Longevity: The Science and Secrets of Living a Long, Healthy Life
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Longevity: The Science and Secrets of Living a Long, Healthy Life

Discover the scientific secrets to longevity and living a long, healthy life. Learn about the key factors that determine lifespan, evidence-based diets and lifestyle changes for increasing longevity, and cutting-edge research aimed at extending the human healthspan.

Introduction

The quest for longevity - living a long and healthy life - has fascinated humans for millennia. In recent decades, scientific research has begun to unravel many of the biological mechanisms that influence aging and lifespan. By understanding the complex interplay of genetics, lifestyle factors, and environment, we can develop evidence-based strategies to maximize our chances of living longer, disease-free lives. This in-depth article explores the science of longevity, including how it is defined and measured, its key determinants, diet and lifestyle approaches to increasing lifespan, and breakthrough research that could extend human healthspan in the future.

Defining and Measuring Longevity

Longevity broadly refers to the length or duration of life. More specifically, it looks at the typical length of life of a species or the average length of life of a population or subgroup. Closely related concepts include life expectancy (the average number of years an individual is expected to live), lifespan (the maximum number of years an individual of a species can live), and healthspan (the number of years lived free of chronic disease and disability).

Longevity is typically measured and expressed as life expectancy from birth. Global life expectancy has increased dramatically in the past century, rising from just 32 years in 1900 to over 72 years today. However, life expectancy varies significantly between countries, from around 54 years in low-income nations to over 81 years in high-income countries like Japan and Hong Kong.

While life expectancy measures length of life, healthspan has emerged as an important metric for quality of life in older age. Healthspan can be quantified in various ways, such as healthy life expectancy (years lived in good health) or disability-free life expectancy. Increasing healthspan - not just lifespan - is a key focus of longevity research.

Key Determinants of Longevity

Longevity is influenced by a complex web of factors, including genetics, lifestyle behaviors, socioeconomic conditions, healthcare, and environment. While estimates vary, studies suggest that genetics account for approximately 20-30% of variation in human lifespan. Specific genes impacting pathways like insulin signaling, cell maintenance, telomere length, and others have been associated with longevity in humans and animal models.

However, research indicates that lifestyle and environmental factors play a larger role than genetics in determining lifespan for most individuals. Behavioral factors like sleep, exercise, nutrition, social connection, and emotional well-being have a major impact on mortality risk and longevity. Studies estimate that adopting healthy habits could increase life expectancy by over a decade.

Socioeconomic factors like income, education, occupation, and access to healthcare are also critical determinants of longevity. Research has consistently found a strong association between socioeconomic status and life expectancy, with individuals of higher status living significantly longer on average. This disparity is likely mediated by a range of factors, including greater access to healthy foods, safe living environments, quality healthcare, and reduced chronic stress.

Environmental exposures, such as air pollution, water contamination, infectious diseases, and climate-related events, can also substantially impact longevity, particularly in low- and middle-income countries. For example, the World Health Organization estimates that air pollution alone causes around 7 million premature deaths globally each year.

Nutrition and Longevity

Dietary habits are one of the most important modifiable risk factors for chronic disease and premature mortality. Extensive research has identified several eating patterns associated with increased longevity and reduced disease risk:

The Mediterranean diet, characterized by high intake of fruits, vegetables, whole grains, legumes, nuts, olive oil, and fish, and low intake of red meat, sugar, and processed foods. Numerous observational studies and clinical trials have linked greater adherence to a Mediterranean diet with increased lifespan and healthspan.

The Okinawan diet, traditional among the long-lived population of Okinawa, Japan. The diet features high amounts of vegetables (particularly sweet potatoes), soy foods, fish, seaweed, and green tea, and low amounts of refined grains, sugar, and animal products. Older Okinawans following this diet have historically had among the world's highest life expectancies and lowest rates of age-related chronic disease.

Plant-based diets, which minimize or eliminate animal products and emphasize whole plant foods like fruits, vegetables, whole grains, legumes, nuts, and seeds. Observational studies have found that vegetarian and vegan diets are associated with significantly lower risks of obesity, type 2 diabetes, cardiovascular disease, and certain cancers. A comprehensive analysis estimated that widespread adoption of plant-based diets could prevent over 10 million premature deaths annually.

Calorie restriction, reducing calorie intake below normal levels without causing malnutrition. Long-term calorie restriction has been shown to extend lifespan in numerous animal models. Observational studies in humans and short-term trials suggest that even moderate calorie restriction may slow biological aging and protect against age-related disease. However, the long-term impacts of calorie restriction in humans are not yet clear.

Emerging research is also probing the longevity potential of dietary approaches like intermittent fasting, low-protein high-carbohydrate diets, and pharmacological agents that mimic the effects of calorie restriction. More studies are needed to clarify the efficacy and safety of these approaches for lifespan extension.

Regardless of the specific diet, research indicates that favoring minimally processed, nutrient-dense foods like fruits, vegetables, whole grains, legumes, nuts, seeds, herbs, and spices can significantly lower disease risk and support healthy longevity. Some specific foods that may have particular longevity benefits include:

  • Leafy green vegetables like spinach, kale, and collards
  • Berries, especially blueberries, blackberries, and raspberries
  • Nuts and seeds, particularly walnuts, almonds, and flax seeds
  • Legumes like beans, lentils, peas, and soy foods
  • Whole grains such as oats, quinoa, brown rice, and whole wheat
  • Fatty fish rich in omega-3s, like salmon, sardines, and anchovies
  • Fermented foods containing beneficial bacteria, such as yogurt, kimchi, and sauerkraut
  • Green tea and coffee, sources of antioxidants and other protective compounds
  • Herbs and spices like turmeric, ginger, garlic, and rosemary

Conversely, high intakes of red and processed meats, refined grains, added sugars, trans fats, and alcohol have been linked to increased chronic disease risk and shorter life expectancy. Limiting these in the diet while favoring whole, minimally processed foods is a key dietary strategy for healthy aging.

The Standard American Diet (SAD), characterized by high intakes of processed foods, refined grains, added sugars, unhealthy fats, and animal-based products, is failing to support the health and well-being of the American population. This dietary pattern, which is low in fruits, vegetables, whole grains, and other nutrient-dense foods, has been linked to increased risks of obesity, type 2 diabetes, cardiovascular disease, certain cancers, and other chronic conditions that contribute to reduced life expectancy and impaired quality of life in older age. The SAD's emphasis on convenience and palatability over nutrition has led to widespread overconsumption of calories and inadequate intake of essential vitamins, minerals, and beneficial plant compounds. Consequently, many Americans are overfed but undernourished, lacking the dietary foundation for optimal health and longevity. Transitioning away from the SAD towards a more whole food, plant-centered eating pattern is a critical step in addressing the nation's chronic disease epidemic and promoting healthy aging.

The impact of the SAD on healthy aging and longevity is particularly concerning given the rapid growth of the older adult population. As more Americans live into their 80s, 90s, and beyond, maintaining good health and functional ability in later life is becoming increasingly important. However, the cumulative effects of poor dietary habits over a lifetime can accelerate the aging process and increase the risk of age-related diseases like heart disease, stroke, cognitive decline, and physical disability. By promoting chronic inflammation, oxidative stress, and metabolic dysfunction, the SAD may be compromising the healthspan and lifespan of older people and preventing more Americans from achieving exceptional longevity.

In contrast, the dietary patterns of centenarians and other long-lived populations around the world, such as the Japanese Okinawans and Italian Sardinians, are rich in nutrient-dense, plant-based foods and low in processed, animal-based products. These eating habits, in combination with other healthy lifestyle factors like regular physical activity and strong social connections, are thought to contribute to their remarkable longevity and extended years of life in good health. While genetics likely play some role in exceptional lifespan, research suggests that lifestyle factors are the key determinants of healthy aging and that even individuals without longevity-associated genes can maximize their potential for a long life through healthy behaviors.

The field of geroscience, which combines insights from gerontology and neuroscience, is beginning to unravel the complex biological mechanisms that link diet to aging and age-related disease. Scientists have identified several key pathways, such as those involved in inflammation, nutrient sensing, and cellular repair, that are modulated by dietary factors and may influence the rate of aging and risk of chronic disease. Emerging research suggests that dietary interventions, such as calorie restriction and intermittent fasting, can target these pathways to promote healthy aging and extend healthspan in animal models. However, more research is needed to translate these findings to humans and develop safe, effective dietary strategies for long-term health promotion.

To address the public health burden of the SAD and support healthy aging, a multifaceted approach is needed. This should include efforts to improve nutrition education, promote healthy food environments, and increase access to affordable, nutritious foods, particularly in underserved communities. Healthcare providers, especially those in geriatrics and primary care, also have an important role to play in assessing dietary habits, providing personalized nutrition guidance, and referring patients to registered dietitians for more intensive counseling. By making healthy eating a national priority and providing the resources and support needed to facilitate dietary change, we can help more Americans achieve a long, healthy life free of chronic disease and disability.

Exercise for Longevity

In addition to diet, a range of lifestyle behaviors can impact lifespan and healthspan:

Physical activity: Regular exercise is one of the most powerful interventions for healthy aging. A wealth of research has shown that physically active individuals have significantly lower risks of chronic diseases, cognitive decline, disability, and premature death compared to inactive individuals. Major health organizations recommend that older adults get at least 150 minutes per week of moderate-intensity aerobic activity (like brisk walking) and twice-weekly strength training. Even light activity like household chores has longevity benefits compared to being sedentary.

Physical activity is a cornerstone of healthy aging, with a robust body of evidence demonstrating its wide-ranging benefits for physical, cognitive, and emotional well-being across the lifespan. Engaging in regular exercise can help prevent or manage many of the chronic diseases that commonly affect older adults, such as heart disease, stroke, type 2 diabetes, osteoporosis, and certain cancers. Moreover, physical activity has been shown to enhance brain health, improve memory and cognitive function, and reduce the risk of dementia and Alzheimer's disease.

The benefits of exercise for older adults extend beyond disease prevention. Regular physical activity can help maintain muscle mass, strength, and flexibility, which are essential for performing daily activities and maintaining independence in later life. Exercise also promotes better balance and coordination, reducing the risk of falls and fall-related injuries, which are a major cause of disability and mortality among older people. Additionally, physical activity has been linked to improved sleep quality, reduced symptoms of depression and anxiety, and enhanced overall quality of life.

Given the numerous health advantages of exercise, major health organizations like the World Health Organization (WHO) and the American College of Sports Medicine (ACSM) have issued specific physical activity guidelines for older adults. These recommendations typically include at least 150 minutes per week of moderate-intensity aerobic activity, such as brisk walking, cycling, or swimming, as well as muscle-strengthening activities involving major muscle groups at least twice per week. Older adults with chronic conditions or functional limitations may need to modify these guidelines based on their individual abilities and health status, emphasizing the importance of personalized exercise prescription.

Importantly, the benefits of physical activity are not limited to structured exercise sessions. Engaging in light activities throughout the day, such as household chores, gardening, or leisurely walks, can also contribute to overall physical activity levels and provide longevity benefits. In fact, studies have shown that even short bouts of movement, like standing up and walking around for a few minutes every hour, can help counteract the negative health effects of prolonged sitting and sedentary behavior.

Despite the well-established benefits of physical activity, many older adults remain insufficiently active. Barriers to exercise participation may include chronic health conditions, functional limitations, lack of motivation, or environmental factors like access to safe and affordable exercise facilities. To promote physical activity among older people, a comprehensive approach is needed that includes education about the benefits of exercise, individualized activity plans, and community-based programs that provide social support and opportunities for engagement.

Healthcare providers, particularly those in geriatrics and primary care, play a crucial role in assessing physical activity levels, providing exercise counseling, and referring patients to appropriate resources like physical therapy or community exercise programs. Innovative approaches, such as the use of wearable activity trackers or mobile health applications, may also help older adults monitor their progress and stay motivated to maintain an active lifestyle.

As the global population ages, promoting regular physical activity among older adults is becoming increasingly important for public health. By prioritizing exercise as a key component of healthy aging, we can help more individuals maintain their physical function, cognitive abilities, and overall well-being into advanced age, ultimately contributing to increased healthspan and lifespan. Through a combination of individual efforts, community-based interventions, and supportive policies, we can create a society that values and enables lifelong physical activity for all.

Sleep and Longevity

Both short and long sleep duration have been linked to increased mortality risk, with 7-8 hours per night appearing optimal for most adults. Sleep quality is also critical, with conditions like insomnia and sleep apnea tied to greater disease risk. Sleep supports longevity through a variety of mechanisms, including memory consolidation, immune system regulation, and clearance of toxic proteins from the brain.

Sleep is a vital biological process that plays a critical role in maintaining physical and mental health throughout the lifespan. Growing evidence suggests that both sleep quantity and quality are important predictors of longevity, with a sleep duration of 7-8 hours per night being optimal for most adults. Studies have consistently shown that individuals who report sleeping less than 6 hours or more than 9 hours per night have a higher risk of all-cause mortality, as well as an increased incidence of chronic diseases such as obesity, type 2 diabetes, cardiovascular disease, and certain cancers.

In addition to sleep duration, sleep quality is a crucial factor in promoting health and longevity. Sleep disorders like insomnia and sleep apnea, which are characterized by difficulty falling or staying asleep and repeated breathing interruptions during sleep, respectively, have been linked to a heightened risk of chronic disease and premature mortality. These conditions can lead to fragmented sleep, reduced time spent in deep, restorative sleep stages, and daytime fatigue, all of which can take a toll on physical and cognitive function over time.

The relationship between sleep and longevity is complex and multifaceted, with sleep influencing a wide range of physiological processes that are essential for health and survival. One key function of sleep is memory consolidation, the process by which new information and experiences are transformed into long-term memories. During sleep, particularly during the rapid eye movement (REM) stage, the brain is highly active and works to strengthen and integrate new neural connections, facilitating learning and cognitive performance. Insufficient or poor-quality sleep can impair this process, leading to memory deficits and cognitive decline over time.

Sleep also plays a critical role in regulating the immune system, with chronic sleep deprivation being associated with increased inflammation, reduced immune cell function, and impaired wound healing. During sleep, the body produces cytokines and other immune-regulating compounds that help fight off infections and repair damaged tissues. Adequate sleep is also important for maintaining a healthy balance of hormones like cortisol and melatonin, which are involved in stress response and sleep-wake regulation, respectively.

Recent research has also highlighted the importance of sleep in clearing toxic proteins from the brain, particularly beta-amyloid and tau, which are associated with the development of Alzheimer's disease and other neurodegenerative disorders. During sleep, the brain's glymphatic system, a network of channels that allows cerebrospinal fluid to flow through the brain and remove waste products, becomes highly active. This process helps prevent the accumulation of neurotoxic proteins and may protect against age-related cognitive decline and dementia.

Given the numerous ways in which sleep supports health and longevity, promoting healthy sleep habits should be a key priority for individuals and public health efforts aimed at increasing healthspan and lifespan. This includes maintaining a consistent sleep schedule, creating a sleep-conducive environment (e.g., dark, quiet, and cool), avoiding stimulating activities before bedtime, and managing stress through relaxation techniques or cognitive-behavioral therapy.

For individuals with sleep disorders like insomnia or sleep apnea, seeking evaluation and treatment from a sleep medicine specialist can be critical for improving sleep quality and reducing associated health risks. Treatment options may include lifestyle modifications, cognitive-behavioral therapy for insomnia (CBT-I), continuous positive airway pressure (CPAP) therapy for sleep apnea, or medications in some cases.

In addition to individual efforts, addressing the societal factors that contribute to sleep deprivation and poor sleep quality, such as long work hours, shift work, and the increasing use of electronic devices, will be important for promoting healthy sleep at a population level. This may involve implementing workplace policies that prioritize work-life balance, providing education about healthy sleep habits, and developing technologies that minimize the impact of electronic devices on sleep.

As our understanding of the complex relationship between sleep and longevity continues to grow, it is becoming increasingly clear that prioritizing healthy sleep is a critical component of any comprehensive strategy for promoting healthy aging and extending healthspan. By recognizing the importance of sleep as a pillar of health alongside diet and physical activity, we can help individuals and communities achieve the goal of living longer, healthier lives.

Social Connection and Longevity

A robust body of research has linked stronger social relationships and social support to lower mortality risk and greater longevity. Conversely, social isolation and loneliness may accelerate aging and disease onset. Cultivating strong, supportive social ties through family, friends, and community appears to be an important component of a longevity-promoting lifestyle.

Social connection is a fundamental human need that plays a critical role in promoting health, well-being, and longevity. A growing body of research has consistently demonstrated that individuals with strong social relationships and support networks have a lower risk of mortality and a greater likelihood of living a long, healthy life. Conversely, social isolation and loneliness have been identified as significant risk factors for premature death, with some studies suggesting that their impact on mortality may be comparable to that of smoking or obesity.

The link between social connection and longevity is thought to be mediated by a variety of psychological, behavioral, and physiological mechanisms. On a psychological level, social relationships provide a sense of belonging, purpose, and meaning in life, which can enhance mental health and resilience in the face of stress and adversity. Socially connected individuals are more likely to engage in health-promoting behaviors, such as exercising regularly, eating a healthy diet, and seeking preventive healthcare, in part because they have the support and encouragement of their loved ones.

Moreover, social support has been shown to buffer the negative effects of stress on the body by modulating the physiological stress response. Chronic stress is associated with increased inflammation, impaired immune function, and accelerated cellular aging, all of which can contribute to the development of age-related diseases like cardiovascular disease, diabetes, and cancer. By providing a sense of security and emotional regulation, social connections can help reduce the burden of stress on the body and promote healthy aging at the cellular level.

Conversely, social isolation and loneliness have been linked to a range of negative health outcomes, including increased risk of depression, cognitive decline, and cardiovascular disease. Loneliness has been associated with elevated levels of stress hormones, inflammation, and impaired immune function, which may accelerate the aging process and increase vulnerability to disease. Additionally, socially isolated individuals may be less likely to engage in health-promoting behaviors and may have limited access to resources and support in times of need.

Given the profound impact of social connection on health and longevity, cultivating strong, supportive relationships should be a key priority for individuals seeking to optimize their healthspan and lifespan. This involves nurturing existing relationships with family and friends, as well as seeking out new opportunities for social engagement through community activities, volunteer work, or shared interests. For older adults, maintaining social connections may be particularly important for preserving cognitive function, preventing disability, and enhancing overall quality of life.

At a societal level, promoting social cohesion and reducing social isolation should be a public health priority. This may involve developing community-based programs that provide opportunities for social interaction and support, particularly for vulnerable populations like older adults, individuals with disabilities, or those living in poverty. Additionally, addressing the social determinants of health, such as access to education, employment, and affordable housing, can help reduce inequalities and promote social connectedness across the lifespan.

In the healthcare setting, assessing social connectedness and addressing loneliness should be a routine part of patient care, particularly for older adults or those with chronic conditions. Healthcare providers can play a key role in identifying individuals at risk for social isolation and referring them to appropriate resources, such as support groups, counseling services, or community-based programs.

As technology continues to evolve, there may be new opportunities to promote social connection and support through digital platforms and interventions. However, it will be important to ensure that these technologies are designed and implemented in a way that enhances, rather than replaces, authentic human interaction and connection.

Ultimately, recognizing the fundamental importance of social connection for health and longevity requires a shift in perspective that values relationships and community as much as individual lifestyle choices. By prioritizing social connectedness as a key component of a healthy, fulfilling life, we can create a society that supports the well-being and longevity of all its members, from infancy to advanced age.

Other Lifestyle Choices and Longevity

Stress management: Chronic psychological stress has been associated with accelerated aging and higher risk of age-related disease. Stress may shorten lifespan by increasing inflammation, impairing immune function, and promoting unhealthy behaviors. Engaging in stress-reducing activities like meditation, yoga, time in nature, and social connection may buffer the negative impacts of stress and support healthy aging.

Alcohol and substance use: Excessive alcohol intake and smoking are major risk factors for chronic disease and early death. However, research suggests that light-to-moderate alcohol consumption (up to 1 drink per day for women and 1-2 for men) may have modest longevity benefits compared to complete abstinence or heavy drinking. The healthiest approach is to avoid smoking entirely and limit alcohol intake.

Preventive medical care: Routine medical check-ups, screenings, and preventive care can help detect and treat potential health issues early, lowering risk of disease progression and premature mortality. Key preventive measures include immunizations, cancer screenings, monitoring of cardiovascular risk factors, and management of chronic conditions like hypertension and type 2 diabetes.

The Blue Zones: Longevity Lessons from the World's Longest-Lived Populations
The Blue Zones are five regions around the world where people live extraordinarily long, healthy lives. These include Okinawa, Japan; Sardinia, Italy; Nicoya, Costa Rica; Ikaria, Greece; and Loma Linda, California. Extensive research in these populations has identified common lifestyle traits that may explain their exceptional longevity:

  • Plant-based diets rich in fruits, vegetables, whole grains, legumes, and nuts
  • Regular, lifelong physical activity, often as part of daily life (walking, gardening, etc.)
  • Strong family and community social connections
  • Daily practices that reduce stress, like napping, prayer, or spending time in nature
  • Sense of purpose in life

While genes likely play some role, studies suggest that environment and lifestyle are the primary drivers of longevity in Blue Zones. These populations demonstrate that a long, healthy lifespan is possible with the right behavioral practices and social conditions.

Advancing Research: The Future of Longevity Science Recent decades have seen major progress in understanding the biological mechanisms of aging and identifying potential targets for lifespan extension. Much of this work has been conducted in animal models like yeast, worms, flies, and mice, with a growing focus on translation to humans. Some key areas of research include:

Genomics and epigenetics: Analyzing the genomes of long-lived individuals and populations to find genes and variants that influence aging and longevity. Epigenetic changes, or modifications to gene expression without altering DNA sequence, are also being studied as potential drivers of aging.

Cellular senescence: As cells age, they can enter a state of senescence where they stop dividing but remain metabolically active and secrete inflammatory factors. Targeting senescent cells with drugs (senolytics) or limiting their accumulation may slow aging and extend healthspan.

Nutrient signaling pathways: Manipulating cellular nutrient-sensing pathways, like insulin/IGF-1 and mTOR, has been shown to extend lifespan in organisms from yeast to primates. Drugs that inhibit these pathways, like rapamycin and metformin, are being studied for potential anti-aging effects in humans.

Mitochondrial dysfunction: Declines in mitochondrial function are thought to be a key driver of aging and age-related disease. Strategies to boost mitochondrial health, such as exercise, calorie restriction, and targeted drugs, may support longevity.

Stem cells: Stem cell function declines with age, impairing tissue regeneration and repair. Approaches to enhance stem cell survival and activity, like systemic factors from young blood, are being explored for healthspan extension.

Other active areas include autophagy (cellular "self-eating" that clears damaged components), proteostasis (maintenance of proper protein folding and function), NAD+ metabolism, and the gut microbiome. By better understanding and developing interventions to target these fundamental aging processes, scientists aim to extend human healthspan by preventing or delaying chronic disease onset.

However, significant challenges remain in translating findings from basic aging research into effective therapies for humans. More clinical trials are needed to establish the safety and efficacy of potential anti-aging compounds and to identify biomarkers of biological aging. Ethical and equitable distribution of future longevity-enhancing therapies is also an important consideration.

Conclusion


The science of longevity has advanced rapidly in recent years, shedding new light on the complex factors that influence human lifespan and healthspan. While our genes play a role, the majority of variation in longevity appears to be determined by modifiable lifestyle and environmental factors. Eating a healthy diet, staying physically active, getting sufficient sleep, managing stress, cultivating social connections, and receiving preventive healthcare can significantly increase the odds of a long, disease-free life. The world's Blue Zones demonstrate that a lifespan of 100+ years is possible with the right behavioral practices and living conditions.

Meanwhile, research is uncovering the cellular and molecular hallmarks of aging and developing interventions to slow their progression. In the coming years, novel therapies that target fundamental aging processes could help further extend human healthspan. By combining a healthy lifestyle with breakthrough treatments, it may eventually be possible to achieve lifespans well beyond what is currently typical.

Ultimately, the goal of longevity science is not just more years lived, but more years spent in good health, free of chronic disease and disability. While immortality remains out of reach, the dream of a longer, healthier life is increasingly becoming a reality. By prioritizing our health, staying informed about scientific advances, and advocating for policies and environments that support healthy aging, we can all maximize our potential for a long, vibrant life.

References:

Olshansky, S. J. (2018). From Lifespan to Healthspan. JAMA, 320(13), 1323-1324. doi:10.1001/jama.2018.12621

Vaupel, J. W. (2010). Biodemography of human ageing. Nature, 464(7288), 536-542. doi:10.1038/nature08984

Kirkwood, T. B. (2005). Understanding the odd science of aging. Cell, 120(4), 437-447. doi:10.1016/j.cell.2005.01.027

Crimmins, E. M. (2015). Lifespan and healthspan: past, present, and promise. The Gerontologist, 55(6), 901-911. doi:10.1093/geront/gnv130

United Nations, Department of Economic and Social Affairs, Population Division (2019). World Population Prospects 2019: Highlights. ST/ESA/SER.A/423.

Robine, J. M., & Ritchie, K. (1991). Healthy life expectancy: evaluation of global indicator of change in population health. BMJ, 302(6774), 457-460. doi:10.1136/bmj.302.6774.457

Olshansky, S. J., Carnes, B. A., & Désesquelles, A. (2001). Prospects for human longevity. Science, 291(5508), 1491-1492. doi:10.1126/science.291.5508.1491

Herskind, A. M., McGue, M., Holm, N. V., Sörensen, T. I., Harvald, B., & Vaupel, J. W. (1996). The heritability of human longevity: a population-based study of 2872 Danish twin pairs born 1870–1900. Human genetics, 97(3), 319-323. doi:10.1007/BF02185763

Skytthe, A., Pedersen, N. L., Kaprio, J., Stazi, M. A., Hjelmborg, J. V., Iachine, I., ... & Christensen, K. (2003). Longevity studies in GenomEUtwin. Twin Research and Human Genetics, 6(5), 448-454. doi:10.1375/twin.6.5.448

Joshi, P. K., Pirastu, N., Kentistou, K. A., Fischer, K., Hofer, E., Schraut, K. E., ... & Wilsoen, J. F. (2017). Genome-wide meta-analysis associates HLA-DQA1/DRB1 and LPA and lifestyle factors with human longevity. Nature communications, 8(1), 1-13. doi:10.1038/s41467-017-00934-5

Kenyon, C. J. (2010). The genetics of aging. Nature, 464(7288), 504-512. doi:10.1038/nature08980

Mathers, J. C. (2015). Impact of nutrition on the aging process. British Journal of Nutrition, 113(S1), S18-S22. doi:10.1017/S0007114514003237

Knoops, K. T., de Groot, L. C., Kromhout, D., Perrin, A. E., Moreiras-Varela, O., Menotti, A., & Van Staveren