Theoretical Perspectives on Age Differences in Brain Activation: HAROLD, PASA, CRUNCH—How Do They STAC Up?
Sara B. Festini, Laura Zahodne, and Patricia A. Reuter-Lorenz
Cognitive neuroimaging studies often report that older adults display more activation of neural networks relative to younger adults, referred to as overactivation. Greater or more widespread activity frequently involves bilateral recruitment of both cerebral hemispheres, especially the frontal cortex. In many reports, overactivation has been associated with superior cognitive performance, suggesting that this activity may reflect compensatory processes that offset age-related decline and maintain behavior. Several theories have been proposed to account for age differences in brain activation, including the Hemispheric Asymmetry Reduction in Older Adults (HAROLD) model, the Posterior-Anterior Shift in Aging (PASA) theory, the Compensation-Related Utilization of Neural Circuits Hypothesis (CRUNCH), and the Scaffolding Theory of Aging and Cognition (STAC and STAC-r). Each model has a different explanatory scope with regard to compensatory processes, and each has been highly influential in the field. HAROLD contrasts the general pattern of bilateral prefrontal activation in older adults with that of more unilateral activation in younger adults. PASA describes both anterior (e.g., frontal) overactivation and posterior (e.g., occipital) underactivation in older adults relative to younger adults. CRUNCH emphasizes that the level or extent of brain activity can change in response to the level of task demand at any age. Finally, STAC and STAC-r take the broadest perspective to incorporate individual differences in brain structure, the capacity to implement functional scaffolding, and life-course neural enrichment and depletion factors to predict cognition and cognitive change across the lifespan. Extant empirical work has documented that compensatory overactivation can be observed in regions beyond the prefrontal cortex, that variations in task difficulty influence the degree of brain activation, and that younger adults can show compensatory overactivation under high mental demands. Additional research utilizing experimental designs (e.g., transcranial magnetic stimulation), longitudinal assessments, greater regional precision, both verbal and nonverbal material, and measures of individual difference factors will continue to refine our understanding of age-related activation differences and adjudicate among these various accounts of neurocognitive aging.
Human visual development is a complex dynamic psychological/neurobiological process, being part of the developing systems for cognition, action, and attention. This article reviews current knowledge and methods of study of human visual development in infancy and childhood, in relation to typical early visual brain development, and how it can change in developmental disorders, both acquired (e.g., related to at-risk births) and genetic disorders. The newborn infant starts life with a functioning subcortical visual system which controls newborn orienting to nearby high contrast objects and faces. Although visual cortex may be active from birth, its characteristic stimulus selectivity and control of visual responses is generally seen to emerge around six to twelve weeks after birth. By age six months the infant has adequate acuity and contrast sensitivity in nearby space, and operating cortical mechanisms for discriminating colors, shapes, faces, movement, stereo depth, and distance of objects, as well as the ability to focus and shift attention between objects of interest. This may include both feedforward and feedback pathways between cortical areas and between cortical and subcortical areas. Two cortical streams start to develop and become interlinked, the dorsal stream underpinning motion, spatial perception and actions, and the ventral stream for recognition of objects and faces. The neural systems developing control and planning of actions include those for directed eye movements, reaching and grasping, and the beginnings of locomotion, with these action systems being integrated into the other developing subcortical and cortical visual networks by one year of age. Analysis of global static form (pattern) and global motion processing allows the development of dorsal and ventral streams to be monitored from infancy through childhood. The development of attention, visuomotor control and spatial cognition in the first years show aspects of function related to the developing dorsal stream, and their integration with the ventral stream.
The milestones of typical visual development can be used to characterize visual and visuo-cognitive disorders early in life, such as in infants with perinatal brain injuries and those born very prematurely. The concept of “dorsal stream vulnerability” is outlined. It was initially based on deficits in global motion sensitivity relative to static form sensitivity, but can be extended to the planning and execution of visuomotor actions and problems of attention, together with visuospatial and numerical cognition. These problems are found in the phenotype of children with both genetic developmental disorders (e.g., Williams syndrome, autism, fragile-X, and dyslexia), and in acquired developmental disorders related to very preterm birth, or in children with abnormal visual input such as congenital cataract, refractive errors, or amblyopia. However, there are subtle differences in the manifestation of these disorders which may also vary considerably across individuals. Development in these clinical conditions illustrates the early, but limited, plasticity of visual brain mechanisms, and provides a challenge for the future in designing successful intervention and treatment.
Working memory as a temporary buffer for cognitive processing is an essential part of the cognitive system. Its capacity and select aspects of its functioning are age sensitive, more so for spatial than verbal material. Assumed causes for this decline include a decline in cognitive resources (such as speed of processing), and/or a breakdown in basic control processes (resistance to interference, task coordination, memory updating, binding, and/or top-down control as inferred from neuroimaging data). Meta-analyses suggest that a decline in cognitive resources explains much more of the age-related variance in true working memory tasks than a breakdown in basic control processes, although the latter is highly implicated in tasks of passive storage. The age-related decline in working memory capacity has downstream effects on more complex aspects of cognition (episodic memory, spatial cognition, and reasoning ability). Working memory remains plastic in old age, and training in working memory and cognitive control processes yields near transfer effects, but little evidence for strong far transfer.