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a total of 130 participants volunteered ; they were divided into physical exercise participants ( pep ; n = 73 ; aged between 16 and 80 ; 36 women ) and not - physical exercise participants ( n - pep ; n = 57 ; aged between 17 and 86 ; 25 women ) based on a self - report questionnaire about physical exercise participation administered at the beginning of each experimental session in accordance with international classification ( 9 ) .
the n - pep group included participants not involved in any structured or planned physical exercise programs .
the pep group included those who reported involvement in regular physical exercise and sports at least 3 days / week , 1 hour / session ( ie , swimming , martial arts , fencing , and low- and high - impact exercise ) .
all of the participants were healthy and without histories of neurological , psychiatric , or chronic somatic diseases .
they were not taking psychoactive or vasoactive medication and had normal or corrected - to - normal vision .
general cognitive functions in older participants were assessed using the mini - mental state examination ( 11 ) ( averaged score was 29/30 , range : 2830 ) .
the older participants were recruited through the vitattiva association in rome , the middle - aged participants were recruited among friends of the authors , and the younger participants were recruited from the local student population .
the two groups were matched for education levels ( years of study : 15.2 , se = 1.3 , for the n - pep group ; 15.0 , se = 1.4 , for the pep group ) .
the participants written consent was obtained ( from parents for minors ) according to the declaration of helsinki after approval by the ethical committee of the irccs santa lucia foundation .
each participant was tested individually in a sound - attenuated dimly lit room after a 64-channel eeg cap was mounted on his / her scalp .
visual stimuli ( ie , four - squared figures made by vertical and horizontal bars ) were randomly displayed for 260ms with equal probability ( p = .25 ) ; the stimulus - onset asynchrony varied from 1 to 2 s to prevent time prediction effects on rt ( for more details on the paradigm ( 2 ) .
the participants performed a discriminative response task , also known as go / no - go task , which is widely used to assess sustained attention , response control , and executive / inhibitory functions ( 12 ) .
the participants had to press a button as fast as possible with the right hand when the target ( go stimuli ; p = .5 ) appeared on the screen and withhold a response when the non target ( no - go stimuli ; p = .5 ) appeared ( see supplementary figure 1 for more details ) .
rts , omissions , and false alarms , as well as the onset latency , peak amplitude , and latency of premotor activity at the afz electrode site ( placed over the medial pfc ) , were separately correlated with age for the n - pep and pep groups .
the data were first fit with linear functions and then with higher order polynomial functions .
the function with the highest correlation value ( pearson s adjusted r 
  coefficient ) was accepted .
a sample - by - sample t test ( two tailed ) was performed between fit functions for each sample ( 100 per function ) using bootstrap statistics .
a total of 130 participants volunteered ; they were divided into physical exercise participants ( pep ; n = 73 ; aged between 16 and 80 ; 36 women ) and not - physical exercise participants ( n - pep ; n = 57 ; aged between 17 and 86 ; 25 women ) based on a self - report questionnaire about physical exercise participation administered at the beginning of each experimental session in accordance with international classification ( 9 ) .
the n - pep group included participants not involved in any structured or planned physical exercise programs .
the pep group included those who reported involvement in regular physical exercise and sports at least 3 days / week , 1 hour / session ( ie , swimming , martial arts , fencing , and low- and high - impact exercise ) .
all of the participants were healthy and without histories of neurological , psychiatric , or chronic somatic diseases .
they were not taking psychoactive or vasoactive medication and had normal or corrected - to - normal vision .
general cognitive functions in older participants were assessed using the mini - mental state examination ( 11 ) ( averaged score was 29/30 , range : 2830 ) .
the older participants were recruited through the vitattiva association in rome , the middle - aged participants were recruited among friends of the authors , and the younger participants were recruited from the local student population .
the two groups were matched for education levels ( years of study : 15.2 , se = 1.3 , for the n - pep group ; 15.0 , se = 1.4 , for the pep group ) .
the participants written consent was obtained ( from parents for minors ) according to the declaration of helsinki after approval by the ethical committee of the irccs santa lucia foundation .
each participant was tested individually in a sound - attenuated dimly lit room after a 64-channel eeg cap was mounted on his / her scalp .
visual stimuli ( ie , four - squared figures made by vertical and horizontal bars ) were randomly displayed for 260ms with equal probability ( p = .25 ) ; the stimulus - onset asynchrony varied from 1 to 2 s to prevent time prediction effects on rt ( for more details on the paradigm ( 2 ) . the participants performed a discriminative response task , also known as go / no - go task , which is widely used to assess sustained attention , response control , and executive / inhibitory functions ( 12 ) .
the participants had to press a button as fast as possible with the right hand when the target ( go stimuli ; p = .5 ) appeared on the screen and withhold a response when the non target ( no - go stimuli ; p = .5 ) appeared ( see supplementary figure 1 for more details ) .
rts , omissions , and false alarms , as well as the onset latency , peak amplitude , and latency of premotor activity at the afz electrode site ( placed over the medial pfc ) , were separately correlated with age for the n - pep and pep groups .
the data were first fit with linear functions and then with higher order polynomial functions .
the function with the highest correlation value ( pearson s adjusted r 
  coefficient ) was accepted .
a sample - by - sample t test ( two tailed ) was performed between fit functions for each sample ( 100 per function ) using bootstrap statistics .
behaviorally , the rts increased with age following a second - order polynomial function ( linear and higher order functions yielded lower correlations ) .
the function was rt = 399  0.068  age + 0.025  age ( adjusted r 
  = .910 , f 
 2,54 = 365.1 , p <
.0001 ) for the n - pep group and rt = 391  0.267  age + 0.022  age ( adjusted r 
  = .911 , f 

.0001 ) for the pep group . the sample - by - sample t - test performed between the fit functions of the two groups showed significant differences from 36.7 years ( p < 0.029 ) up to the end of the comparable fit functions ( figure 1a ) .
.128 ) were found ( figure 1b ) . in line with previous investigations , we confirmed on a large sample that although response speed decreases with age , response accuracy does not change across life span .
( b ) lack of correlation between response accuracy ( expressed as percentage of the false alarms ) and age .
pfc activity was analyzed in the premotor preparation phase ( supplementary materials ) on the medial prefrontal electrode ( afz ) , and the peak latency was used as a dependent variable ( hereafter called pfc_pl ) .
the function was pfc_pl = 373  0.431  age + 0.099  age ( adjusted r 
  = .930 , f 
 2,54 = 373.1 , p <
.0001 ) for the n - pep and pfc_pl = 394  1.839  age + 0.082  age ( adjusted r 
  = .860 , f 
 2,70 = 222.9 , p <
.0001 ) for the pep . the sample - by - sample t test performed between the fit functions of the two groups showed significant differences from the age of 38.0 years ( p = .024 ) up to the end of the comparable fit functions ( figure 2a ) . although the onset latency and peak amplitude of pfc activity were significantly correlated with age and physical exercise , the determination coefficients were lower .
furthermore , the correlations between pfc_pls and rts were also significant ( p < .0001 ) in both the n - pep ( adjusted r 
  = .835 ; slope = 0.245 ) and pep ( adjusted r 
  = .846 ; slope = 0.253 ) groups , indicating that anticipation in the pfc_pl corresponded with slow rts ( figure 2b ) .
( a ) correlation between age and premotor activity in the prefrontal cortex ( pfc ) .
( b ) linear correlation between motor response time and premotor activity in the pfc . 

figure 3a shows mrcp waveforms over the medial prefrontal areas in the n - pep and pep groups divided into two age cohorts ( n = 25 ) : younger ( mean age : 25 years ) and older ( mean age : 65 years ) .
activity started earlier and was larger in the n - pep than in the pep group .
the minimum norm estimation ( supplementary materials ) of the cerebral sources that generate prefrontal activity at its peak latency were consistently bilaterally localized within the pfc in the frontal pole ( figure 3b ) and showed an increasing gradient of intensity from the younger pep ( the lowest ) to the older n - pep ( the highest ) participants .
this localization agrees well with previous studies that used the same paradigm but a different source localization method ( 2 ) .
( a ) mrcp waveforms over the medial prefrontal cortex ( pfc ; recorded on afz ) across the four cohorts .
( b ) source localization of the aforementioned activity using mne rendered on a realistic template of the brain ( frontal point of view ) at pfc_pl .
the activation foci were within the mid - dorsal pfc in the middle - frontal gyrus ( bas 910 ) .
the issue of population aging will be a challenge in the coming decades in both advanced and developing countries .
although the clinical literature has recognized for years that exercise affects overall health and brain function ( 13 ) , less is known about the benefits of physical exercise on executive functions mediated by the pfc across the adult life span .
we extended previous investigations by studying a population with a greater age range ( 1586 years ) . using high - resolution 64-channel eeg , we showed novel results concerning the timing of pfc activity during motor planning , its modulation by physical activity and its relationship with behavioral parameters .
physical activity did not affect pfc activity during young adulthood , but its effect was evident in participants older than 38 years of age : active individuals engaged the pfc less and were faster than less active individuals .
we did not observe significant differences in the executive processes of the young participants between the two groups .
this suggests that compared with middle - aged and older participants , the young participants were easily able to accomplish the task , showing decreased effortful task preparation , especially in the pfc .
the beneficial effects of physical exercise on cognitive functions in young people were not detected during the performance of the present task , but they can not be excluded .
we observed that physical exercise is particularly important from middle age onwards , when the brain faces a series of challenges that can include neurodegenerative diseases .
we extended previous fmri findings , which revealed lower activation of the pfc in older adults during executive control tasks following an exercise intervention ( 14 ) .
the present findings reveal that in middle - aged and older individuals , moderate - to - high levels of physical exercise has beneficial effects on the planning and execution of a response , as well as on the executive functions mediated by the pfc ; thus , public policy should focus on ways of increasing volitional activity in adults , starting from middle age .
there are accumulating animal and human studies explaining the metabolic and molecular changes induced by physical exercise that promote cognitive health in older people .
physical exercise has been shown to promote angiogenesis ( 15,16 ) , neurogenesis ( 17 ) ( see also the neurogenic reserve hypothesis ) ( 18 ) , and synaptogenesis ( 19 ) and to modulate central and peripheral levels of key neurochemicals , such as insulin like growth factor 1 and brain - derived neurotrophic factor ( 20 ) .
physical exercise also induces structural and functional changes across different brain regions , as revealed by neuroimaging findings in response to specific physical exercise interventions ( 2123 ) .
this study reports the first evidence of changes in the electrophysiology of the pfc induced by physical exercise .
further studies combining different techniques and behavioral measures would contribute to improving our understanding of the mechanisms underpinning cortical overrecruitment in older adults and how it might be mitigated .
furthermore , further studies should identify methods to choose the most appropriate tools to investigate the aging brain and to propose individually tailored interventions using a combination of physical and cognitive programs for successful aging .
a critical issue of this study is its correlational nature and the lack of a programmed intervention .
future work should seek to replicate the present findings with controlled longitudinal intervention programs . in summary
, these findings are of great importance to researchers in different related fields for fully understanding the aging process and developing effective multidisciplinary therapeutic and prevention strategies to counteract brain diseases .
motor planning in older adults likely relies on the overengagement of the prefrontal cortex ( pfc ) and is associated with slowness of movement and responses . does a physically active lifestyle 
 counteract the overrecruitment of the pfc during action preparation ? 
 this study used high - resolution electroencephalography to measure the effect of physical exercise on the executive functions of the pfc preceding a visuomotor discriminative task . 
 a total of 130 participants aged 1586 were divided into two groups based on physical exercise participation . 
 the response times and accuracy and the premotor activity of the pfc were separately correlated with age for the two groups . 
 the data were first fit with a linear function and then a higher order polynomial function . 
 we observed that after 3540 years of age , physically active individuals have faster response times than their less active peers and showed no signs of pfc hyperactivity during motor planning . 
 the present findings show that physical exercise could speed up the response of older people and reveal that also in middle - aged people , moderate - to - high levels of physical exercise benefits the planning / execution of a response and the executive functions mediated by the pfc , counteracting the neural overactivity often observed in the elderly adults .