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1.  Corrigendum: Cognitive training with casual video games: points to consider 
PMCID: PMC3960918
attention; working memory; reasoning; fluid intelligence; video games; cognitive training; casual games; transfer of training
2.  Do Athletes Excel at Everyday Tasks? 
Cognitive enhancements are associated with sport training. We extended the sport-cognition literature by using a realistic street crossing task to examine the multitasking and processing speed abilities of collegiate athletes and nonathletes.
Pedestrians navigated trafficked roads by walking on a treadmill in a virtual world, a challenge that requires the quick and simultaneous processing of multiple streams of information.
Athletes had higher street crossing success rates than nonathletes, as reflected by fewer collisions with moving vehicles. Athletes also showed faster processing speed on a computer-based test of simple reaction time, and shorter reaction times were associated with higher street crossing success rates.
The results suggest that participation in athletics relates to superior street crossing multitasking abilities and that athlete and nonathlete differences in processing speed may underlie this difference. We suggest that cognitive skills trained in sport may transfer to performance on everyday fast-paced multitasking abilities.
PMCID: PMC3953501  PMID: 21407125
3.  A neuroimaging investigation of the association between aerobic fitness, hippocampal volume, and memory performance in preadolescent children 
Brain research  2010;1358:172-183.
Because children are becoming overweight, unhealthy, and unfit, understanding the neurocognitive benefits of an active lifestyle in childhood has important public health and educational implications. Animal research has indicated that aerobic exercise is related to increased cell proliferation and survival in the hippocampus as well as enhanced hippocampal-dependent learning and memory. Recent evidence extends this relationship to elderly humans by suggesting that high aerobic fitness levels in older adults are associated with increased hippocampal volume and superior memory performance. The present study aimed to further extend the link between fitness, hippocampal volume, and memory to a sample of preadolescent children. To this end, magnetic resonance imaging was employed to investigate whether higher- and lower-fit 9- and 10-year-old children showed differences in hippocampal volume and if the differences were related to performance on an item and relational memory task. Relational but not item memory is primarily supported by the hippocampus. Consistent with predictions, higher-fit children showed greater bilateral hippocampal volumes and superior relational memory task performance compared to lower-fit children. Hippocampal volume was also positively associated with performance on the relational but not the item memory task. Furthermore, bilateral hippocampal volume was found to mediate the relationship between fitness level (VO2 max) and relational memory. No relationship between aerobic fitness, nucleus accumbens volume, and memory was reported, which strengthens the hypothesized specific effect of fitness on the hippocampus. The findings are the first to indicate that aerobic fitness may relate to the structure and function of the preadolescent human brain.
PMCID: PMC3953557  PMID: 20735996
Brain; Children; Exercise; Hippocampus; MRI; Physical activity
4.  Neurobiological markers of exercise-related brain plasticity in older adults 
The current study examined how a randomized one-year aerobic exercise program for healthy older adults would affect serum levels of brain-derived neurotrophic factor (BDNF), insulin-like growth factor type 1 (IGF-1), and vascular endothelial growth factor (VEGF) - putative markers of exercise-induced benefits on brain function. The study also examined whether (a) change in the concentration of these growth factors was associated with alterations in functional connectivity following exercise, and (b) the extent to which pre-intervention growth factor levels were associated with training-related changes in functional connectivity. In 65 participants (mean age = 66.4), we found that although there were no group-level changes in growth factors as a function of the intervention, increased temporal lobe connectivity between the bilateral parahippocampus and the bilateral middle temporal gyrus was associated with increased BDNF, IGF-1, and VEGF for an aerobic walking group but not for a non-aerobic control group, and greater pre-intervention VEGF was associated with greater training-related increases in this functional connection. Results are consistent with animal models of exercise and the brain, but are the first to show in humans that exercise-induced increases in temporal lobe functional connectivity are associated with changes in growth factors and may be augmented by greater baseline VEGF.
PMCID: PMC3544982  PMID: 23123199
exercise; aging; functional connectivity; fMRI; default mode network; aerobic fitness; growth factors
5.  Cognitive training with casual video games: points to consider 
Frontiers in Psychology  2014;4:1010.
Brain training programs have proliferated in recent years, with claims that video games or computer-based tasks can broadly enhance cognitive function. However, benefits are commonly seen only in trained tasks. Assessing generalized improvement and practicality of laboratory exercises complicates interpretation and application of findings. In this study, we addressed these issues by using active control groups, training tasks that more closely resemble real-world demands and multiple tests to determine transfer of training. We examined whether casual video games can broadly improve cognition, and selected training games from a study of the relationship between game performance and cognitive abilities. A total of 209 young adults were randomized into a working memory–reasoning group, an adaptive working memory–reasoning group, an active control game group, and a no-contact control group. Before and after 15 h of training, participants completed tests of reasoning, working memory, attention, episodic memory, perceptual speed, and self-report measures of executive function, game experience, perceived improvement, knowledge of brain training research, and game play outside the laboratory. Participants improved on the training games, but transfer to untrained tasks was limited. No group showed gains in reasoning, working memory, episodic memory, or perceptual speed, but the working memory–reasoning groups improved in divided attention, with better performance in an attention-demanding game, a decreased attentional blink and smaller trail-making costs. Perceived improvements did not differ across training groups and those with low reasoning ability at baseline showed larger gains. Although there are important caveats, our study sheds light on the mixed effects in the training and transfer literature and offers a novel and potentially practical training approach. Still, more research is needed to determine the real-world benefits of computer programs such as casual games.
PMCID: PMC3882717  PMID: 24432009
attention; working memory; reasoning; fluid intelligence; video games; cognitive training; casual games; transfer of training
6.  Selling points: What cognitive abilities are tapped by casual video games? 
Acta psychologica  2012;142(1):74-86.
The idea that video games or computer-based applications can improve cognitive function has led to a proliferation of programs claiming to “train the brain.” However, there is often little scientific basis in the development of commercial training programs, and many research-based programs yield inconsistent or weak results. In this study, we sought to better understand the nature of cognitive abilities tapped by casual video games and thus reflect on their potential as a training tool. A moderately large sample of participants (n=209) played 20 web-based casual games and performed a battery of cognitive tasks. We used cognitive task analysis and multivariate statistical techniques to characterize the relationships between performance metrics. We validated the cognitive abilities measured in the task battery, examined a task analysis-based categorization of the casual games, and then characterized the relationship between game and task performance. We found that games categorized to tap working memory and reasoning were robustly related to performance on working memory and fluid intelligence tasks, with fluid intelligence best predicting scores on working memory and reasoning games. We discuss these results in the context of overlap in cognitive processes engaged by the cognitive tasks and casual games, and within the context of assessing near and far transfer. While this is not a training study, these findings provide a methodology to assess the validity of using certain games as training and assessment devices for specific cognitive abilities, and shed light on the mixed transfer results in the computer-based training literature. Moreover, the results can inform design of a more theoretically-driven and methodologically-sound cognitive training program.
PMCID: PMC3679476  PMID: 23246789
Working memory; Reasoning; Fluid intelligence; Video games; Cognitive training; Casual games
7.  Parietal plasticity after training with a complex video game is associated with individual differences in improvements in an untrained working memory task 
Researchers have devoted considerable attention and resources to cognitive training, yet there have been few examinations of the relationship between individual differences in patterns of brain activity during the training task and training benefits on untrained tasks (i.e., transfer). While a predominant hypothesis suggests that training will transfer if there is training-induced plasticity in brain regions important for the untrained task, this theory lacks sufficient empirical support. To address this issue we investigated the relationship between individual differences in training-induced changes in brain activity during a cognitive training videogame, and whether those changes explained individual differences in the resulting changes in performance in untrained tasks. Forty-five young adults trained with a videogame that challenges working memory, attention, and motor control for 15 2-h sessions. Before and after training, all subjects received neuropsychological assessments targeting working memory, attention, and procedural learning to assess transfer. Subjects also underwent pre- and post-functional magnetic resonance imaging (fMRI) scans while they played the training videogame to assess how these patterns of brain activity change in response to training. For regions implicated in working memory, such as the superior parietal lobe (SPL), individual differences in the post-minus-pre changes in activation predicted performance changes in an untrained working memory task. These findings suggest that training-induced plasticity in the functional representation of a training task may play a role in individual differences in transfer. Our data support and extend previous literature that has examined the association between training related cognitive changes and associated changes in underlying neural networks. We discuss the role of individual differences in brain function in training generalizability and make suggestions for future cognitive training research.
PMCID: PMC3968753
cognitive training; neuroplasticity; tranfser; working memory; video games
8.  Training and Transfer of Training in Rapid Visual Search for Camouflaged Targets 
PLoS ONE  2013;8(12):e83885.
Previous examinations of search under camouflage conditions have reported that performance improves with training and that training can engender near perfect transfer to similar, but novel camouflage-type displays [1]. What remains unclear, however, are the cognitive mechanisms underlying these training improvements and transfer benefits. On the one hand, improvements and transfer benefits might be associated with higher-level overt strategy shifts, such as through the restriction of eye movements to target-likely (background) display regions. On the other hand, improvements and benefits might be related to the tuning of lower-level perceptual processes, such as figure-ground segregation. To decouple these competing possibilities we had one group of participants train on camouflage search displays and a control group train on non-camouflage displays. Critically, search displays were rapidly presented, precluding eye movements. Before and following training, all participants completed transfer sessions in which they searched novel displays. We found that search performance on camouflage displays improved with training. Furthermore, participants who trained on camouflage displays suffered no performance costs when searching novel displays following training. Our findings suggest that training to break camouflage is related to the tuning of perceptual mechanisms and not strategic shifts in overt attention.
PMCID: PMC3873983  PMID: 24386301
9.  Aging Brain from a Network Science Perspective: Something to Be Positive About? 
PLoS ONE  2013;8(11):e78345.
To better understand age differences in brain function and behavior, the current study applied network science to model functional interactions between brain regions. We observed a shift in network topology whereby for older adults subcortical and cerebellar structures overlapping with the Salience network had more connectivity to the rest of the brain, coupled with fragmentation of large-scale cortical networks such as the Default and Fronto-Parietal networks. Additionally, greater integration of the dorsal medial thalamus and red nucleus in the Salience network was associated with greater satisfaction with life for older adults, which is consistent with theoretical predictions of age-related increases in emotion regulation that are thought to help maintain well-being and life satisfaction in late adulthood. In regard to cognitive abilities, greater ventral medial prefrontal cortex coherence with its topological neighbors in the Default Network was associated with faster processing speed. Results suggest that large-scale organizing properties of the brain differ with normal aging, and this perspective may offer novel insight into understanding age-related differences in cognitive function and well-being.
PMCID: PMC3819386  PMID: 24223147
10.  Functional connectivity: a source of variance in the association between cardiorespiratory fitness and cognition? 
Neuropsychologia  2010;48(5):1394-1406.
Over the next twenty years the number of Americans diagnosed with dementia is expected to more than double (CDC 2007). It is, therefore, an important public health initiative to understand what factors contribute to the longevity of a healthy mind. Both default mode network (DMN) function and increased aerobic fitness have been associated with better cognitive performance and reduced incidence of Alzheimer’s disease among older adults. Here we examine the association between aerobic fitness, functional connectivity in the DMN, and cognitive performance. Results showed significant age-related deficits in functional connectivity in both local and distributed DMN pathways. However, in a group of healthy elderly adults, almost half of the age-related disconnections showed increased functional connectivity as a function of aerobic fitness level. Finally, we examine the hypothesis that functional connectivity in the DMN is one source of variance in the relationship between aerobic fitness and cognition. Results demonstrate instances of both specific and global DMN connectivity mediating the relationship between fitness and cognition. We provide the first evidence for functional connectivity as a source of variance in the association between aerobic fitness and cognition, and discuss results in the context of neurobiological theories of cognitive aging and disease.
PMCID: PMC3708614  PMID: 20079755
cognitive aging; fMRI; functional connectivity; aerobic exercise; executive function; spatial memory
11.  The association between aerobic fitness and executive function is mediated by prefrontal cortex volume 
Brain, Behavior, and Immunity  2011;26(5):811-819.
Aging is marked by a decline in cognitive function, which is often preceded by losses in gray matter volume. Fortunately, higher cardiorespiratory fitness (CRF) levels are associated with an attenuation of age-related losses in gray matter volume and a reduced risk for cognitive impairment. Despite these links, we have only a rudimentary understanding of whether fitness-related increases in gray matter volume lead to elevated cognitive function. In this cross-sectional study, we examined whether the association between higher aerobic fitness levels and elevated executive function was mediated by greater gray matter volume in the prefrontal cortex (PFC). One hundred and forty-two older adults (mean age = 66.6 years) completed structural magnetic resonance imaging (MRI) scans, CRF assessments, and performed Stroop and spatial working memory (SPWM) tasks. Gray matter volume was assessed using an optimized voxel-based morphometry approach. Consistent with our predictions, higher fitness levels were associated with (a) better performance on both the Stroop and SPWM tasks, and (b) greater gray matter volume in several regions, including the dorsolateral PFC (DLPFC). Volume of the right inferior frontal gyrus and precentral gyrus mediated the relationship between CRF and Stroop interference while a non-overlapping set of regions bilaterally in the DLPFC mediated the association between CRF and SPWM accuracy. These results suggest that specific regions of the DLPFC differentially relate to inhibition and spatial working memory. Thus, fitness may influence cognitive function by reducing brain atrophy in targeted areas in healthy older adults.
PMCID: PMC3321393  PMID: 22172477
cardiorespiratory fitness; executive function; voxel-based morphometry; cortical volume; prefrontal cortex; mediation
12.  Perceptual-Cognitive Expertise in Elite Volleyball Players 
The goal of the current study was to investigate the relationship between sport expertise and perceptual and cognitive skills, as measured by the component skills approach. We hypothesized that athletes would outperform non-athlete controls in a number of perceptual and cognitive domains and that sport expertise would minimize gender differences. A total of 154 individuals (87 professional volleyball players and 67 non-athlete controls) participated in the study. Participants performed a cognitive battery, which included tests of executive control, memory, and visuo-spatial attention. Athletes showed superior performance speed on three tasks (two executive control tasks and one visuo-spatial attentional processing task). In a subset of tasks, gender effects were observed mainly in the control group, supporting the notion that athletic experience can reduce traditional gender effects. The expertise effects obtained substantiate the view that laboratory tests of cognition may indeed enlighten the sport-cognition relationship.
PMCID: PMC3590639  PMID: 23471100
cognition; expertise; sport
13.  Physical Activity Improves Verbal and Spatial Memory in Older Adults with Probable Mild Cognitive Impairment: A 6-Month Randomized Controlled Trial 
Journal of Aging Research  2013;2013:861893.
We report secondary findings from a randomized controlled trial on the effects of exercise on memory in older adults with probable MCI. We randomized 86 women aged 70–80 years with subjective memory complaints into one of three groups: resistance training, aerobic training, or balance and tone (control). All participants exercised twice per week for six months. We measured verbal memory and learning using the Rey Auditory Verbal Learning Test (RAVLT) and spatial memory using a computerized test, before and after trial completion. We found that the aerobic training group remembered significantly more items in the loss after interference condition of the RAVLT compared with the control group after six months of training. In addition, both experimental groups showed improved spatial memory performance in the most difficult condition where they were required to memorize the spatial location of three items, compared with the control group. Lastly, we found a significant correlation between spatial memory performance and overall physical capacity after intervention in the aerobic training group. Taken together, our results provide support for the prevailing notion that exercise can positively impact cognitive functioning and may represent an effective strategy to improve memory in those who have begun to experience cognitive decline.
PMCID: PMC3595715  PMID: 23509628
14.  The effects of physical activity on functional MRI activation associated with cognitive control in children: a randomized controlled intervention 
This study used functional magnetic resonance imaging (fMRI) to examine the influence of a 9-month physical activity program on task-evoked brain activation during childhood. The results demonstrated that 8- to 9-year-old children who participated in 60+ min of physical activity, 5 days per week, for 9 months, showed decreases in fMRI brain activation in the right anterior prefrontal cortex coupled with within-group improvements in performance on a task of attentional and interference control. Children assigned to a wait-list control group did not show changes in brain function. Furthermore, at post-test, children in the physical activity group showed similar anterior frontal brain patterns and incongruent accuracy rates to a group of college-aged young adults. Children in the wait-list control group still differed from the young adults in terms of anterior prefrontal activation and performance at post-test. There were no significant changes in fMRI activation in the anterior cingulate cortex (ACC) for either group. These results suggest that physical activity during childhood may enhance specific elements of prefrontal cortex function involved in cognitive control.
PMCID: PMC3594762  PMID: 23487583
activation; brain; children; fitness; fMRI; physical activity
15.  History of mild traumatic brain injury is associated with deficits in relational memory, reduced hippocampal volume, and less neural activity later in life 
Evidence suggests that a history of head trauma is associated with memory deficits later in life. The majority of previous research has focused on moderate-to-severe traumatic brain injury (TBI), but recent evidence suggests that even a mild TBI (mTBI) can interact with the aging process and produce reductions in memory performance. This study examined the association of mTBI with memory and the brain by comparing young and middle-aged adults who have had mTBI in their recent (several years ago) and remote (several decades ago) past, respectively, with control subjects on a face-scene relational memory paradigm while they underwent functional magnetic resonance imaging (fMRI). Hippocampal volumes were also examined from high-resolution structural images. Results indicated middle-aged adults with a head injury in their remote past had impaired memory compared to gender, age, and education matched control participants, consistent with previous results in the study of memory, aging, and TBI. The present findings extended previous results by demonstrating that these individuals also had smaller bilateral hippocampi, and had reduced neural activity during memory performance in cortical regions important for memory retrieval. These results indicate that a history of mTBI may be one of the many factors that negatively influence cognitive and brain health in aging.
PMCID: PMC3749487  PMID: 23986698
mTBI; hippocampus; aging; relational memory; fMRI
16.  Aerobic fitness is associated with greater efficiency of the network underlying cognitive control in preadolescent children 
Neuroscience  2011;199:166-176.
This study examined whether individual differences in aerobic fitness are associated with differences in activation of cognitive control brain networks in preadolescent children. As expected, children performed worse on a measure of cognitive control compared to a group of young adults. However, individual differences in aerobic fitness were associated with cognitive control performance among children. Lower-fit children had disproportionate performance cost in accuracy with increasing task difficulty, relative to higher-fit children. Brain activation was compared between performance-matched groups of lower- and higher-fit children. Fitness groups differed in brain activity for regions associated with response execution and inhibition, task set maintenance, and top-down regulation. Overall, differing activation patterns coupled with different patterns of brain-behavior correlations suggest an important role of aerobic fitness in modulating task strategy and the efficiency of neural networks that implement cognitive control in preadolescent children.
PMCID: PMC3237764  PMID: 22027235
exercise; physical activity; aerobic fitness; executive control; fMRI; development
17.  Caudate Nucleus Volume Mediates the Link between Cardiorespiratory Fitness and Cognitive Flexibility in Older Adults 
Journal of Aging Research  2012;2012:939285.
The basal ganglia play a central role in regulating the response selection abilities that are critical for mental flexibility. In neocortical areas, higher cardiorespiratory fitness levels are associated with increased gray matter volume, and these volumetric differences mediate enhanced cognitive performance in a variety of tasks. Here we examine whether cardiorespiratory fitness correlates with the volume of the subcortical nuclei that make up the basal ganglia and whether this relationship predicts cognitive flexibility in older adults. Structural MRI was used to determine the volume of the basal ganglia nuclei in a group of older, neurologically healthy individuals (mean age 66 years, N = 179). Measures of cardiorespiratory fitness (VO2max), cognitive flexibility (task switching), and attentional control (flanker task) were also collected. Higher fitness levels were correlated with higher accuracy rates in the Task Switching paradigm. In addition, the volume of the caudate nucleus, putamen, and globus pallidus positively correlated with Task Switching accuracy. Nested regression modeling revealed that caudate nucleus volume was a significant mediator of the relationship between cardiorespiratory fitness, and task switching performance. These findings indicate that higher cardiorespiratory fitness predicts better cognitive flexibility in older adults through greater grey matter volume in the dorsal striatum.
PMCID: PMC3415086  PMID: 22900181
18.  Aerobic Fitness and Response Variability in Preadolescent Children Performing a Cognitive Control Task 
Neuropsychology  2011;25(3):333-341.
To investigate the relationship between aerobic fitness and cognitive variability in preadolescent children.
Forty-eight preadolescent children (25 males, 23 females, mean age = 10.1 years) were grouped into higher- and lower-fit groups according to their performance on a test of aerobic capacity (VO2max). Cognitive function was measured via behavioral responses to a modified flanker task. The distribution in reaction time was calculated within each participant to assess intra-individual variability of performance. Specifically, the standard deviation and coefficient variation of reaction time were used to represent cognitive variability.
Preadolescent children, regardless of fitness, exhibited longer reaction time, increased response variability, and decreased response accuracy to incongruent compared to congruent trials. Further, higher-fit children were less variable in their response time and more accurate in their responses across conditions of the flanker task, while no group differences were observed for response speed.
These findings suggest that fitness is associated with better cognitive performance during a task that varies cognitive control demands, and extends this area of research to suggest that intra-individual variability may be a useful measure to examine the relationship between fitness and cognition during preadolescence.
PMCID: PMC3086950  PMID: 21443340
Physical Activity; Executive Control; Standard Deviation; Coefficient of Variation; Reaction Time
19.  Beyond vascularization: aerobic fitness is associated with N-acetylaspartate and working memory 
Brain and Behavior  2012;2(1):32-41.
Aerobic exercise is a promising form of prevention for cognitive decline; however, little is known about the molecular mechanisms by which exercise and fitness impacts the human brain. Several studies have postulated that increased regional brain volume and function are associated with aerobic fitness because of increased vascularization rather than increased neural tissue per se. We tested this position by examining the relationship between cardiorespiratory fitness and N-acetylaspartate (NAA) levels in the right frontal cortex using magnetic resonance spectroscopy. NAA is a nervous system specific metabolite found predominantly in cell bodies of neurons. We reasoned that if aerobic fitness was predominantly influencing the vasculature of the brain, then NAA levels should not vary as a function of aerobic fitness. However, if aerobic fitness influences the number or viability of neurons, then higher aerobic fitness levels might be associated with greater concentrations of NAA. We examined NAA levels, aerobic fitness, and cognitive performance in 137 older adults without cognitive impairment. Consistent with the latter hypothesis, we found that higher aerobic fitness levels offset an age-related decline in NAA. Furthermore, NAA mediated an association between fitness and backward digit span performance, suggesting that neuronal viability as measured by NAA is important in understanding fitness-related cognitive enhancement. Since NAA is found exclusively in neural tissue, our results indicate that the effect of fitness on the human brain extends beyond vascularization; aerobic fitness is associated with neuronal viability in the frontal cortex of older adults.
PMCID: PMC3343297  PMID: 22574272
Aging; brain; exercise; fitness; human; N-acetylaspartate; working memory
20.  Examining neural correlates of skill acquisition in a complex videogame training program 
Acquisition of complex skills is a universal feature of human behavior that has been conceptualized as a process that starts with intense resource dependency, requires effortful cognitive control, and ends in relative automaticity on the multi-faceted task. The present study examined the effects of different theoretically based training strategies on cortical recruitment during acquisition of complex video game skills. Seventy-five participants were recruited and assigned to one of three training groups: (1) Fixed Emphasis Training (FET), in which participants practiced the game, (2) Hybrid Variable-Priority Training (HVT), in which participants practiced using a combination of part-task training and variable priority training, or (3) a Control group that received limited game play. After 30 h of training, game data indicated a significant advantage for the two training groups relative to the control group. The HVT group demonstrated enhanced benefits of training, as indexed by an improvement in overall game score and a reduction in cortical recruitment post-training. Specifically, while both groups demonstrated a significant reduction of activation in attentional control areas, namely the right middle frontal gyrus, right superior frontal gyrus, and the ventral medial prefrontal cortex, participants in the control group continued to engage these areas post-training, suggesting a sustained reliance on attentional regions during challenging task demands. The HVT group showed a further reduction in neural resources post-training compared to the FET group in these cognitive control regions, along with reduced activation in the motor and sensory cortices and the posteromedial cortex. Findings suggest that training, specifically one that emphasizes cognitive flexibility can reduce the attentional demands of a complex cognitive task, along with reduced reliance on the motor network.
PMCID: PMC3351675  PMID: 22615690
skill acquisition; training strategies; attentional control; functional MRI
21.  Striatal Volume Predicts Level of Video Game Skill Acquisition 
Cerebral Cortex (New York, NY)  2010;20(11):2522-2530.
Video game skills transfer to other tasks, but individual differences in performance and in learning and transfer rates make it difficult to identify the source of transfer benefits. We asked whether variability in initial acquisition and of improvement in performance on a demanding video game, the Space Fortress game, could be predicted by variations in the pretraining volume of either of 2 key brain regions implicated in learning and memory: the striatum, implicated in procedural learning and cognitive flexibility, and the hippocampus, implicated in declarative memory. We found that hippocampal volumes did not predict learning improvement but that striatal volumes did. Moreover, for the striatum, the volumes of the dorsal striatum predicted improvement in performance but the volumes of the ventral striatum did not. Both ventral and dorsal striatal volumes predicted early acquisition rates. Furthermore, this early-stage correlation between striatal volumes and learning held regardless of the cognitive flexibility demands of the game versions, whereas the predictive power of the dorsal striatal volumes held selectively for performance improvements in a game version emphasizing cognitive flexibility. These findings suggest a neuroanatomical basis for the superiority of training strategies that promote cognitive flexibility and transfer to untrained tasks.
PMCID: PMC3841463  PMID: 20089946
basal ganglia; caudate nucleus; cognitive flexibility; nucleus accumbens; procedural learning
22.  Basal Ganglia Volume Is Associated with Aerobic Fitness in Preadolescent Children 
Developmental Neuroscience  2010;32(3):249-256.
The present investigation is the first to explore the association between childhood aerobic fitness and basal ganglia structure and function. Rodent research has revealed that exercise influences the striatum by increasing dopamine signaling and angiogenesis. In children, higher aerobic fitness levels are associated with greater hippocampal volumes, superior performance on tasks of attentional and interference control, and elevated event-related brain potential indices of executive function. The present study used magnetic resonance imaging to investigate if higher-fit and lower-fit 9- and 10-year-old children exhibited differential volumes of other subcortical brain regions, specifically the basal ganglia involved in attentional control. The relationship between aerobic fitness, dorsal and ventral striatum volumes and performance on an attention and inhibition Eriksen flanker task was also examined. The results indicated that higher-fit children showed superior flanker task performance compared to lower-fit children. Higher-fit children also showed greater volumes of the dorsal striatum, and dorsal striatum volume was negatively associated with behavioral interference. The results support the claim that the dorsal striatum is involved in cognitive control and response resolution and that these cognitive processes vary as a function of aerobic fitness. No relationship was found between aerobic fitness, the volume of the ventral striatum and flanker performance. The findings suggest that increased childhood aerobic fitness is associated with greater dorsal striatal volumes and that this is related to enhanced cognitive control. Because children are becoming increasingly overweight, unhealthy and unfit, understanding the neurocognitive benefits of an active lifestyle during childhood has important public health and educational implications.
PMCID: PMC3696376  PMID: 20693803
Brain; Development; Exercise; MRI; Physical activity; Neurocognition; Neuroimaging; Striatum
23.  Aerobic Fitness is Associated With Hippocampal Volume in Elderly Humans 
Hippocampus  2009;19(10):1030-1039.
Deterioration of the hippocampus occurs in elderly individuals with and without dementia, yet individual variation exists in the degree and rate of hippocampal decay. Determining the factors that influence individual variation in the magnitude and rate of hippocampal decay may help promote lifestyle changes that prevent such deterioration from taking place. Aerobic fitness and exercise are effective at preventing cortical decay and cognitive impairment in older adults and epidemiological studies suggest that physical activity can reduce the risk for developing dementia. However, the relationship between aerobic fitness and hippocampal volume in elderly humans is unknown. In this study, we investigated whether individuals with higher levels of aerobic fitness displayed greater volume of the hippocampus and better spatial memory performance than individuals with lower fitness levels. Furthermore, in exploratory analyses, we assessed whether hippocampal volume mediated the relationship between fitness and spatial memory. Using a region-of-interest analysis on magnetic resonance images in 165 nondemented older adults, we found a triple association such that higher fitness levels were associated with larger left and right hippocampi after controlling for age, sex, and years of education, and larger hippocampi and higher fitness levels were correlated with better spatial memory performance. Furthermore, we demonstrated that hippocampal volume partially mediated the relationship between higher fitness levels and enhanced spatial memory. Our results clearly indicate that higher levels of aerobic fitness are associated with increased hippocampal volume in older humans, which translates to better memory function.
PMCID: PMC3072565  PMID: 19123237
aging; MRI; spatial memory; cognition; brain
24.  BDNF is Associated With Age-Related Decline in Hippocampal Volume 
Hippocampal volume shrinks in late adulthood, but the neuromolecular factors that trigger hippocampal decay in aging humans remains a matter of speculation. In rodents, brain derived neurotrophic factor (BDNF) promotes the growth and proliferation of cells in the hippocampus and is important in long-term potentiation and memory formation. In humans, circulating levels of BDNF decline with advancing age and a genetic polymorphism for BDNF has been related to gray matter volume loss in old age. In this study, we tested whether age-related reductions in serum levels of BDNF would be related to shrinkage of the hippocampus and memory deficits in older adults. Hippocampal volume was acquired by automated segmentation of magnetic resonance images in 142 older adults without dementia. The caudate nucleus was also segmented and examined in relation to levels of serum BDNF. Spatial memory was tested using a paradigm in which memory load was parametrically increased. We found that increasing age was associated with smaller hippocampal volumes, reduced levels of serum BDNF, and poorer memory performance. Lower levels of BDNF were associated with smaller hippocampi and poorer memory, even when controlling for the variation related to age. In an exploratory mediation analysis, hippocampal volume mediated the age-related decline in spatial memory and BDNF mediated the age-related decline in hippocampal volume. Caudate nucleus volume was unrelated to BDNF levels or spatial memory performance. Our results identify serum BDNF as a significant factor related to hippocampal shrinkage and memory decline in late adulthood.
PMCID: PMC3069644  PMID: 20392958
brain-derived neurotrophic factor; hippocampus; human; brain; caudate nucleus; aging
25.  Resting Hippocampal Blood Flow, Spatial Memory and Aging 
Brain research  2009;1315C:119.
Aging is accompanied by a general deterioration of fluid cognitive processes and a reduction in resting cerebral blood flow (CBF). While the two phenomena have been observed independently, it is uncertain whether individual differences in cerebral blood flow are reliably associated with cognitive functioning in older adults. Furthermore, previous studies have concentrated primarily on gross measures of cognition and global gray matter CBF, leaving open the possibility that perfusion of specific brain regions may relate differentially to distinct cognitive domains. The present study sought to provide a more focused treatment of CBF and cognitive function in the context of aging by investigating the relationships among aging, spatial memory and resting hippocampal blood flow, both between and within younger and older adult groups. Blood flow was quantified using a novel Flow-Enhanced Signal Intensity (FENSI) technique which provides a localized, functionally-relevant measure of volumetric flow across a given unit area. As expected, we found that aging was associated with poorer spatial memory and reduced resting CBF. Moreover, hippocampal blood flow was positively correlated with spatial memory performance in the older adult group, suggesting that increased blood flow to the hippocampus is associated with superior memory performance in older adults. These results demonstrate a region-specific CBF—cognition relationship and thereby offer new insight into the complex connection between the aging brain and behavior.
PMCID: PMC2822086  PMID: 20026320
hippocampus; cerebral blood flow; brain perfusion; cognition; memory; aging

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