Can attention selectively bias bistable perception? Differences between binocular rivalry and ambiguous figures

“…competition model of selective attention, concurrent visual inputs compete for representation across a network of visual areas, and attention serves to enhance the neural representation of attended stimuli and to suppress the neural representation of unattended stimuli (Desimone & Duncan, 1995).”

“A growing number of psychophysical, physiological, and neuroimaging studies have relied on bistable phenomena to investigate perceptual selection and the neural correlates of visual awareness (for reviews, see Blake & Logothetis, 2002; Tong, 2003).”

“Feature and object-based attention can effectively bias perception under normal, dioptic viewing conditions (for a review see Kanwisher & Wojciulik, 2000), and could conceivably facilitate attentional selection during rivalrous dichoptic viewing.”

“…evidence suggests that binocular rivalry also involves competition between high-level pattern representations, similar to what is presumed to occur in ambiguous figure reversal (Kovacs, Papathomas, Yang, & Feher, 1996; Logothetis, Leopold, & Sheinberg, 1996; Sheinberg & Logothetis, 1997).”

“There is also considerable psychophysical and neuroimaging evidence to suggest that binocular rivalry results from low-level interocular competition between monocular channels (Blake, 1989; Blake, Westendorf, & Overton, 1980; Nguyen, Freeman, & Wenderoth, 2001; Polonsky, Blake, Braun, & Heeger, 2000; Tong & Engel, 200…”

“A growing number of psychophysical, physiological, and neuroimaging studies have relied on bistable phenomena to investigate perceptual selection and the neural correlates of visual awareness (for reviews, see Blake & Logothetis, 2002; Tong, 2003).”

“…psychophysical and neuroimaging evidence to suggest that binocular rivalry results from low-level interocular competition between monocular channels (Blake, 1989; Blake, Westendorf, & Overton, 1980; Nguyen, Freeman, & Wenderoth, 2001; Polonsky, Blake, Braun, & Heeger, 2000; Tong & Engel, 2001).”

“…psychophysical and neuroimaging evidence to suggest that binocular rivalry results from low-level interocular competition between monocular channels (Blake, 1989; Blake, Westendorf, & Overton, 1980; Nguyen, Freeman, & Wenderoth, 2001; Polonsky, Blake, Braun, & Heeger, 2000; Tong & Engel, 2001).”

“In contrast, binocular rivalry occurs automatically for most observers with normal binocular vision (Blake, 2001).”

“Our observers could more than double their rate of fast alternations as compared to slow alternations when viewing highcontrast rivalry stimuli, matching the level of control found in previous studies (Lack, 1971; Lack, 1978; Meredith, 1962).”

“Previous studies have shown evidence of selective attentional control over ambiguous figure reversal (Gomez, Argandona, Solier, Angulo, & Vazquez, 1995; Horlitz & O'Leary, 1993; Liebert & Burk, 1985; Peterson, 1986; Toppino, 2003).”

“Moreover, people show less attentional control of alternation rates for the Necker cube than for reversible figures that have multiple object interpretations (e.g., duck/rabbit, chef/dog) (Struber & Stadler, 1999).”

“Rock and Mitchener (1992) found that only one third of naïve observers reported spontaneous reversals for ambiguous figures, and that knowledge of reversibility helped naïve observers to perceive ambiguous figure reversals (see also Girgus, Rock, & Egatz, 1977).”

“Binocular rivalry involves fluctuations in the phenomenal visibility of low-level features, whereas in ambiguous figure reversal, the low-level features remain intact while the high-level organization of those features changes over time (Tong, 2001).”

“The Necker cube was chosen because it is one of the most commonly studied ambiguous figures (George, 1936; Gomez et al. 1995; Horlitz & O'Leary, 1993; Kawabata, Yamagami, & Noaki, 1978; Long & Toppino, 1981; Long, Toppino, & Mondin, 1992; Necker, 1832; Toppino, 2003).”

“Previous studies have shown evidence of selective attentional control over ambiguous figure reversal (Gomez, Argandona, Solier, Angulo, & Vazquez, 1995; Horlitz & O'Leary, 1993; Liebert & Burk, 1985; Peterson, 1986; Toppino, 2003).”

“Along these lines, more recent studies have found evidence of selective attentional control over ambiguous figure reversal (Gomez et al., 1995; Horlitz & O'Leary, 1993; Liebert & Burk, 1985; Peterson, 1986; Toppino, 2003), consistent with the present findings.”

“perception of the Necker cube to a small extent (Kawabata et al., 1978; Toppino, 2003).”

“However, a significant interaction was found for the top view (F(4, 20) = 4.99, p .01), similar to what a previous study found in bistable apparent motion (Suzuki & Peterson, 2000 ).”

“The Necker cube was chosen because it is one of the most commonly studied ambiguous figures (George, 1936; Gomez et al. 1995; Horlitz & O'Leary, 1993; Kawabata, Yamagami, & Noaki, 1978; Long & Toppino, 1981; Long, Toppino, & Mondin, 1992; Necker, 1832; Toppino, 2003).”

“Previous studies have shown evidence of selective attentional control over ambiguous figure reversal (Gomez, Argandona, Solier, Angulo, & Vazquez, 1995; Horlitz & O'Leary, 1993; Liebert & Burk, 1985; Peterson, 1986; Toppino, 2003).”

“Our observers could more than double their rate of fast alternations as compared to slow alternations when viewing highcontrast rivalry stimuli, matching the level of control found in previous studies (Lack, 1971; Lack, 1978; Meredith, 1962).”

“But can observers control subsequent alternations of rivalry following initial dominance? Observers can indeed exert some selective attentional control over what they perceive during steady rivalry viewing [ 65 ,69], and this control can be further amplified if observers are given a challenging visual task requiring f…”

“It is known that both phenomena are largely independent of attention: Rivalry switches cannot be fully suppressed at will (Meng & Tong, 2004 ), and visual illusions are the prime example of the cognitive impenetrability of visual perception (Pylyshyn, 1999 ).”

“…analyzing and synthesizing basic components of objects (Kosslyn & Koenig, 1992; Michelon & Koenig, 2002), including focal areas, critical features (Long & Olszweski, 1999), fixation points (Meng & Tong, 2004; Toppino, 2003), and spatial proximity or crowding (Pelli, Palomares, & Majaj, 2004).”

“figures, it is difficult to wilfully influence perceptual alternations in binocular rivalry [ 77 ].”

“It has been shown that subjects can willfully change the switching rate in rivalry (Lack 1969) and that attention to one of the monocular stimuli producing rivalry slightly shortens the intervals for which the unattended stimulus is dominant (Meng and Tong 2004).”

“The number of switches [13], proportion of Grouped percept, and effect of volitional control [30] were compared for each subject across the two modalities in the present data.”

“Using a different approach, Meng and Tong (2004) compared the perception of ambiguous figures to the more stimulus-driven processing involved in other forms of bistable perception (e.g.”

“In the visual domain , the alternation of some bistable percepts can be influenced by selec - tive attention ( Meng and Tong , 2004 ) , but spontaneous switches still occur .”

“Effortful attempts to control rivalry, for example paying more attention to one percept in order to prevent its alternating with the rival percept, fail during sustained rivalry (Meng and Tong, 2004).”

“This latter option would be consistent with psychophysical findings showing that directed attention modulates the frequency of perceptual alternations (Meng and Tong, 2004; van Ee et al., 2005; Chong et al., 2005; Paffen et al., 2006).”

“6 After completion of this paper we learned about a recent useful paper ( Meng & Tong, in press ).”

“Selective attention is also necessary to see the alternative image in an ambiguous figure (see Leopold & Logothetis, 1999, for a review; Meng & Tong, 2004).”

“It has been shown that subjects can willfully change the switching rate in rivalry (Lack 1969) and that attention to one of the monocular stimuli producing rivalry slightly shortens the intervals for which the unattended stimulus is dominant (Meng and Tong 2004).”

“2005), and between 5 and 10 s in the ambiguous depth perception (Meng and Tong 2004; Zhou et al. 2004; Moreno-Bote et al. 2008) and ambiguous motion displays (Hupe and Rubin 2003).”

“Increasing the difference in stimulus strength between the two eyes will primarily act to increase the average perceptual dominance duration of the stronger stimulus.”

“Recently, Meng & Tong (2004) showed that although attention can change alternation rates, its effect on dominance duration during rivalry is negligible.”

“(Meng and Tong 2004), which makes them only conditionally comparable.”

“The extent to which all these forms of rivalry exhibit the same phenomenology is subject to debate [13,14,15,16,17] as is the neural origin of rivalry.”

“…to the passive conditions (from 2.6 0.2 s to 3.6 0.3 for looming, and from 2.3 0.2 s to 3.0 0.3 s for radial patterns, both p 0.001, paired t test), replicating previous work (Lack, 1978; Toppino, 2003; Meng and Tong, 2004; Chong et al., 2005; Slotnick and Yantis, 2005; van Ee et al., 2005).”

“Each object (average width: 2.79°; average height: 2.31°) was displayed within a tinted square (green or red; width: 3.5°), which was surrounded by a black circle to support binocular fusion (radius: 4.95°; Meng and Tong, 2004).”

“In fact, such differences have been found in the extent to which subjects can voluntarily control the rivalry process (Meng & Tong, 2004; Van Ee et al., 2005).”

“During continuous presentation, one aspect lasts on average for around 3 s before a switch in perspectives occurs, with some inter-individual variability and variance attributable to stimulus characteristics of the particular ambiguous figure (Gómez et al., 1995; von Steinbüchel et al., 1999; Meng and Tong, 2004; Korn…”

“We know, for example, that attention can influence the rate of alternations during rivalry (Chong, Tadin, & Blake, in press; Lack, 1978; Meng & Tong, 2004; Meredith & Meredith, www.elsevier.com/locate/visres Vision Research 46 (2006) 1794–1803 0042-6989/$ -see front matter Ó 2005 Elsevier Ltd.”

“Several studies have examined the role of attention in binocular rivalry (Lack, 1978; Meng & Tong, 2004; Mitchell, Stoner, & Reynolds, 2004; Ooi & He, 1999; van Ee, 2005).”

“For example, a previous psychophysical study found that bistable perception elicited by ambiguous images is more susceptible to attentional modulation than that elicited by binocular rivalry and thus may have a stronger top-down component (89).”

“However, binocular rivalry can also be modified by shifts in attention (Meng & Tong, 2004): if participants focus their attention on specific stimuli, these stimuli are slightly more likely to enter awareness than unattended stimuli.”

“, 2006), with attention (Meng and Tong, 2004; Mitchell et al., 2004; Paffen et al., 2006; Pastukhov and Braun, 2007), and when a display is periodically interrupted (Orbach et al.”

“Whereas voluntary control over continuously presented bistable stimuli has been known for some time (Meng & Tong, 2004; van Ee et al., 2005), it has never been shown for percept choices at stimulus onset.”

“Interestingly, the influences of feedback that we find mirror those found previously for attentional control (Chong, Tadin, & Blake, 2005; Meng & Tong, 2004; van Ee, van Dam, & Brouwer, 2005), in that attention increases dominance durations of the attended stimulus, while leaving the dominance…”

“paper ( Meng & Tong, in press ) that focused on the differences in data produced by three rivalry paradigms: grating, house-face and Necker cube.”

“…by local adaptation (Chen & He, 2004; Blake, Sobel, & Gilroy, 2003), and presumably by feature- or object-based attention (Chong & Blake, 2005; Meng & Tong, 2004; Mitchell, Stoner, & Reynolds, 2004) and other less well-explored factors such as slow drifts in contrast sensitivity and…”

“If confirmed to be true in future studies, this would be a serious advantage for the discrete method because different bistable stimuli are affected differently by attention (e.g., Lack, 1978; Meng & Tong, 2004; van Ee, van Dam, & Brouwer, 2005).”

“For example, variables that influence perceptual stability like eye position (Einhauser, Martin and Konig, 2004; Ellis and Stark, 1978; Kawabata, 1978) and attention (van Ee, Noest, Brascamp and van den Berg, 2006; Hol, Koene and van Ee, 2003; van Ee, van Dam and Brower , 2005; Toppino, 2003; Meng and Tong, 2004) are…”

“Some bistable phenomena, such as the perceptual flipping in binocular rivalry appear not to be mod- ulated by selective attentional control, whereas others, such as the Necker cube, can be modulated by voluntary selective attention (Meng & Tong, 2004).”

“Keywords: Perceptual rivalry; Binocular rivalry; Voluntary perceptual control; Saccades”

“In recent years, studies by Meng and Tong (2004) and van Ee et al. (2005) have revived the issue of voluntary control and reach a similar conclusion to that of Lack (1978): control over binocular rivalry is possible.”

“Although binocular rivalry is highly resistant to volitional control, percept dominance and alternation rate are both sensitive to endogenous attentional allocation (Lack 1974; Ooi and He 1999; Meng and Tong 2004; Chong et al. 2005; van Ee et al. 2005; Paffen et al. 2006; Alais et al. 2010).”

“Subtle experimental manipulations, such as attention (Meng and Tong, 2004; Mitchell et al., 2004; Chong and Blake, 2006; Kamphuisen et al., 2007), sensory memory (Pearson and Brascamp, 2008), or imagery (Pearson et al., 2008), can bring about a slight imbalance in the neural states, creating a bias…”

“Even for the same observer and same display, dominance times vary substantially with stimulus intensity [16,17], with attention [18–21], and when a display is periodically interrupted [22–24].”

“All patients and control participants were included in an analysis that compared the relative percentage of partic- ipants who were able to recognize the second views of the ambiguous stimulus patterns within 60 sec.”

“In the average PD analysis, to resolve inter- and intraindividual variations we normalized the average PDs (see, e.g., Meng and Tong 2004) to the third SFM trial per task block (see Materials and Methods).”

“Indeed, those kinds of stimulus influences are considerably more robust than are influences attributable to nonsensory factors, such as attention (e.g., Lack, 1978; Meng & Tong, 2004 ), meaning (e.g., Blake, 1988), and affective connotation (e.g., Alpers & Pauli, 2006).”

“Consistent with this possibility, shifts of attention have been found to influence dominance (Meng & Tong, 2004; Mitchell et al., 2004) and to modulate overall reversal rates (van Ee et al., 2005), although the degree of attentional influence varied between studies.”

“Previous research suggests that binocular rivalry is less susceptible to top–down modulation than ambiguous figure perception [Meng and Tong, 2004]: subjects can more easily control their perception of ambiguous figures than of binocularly rivaling stimuli when instructed to do so.”

“The stimuli employed during binocular rivalry have been shown to fall under the category of ambiguous figures [36], and the alternation rate of binocular rivalry is under voluntary control, albeit to a substantially lesser degree than bistable perceptual figures [41].”

“…differences (Blake & Logothetis, 2002), stimulus familiarity (Jiang et al. 2007), cross-modal sensory influences (Zhou et al. 2010) and endogenous factors directly related to the task currently at hand, such as endogenous attention or learned expectations (Meng & Tong, 2004 ; Sterzer et al. 2008a).”

“In contrast, when the same subjects were asked to perform an analogous task with the bistable Necker cube, strong attentional modulation was observed (Figure 4; Meng and Tong, 2004; also see Toppino, 2003).”

“This has been demonstrated previously for various distinct ambiguous stimuli (e.g., Hol et al., 2003; Lack, 1978; Meng & Tong, 2004; Peterson, 1986; Suzuki & Peterson, 2000; Toppino, 2003; van Ee et al., 2005).”

“Two visual factors, one high-level (selective attention) and one lowlevel (stimulus contrast) which have been shown to modulate visual perception in the binocular rivalry situation (Meng and Tong, 2004), are used to probe behaviorally the underlying mechanisms by which the auditory semantic context modulating visual a…”

“, 2007 ; Meng & Tong , 2004 ) .”

“, attention, semantics, context) modulate only the perceptually dominant stimulus (Sobel and Blake, 2002; Meng and Tong, 2004; Chong et al., 2005; but see Hancock and Andrews, 2007; Paffen and Alais, 2012).”

“are weaker for binocular rivalry than for the Necker cube (Meng and Tong 2004) and that the degree of volitional control that biases perceptual dominance is different for auditory”

“Voluntary control and controlled attention do not influence which image is consciously seen (Meng & Tong, 2004), although imagining an object increases its dominance (Pearson, Clifford, & Tong, 2008).”

“Of course, there are likely to be some differences between the rivalry that relies on monocularly incompatible images, and ambiguous images that rely on incompatible binocular percepts, so results may differ between these phenomena in terms of lower-tier areas such as V1, in which the monocular images are combined for…”

“…an ambiguous display the rate of perceptual alternations may be modulated by stimulus parameters (e.g. Brouwer & van Ee, 2006; Klink, van Ee, & van Wezel, 2008; Levelt, 1966) and also by factors such as attention (Chong, Tadin, & Blake, 2005; Meng & Tong, 2004; van Ee, van Dam, & Brouwer, 2005).”

“Top–down signals, such as those associated with selective attention, can bias this neural competition in favor of one perceptual interpretation over another (Meng and Tong, 2004).”

“This effect is similar of that of attention on binocular rivalry [19,20] in which attentional selection of one image tends to prolong dominance of the attended stimulus but generally does not shorten the suppression period ([19,21]; but see also [22]).”

“…caused by mechanisms acting in a feedback fashion on lower level sensory mechanisms (e.g., Georgiades & Harris, 1997; Horlitz & O'Leary, 1993; Kawabata, 1986; Leopold, 2003; Meng & Tong, 2004; Pelton & Solley, 1968; Rock, Hall, & Davis, 1994; Shulman, 1993; Struber & Stadler, 1999; Toppino, 2003).”

“…and Chow, 2011) and, consistently with the outcome of Scocchia and colleagues, previous studies comparing binocular rivalry to other instances of bistable perception showed less amenability of rivalry to top-down modulation (i.e., Washburn and Gillette, 1933; George, 1936; Meng and Tong, 2004).”

“2b in terms of sensitivity (d0), for the ‘easy’ and ‘difficult’ levels.”

“, 2007), an increased intentional effort to speed perceptual switches (e.g., Lack, 1974, 1978; Meng and Tong, 2004; van Ee et al., 2005), or a combination of these factors (potentially mediated by feedback signals from frontal and prefrontal cortexes; e.”

“, 2011), binocular rivalry is often only weakly susceptible to cognitive control (Meng and Tong, 2004).”

“In their study, Meng and Tong (2004) failed to find a reliable effect of attention with an AM of between 5% and 13%.”

“…field, context of stimuli, etc.; Blake & Logothetis, 2002) and, less clearly understood, on factors intrinsic to observers, such as attention (Meng & Tong, 2004; Paffen, Alais, & Verstraten, 2006) and serotonin levels in the brain (Carter, Pettigrew et al., 2005; Nagamine, Yoshino, Miyazaki,…”

“…dominance during binocular rivalry, meaning that subjects are able to exert some attentional control over what they perceive during rivalry (Meng & Tong, 2004; Mitchell, Stoner, & Reynolds, 2004; Ooi & He, 1999), in particular if they are given a visual task that requires focused attention…”

“…can be controlled voluntarily (e.g. Horlitz & O’Leary, 1993; Kawabata, 1986; Leopold & Logothetis, 1999; Liebert & Burk, 1985; Long & Toppino, 2004; Meng & Tong, 2004; Pelton & Solley, 1968; Rock et al., 1994; Shulman, 1993; Struber & Stadler, 1999; Toppino, 2003; van Ee, 2005; van Ee et al.,…”

“C and Video S1): KD in a two-dimensional projection of a rotating cloud of dots (Wallach and O’Connell, 1953), BR between two gratings of different color and orientation (Wheatstone, 1838; Meng and Tong, 2004), and the NC (Necker, 1832).”

“…different, kinds of multistable displays (Figure 1A and Movie 1): kinetic depth (KD) in a two-dimensional projection of a rotating cloud of dots (Wallach & O’Connell, 1953) and binocular rivalry (BR) between two gratings of different color and orientation (Meng & Tong, 2004; Wheatstone, 1838).”

“These include attention (Meng and Tong, 2004; van Ee, 2005), context (Fukuda and Blake, 1992; Carter et al., 2004; Paffen et al., 2004) and manipulating stimulus strength by increasing the contrast (Mueller and Blake, 1989), motion (Breese, 1909) and spatial frequency (Fahle, 1982) of one of the…”

“…phenomena including perception of ambiguous auditory stimuli (e.g., Einhäuser, Stout, Koch, & Carter , 2008), perception of traditional visual ambiguous figures such as the Necker cube (e.g., Meng & Tong, 2004), perception of illusory 0042-6989/$ -see front matter Ó 2009 Elsevier Ltd.”

“In the case of binocular rivalry, the dominance durations are only minimally affected by voluntary control (Meng and Tong, 2004; Chong et al., 2005; Klink et al., 2008), yet attention is critical for its manifestation (Chong and Blake, 2006; Kohler et al., 2008; Dieter and Tadin, 2011; Zhang et…”

“Meng and Tong (2004) compared the amount of control for different types of binocular rivalry and the Necker cube.”

“trol or attention (Meng and Tong, 2004).”

“Selective attentional control over perceptual alternations in binocular rivalry is considerably weaker than control over alternations in other forms of bistable perception, such as Necker cube or face/house reversals (Meng & Tong, 2004; van Ee, van Dam, & Brouwer, 2005).”

“For instance, voluntary control of subjects over reversal rates seems to be pronounced for ambiguous stimuli (van Ee et al. 2005) but quite limited for binocular rivalry (van Ee 2005, Meng and Tong 2004).”

“But there appears to be also a role of specific binocular rivalry based mechanisms in that it seems to be harder to control binocular rivalry than figural rivalry (cf. Blake, 1988; George, 1936; Meng & Tong, 2004; van Ee et al., 2005; Washburn & Gillette, 1933).”

“…immediate effect on binocular rivalry : Attention can bias the competition toward one of the rival targets ( Chong , Tadin , & Blake , 2005 ; Lack , 1978 ; Meng & Tong , 2004 ; van Ee , van Dam , & Brouwer , 2005 ) as well as change its temporal dynamics ( Paffen , Alais , & Verstraten , 2006 ) .”