Mathematics bridging cognition and single-cell physiology.
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Biomedical subjects
Publications and source records attributed to Claus Bundesen.
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Illusory motion can be generated by successively flashing a stationary visual stimulus in two spatial locations separated by several degrees of visual angle. In appropriate conditions, the apparent motion is indistinguishable from real motion: The observer experiences a luminous object traversing a continuous path from one stimulus location to the other through intervening positions where no physical stimuli exist. The phenomenon has been extensively investigated for nearly a century but little is known about its neurophysiological foundation. Here we present images of activations in the primary visual cortex in response to real and apparent motion. The images show that during apparent motion, a path connecting the cortical representations of the stimulus locations is filled in by activation. The activation along the path of apparent motion is similar to the activation found when a stimulus is presented in real motion between the two locations.
A model of normal attentional function, based on the concept of competitive parallel processing, is used to compare attentional deficits following parietal and frontal lobe lesions. Measurements are obtained for visual processing speed, capacity of visual short-term memory (VSTM), spatial bias (bias to left or right hemifield) and top-down control (selective attention based on task relevance). The results show important differences, but also surprising similarities, in parietal and frontal lobe patients. For processing speed and VSTM, deficits are selectively associated with parietal lesions, in particular lesions of the temporoparietal junction. We discuss explanations based on either grey matter or white matter lesions. In striking contrast, measures of attentional weighting (spatial bias and top-down control) are predicted by simple lesion volume. We suggest that attentional weights reflect competition between broadly distributed object representations. Parietal and frontal mechanisms work together, both in weighting by location and weighting by task context.
Previous studies have shown that MT (i.e., the MT-complex) is activated during visual perception of apparent motion. To further explore the function of MT, we measured activation in MT by positron emission tomography (PET) using a broad range of stroboscopic stimulus events in which (a) the frame rate was so fast that observers perceived stimulus frames as simultaneous, (b) the frame rate was slower and generated compelling impressions of apparent motion, or (c) the frame rate was so slow that observers perceived temporal succession (successive views of the same objects at different locations) instead of motion. As expected, the simultaneity condition showed no activation (reliable increase in regional cerebral blood flow, rCBF) in MT whereas the motion condition showed activation in both left and right MT. However, the succession condition showed even stronger activation in left and right MT than did the motion condition. MT seems implicated in perception of retinal stimuli as successive views of the same object at different locations whether or not the views are connected by apparent motion.
A neural theory of visual attention (NTVA) is presented. NTVA is a neural interpretation of C. Bundesen's (1990) theory of visual attention (TVA). In NTVA, visual processing capacity is distributed across stimuli by dynamic remapping of receptive fields of cortical cells such that more processing resources (cells) are devoted to behaviorally important objects than to less important ones. By use of the same basic equations used in TVA, NTVA accounts for a wide range of known attentional effects in human performance (reaction times and error rates) and a wide range of effects observed in firing rates of single cells in the primate visual system. NTVA provides a mathematical framework to unify the 2 fields of research--formulas bridging cognition and neurophysiology.
In two experiments, subjects were given arbitrary letter cues or meaningful word cues that specified the task to be performed on a subsequent target stimulus. Letter and word cues were presented in separate blocks. There were two cues of each type for each task. Three kinds of transitions separated tasks: cue repetitions, in which both the cue and the task repeated; task repetitions, in which the cue changed but the task repeated; and task alternations, in which both the cue and the task changed. Responses were faster for cue than for task repetitions for both cue types. With word cues, task repetitions were not reliably faster than task alternations. With letter cues, task repetitions were reliably faster than task alternations in the first block but not in the second block. The results suggest that subjects responded to the compound of the cue and the target rather than switching task set between trials.
We report on a patient who complained of reduced awareness in the left visual field, but showed no visual neglect or extinction in clinical testing. By MR scanning, the brain damage was localized to the right basal ganglia, also involving structures in right frontal cortex. Using psychophysical testing and mathematical modeling based on Bundesen's theory of visual attention [TVA; Psychol. Rev. 97 (1990) 523], the patient's subjective experience of attentional disturbance was confirmed, and the deficit was specified into several components. At very short exposure durations, two effects were shown. The detection threshold was elevated, particularly in the left visual field, and stimuli in this side were given less attentional weight. In addition, the capacity of visual short-term memory (VSTM) was markedly reduced in both visual fields. The robustness of the test results was evaluated by bootstrap analysis. The study demonstrates the sensitivity and specificity gained by combining psychophysical testing with TVA modeling in the analysis of visual attention disorders. Extending the results of a pioneer study of parietal neglect patients by Duncan et al. [J. Exp. Psychol.: Gen. 128 (1999) 450], this study demonstrates the strength of the method in a single case, with a lesion outside parietal cortex, and only minor clinical symptoms.
Does the explicit task-cuing procedure require an endogenous a act of control? In 5 experiments, cues indicating which task to perform preceded targets by several stimulus onset asynchronies (SOAs). Two models were developed to account for changes in reaction time (RT) with SOA. Model 1 assumed an endogenous act of task switching for cue alterations but not for cue repetitions. Model 2 assumed no such act. In Experiments 1 and 2, the cue was masked or not masked. Masking interacted underadditively with repetition and alternation, consistent with Model 2 but not Model 1. In Experiments 3 and 4, 2 cues were used for each task. RT was slower for task repetition than for cue repetition and about the same as RT for task alternation, consistent with Model 2 but not Model 1. The results suggest that the explicit task-cuing procedure does not require an endogenous act of control.
Observers were given brief presentations of pairs of simultaneous stimuli consisting of a visual and a spoken letter. In the visual focused-attention conditon, only the visual letter should be reported; in the auditory focused-attention condition, only the spoken letter should be reported; in the divided-attention condition, both letters, as well as their respective modalities, should be reported (forced choice). The proportions of correct reports were nearly the same in the three conditions (no significant divided-attention decrement), and in the divided-attention condition, the probability that the visual letter was correctly reported was independent of whether the auditory letter was correctly reported. However, with a probability much higher than chance, the observers reported hearing the visual stimulus letter or seeing the spoken stimulus letter (modality confusions). The strength of the effect was nearly the same with focused as with divided attention. We also discovered a crossmodal congruity effect: Performance was better when the two letters in a stimulus pair were the same than when they differed in type.
Observers were presented with brief exposures of pairs of colored objects (letters) and asked to report both the color and the shape of each object. Several observers showed strikingly clear evidence of nearly perfect stochastic independence between reports of the four features (two colors and two shapes). For instance, the probability that the shape of a given object could be reported seemed independent of (a) whether the color of the object could be reported and (b) whether features of the other object could be reported. Such stochastic independence is predicted by many parallel-processing models (e.g., Bundesen, 1990). However, the results are difficult to reconcile with simple serial models in which the encoding of one object is completed before the encoding of another object is begun.
Attentional effects in the visual pathways were investigated by contrasting the distribution of regional cerebral blood flow (rCBF) measured by H(2)(15)O positron emission tomography (PET) during performance of a shape-matching task with the distribution of rCBF during a less demanding color-matching task. The two tasks were performed using the same stimuli: pairs of colored random shapes shown at a fixed rate (2 s per pair). In the shape-matching task, the subjects determined whether the two stimuli were the same in shape regardless of differences in size or color. In the color-matching task, the subjects determined whether the two stimuli were the same in color regardless of differences in size or shape. Mean reaction time for shape-matching exceeded mean reaction time for color-matching by nearly 200 ms. The corresponding shape-color comparison showed extensive bilateral increases in rCBF in visual areas in the occipital and parietal lobes, including the primary visual cortex. Subcortical activations were found in cerebellum (particularly the vermis) and in the thalamus with the focus in a region comprising the lateral geniculate nucleus, the pulvinar, and adjacent parts of the reticular nucleus. Frontal activations were found in a region that seems implicated in visual short-term memory (posterior parts of the superior sulcus and the middle gyrus). The reverse, color-shape comparison showed bilateral increases in rCBF in the anterior cingulate gyri, superior frontal gyri, and superior and middle temporal gyri. The attentional effects found by the shape-color comparison in the thalamus and the primary visual cortex may have been generated by feedback signals preserving visual representations of selected stimuli in short-term memory.
In visual search, inefficient performance of human observers is typically characterized by a steady increase in reaction time with the number of array elements-the so-called set-size effect. In general, set-size effects are taken to indicate that processing of the array elements depends on limited-capacity resources, that is, it involves attention. Contrasting theories have been proposed to account for this attentional involvement, however. While some theories have attributed set-size effects to the intervention of serial attention mechanisms, others have explained set-size effects in terms of parallel, competitive architectures. Conclusive evidence in favor of one or the other notion is still lacking. Especially in view of the wide use of visual search paradigms to explore the functional neuroanatomy of attentional mechanisms in the primate brain, it becomes essential that the nature of the attentional involvement in these paradigms be clearly defined at the behavioral level. Here we report a series of experiments showing that highly inefficient search indeed recruits serial attention deployment to the individual array elements. In addition, we describe a number of behavioral signatures of serial attention in visual search that can be used in future investigations to attest a similar involvement of serial attention in other search paradigms. We claim that only after having recognized these signatures can one be confident that truly serial mechanisms are engaged in a given visual search task, thus making it amenable for exploring the functional neuroanatomy underlying its performance.
Numerous brain functions, such as awareness of surroundings, control of movements, thoughts, and memory have always been a kind of mystery, which has provoked human curiosity and thus inspired research in many areas. The present paper gives an overview on imaging research of the human brain's functions. The historical progress is briefly reviewed with the emphasis on major Danish contributions. The major methods of mapping brain functions and their biological basis are mentioned. These methods are positron emission tomography (PET) and functional magnetic resonance imaging (fMRI). They are highly technological and generate enormous amounts of data. Thus, data analysis will per se be a research area in brain imaging. Finally, several examples of the authors' results on functional activation are dealt with. These examples aim to illustrate the research area and its contribution to our increased knowledge of the working brain.