Bruno Laeng

Pupillometry A Window to the Preconscious

The measurement of pupil diameter in psychology (in short, “pupillometry”) has just celebrated 50 years. The method established itself after the appearance of three seminal studies (Hess & Polt, 1960, 1964; Kahneman & Beatty, 1966). Since then, the method has continued to play a significant role within the field, and pupillary responses have been successfully used to provide an estimate of the “intensity” of mental activity and of changes in mental states, particularly changes in the allocation of attention and the consolidation of perception. Remarkably, pupillary responses provide a continuous measure regardless of whether the participant is aware of such changes. More recently, research in neuroscience has revealed a tight correlation between the activity of the locus coeruleus (i.e., the “hub” of the noradrenergic system) and pupillary dilation. As we discuss in this short review, these neurophysiological findings provide new important insights to the meaning of pupillary responses for mental activity. Finally, given that pupillary responses can be easily measured in a noninvasive manner, occur from birth, and can occur in the absence of voluntary, conscious processes, they constitute a very promising tool for the study of preverbal (e.g., infants) or nonverbal participants (e.g., animals, neurological patients).

Lateralization of categorical and coordinate spatial functions: A study of unilateral stroke patients

Sixty patients with unilateral stroke (half with left hemisphere damage and half with right hemisphere damage) and a control group (N = 15) matched for age and educational level were tested in two experiments. In one experiment they were first shown, on each trial, a sample drawing depicting one or more objects. Following a short delay, they were asked to identify the drawing when it was paired with a drawing in which the same object(s) was transformed in categorical or coordinate spatial relations. In the other experiment, the same subjects first were shown, on each trial, a sample drawing. They then judged which of two variants (each in one type of spatial relation) looked more similar to the sample drawing. Typically, patients with left-sided stroke mistakenly identified the categorical transformation for the sample drawing in the first task; in the second task, they judged the categorical transformation as more similar to the sample drawing. Patients with right-sided stroke mistakenly identified the coordinate transformations for the sample drawing in the first task, and, in the second task, typically judged the drawings transformed along coordinate spatial relations as more similar to the sample drawing. These findings provide evidence for complementary lateralization of the two types of spatial perception. It can therefore be inferred that separate functional subsystems process the two types of spatial relations.

Pupil size signals mental effort deployed during multiple object tracking and predicts brain activity in the dorsal attention network and the locus coeruleus

Attentional effort relates to the allocation of limited-capacity attentional resources to meet current task demands and involves the activation of top-down attentional systems in the brain. Pupillometry is a sensitive measure of this intensity aspect of top-down attentional control. Studies relate pupillary changes in response to cognitive processing to activity in the locus coeruleus (LC), which is the main hub of the brains noradrenergic system and it is thought to modulate the operations of the brains attentional systems. In the present study, participants performed a visual divided attention task known as multiple object tracking (MOT) while their pupil sizes were recorded by use of an infrared eye tracker and then were tested again with the same paradigm while brain activity was recorded using fMRI. We hypothesized that the individual pupil dilations, as an index of individual differences in mental effort, as originally proposed by Kahneman (1973), would be a better predictor of LC activity than the number of tracked objects during MOT. The current results support our hypothesis, since we observed pupil-related activity in the LC. Moreover, the changes in the pupil correlated with activity in the superior colliculus and the right thalamus, as well as cortical activity in the dorsal attention network, which previous studies have shown to be strongly activated during visual tracking of multiple targets. Follow-up pupillometric analyses of the MOT task in the same individuals also revealed that individual differences to cognitive load can be remarkably stable over a lag of several years. To our knowledge this is the first study using pupil dilations as an index of attentional effort in the MOT task and also relating these to functional changes in the brain that directly implicate the LC-NE system in the allocation of processing resources.

Pupillary Stroop effects

We recorded the pupil diameters of participants performing the words’ color-naming Stroop task (i.e., naming the color of a word that names a color). Non-color words were used as baseline to firmly establish the effects of semantic relatedness induced by color word distractors. We replicated the classic Stroop effects of color congruency and color incongruency with pupillary diameter recordings: relative to non-color words, pupil diameters increased for color distractors that differed from color responses, while they reduced for color distractors that were identical to color responses. Analyses of the time courses of pupil responses revealed further differences between color-congruent and color-incongruent distractors, with the latter inducing a steep increase of pupil size and the former a relatively lower increase. Consistent with previous findings that have demonstrated that pupil size increases as task demands rise, the present results indicate that pupillometry is a robust measure of Stroop interference, and it represents a valuable addition to the cognitive scientist’s toolbox.

CEREBRAL LATERALIZATION FOR THE PROCESSING OF SPATIAL COORDINATES AND CATEGORIES IN LEFT- AND RIGHT-HANDERS

Subjects judged whether a tachistoscopially lateralized drawing was identical or different to a drawing seen immediately before in free vision. The drawings depicted natural objects (e.g. animals). On half of the trials the tachistoscopic drawing presented the same objects but either the categorical or the coordinate spatial relations (according to Kosslyns definitions [23]) between the objects were transformed. In the first experiment 38 right-handed subjects (half males and half females) were tested. Categorical judgements were faster when the match drawing appeared in the right visual field, whereas coordinate judgements were faster when the match drawing appeared in the left visual field. In the second experiment 26 right-handed and 40 left-handed subjects participated. Almost all the subjects were female. Right-handed subjects replicated the findings of the subjects in the first experiment. However, the LHs did not show any difference in response times between spatial conditions and visual fields. These findings support Kosslyns hypothesis that the left and right hemispheres are specialized respectively for processing categorical and coordinate spatial relations. Moreover, they also suggest that this lateralization pattern is not typical of left-handers.

Does Color Synesthesia Pose a Paradox for Early-Selection Theories of Attention?

P.M. is a synesthete who experiences colors when viewing alphanumeric symbols. Her search for a target differing from distractors by a synesthetic color feature takes the form of a pop-out search. Thus, it would seem that synesthesia can occur preattentively. However, discrepancies between the regression functions of response times observed in target-present trials and target-absent trials, and the fact that fast response times occur only when the target is within a few degrees of visual angle from fixation, indicate that P.M.s synesthesia does not occur preattentively, but rather is within the focus of attention. We conclude that synesthesia is a genuine perceptual phenomenon that can have substantial influence on visual processing.

The Eye Pupil Adjusts to Imaginary Light

If a mental image is a rerepresentation of a perception, then properties such as luminance or brightness should also be conjured up in the image. We monitored pupil diameters with an infrared eye tracker while participants first saw and then generated mental images of shapes that varied in luminance or complexity, while looking at an empty gray background. Participants also imagined familiar scenarios (e.g., a “sunny sky” or a “dark room”) while looking at the same neutral screen. In all experiments, participants’ eye pupils dilated or constricted, respectively, in response to dark and bright imagined objects and scenarios. Shape complexity increased mental effort and pupillary sizes independently of shapes’ luminance. Because the participants were unable to voluntarily constrict their eyes’ pupils, the observed pupillary adjustments to imaginary light present a strong case for accounts of mental imagery as a process based on brain states similar to those that arise during perception.

Bright illusions reduce the eye's pupil.

We recorded by use of an infrared eye-tracker the pupil diameters of participants while they observed visual illusions of lightness or brightness. Four original illusions {based on Gaetano Kaniszas [Kanizsa G (1976) Subjective contours. Sci Am 234:48–52] and Akiyoshi Kitaokas [Kitaoka A. (2005) Trick Eyes (Barnes & Noble, New Providence, NJ).] examples} were manipulated to obtain control conditions in which the perceived illusory luminance was either eliminated or reduced. All stimuli were equiluminant so that constrictions in pupillary size could not be ascribed to changes in light energy. We found that the pupillary diameter rapidly varied according to perceived brightness and lightness strength. Differences in local contrast information could be ruled out as an explanation because, in a second experiment, the observers maintained eye fixation in the center of the display; thus, differential stimulation of the fovea by local contrast changes could not be responsible for the pupillary differences. Hence, the most parsimonious explanation for the present findings is that pupillary responses to ambient light reflect the perceived brightness or lightness of the scene and not simply the amount of physical light energy entering the eye. Thus, the pupillary physiological response reflects the subjective perception of light and supports the idea that the brains visual circuitry is shaped by visual experience with images and their possible sources.

Scrutinizing visual images: the role of gaze in mental imagery and memory.

Gaze was monitored by use of an infrared remote eye-tracker during perception and imagery of geometric forms and figures of animals. Based on the idea that gaze prioritizes locations where features with high information content are visible, we hypothesized that eye fixations should focus on regions that contain one or more local features that are relevant for object recognition. Most importantly, we predicted that when observers looked at an empty screen and at the same time generated a detailed visual image of what they had previously seen, their gaze would probabilistically dwell within regions corresponding to the original positions of salient features or parts. Correlation analyses showed positive relations between gazes dwell time within locations visited during perception and those in which gaze dwelled during the imagery generation task. Moreover, the more faithful an observers gaze enactment, the more accurate was the observers memory, in a separate test, of the dimension or size in which the forms had been perceived. In another experiment, observers saw a series of pictures of animals and were requested to memorize them. They were then asked later, in a recall phase, to answer a question about a property of one of the encoded forms; it was found that, when retrieving from long-term memory a previously seen picture, gaze returned to the location of the part probed by the question. In another experimental condition, the observers were asked to maintain fixation away from the original location of the shape while thinking about the answer, so as to interfere with the gaze enactment process; such a manipulation resulted in measurable costs in the quality of memory. We conclude that the generation of mental images relies upon a process of enactment of gaze that can be beneficial to visual memory.

Do separate processes identify objects as exemplars versus members of basic-level categories? Evidence from hemispheric specialization

When an object is identified as a specific exemplar, is it analyzed differently than when it is identified at the basic level? On the basis of a previous theory, we predicted that the left hemisphere (LH) is specialized for classifying objects at the basic level and the right hemisphere (RH) is specialized for classifying objects as specific exemplars. To test this prediction, participants were asked to view lateralized pictures of animals, artifacts, and faces of famous people; immediately after each picture was presented, a label was read aloud by the computer, and the participants decided whether the label was correct for that picture. A label could name the object at either the basic level (e.g., bird) or as an exemplar (e.g., robin). As predicted, we found that basic-level labels were matched faster when pictures were presented in the right visual field (and hence encoded initially in the LH), whereas exemplar labels were matched faster when pictures were presented in the left visual field (and hence encoded initially in the RH).