Maximum life span is a measure of the maximum amount of time one or more members of a group has been observed to survive between birth and death.

Definition

In animal studies, maximum life span is often taken to be the mean life span of the most long-lived 10% of a given cohort A cohort study or panel study is a form of longitudinal study used in medicine, social science and ecology. It is one type of study design and should be compared with a cross-sectional study. By another definition, however, maximum life span corresponds to the age at which the oldest known member of a species In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are often used, such as based on similarity of DNA or or experimental group has died. Calculation of the maximum life span in the former sense depends upon initial sample size.^[1]

Maximum life span is in contrast with mean life span (average life span Life expectancy is the expected number of years of life remaining at a given age. It is denoted by ex, which means the average number of subsequent years of life for someone now aged x, according to a particular mortality experience. (In technical literature, this symbol means the average number of complete years of life remaining, IE excluding or life expectancy Life expectancy is the expected number of years of life remaining at a given age. It is denoted by ex, which means the average number of subsequent years of life for someone now aged x, according to a particular mortality experience. (In technical literature, this symbol means the average number of complete years of life remaining, ie excluding). Mean life span varies with susceptibility to disease, accident An accident is a specific, unidentifiable, unexpected, unusual and unintended external action which occurs in a particular time and place, with no apparent and deliberate cause but with marked effects. It implies a generally negative outcome which may have been avoided or prevented had circumstances leading up to the accident been recognized, and, suicide Suicide is the term used for the deliberate self-destruction by a living being, resulting in their own death. Such actions are typically characterised as being made out of despair, or attributed to some underlying mental disorder which includes depression, bipolar disorder, schizophrenia, alcoholism and drug abuse. Financial difficulties, and homicide Murder, as defined in common law countries, is the unlawful killing of another human being with intent , and generally this state of mind distinguishes murder from other forms of unlawful homicide (such as manslaughter). As the loss of a human being inflicts enormous grief upon the individuals close to the victim, as well as the fact that the, whereas maximum life span is determined by "rate of aging".^[2]

In humans

The oldest recognized person The following tables list only the verified oldest people in world in ordinal rank, such as oldest person or oldest man. A supercentenarian is considered verified if their claim has been accepted by an international body that specifically deals in longevity research, such as the Gerontology Research Group or the Guinness Book of World Records. The on record is Jeanne Calment Jeanne Louise Calment had the longest confirmed human life span in history, living 122 years and 164 days (44,724 days total). She lived in Arles, France, for her entire life, and outlived both her daughter and grandson. She became especially well known from the age of 113, when the centenary of Vincent van Gogh's visit brought reporters to Arles,, a French woman To be French, according to the first article of the Constitution, is to be a citizen of France, regardless of one's origin, race, or religion . According to its principles, France has devoted herself the destiny of a proposition nation, a generic territory where people are bounded only by the French language and the assumed willingness to live who lived for 122 years and 164 days. Maximum recorded life span for humans has remained about 105−122 calendar years throughout recorded history, despite steady improvements in life expectancy Life expectancy is the expected number of years of life remaining at a given age. It is denoted by ex, which means the average number of subsequent years of life for someone now aged x, according to a particular mortality experience. (In technical literature, this symbol means the average number of complete years of life remaining, ie excluding. Reduction of infant mortality has accounted for most of this increased average longevity The word "longevity" is sometimes used as a synonym for "life expectancy" in demography or to connote "long life", especially when it concerns someone or something lasting longer than expected, but since the 1960s mortality rates Mortality rate is a measure of the number of deaths in some population, scaled to the size of that population, per unit time. Mortality rate is typically expressed in units of deaths per 1000 individuals per year; thus, a mortality rate of 9.5 in a population of 100,000 would mean 950 deaths per year in that entire population. It is distinct from among those over 80 years have decreased by about 1.5% per year. Advances in medicine,^[which?] calorie restriction Calorie restriction, or caloric restriction , is a dietary regimen that restricts calorie intake, where the baseline for the restriction varies, usually being the previous, unrestricted, intake of the subjects. CR when not associated with malnutrition, is thought to improve age-related health and to slow the aging process in some animals and fungi with adequate nutrition, or other interventions are said to have slowed the aging process Senescence or biological aging is the change in the biology of an organism as it ages after its maturity. Such changes range from those affecting its cells and their function to that of the whole organism. There are a number of theories as to why senescence occurs, including ones that claim it is programmed by gene expression changes and that it. Although calorie restriction has not been proven to extend the maximum human life span, as of 2006, results in ongoing primate studies are promising.^[3]

In other animals

Small animals such as birds Birds are winged, bipedal, endothermic (warm-blooded), egg-laying, vertebrate animals. There are around 10,000 living species, making them the most varied of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from the 5 cm (2 in) Bee Hummingbird to the 2.75 m (9 ft) Ostrich and squirrels Squirrels belong to a large family of small or medium-sized rodents called the Sciuridae. The family includes tree squirrels, ground squirrels, chipmunks, marmots , flying squirrels, and prairie dogs. Squirrels are indigenous to the Americas, Eurasia, and Africa and have been introduced to Australia. Squirrels are first attested in the Eocene, rarely live to their maximum life span, usually dying of accidents An accident is a specific, unidentifiable, unexpected, unusual and unintended external action which occurs in a particular time and place, with no apparent and deliberate cause but with marked effects. It implies a generally negative outcome which may have been avoided or prevented had circumstances leading up to the accident been recognized, and, disease A disease is an abnormal condition affecting the body of an organism. It is often construed to be a medical condition associated with specific symptoms and signs. It may be caused by external factors, such as infectious disease, or it may be caused by internal disfunctions, such as autoimmune diseases or predation In ecology, predation describes a biological interaction where a predator feeds on its prey (the organism that is attacked). Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption. The other main. Grazing animals accumulate wear and tear to their teeth to the point where they can no longer eat, and they die of starvation Starvation is a severe reduction in vitamin, nutrient and energy intake. It is the most extreme form of malnutrition. In humans, prolonged starvation can cause permanent organ damage[citation needed] and eventually, death. The term inanition refers to the symptoms and effects of starvation.

The maximum life span of most species has not been accurately determined, because the data collection has been minimal and the number of species studied in captivity (or by monitoring in the wild) has been small.

Maximum life span is usually longer for species that are larger or have effective defenses against predation, such as bird flight, tortoise shells, porcupine quills, or large primate brains. When compared to primates, of the approximately 20,000 to 25,000 genes in the human genome In modern molecular biology, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA, it is estimated that 6% of these are different from those of a chimpanzee which has an average lifespan of only 52 years, in contrast to the human lifespan. The difference in longevity The word "longevity" is sometimes used as a synonym for "life expectancy" in demography or to connote "long life", especially when it concerns someone or something lasting longer than expected between humans and chimps could be due to as few as a hundred genes A gene is a unit of heredity in a living organism. It is normally a stretch of DNA that codes for a type of protein or for an RNA chain that has a function in the organism. All living things depend on genes, as they specify all proteins and functional RNA chains. Genes hold the information to build and maintain an organism's cells and pass genetic or fewer; however there may be other factors that shorten the life span of chimpanzees.

The differences in life span between species demonstrate the role of genetics Genetics , a discipline of biology, is the science of heredity and variation in living organisms. The fact that living things inherit traits from their parents has been used since prehistoric times to improve crop plants and animals through selective breeding. However, the modern science of genetics, which seeks to understand the process of in determining maximum life span ("rate of aging"). The records (in years) are these:

• for common house mouse The house mouse is a small rodent, a mouse, one of the most numerous species of the genus Mus, 4^[4]
• for dogs The dog is a domesticated form of the gray wolf, a member of the Canidae family of the order Carnivora. The term is used for both feral and pet varieties. The domestic dog has been one of the most widely kept working and companion animals in human history, 29, in Australia Canine conditions such as temperature, change, hearing, and skin condition often degrade with geriatric age, and medical conditions such as cancer, renal failure, arthritis and joint conditions, and other signs of old age may appear
• for cats The cat , also known as the domestic cat or housecat to distinguish it from other felines and felids, is a small domesticated carnivorous mammal that is valued by humans for its companionship and ability to hunt vermin and household pests. Cats have been associated with humans for at least 9,500 years, and are currently the most popular pet in the, 36^[5] (England, unconfirmed)
• for polar bears The polar bear is a bear native largely within the Arctic circle encompassing the Arctic Ocean, its surrounding seas and surrounding land masses. It is the world's largest land carnivore and also the largest bear, together with the omnivorous Kodiak bear, which is approximately the same size. An adult male weighs around 350–680 kg (770–1,500, 42^[6] (Debby Debby was the world's oldest polar bear. She lived in the Assiniboine Park Zoo in Winnipeg. In August 2008, the Guinness Book of World Records certified her as not only the oldest polar bear, but one of the three oldest individuals ever recorded of all bear species)
• for goldfish The goldfish is a freshwater fish in the family Cyprinidae of order Cypriniformes. It was one of the earliest fish to be domesticated, and is one of the most commonly kept aquarium fish, 49^[7]
• for horses The horse is a hooved (ungulate) mammal, a subspecies of the family Equidae. The horse has evolved over the past 45 to 55 million years from a small multi-toed creature into the large, single-toed animal of today. Humans began to domesticate horses around 4000 BCE, and their domestication is believed to have been widespread by 3000 BCE. Although, 62
• for common chimpanzees The Common Chimpanzee , also known as the Robust Chimpanzee, is a great ape. The name troglodytes, Greek for 'cave-dweller', was coined by Johann Friedrich Blumenbach in his Handbuch der Naturgeschichte (Handbook of Natural History) published in 1779. Colloquially, it is often called the chimpanzee (or simply 'chimp'), though technically this term, 59.4^[8]
• for Asian elephants The Asian or Asiatic Elephant , sometimes known by the name of one of its subspecies, the Indian Elephant, is one of the three living species of elephant, and the only living species of the genus Elephas. It is the largest living land animal in Asia. The species is found primarily in Bangladesh, India, Sri Lanka, Indochina and parts of Nepal and, 86^[9]

The longest-lived vertebrates have been variously described as

• koi Koi (English: /ˈkɔɪ/), or more specifically nishikigoi (錦鯉?, [niɕi̥kiꜜɡo.i], literally "brocaded carp"), are ornamental domesticated varieties of the common carp (Cyprinus carpio) that are kept for decorative purposes in an outdoor koi pond or water gardens. They are also called Japanese carp (A Japanese species of fish, 200+ years, though generally not exceeding 25) ^[10]
• Greenland Sharks (A species of shark native to the North Atlantic, believed to be about 200 years)
• tortoises Tortoises or land turtles are land-dwelling reptiles of the family of Testudinidae, order Testudines. Like their marine cousins, the sea turtles, tortoises are shielded from predators by a shell. The top part of the shell is the carapace, the underside is the plastron, and the two are connected by the bridge. The tortoise has both an endoskeleton (Galápagos tortoise The Galápagos tortoise or Galápagos giant tortoise is the largest living tortoise, native to seven islands of the Galápagos archipelago. Fully grown adults can weigh over 300 kilograms (661 lb) and measure 1.2 meters (4 ft) long. They are long-lived with a life expectancy in the wild estimated to be 100-150 years. Populations fell dramatically) (190 years)^[11]
• tuataras The tuatara is a reptile endemic to New Zealand which, though it resembles most lizards, is actually part of a distinct lineage, order Sphenodontia. The two species of tuatara are the only surviving members of its order, which flourished around 200 million years ago. Their most recent common ancestor with any other extant group is with the (a New Zealand reptile species, 100-200+ years^[12])
• eels Eels are an order of fish, which consists of four suborders, 19 families, 110 genera and approximately 800 species. Most eels are predators. The term "eel" is also used for some other similarly shaped fish, such as electric eels and spiny eels, but these are not members of the Anguilliformes order, the so called Brantevik eel (Swedish: Branteviksålen) is thought to have lived in a water well in southern Sweden since 1859, which makes it over 150 years old.^[13]
• whales Whale is the common name for various marine mammals of the order Cetacea. The term whale sometimes refers to all cetaceans, but more often it excludes dolphins and porpoises, which belong to suborder Odontoceti (toothed whales). This suborder also includes the sperm whale, killer whale, pilot whale, and beluga whale. The other Cetacean suborder (Bowhead Whale The bowhead whale is a baleen whale of the right whale family Balaenidae in suborder Mysticeti. A stocky dark-colored whale without a dorsal fin, it can grow to 20 meters (66 ft) in length. Estimated maximum weight of this thick-bodied species is 136 tonnes (134 LT; 150 ST), second only to the blue whale, although the bowhead's maximum length is) (about 200 years)

Although this idea was unproven for a time, recent research has indicated that bowhead whales The bowhead whale is a baleen whale of the right whale family Balaenidae in suborder Mysticeti. A stocky dark-colored whale without a dorsal fin, it can grow to 20 meters (66 ft) in length. Estimated maximum weight of this thick-bodied species is 136 tonnes (134 LT; 150 ST), second only to the blue whale, although the bowhead's maximum length is recently killed still had harpoons A harpoon is a long spear-like instrument used in fishing to catch fish or large marine mammals such as whales. It accomplishes this task by impaling the target animal, allowing the fishermen to use a rope or chain attached to the butt of the projectile to catch the animal. A harpoon can also be used as a weapon in their bodies from the 1790s, which, along with analysis of amino acids Amino acids are molecules containing an amine group, a carboxylic acid group and a side chain that varies between different amino acids. These molecules contain the key elements of carbon, hydrogen, oxygen, and nitrogen. These molecules are particularly important in biochemistry, where this term refers to alpha-amino acids with the general formula, has indicated a maximum life span, so far, of at least 211 years ^[14].

Invertebrate species which continue to grow as long as they live (e.g., certain clams, some coral species) can on occasion live hundreds of years:

• A bivalve mollusc Bivalvia is a class of marine and freshwater mollusks known for some time as Pelecypoda, but now commonly referred to simply as bivalves. As with Gastropoda and Cephalopoda, the term Pelecypoda is in reference to the animal itself while Bivalvia simply describes the shell. Other names for the class include Acephala, Bivalva, and Lamellibranchia (Arctica islandica Arctica islandica, commonly known as the ocean quahog, is an edible marine bivalve mollusk native to the North Atlantic ocean, which is harvested commercially. This species is also known by a number of different common names, including Icelandic cyprine, mahogany clam, mahogany quahog, black quahog, and black clam) (between 405–410 years^[15])

One species of jellyfish Jellyfish are free-swimming members of the phylum Cnidaria. Jellyfish have several different morphologies that represent several different cnidarian classes including the Scyphozoa (over 200 species), Staurozoa (about 50 species), Cubozoa (about 20 species), and Hydrozoa (about 1000–1500 species that make jellyfish and many more that do not), Turritopsis nutricula Turritopsis nutricula or immortal jellyfish is a hydrozoan whose medusa, or jellyfish, form can revert to the polyp stage after becoming sexually mature. It is the only known case of a metazoan capable of reverting completely to a sexually immature, colonial stage after having reached sexual maturity as a solitary stage. It does this through the, reverts to a sexually immature stage after reproducing, rather than dying as in other jellyfish. Consequently the species is considered biologically immortal Biological immortality is the absence of a sustained increase in rate of mortality as a function of chronological age. A cell or organism that does not age, or which at some point in its life will cease to age, is one which is deemed to be biologically immortal. However, this definition of immortality has been challenged in the new "Handbook and has no maximum lifespan.

However, with the possible exception of the Bowhead whale, the claims of lifespans >100 year must be taken with some skepticism as they rely on conjecture (e.g. counting otoliths) rather than empirical, continuous documentation.

In plants

Plants are referred to as annuals which live only one year, biennials which live two years, and perennials which live longer than that. The longest-lived perennials, woody-stemmed plants such as trees and bushes, often live for hundreds and even thousands of years (one may question whether or not they may die of old age). A giant sequoia, General Sherman is alive and well in its third millennium. A Great Basin Bristlecone Pine called Methuselah is 4,838 years old and the Bristlecone Pine called Prometheus was a little older still, 4,844 years, when it was cut down in 1964. The oldest known plant (probably oldest living thing) is a creosote bush (Larrea tridentata) in the Mojave Desert called King Clone at about 11,700 years.

Increasing maximum life span

Currently, the only (non-transgenic) method of increasing maximum life span that is recognized by biogerontologists is calorie restriction with adequate nutrition. "Maximum life span" here means the mean life span of the most long-lived 10% of a given cohort, as caloric restriction has not yet been shown to break mammalian world records for longevity. Rats, mice, and hamsters experience maximum life-span extension from a diet that contains 40–60% of the calories (but all of the required nutrients) that the animals consume when they can eat as much as they want. Mean life span is increased 65% and maximum life span is increased 50%, when caloric restriction is begun just before puberty.^[16]). For fruit flies the life extending benefits of calorie restriction are gained immediately at any age upon beginning calorie restriction and ended immediately at any age upon resuming full feeding^[17]).

Mammals fed antioxidants show up to a 30% increase in mean life span, but no increase in maximum life span (though even that is controversial: many studies report no increase in lifespan at all). Antioxidants are most valuable for animals that are cancer-prone or subjected to radiation or chemical toxins. There are evidently homeostatic mechanisms in cells that govern the amount of allowable antioxidant activity. Many life-extensionists have dismissed the value of antioxidants simply because they have not been shown to increase maximum life span, but such a view neglects the significance of an extended mean life span.

A few transgenic species of mice have been created that have maximum life spans greater than that of wild-type or laboratory mice. The Ames and Snell mice, which have mutations in pituitary transcription factors and hence are deficient in Gh, LH, TSH, and secondarily IGF1, have extensions in maximal lifespan of up to 65%. To date, both in absolute and relative terms, these Ames and Snell mice have the maximum lifespan of any mouse not on caloric restriction (see below on GhR). Mutations/knockout of other genes affecting the GH/IGF1 axis, such as Lit, Ghr and Irs1 have also shown extension in lifespan, but much more modest both in relative and absolute terms. The longest lived laboratory mouse ever was a Ghr knockout mouse on caloric restriction, which lived to ~1800 days (maximum for normal B6 mice under ideal conditions is 1200 days) in the lab of A. Bartke at Southern Illinois University.

Most biomedical gerontologists (gerontologists who search for ways to extend maximum life span) believe that biomedical molecular engineering will eventually extend maximum lifespan and even bring about rejuvenation.

While most aging researchers are rather cautious, fearing a vitalist public backlash, one theoretical gerontologist not shy of expressing opinions on the extension of human lifespan is Aubrey de Grey. His theoretical project to reverse the damage called aging is called SENS (Strategies for Engineered Negligible Senescence). Dr. de Grey has established The Methuselah Mouse Prize to award money to researchers who can extend the maximum life span of mice. A. Bartker collected the prize for the GhR knockout mouse and Speakman collected the prize for extending the maximum lifespan of an adult mouse, using caloric restriction initiated late in life.

Research data concerning maximum life span

• A comparison of the heart mitochondria in rats (4-year maximum life span) and pigeons (35-year maximum life span) showed that pigeon mitochondria leak fewer free-radicals than rat mitochondria, despite the fact that both animals have similar metabolic rate and cardiac output^[18]
• For mammals there is a direct relationship between mitochondrial membrane fatty acid saturation and maximum life span^[19]
• Studies of the liver lipids of mammals and a bird (pigeon) show an inverse relationship between maximum life span and number of double bonds^[20]
• Selected species of birds and mammals show an inverse relationship between telomere rate of change (shortening) and maximum life span^[21]
• Maximum life span correlates negatively with antioxidant enzyme levels and free-radicals production and positively with rate of DNA repair^[22]
• Female mammals express more Mn−SOD and glutathione peroxidase antioxidant enzymes than males. This has been hypothesized as the reason they live longer^[23] However, mice entirely lacking in Glutathione peroxidase 1 do not show a reduction in lifespan.
• The maximum life span of transgenic mice has been extended about 20% by overexpression of human catalase targeted to mitochondria^[24]
• A comparison of 7 non-primate mammals (mouse, hamster, rat, guinea-pig, rabbit, pig and cow) showed that the rate of mitochondrial superoxide and hydrogen peroxide production in heart and kidney were inversely correlated with maximum life span^[25]
• A study of 8 non-primate mammals showed an inverse correlation between maximum life span and oxidative damage to mtDNA (Mitochondrial DNA) in heart & brain^[26]
• A study of several species of mammals and a bird (pigeon) indicated a linear relationship between oxidative damage to protein and maximum life span^[27]
• There is a direct correlation between DNA repair and maximum life span for mammalian species^[28]
• Drosophila (fruit-flies) bred for 15 generations by only using eggs that were laid toward the end of reproductive life achieved maximum life spans 30% greater than that of controls^[29]
• Overexpression of the enzyme which synthesizes glutathione in long-lived transgenic Drosophila (fruit-flies) extended maximum lifespan by nearly 50%^[30]
• A mutation in the age−1 gene of the nematode worm Caenorhabditis elegans increased mean life span 65% and maximum life span 110%^[31]. However, the degree of lifespan extension in relative terms by both the age-1 and daf-2 mutations is strongly dependent on ambient temperature, with ~10% extension at 16 °C and 65% extension at 27 °C.
• Fat-specific Insulin Receptor KnockOut (FIRKO) mice have reduced fat mass, normal calorie intake and an increased maximum life span of 18%.^[32]
• The capacity of mammalian species to detoxify the carcinogenic chemical benzo(a)pyrene to a water-soluble form also correlates well with maximum life span.^[33]
• Short-term induction of oxidative stress due to calorie restriction increases life span in Caenorhabditis elegans by promoting stress defense, specifically by inducing an enzyme called catalase. As shown by Michael Ristow and co-workers nutritive antioxidants completely abolish this extension of life span by inhibiting a process called mitohormesis.^[34]

See also

• Ageing
• American Aging Association
• Aubrey de Grey
• Biodemography
• Biological immortality
• Calorie restriction
• Compression of morbidity
• DNA damage theory of aging
• Extreme longevity tracking
• Gerontology
• Hayflick limit
• Indefinite lifespan
• Life expectancy
• Life extension
• List of long-living organisms
• Longevity
• Methuselah Mouse Prize
• Michael Ristow
• Mitohormesis
• Oldest people
• Senescence
• Strategies for Engineered Negligible Senescence (SENS)

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34. ^ Publication demonstrating that oxidative stress is promoting life span

External links

• AnAge Database
• Centenarians’ Road
• Calorie Restriction Society
• Informational website on the biology of aging.
• Living Longer, Living Stronger
• Mechanisms of Aging
• Strategies for Engineered Negligible Senescence (SENS)
• The animals with the longest and shortest life spans
• The Longevity Meme (Longevity Activism)
• The Secrets of Long Life (National Geographic magazine)

Categories: Actuarial science | Aging | Demography | Gerontology

See Also:

• Maximum Life Span: Encyclopedia
• Indefinite Lifespan: Encyclopedia

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