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ABSTRACT: Many everyday situations require focusing on visual or auditory information while ignoring the other modality. Previous findings suggest an attentional mechanism that operates between sensory modalities and governs such states. To date, evidence is equivocal as to whether this 'intermodal' attention relies on a distribution of resources either common or specific to sensory modalities. We provide new insights by investigating consequences of a shift from simultaneous ('bimodal') attention to vision and audition to unimodal selective attention. Concurrently presented visual and auditory stimulus streams were frequency-tagged to elicit steady-state responses (SSRs) recorded simultaneously in electro- and magnetoencephalograms (EEG/MEG). After the shift, decreased amplitudes of the SSR corresponding to the unattended sensory stream indicated reduced processing. We did not observe an amplitude increase of the SSR corresponding to the attended sensory stream. These findings are incompatible with a common-resources account. A redistribution of attentional resources between vision and audition would result in simultaneous processing gain in the attended sensory modality and reduction in the unattended sensory modality. Our results favor a modality-specific-resources account, which allows for independent modulation of early cortical processing in each sensory modality.
NeuroImage 12/2012; · 5.89 Impact Factor
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ABSTRACT: Orienting attention to locations in mnemonic representations engages processes that functionally and anatomically overlap the neural circuitry guiding prospective shifts of spatial attention. The attention-based rehearsal account predicts that the requirement to withdraw attention from a memorized location impairs memory accuracy. In a dual-task study, we simultaneously presented retro-cues and pre-cues to guide spatial attention in short-term memory (STM) and perception, respectively. The spatial direction of each cue was independent of the other. The locations indicated by the combined cues could be compatible (same hand) or incompatible (opposite hands). Incompatible directional cues decreased lateralized activity in brain potentials evoked by visual cues, indicating interference in the generation of prospective attention shifts. The detection of external stimuli at the prospectively cued location was impaired when the memorized location was part of the perceptually ignored hand. The disruption of attention-based rehearsal by means of incompatible pre-cues reduced memory accuracy and affected encoding of tactile test stimuli at the retrospectively cued hand. These findings highlight the functional significance of spatial attention for spatial STM. The bidirectional interactions between both tasks demonstrate that spatial attention is a shared neural resource of a capacity-limited system that regulates information processing in internal and external stimulus representations.
Cerebral Cortex 11/2012; · 6.54 Impact Factor
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ABSTRACT: Previous studies of feature-selective attention have focused on situations in which attention is directed to one of two spatially superimposed stimuli of equal salience. While such overlapping stimuli should maximize stimulus interactions, it is still unknown how bottom-up biases favoring one or the other stimulus influence the efficiency of feature-selective attention. We examined the integration of bottom-up contrast and top-down feature-selection biases on stimulus processing. Two fully overlapping random dot kinematograms (RDKs) of light and dark dots were presented on a gray background of intermediate luminance. On each trial, human participants attended one RDK to detect brief coherent motion targets, while ignoring any events in the unattended RDK. Concurrently, through changes in background luminance, stimulus contrast could be set to five different levels: the stimuli could either be equal, or one of the two stimuli could have twice or four times the contrast of the other stimulus. This manipulation introduced a bottom-up bias toward the stimulus with the higher contrast while keeping the difference between the stimuli constant. Stimulus processing was measured by means of steady-state visual evoked potentials (SSVEPs). SSVEP amplitudes generally increased with higher contrast of the driving stimulus. At earlier levels of processing, attention increased the slope of this linear relation, i.e., attention multiplicatively enhanced SSVEP amplitudes. However, at later levels of processing, attention had an additive effect. These effects of attention can be attributed to the differential integration of gain enhancement and inhibitory stimulus competition at different levels of the visual processing hierarchy.
Journal of Neuroscience 11/2012; 32(47):16953-8. · 7.11 Impact Factor
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ABSTRACT: A better understanding of age-related change in the attentional modulation of perceptual processing may help elucidate cognitive change. For example, increased cognitive interference due to inappropriate processing of irrelevant information has been suggested to contribute to cognitive decline. However, it is not yet clear whether interference effects observed at later stages, such as executive function or response selection, are caused by leaky attentional selection at early, sensory stages of processing. Here, we investigated attentional control of sensory selection by comparing younger and older adults' ability to sustain spatial selective attention to one of two centrally presented, overlapping rapid serial visual presentation (RSVP) letter sequences, one large and one small. These stimuli elicited separable steady-state visual evoked potentials (SSVEP), which provide an index of early visual processing for each stimulus separately and are known to be modulated by selective attention. The condition of most interest required participants to attend to the larger letters while ignoring the smaller letters, as these foveally presented irrelevant stimuli were expected to present the strongest interference. Although the rapid presentation rates made the task demanding, detection ability did not differ between young and older adults. Accordingly, attentional modulation of SSVEP amplitudes was found in both age groups. Neither the magnitude nor the cortical sources of these SSVEP attention effects differed between age groups. Our results thus suggest that in the current task, the effect of voluntary spatial attention on sustained sensory processing in early visual areas is maintained in healthy old age.
Brain research 07/2012; 1469:24-34. · 2.46 Impact Factor
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ABSTRACT: Attention filters behaviorally relevant stimuli from the constant stream of sensory information comprising our environment. Research into underlying neural mechanisms in humans suggests that visual attention biases mutual suppression between stimuli resulting from competition for limited processing resources. As a consequence, processing of an attended stimulus is facilitated. This account makes 2 assumptions: 1) An attended stimulus is released from mutual suppression with competing stimuli and 2) an attended stimulus experiences greater gain in the presence of competing stimuli than when it is presented alone. Here, we tested these assumptions by recording frequency-tagged potentials elicited in early visual cortex that index stimulus-specific processing. We contrasted the processing of a given stimulus when its location was attended or unattended and in the presence or the absence of a nearby competing stimulus. At variance with previous findings, competition similarly suppressed processing of attended and unattended stimuli. Moreover, the magnitude of attentional gain was comparable in the presence or the absence of competing stimuli. We conclude that visuospatial selective attention does not directly modulate mutual suppression between stimuli but instead acts as a signal gain, which biases processing toward attended stimuli independent of competition.
Cerebral Cortex 04/2012; · 6.54 Impact Factor
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ABSTRACT: The focus of attention can be flexibly altered in mnemonic representations of past sensory events. We investigated the neural mechanisms of selection in tactile STM by applying vibrotactile sample stimuli of different intensities to both hands, followed by a symmetrically shaped visual retro-cue. The retro-cue indicated whether the weak or strong sample was relevant for subsequent comparison with a single tactile test stimulus. Locations of tactile stimuli were randomized, and the required response did not depend upon the spatial relation between cued sample and test stimulus. Selection between spatially segregated items in tactile STM was mirrored in lateralized activity following visual retro-cues (N2pc) and influenced encoding of task-irrelevant tactile probe stimuli (N140). Our findings support four major conclusions. First, retrospective selection results in transient shifts of spatial attention. Second, retrospective selection is functionally dissociable from attention-based rehearsal of locations. Third, selection mechanisms are linked across processing stages, as attention shifts in STM influence encoding of sensory signals. Fourth, selection in tactile STM recruits attentional control mechanisms that are, at least partially, supramodal.
Journal of Cognitive Neuroscience 03/2012; 24(7):1596-609. · 5.18 Impact Factor
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ABSTRACT: A number of studies have shown that emotionally arousing stimuli are preferentially processed in the human brain. Whether or not this preference persists under increased perceptual load associated with a task at hand remains an open question. Here we manipulated two possible determinants of the attentional selection process, perceptual load associated with a foreground task and the emotional valence of concurrently presented task-irrelevant distractors. As a direct measure of sustained attentional resource allocation in early visual cortex we used steady-state visual evoked potentials (SSVEPs) elicited by distinct flicker frequencies of task and distractor stimuli. Subjects either performed a detection (low load) or discrimination (high load) task at a centrally presented symbol stream that flickered at 8.6 Hz while task-irrelevant neutral or unpleasant pictures from the International Affective Picture System (IAPS) flickered at a frequency of 12 Hz in the background of the stream. As reflected in target detection rates and SSVEP amplitudes to both task and distractor stimuli, unpleasant relative to neutral background pictures more strongly withdrew processing resources from the foreground task. Importantly, this finding was unaffected by the factor 'load' which turned out to be a weak modulator of attentional processing in human visual cortex.
PLoS ONE 01/2012; 7(5):e37186. · 4.09 Impact Factor
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ABSTRACT: Numerous studies have found that emotionally arousing faces or scenes capture visual processing resources. Here we investigated whether emotional distractor words capture attention in an analogous way. Participants detected brief intervals of coherent motion in an array of otherwise randomly moving squares superimposed on words of positive, neutral or negative valence. Processing of the foreground task was assessed by behavioural responses and steady-state visual evoked potentials (SSVEPs) elicited by the squares flickering at 15 Hz. Although words were task-irrelevant, P2 and N400 deflections to negative words were enhanced, indicating that emotionally negative word content modulated lexico-semantic processing and that emotional significance was detected. In contrast, the time course of behavioural data and SSVEP amplitudes revealed no interference with the task regardless of the emotional connotation of distractor words. This dissociation of emotion effects on early perceptual versus lexical stages of processing suggests that written emotional words do not inevitably lead to attentional modulation in early visual areas. Prior studies have shown a distraction effect of emotional pictures on a similar task. Thus, our results indicate the specificity of emotion effects on sensory processing and semantic encoding dependent on the information channel that emotional significance is derived from.
NeuroImage 12/2011; 60(1):130-8. · 5.89 Impact Factor
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ABSTRACT: A central controversy in the field of attention is how the brain deals with emotional distractors and to what extent they capture attentional processing resources reflexively due to their inherent significance for guidance of adaptive behavior and survival. Especially, the time course of competitive interactions in early visual areas and whether masking of briefly presented emotional stimuli can inhibit biasing of processing resources in these areas is currently unknown. We recorded frequency-tagged potentials evoked by a flickering target detection task in the foreground of briefly presented emotional or neutral pictures that were followed by a mask in human subjects. We observed greater competition for processing resources in early visual cortical areas with shortly presented emotional relative to neutral pictures ~275 ms after picture offset. This was paralleled by a reduction of target detection rates in trials with emotional pictures ~400 ms after picture offset. Our finding that briefly presented emotional distractors are able to bias attention well after their offset provides evidence for a rather slow feedback or reentrant neural competition mechanism for emotional distractors that continues after the offset of the emotional stimulus.
Journal of Neuroscience 11/2011; 31(44):15914-8. · 7.11 Impact Factor
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ABSTRACT: According to the attention-based rehearsal hypothesis, maintenance of spatial information is mediated by covert orienting towards memorized locations. In a somatosensory memory task, participants simultaneously received bilateral pairs of mechanical sample pulses. For each hand, sample stimuli were randomly assigned to one of three locations (fingers). A subsequent visual retro-cue determined whether the left or right hand sample was to be memorized. The retro-cue elicited lateralized activity reflecting the location of the relevant sample stimulus. Sensory processing during the retention period was probed by task-irrelevant pulses randomized to locations at the cued and uncued hand. The somatosensory N140 was enhanced for probes delivered to the cued hand, relative to uncued. Probes presented shortly after the retro-cue showed greatest attentional modulations. This suggests that transient contributions from retrospective selection overlapped with the sustained effect of attention-based rehearsal. In conclusion, focal attention shifts within tactile mnemonic content occurred after retro-cues and guided sensory processing during retention.
Biological psychology 09/2011; 89(1):39-46. · 4.36 Impact Factor
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ABSTRACT: Intermodal attention (IA) is assumed to allocate limited neural processing resources to input from one specific sensory modality. We investigated effects of sustained IA on the amplitude of a 40-Hz auditory (ASSR) and a 4.3-Hz visual steady-state response (VSSR). To this end, we concurrently presented amplitude-modulated multi-speech babble and a stream of nonsense letter sets to elicit the respective brain responses. Subjects were cued trialwise to selectively attend to one of the streams for several seconds where they had to perform a lexical decision task on occasionally occurring words and pseudowords. Attention to the auditory stream led to greater ASSR amplitudes than attention to the visual stream. Vice versa, the VSSR amplitude was greater when the visual stream was attended. We demonstrate that IA research by means of frequency tagging can be extended to complex stimuli as used in the current study. Furthermore, we show not only that IA selectively modulates processing of concurrent multisensory input but that this modulation occurs during trial-by-trial cueing of IA. The use of frequency tagging may be suitable to study the role of IA in more naturalistic setups that comprise a larger number of multisensory signals.
Experimental Brain Research 04/2011; 213(2-3):321-7. · 2.39 Impact Factor
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ABSTRACT: Selective attention to a primary task can be biased by the occurrence of emotional distractors that involuntary attract attention due to their intrinsic stimulus significance. What is largely unknown is the time course and magnitude of competitive interactions between a to-be-attended foreground task and emotional distractors. We used pleasant, unpleasant and neutral pictures from the International Affective Picture System (IAPS) that were either presented in intact or phase-scrambled form. Pictures were superimposed by a flickering display of moving random dots, which constituted the primary task and enabled us to record steady-state visual evoked potentials (SSVEPs) as a continuous measure of attentional resource allocation directed to the task. Subjects were required to attend to the dots and to detect short intervals of coherent motion while ignoring the background pictures. We found that pleasant and unpleasant relative to neutral pictures more strongly influenced task-related processing as reflected in a significant decrease in SSVEP amplitudes and target detection rates, both covering a time window of several hundred milliseconds. Strikingly, the effect of semantic relative to phase-scrambled pictures on task-related activity was much larger, emerged earlier and lasted longer in time compared to the specific effect of emotion. The observed differences in size and duration of time courses of semantic and emotional picture processing strengthen the assumption of separate functional mechanisms for both processes rather than a general boosting of neural activity in favor of emotional stimulus processing.
NeuroImage 12/2010; 53(4):1326-33. · 5.89 Impact Factor
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ABSTRACT: Multiple concurrently presented stimuli are thought to compete for neuronal processing resources. Such competitive stimulus interactions can be investigated by "frequency tagging" each stimulus with an individual temporal frequency. In this case, all stimuli will drive distinct steady-state visual evoked potentials (SSVEPs), hence allowing for an assessment of the distribution of processing resources. Here, we investigated whether competitive effects on SSVEP amplitudes are dependent upon the choice of tagging frequency of either the driving stimulus or a close-by competing stimulus. In particular, we were interested whether changes in amplitude are specific to a 10-Hz SSVEP, as it has been suggested that tagging frequencies within the alpha band drive uniquely characterized neural networks. If this was the case, an additional competition might be introduced when two stimuli are tagged with frequencies within the alpha band and thus compete for processing resources in similar networks. Additionally, we tested whether effects on SSVEP amplitude differ when the competing stimulus is tagged with a frequency of 12 Hz that produces a perceptible flicker when compared to an imperceptible 60-Hz flicker. We found a significant decrease in amplitude of 10- and 15-Hz SSVEPs upon presentation of the competing stimulus regardless of its tagging frequency. Our results clearly indicate that an SSVEP with a frequency within the alpha band and a 15-Hz SSVEP show similar sensitivity to effects of competition. Furthermore, the observed effects of competition on SSVEP amplitude occur independently of flicker perceptibility.
Experimental Brain Research 09/2010; 205(4):489-95. · 2.39 Impact Factor
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ABSTRACT: Recent studies have suggested that the extent to which primary task demands draw on attentional resources determines whether or not task-irrelevant emotional stimuli are processed. Another important factor that can bias task-relevant and task-irrelevant stimulus competition is the bottom-up factor of stimulus salience. Here, we investigated the effect of stimulus salience associated with a primary motion task on the processing of emotional face distractors. Faces of different emotional valences were presented within a context of randomly moving dots. Subjects had to detect short intervals of coherent motion while ignoring the background faces. Task salience was manipulated by the level of motion coherence of the dots with high motion coherence being associated with high salience. Using functional magnetic resonance imaging, we show that emotional faces, compared with neutral faces, more strongly interfered with the primary task, as reflected in significant signal decreases in task-related motion area V5/hMT+. In addition, these faces elicited significant signal increases in the left amygdala. Most importantly, task salience was found to further increase amygdala's activity when presented together with an emotional face. Our data support a more general role of the amygdala as a behavioral relevance detector, which flexibly integrates behavioral relevant, salient context information to decode the emotional content of a visual scene.
Journal of Neuroscience 04/2010; 30(15):5204-10. · 7.11 Impact Factor
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ABSTRACT: Although attention in older adults is an active research area, feature-selective aspects have not yet been explicitly studied. Here we report the results of an exploratory study involving directed changes in feature-selective attention. The stimuli used were two random dot kinematograms (RDKs) of different colours, superimposed and centrally presented. A colour cue with random onset after the beginning of each trial instructed young and older subjects to attend to one of the RDKs and detect short intervals of coherent motion while ignoring analogous motion events in the non-cued RDK. Behavioural data show that older adults could detect motion, but discriminated target from distracter motion less reliably than young adults. The method of frequency tagging allowed us to separate the EEG responses to the attended and ignored stimuli and directly compare steady-state visual evoked potential (SSVEP) amplitudes elicited by each stimulus before and after cue onset. We found that younger adults show a clear attentional enhancement of SSVEP amplitude in the post-cue interval, while older adults' SSVEP responses to attended and ignored stimuli do not differ. Thus, in situations where attentional selection cannot be spatially resolved, older adults show a deficit in selection that is not shared by young adults.
Neuroscience Letters 02/2010; 474(1):5-8. · 2.11 Impact Factor
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ABSTRACT: This study examined the latency and amplitude of cortical processes associated with feature-based visual selective attention, using frequency-domain and time-domain measures derived from dense-array electroencephalography. Participants were asked to identify targets based on conjunctions of three types of object features (color, size, and completeness). This procedure aimed to examine (1) the modulation of sensory responses to one or more stimulus features characterizing an object and (2) the facilitation and reduction effects associated with competing features, attended and unattended, in the same object. The selection negativity, an event-related potential measure of sensory amplification for attended features, showed a parametric increase of amplitude as a function of the number of attended features. Late oscillations in the gamma band range were also smaller for stimuli with one or more non-attended visual features but were enhanced for stimuli sharing the overall gestalt with the target. The latency of this late gamma modulation was delayed when two target features were combined, compared to one single discriminative feature. Latency analyses also showed that late bursts of induced high-frequency oscillatory activity peaked around 60 ms later than the selection negativity. Oscillatory activity reflected both selective amplification and competition between object features. These results suggest that sensory amplification of selected features is followed by integrative processing in more widespread networks. Oscillatory activity in these networks is reduced by distraction and is enhanced when attended features can be mapped to specific action.
Brain research 12/2009; 1313:172-84. · 2.46 Impact Factor
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ABSTRACT: We investigated mechanisms of concurrent attentional selection of location and color using electrophysiological measures in human subjects. Two completely overlapping random dot kinematograms (RDKs) of two different colors were presented on either side of a central fixation cross. On each trial, participants attended one of these four RDKs, defined by its specific combination of color and location, in order to detect coherent motion targets. Sustained attentional selection while monitoring for targets was measured by means of steady-state visual evoked potentials (SSVEPs) elicited by the frequency-tagged RDKs. Attentional selection of transient targets and distractors was assessed by behavioral responses and by recording event-related potentials to these stimuli. Spatial attention and attention to color had independent and largely additive effects on the amplitudes of SSVEPs elicited in early visual areas. In contrast, behavioral false alarms and feature-selective modulation of P3 amplitudes to targets and distractors were limited to the attended location. These results suggest that feature-selective attention produces an early, global facilitation of stimuli having the attended feature throughout the visual field, whereas the discrimination of target events takes place at a later stage of processing that is only applied to stimuli at the attended position.
Journal of Cognitive Neuroscience 09/2009; 23(1):238-46. · 5.18 Impact Factor
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ABSTRACT: We investigated the impact of task difficulty on neuronal facilitation of to-be-attended somatosensory vibrotactile stimuli and transient tactile distracters during sustained attention. In Experiment 1, we employed an easy detection task and in Experiment 2 a challenging discrimination task. Sustained attention was manipulated by presenting vibrotactile stimuli simultaneously to the index fingers of both hands for a period of 3s. These stimuli elicited the steady-state somatosensory evoked potential (SSSEP). Subjects attended to one body side and had to detect target-stimuli that were embedded in the ongoing vibratory streams. When subjects discriminated target-stimuli we found increased SSSEP amplitudes of the to-be-attended vibrotactile stream and greater N140 amplitudes of the somatosensory evoked potential (SEP). During stimulus detection no such facilitation of the SSSEP was found. Greater N140 amplitudes elicited by to-be-ignored distracters indicated that they pull attention to that body location under conditions of low load.
Biological psychology 06/2009; 81(2):77-85. · 4.36 Impact Factor
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ABSTRACT: The purpose of this study was to test whether color-motion correlations carried by a pure color difference (S-cone component only) can be used to improve global motion extraction. We also examined the neural markers of color-motion correlation processing in event-related potentials. Color and motion information was dissociated using a two-colored random dot kinematogram, wherein coherent motion and motion noise differed from each other only in their S-cone component, with spatial and temporal parameters set so that global motion processing relied solely on a constant L-M component. Hence, when color and the local motion direction are correlated, more efficient segregation of coherent motion can only be brought about by the S-cone difference, and crucially, this S-cone component does not provide any effective input to a global motion mechanism but only changes the color appearance of the moving dots. The color contrasts (vector length in the S vs. L-M plane) of both the dots carrying coherent motion and the dots moving randomly were fixed at motion discrimination threshold to ensure equal effectiveness for motion extraction. In the behavioral experiment, participants were asked to discriminate between coherent and random motion, and d' was determined for three different conditions: uncorrelated, uncued correlated, and cued correlated. In the electroencephalographic experiment, participants discriminated direction of motion for uncued correlated and cued correlated conditions. Color-motion correlations were found to improve performance. Cueing a specific color also modulated the N1 component of the event-related potential, with sources in visual area middle temporal. We conclude that S-cone signals "invisible" to the motion system can influence the analysis by direction-selective motion mechanisms through grouping of local motion signals by color. This grouping mechanism must precede motion processing and is likely to be under attentional control.
Visual Neuroscience 05/2009; 26(2):237-48. · 2.23 Impact Factor
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ABSTRACT: The aim of the present study was to simultaneously measure and compare intermodal attention effects in event-related brain potentials (ERPs) and auditory steady-state responses (ASSRs). For this purpose, 40-Hz amplitude modulated tones and a visual fixation cross were presented concurrently. By means of target detection tasks either on the sounds or on the fixation cross, participants' attention was directed to the respective modality. Attended sounds elicited a negative difference (Nd) in the ERP relative to unattended sounds. Nd was divided into an early and a late part as often observed for intramodal attention. Moreover, attention to the sounds led to a significant enhancement of the ASSR. This modulation of the ASSR by intermodal attention is demonstrated for the first time in the EEG. The present data suggest that ASSRs could provide a useful tool for the investigation of the neural dynamics of intermodal attentional processes.
Psychophysiology 02/2009; 46(2):321-7. · 3.29 Impact Factor
