Life

For other uses, see Life (disambiguation) and Life on Earth (disambiguation).

Life (Biota / Vitae / Eobionti)
Plants in the Rwenzori Mountains, Uganda
Scientific classification
Domains and kingdoms
Life on Earth: Non-cellular life Viruses Viroids Cellular life Prokaryotes Domain Bacteria Domain Archaea Domain Eukarya Kingdom Protista Kingdom Plantae Kingdom Fungi Kingdom Animalia

This article is one of a series on:
Life in the Universe
Astrobiology
Life in the Solar System
Life on Venus
Life on Earth
Life on Mars
Life on Europa
Life on Titan
Life outside the Solar System
SETI
Exoplanetology
Planetary habitability
Circumstellar habitable zone

Life is a characteristic distinguishing physical entities having biological processes (such as signaling and self-sustaining processes) from those that do not, either because such functions have ceased (death), or because they lack such functions and are classified as inanimate. Various forms of life exist such as plants, animals, fungi, protists, archaea, and bacteria. The criteria can at times be ambiguous and may or may not define viruses, viroids or potential artificial life as living. Biology is the primary science concerned with the study of life, although many other sciences are involved.

The smallest contiguous unit of life is called an organism. Organisms are composed of one or more cells, undergo metabolism, maintain homeostasis, can grow, respond to stimuli, reproduce (either sexually or asexually) and, through evolution, adapt to their environment in successive generations. A diverse array of living organisms can be found in the biosphere of Earth, and the properties common to these organisms—plants, animals, fungi, protists, archaea, and bacteria—are a carbon- and water-based cellular form with complex organization and heritable genetic information.

Abiogenesis is the natural process of life arising from non-living matter, such as simple organic compounds. The age of the Earth is about 4.54 billion years. The earliest life on Earth arose at least 3.5 billion years ago, during the Eoarchean Era when sufficient crust had solidified following the molten Hadean Eon. The earliest physical evidence of life on Earth is biogenic graphite from 3.7 billion-year-old metasedimentary rocks found in Western Greenland and microbial mat fossils in 3.48 billion-year-old sandstone found in Western Australia. Some theories, such as the Late Heavy Bombardment theory, suggest that life on Earth may have started even earlier, and may have begun as early as 4.1 billion years ago according to a 2015 study, or even, 4.25 billion years ago, or even earlier yet, 4.4 billion years ago, according to another study. According to one of the researchers, "If life arose relatively quickly on Earth ... then it could be common in the universe."

The mechanism by which life began on Earth is unknown, although many hypotheses have been formulated. Since emerging, life has evolved into a variety of forms, which have been classified into a hierarchy of taxa. Life can survive and thrive in a wide range of conditions. Nonetheless, more than 99 percent of all species, amounting to over five billion species, that ever lived on Earth are estimated to be extinct. Estimates on the number of Earth's current species range from 10 million to 14 million, of which about 1.2 million have been documented and over 86 percent have not yet been described.

The chemistry leading to life may have begun shortly after the Big Bang, 13.8 billion years ago, during a habitable epoch when the Universe was only 10–17 million years old. Though life is confirmed only on the Earth, many think that extraterrestrial life is not only plausible, but probable or inevitable. Other planets and moons in the Solar System and other planetary systems are being examined for evidence of having once supported simple life, and projects such as SETI are trying to detect radio transmissions from possible alien civilizations.

The meaning of life—its significance, origin, purpose, and ultimate fate—is a central concept and question in philosophy and religion. Both philosophy and religion have offered interpretations as to how life relates to existence and consciousness, and on related issues such as life stance, purpose, conception of a god or gods, a soul or an afterlife. Different cultures throughout history have had widely varying approaches to these issues.

Early theories

Materialism

Plant growth in the Hoh Rainforest

Herds of zebra and impala gathering on the Maasai Mara plain

An aerial photo of microbial mats around the Grand Prismatic Spring of Yellowstone National Park

Some of the earliest theories of life were materialist, holding that all that exists is matter, and that life is merely a complex form or arrangement of matter. Empedocles (430 BC) argued that every thing in the universe is made up of a combination of four eternal "elements" or "roots of all": earth, water, air, and fire. All change is explained by the arrangement and rearrangement of these four elements. The various forms of life are caused by an appropriate mixture of elements.

Democritus (460 BC) thought that the essential characteristic of life is having a soul (psyche). Like other ancient writers, he was attempting to explain what makes something a living thing. His explanation was that fiery atoms make a soul in exactly the same way atoms and void account for any other thing. He elaborates on fire because of the apparent connection between life and heat, and because fire moves.

Plato's world of eternal and unchanging Forms, imperfectly represented in matter by a divine Artisan, contrasts sharply with the various mechanistic Weltanschauungen, of which atomism was, by the fourth century at least, the most prominent... This debate persisted throughout the ancient world. Atomistic mechanism got a shot in the arm from Epicurus... while the Stoics adopted a divine teleology... The choice seems simple: either show how a structured, regular world could arise out of undirected processes, or inject intelligence into the system.

— R. J. Hankinson, Cause and Explanation in Ancient Greek Thought

The mechanistic materialism that originated in ancient Greece was revived and revised by the French philosopher René Descartes, who held that animals and humans were assemblages of parts that together functioned as a machine. In the 19th century, the advances in cell theory in biological science encouraged this view. The evolutionary theory of Charles Darwin (1859) is a mechanistic explanation for the origin of species by means of natural selection.

Hylomorphism

Hylomorphism is a theory, originating with Aristotle (322 BC), that all things are a combination of matter and form. Biology was one of his main interests, and there is extensive biological material in his extant writings. In this view, all things in the material universe have both matter and form, and the form of a living thing is its soul (Greek psyche, Latin anima). There are three kinds of souls: the vegetative soul of plants, which causes them to grow and decay and nourish themselves, but does not cause motion and sensation; the animal soul, which causes animals to move and feel; and the rational soul, which is the source of consciousness and reasoning, which (Aristotle believed) is found only in man. Each higher soul has all the attributes of the lower one. Aristotle believed that while matter can exist without form, form cannot exist without matter, and therefore the soul cannot exist without the body.

This account is consistent with teleological explanations of life, which account for phenomena in terms of purpose or goal-directedness. Thus, the whiteness of the polar bear's coat is explained by its purpose of camouflage. The direction of causality (from the future to the past) is in contradiction with the scientific evidence for natural selection, which explains the consequence in terms of a prior cause. Biological features are explained not by looking at future optimal results, but by looking at the past evolutionary history of a species, which led to the natural selection of the features in question.

Vitalism

Vitalism is the belief that the life-principle is non-material. This originated with Stahl (17th century), and held sway until the middle of the 19th century. It appealed to philosophers such as Henri Bergson, Nietzsche, Wilhelm Dilthey, anatomists like Bichat, and chemists like Liebig. Vitalism included the idea that there was a fundamental difference between organic and inorganic material, and the belief that organic material can only be derived from living things. This was disproved in 1828, when Friedrich Wöhler prepared urea from inorganic materials. This Wöhler synthesis is considered the starting point of modern organic chemistry. It is of historical significance because for the first time an organic compound was produced in inorganic reactions.

During the 1850s, Helmholtz, anticipated by Mayer, demonstrated that no energy is lost in muscle movement, suggesting that there were no "vital forces" necessary to move a muscle. These results led to the abandonment of scientific interest in vitalistic theories, although the belief lingered on in pseudoscientific theories such as homeopathy, which interprets diseases and sickness as caused by disturbances in a hypothetical vital force or life force.

Definitions

It is a challenge for scientists and philosophers to define life. This is partially because life is a process, not a substance. Any definition must be general enough to both encompass all known life and any unknown life that may be different from life on Earth. Some may even consider that life is not real at all, but a concept instead.

Biology

Since there is no unequivocal definition of life, the current understanding is descriptive. Life is considered a characteristic of something that exhibits all or most of the following traits:

1. Homeostasis: Regulation of the internal environment to maintain a constant state; for example, sweating to reduce temperature.
2. Organization: Being structurally composed of one or more cells — the basic units of life.
3. Metabolism: Transformation of energy by converting chemicals and energy into cellular components (anabolism) and decomposing organic matter (catabolism). Living things require energy to maintain internal organization (homeostasis) and to produce the other phenomena associated with life.
4. Growth: Maintenance of a higher rate of anabolism than catabolism. A growing organism increases in size in all of its parts, rather than simply accumulating matter.
5. Adaptation: The ability to change over time in response to the environment. This ability is fundamental to the process of evolution and is determined by the organism's heredity, diet, and external factors.
6. Response to stimuli: A response can take many forms, from the contraction of a unicellular organism to external chemicals, to complex reactions involving all the senses of multicellular organisms. A response is often expressed by motion; for example, the leaves of a plant turning toward the sun (phototropism), and chemotaxis.
7. Reproduction: The ability to produce new individual organisms, either asexually from a single parent organism, or sexually from two parent organisms, "with an error rate below the sustainability threshold."

These complex processes, called physiological functions, have underlying physical and chemical bases, as well as signaling and control mechanisms that are essential to maintaining life.

Alternatives

See also: Entropy and life

To reflect the minimum phenomena required, other biological definitions of life have been proposed, many of these are based upon chemical systems. Biophysicists have commented that living things function on negative entropy. In other words, living processes can be viewed as a delay of the spontaneous diffusion or dispersion of the internal energy of biological molecules towards more potential microstates. In more detail, according to physicists such as John Bernal, Erwin Schrödinger, Eugene Wigner, and John Avery, life is a member of the class of phenomena that are open or continuous systems able to decrease their internal entropy at the expense of substances or free energy taken in from the environment and subsequently rejected in a degraded form. At a higher level, living beings are thermodynamic systems that have an organized molecular structure. That is, life is matter that can reproduce itself and evolve as survival dictates. Hence, life is a self-sustained chemical system capable of undergoing Darwinian evolution.

Others take a systemic viewpoint that does not necessarily depend on molecular chemistry. One systemic definition of life is that living things are self-organizing and autopoietic (self-producing). Variations of this definition include Stuart Kauffman's definition as an autonomous agent or a multi-agent system capable of reproducing itself or themselves, and of completing at least one thermodynamic work cycle.

Viruses

Electron micrograph of adenovirus with a cartoon to demonstrate its icosahedral structure

Viruses are most often considered replicators rather than forms of life. They have been described as "organisms at the edge of life," since they possess genes, evolve by natural selection, and replicate by creating multiple copies of themselves through self-assembly. However, viruses do not metabolize and they require a host cell to make new products. Virus self-assembly within host cells has implications for the study of the origin of life, as it may support the hypothesis that life could have started as self-assembling organic molecules.

Living systems theories

The idea that the Earth is alive is found in philosophy and religion, but the first scientific discussion of it was by the Scottish scientist James Hutton. In 1785, he stated that the Earth was a superorganism and that its proper study should be physiology. Hutton is considered the father of geology, but his idea of a living Earth was forgotten in the intense reductionism of the 19th century. The Gaia hypothesis, proposed in the 1960s by scientist James Lovelock, suggests that life on Earth functions as a single organism that defines and maintains environmental conditions necessary for its survival.

The first attempt at a general living systems theory for explaining the nature of life was in 1978, by American biologist James Grier Miller. Such a general theory, arising out of the ecological and biological sciences, attempts to map general principles for how all living systems work. Instead of examining phenomena by attempting to break things down into component parts, a general living systems theory explores phenomena in terms of dynamic patterns of the relationships of organisms with their environment. Robert Rosen (1991) built on this by defining a system component as "a unit of organization; a part with a function, i.e., a definite relation between part and whole." From this and other starting concepts, he developed a "relational theory of systems" that attempts to explain the special properties of life. Specifically, he identified the "nonfractionability of components in an organism" as the fundamental difference between living systems and "biological machines."

A systems view of life treats environmental fluxes and biological fluxes together as a "reciprocity of influence", and a reciprocal relation with environment is arguably as important for understanding life as it is for understanding ecosystems. As Harold J. Morowitz (1992) explains it, life is a property of an ecological system rather than a single organism or species. He argues that an ecosystemic definition of life is preferable to a strictly biochemical or physical one. Robert Ulanowicz (2009) highlights mutualism as the key to understand the systemic, order-generating behavior of life and ecosystems.

Complex systems biology (CSB) is a field of science that studies the emergence of complexity in functional organisms from the viewpoint of dynamic systems theory. The latter is often called also systems biology and aims to understand the most fundamental aspects of life. A closely related approach to CSB and systems biology, called relational biology, is concerned mainly with understanding life processes in terms of the most important relations, and categories of such relations among the essential functional components of organisms; for multicellular organisms, this has been defined as "categorical biology", or a model representation of organisms as a category theory of biological relations, and also an algebraic topology of the functional organization of living organisms in terms of their dynamic, complex networks of metabolic, genetic, epigenetic processes and signaling pathways.

It has also been argued that the evolution of order in living systems and certain physical systems obey a common fundamental principle termed the Darwinian dynamic. The Darwinian dynamic was formulated by first considering how macroscopic order is generated in a simple non-biological system far from thermodynamic equilibrium, and then extending consideration to short, replicating RNA molecules. The underlying order generating process for both types of system was concluded to be basically similar.

Another systemic definition, called the Operator theory, proposes that 'life is a general term for the presence of the typical closures found in organisms; the typical closures are a membrane and an autocatalytic set in the cell', and also proposes that an organism is 'any system with an organisation that complies with an operator type that is at least as complex as the cell. Life can also be modeled as a network of inferior negative feedbacks of regulatory mechanisms subordinated to a superior positive feedback formed by the potential of expansion and reproduction.

Origin

Main article: Abiogenesis

Evidence suggests that life on Earth has existed for at least 3.5 billion years, with the oldest physical traces of life dating back 3.7 billion years. All known life forms share fundamental molecular mechanisms, reflecting their common descent; based on these observations, hypotheses on the origin of life attempt to find a mechanism explaining the formation of a universal common ancestor, from simple organic molecules via pre-cellular life to protocells and metabolism. Models have been divided into "genes-first" and "metabolism-first" categories, but a recent trend is the emergence of hybrid models that combine both categories.

There is no current scientific consensus as to how life originated. However, most accepted scientific models build on the following observations:

• The Miller–Urey experiment, and the work of Sidney Fox, show that conditions on the primitive Earth favored chemical reactions that synthesize amino acids and other organic compounds from inorganic precursors.
• Phospholipids spontaneously form lipid bilayers, the basic structure of a cell membrane.

Living organisms synthesize proteins, which are polymers of amino acids using instructions encoded by deoxyribonucleic acid (DNA). Protein synthesis entails intermediary ribonucleic acid (RNA) polymers. One possibility for how life began is that genes originated first, followed by proteins; the alternative being that proteins came first and then genes.

However, since genes and proteins are both required to produce the other, the problem of considering which came first is like that of the chicken or the egg. Most scientists have adopted the hypothesis that because of this, it is unlikely that genes and proteins arose independently.

Therefore, a possibility, first suggested by Francis Crick, is that the first life was based on RNA, which has the DNA-like properties of information storage and the catalytic properties of some proteins. This is called the RNA world hypothesis, and it is supported by the observation that many of the most critical components of cells (those that evolve the slowest) are composed mostly or entirely of RNA. Also, many critical cofactors (ATP, Acetyl-CoA, NADH, etc.) are either nucleotides or substances clearly related to them. The catalytic properties of RNA had not yet been demonstrated when the hypothesis was first proposed, but they were confirmed by Thomas Cech in 1986.

One issue with the RNA world hypothesis is that synthesis of RNA from simple inorganic precursors is more difficult than for other organic molecules. One reason for this is that RNA precursors are very stable and react with each other very slowly under ambient conditions, and it has also been proposed that living organisms consisted of other molecules before RNA. However, the successful synthesis of certain RNA molecules under the conditions that existed prior to life on Earth has been achieved by adding alternative precursors in a specified order with the precursor phosphate present throughout the reaction. This study makes the RNA world hypothesis more plausible.

Geological findings in 2013 showed that reactive phosphorus species (like phosphite) were in abundance in the ocean before 3.5 Ga, and that Schreibersite easily reacts with aqueous glycerol to generate phosphite and glycerol 3-phosphate. It is hypothesized that Schreibersite-containing meteorites from the Late Heavy Bombardment could have provided early reduced phosphorus, which could react with prebiotic organic molecules to form phosphorylated biomolecules, like RNA.

In 2009, experiments demonstrated Darwinian evolution of a two-component system of RNA enzymes (ribozymes) in vitro. The work was performed in the laboratory of Gerald Joyce, who stated, "This is the first example, outside of biology, of evolutionary adaptation in a molecular genetic system."

Prebiotic compounds may have extraterrestrial origin. NASA findings in 2011, based on studies with meteorites found on Earth, suggest DNA and RNA components (adenine, guanine and related organic molecules) may be formed in outer space.

In March 2015, NASA scientists reported that, for the first time, complex DNA and RNA organic compounds of life, including uracil, cytosine and thymine, have been formed in the laboratory under outer space conditions, using starting chemicals, such as pyrimidine, found in meteorites. Pyrimidine, like polycyclic aromatic hydrocarbons (PAHs), the most carbon-rich chemical found in the Universe, may have been formed in red giants or in interstellar dust and gas clouds, according to the scientists.

According to the panspermia hypothesis, microscopic life—distributed by meteoroids, asteroids and other small Solar System bodies—may exist throughout the universe.

Environmental conditions

Cyanobacteria dramatically changed the composition of life forms on Earth by leading to the near-extinction of oxygen-intolerant organisms.

The diversity of life on Earth is a result of the dynamic interplay between genetic opportunity, metabolic capability, environmental challenges, and symbiosis. For most of its existence, Earth's habitable environment has been dominated by microorganisms and subjected to their metabolism and evolution. As a consequence of these microbial activities, the physical-chemical environment on Earth has been changing on a geologic time scale, thereby affecting the path of evolution of subsequent life. For example, the release of molecular oxygen by cyanobacteria as a by-product of photosynthesis induced global changes in the Earth's environment. Since oxygen was toxic to most life on Earth at the time, this posed novel evolutionary challenges, and ultimately resulted in the formation of Earth's major animal and plant species. This interplay between organisms and their environment is an inherent feature of living systems.

All life forms require certain core chemical elements needed for biochemical functioning. These include carbon, hydrogen, nitrogen, oxygen, phosphorus, and sulfur—the elemental macronutrients for all organisms—often represented by the acronym CHNOPS. Together these make up nucleic acids, proteins and lipids, the bulk of living matter. Five of these six elements comprise the chemical components of DNA, the exception being sulfur. The latter is a component of the amino acids cysteine and methionine. The most biologically abundant of these elements is carbon, which has the desirable attribute of forming multiple, stable covalent bonds. This allows carbon-based (organic) molecules to form an immense variety of chemical arrangements. Alternative hypothetical types of biochemistry have been proposed that eliminate one or more of these elements, swap out an element for one not on the list, or change required chiralities or other chemical properties.

Range of tolerance

The inert components of an ecosystem are the physical and chemical factors necessary for life — energy (sunlight or chemical energy), water, temperature, atmosphere, gravity, nutrients, and ultraviolet solar radiation protection. In most ecosystems, the conditions vary during the day and from one season to the next. To live in most ecosystems, then, organisms must be able to survive a range of conditions, called the "range of tolerance." Outside that are the "zones of physiological stress", where the survival and reproduction are possible but not optimal. Beyond these zones are the "zones of intolerance", where survival and reproduction of that organism is unlikely or impossible. Organisms that have a wide range of tolerance are more widely distributed than organisms with a narrow range of tolerance.

Deinococcus radiodurans is an extremophile that can resist extremes of cold, dehydration, vacuum, acid, and radiation exposure.

To survive, selected microorganisms can assume forms that enable them to withstand freezing, complete desiccation, starvation, high levels of radiation exposure, and other physical or chemical challenges. These microorganisms may survive exposure to such conditions for weeks, months, years, or even centuries. Extremophiles are microbial life forms that thrive outside the ranges where life is commonly found. They excel at exploiting uncommon sources of energy. While all organisms are composed of nearly identical molecules, evolution has enabled such microbes to cope with this wide range of physical and chemical conditions. Characterization of the structure and metabolic diversity of microbial communities in such extreme environments is ongoing.

Microbial life forms thrive even in the Mariana Trench, the deepest spot on the Earth. Microbes also thrive inside rocks up to 1900 feet below the sea floor under 8500 feet of ocean.

Investigation of the tenacity and versatility of life on Earth, as well as an understanding of the molecular systems that some organisms utilize to survive such extremes, is important for the search for life beyond Earth. For example, lichen could survive for a month in a simulated Martian environment.

Form and function

Cells are the basic unit of structure in every living thing, and all cells arise from pre-existing cells by division. Cell theory was formulated by Henri Dutrochet, Theodor Schwann, Rudolf Virchow and others during the early nineteenth century, and subsequently became widely accepted. The activity of an organism depends on the total activity of its cells, with energy flow occurring within and between them. Cells contain hereditary information that is carried forward as a genetic code during cell division.

There are two primary types of cells. Prokaryotes lack a nucleus and other membrane-bound organelles, although they have circular DNA and ribosomes. Bacteria and Archaea are two domains of prokaryotes. The other primary type of cells are the eukaryotes, which have distinct nuclei bound by a nuclear membrane and membrane-bound organelles, including mitochondria, chloroplasts, lysosomes, rough and smooth endoplasmic reticulum, and vacuoles. In addition, they possess organized chromosomes that store genetic material. All species of large complex organisms are eukaryotes, including animals, plants and fungi, though most species of eukaryote are protist microorganisms. The conventional model is that eukaryotes evolved from prokaryotes, with the main organelles of the eukaryotes forming through endosymbiosis between bacteria and the progenitor eukaryotic cell.

The molecular mechanisms of cell biology are based on proteins. Most of these are synthesized by the ribosomes through an enzyme-catalyzed process called protein biosynthesis. A sequence of amino acids is assembled and joined together based upon gene expression of the cell's nucleic acid. In eukaryotic cells, these proteins may then be transported and processed through the Golgi apparatus in preparation for dispatch to their destination.

Cells reproduce through a process of cell division in which the parent cell divides into two or more daughter cells. For prokaryotes, cell division occurs through a process of fission in which the DNA is replicated, then the two copies are attached to parts of the cell membrane. In eukaryotes, a more complex process of mitosis is followed. However, the end result is the same; the resulting cell copies are identical to each other and to the original cell (except for mutations), and both are capable of further division following an interphase period.

Multicellular organisms may have first evolved through the formation of colonies of like cells. These cells can form group organisms through cell adhesion. The individual members of a colony are capable of surviving on their own, whereas the members of a true multi-cellular organism have developed specializations, making them dependent on the remainder of the organism for survival. Such organisms are formed clonally or from a single germ cell that is capable of forming the various specialized cells that form the adult organism. This specialization allows multicellular organisms to exploit resources more efficiently than single cells.

Cells have evolved methods to perceive and respond to their microenvironment, thereby enhancing their adaptability. Cell signaling coordinates cellular activities, and hence governs the basic functions of multicellular organisms. Signaling between cells can occur through direct cell contact using juxtacrine signalling, or indirectly through the exchange of agents as in the endocrine system. In more complex organisms, coordination of activities can occur through a dedicated nervous system.

Classification

Main article: Biological classification

The hierarchy of biological classification's eight major taxonomic ranks. Life is divided into domains, which are subdivided into further groups. Intermediate minor rankings are not shown.

The first known attempt to classify organisms was conducted by the Greek philosopher Aristotle (384–322 BC), who classified all living organisms known at that time as either a plant or an animal, based mainly on their ability to move. He also distinguished animals with blood from animals without blood (or at least without red blood), which can be compared with the concepts of vertebrates and invertebrates respectively, and divided the blooded animals into five groups: viviparous quadrupeds (mammals), oviparous quadrupeds (reptiles and amphibians), birds, fishes and whales. The bloodless animals were also divided into five groups: cephalopods, crustaceans, insects (which included the spiders, scorpions, and centipedes, in addition to what we define as insects today), shelled animals (such as most molluscs and echinoderms) and "zoophytes." Though Aristotle's work in zoology was not without errors, it was the grandest biological synthesis of the time and remained the ultimate authority for many centuries after his death.

The exploration of the American continent revealed large numbers of new plants and animals that needed descriptions and classification. In the latter part of the 16th century and the beginning of the 17th, careful study of animals commenced and was gradually extended until it formed a sufficient body of knowledge to serve as an anatomical basis for classification. In the late 1740s, Carl Linnaeus introduced his system of binomial nomenclature for the classification of species. Linnaeus attempted to improve the composition and reduce the length of the previously used many-worded names by abolishing unnecessary rhetoric, introducing new descriptive terms and precisely defining their meaning. By consistently using this system, Linnaeus separated nomenclature from taxonomy.

The fungi were originally treated as plants. For a short period Linnaeus had classified them in the taxon Vermes in Animalia, but later placed them back in Plantae. Copeland classified the Fungi in his Protoctista, thus partially avoiding the problem but acknowledging their special status. The problem was eventually solved by Whittaker, when he gave them their own kingdom in his five-kingdom system. Evolutionary history shows that the fungi are more closely related to animals than to plants.

As new discoveries enabled detailed study of cells and microorganisms, new groups of life were revealed, and the fields of cell biology and microbiology were created. These new organisms were originally described separately in protozoa as animals and protophyta/thallophyta as plants, but were united by Haeckel in the kingdom Protista; later, the prokaryotes were split off in the kingdom Monera, which would eventually be divided into two separate groups, the Bacteria and the Archaea. This led to the six-kingdom system and eventually to the current three-domain system, which is based on evolutionary relationships. However, the classification of eukaryotes, especially of protists, is still controversial.

As microbiology, molecular biology and virology developed, non-cellular reproducing agents were discovered, such as viruses and viroids. Whether these are considered alive has been a matter of debate; viruses lack characteristics of life such as cell membranes, metabolism and the ability to grow or respond to their environments. Viruses can still be classed into "species" based on their biology and genetics, but many aspects of such a classification remain controversial.

In the 1960s a trend called cladistics emerged, arranging taxa based on clades in an evolutionary or phylogenetic tree.

Linnaeus 1735	Haeckel 1866	Chatton 1925	Copeland 1938	Whittaker 1969	Woese et al. 1990	Cavalier-Smith 1998
2 kingdoms	3 kingdoms	2 empires	4 kingdoms	5 kingdoms	3 domains	6 kingdoms
(not treated)	Protista	Prokaryota	Monera	Monera	Bacteria	Bacteria
					Archaea
		Eukaryota	Protoctista	Protista	Eucarya	Protozoa
						Chromista
Vegetabilia	Plantae		Plantae	Plantae		Plantae
				Fungi		Fungi
Animalia	Animalia		Animalia	Animalia		Animalia

Main article: Kingdom (biology) § Summary

Extraterrestrial life

Main articles: Extraterrestrial life, Astrobiology, Astroecology and Life in the solar system

Earth is the only planet known to harbor life. Other locations within the Solar System that may host microbial life include subsurface Mars, the atmosphere of Venus, and subsurface oceans on some of the moons of the gas giant planets. The variables of the Drake equation are used to discuss the conditions in solar systems where civilization is most likely to exist.

Beyond the solar system, the region around another main sequence star that could support Earth-like life on an Earth-like planet is known as the habitable zone. The inner and outer radii of this zone vary with the luminosity of the star, as does the time interval during which the zone survives. Stars more massive than the Sun have a larger habitable zone, but remain on the main sequence for a shorter time interval. Small red dwarf stars have the opposite problem, with a smaller habitable zone that is subject to higher levels of magnetic activity and the effects of tidal locking from close orbits. Hence, stars in the intermediate mass range such as the Sun may have a greater likelihood for Earth-like life to develop. The location of the star within a galaxy may also have an impact on the likelihood of life forming. Stars in regions with a greater abundance of heavier elements that can form planets, in combination with a low rate of potentially habitat-damaging supernova events, are predicted to have a higher probability of hosting planets with complex life.

Death

Main article: Death

Animal corpses, like this African buffalo, are recycled by the ecosystem, providing energy and nutrients for living creatures

Death is the permanent termination of all vital functions or life processes in an organism or cell. It can occur as a result of an accident, medical conditions, biological interaction, malnutrition, poisoning, senescence, or suicide. After death, the remains of an organism re-enter the biogeochemical cycle. Organisms may be consumed by a predator or a scavenger and leftover organic material may then be further decomposed by detritivores, organisms that recycle detritus, returning it to the environment for reuse in the food chain.

One of the challenges in defining death is in distinguishing it from life. Death would seem to refer to either the moment life ends, or when the state that follows life begins. However, determining when death has occurred requires drawing precise conceptual boundaries between life and death. This is problematic, however, because there is little consensus over how to define life. The nature of death has for millennia been a central concern of the world's religious traditions and of philosophical inquiry. Many religions maintain faith in either a kind of afterlife or reincarnation for the soul, or resurrection of the body at a later date.

Extinction is the process by which a group of taxa or species dies out, reducing biodiversity. The moment of extinction is generally considered the death of the last individual of that species. Because a species' potential range may be very large, determining this moment is difficult, and is usually done retrospectively after a period of apparent absence. Species become extinct when they are no longer able to survive in changing habitat or against superior competition. In Earth's history, over 99% of all the species that have ever lived have gone extinct; however, mass extinctions may have accelerated evolution by providing opportunities for new groups of organisms to diversify.

Fossils are the preserved remains or traces of animals, plants, and other organisms from the remote past. The totality of fossils, both discovered and undiscovered, and their placement in fossil-containing rock formations and sedimentary layers (strata) is known as the fossil record. A preserved specimen is called a fossil if it is older than the arbitrary date of 10,000 years ago. Hence, fossils range in age from the youngest at the start of the Holocene Epoch to the oldest from the Archaean Eon, up to 3.4 billion years old.

Artificial life

Main articles: Artificial life and Synthetic biology

Artificial life is a field of study that examines systems related to life, its processes, and its evolution through simulations using computer models, robotics, and biochemistry. The study of artificial life imitates traditional biology by recreating some aspects of biological phenomena. Scientists study the logic of living systems by creating artificial environments—seeking to understand the complex information processing that defines such systems. While life is, by definition, alive, artificial life is generally referred to as data confined to a digital environment and existence.

Synthetic biology is a new area of biotechnology that combines science and biological engineering. The common goal is the design and construction of new biological functions and systems not found in nature. Synthetic biology includes the broad redefinition and expansion of biotechnology, with the ultimate goals of being able to design and build engineered biological systems that process information, manipulate chemicals, fabricate materials and structures, produce energy, provide food, and maintain and enhance human health and the environment.

Phanerozoic Eon

Main article: Phanerozoic

The Phanerozoic Eon is the current eon in Earth's history. The Phanerozoic began 540 million years ago and continues to the present. The Phanerozoic comprises the Paleozoic Era, the Mesozoic Era, and the Cenozoic Era. This eon is when the diversity of life increases dramatically, starting with the Cambrian Explosion.

Paleozoic Era

Main article: Paleozoic

The Paleozoic is a time in earth's history when complex life forms evolve, take their first breath of oxygen on dry land, and when the forerunner of all life on earth begin to diversify. There are six periods in the Paleozoic eras: the Cambrian, the Ordovician, the Silurian, the Devonian, the Carboniferous and the Permian.

Cambrian

Trilobites

The Cambrian spans from 540 million years to 485 million years ago and is the first period of the Paleozoic and of the Phanerozoic Eon. The Cambrian sparks a boom in evolution in an event known as the Cambrian Explosion in which the largest number of creatures evolve in the history of Earth during one period. Creatures like algae evolve, but most of the water is populated by armored arthropods, like trilobites. Almost all marine phyla evolved in this period. During this time, the super-continent Rodinia begins to break up, most of which becomes the super-continent Gondwana.

Ordovician

Cephalaspis (a jaw-less fish)

The Ordovician spans from 485 million years to 440 million years ago. The Ordovician is a time in earths history in which many species still prevalent today evolved, such as primitive fish, cephalopods, and coral. The most common forms of life, however, were trilobites, snails and shellfish. More importantly, the first arthropods went ashore to colonize the empty continent of Gondwana. By the end of the period, Gondwana was at the south pole, early North America had collided with Europe, closing the Atlantic Ocean. Glaciation of Africa resulted in a major drop in sea level, killing off all life that staked a claim along coastal Gondwana. Glaciation caused a snowball earth, and the Ordovician-Silurian extinction in which 60% of marine invertebrates and 25% of families went extinct, and is considered the first mass extinction and the second deadliest extinction.

Silurian

The Silurian spans from 440 million years to 415 million years ago. The Silurian saw the healing of the earth that recovered from the snowball earth. This period saw the mass evolution of fish, as jaw-less fish became more numerous, jawed fish evolved, and the first freshwater fish evolved, though arthropods, such as sea scorpions, were still apex predators. Fully terrestrial life evolved, which included early arachnids, fungi, and centipedes. Also, the evolution of vascular plants (Cooksonia) allowed plants to gain a foothold on land. These early plants are the forerunners of all plant life on land. During this time, there are four continents: Gondwana (Africa, South America, Australia, Antarctica, Siberia), Laurentia (North America), Baltica (Northern Europe), and Avalonia (Western Europe). The recent rise in sea levels provided many new species to thrive in water.

Devonian

Eogyrinus (an amphibian) of the Carboniferous

The Devonian spans from 415 million years to 360 million years ago. Also known as "The Age of the Fish", the Devonian features a huge diversification of fish, including armored fish like Dunkleosteus and lobe-finned fish which eventually evolved into the first tetrapods. On land, plant groups diversified incredibly in an event known as the Devonian Explosion where the first trees evolved, as well as seeds. This event also diversified arthropod life. The first amphibians also evolved, and the fish were now at the top of the food chain. Near the end of the Devonian, 70% of all species went extinct in an event known as the Late Devonian extinction and is the second mass extinction event the world has seen.

Carboniferous

Dimetrodon

The Carboniferous spans from 360 million to 300 million years ago. During this time, average global temperatures were exceedingly high; the early Carboniferous averaged at about 20 degrees Celsius (but cooled down to 10 degrees during the Middle Carboniferous). Tropical swamps dominated the earth, and the large amounts of trees created much of the carbon for the coal that is used today (hence the name "Carbon-iferous"). Perhaps the most important evolutionary development of the time was the evolution of amniotic eggs, which allowed amphibians to head farther inland and remained the dominant vertebrates throughout the duration of this period. Also, the first reptiles and synapsids evolved in the swamps. Throughout the Carboniferous, there was a cooling pattern, which eventually led to the glaciation of Gondwana as much of it was situated around the south pole in an event known as the Permo-Carboniferous glaciation or the Carboniferous Rainforest Collapse.

Permian

The Permian spans from 300 million to 250 million years ago and was the last period of the Paleozoic. At the beginning, all continents formed together to form the super-continent Pangaea and had one ocean called Panthalassa. The earth was very dry during this time, with harsh seasons as the climate of the interior of Pangaea wasn't regulated by large bodies of water. Reptiles and synapsids flourished in the new dry climate. Creatures such as Dimetrodon and Edaphosaurus ruled the new continent. The first conifers evolve, and dominate the terrestrial landscape. Nearing the end of the Permian, however, Pangaea got drier and drier. The interior was nothing but dry deserts, and new species such as Scutosaurus and Gorgonopsid filled the empty desert. Eventually, they disappeared, along with 95% of all life on earth in an event simply known as "the Great Dying", and is the third mass extinction event of the world.

Mesozoic Era

Main article: Mesozoic

Also known as "the Age of the dinosaurs", the Mesozoic features the rise of reptiles on their 150 million year conquest to rule the earth from the seas, the land, and even in the air. There are 3 periods in the Mesozoic: the Triassic, the Jurassic, and the Cretaceous.

Triassic

The Triassic ranges from 250 million to 200 million years ago. The Triassic is a desolate transitional state in Earth's history between the Permian Extinction and the lush Jurassic Period. It has three major epochs: the Early Triassic, the Middle Triassic and the Late Triassic.

The Early Triassic lived between 250 million to 247 million years ago and was dominated by deserts as Pangaea had not yet broken up, thus the interior was nothing but arid. The Earth had just witnessed a massive die-off in which 95% of all life went extinct. The most common life on earth were Lystrosaurus, Labyrinthodont, and Euparkeria along with many other creatures that managed to survive the Great Dying. Temnospondyli evolved during this time and would be the dominant predator for much of the Triassic.

Plateosaurus (a prosauropod)

The Middle Triassic spans from 247 million to 237 million years ago. The Middle Triassic featured the beginnings of the breakup of Pangaea, and the beginning of the Tethys Sea. The ecosystem had recovered from the devastation that was the Great Dying. Phytoplankton, coral, and crustaceans all had recovered, and the reptiles began to get bigger and bigger. New aquatic reptiles evolved such as Ichthyosaurs and Nothosaurs. Meanwhile, on land, Pine forests flourished, bringing along mosquitoes and fruit flies. The first ancient crocodilians evolved, which sparked competition with the large amphibians that had since rule the freshwater world.

The Late Triassic spans from 237 million to 200 million years ago. Following the bloom of the Middle Triassic, the Late Triassic featured frequent heat spells, as well as moderate precipitation (10-20 inches per year). The recent warming led to a boom of reptilian evolution on land as the first true dinosaurs evolve, as well as pterosaurs. All this climactic change, however, resulted in a large die-out known as the Triassic-Jurassic extinction event, in which all archosaurs (excluding ancient crocodiles), synapsids, and almost all large amphibians went extinct, as well as 34% of marine life in the fourth mass extinction event of the world. The cause is debatable.

Jurassic

Rhamphorhynchus

The Jurassic ranges from 200 million years to 145 million years ago and features 3 major epochs: The Early Jurassic, the Middle Jurassic, and the Late Jurassic.

The Early Jurassic spans from 200 million years to 175 million years ago. The climate was much more humid than the Triassic, and as a result, the world was very tropical. In the oceans, Plesiosaurs, Ichthyosaurs and Ammonites fill waters as the dominant races of the seas. On land, dinosaurs and other reptiles stake their claim as the dominant race of the land, with species such as Dilophosaurus at the top. The first true crocodiles evolved, pushing out the large amphibians to near extinction. All-in-all, reptiles rise to rule the world. Meanwhile, the first true mammals evolve, but never exceed the height of a shrew.

The Middle Jurassic spans from 175 million to 163 million years ago. During this epoch, reptiles flourished as huge herds of sauropods, such as Brachiosaurus and Diplodocus, filled the fern prairies of the Middle Jurassic. Many other predators rose as well, such as Allosaurus. Conifer forests made up a large portion of the forests. In the oceans, Plesiosaurs were quite common, and Ichthyosaurs were flourishing. This epoch was the peak of the reptiles.

(Inaccurately portrayed) Stegosaurus

The Late Jurassic spans from 163 million to 145 million years ago. The Late Jurassic featured a massive extinction of sauropods and Ichthyosaurs due to the separation of Pangaea into Laurasia and Gondwana in an extinction known as the Jurassic-Cretaceous extinction. Sea levels rose, destroying fern prairies and creating shallows in its wake. Ichthyosaurs went extinct whereas sauropods, as a whole, did not die out in the Jurassic; in fact, some species, like the Titanosaurus, lived up to the K-T extinction. The increase in sea-levels opened up the Atlantic sea way which would continue to get larger over time. The divided world would give opportunity for the diversification of new dinosaurs.

Cretaceous

The Cretaceous is the longest era in the Mesozoic, but has only two periods: the Early Cretaceous, and the Late Cretaceous.

Tylosaurus (a mosasaur) hunting Xiphactinus

The Early Cretaceous spans from 145 million to 100 million years ago. The Early Cretaceous saw the expansion of seaways, and as a result, the decline and extinction of sauropods (except in South America). Many coastal shallows were created, and that caused Ichthyosaurs to die out. Mosasaurs evolved to replace them as head of the seas. Some island-hopping dinosaurs, like Eustreptospondylus, evolved to cope with the coastal shallows and small islands of ancient Europe. Other dinosaurs rose up to fill the empty space that the Jurassic-Cretaceous extinction left behind, such as Carcharodontosaurus and Spinosaurus. Of the most successful would be the Iguanodon which spread to every continent. Seasons came back into effect an the poles got seasonally colder, but dinosaurs still inhabited this area like the Leaellynasaura which inhabited the polar forests year-round, and many dinosaurs migrated there during summer like Muttaburrasaurus. Since it was too cold for crocodiles, it was the last stronghold for large amphibians, like Koolasuchus. Pterosaurs got larger as species like Tapejara and Ornithocheirus evolved. More importantly, the first true birds evolved which sparked competition between them and the pterosaurs.

The Late Cretaceous spans from 100 million to 65 million years ago. The Late Cretaceous featured a cooling trend that would continue on in the Cenozoic period. Eventually, tropics were restricted to the equator and areas beyond the tropic lines featured extreme seasonal changes in weather. Dinosaurs still thrived as new species such as Tyrannosaurus, Ankylosaurus, Triceratops and Hadrosaurs dominated the food web. Pterosaurs, however, were going into a decline as birds took to the skies. The last pterosaur to die off was Quetzalcoatlus. Marsupials evolved within the large conifer forests as scavengers. In the oceans, Mosasaurs ruled the seas to fill the role of the Ichthyosaurs, and huge plesiosaurs, such as Elasmosaurus, evolved. Also, the first flowering plants evolved. At the end of the Cretaceous, the Deccan traps and other volcanic eruptions were poisoning the atmosphere. As this was continuing, it is thought that a large meteor smashed into earth, creating the Chicxulub Crater in an event known as the K-T Extinction, the fifth and most recent mass extinction event, in which 75% of life on earth went extinct, including all non-avian dinosaurs. Everything over 10 kilograms went extinct. The age of the dinosaurs was officially over.

Cenozoic Era

Main article: Cenozoic

The Cenozoic features the rise of mammals on their conquest to rule the land, as the dinosaurs have now left a huge opening as top dog. There are three division of the Cenozoic: the Paleogene, the Neogene and Quaternary.

Paleogene

The Paleogene spans from the extinction of the dinosaurs, some 65 million years ago, to the dawn of the Neogene twenty three million years ago. It features three epochs: the Paleocene, Eocene and Oligocene.

Basilosaurus

The Paleocene ranged from 65 million to 55 million years ago. The Paleocene is a transitional point between the devastation that is the K-T extinction, to the rich jungles environment that is the Early Eocene. The Early Paleocene saw the recovery of the earth. The continents began to take their modern shape, but all continents (and India) were separated from each other. Afro-Eurasia is separated by the Tethys Sea, and the Americas are separated by the strait of Panama, as the isthmus has not yet formed. This epoch features a general warming trend, with jungles eventually reaching the poles. The oceans were dominated by sharks as the large reptiles that had once ruled went extinct. Archaic mammals filled the world such as creodonts and early primates that evolved during the Mesozoic, and as a result, there was nothing over 10 kilograms. Mammals are still quite small.

The Eocene Epoch ranged from 55 million years to 33 million years ago. In the Early-Eocene, life was small and living in cramped jungles, much like the Paleocene. There was nothing over the weight of 10 kilograms. Among them were early primates, whales and horses along with many other early forms of mammals. At the top of the food chains were huge birds, such as Gastornis. It is the only time in recorded history that birds ruled the world (excluding their ancestors, the dinosaurs). The temperature was 30 degrees Celsius with little temperature gradient from pole to pole. In the Mid-Eocene, the circum-Antarctic current between Australia and Antarctica formed which disrupted ocean currents worldwide and as a result caused a global cooling effect, shrinking the jungles. This allowed mammals to grow to mammoth proportions, such as whales which are, by now, almost fully aquatic. Mammals like Andrewsarchus were now at the top of the food-chain and sharks were replaced by whales such as Basilosaurus as rulers of the seas. The Late-Eocene saw the rebirth of seasons, which caused the expansion of savanna-like areas, along with the evolution of grass.

The Oligocene Epoch spans from 33 million to 23 million years ago. The Oligocene feature the expansion of grass which had led to many new species to evolve, including the first elephants, cats, dogs, marsupials and many other species still prevalent today. Many other species of plants evolved in this period too, such as the evergreen trees. A cooling period was still in effect and seasonal rains were as well. Mammals still continued to grow larger and larger. Paraceratherium, the largest land mammal to ever live evolved during this period, along with many other perissodactyls in an event known as the Grand coupre.

Neogene

Animals of the Miocene (Chalicotherium, Hyenadon, Entelodont...)

The Neogene spans from 23 million to 3 million years ago, and is the shortest geological period in the Phanerozoic Eon. It features 2 epochs: the Miocene, and the Pliocene.

The Miocene spans from 23 to 5 million years ago and is a period in which grass spreads further across, effectively dominating a large portion of the world, diminishing forests in the process. Kelp forests evolved, leading to new species such as sea otters to evolve. During this time, perissodactyls thrived, and evolved into many different varieties. Alongside them were the apes, which evolved into a staggering 30 species. Overall, arid and mountainous land dominated most of the world, as did grazers. The Tethys Sea finally closed with the creation of the Arabian Peninsula and in its wake left the Black, Red, Mediterranean and Caspian Seas. This only increased aridity. Many new plants evolved, and 95% of modern seed plants evolved in the mid-Miocene.

The Pliocene ranges from 5 to 2 million years ago. The Pliocene features dramatic climactic changes, which ultimately leads to modern species and plants. The most dramatic are the formation of Panama, and the accumulation of ice at the poles, leading to a massive die-off, India and Asia collide forming the Himalayas, the Rockies and Appalachian mountain ranges were formed, and the Mediterranean Sea dried up for the next several million years. Along with these major geological events, the Australopithecus evolves in Africa, beginning the human branch. Also, with the isthmus of Panama, animals migrate across North and South America, wreaking havoc on the local ecology. Climactic changes bring along savannas that are still continuing to spread across the world, Indian monsoons, deserts in East Asia, and the beginnings of the Sahara desert. The earth's continents and seas move into their present shapes, and the world map hasn't changed much since.

Quaternary

The Quaternary ranges from 3 million to present day, and features modern animals, and dramatic climate changes and features two epochs: the Pleistocene and the Holocene.

Mega-fauna of the Pleistocene (Mammoths, Cave lions, Woolly Rhino, Megaloceros, American Horses

The Pleistocene lasted from 3 million to 12,000 years ago. This epoch features the ice ages which is a result from the cooling effect that started in the Mid-Eocene. As the ice progressively migrated towards the equator, the areas north and south of the tropic line featured intense winters yet mild summers. Meanwhile, Africa experienced terrible droughts which resulted in the creation of the Sahara, Namib, and Kalahari deserts. To cope, many animals evolved including Mammoths, Giant ground sloths, Dire wolves and most famously Homo sapiens. 100,000 years ago marked the end of one of the worst droughts of Africa, and the expansion of primitive man. As the Pleistocene draws to a close, one of the largest die-outs causes many mega-fauna to die off, including the last hominid species (excluding Homo sapiens). All continents are effected, but Africa isn't hit quite as hard.

The Holocene ranges from 12,000 years ago to present day. Also known as "the Age of Man", the Holocene features the rise of man on his path to sentience. All recorded history and "the history of the world" lies within the boundaries of the Holocene epoch. Human activity, however, is being blamed for a die-out that has been going on since 10,000 B.C.E. commonly referred to as "the Sixth Extinction" with an estimated extinction rate of 140,000 species per year.

See also

• Alpha taxonomy
• Astroecology
• Biological organisation
• Biosignature
• Chronobiology
• Evolutionary history of life
• Life on Mars
• Life on Titan
• Life on Venus
• Lists of organisms by population
• Non-cellular life
• Phylogenetics

Further reading

• Kauffman, Stuart. The Adjacent Possible: A Talk with Stuart Kauffman
• Seeding the Universe With Life Legacy Books, Washington D. C., 2000, ISBN 0-476-00330-X
• Walker, Martin G. LIFE! Why We Exist...And What We Must Do to Survive Dog Ear Publishing, 2006, ISBN 1-59858-243-7

External links

• Wikispecies – a free directory of life
• Resources for life in the Solar System and in galaxy, and the potential scope of life in the cosmological future
• "The Adjacent Possible: A Talk with Stuart Kauffman"
• Stanford Encyclopedia of Philosophy entry
• The Kingdoms of Life