Rewards and Cognitive Control in the Human Prefrontal Cortex.

The human prefrontal cortex (PFC) subserves cognitive control, that is, the ability to form behavioral strategies that coordinate actions and thoughts in relation to internal goals. Cognitive control involves the medial and lateral PFC but we still poorly understand how these regions guide strategy selection according to expected rewards. We addressed this issue using neuroimaging, computational modeling and model-based analyses of information flows between medial and lateral PFC. We show here that the (dorsal) medial PFC encodes and conveys to lateral PFC reward expectations driving strategy selection, while strategy selection originates in lateral PFC and propagates backward to medial PFC. This functional loop through lateral PFC enables strategy selection to further comply with learned rules encoded in lateral PFC rather than with reward expectations conveyed from medial PFC. Thus, the medial and lateral PFC are functionally coupled in cognitive control for integrating expected rewards and learned rules in strategy selection.

Experimental evidence for circular inference in schizophrenia.

Schizophrenia (SCZ) is a complex mental disorder that may result in some combination of hallucinations, delusions and disorganized thinking. Here SCZ patients and healthy controls (CTLs) report their level of confidence on a forced-choice task that manipulated the strength of sensory evidence and prior information. Neither group's responses can be explained by simple Bayesian inference. Rather, individual responses are best captured by a model with different degrees of circular inference. Circular inference refers to a corruption of sensory data by prior information and vice versa, leading us to 'see what we expect' (through descending loops), to 'expect what we see' (through ascending loops) or both. Ascending loops are stronger for SCZ than CTLs and correlate with the severity of positive symptoms. Descending loops correlate with the severity of negative symptoms. Both loops correlate with disorganized symptoms. The findings suggest that circular inference might mediate the clinical manifestations of SCZ.

Effects of age on the neural correlates of retrieval cue processing are modulated by task demands.

The electrophysiological correlates of retrieval orientation--the differential processing of retrieval cues according to the nature of the sought-for information--were investigated in healthy young (18-20 years old) and older (63-77 years old) adults. In one pair of study-test cycles, subjects studied either words or pictures presented in one of two visually distinct contexts, and then performed a yes/no recognition task with words as test items. In another pair of study-test cycles, subjects again made recognition judgments, but were required, in addition, to signal the study context for each item judged "old." Young subjects' event-related potentials (ERPs) for new (unstudied) test items were more negative-going when the study material was pictures rather than words, and this effect varied little between the two retrieval tasks. Replicating a previous report [Morcom, A. M., & Rugg, M. D. Effects of age on retrieval cue processing as revealed by ERPs. Neuropsychologia, 42, 1525-1542, 2004], the effects of study material on the ERPs of the older subjects were attenuated and statistically nonsignificant in the recognition task. In the source retrieval task, however, material effects in the older group were comparable in both onset latency and magnitude with those of the young subjects. Thus, the failure of older adults to demonstrate differential cue processing in tests of recognition memory likely reflects the adoption of a specific retrieval strategy rather than the incapacity to process retrieval cues in a goal-directed manner.

The relationship between aging, performance, and the neural correlates of successful memory encoding.

The present functional magnetic resonance imaging (fMRI) study investigated whether age-related differences in the neural correlates of successful memory encoding are modulated by memory performance. Young (mean age 22 years; N = 16) and older (mean age 69 years; N = 32) subjects were scanned while making animacy decisions on visually presented words. Memory for the words was later assessed in a recognition test, allowing fMRI activity elicited by study words to be contrasted according to subsequent memory performance. Young and older adults exhibited equivalent subsequent memory effects (enhanced activity for later remembered items) in an extensive network that included left inferior prefrontal cortex and anterior hippocampus. In posterior cingulate cortex, reversed subsequent memory effects (greater activity for later forgotten items) were of greater magnitude in young subjects. A voxel-of-interest analysis conducted on left and right prefrontal subsequent memory effects revealed that the effects were distributed more bilaterally in older than in young subjects, replicating previous findings. This age-related difference was confined to older subjects with relatively poor recognition performance, who were also the only group to demonstrate statistically significant right prefrontal subsequent memory effects. The findings suggest that relative preservation of memory performance with increasing age does not depend upon right prefrontal "over-recruitment."

Do working-memory executive components mediate the effects of age on strategy selection or on strategy execution? Insights from arithmetic problem solving.

Younger and older adults performed an inequality verification task (7+6 < 15, Yes/No?) in a control condition and in a dual-task condition where they simultaneously performed an executive-component task. Arithmetic-problem characteristics were manipulated in order to test strategy selection (i.e., choice of appropriate strategies in order to improve performance) and strategy execution (i.e., performance of the cognitive processes involved in each strategy). Results revealed that strategy selection changes with age: Older adults mainly selected one type of strategies in contrast to younger adults who used several types of strategies. These age-related changes were similar in the control and dual-task conditions. Strategy execution also changed with age, as shown by larger age-related differences on hardest problems. These age-related changes were larger in the dual-task condition, compared to the control condition. This impact of executive components as mediator of age-related changes depended on general age-related slowing. We discuss these findings in order to further understand the effects of age on arithmetic performance.

Working memory, strategy execution, and strategy selection in mental arithmetic.

A total of 72 participants estimated products of complex multiplications of two-digit operands (e.g., 63 x 78), using two strategies that differed in complexity. The simple strategy involved rounding both operands down to the closest decades (e.g., 60 x 70), whereas the complex strategy required rounding both operands up to the closest decades (e.g., 70 x 80). Participants accomplished this estimation task in two conditions: a no-load condition and a working-memory load condition in which executive components of working memory were taxed. The choice/no-choice method was used to obtain unbiased strategy execution and strategy selection data. Results showed that loading working-memory resources led participants to poorer strategy execution. Additionally, participants selected the simple strategy more often under working-memory load. We discuss the implications of the results to further our understanding of variations in strategy selection and execution, as well as our understanding of the impact of working-memory load on arithmetic performance and other cognitive domains.

Arithmetic split effects reflect strategy selection: an adult age comparative study in addition comparison and verification tasks.

We tested whether split effects in arithmetic (i.e., better performance on large-split problems, like 3 + 8 = 16, than on small-split problems, like 3 + 8 = 12) reflect decision processing or strategy selection. To achieve this end, we tested performance of younger and older adults, matched on arithmetic skills, on two arithmetic tasks: the addition/number comparison task (e.g., 4 + 8, 13; which item is the larger?) and in the inequality verification task (e.g., 4 + 8 < 13; Yes/No?). In both tasks, split between additions and proposed numbers were manipulated. We also manipulated the difficulty of the additions, which represents an index of arithmetic fact calculation (i.e., hard problems, like 6 + 8 < 15, are solved more slowly than easy problems, like 2 + 4 < 07, suggesting that calculation takes longer). Analyses of latencies revealed three main results: First, split effects were of smaller magnitude in older adults compared to younger adults, whatever the type of arithmetic task; second, split effects were of smaller magnitude on easy problems; and third, calculation processes were well maintained in older adults with high level of arithmetic skills. This set of results improves our understanding of cognitive aging and strategy selection in arithmetic.

Age-related differences in arithmetic problem-verification strategies.

To test age-related differences in split and problem-difficulty effects, adults between the ages of 20 and 80 years (N = 138) performed a simple and a complex inequality verification task (e.g., 6 + 3 < 11, 271 + 182 < 458; true or false?). Split effects in verification tasks (i.e., better performance for large-split than for small-split problems) reflect strategy selection between nonexhaustive verification (e.g., evaluation of plausibility; estimation) and exhaustive verification (e.g., retrieval; calculation). Problem-difficulty effects (i.e., better performance for easy than hard problems) reflect calculation processing. Results showed decreased split effects across age groups, particularly in the complex task. Moreover, problem-difficulty effects did not vary across age groups. Age-related changes were mostly mediated by age-related declines in processing speed.

Alzheimer's disease disrupts arithmetic fact retrieval processes but not arithmetic strategy selection.

Three groups of healthy younger adults, healthy older adults, and probable AD patients, performed an addition/number comparison task. They compared 128 couples of additions and numbers (e.g., 4 + 9 15) and had to identify the largest item for each problem by pressing one of two buttons located under each item. Manipulations of problem characteristics (i.e., problem difficulty and splits between correct sums and proposed numbers) enabled us to examine strategy selection and specific arithmetic fact retrieval processes. Results showed that arithmetic facts retrieval processes, which were spared with aging, were impaired in AD patients. However, AD patients were able to switch between strategies across trials according to problem characteristics as well as healthy older adults, and less systematically than healthy younger adults. We discuss implications of these findings for further understanding AD-related differences in arithmetic in particular, and problem solving in general.