How Exercise Improves Brain Health: The Complete Guide

Few interventions in the history of neuroscience have produced evidence as broad, as deep, and as consistent as physical exercise. Over the past three decades, researchers have moved from curious observations in mice to massive human trials involving millions of participants — and the findings are remarkable. Regular physical activity grows new brain cells, increases your brain's volume, elevates the proteins that keep neurons alive, and reduces your risk of dementia by as much as 60%.

This isn't hype. It's the conclusion of over 2,700 randomized controlled trials and hundreds of meta-analyses spanning every age group and cognitive domain. But the story is also more nuanced than most headlines suggest, and at Brain Zone, we think you deserve the full picture — including the uncertainties.

This guide walks through exactly what happens in your brain when you exercise, which cognitive functions benefit most, how exercise compares to psychiatric medication for depression, what it means for aging, and how much you actually need to do. Every claim is grounded in peer-reviewed research, and we'll tell you where the evidence is strong, where it's emerging, and where honest scientists still disagree.

Your brain on exercise: the biology behind the benefits

Exercise doesn't improve your brain through a single pathway. It works through at least seven reinforcing biological mechanisms, which helps explain why the cognitive benefits are so broad and so durable. Understanding these mechanisms — even at a high level — gives you a much clearer picture of why movement matters.

BDNF: the growth factor your brain depends on

Brain-Derived Neurotrophic Factor, or BDNF, is a protein that keeps existing neurons healthy and helps new ones grow. Think of it as fertilizer for your brain. Exercise is the most reliable way to increase it.

When you exercise, your brain produces the majority of the BDNF circulating in your bloodstream — roughly 70–80%, according to research by Rasmussen and colleagues published in Experimental Physiology in 2009. A single workout increases BDNF levels two- to threefold, and this effect gets stronger with regular training. A 2015 meta-analysis pooling 29 studies found that a single exercise bout produces a moderate BDNF increase, while regular training amplifies this acute response even further.

The long-term effects are especially striking. Seifert and colleagues demonstrated that three months of endurance training boosted resting BDNF release from the brain by 3.5-fold. And the mechanism is elegant: Sleiman et al. showed in eLife that exercise triggers a ketone body called β-hydroxybutyrate to accumulate in the hippocampus, which unlocks the gene that produces BDNF. The result is a cascade that supports synaptic plasticity, neuronal growth, and memory consolidation.

Exercise literally grows new brain cells

For most of the twentieth century, neuroscientists believed you were born with all the neurons you'd ever have. That turned out to be wrong. In a landmark 1999 paper in Nature Neuroscience, van Praag, Kempermann, and Gage showed that mice with access to running wheels dramatically increased the production of new neurons in the hippocampus — the brain's memory center. A companion paper in PNAS confirmed these new cells were fully functional, enhancing learning and the formation of new memories.

This neurogenesis extends across the lifespan. Even in aged mice, voluntary exercise reversed neurogenesis decline to half of young control levels and improved maze learning. But not all exercise is equal here. Nokia and colleagues compared exercise types in 2016 and found that only sustained aerobic exercise significantly boosted hippocampal neurogenesis — producing two to three times more new neurons than sedentary controls. High-intensity interval training had modest effects, and resistance training had none.

In humans, directly counting new neurons isn't yet possible, but Pereira et al. used MRI to show that 12 weeks of aerobic exercise increased blood volume in the dentate gyrus — a validated proxy for neurogenesis — in the exact brain region where new neurons are born. The dentate gyrus is critical for pattern separation, your ability to distinguish between similar memories. This likely explains why exercise so reliably helps with memory.

New blood vessels, stronger connections

Exercise increases blood flow to the brain and promotes the growth of new blood vessels — a process called angiogenesis. Morland and colleagues revealed the mechanism in Nature Communications: lactate released from working muscles activates a receptor on cells surrounding brain blood vessels, stimulating the growth factor VEGF and increasing capillary density throughout the cortex and hippocampus. When researchers knocked out this receptor in mice, exercise-induced angiogenesis disappeared entirely.

Structural changes follow. Colcombe et al. showed in 2006 that six months of aerobic exercise increased both gray matter and white matter volume in the prefrontal and temporal cortices of older adults. These regions govern planning, decision-making, and memory — precisely the functions that decline most with age. And these structural improvements depend on BDNF: when Vaynman and colleagues blocked BDNF signaling in exercising rats, the cognitive benefits vanished completely.

The real science behind the runner's high

Exercise boosts every major neurotransmitter system in the brain. It increases serotonin synthesis, elevates dopamine in reward circuits, and activates norepinephrine pathways that sharpen attention and alertness.

But the most surprising recent discovery concerns the runner's high. For decades, endorphins got the credit. That story has been rewritten. In a 2015 study published in PNAS, Fuss and colleagues demonstrated that the euphoria and anxiety reduction following exercise depend on cannabinoid receptors, not opioid receptors. Exercise increases your body's own cannabis-like molecules — particularly anandamide — and blocking cannabinoid receptors eliminated the runner's high, while blocking opioid receptors had no effect. Siebers et al. confirmed this in humans in 2021: participants who took an opioid blocker before exercising still experienced euphoria. The runner's high is an endocannabinoid phenomenon, not an endorphin one.

Calming inflammation and taming the stress response

Chronic inflammation in the brain is a major driver of cognitive decline. Exercise is one of the most effective ways to reduce it. As Gleeson and colleagues described in Nature Reviews Immunology, regular exercise lowers inflammatory markers like IL-6, TNF-α, and CRP while boosting anti-inflammatory signals. Intriguingly, the IL-6 released from muscles during exercise acts as an anti-inflammatory molecule, triggering a compensatory cascade that quiets systemic inflammation.

Exercise also recalibrates your stress response. While a single workout temporarily raises cortisol, chronic training does the opposite — it restores healthy cortisol rhythms, improves your brain's ability to shut down the stress response, and reduces how strongly you react to psychological stressors. Researchers call this the cross-stressor adaptation hypothesis: regular exercisers develop a physiological resilience that extends far beyond the gym, dampening the stress response to everyday challenges.

What exercise does for your thinking

The biological mechanisms above translate into measurable improvements across virtually every cognitive domain that scientists can test. Executive function — your ability to plan, make decisions, and resist distractions — benefits the most.

Planning, focus, and mental flexibility

The landmark meta-analysis by Colcombe and Kramer in Psychological Science (2003) analyzed 18 intervention studies and found that executive function showed the strongest response to exercise, with an effect size of 0.68. Programs combining aerobic and resistance training outperformed aerobic exercise alone.

More rigorous analyses have tempered these numbers without changing the conclusion. Smith et al. applied stricter criteria to 29 RCTs involving 2,049 participants and found smaller but still statistically significant improvements in attention, processing speed, and executive function. Northey and colleagues confirmed the pattern in adults over 50, with optimal results from 45–60 minute sessions at moderate-to-vigorous intensity. Aerobic exercise, resistance training, multicomponent programs, and tai chi all produced significant benefits.

Memory: the hippocampus can actually grow

The most celebrated study in this field is the 2011 trial by Erickson and colleagues published in PNAS. In this randomized controlled trial, 120 sedentary older adults (ages 55–80) were assigned to either a moderate walking program or a stretching control group for one year. The walkers exercised for 40 minutes, three times weekly. After twelve months, their anterior hippocampal volume had increased by 2% — effectively reversing one to two years of normal age-related shrinkage. The stretching group, as expected, showed a 1.4% decline. The volume increases correlated with both elevated BDNF and improved spatial memory.

A 2022 meta-analysis of 36 RCTs (2,750 participants) confirmed that aerobic exercise improves episodic memory overall, with stronger effects in adults aged 55–68 and in women. Working memory improvements are less consistent on their own but appear strongest when aerobic and resistance training are combined.

Creativity gets a boost — with a catch

Exercise also enhances creative thinking, though selectively. In a clever series of experiments, Oppezzo and Schwartz at Stanford found that walking increased divergent thinking — the kind of open-ended, brainstorming-style creativity — in 81% of participants. But it didn't improve convergent thinking (finding a single correct answer). The effect held even on a treadmill facing a blank wall, confirming that physical movement itself, not a change of scenery, drives creative ideation.

Exercise as mental health treatment

The evidence for exercise as a treatment for depression and anxiety has reached the point where it rivals front-line psychiatric medication — a claim that would have been controversial a decade ago but is now supported by the largest evidence syntheses ever conducted in the field.

Depression: exercise works as well as medication

The study that first made headlines was the SMILE trial by Blumenthal and colleagues, published in 1999. The team randomized 156 patients with major depressive disorder to aerobic exercise, the antidepressant sertraline, or both. After 16 weeks, all three groups improved equally. But the six-month follow-up told a more dramatic story: only 8% of the exercise group relapsed, compared to 38% of those on medication alone. Exercise appeared not only to treat depression but to build resilience against its return.

Since then, the evidence base has exploded. A 2024 network meta-analysis in the BMJ, drawing on 218 studies and over 14,000 participants, provided the most granular picture yet. Walking and jogging showed the strongest antidepressant effects, followed by yoga, strength training, and tai chi. The 2023 umbrella review by Singh et al. in the British Journal of Sports Medicine synthesized 97 systematic reviews encompassing 1,039 RCTs and 128,119 participants. Their conclusion: the effects of exercise on depression are "comparable to or slightly greater than" those observed for psychotherapy and pharmacotherapy.

This doesn't mean exercise should replace medication for everyone. Severe depression often requires pharmaceutical treatment, and combining exercise with medication may offer the best outcomes. But for mild to moderate depression, and for long-term relapse prevention, the case for exercise is now difficult to dispute.

Anxiety, stress, and better sleep

Exercise reduces anxiety through several converging pathways — from boosting calming neurotransmitters to a fascinating process called interoceptive habituation, where the physical arousal during exercise (racing heart, heavy breathing) mimics anxiety symptoms and gradually desensitizes you to them.

The scale of the mental health benefit is striking. Chekroud and colleagues analyzed data from 1.2 million U.S. adults in The Lancet Psychiatry and found that exercisers reported 43.2% fewer days of poor mental health than non-exercisers. The sweet spot was roughly 45 minutes per session, three to five times weekly. Exercising beyond 90 minutes per session or more than 23 times per month showed diminishing returns — a reminder that more isn't always better.

Sleep improvements add another layer of benefit. A meta-analysis of 66 studies found that regular exercise moderately improves sleep quality and reduces the time it takes to fall asleep, effects that cascade into better cognitive function and emotional regulation the following day.

Protecting your brain as you age

If exercise has a single most consequential benefit for public health, it may be its ability to protect the aging brain against cognitive decline and dementia.

Dementia risk drops dramatically

The longitudinal evidence is powerful. Rovio and colleagues followed 1,449 Finnish adults for 21 years and found that physical activity in midlife — at least twice weekly — was associated with 50% lower odds of developing dementia and 60% lower odds of Alzheimer's disease. The protective effect was strongest in people carrying the APOE-ε4 gene variant, the most significant known genetic risk factor for Alzheimer's. Larson et al. found a similar ~40% reduction in dementia risk among adults who exercised three or more times weekly, with the frailest participants benefiting the most.

A 2014 meta-analysis of 47 cohort studies confirmed the pattern: higher physical activity reduced cognitive decline risk by 35%. And in 2025, a study by Yau et al. in Nature Medicine revealed a specific mechanism — physical activity slowed the accumulation of tau protein, one of the hallmark pathologies of Alzheimer's disease, with benefits plateauing at around 5,000–7,500 steps per day.

The FINGER trial: the most successful dementia prevention study

The Finnish Geriatric Intervention Study to Prevent Cognitive Impairment and Disability — the FINGER trial — is the closest thing we have to a proven dementia prevention program. Published in The Lancet in 2015, this two-year trial enrolled 1,260 at-risk adults aged 60–77 and tested a combined intervention of exercise, cognitive training, nutritional guidance, and cardiovascular risk management.

The results were striking: the intervention group showed 25% greater total cognitive improvement than controls, 83% greater improvement in executive function, and 150% greater improvement in processing speed. The model has since been adopted worldwide through the World-Wide FINGERS network, now spanning over 60 countries.

It's worth noting that not all similar trials have succeeded. The MAPT and PreDIVA trials produced null results on their primary outcomes, possibly due to less intensive interventions. This suggests that the combination and intensity of lifestyle components matters enormously — and that exercise alone, while beneficial, works best as part of a comprehensive approach.

Your brain literally stays bigger

Exercise preserves brain tissue. Erickson et al. showed in the Cardiovascular Health Cognition Study that higher physical activity levels predicted larger volumes of frontal, temporal, and hippocampal brain regions nine years later. Ten Brinke et al. found that six months of aerobic training increased hippocampal volume by 4% in women with mild cognitive impairment. White matter — the wiring that connects brain regions — also benefits: a 2024 meta-analysis of 9 RCTs confirmed structural white matter improvements in older adults who exercised.

Exercise and the developing brain

The effects aren't limited to aging adults. Exercise profoundly shapes the developing brains of children and adolescents, influencing everything from academic performance to brain maturation.

Fitter kids perform better in school

The California Department of Education analyzed data from nearly 885,000 students and found a consistent positive relationship between physical fitness and standardized test scores in both reading and mathematics. The pattern held across 5th, 7th, and 9th graders, and the relationship was specifically tied to aerobic fitness — not muscular strength or flexibility.

Physical activity changes children's brain structure

The FITKids trial provides the strongest experimental evidence. Hillman and colleagues randomized 221 children ages 7–9 to a nine-month after-school physical activity program or waitlist control. The exercise group showed improved inhibitory control, cognitive flexibility, and enhanced neural processing. Benefits were dose-dependent: kids who attended more sessions showed larger brain changes.

Neuroimaging substudies revealed something remarkable — the physical activity program increased white matter quality in the corpus callosum, the tract connecting the brain's two hemispheres. Higher-fit children also showed cortical thinning patterns that mirror healthy brain maturation, suggesting exercise accelerates normal synaptic pruning and myelination.

Particular promise for ADHD

Exercise shows especially strong potential for attention disorders. Pontifex et al. found that a single 20-minute session of moderate walking improved response accuracy, inhibitory control, and academic performance in children with ADHD — the very cognitive functions most impaired by the condition. Meta-analyses of RCTs confirm that aerobic exercise significantly enhances selective attention, working memory, and impulse control in children with ADHD.

Which type of exercise is best for your brain?

Different exercise types activate different biological pathways and produce different cognitive outcomes. The research increasingly suggests that variety may be your best strategy.

Aerobic exercise: the gold standard

Running, walking, swimming, and cycling remain the most extensively studied forms of exercise for brain health, and they show the broadest cognitive benefits. The Erickson hippocampal volume study used nothing more than moderate-intensity walking — proof that you don't need to be an athlete for structural brain changes.

Resistance training: surprisingly powerful for thinking skills

Weight training has emerged as unexpectedly potent for executive function. Liu-Ambrose and colleagues showed that even once-weekly resistance training significantly improved selective attention and conflict resolution in women aged 65–75. A 2025 network meta-analysis found that resistance training actually produced the greatest improvement in global cognition among all exercise types studied. If you've been told that only cardio matters for your brain, the evidence suggests otherwise.

HIIT: the biggest BDNF spike

High-intensity interval training generates the largest acute increase in BDNF of any exercise modality. The mechanism likely involves elevated blood lactate, which crosses the blood-brain barrier and directly stimulates BDNF expression in the hippocampus. A 2023 network meta-analysis confirmed HIIT as the best modality for acutely boosting BDNF.

Dance: the two-for-one brain workout

Dance may offer unique benefits because it combines physical exertion with continuous cognitive demands — learning new steps, coordinating with music, navigating space. Rehfeld et al. conducted an 18-month trial comparing a constantly-changing dance program with conventional fitness training in older adults. Dancing was the only group that increased volume in the parahippocampal region, a brain area linked to working memory and episodic memory, and the only group showing significant BDNF increases at six months. The key: choreographies were constantly novel, preventing the brain from going on autopilot.

Yoga: brain benefits through a different door

Yoga affects the brain primarily through stress regulation rather than cardiovascular fitness. Gothe and Damoiseaux reviewed 11 neuroimaging studies and found yoga increases hippocampal and prefrontal cortex volume — the same regions that benefit from aerobic exercise, despite a completely different physiological pathway. Streeter et al. showed that yoga increases GABA levels in the thalamus by 27%, an effect comparable to anti-anxiety medication.

The muscle-brain axis: how your body talks to your brain

One of the most exciting frontiers in neuroscience is discovering exactly how exercising muscles communicate with the brain across the blood-brain barrier. Several molecular messengers have now been identified.

Irisin: the exercise hormone

In 2012, Boström and colleagues published a breakthrough paper in Nature describing irisin, a hormone released from muscles during exercise. Irisin travels through the bloodstream and drives BDNF expression in the hippocampus. It's present in human cerebrospinal fluid and appears to promote neuronal growth, though some scientific debate continues about its precise role in humans.

Lactate: from waste product to brain fuel

Lactate — the molecule responsible for the burn you feel during intense exercise — turns out to be a powerful signaling molecule. El Hayek and colleagues mapped the complete pathway in 2019: lactate crosses the blood-brain barrier, activates the enzyme SIRT1, and ultimately drives BDNF expression in the hippocampus. When the researchers blocked lactate transport during exercise, hippocampal BDNF induction was abolished entirely. And when they simply injected lactate into sedentary animals, it mimicked the memory benefits of exercise.

Cathepsin B: the 15-minute messenger

Moon and colleagues identified cathepsin B as a muscle-secreted protein that crosses the blood-brain barrier within 15 minutes and enhances both BDNF production and the birth of new hippocampal neurons. Mice lacking cathepsin B showed no exercise-induced neurogenesis or memory improvement. In humans, four months of treadmill exercise increased plasma cathepsin B, and the increase correlated directly with memory performance.

The kynurenine pathway: why exercise fights depression

Perhaps the most elegant discovery links exercise to depression resistance at the molecular level. Agudelo et al. showed in Cell that exercise-activated muscles convert a neurotoxic substance called kynurenine into a harmless form that cannot enter the brain. By filtering kynurenine from the bloodstream, exercise protects the brain from stress-induced inflammation. Mice engineered with elevated levels of the responsible muscle enzyme were completely resistant to depression — even under chronic stress.

Exercise rewrites your genetic code

Exercise doesn't just change your brain chemistry temporarily — it modifies which genes are active. Gomez-Pinilla and colleagues showed that exercise alters DNA methylation at the BDNF gene in the hippocampus, creating stable, long-lasting increases in BDNF production. Even more remarkably, Fernandes et al. reviewed evidence suggesting these epigenetic changes can be inherited — offspring of exercising parents show increased neuroplasticity markers, hinting at transgenerational benefits.

How much exercise does your brain actually need?

The evidence converges on practical guidelines that are achievable for most people.

The evidence-based minimum

Approximately 150 minutes per week of moderate-intensity aerobic exercise is the threshold where benefits consistently appear. This is what the World Health Organization recommends, and it's close to the protocol that grew hippocampal volume in the Erickson trial — 120 minutes weekly of moderate walking. The DOSE study found that this level of exercise reduced depression scores by 47%, while a lower dose performed no better than placebo. Interestingly, splitting the time into three sessions or five didn't significantly affect outcomes — total weekly volume matters more than frequency.

The optimal dose for cognitive performance

For executive function specifically, Northey and colleagues identified 45–60 minutes per session at moderate-to-vigorous intensity as optimal. Combining aerobic and resistance training outperforms either alone. For older adults with existing cognitive concerns, the evidence suggests resistance training two to three times weekly alongside aerobic sessions.

Timing and the law of diminishing returns

Morning exercise may offer particular advantages. Research by Wheeler et al. showed that morning exercise elevated BDNF for eight hours and improved decision-making across the entire day. The acute cognitive boost from a single workout lasts roughly one to two hours, making pre-work or pre-study exercise a strategic choice.

But the relationship between exercise and brain benefit isn't strictly linear. Chekroud et al.'s analysis of 1.2 million adults found that benefits peaked at around 45 minutes per session, three to five times weekly. Sessions beyond 90 minutes and exercising more than five times per week showed diminishing returns. Excessive exercise can even raise cortisol to levels that counteract BDNF production — a reminder that balance matters.

Putting it into practice

The single most important finding in this entire body of research is that the biggest cognitive gain comes from the transition between doing nothing and doing something. You don't need to become a marathon runner. The Erickson study enrolled previously sedentary older adults and used a gradual seven-week ramp-up from 10 to 40 minutes of walking. If you're currently inactive, that's your starting point.

For children and adolescents, the WHO recommends at least 60 minutes daily of moderate-to-vigorous activity. Coordinative activities — dance, team sports, active games — may produce superior cognitive benefits because they engage the brain and body simultaneously.

For adults aged 18–64, aim for at least 150 minutes weekly of moderate aerobic exercise plus two resistance training sessions. This combination provides the broadest cognitive benefits, strengthening both memory and executive function.

For older adults, multicomponent programs that combine aerobic exercise, resistance training, and balance work are optimal. Dance programs deserve special consideration given their unique dual-benefit profile for both physical fitness and neuroplasticity.

For those with mild cognitive impairment, structured exercise is supported as a frontline intervention, with aerobic exercise showing particularly large effects on global cognition.

Several lifestyle factors amplify the brain benefits of exercise. Sleep enables memory consolidation, and exercise improves sleep quality — creating a virtuous cycle. Proper nutrition provides the raw materials for BDNF synthesis. Social engagement, which is naturally built into group exercise, dance, and team sports, independently supports cognitive function. Chekroud et al.'s analysis found that team sports were associated with the lowest mental health burden of any exercise type — likely because they combine physical activity with social connection.

What we still don't know

Honesty about uncertainty is central to how we approach science at Brain Zone, and there are genuine open questions in this field.

Higher-quality studies consistently produce more modest effect sizes than lower-quality ones. A 2025 umbrella review of 133 meta-analyses emphasizes that while the direction of evidence is clear, methodological rigor tempers the magnitude of effects. Cognitive improvements in healthy young adults, specifically, remain modest and sometimes inconsistent under the most rigorous study designs.

Individual variability in response to exercise is substantial and poorly understood. Genetic differences — including a common variant in the BDNF gene called Val66Met — may influence how much your brain responds to training. Sex differences are also emerging, with women showing greater episodic memory benefits but sometimes smaller BDNF responses.

Exercise studies face an inherent blinding problem: participants know whether they're exercising. Active control groups help, but they're imperfect. And in long-term epidemiological studies, there's always the question of reverse causation — people in the earliest stages of neurodegeneration may reduce their activity years before diagnosis, making exercise appear more protective than it truly is.

These limitations are real, but they don't undermine the core conclusion. The consistency of findings across thousands of studies, multiple research methodologies, and diverse populations provides a level of confidence that few health interventions can match.

Emerging research is now pursuing exercise-derived blood factors that could potentially be delivered as pharmaceuticals. Horowitz et al. demonstrated in Science that plasma from exercised mice transferred neurogenic benefits to sedentary mice, suggesting that specific molecules in the blood carry exercise's brain benefits. The muscle-brain axis — irisin, cathepsin B, lactate, and the kynurenine pathway — represents the frontier where molecular biology meets the possibility of exercise in a pill.

The bottom line

The evidence is clear in its direction, even as precise effect sizes continue to be refined. Exercise improves brain health through at least seven reinforcing biological mechanisms. It enhances every major cognitive domain. It treats depression with efficacy comparable to first-line medications. And it reduces dementia risk by 40–60% in the best longitudinal studies.

Three findings stand out as particularly surprising. First, resistance training may rival aerobic exercise for executive function — the cognitive domain most relevant to daily decision-making and productivity. Second, the runner's high is an endocannabinoid phenomenon, not an endorphin one, rewriting a 40-year scientific narrative. Third, the epigenetic changes triggered by exercise may transmit cognitive benefits to the next generation, suggesting that the effects of physical activity extend far beyond the individual.

But the most consequential takeaway is also the simplest. A moderate walking program — the most accessible, lowest-barrier form of exercise — is sufficient to grow new hippocampal neurons, elevate BDNF, reduce neuroinflammation, and protect against cognitive decline. You don't need a gym membership, expensive equipment, or athletic ability. The best exercise for your brain is the one you'll actually do.

---

Sources:

Agudelo, L.Z., et al. (2014). Skeletal muscle PGC-1α1 modulates kynurenine metabolism and mediates resilience to stress-induced depression. Cell, 159(1), 33–45. doi:10.1016/j.cell.2014.07.051

Aghjayan, S.L., et al. (2022). Aerobic exercise, cardiorespiratory fitness, and the human hippocampus. Communications Medicine, 2, 75. doi:10.1038/s43856-022-00079-7

Blondell, S.J., et al. (2014). Does physical activity prevent cognitive decline and dementia? BMC Public Health, 14, 510. doi:10.1186/1471-2458-14-510

Blumenthal, J.A., et al. (1999). Effects of exercise training on older patients with major depression. Archives of Internal Medicine, 159(19), 2349–2356. doi:10.1001/archinte.159.19.2349

Boström, P., et al. (2012). A PGC1-α-dependent myokine that drives brown-fat-like development of white fat and thermogenesis. Nature, 481, 463–468. doi:10.1038/nature10777

Chekroud, S.R., et al. (2018). Association between physical exercise and mental health in 1.2 million individuals in the USA. The Lancet Psychiatry, 5(9), 739–746. doi:10.1016/S2215-0366(18)30227-X

Colcombe, S., & Kramer, A.F. (2003). Fitness effects on the cognitive function of older adults: A meta-analytic study. Psychological Science, 14(2), 125–130. doi:10.1111/1467-9280.t01-1-01430

Colcombe, S.J., et al. (2006). Aerobic exercise training increases brain volume in aging humans. Journal of Gerontology A, 61(11), 1166–1170. doi:10.1093/gerona/61.11.1166

El Hayek, L., et al. (2019). Lactate mediates the effects of exercise on learning and memory through SIRT1-dependent activation of hippocampal brain-derived neurotrophic factor. Journal of Neuroscience, 39(13), 2369–2382. doi:10.1523/JNEUROSCI.1661-18.2019

Erickson, K.I., et al. (2010). Physical activity predicts gray matter volume in late adulthood. Neurology, 75(16), 1415–1422. doi:10.1212/WNL.0b013e3181f88359

Erickson, K.I., et al. (2011). Exercise training increases size of hippocampus and improves memory. PNAS, 108(7), 3017–3022. doi:10.1073/pnas.1015950108

Fernandes, J., et al. (2017). Running throughout middle-age keeps old adult-born neurons wired. Neuroscience & Biobehavioral Reviews, 77, 143–159. doi:10.1016/j.neubiorev.2017.03.007

Fuss, J., et al. (2015). A runner's high depends on cannabinoid receptors in mice. PNAS, 112(42), 13105–13108. doi:10.1073/pnas.1514996112

Gleeson, M., et al. (2011). The anti-inflammatory effects of exercise. Nature Reviews Immunology, 11, 607–615. doi:10.1038/nri3041

Gomez-Pinilla, F., et al. (2011). Exercise impacts brain-derived neurotrophic factor plasticity by engaging mechanisms of epigenetic regulation. European Journal of Neuroscience, 33(3), 383–390. doi:10.1111/j.1460-9568.2010.07508.x

Hillman, C.H., et al. (2014). Effects of the FITKids randomized controlled trial on executive control and brain function. Pediatrics, 134(4), e1063–e1071. doi:10.1542/peds.2013-3219

Horowitz, A.M., et al. (2020). Blood factors transfer beneficial effects of exercise on neurogenesis and cognition to the aged brain. Science, 369(6500), 167–173. doi:10.1126/science.aaw2622

Moon, H.Y., et al. (2016). Running-induced systemic cathepsin B secretion is associated with memory function. Cell Metabolism, 24(2), 332–340. doi:10.1016/j.cmet.2016.05.025

Morland, C., et al. (2017). Exercise induces cerebral VEGF and angiogenesis via the lactate receptor HCAR1. Nature Communications, 8, 15557. doi:10.1038/ncomms15557

Ngandu, T., et al. (2015). A 2 year multidomain intervention of diet, exercise, cognitive training, and vascular risk monitoring versus control to prevent cognitive decline in at-risk elderly people (FINGER). The Lancet, 385(9984), 2255–2263. doi:10.1016/S0140-6736(15)60461-5

Noetel, M., et al. (2024). Effect of exercise for depression: systematic review and network meta-analysis of randomised controlled trials. BMJ, 384, e075847. doi:10.1136/bmj-2023-075847

Nokia, M.S., et al. (2016). Physical exercise increases adult hippocampal neurogenesis in male rats provided it is aerobic and sustained. The Journal of Physiology, 594(7), 1855–1873. doi:10.1113/JP271552

Northey, J.M., et al. (2018). Exercise interventions for cognitive function in adults older than 50. British Journal of Sports Medicine, 52(3), 154–160. doi:10.1136/bjsports-2016-096587

Oppezzo, M., & Schwartz, D.L. (2014). Give your ideas some legs: The positive effect of walking on creative thinking. Journal of Experimental Psychology: Learning, Memory, and Cognition, 40(4), 1142–1152. doi:10.1037/a0036577

Pereira, A.C., et al. (2007). An in vivo correlate of exercise-induced neurogenesis in the adult dentate gyrus. PNAS, 104(13), 5638–5643. doi:10.1073/pnas.0611721104

Rehfeld, K., et al. (2018). Dance training is superior to repetitive physical exercise in inducing brain plasticity in the elderly. PLOS ONE, 13(7), e0196636. doi:10.1371/journal.pone.0196636

Rovio, S., et al. (2005). Leisure-time physical activity at midlife and the risk of dementia and Alzheimer's disease. The Lancet Neurology, 4(11), 705–711. doi:10.1016/S1474-4422(05)70198-8

Seifert, T., et al. (2010). Endurance training enhances BDNF release from the human brain. American Journal of Physiology — Regulatory, Integrative and Comparative Physiology, 298(2), R372–R377. doi:10.1152/ajpregu.00525.2009

Siebers, M., et al. (2021). Exercise-induced euphoria and anxiolysis do not depend on endogenous opioids in humans. Psychoneuroendocrinology, 126, 105173. doi:10.1016/j.psyneuen.2021.105173

Singh, B., et al. (2023). Effectiveness of physical activity interventions for improving depression, anxiety and distress: an overview of systematic reviews. British Journal of Sports Medicine, 57(18), 1203–1209. doi:10.1136/bjsports-2022-106195

Sleiman, S.F., et al. (2016). Exercise promotes the expression of brain derived neurotrophic factor (BDNF) through the action of the ketone body β-hydroxybutyrate. eLife, 5, e15092. doi:10.7554/eLife.15092

Szuhany, K.L., et al. (2015). A meta-analytic review of the effects of exercise on brain-derived neurotrophic factor. Journal of Psychiatric Research, 60, 56–64. doi:10.1016/j.jpsychires.2014.10.003

van Praag, H., et al. (1999). Running enhances neurogenesis, learning, and long-term potentiation in mice. PNAS, 96(23), 13427–13431. doi:10.1073/pnas.96.23.13427

van Praag, H., et al. (1999). Running increases cell proliferation and neurogenesis in the adult mouse dentate gyrus. Nature Neuroscience, 2(3), 266–270. doi:10.1038/6368