Early Event-Related Potential Latency Shifts Underlying Prior Entry in A
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Temporal Order is Coded Temporally in the Brain: Early Event-related Potential Latency Shifts Underlying Prior Entry in a Cross-modal Temporal Order Judgment Task Downloaded from http://mitprc.silverchair.com/jocn/article-pdf/19/1/109/1756371/jocn.2007.19.1.109.pdf by guest on 18 May 2021 J. Vibell, C. Klinge, M. Zampini, C. Spence, and A. C. Nobre Abstract & The speeding-up of neural processing associated with at- potentially lacking the sensitivity to reveal putative modula- tended events (i.e., the prior-entry effect) has long been tions in the timing of neural processing. In the present study, proposed as a viable mechanism by which attention can pri- we used a cross-modal temporal order judgment task while oritize our perception and action. In the brain, this has been shifting attention between the visual and tactile modalities thought to be regulated through a sensory gating mechanism, to investigate the mechanisms underlying selective attention increasing the amplitudes of early evoked potentials while electrophysiologically. Our results indicate that attention can leaving their latencies unaffected. However, the majority of indeed speed up neural processes during visual perception, previous research has emphasized speeded responding and thereby providing the first electrophysiological support for has failed to emphasize fine temporal discrimination, thereby the existence of prior entry. & INTRODUCTION reach consciousness earlier (e.g., see Zampini, Shore, In his attempt to summarize the most important ques- & Spence, 2005; Schneider & Bavelier, 2003; Spence, tions in attention research nearly a century ago, Titchener Shore, & Klein, 2001). (1908) formulated seven fundamental laws of atten- The law of prior entry has typically been investigated tion. The fourth law, the law of prior entry, states that using unspeeded temporal order judgment (TOJ) tasks attended stimuli come into consciousness more rapidly (see Shore & Spence, 2005, for a recent review). Unlike than unattended stimuli. The law was based on Geiger’s reaction time (RT) studies, TOJs do not require a (1903) pendulum experiment, whereby participants per- speeded response, but instead typically involve partic- ceived a pendulum at different locations when a sound ipants making nonspeeded perceptual judgments. A was presented depending on whether their attention typical TOJ trial consists of two stimuli separated in time was directed to the sound or to the sight of the by a small but variable stimulus onset asynchrony (SOA). pendulum itself (see also Mollon & Perkins, 1996). On the basis of a participant’s decisions about which Geiger’s experiment generated much debate and several stimulus appeared first at each SOA, the mean point different explanations have been put forward. Most of subjective simultaneity (PSS) can be calculated. The notably, contributions of an anticipation bias or physio- PSS indicates by how much time one stimulus has to logical factors (auditory processing being faster than lead the other in order for the two to be judged as oc- visual processing) were proposed, which were refuted curring simultaneously (i.e., the average SOA at which by both Titchener and James (see Titchener, 1908). participants make each response equally often). Because Instead, Titchener suggested that the time of conscious the accuracy of temporal discrimination performance is perception of a stimulus could vary within a brief tem- stressed in TOJ studies rather than the speed of re- poral interval depending upon the current focus of an sponding, behavioral modulations in the PSS from cuing observer’s attention. In the century since Titchener are thought to result from early perceptual modulations formulated his law of prior entry, a great deal of con- of sensory information processing by attention, making troversy has been generated as to whether attention them ideal for the study of prior entry. Early behavioral actually speeds up the processing of stimuli so that they studies provided mixed evidence for the law of prior entry (see Spence, Shore, et al., 2001, for a comprehen- sive review). Early research faced extensive criticism, University of Oxford most notably that the prior-entry effects in many studies D 2007 Massachusetts Institute of Technology Journal of Cognitive Neuroscience 19:1, pp. 109–120 Downloaded from http://www.mitpressjournals.org/doi/pdf/10.1162/jocn.2007.19.1.109 by guest on 01 October 2021 may have stemmed from response biases, changes in was directed to either side by means of a brief non- decision criteria, or low-level sensory interactions be- predictive tone presented 100–300 msec before the on- tween the attention-modulating cue and target stimuli set of the TOJ stimuli, and participants had to judge (e.g., Schneider & Bavelier, 2003; see also Zampini et al., which of two lateralized differently colored light flashes 2005; Spence, Shore, et al., 2001). occurred first. The results showed a behavioral prior- Spence, Shore, et al. (2001) highlighted and evaluated entry effect of 68.5 msec, but exogenous spatial atten- a number of the confounds and ambiguities that were tion did not lead to any shift in the latency of early visual present in previous research and reported a series of potentials. Instead, their gain was modulated, as has experiments designed to test for attentional effects upon commonly been observed in any number of other spa- Downloaded from http://mitprc.silverchair.com/jocn/article-pdf/19/1/109/1756371/jocn.2007.19.1.109.pdf by guest on 18 May 2021 TOJs while accounting for these confounds. First and tial attention experiments (Anllo-Vento, Schoenfeld, & foremost, the effects of response bias were identified— Hillyard, 2004; Eimer, 1998, 2004; Luck, Woodman, & the tendency by participants to respond to the stimulus Vogel, 2000; Hillyard & Anllo-Vento, 1998; Mangun, to which they had been instructed to attend. To coun- 1995). The authors concluded that the behavioral shifts teract this bias, Spence and his colleagues used a cuing in the PSS were not caused by perceptual prior entry, design in which the participants perceived stimuli from but that they arose instead from modulations of signal different sensory modalities in different spatial locations, strength in early visual–cortical pathways. and responded on a dimension that was orthogonal to The current experiment revisited the possibility that that to which they were instructed to attend (e.g., perceptual prior-entry effects may occur under different responding on the basis of the stimulus location that manipulations of selective attention. Closer to the orig- came first while being instructed to attend to a particular inal observations in Geiger’s (1903) pendulum experi- sensory modality). While accounting for these potential ment, we manipulated the focus of attention to one biases,Spenceandcolleagueswereabletogarner sensory modality versus another. We used visual and support for the veracity of the prior-entry hypothesis. tactile peripheral stimuli, given that this is the modality A follow-up experiment by Zampini et al. (2005) also pairing that has given rise to the biggest prior-entry ef- confirmed that attention leads to a shift in the PSS when fect in previous behavioral work (Spence, Nicholls, et al., using a simultaneity judgment task instead. 2001; Spence, Shore, et al., 2001). Using an orthogonal Although it has proved difficult to arbitrate unequiv- cuing task (based on Spence, Shore, et al., 2001), partic- ocally between accounts invoking cognitive biases ipants were cued to attend to either vision or touch and (Schneider & Bavelier, 2003) or attentional prior-entry to respond according to the side on which the first stim- effects (Zampini et al., 2005; Spence, Shore, et al., 2001) ulus occurred. The orienting of attention was manipu- based solely on the available behavioral evidence, event- lated in a sustained manner by increasing the proportion related potentials (ERPs) should provide more direct of stimuli presented in one modality (attended) and evidence with which to address the issue. Indeed, by task instruction. Sustained attention manipulations Dennett (1991, p. 162) suggested that our brains may are often more effective than trial-by-trial manipulations, represent temporal order with primary evoked poten- and therefore might provide a better platform for un- tials as a reference point (see also Kelly, 2005; Roache, veiling mechanisms of attentional modulation. 1999; Ko¨hler, 1947). ERPs provide a readout of brain The task allowed us to identify the stages of percep- activity with real-time resolution, and thus, can reveal the tual processing of visual stimuli and to investigate their transient modulation of information processing during modulation. In particular, we were interested in inves- the perceptual analysis of a stimulus, such as changes in tigating whether differences in the relative latencies of the timing of a particular stage of analysis or in the early visual potentials could be observed as a function of amount of processing during a particular stage. There- attention being directed to a particular sensory modality fore, we recorded ERPs during a visuotactile TOJ task in a TOJ task. Latency shifts in early visual potentials using a design based on the work of Spence, Shore, and would provide the strongest evidence yet in support Klein (2001). Schneider and Bavelier (2003, p. 334) of the prior-entry hypothesis and would rule out ex- suggest that evaluating the prior-entry hypothesis re- planations based solely on any direct