Phenoptosis (pheno – showing or demonstrating, ptosis – programmed death, "falling off"), the term was proposed by V.P. Skulachev in 1999. It signifies the phenomenon of programmed
death
Death is the irreversible cessation of all biological functions that sustain an organism. For organisms with a brain, death can also be defined as the irreversible cessation of functioning of the whole brain, including brainstem, and brain ...
of an
organism
In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and ...
, i.e. that an organism's
genes
In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
include features that under certain circumstances will cause the organism to rapidly degenerate and die off. Recently this has been referred to as "fast phenoptosis" as aging is being explored as "slow phenoptosis". Phenoptosis is a common feature of living
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, whose ramifications for humans is still being explored. The concept of
programmed cell death
Programmed cell death (PCD; sometimes referred to as cellular suicide) is the death of a cell as a result of events inside of a cell, such as apoptosis or autophagy. PCD is carried out in a biological process, which usually confers advantage durin ...
was used before, by
Lockshin & Williams
[
] in 1964 in relation to
insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
tissue development, around eight years before "apoptosis" was coined.
Evolutionary significance
In multicellular organisms, worn-out and ineffective cells are dismantled and recycled for the greater good of the whole organism in a process called
apoptosis
Apoptosis (from grc, ἀπόπτωσις, apóptōsis, 'falling off') is a form of programmed cell death that occurs in multicellular organisms. Biochemical events lead to characteristic cell changes (morphology) and death. These changes incl ...
. It is believed that phenoptosis is an
evolutionary mechanism that culls out the damaged, aged, infectious, or those in direct competition with their own offspring
for the good of the species. Special circumstances need to exist for the "phenoptosis" strategy to be an
evolutionarily stable strategy
An evolutionarily stable strategy (ESS) is a strategy (or set of strategies) that is ''impermeable'' when adopted by a population in adaptation to a specific environment, that is to say it cannot be displaced by an alternative strategy (or set of ...
(ESS), let alone the only ESS. Examples of "phenoptosis" given below are really examples of semelpary - a life history with a single reproduction followed by death, which evolves not "for the good of the species" but as the ESS in the conditions of high adult-to-juvenile mortality ratio. The elimination of parts detrimental to the organism or individuals detrimental to the species has been deemed "The samurai law of biology" – it is better to die than to be wrong.
Stress-induced, acute, or fast phenoptosis is the rapid deterioration of an organism induced by a life event such as breeding. Elimination of the parent provides space for fitter offspring. As a species this has been advantageous particularly to species that die immediately after spawning.
Age-induced, soft, or slow phenoptosis is the slow deterioration and death of an organism due to accumulated stresses over long periods of time. In short, it has been proposed that aging, heart disease, cancer, and other age related ailments are means of phenoptosis. "Death caused by aging clears the population of ancestors and frees space for progeny carrying new useful traits."
It has also been proposed that age provides a selective advantage to brains over brawn. An example made by V. P. Skulachev provides that of two hares, one faster and one smarter, the faster hare may have a selective advantage in youth but as aging occurs and muscles deteriorate it is the smarter hare that now has the selective advantage.
Examples in nature
* ''
E. coli
''Escherichia coli'' (),Wells, J. C. (2000) Longman Pronunciation Dictionary. Harlow ngland Pearson Education Ltd. also known as ''E. coli'' (), is a Gram-negative, facultative anaerobic, rod-shaped, coliform bacterium of the genus ''Escher ...
'' – programmed death is initiated by
infection
An infection is the invasion of tissues by pathogens, their multiplication, and the reaction of host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmissible disease or communicable dise ...
by
phage
A bacteriophage (), also known informally as a ''phage'' (), is a duplodnaviria virus that infects and replicates within bacteria and archaea. The term was derived from "bacteria" and the Greek φαγεῖν ('), meaning "to devour". Bacterio ...
. This prevents further spread of phage to the remaining population.
* ''
Saccharomyces cerevisiae
''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have been o ...
'' – Under stress the yeast mitochondria produce
reactive oxygen species
In chemistry, reactive oxygen species (ROS) are highly reactive chemicals formed from diatomic oxygen (). Examples of ROS include peroxides, superoxide, hydroxyl radical, singlet oxygen, and alpha-oxygen.
The reduction of molecular oxygen () p ...
ROS, leading to loss of membrane potential within the mitochondria and death of the cell.
*
Amoeba
An amoeba (; less commonly spelled ameba or amœba; plural ''am(o)ebas'' or ''am(o)ebae'' ), often called an amoeboid, is a type of Cell (biology), cell or unicellular organism with the ability to alter its shape, primarily by extending and ret ...
''
Dictyostelium
''Dictyostelium'' is a genus of single- and multi-celled eukaryotic, phagotrophic bacterivores. Though they are Protista and in no way fungal, they traditionally are known as "slime molds". They are present in most terrestrial ecosystems as ...
'' – Under stress amoeba form multicellular
fruiting bodies
The sporocarp (also known as fruiting body, fruit body or fruitbody) of fungi is a multicellular structure on which spore-producing structures, such as basidia or asci, are borne. The fruitbody is part of the sexual phase of a fungal life cyc ...
. The better nourished cells
differentiate into spores. The less healthy cells differentiate into the stalks of the fruiting body. After maturation of the spores, the stalk cells undergo phenoptosis.
*
Nematode
The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
''
Caenorhabditis elegans
''Caenorhabditis elegans'' () is a free-living transparent nematode about 1 mm in length that lives in temperate soil environments. It is the type species of its genus. The name is a blend of the Greek ''caeno-'' (recent), ''rhabditis'' (ro ...
'' – Under normal conditions ''Caenorhabditis elegans'' display a normal aging life cycle. However, if there is increased stress after breeding they undergo phenoptosis, like in yeast, induced by the mitochondria.
*
Mayfly
Mayflies (also known as shadflies or fishflies in Canada and the upper Midwestern United States, as Canadian soldiers in the American Great Lakes region, and as up-winged flies in the United Kingdom) are aquatic insects belonging to the ord ...
– Adult mayflies have no functional mouth and die from malnutrition.
*
Mite
Mites are small arachnids (eight-legged arthropods). Mites span two large orders of arachnids, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari, but genetic analysis does not show clear evid ...
''
Adactylidium
''Adactylidium'' is a genus of mites known for its unusual life cycle. An impregnated female mite feeds upon a single egg of a thrips, rapidly growing five to eight female offspring and one male in her body. The single male mite mates with all t ...
'' – The initial food source of ''Adactylidium'' mite
larvae
A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle.
The ...
is the body tissues of their mother resulting in her death.
*
Squid
True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting t ...
– Some male squid die immediately after mating. This provides an abundant food source for those predators that would prey on the eggs.
*
Marsupial mice
The Dasyuridae are a family of marsupials native to Australia and New Guinea, including 71 extant species divided into 17 genera. Many are small and mouse-like or shrew-like, giving some of them the name marsupial mice or marsupial shrews, but th ...
– Males die two weeks after reproducing from an overabundance of their own
pheromone
A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s.
*
Salmon
Salmon () is the common name for several list of commercially important fish species, commercially important species of euryhaline ray-finned fish from the family (biology), family Salmonidae, which are native to tributary, tributaries of the ...
– Die soon after
spawning
Spawn is the eggs and sperm released or deposited into water by aquatic animals. As a verb, ''to spawn'' refers to the process of releasing the eggs and sperm, and the act of both sexes is called spawning. Most aquatic animals, except for aquati ...
.
*
Septic shock
Septic shock is a potentially fatal medical condition that occurs when sepsis, which is organ injury or damage in response to infection, leads to dangerously low blood pressure and abnormalities in cellular metabolism. The Third International Con ...
– Severe infection by
pathogen
In biology, a pathogen ( el, πάθος, "suffering", "passion" and , "producer of") in the oldest and broadest sense, is any organism or agent that can produce disease. A pathogen may also be referred to as an infectious agent, or simply a germ ...
s often results in death by
sepsis
Sepsis, formerly known as septicemia (septicaemia in British English) or blood poisoning, is a life-threatening condition that arises when the body's response to infection causes injury to its own tissues and organs. This initial stage is follo ...
. Sepsis, however, is not a result of
toxin
A toxin is a naturally occurring organic poison produced by metabolic activities of living cells or organisms. Toxins occur especially as a protein or conjugated protein. The term toxin was first used by organic chemist Ludwig Brieger (1849– ...
s activated by the pathogen, rather it is directed by the organism itself. Similar to phenoptosis of ''E. coli'', this has been suggested to be a means to separate dangerously infected individuals from healthy ones.
Proposed mechanisms
Mitochondrial ROS – The production of
ROS by the mitochondria. This causes
oxidative damage
Oxidative stress reflects an imbalance between the systemic manifestation of reactive oxygen species and a biological system's ability to readily detoxify the reactive intermediates or to repair the resulting damage. Disturbances in the normal r ...
to the
inner compartment of the
mitochondria
A mitochondrion (; ) is an organelle found in the Cell (biology), cells of most Eukaryotes, such as animals, plants and Fungus, fungi. Mitochondria have a double lipid bilayer, membrane structure and use aerobic respiration to generate adenosi ...
and destruction of the mitochondria.
Clk1 gene – the gene thought to be responsible to aging due to mitochondrial ROS.
EF2 kinase – Blocks
phosphorylation
In chemistry, phosphorylation is the attachment of a phosphate group to a molecule or an ion. This process and its inverse, dephosphorylation, are common in biology and could be driven by natural selection. Text was copied from this source, wh ...
of
elongation factor 2 thus blocking protein synthesis.
Glucocorticoid
Glucocorticoids (or, less commonly, glucocorticosteroids) are a class of corticosteroids, which are a class of steroid hormones. Glucocorticoids are corticosteroids that bind to the glucocorticoid receptor that is present in almost every vertebr ...
regulation
Regulation is the management of complex systems according to a set of rules and trends. In systems theory, these types of rules exist in various fields of biology and society, but the term has slightly different meanings according to context. For ...
– A common route for phenoptosis is breakdown of glucocorticoid regulation and inhibition, leading to massive excess of these
corticosteroids
Corticosteroids are a class of steroid hormones that are produced in the adrenal cortex of vertebrates, as well as the synthetic analogues of these hormones. Two main classes of corticosteroids, glucocorticoids and mineralocorticoids, are involve ...
in the body.
Other examples
Robert Sapolsky
Robert Morris Sapolsky (born April 6, 1957) is an American neuroendocrinology researcher and author. He is a professor of biology, and professor of neurology and neurological sciences and, by courtesy, neurosurgery, at Stanford University. In ad ...
discusses phenoptosis in his book ''
Why Zebras Don't Get Ulcers'', 3rd Ed., p. 245-247. He states that:
See also
*
Apoptosis
Apoptosis (from grc, ἀπόπτωσις, apóptōsis, 'falling off') is a form of programmed cell death that occurs in multicellular organisms. Biochemical events lead to characteristic cell changes (morphology) and death. These changes incl ...
*
Evolution of ageing
Enquiry into the evolution of ageing, or aging, aims to explain why a detrimental process such as ageing would evolve, and why there is so much variability in the lifespans of organisms. The classical theories of evolution (mutation accumulation, ...
*
Programmed cell death
Programmed cell death (PCD; sometimes referred to as cellular suicide) is the death of a cell as a result of events inside of a cell, such as apoptosis or autophagy. PCD is carried out in a biological process, which usually confers advantage durin ...
*
Sociobiology
Sociobiology is a field of biology that aims to examine and explain social behavior in terms of evolution. It draws from disciplines including psychology, ethology, anthropology, evolution, zoology, archaeology, and population genetics. Within t ...
*
Survival of the fittest
"Survival of the fittest" is a phrase that originated from Darwinian evolutionary theory as a way of describing the mechanism of natural selection. The biological concept of fitness is defined as reproductive success. In Darwinian terms, th ...
References
{{Death
Programmed cell death
Senescence