We review evidence for partially segregated networks of brain areas that carry out different attentional functions. One system, which includes parts of the intraparietal cortex and superior frontal cortex, is involved in preparing and applying goal-directed (top-down) selection for stimuli and responses. This system is also modulated by the detection of stimuli. The other system, which includes the temporoparietal cortex and inferior frontal cortex, and is largely lateralized to the right hemisphere, is not involved in top-down selection. Instead, this system is specialized for the detection of behaviourally relevant stimuli, particularly when they are salient or unexpected. This ventral frontoparietal network works as a 'circuit breaker' for the dorsal system, directing attention to salient events. Both attentional systems interact during normal vision, and both are disrupted in unilateral spatial neglect.

Control of goal-directed and stimulus-driven attention in the brain

A baseline or control state is fundamental to the understanding of most complex systems. Defining a baseline state in the human brain, arguably our most complex system, poses a particular challenge. Many suspect that left unconstrained, its activity will vary unpredictably. Despite this prediction we identify a baseline state of the normal adult human brain in terms of the brain oxygen extraction fraction or OEF. The OEF is defined as the ratio of oxygen used by the brain to oxygen delivered by flowing blood and is remarkably uniform in the awake but resting state (e.g., lying quietly with eyes closed). Local deviations in the OEF represent the physiological basis of signals of changes in neuronal activity obtained with functional MRI during a wide variety of human behaviors. We used quantitative metabolic and circulatory measurements from positron-emission tomography to obtain the OEF regionally throughout the brain. Areas of activation were conspicuous by their absence. All significant deviations from the mean hemisphere OEF were increases, signifying deactivations, and resided almost exclusively in the visual system. Defining the baseline state of an area in this manner attaches meaning to a group of areas that consistently exhibit decreases from this baseline, during a wide variety of goal-directed behaviors monitored with positron-emission tomography and functional MRI. These decreases suggest the existence of an organized, baseline default mode of brain function that is suspended during specific goal-directed behaviors.

/pdf/a-default-mode-of-brain-function-zwenkbmjw7.pdf

A default mode of brain function.

Publisher Summary This chapter focuses on the modern notion of short-term memory, called working memory. Working memory refers to the temporary maintenance of information that was just experienced or just retrieved from long-term memory but no longer exists in the external environment. These internal representations are short-lived, but can be maintained for longer periods of time through active rehearsal strategies, and can be subjected to various operations that manipulate the information in such a way that makes it useful for goal-directed behavior. Working memory is a system that is critically important in cognition and seems necessary in the course of performing many other cognitive functions, such as reasoning, language comprehension, planning, and spatial processing. This chapter demonstrates the functional importance of dopamine to working memory function in several ways. Elucidation of the cognitive and neural mechanisms underlying human working memory is an important focus of cognitive neuroscience and neurology for much of the past decade. One conclusion that arises from research is that working memory, a faculty that enables temporary storage and manipulation of information in the service of behavioral goals, can be viewed as neither a unitary, nor a dedicated system. Data from numerous neuropsychological and neurophysiological studies in animals and humans demonstrates that a network of brain regions, including the prefrontal cortex, is critical for the active maintenance of internal representations.

Chapter 11 Working memory

Recent developments in the quantitative analysis of complex networks, based largely on graph theory, have been rapidly translated to studies of brain network organization. The brain's structural and functional systems have features of complex networks--such as small-world topology, highly connected hubs and modularity--both at the whole-brain scale of human neuroimaging and at a cellular scale in non-human animals. In this article, we review studies investigating complex brain networks in diverse experimental modalities (including structural and functional MRI, diffusion tensor imaging, magnetoencephalography and electroencephalography in humans) and provide an accessible introduction to the basic principles of graph theory. We also highlight some of the technical challenges and key questions to be addressed by future developments in this rapidly moving field.

Complex brain networks: graph theoretical analysis of structural and functional systems

Co-Planar Stereotaxic Atlas of the Human Brain—3-Dimensional Proportional System: An Approach to Cerebral Imaging, J. Talairach, P. Tournoux. Georg Thieme Verlag, New York (1988), 122 pp., 130 figs. DM 268

The Reorienting System of the Human Brain: From Environment to Theory of Mind

Medial prefrontal cortex (MPFC) is among those brain regions having the highest baseline metabolic activity at rest and one that exhibits decreases from this baseline across a wide variety of goal-directed behaviors in functional imaging studies. This high metabolic rate and this behavior suggest the existence of an organized mode of default brain function, elements of which may be either attenuated or enhanced. Extant data suggest that these MPFC regions may contribute to the neural instantiation of aspects of the multifaceted “self.” We explore this important concept by targeting and manipulating elements of MPFC default state activity. In this functional magnetic resonance imaging (fMRI) study, subjects made two judgments, one self-referential, the other not, in response to affectively normed pictures: pleasant vs. unpleasant (an internally cued condition, ICC) and indoors vs. outdoors (an externally cued condition, ECC). The ICC was preferentially associated with activity increases along the dorsal MPFC. These increases were accompanied by decreases in both active task conditions in ventral MPFC. These results support the view that dorsal and ventral MPFC are differentially influenced by attentiondemanding tasks and explicitly self-referential tasks. The presence of self-referential mental activity appears to be associated with increases from the baseline in dorsal MPFC. Reductions in ventral MPFC occurred consistent with the fact that attention-demanding tasks attenuate emotional processing. We posit that both self-referential mental activity and emotional processing represent elements of the default state as represented by activity in MPFC. We suggest that a useful way to explore the neurobiology of the self is to explore the nature of default state activity.

Medial prefrontal cortex and self-referential mental activity: Relation to a default mode of brain function

Nine previous positron emission tomography (PET) studies of human visual information processing were reanalyzed to determine the consistency across experiments of blood flow decreases during active tasks relative to passive viewing of the same stimulus array. Areas showing consistent decreases during active tasks included posterior cingulate/precuneous (Brodmann area, BA 31/7), left (BAS 40 and 39/19) and right (BA 40) inferior parietal cortex, left dorsolateral frontal cortex (BA S), left lateral inferior frontal cortex (BA 10/47), left inferior temporal gyrus @A 20), a strip of medial frontal regions running along a dorsal-ventral axis (BAs 8, 9, 10, and 32), and the right amygdala.

Experiments involving language-related processes tended to show larger decreases than nonlanguage experiments. This trend mainly reflected blood flow increases at certain areas in the passive conditions of the language experiments (relative to a fixation control in which no task stimulus was present) and slight blood flow decreases in the passive conditions of the nonlanguage experiments. When the active tasks were referenced to the fixation condition, the overall size of blood flow decreases in language and nonlanguage tasks were the same, but differences were found across cortical areas. Decreases were more pronounced in the posterior cingulate/precuneous (BAS 31/7) and right inferior parietal cortex (BA 40) during language-related tasks and more pronounced in left inferior frontal cortex (BA 10/47) during nonlanguage tasks.

Blood flow decreases did not generally show significant differences across the active task states within an experiment, but a verb-generation task produced larger decreases than a read task in right and left inferior parietal lobe (BA 40) and the posterior cingulate/precuneous (BA 31/7), while the read task produced larger decreases in left lateral inferior frontal cortex (BA 10/47). These effects mirrored those found between experiments in the language-nonlanguage comparison.

Consistent active minus passive decreases may reflect decreased activity caused by active task processes that generalize over tasks or increased activity caused by passive task processes that are suspended during the active tasks. Increased activity during the passive condition might reflect ongoing processes, such as unconstrained verbally mediated thoughts and monitoring of the external environment, body, and emotional state.

/pdf/common-blood-flow-changes-across-visual-tasks-ii-decreases-4r2u5i1njk.pdf

Gordon L. Shulman

Papers

Control of goal-directed and stimulus-driven attention in the brain

A default mode of brain function.

The Reorienting System of the Human Brain: From Environment to Theory of Mind

Medial prefrontal cortex and self-referential mental activity: Relation to a default mode of brain function

Common blood flow changes across visual tasks: Ii. decreases in cerebral cortex