a lack
Corresponding author. Tel.: +49 0 89 218072520.
E-mail address: nke@psy.lmu.de (K. Finke).
of awareness for those in the left hemi-eld is the most promi-
nent feature of neglect (Bartolomeo & Chokron, 2002; Bisiach
& Vallar, 1988; Heilman, Watson, & Valenstein, 2003; Karnath,
1988), patients also show non-spatial decits such as, for exam-
ple, a prolonged attentional blink (Husain, Shapiro, Martin, &
Kennard, 1997), impaired working memory (Danckert & Ferber,
2006; Wojciulik et al., 2001), and decient vigilance (Hjaltason,
Tegnr, Tham, Levander, & Ericson, 1996; Robertson et al., 1997).
In fact, these non-spatial symptoms may even be more important
for the severity and persistence of hemi-neglect than the spatial
bias itself (Husain & Rorden, 2003). They may result from reduced
alertness, a foundational form of attention or processing capacity,
which more complex cognitive functions draw from (Raz & Buhle,
2006).

0028-3932/$ see front matter 2012 Elsevier Ltd. All rights reserved.
doi:10.1016/j.neuropsychologia.2012.02.008
 K. Finke et al. / Neuropsychologia 50 (2012) 11781189
Lowered alertness, i.e. an impeded ability to achieve and main-
tain a state of high sensitivity to an impending stimulus (Posner,
2008; Raz & Buhle, 2006), is a common observation in neglect
patients (Heilman et al., 1978). Whilst tonic alertness refers to
the intrinsic and more long-term control of the arousal level inde-
pendent from external warning cues, phasic alertness denotes the
brief adaptive increase of the arousal level pending on an upcom-
ing warning stimulus (Posner, 2008; Raz & Buhle, 2006; Sturm
& Willmes, 2001). Functional brain imaging studies have shown
a substantial overlap between the cerebral networks underlying
tonic and phasic alertness on the one hand, and those engaged in
governing spatial attention on the other, with the right inferior pari-
etal cortex, the most frequent lesion site in neglect patients, being
a key node in both (Coull, Frackowiak, & Frith, 1998; Kinomura,
Larsson, Gulyas, & Roland, 1996; Robertson, Mattingley, Rorden, &
Driver, 1998; Sturm & Willmes, 2001; Sturm et al., 1999, 2006).
In congruence with these overlapping brain networks of attention,
less alert neglect patients have been shown to have greater leftward
inattention (Bartolomeo & Chokron, 2002; Funk, Finke, Mller, Utz,
& Kerkhoff, 2010; Heilman et al., 2003; von Cramon & Kerkhoff,
1993). The alertness and spatial attention networks represent dis-
sociable brain systems (Husain & Rorden, 2003), however, and the
current knowledge about the nature and dynamics of their interac-
tion is only sparse. The present study is aimed at investigating this
question.
Robertson et al. (1998) were the rst to directly examine in
neglect patients, on a trial-by-trial basis, the causal relationship
between alertness changes induced by sudden onset tones and
modications of the rightward spatial bias. The authors found that
the pathological asymmetry of spatial attentional selection indeed
was alleviated by such a phasic enhancement of alertness. The alert-
ing cues temporarily reduced the tendency of neglect patients to
report the rightmost of two bars as coming rst in a temporal order
judgement task. This result demonstrates, that even the severely
damaged attentional system of neglect patients bears the potential
to rapidly adapt in response to a warning cue so that their chronic
inattention to the left hemi-eld can be overcome, resulting in a
more balanced spatial lateralization.
However, the underlying mechanisms remain unclear. A closer
understanding of the nature and time course of the effects of alert-
ness changes on non-spatial and spatial attention could, though,
provide important clues as to the cognitive architecture of attention
networks. For example, the exact relationship between phasic and
intrinsic alertness is still under debate (Posner, 2008; Raz & Buhle,
2006) and the question whether they are supported by the same
or by different systems remains unresolved. Furthermore, closer
knowledge about alertness effects could also have implications for
therapeutic intervention. Current treatment approaches suggest to
be based on the modulation of intrinsic alertness (Thimm, Fink,
Kust, Karbe, & Sturm, 2006) or both phasic and intrinsic aspects
(DeGutis & van Vleet, 2010), whilst clear evidence about the rele-
vant mechanisms mediating improvement in neglect is still lacking.
To investigate this issue, we used an approach based on a for-
mal theory of visual attention (TVA, Bundesen, 1990, 1998) that
allows simultaneous assessment of non-spatial and spatial aspects
of attention within the same subjects and with the same task. In
neglect patients, such a method had been applied for the rst time
by Duncan et al. (1999). These authors used whole and partial report
of briey presented letter arrays to obtain parametric estimates of
perceptual processing speed as a non-spatial component and lat-
eral bias as a spatial component of visual selective attention. They
found, conrming the presence of non-spatial attentional decits
in neglect, processing speed to be signicantly reduced in patients
within both visual hemi-elds.
In a recent study, Matthias et al. (2010) used an analo-
gous methodology to assess the nature and time course of the
1179
interaction between alertness, non-spatial and spatial attention
in healthy subjects. To that end, whole-report performance was
compared between two conditions: a no-cue condition, assumed
to reect possible changes in intrinsic alertness over the course of
a trial (Coull & Nobre, 1998; Posner, 1978; Sturm et al., 1999), and
an alerting-cue condition, assumed to reect modulations of phasic
alertness. In this way, it was possible to examine whether different
levels of alertness would affect non-spatial (i.e. processing speed)
and spatial attentional components differentially or, alternatively,
in a global manner. In addition, by introducing different cue-target
SOAs, it was also possible to map the time course of these effects,
so as to ascertain whether alertness-related effects on non-spatial
and spatial attention develop in a parallel or a sequential manner.
Matthias et al. (2010) found that an increase in the level of alert-
ness irrespective of whether this was phasically or intrinsically
induced rst enhanced general processing speed and was then
followed by a leftward shift of spatial attention. They interpreted
these results as indicating that intrinsic and phasic alertness effects
involve the same processing route, on which spatial and non-spatial
mechanisms of attention are mediated by independent processing
systems which are activated, as a result of enhanced alertness, in
temporal succession.
Based on these preceding results in healthy young subjects, we
applied the same procedure in neglect patients. Six patients with
left visual hemi-spatial neglect following right temporo-parietal
lesions and six healthy normal control subjects were assessed with
a whole-report task, in which they had to name stimuli presented
either unilaterally, on either side of the visual eld, or bilaterally, in
the left and the right hemi-eld. As in the prior study on normal sub-
jects (Matthias et al., 2010), the whole-report task was presented in
a no-cue and an alerting-cue condition. SOA variations were used
to map the time course of the effects of the alertness level on an
index of processing speed (i.e. sensory effectiveness) and spatial
attentional weighting. We expected to nd similar results in normal
control participants as in our preceding study and wanted to anal-
yse, which of the processes and mechanisms prevalent in normal
subjects would be altered or impaired in neglect patients.
Before presenting our method and the results, however, we pro-
vide, for those readers unfamiliar with TVA, a brief description of
its main ideas and concepts.
1.1. Theory of visual attention (TVA)
In TVA (for a detailed mathematical description see Bundesen,
1990, 1998; Duncan et al., 1999; Kyllingsbaek, 2006), visual objects
and their features are assumed to be processed in parallel and
compete for selection. The race amongst objects and features
is decided according to a speed criterion: features which enter
visual short term memory (VSTM) rst cause the complete encod-
ing of these objects into VSTM. Biases can arise if some objects
receive higher attentional weights than others due to either auto-
matic (bottom-up) or intentional (top-down) factors, conferring
a speed advantage in the race for selection. According to TVA, the
probability with which an object is selected (when VSTM is not yet
lled up to capacity) is thus determined by (i) the general process-
ing capacity available reected by an objects basic processing
rate (speed) and (ii) the attentional weight assigned to an object.
Independent quantitative estimates of these two atten-
tional components general processing capacity and attentional
weight are derived from the patients whole-report performance.
One aspect of weighting in TVA, which is especially relevant to
the present study, concerns the lateral distribution of attentional
weights within the visual eld. In TVA, the spatial distribution of
attention across the visual eld is indicated by a spatial laterality
index (parameter w ), relating the attentional weights for objects
in the left and the right visual hemi-eld (wleft and wright ). These
1180 K. Finke et al. / Neuropsychologia 50 (2012) 11781189

weights can be derived from performance accuracy in conditions Table 1

Demographic and clinical details for each participant. Abbreviations: Age, age in
in which patients have to report a stimulus that is accompanied
years; School, school education in years; Vision, visual eld impairment; TSI, time
by another stimulus in either the same or the opposite hemi- since injury in months; F, female; M, male; H (l), hemianopia in the left visual
eld (see Kyllingsbaek, 2006). In case of a rightward attentional hemi-eld; Q (ll), quadrantanopia in the lower left visual hemi-eld; SD, standard
bias, as in neglect, stimuli in the right hemi-eld receive a higher deviation.
attentional weight than competing stimuli in the left eld, so that Gender Age School Vision TSI
accuracy of report for the left hemi-eld will decline when stim-
Patient
uli are presented bilaterally (Duncan et al., 1999). Since such an
PB F 76 10 H (l) 2
unbalanced competition favouring rightward stimuli would have OB F 79 10 H (l) 2
no effect with unilateral, but a strong effect with bilateral target KKL M 65 13 Q (ll) 3
displays, the biased competition account provides a suitable expla- ML M 73 10 4
FP M 72 10 2
nation why contralesional extinction is a characteristic neglect
EW M 71 13 6
symptom (Desimone & Duncan, 1995). In extinction, a contrale-
sionally presented stimulus is detected or identied relatively well Mean 72.67 11.00 3.17
SD 4.76 1.55 1.60
when presented alone (i.e. without competing stimuli in the ipsile-
Control
sional eld), but the same stimulus is disregarded (extinguished) C1 F 71 10
or only poorly identied in the presence of simultaneously pre- C2 M 75 10
sented ipsilesional input (Bender, 1952). C3 M 71 13
C4 F 81 10
However, the probability with which an object is identied does
C5 F 74 10
not only depend on its attentional weight, but also on the sensory C6 M 69 13
effectiveness, which is inuenced by stimulus properties (such as
Mean 73.50 11.00
discriminability, luminance, contrast, retinal eccentricity) and by
SD 4.28 1.55
the observers rate of information uptake. In TVA, the parameter
for sensory effectiveness, si (which is assumed to be independent
of spatial attentional weighting), refers to the accuracy of reporting
Table 2
a single element presented alone, under conditions of no stimu-
Scores in the subtests and total score of the conventional part of the Behavioural
lus competition. Thus, s reects the total processing rate, rather Inattention Test (BIT, Wilson et al., 1987) for each neglect patient, together with cut-
than how capacity is divided between the various objects of a dis- off scores. Scores below the cut-offs are highlighted in bold. Abbreviations: Lines, line
play, providing a further TVA-based indicator of processing speed crossing; letters, letter cancellation, stars, star cancellation; copying, gure/shape
copying; bisection, line bisection; drawing, representational drawing; sum, sum of
(Duncan et al., 1999). Whilst parameter si can only be estimated by
scores.
using a variety of exposure durations, parameter Ai , a more indirect
measure of sensory effectiveness, can be derived from an experi- Lines Letters Stars Copying Bisection Drawing Sum
ment with constant exposure duration. It is dened as a compound Patient
of sensory effectiveness multiplied by the effective exposure dura- PB 36 23 53 0 7 1 120
tion (i.e. physical exposure time minus the perception threshold, OB 36 32 49 2 5 3 127
KKL 36 35 40 3 6 3 123
see Duncan et al., 1999). If, as in the given study, stimulus fea-
ML 36 31 48 4 6 3 128
tures and the effective exposure duration are held constant within FP 36 33 45 1 6 3 124
a given participant, Ai is proportional to si . Variations in A-values EW 36 32 45 1 6 3 123
can be therefore taken as (indirect1 ) measures of changes in sen- Cut off 34 32 51 3 7 2 129
sory effectiveness of that participant across different experimental
conditions, such as stimulus position and alerting conditions (see
Matthias et al., 2010). letter, and star cancellation, gure and shape copying, line bisection and repre-
sentational drawing. The results of each patient in the BIT are listed in Table 2.
2. Methods Patients were tested by an orthoptician in order to measure visual-eld decits.
Patients with visual eld decits were only included when, due to macular sparing,
2.1. Participants the region of the central 5-degrees stimulus array used in our study proved to be
unaffected.
Six right-handed patients with unilateral right hemisphere lesions from stroke All participants had normal or corrected-to-normal visual acuity and none of
affecting the medial cerebral artery (MCA) and six healthy control subjects were them suffered from colour blindness or any psychiatric or neurological disease (apart
studied. Age, gender distribution and education did not differ signicantly between from the current event in the neglect group). They were nave as to the purpose of
patients and control subjects (all ps > .70). In Table 1, the demographic characteristics the experiment. Written informed consent according to the Declaration of Helsinki
are listed for all subjects, and, additionally in neglect patients, the presence of visual II was obtained from all participants and the study was approved by the local ethical
eld decits and the time since the brain lesion occurred are listed. committee.
Brain damage in each patient, as demonstrated by CT or MRI scans that
were taken mostly in the acute state, was plotted by a neuroradiological 2.2. Apparatus and stimuli, design and procedure
expert blinded to the behavioural results of the patients, using MRICro soft-
ware (http://www.mccauslandcenter.sc.edu/mricro/), and a lesion overlap was 2.2.1. Whole report task
constructed (see Fig. 1). As can be seen, all patients exhibited a temporo-parietal Following previous studies of whole report tasks (e.g. Finke et al., 2005), on each
lesion in the right hemisphere. trial, either a single target letter (0.5 high 0.4 wide) or two target letters were
At the time of investigation, all patients had mild visuo-spatial neglect accord- presented. Targets appeared with equal frequency at each of the possible stimulus
ing to their performance in the conventional part of the Behavioural Inattention locations in the corners of an imaginary square (with an edge length of 5 ): upper
Test (BIT, Wilson et al., 1987), a standard test battery, including line crossing, left, lower left, upper right, lower right corner (see Fig. 2, bottom panel). Thus, tar-
gets were presented 2.5 away from the xation cross in the parafoveal elds on
both sides. Dual targets were placed either vertically (column display) or horizon-
tally (row display), but never diagonally. All target stimuli were masked. The masks
1
Although parameter A is (unlike the direct measure s) only an indirect esti- consisted of letter-sized squares (of 0.5 ) lled with a + and an and presented
mate of sensory effectiveness under constant exposure conditions, we will use the for 500 ms at each letter location. The letters for a given trial display were chosen
term sensory effectiveness A throughout the article up from now. We will, how- randomly from the set (ABEFHJKLMNPRSTWXYZ), with a particular letter appearing
ever, restrain from making comparisons between individuals who have been given only once at a time.
somewhat different exposure durations and who presumably have also differences The participants task was to verbally report the letters they had recognized with
in visual thresholds. certainty. The target letters could be named in any, arbitrary order, and there was
 K. Finke et al. / Neuropsychologia 50 (2012) 11781189 1181

Fig. 1. Lesion locations for neglect patients (PB, OB, KKL, ML, FP, and EW, from top to bottom) reconstructed for 8 transversal slices (left-hand side) and their positions in
sagittal orientation (right-hand side); numbers above each slice document the z-score in Talairach coordinates and lesion overlap (at the very bottom); the highest overlap
of patient groups lesions is shown in light red, the lowest overlap in dark red.

no emphasis on reporting speed. The experimenter entered the reported letter(s)
using the computer keyboard and initiated the next trial after the participants had
indicated that they were ready. The intertrial interval was 1000 ms.
The target exposure duration was individually determined for each participant
in a pretest part and then introduced into the experimental phase. The pretest con-
sisted of 72 single target trials (without alerting cues), presented for an exposure
duration of 171 ms. It was used to determine whether a participant was able to
reach an accuracy of 6080% for single-target report. If the participant performed
outside this range, the exposure duration in the experimental phase was adjusted
accordingly. The individual exposure durations of each participant are given
in Table 3.
2.2.2. Alertness conditions
We compared an alerting-cue condition (assumed to involve a phasic alert-
ness enhancement for a period of a few hundred milliseconds) with a no-cue
condition (purely involving intrinsic alertness). In the alerting-cue condition,
1182 K. Finke et al. / Neuropsychologia 50 (2012) 11781189

Fig. 2. Sequence of frames presented on a no-cue trial (top panel) and an alerting-cue trial (middle panel), together with the eight possible target displays: single targets,
dual targets in the same hemi-eld and dual targets presented in opposite hemi-elds (bottom panel; the T symbols denote target locations).

a white outline frame (5 5 ) ashed briey around the whole (potential) dis- location is likely to have discouraged patients from making eye movements. In any
play array. In the no-cue condition, the screen remained blank for the same length case, because the stimulus exposure durations were relatively short (200 ms), eye
of time. movements were unlikely to affect performance systematically. However, espe-
The alerting cue was non-informative as to the location of the upcoming target cially in neglect patients, suboptimal xation might occur already at the beginning
letters. Thus, although alerting the participants to the imminent appearance of the of a trial. Thus, to better control for eye- and head-movements during testing,
target array, this warning signal was designed to induce a spatially diffuse distri- we used a light-sensitive web-cam. With this device, eye and head movements
bution of attentional weighting across the (potential) stimulus display (i.e. it could could be observed on-line by the experimenter with minimal distraction of the
not be used to systematically orient spatial attention to the stimulus locations). participant. When eye or head movements occurred, participants were reminded
The non-informativeness of the alerting-cue with regard to the upcoming target to hold the xation and to try to avoid further movements.

2.2.3. SOA variation

Table 3 Three cue-target stimulus onset asynchronies2 were used: 80, 200, or 650 ms.
Target exposure durations (in ms) for each participant. Because the SOAs were randomized, the alerting cue was expected to primarily

Neglect patients Control subjects

Participant Exposure duration Participant Exposure duration

2
In the no-cue condition, the time frame within a trial was identical to the cued
PB 200 C1 157
condition except for the presentation of the cue itself. Because, in the no-cue condi-
OB 200 C2 171
tion, the screen remained blank when, in the cued condition, the cue appeared, the
KKL 171 C3 157
given SOAs of 80, 200 and 650 ms between cue and target presentation are, strictly
ML 157 C4 171
speaking, only valid for the cued condition. However, we will use the term SOA for
FP 157 C5 157
both the cued and the no-cue condition, in order to compare performance of subjects
EW 200 C6 128
at the same time point within a given trial.
 K. Finke et al. / Neuropsychologia 50 (2012) 11781189
induce a more general alerting/arousing effect, rather than supporting any specic
temporal expectations about the onset of the stimulus array.
2.2.4. Procedure
The PC-controlled experiment was conducted in a dimly lit room. Stimuli were
presented on a 17 in. monitor (1024 768 pixel screen resolution, 70 Hz refresh
rate). Participants viewed the monitor from a distance of 50 cm, controlled with the
aid of a head- and chinrest.
Fig. 2 illustrates the sequence of frames presented on a no-cue (top panel) and
an alerting-cue (bottom panel) trial. Following initiation of a trial sequence (by the
experimenter), participants xated a white xation cross (0.3 0.3 ) which was mixed ANOVAS were carried out on the TVA parameters in ques-
presented for the entire trial duration in the centre of the screen, on a black back-
ground. Then, after 1100 ms, either the alerting cue appeared for 50 ms or the screen
remained blank for the same length of time. After the variable cue-target SOA, the
whole-report display was presented for the pre-set exposure duration. Following
registration of the participants report, the next trial started after an intertrial inter-
val of 1000 ms. Cue/no-cue conditions as well as the three SOAs were presented in
random order within the same block.
In the rst session (lasting around 30 min), the whole-report pre-test was
applied to determine the individual presentation times. In neglect patients, fur-
thermore, the BIT was performed. The second to the fourth session (lasting around
45 min each) consisted of the experimental phase of the whole report. It com-
prised of eight different target conditions (four single-target and four dual-target
conditions) for each SOA (80, 200, 650 ms) and each of the two cueing conditions
(no-cue, alerting-cue). In total, the experiment comprised 846 trials per subject.
Standardized verbal instructions were given before each block of trials. In order to
avoid or minimize a possible inuence of the alerting cue on the no-cue condition,
alerting-cue and no-cue trials were presented in different blocks. The order of the
blocks and sessions was counterbalanced across subjects to control for sequence
effects. Hence, for three patients and three control subjects, the sequence was AB,
BA, AB, and for the other three subjects of each group BA, AB, BA (with A = block
with no-cue trials, and B = block with alerting-cue trials). To control for time-
on-day effects, the participants completed each of the four sessions at the same
time of day and within one week. A 5-min break was included in the middle of
each session.
2.2.5. Estimation of the TVA parameters
The whole-report task permitted quantitative estimates to be derived of visual
sensory effectiveness A and of attentional weight w, applying the algorithms
described in detail by Kyllingsbaek (2006) and used in several recent studies (e.g.
Bublak et al., 2005; Duncan et al., 1999; Finke et al., 2005; Habekost & Bundesen,
2003). These parameters were estimated separately for the four positions used in
the task (upper left, upper right, lower right, lower left): parameters Ai and wi ,
respectively, where i is the stimulus position.
Enhancement of global processing capacity of a given participant by an alert-
ing cue would be reected in higher estimates of A across all stimulus positions.
Indices of sensory effectiveness were calculated for each hemi-eld (by averaging
across the respective values for the upper and lower positions within each hemi-
eld): Aleft and Aright . This permitted us to examine whether alerting cues enhanced
sensory processing globally (across hemi-elds) in neglect patients, or whether,
possibly, sensory processing in the neglected contralesional hemi-eld would par-
ticularly benet (and, thus, would specically impact the lateralized disturbances
in the syndrome).
The parameter attentional weight is related to performance losses in multi-
element displays. Stimuli with higher weight are stronger competitors and, thus,
disturb report accuracy for further stimuli more than stimuli with lower attentional
weight. The estimated attentional weights for upper and lower stimulus positions
were averaged in order to receive separate indices of attentional weighting for each
hemi-eld, wleft and wright . However, the relative, rather than the absolute, amount
of attentional weight attributed to each hemi-eld provides useful information on
spatial attentional selection. Thus, a laterality index the parameter spatial distri-
bution of attentional weighting (w ) was computed. This index (which is related
to performance losses with (row) multi-element- compared to single-element dis-
plays) indicates whether one hemi-eld is favoured in the spatial selection process.
If attentional weights are biased towards one hemi-eld, performance in bilateral
(compared to unilateral) presentation conditions will suffer more for targets pre-
sented in the hemi-eld with relatively low attentional weight, compared to targets
in the hemi-eld with high weight. Formally, this index is dened as the ratio of
values across the two hemi-elds, w = wleft /(wleft + wright ). Hence, a value of w = .5
indicates a balanced distribution of weights, values of w > .5 indicate a leftward,
and values of w < .5 a rightward spatial-attentional bias. If attentional weights are
biased towards one hemi-eld, performance in bilateral (compared to unilateral)
presentation conditions will suffer more for targets presented in the hemi-eld
with relatively low attentional weight, compared to targets in the hemi-eld with
high weight. Note, though, that under bilateral stimulus conditions, it is in prin-
ciple also possible that a difference in performance between left- and right-side
targets is due to reduced sensory effectiveness in target discrimination on one
side. This factor was controlled for by the hemi-eld-specic analyses of sensory
effectiveness A.
1183
3. Results
In what follows, we will rst describe the qualitative accuracy
pattern (correctly identied letters) obtained across the different
groups and experimental conditions. Second, we will present the
parameter estimates derived in TVA-based model ts to the data,
that is, sensory effectiveness and spatial attentional weighting.
Next, in order to compare sensory effectiveness and spatial atten-
tional weighting effects across groups, SOAs and cueing conditions,
tion.
3.1. Report accuracy: raw data indicators of sensory
enhancement and of changes in the spatial distribution of
attention
To illustrate sensory effectiveness and attentional weighting
across the two hemi-elds, Fig. 3 shows separately the mean pro-
portion of target letters correctly identied by patients and controls
in each hemi-eld and at each SOA for ve experimental conditions:
single-target letter; target accompanied by a target in the ipsilat-
eral eld (column displays) and target accompanied by a target in
the contralateral eld (row displays). In general, the performance
levels of both groups are in accordance with the prediction from
the TVA model: accuracy was highest in single-target conditions
and decreased in dual target conditions.
Accuracy in single target conditions (rst two bars of each g-
ure) can be taken as a raw data indicator of how well a stimulus is
basically processed, at a given exposure duration, under conditions
of no stimulus competition, that is, of pure sensory effectiveness, in
each hemi-eld. Due to individually adjusted exposure durations
for each participant, between-group comparisons do not make
sense with respect to the general level of accuracy. However, accu-
racy changes across alerting conditions and hemi-elds can be
taken as indicators of changes of sensory effectiveness and of spa-
tial weighting. Fig. 3a shows that in neglect patients an alerting
cue seems to enhance accuracy for single targets at the 80-ms and
200-ms SOAs. However, this improvement does mainly occur in the
right hemi-eld; only at the longest, 650-ms SOA, an improvement
is also manifest in the left eld. In control participants, an alertness
cue does not seem to enhance single-target report in general. How-
ever, also in this group, an enhancement is evident for left visual
eld single targets at the longest SOA (Fig. 3b).
A comparison between accuracy losses in column displays on
the one hand and row displays on the other indicates possible
changes in spatial attentional weighting processes. As would be
expected in neglect patients with a reduced awareness of the
left side, in the no-cue condition, report accuracy was lowest for
row displays (i.e. in conditions with competition between the two
hemi-elds), especially for the left side. However, this pattern was
modied in the alerting-cue conditions. Especially at the shortest
SOA, report accuracy for the left side even exceeded that for the
right side (Fig. 3a).
3.2. TVA parameter estimates
For each participant, cueing condition and SOA, sensory effec-
tiveness and attentional weights were quantied for each of the
four possible target locations. The empirically assessed mean accu-
racy scores for the different experimental conditions and the values
theoretically predicted (based on the best ts of the TVA model
parameters) showed a close correspondence: the average corre-
lation between the observed and the predicted scores across all
SOAs in the no-cue condition was .92 (neglect patients, SD = .04) and
.86 (control participants, SD = .07). In the alerting-cue condition, it
was .88 (neglect patients, SD = .09) and .83 (control participants,
1184 K. Finke et al. / Neuropsychologia 50 (2012) 11781189

(a) Neglect, no-cue, 80 ms SOA Neglect, cue, 80 ms SOA

100 100

Accuracy (%)

Accuracy (%)
80 80
L L
60 60
R R
40 40
20 20

0 0

ST DT column DT row ST DT column DT row

Neglect, no-cue, 200 ms SOA Neglect, cue, 200 ms SOA

100 100
Accuracy (%)

Accuracy (%)
80 80
60
L L
60
R R
40 40
20 20
0 0
ST DT column DT row ST DT column DT row

Neglect, no-cue, 650 ms SOA Neglect, cue, 650 ms SOA

100
100
Accuracy (%)

Accuracy (%)
80 80
60
L L
60
R R
40 40
20 20

0 0

ST DT column DT row ST DT column DT row

(b) Control, no-cue, 80 ms SOA Control, cue, 80 ms SOA

100 100
Accuracy (%)

Accuracy (%)

80 80
60
L 60
L
R R
40 40
20 20
0 0
ST DT column DT row ST DT column DT row

Control, no-cue, 200 ms SOA Control, cue, 200 ms SOA

100 100
Accuracy (%)

Accuracy (%)

80 80
60
L L
60
R R
40 40
20 20
0 0
ST DT column DT row ST DT column DT row

Control, no-cue, 650 ms SOA Control, cue, 650 ms SOA

100 100
Accuracy (%)

Accuracy (%)

80 80
60
L 60
L
R R
40 40
20 20
0 0
ST DT column DT row ST DT column DT row

Fig. 3. (a and b) Mean proportions of correctly identied target letters (% correct) presented in either the left or the right hemi-eld in the different experimental conditions,
separately for neglect patients (a) and for control participants (b). Each bar represents the mean in one condition: single target (ST), target accompanied by a target in the
same hemi-eld/dual targets in column displays (DT column), target accompanied by a target in the opposite hemi-eld/dual targets in row displays (dual target row). For
each group, separate gures are shown for the two cueing conditions (no-cue, alerting-cue) and the three SOAs (80, 200, 650 ms). Error bars give the standard errors.
 K. Finke et al. / Neuropsychologia 50 (2012) 11781189 1185

Neglect, left hemi-field Neglect, right hemi-field

Sensory effectiveness A

Sensory effectiveness A
5 5
4 4
3 no-cue 3 no-cue
2 cue 2 cue

1 1
0 0
80 200 650 80 200 650
SOA SOA

Control, left hemi-field Control, right hemi-field

Sensory effectiveness A

Sensory effectiveness A
5 5
4 4
3 no-cue 3 no-cue
2 cue 2 cue

1 1
0 0
80 200 650 80 200 650
SOA SOA
Fig. 4. Parameter A (sensory effectiveness) for the left (left half) and the right hemi-eld (right half), separately for neglect patients (upper half) and control participants
(lower half). The different lines depict sensory effectiveness for the two cueing conditions (no-cue, alerting-cue) as a function of SOA (80, 200, 650 ms). Error bars give the
standard errors.

SD = .04). Thus, in the no-cue condition, the model explained 84% separately for the two hemi-elds. As can be seen, the temporary
(neglect patients) and, respectively, 74% (control participants), and sensory enhancement in the right eld is reliably observed at the
in the alerting-cue condition it explained 77% (neglect patients) 80-ms and 200-ms SOAs, in each single case. Cue-induced sensory
and, respectively, 68% (control participants) of the variance. effects in the left, neglected, eld are generally more variable across

3.3. TVA parameter sensory effectiveness A: quantitative

Neglect, single cases, left hemi-field PB no-cue
estimation of global and/or lateral processing capacity
6 OB no-cue
Sensory effectiveness A

enhancement KKL no-cue

5 ML no-cue
Fig. 4 presents the time course of the effect of the alerting cue, FP no-cue
4
compared to the no-cue condition, on sensory effectiveness A in EW no-cue
PB cue
each hemi-eld, for each group. 3
OB cue
Separate ANOVAs were conducted for the neglect patients and KKL cue
2
for the control participants, each with the within-subject factors ML cue
SOA (80, 200, 650 ms), cue (no-cue, alerting-cue), and side (left 1 FP cue
hemi-eld, right hemi-eld). EW cue
0
80 200 650
3.3.1. Neglect patients
SOA
In neglect patients, all main effects were signicant [cue: F(1,
5) = 11.13; p < .05; SOA: F(2, 4) = 6.78; p < .05; side: F(1, 5) = 7.19; Neglect, single cases, right hemi-field PB no-cue
p < .05]. As can be seen in Fig. 4, the alerting cue did indeed induce OB no-cue
6
Sensory effectiveness A

the global attentional resource enhancement in question (main KKL no-cue

effect of cue). However, signicant interactions of SOA side [F(2, 5 ML no-cue

FP no-cue
4) = 4.62; p < .05] and, more importantly, of SOA cue side [F(2, 4 EW no-cue
4) = 4.31; p < .05] showed that the time course of sensory changes
3 PB cue
and the impact of cueing on these changes differed between hemi- OB cue
elds. In accordance with the accuracy pattern for single targets, 2 KKL cue
the three-way interaction reects the fact that, in the right hemi- ML cue
1
eld, the alerting cue induced a signicant enhancement of sensory FP cue
effectiveness for a relatively short period of time. Sensory effective- 0 EW cue
ness was higher following an alerting cue compared to the no-cue 80 200 650
condition at the SOAs of 80 ms [t(5) = 5.57; p < .01] and of 200 ms SOA
[t(5) = 3.26; p < .05], but not at the SOA of 650 ms (p > .75). In con-
trast, in the left hemi-eld, sensory effectiveness was enhanced Fig. 5. Single case data of parameter A (sensory effectiveness) for each of the neglect
patients, separately for the left (upper gure) and the right hemi-eld (lower g-
only at the longest SOA of 650 ms (t(5) = 2.48; p < .05).
ure). For each participant, solid and dashed lines depict sensory effectiveness in the
In Fig. 5, the time course of cue-induced sensory effectiveness no-cue and the alerting-cue condition, respectively, as a function of SOA (80, 200,
changes is illustrated for each neglect patient (single-case data), 650 ms).
1186 K. Finke et al. / Neuropsychologia 50 (2012) 11781189

patients, and no systematic enhancement effect seems to prevail. Neglect

Notably, the three patients with visual eld defects (PB, OB, KKL)
showed values that were comparable to those of patients without 0.8 leftward bias
visual eld defects in each condition, and, furthermore, they also

w Left/(w Left + w Rright)

0.7
showed comparable SOA- and cue-induced effects in both hemi-
elds. 0.6
To summarise, with respect to sensory effectiveness, neglect no-cue
patients showed a marked cue-induced enhancement at the short- 0.5
est SOA and, too a lesser degree, also at the medium SOA within
cue
0.4
the right hemi-eld. In contrast, within the left hemi-eld, sen-
sory effectiveness increased at the longest SOA, only, in the cue 0.3
condition. Notably, neglect patients did not show any dynamics in rightward bias
sensory effectiveness whatsoever in the no-cue condition, in both 0.2
hemi-elds. 80 200 650
SOA

3.3.2. Control participants

Control
In control participants, the main effect of side [F(1, 5) = 7.09;
0.8 leftward bias
p < .05] and the SOA cue side interaction [F(2, 4) = 12.56; p < .01]

w Left/(w Left + w Rright)

were signicant. Mirroring somewhat the results for neglect
patients in the left hemi-eld, sensory effectiveness was enhanced
by an alerting cue at the longest SOA of 650 ms in the right hemi-
eld in control subjects (t(5) = 2.16; p < .05). For the left hemi-eld,
sensory effectiveness increased from the 200-ms to the 650-ms
SOA [t(5) = 3.16; p < .05] in the alerting-cue condition,; and only at
the latter (the longest SOA) was there a tendency towards a higher
sensory effectiveness for the alerting-cue compared to the no-cue
condition [t(5) = 2.34; p < .07]. Another (non-signicant) tendency
suggested sensory processing to be somewhat higher at the 200-
ms compared to the 80-ms SOA in the no-cue condition [t(5) = 2.36;
p < .07].
In sum, in control subjects, the enhancement of sensory effec-
tiveness at the longest SOA in the right hemi-eld following a cue
mirrored the performance of neglect patients within the left hemi-
eld. And the same result was obtained in control subjects also
within the left hemi-eld, with even a more marked increase of
sensory effectiveness at the longest SOA. In the no-cue condition,
control subjects showed the largest differences of sensory effective-
ness within the left hemi-eld: There was a strong increase from
the shortest to the medium SOA, where sensory effectiveness even
exceeded that in the cueing condition, and then a decline to the
longest SOA. This pattern was quite different from the right hemi-
eld, where sensory effectiveness achieved its greatest value at the
shortest SOA.
3.4. TVA parameter spatial distribution of attentional weighting
w : quantitative estimation of laterality changes in selection
processes
Fig. 6 illustrates the average SOA-dependent time course of the
spatial distribution of the attentional weighting parameter w for
the no-cue and the alerting-cue condition. As can be seen, the
cue induced changes of spatial attentional weighting mainly in the
patient group and mainly at the shorter SOAs.
A mixed ANOVA with the factors group, SOA and cue and with
w as dependent variable conrmed this pattern statistically. This
analysis revealed all main effects [group: F(1, 10) = 55.76; p < .01;
SOA: F(2, 9) = 5.18; p < .05; cue: F(1, 10) = 22.17; p < .01] and two-
way interactions [group SOA: F(2, 9) = 4.95; p < .05; group cue:
F(1, 10) = 6.35; p < .05; SOA cue: F(2, 9) = 3.89; p < .05], and the
three-way interaction group SOA cue [F(2, 9) = 5.57; p < .05] to
be signicant. Separate repeated-measures ANOVAs were con-
ducted subsequently in order to examine the effects of cue and
SOA within both groups.
0.7
0.6
no-cue
0.5
cue
0.4
0.3
rightward bias
0.2
80 200 650
SOA
Fig. 6. Parameter w (spatial distribution of attentional weighting) for neglect
patients (upper gure) and for control participants (lower gure). Separate lines
depict the laterality of spatial weighting for the two cueing conditions (no-cue,
alerting-cue), as a function of SOAs (80, 200, 650 ms). Error bars give the standard
errors. Values of w > .50 = leftward attentional bias; w < .50 = rightward attentional
bias; w = .50 = no bias.
3.4.1. Neglect patients
In neglect patients, the main effect of cue [F(1, 5) = 51.26; p < .01]
was signicant, whilst that of SOA [F(2, 4) = 3.52; p > .10] was not.
Furthermore, a signicant SOA cue interaction [F(2, 4) = 5.61;
p < .05] was obtained. As illustrated in Fig. 7, compared to the no-cue
condition, the alerting cues induced a signicant re-distribution of
attentional weights towards the left hemi-eld at the SOAs of 80 ms
[t(5) = 3.92; p < .01] and 250 ms, [t(5) = 6.48; p < .01]; whilst this
effect disappeared at the longest SOA of 650 ms [p > .70].
Additional analyses examined whether the spatial distribution
of attentional weighting was signicantly biased towards either
side, compared to the optimal, balanced value of .5. As illustrated in
Fig. 6, in no-cue conditions, neglect patients were generally biased
towards the right (w < .05). This was conrmed by signicant or
borderline signicant effects at all SOAs [80 ms: t(5) = 3.15; p < .05;
200 ms: t(5) = 6.86; p < .01; 650 ms: t(5) = 1.93; p < .07]. Follow-
ing an alerting cue, a signicant leftward bias (w > .5) appeared
at the shortest, 80-ms SOA [t(5) = 5.93; p < .01], followed again by a
signicantly rightward biased distribution of weights at the longer,
200-ms [t(5) = 3.05; p < .05] and 650-ms [t(5) = 6.00; p < .01] SOAs.
From Fig. 7, the single case neglect patient data can be seen
to conrm the changes of attentional selection laterality induced
by the alerting cue and also its time course (again, there was
no indication of any difference between patients PB, OB, and
KKL with visual eld decits and patients without visual eld
decits). In the no-cue conditions, a general, more or less pro-
nounced rightward spatial attentional bias (w < .5) is evident
 K. Finke et al. / Neuropsychologia 50 (2012) 11781189
Fig. 7. Single case data of parameter w (spatial distribution of attentional weight-
ing) for each of the neglect patients in the no-cue condition (upper gure) and
the alerting-cue condition (lower gure), as a function of SOA (80, 200, 650 ms).
Error bars give the standard errors. Values of w > .50 = leftward attentional bias;
w < .50 = rightward attentional bias; w = .50 = no bias.
in each patient and cueing condition. Following an alerting cue,
the overall pattern described above at the group level was con-
sistently present in each single case. Each patient showed a
lateral redistribution of attentional weights towards the left side,
and even a slight leftward bias in each single case (w > .5)
at the shortest, 80-ms SOA which gradually disappears as SOA
increases.
In summary, cueing had a marked impact on the spatial bias of
attention in neglect patients. In fact, at the shortest SOA, the right-
ward bias disappeared and turned into a leftward spatial weighting
of attention. This effect dropped linearly at the later SOAs, so that at
the longest SOA, a rightward bias was again obtained. In contrast,
more or less the same rightward bias prevailed across all SOAs in
the no-cue condition.
3.4.2. Control participants
The main effect of cue was not signicant [F(1, 5) = 1.61; p < .01],
but the ANOVA revealed a signicant main effect of SOA [F(2,
4) = 8.31; p < .05] and a tendency for an SOA cue interaction [F(2,
4) = 3.32; p < .08]. Mainly driven by the no-cue condition (see Fig. 6),
a slight but signicant shift of spatial attentional weights from the
left to the right side occurred from the shortest SOA of 80 ms to that
of 200 ms (t(5) = 3.60; p < .05) and another signicant shift back to
the left side from the SOA of 200 ms to that of 650 ms (t(5) = 4.27;
p < .01). The spatial distribution of attention did not differ signif-
icantly between the shortest and the longest SOA (p > .75). In the
cue condition, there was a slight leftward shift of spatial attentional
1187
weighting as a function of SOA, but this effect was not signicant
(t(5) = 1.56; p > .15).
Comparisons against the optimal value of 0.5 did not indicate
a lateralization in any of the cueing conditions at the shortest and
the intermediate SOA (all t < 2.00, all p > .10). At the longest SOA
of 650 ms, control participants were signicantly biased towards
the left in both cueing conditions [no-cue: t(5) = 3.66; p < .05; cue:
t(5) = 4.20; p < .01].
To summarise, concerning the bias of spatial attention, control
subjects showed a complementary pattern compared to the results
of sensory effectiveness in the no-cue condition: At the medium
SOA, a slight rightward shift occurred, whereas a leftward bias
prevailed at both the shortest and the longest SOA. In the cue-
ing condition, there was a slight leftward bias, which was largely
consistent across SOAs.
4. Discussion
The aim of the present study was to investigate the nature and
time course of the effect of a phasic alerting cue on spatial and
non-spatial aspects of attention, as compared to a no-cue condi-
tion, in patients with unilateral neglect for contralesional space. By
using a TVA-based whole-report paradigm our study was designed
to independently and separately examine the inuence of cue-
induced phasic alertness on processing capacity (i.e. on sensory
effectiveness as an indicator of processing speed) and on the spa-
tial distribution of attentional weights across the two hemi-elds
(i.e. on the laterality of selection). We assessed (1) whether cueing
would affect sensory and/or spatial attentional-components and, if
so, (2) in what time ranges these various effects would occur and
(3) whether they would differ from those found in normal control
participants.
As a rst main result of our study, control subjects showed
the greatest effects in the no-cue condition, with complemen-
tary, even opposing patterns for sensory effectiveness (left-eld
enhancement) and spatial weighting of attention (rightward shift),
especially at the medium SOA. This result is in close correspondence
with the data presented by Matthias et al. (2010) in their Experi-
ment 2, which had applied the same paradigm as in the present
study, in particular using a randomised variation of SOA, in young
healthy subjects. Matthias et al. (2010) interpreted this result as
reecting the effects of top-down mediated modulations of intrin-
sic alertness and conrmed this assumption by showing that the
pattern disappeared when SOAs were presented blockwise, so that
adaptation of a mental set on a trial-by-trial basis was no longer
necessary. The seemingly contradictory overlay between enhanced
intrinsic alertness and a, in relation to the other SOAs, more right-
ward spatial bias at the medium SOA, was interpreted as indicating
that enhanced alertness inuences two separate systems one sup-
porting processing speed, the other governing spatial attention
that act independently from each other, but on a different time
scale. The enhancement of processing speed occurs rst, and is
followed by a leftward shift of spatial attention, observable at the
longest SOA.
Following this assumption, the results of the control subjects in
the present study can be interpreted as further evidence that spatial
and non-spatial mechanisms of attention are mediated by inde-
pendent processing systems which exert their effects, as a result of
enhanced intrinsic alertness, in temporal succession. According to
the neuro-cognitive model proposed by Heilman et al. (2003) and
Posner and Petersen (1990), modulations of intrinsic alertness are
mediated by a right-hemispheric network in which frontal regions
exert top-down control, via thalamic nuclei, on posterior cortical
areas related to perceptual processing, by activating noradrenergic
nuclei of the ponto-mesencephalic part of the brainstem (see also
1188 K. Finke et al. / Neuropsychologia 50 (2012) 11781189
Mesulam, 2000, for a similar idea). Consistent with right temporo-
parietal damage interfering with this system, and in congruence
with empirical evidence that regulation of intrinsic alertness is de-
cient in neglect patients (Bartolomeo & Chokron, 2002; Heilman
et al., 2003), there was almost no sign of any modulation of the
performance of neglect patients in the no-cue condition of our
study.
In contrast, and as the second main result of our study,
neglect patients showed the greatest effects in the cue condi-
tion, especially at the shortest SOA. Interestingly, as for control
subjects though complementary concerning the hemi-eld the
pattern for sensory effectiveness (right-eld enhancement) and
spatial bias (leftwards) was opposing. Especially because the sam-
ple size was relatively small in our study, it is noteworthy that this
result was highly consistent across patients at the single case level.
This pattern is compatible with the above presented interpretation
of separate systems for speed and spatial attention. However, both
systems bring about their effects not in temporal succession, but in
parallel, already shortly after cue-induced phasic enhancement of
alertness. Moreover, the switching of the hemi-eld, in which the
effects are observed for patients as compared to control subjects,
requires an explanation. After all, if a right-hemispheric dominance
for alertness is assumed, the greatest effects would be expected to
be observed within the left hemi-eld, as was the case in control
participants.
Taking into account the damage to the right-sided alerting net-
work in neglect patients (indicated by the lack of intrinsic alertness
modulations), and the fact that a phasic alerting cue exerted effects
in both hemi-elds (although predominantly on the right side), the
greater enhancement of sensory effectiveness in the ipsilesional
eld can be explained in the following way: Phasic alertness might
activate a system supporting processing speed (indicated by sen-
sory effectiveness in our study) that leads to a global stimulation
within both hemispheres. When only a single stimulus is present
within one hemi-eld, then the intact hemisphere (the left one
in the neglect patients) will always outperform the damaged one,
giving rise to a right-hemi-eld advantage in single target displays.
The simultaneous effect of phasic alertness on the spatial bias
points to the additional cue-induced activation of a second system,
devoted to spatial attentional weighting, which exerts a specically
right-hemispheric effect. This system would become especially rel-
evant in situations of inter-hemispheric competition, as in the
bilateral target displays of the whole-report paradigm of our study,
and, thus, would lead to a more balanced spatial weighting in these
conditions.
The rapidly evolving effects on sensory effectiveness and spa-
tial bias following the cue suggest that the two simultaneously
activated systems mediating speed and spatial weighting rather
represent lower-level centres. Therefore, one plausible neural can-
didate structure underlying these effects is the thalamus, which
is known to be relevant both in the ascending thalamic mes-
encephalic phasic alertness system (Fan, McCandliss, Fossella,
Flombaum, & Posner, 2005; Posner & Petersen, 1990; Sturm &
Willmes, 2001; Thiel & Fink, 2007) and which has also been
proposed to be critical for visual extinction by several authors
(Fimm et al., 2001; Karnath, Himmelbach, & Rorden, 2002; Rafal
& Posner, 1987). For instance, Habekost and Rostrup (2006)
reported a rightward spatial bias in a comparable TVA-based
paradigm following stroke in the right-sided pulvinar nucleus of
the thalamus. In fact, on the neural interpretation of the TVA
(NTVA) proposed by Bundesen, Habekost, & Kyllingsbk (2005),
the pulvinar nucleus of the thalamus is assumed as the critical
region mediating spatial-attentional weighting across the visual
eld.
Based on our results, we would suggest that both thala-
mic subsystems are simultaneously activated by another system
which is directly related to phasic alertness and responds to
the alerting cue. A possible candidate for this would be the
ascending reticular activating system, which receives direct sen-
sory input (Jones, 2003). These assumptions would predict that
neglect patients with thalamic lesions do not show effects of
cue-induced enhancement of phasic alerting, and, therefore, no
increase of sensory effectiveness and no balanced spatial weighting
should be observed in these patients after presenting an alertness
cue.
Our data clearly show that enhanced phasic alerting brings
about a re-distribution of attentional weights in neglect patients.
Whilst a rightward spatial bias prevailed in the no-cue condition
and also at the longest SOA in the cue condition, a more balanced, in
fact even slightly leftward spatial bias was found 80 ms and 200 ms
after cue presentation. This effect was very consistent on the single
case level and was found in each of the six patients assessed (see
Fig. 7). This result is in congruence with the earlier data presented
by Robertson et al. (1998) for a different paradigm, and demon-
strates the adaptive potential of a damaged attentional network.
However, it also raises the question of the clinical implications of
such a short-lived effect. After all, at the longest SOA, the patho-
logical rightward bias reappeared and the neglect pattern was fully
re-established.
It should be taken into account, however, that a core function
of an alertness cue is to initiate an orienting response (Posner,
2008). In reality settings, this is typically accomplished by eye
or head movements which support the aim to foveate a salient
stimulus that is available afterwards for close examination. In our
study, when such a stimulus did not appear within the 650 ms fol-
lowing the alertness cue, the neglect pattern was re-established.
However, this does not imply that, in case visual stimulation is
indeed provided, the cue-induced benet resulting is necessarily
only shortlived. In our study we only used briey presented stim-
uli. It remains an open empirical issue whether, under conditions
of enduring sensory stimulation (such as, e.g. the unlimited pre-
sentation of a visual search display), the alertness-induced effect
might be more stable and even remain signicant for the dura-
tion of any target scene presentation. Furthermore, it should be
tested whether, using a systematic alertness training, the effec-
tive period, i.e. the gap between cue and target, without sensory
stimulation, can be prolonged. Nevertheless, should alerting cues
be used as a therapeutic tool in a clinical setting, then the SOA
between an alerting cue and a target should be adjusted to the
effective short-time window documented in our study in order to
reach the maximum benet, at least at the beginning of an alertness
training.
In summary, based on the present results, it can be concluded
that higher levels of alertness can help overcome the typical neglect
symptoms, such as rightward lateralization and unilateral extinc-
tion, indicative of the relevance of alertness in disturbed attentional
competition and, thus, spatial-attentional asymmetries. Notably,
a quick redistribution of attentional weights across both hemi-
elds has been identied as the major source of the performance
benet even at the single case level, i.e. in each of the patients
tested. Our results are in accordance with previous studies showing
a rightward bias in exploration behaviour and a unilateral right-
ward lateralization in neglect (e.g. Bartolomeo & Chokron, 2002;
Bublak et al., 2005; Duncan et al., 1999; e.g. Heilman et al., 2003;
Karnath, 1988). In addition, they provide important clues as to
both the time course and the effective mechanism responsible for
the alleviation of these decits. Due to our low sample size and
the limitations of our neuroanatomical evidence, these clues have
to be further corroborated by future studies with larger sample
sizes also aiming at a closer examination of longer-term training
effects and a better understanding of the precise neuroanatomical
correlates
 K. Finke et al. / Neuropsychologia 50 (2012) 11781189
References
Bartolomeo, P., & Chokron, S. (2002). Orienting of attention in left unilateral neglect.
Neuroscience and Biobehavioral Reviews, 26, 217234.
Bender, M. B. (1952). Disorders in perception. Springeld, IL: Charles C Thomas.
Bisiach, E., & Vallar, G. (1988). Hemineglect in humans. In F. Boller, J. Grafman, G.
Rizzolatti, & H. Goodglass (Eds.), Handbook of neuropsychology (pp. 195222).
New York, NY, US: Elsevier Science.
Bublak, P., Finke, K., Krummenacher, J., Preger, R., Kyllingsbaek, S., Mller, H. J., et al.
(2005). Usability of a theory of visual attention (TVA) for parameter-based mea-
surement of attention II: Evidence from two patients with frontal or parietal
damage. Journal of the International Neuropsychological Society, 11, 843854.
Bundesen, C. (1990). A theory of visual attention. Psychological Review, 97, 523
547.
Bundesen, C. (1998). A computational theory of visual attention. Philosophical
Transactions of the Royal Society of London: Series B Biological Sciences, 353,
12711281.
Bundesen, C., Habekost, T., & Kyllingsbk, S. (2005). A neural theory of visual
attention: Bridging cognition and neurophysiology. Psychological Review, 112,
291328.
Coull, J. T., & Nobre, A. C. (1998). Where and when to pay attention: The neural sys-
tems for directing attention to spatial locations and to time intervals as revealed
by both PET and fMRI. Journal of Neuroscience, 18, 74267435.
Coull, J. T., Frackowiak, R. S., & Frith, C. D. (1998). Monitoring for target objects:
Activation of right frontal and parietal cortices with increasing time on task.
Neuropsychologia, 36, 13251334.
Danckert, J., & Ferber, S. (2006). Revisiting unilateral neglect. Neuropsychologia, 44,
9871006.
DeGutis, J. M., & van Vleet, T. M. (2010). Tonic and phasic alertness training: a novel
behavioral therapy to improve spatial and non-spatial attention in patients with
hemispatial neglect. Frontiers in Human Neuroscience, 4, pii: 60
Desimone, R., & Duncan, J. (1995). Neural mechanisms of selective visual attention.
Annual Review of Neuroscience, 18, 193222.
Duncan, J., Bundesen, C., Olson, A., Humphreys, G., Chavda, S., & Shibuya, H. (1999).
Systematic analysis of decits in visual attention. Journal of Experimental Psy-
chology: General, 128, 450478.
Fan, J., McCandliss, B. D., Fossella, J., Flombaum, J. I., & Posner, M. I. (2005). The
activation of attentional networks. Neuroimage, 26, 471479.
Fimm, B., Zahn, R., Mull, M., Kemeny, S., Buchwald, F., Block, F., et al. (2001). Asymme-
tries of visual attention after circumscribed subcortical vascular lesions. Journal
of Neurology, Neurosurgery and Psychiatry, 71, 652657.
Finke, K., Bublak, P., Krummenacher, J., Kyllingsbaek, S., Muller, H. J., & Schneider,
W. X. (2005). Usability of a theory of visual attention (TVA) for parameter-
based measurement of attention I: Evidence from normal subjects. Journal of
the International Neuropsychological Society, 11, 832842.
Funk, J., Finke, K., Mller, H. J., Utz, K. S., & Kerkhoff, G. (2010). Effects of lateral
head inclination on multimodal spatial orientation judgements in neglect: Evi-
dence for impaired spatial orientation constancy. Neuropsychologia, 48, 1616
1627.
Habekost, T., & Bundesen, C. (2003). Patient assessment based on a theory of visual
attention (TVA): Subtle decits after a right frontal-subcortical lesion. Neuropsy-
chologia, 41, 11711188.
Habekost, T., & Rostrup, E. (2006). Persisting asymmetries of vision after right side
lesions. Neuropsychologia, 44, 876895.
Heilman, K. M., Schwartz, H. D., & Watson, R. T. (1978). Hypoarousal in patients with
neglect syndrome and emotional indifference. Neurology, 28, 229232.
Heilman, K. M., Watson, R. T., & Valenstein, E. (2003). Neglect and related disorders.
In K. M. Heilman, & E. Valenstein (Eds.), Clinical neuropsychology (pp. 296346).
Oxford: Oxford University Press.
Hjaltason, H., Tegnr, R., Tham, K., Levander, M., & Ericson, K. (1996). Sustained
attention and awareness of disability in chronic neglect. Neuropsychologia, 34,
12291233.
Husain, M., & Rorden, C. (2003). Non-spatially lateralized mechanisms in hemispatial
neglect. Nature Reviews Neuroscience, 4, 2636.
Husain, M., Shapiro, K., Martin, J., & Kennard, C. (1997). Abnormal temporal dynamics
of visual attention in spatial neglect patients. Nature, 385, 154156.
1189
Jones, B. E. (2003). Arousal systems. Frontiers in Bioscience, 8, 438451.
Karnath, H. O. (1988). Decits of attention in acute and recovered visual hemi-
neglect. Neuropsychologia, 26, 2743.
Karnath, H. O., Himmelbach, M., & Rorden, C. (2002). The subcortical anatomy
of human spatial neglect: Putamen, caudate nucleus and pulvinar. Brain, 125,
350360.
Kinomura, S., Larsson, J., Gulyas, B., & Roland, P. E. (1996). Activation by attention
of the human reticular formation and thalamic intralaminar nuclei. Science, 271,
512515.
Kyllingsbaek, S. (2006). Modeling visual attention. Behavior Research Methods, 38,
123133.
Matthias, E., Bublak, P., Mller, H. J., Schneider, W. X., Krummenacher, J., & Finke,
K. (2010). The inuence of alertness on spatial and non-spatial components
of visual attention. Journal of Experimental Psychology: Human Perception and
Performance, 36, 3856.
Mesulam. (2000). Attentional networks, confusional states, and neglect syndromes.
In M. M. Mesulam (Ed.), Principles of behavioral and cognitive neurology (pp.
174256). New York: Oxford University Press.
Posner, M. I. (1978). Chronometric explorations of mind. Hillsdale, NJ: Lawrenc Erl-
baum Associates.
Posner, M. I. (2008). Measuring alertness. Annual of the New York Academy of Sciences,
1129, 193199.
Posner, M. I., & Petersen, S. E. (1990). The attention system of the human brain.
Annual Review of Neuroscience, 13, 2542.
Rafal, R. D., & Posner, M. I. (1987). Decits in human visual spatial attention following
thalamic lesions. Proceedings of the National Academy of Sciences of the United
States of America, 84, 73497735.
Raz, A., & Buhle, J. (2006). Typologies of attentional networks. Nature Reviews Neu-
roscience, 7, 367379.
Robertson, I. H., Mattingley, J. B., Rorden, C., & Driver, J. (1998). Phasic alerting of
neglect patients overcomes their spatial decit in visual awareness. Nature, 395,
169172.
Robertson, I. H., Manly, T., Beschin, N., Daini, R., Haeske-Dewick, H., Hmberg, V., et al.
(1997). Auditory sustained attention is a marker of unilateral spatial neglect.
Neuropsychologia, 35, 15271532.
Samuelsson, H., Hjelmquist, E. K., Jensen, C., Ekholm, S., & Blomstrand, C. (1998).
Nonlateralized attentional decits: An important component behind persisting
visuospatial neglect? Journal of Clinical and Experimental Neuropsychology, 20,
7388.
Sturm, W., & Willmes, K. (2001). On the functional neuroanatomy of intrinsic and
phasic alertness. Neuroimage, 14, S76S84.
Sturm, W., de Simone, A., Krause, B. J., Specht, K., Hesselmann, V., Raderma-
cher, I., et al. (1999). Functional anatomy of intrinsic alertness: Evidence for a
fronto-parietal-thalamic-brainstem network in the right hemisphere. Neuropsy-
chologia, 37, 797805.
Sturm, W., Schmenk, B., Fimm, B., Specht, K., Weis, S., Thron, A., et al. (2006). Spa-
tial attention: More than intrinsic alerting? Experimental Brain Research, 171,
1625.
Thiel, C. M., & Fink, G. R. (2007). Visual and auditory alertness: Modality-specic
and supramodal neural mechanisms and their modulation by nicotine. Journal
of Neurophysiology, 97, 27582768.
Thimm, M., Fink, G. R., Kust, J., Karbe, H., & Sturm, W. (2006). Impact of alertness
training on spatial neglect: A behavioural and fMRI study. Neuropsychologia, 44,
12301246.
van Kessel, M. E., van Nes, I. J., Brouwer, W. H., Geurts, A. C., & Fasotti, L. (2010).
Visuospatial asymmetry and non-spatial attention in subacute stroke patients
with and without neglect. Cortex, 46, 602612.
von Cramon, D., & Kerkhoff, G. (1993). On the cerebral organization of elementary
visuo-spatial perception. In B. Bulyas, D. Ottoson, & P. Roland (Eds.), Functional
organization of the human visual cortex. Pergamon Press: Oxford.
Wilson, B. A., Cockburn, J., & Halligan, P. (1987). Behavioral Inattention Test (BIT).
Titcheld: Thames Valley Test Company.
Wojciulik, E., Rorden, C., Clarke, K., Husain, M., & Driver, J. (2001). Group study of
an undercover test for visuospatial neglect: Invisible cancellation can reveal
more neglect than standard cancellation. Journal of Neurology, Neurosurgery and
Psychiatry, 75, 13561358.