An organism is any living thing that functions as an individual. Such a definition raises more problems than it solves, not least because the concept of an individual is also difficult. Many criteria, few of them widely accepted, have been proposed to define what an organism is. Among the most common is that an organism has autonomous reproduction, growth, and metabolism. This would exclude viruses, despite the fact that they evolve like organisms. Other problematic cases include colonial organisms; a colony of eusocial insects is organised adaptively, and has germ-soma specialisation, with some insects reproducing, others not, like cells in an animal's body. The body of a siphonophore, a jelly-like marine animal, is composed of organism-like zooids, but the whole structure looks and functions much like an animal such as a jellyfish, the parts collaborating to provide the functions of the colonial organism.
The evolutionary biologists David Queller and Joan Strassmann state that "organismality", the qualities or attributes that define an entity as an organism, has evolved socially as groups of simpler units (from cells upwards) came to cooperate without conflicts. They propose that cooperation should be used as the "defining trait" of an organism. This would treat many types of collaboration, including the fungus/alga partnership of different species in a lichen, or the permanent sexual partnership of an anglerfish, as an organism.
Etymology
The term "organism" (from the Ancient Greek ὀργανισμός, derived from órganon, meaning 'instrument, implement, tool', 'organ of sense', or 'apprehension') first appeared in the English language in the 1660s with the now-obsolete meaning of an organic structure or organization. It is related to the verb "organize". In his 1790 Critique of Judgment, Immanuel Kant defined an organism as "both an organized and a self-organizing being".
Whether criteria exist, or are needed
Among the criteria that have been proposed for being an organism are:
- autonomous reproduction, growth, and metabolism
- noncompartmentability – structure cannot be divided without losing functionality.Richard Dawkins stated this as "the quality of being sufficiently heterogeneous in form to be rendered non-functional if cut in half". However, many organisms can be cut into pieces which then grow into whole organisms.
- individuality – the entity has simultaneous holdings of genetic uniqueness, genetic homogeneity and autonomy
- an immune response, separating self from foreign
- "anti-entropy", the ability to maintain order, a concept first proposed by Erwin Schrödinger; or in another form, that Claude Shannon's information theory can be used to identify organisms as capable of self-maintaining their information content
Other scientists think that the concept of the organism is inadequate in biology; that the concept of individuality is problematic; and from a philosophical point of view, question whether such a definition is necessary.
Problematic cases include colonial organisms: for instance, a colony of eusocial insects fulfills criteria such as adaptive organisation and germ-soma specialisation. If so, the same argument, or a criterion of high co-operation and low conflict, would include some mutualistic (e.g. lichens) and sexual partnerships (e.g. anglerfish) as organisms. If group selection occurs, then a group could be viewed as a superorganism, optimized by group adaptation.
Another view is that attributes like autonomy, genetic homogeneity and genetic uniqueness should be examined separately, rather than requiring that an organism possess all of them. On this view, there are multiple dimensions to biological individuality, resulting in several types of organism.
Organisms at differing levels of biological organisation
Differing levels of biological organisation give rise to potentially different understandings of the nature of organisms. A unicellular organism is a microorganism such as a protist, bacterium, or archaean, composed of a single cell, which may contain functional structures called organelles. A multicellular organism such as an animal, plant, fungus, or alga is composed of many cells, often specialised. A colonial organism such as a siphonophore is a being which functions as an individual but is composed of communicating individuals. A superorganism is a colony, such as of ants, consisting of many individuals working together as a single functional or social unit. A mutualism is a partnership of two or more species which each provide some of the needs of the other. A lichen consists of fungi and algae or cyanobacteria, with a bacterial microbiome; together, they are able to flourish as a kind of organism, the components having different functions, in habitats such as dry rocks where neither could grow alone. The evolutionary biologists David Queller and Joan Strassmann state that "organismality" has evolved socially, as groups of simpler units (from cells upwards) came to cooperate without conflicts. They propose that cooperation should be used as the "defining trait" of an organism.
| Level | Example | Composition | Metabolism, growth, reproduction | Co-operation |
|---|---|---|---|---|
| Virus | Tobacco mosaic virus | Nucleic acid, protein | No | No metabolism, so not living, not an organism, say many biologists; but they evolve, their genes collaborating to manipulate the host |
| Unicellular organism | Paramecium | One cell, with organelles e.g. cilia for specific functions | Yes | Inter-cellular (inter-organismal) signalling |
| Swarming protistan | Dictyostelium (cellular slime mould) | Unicellular amoebae | Yes | Free-living unicellular amoebae for most of lifetime; swarm and aggregate to a multicellular slug, cells specialising to form a dead stalk and a fruiting body |
| Multicellular organism | Mushroom-forming fungus | Cells, grouped into organs for specific functions (e.g. reproduction) | Yes | Cell specialisation, communication |
| Permanent sexual partnership | Anglerfish | Male and female permanently fastened together | Yes | Male provides male gametes; female provides all other functions |
| Mutualism | Lichen | Organisms of different species | Yes | Fungus provides structure, absorbs water and minerals; alga photosynthesises |
| Joined colony | Siphonophore | Zooids joined together | Yes | Organism specialisation; inter-organism signalling |
| Superorganism | Ant colony | Individuals living together | Yes | Organism specialisation (many ants do not reproduce); inter-organism signalling |
Samuel Díaz‐Muñoz and colleagues (2016) accept Queller and Strassmann's view that organismality can be measured wholly by degrees of cooperation and of conflict. They state that this situates organisms in evolutionary time, so that organismality is context dependent. They suggest that highly integrated life forms, which are not context dependent, may evolve through context-dependent stages towards complete unification.
Boundary cases
Viruses
Viruses are not typically considered to be organisms, because they are incapable of autonomous reproduction, growth, metabolism, or homeostasis. Although viruses have a few enzymes and molecules like those in living organisms, they have no metabolism of their own; they cannot synthesize the organic compounds from which they are formed. In this sense, they are similar to inanimate matter. Viruses have their own genes, and they evolve. Thus, an argument that viruses should be classed as living organisms is their ability to undergo evolution and replicate through self-assembly. However, some scientists argue that viruses neither evolve nor self-reproduce. Instead, viruses are evolved by their host cells, meaning that there was co-evolution of viruses and host cells. If host cells did not exist, viral evolution would be impossible. As for reproduction, viruses rely on hosts' machinery to replicate. The discovery of viruses with genes coding for energy metabolism and protein synthesis fuelled the debate about whether viruses are living organisms, but the genes have a cellular origin. Most likely, they were acquired through horizontal gene transfer from viral hosts.
| Capability | Cellular organism | Virus |
|---|---|---|
| Metabolism | Yes | No, rely entirely on host cell |
| Growth | Yes | No, just self-assembly |
| Reproduction | Yes | No, rely entirely on host cell |
| Store genetic information about themselves | DNA | DNA or RNA |
| Able to evolve | Yes: mutation, recombination, natural selection | Yes: high mutation rate, natural selection |
There is an argument for viewing viruses as cellular organisms. Some researchers perceive viruses not as virions alone, which they believe are just spores of an organism, but as a virocell - an ontologically mature viral organism that has cellular structure. Such virus is a result of infection of a cell and shows all major physiological properties of other organisms: metabolism, growth, and reproduction, therefore, life in its effective presence.
Organism-like colonies
The philosopher Jack A. Wilson examines some boundary cases to demonstrate that the concept of organism is not sharply defined. In his view, sponges, lichens, siphonophores, slime moulds, and eusocial colonies such as those of ants or naked molerats, all lie in the boundary zone between being definite colonies and definite organisms (or superorganisms).
| Function | Colonial siphonophore | Jellyfish |
|---|---|---|
| Buoyancy | Top of colony is gas-filled | Jelly |
| Propulsion | Nectophores co-ordinate to pump water | Body pulsates to pump water |
| Feeding | Palpons and gastrozooids ingest prey, feed other zooids | Tentacles trap prey, pass it to mouth |
| Functional structure | Single functional individual | Single functional individual |
| Composition | Many zooids, possibly individuals | Many cells |
Synthetic organisms
Scientists and bio-engineers are experimenting with different types of synthetic organism, from chimaeras composed of cells from two or more species, cyborgs including electromechanical limbs, hybrots containing both electronic and biological elements, and other combinations of systems that have variously evolved and been designed.
An evolved organism takes its form by the partially understood mechanisms of evolutionary developmental biology, in which the genome directs an elaborated series of interactions to produce successively more elaborate structures. The existence of chimaeras and hybrids demonstrates that these mechanisms are "intelligently" robust in the face of radically altered circumstances at all levels from molecular to organismal.
Synthetic organisms already take diverse forms, and their diversity will increase. What they all have in common is a teleonomic or goal-seeking behaviour that enables them to correct errors of many kinds so as to achieve whatever result they are designed for. Such behaviour is reminiscent of intelligent action by organisms; intelligence is seen as an embodied form of cognition.
References
- Mosby's Dictionary of Medicine, Nursing and Health Professions (10th ed.). St. Louis, Missouri: Elsevier. 2017. p. 1281. ISBN 978-0-3232-2205-1.
- ὄργανον. Liddell, Henry George; Scott, Robert; A Greek–English Lexicon at the Perseus Project
- "organism (n.)". Online Etymology Dictionary. Retrieved 11 April 2024.
- Kant, Immanuel (1790). Critique of Judgment. Lagarde und Friederich. §65 5:374.
- Huneman, Philippe (2017). "Kant's Concept of Organism Revisited: A Framework for a Possible Synthesis between Developmentalism and Adaptationism?". The Monist. 100 (3): 373–390. doi:10.1093/monist/onx016. JSTOR 26370801.
- Rosen, Robert (September 1958). "A relational theory of biological systems". The Bulletin of Mathematical Biophysics. 20 (3): 245–260. doi:10.1007/BF02478302. ISSN 0007-4985.
- Moreira, D.; López-García, P.N. (April 2009). "Ten reasons to exclude viruses from the tree of life". Nature Reviews Microbiology. 7 (4): 306–311. doi:10.1038/nrmicro2108. PMID 19270719. S2CID 3907750.
- Wilson, Jack A. (2000). "Ontological butchery: organism concepts and biological generalizations". Philosophy of Science. 67: 301–311. doi:10.1086/392827. JSTOR 188676. S2CID 84168536.
- Santelices, Bernabé (April 1999). "How many kinds of individual are there?". Trends in Ecology & Evolution. 14 (4): 152–155. doi:10.1016/S0169-5347(98)01519-5. PMID 10322523.
- Pradeu, T. (2010). "What is an organism? An immunological answer". History and Philosophy of the Life Sciences. 32 (2–3): 247–267. PMID 21162370.
- Bailly, Francis; Longo, Giuseppe (2009). "Biological Organization and Anti-entropy". Journal of Biological Systems. 17 (1): 63–96. doi:10.1142/S0218339009002715. ISSN 0218-3390.
- Piast, Radosław W. (June 2019). "Shannon's information, Bernal's biopoiesis and Bernoulli distribution as pillars for building a definition of life". Journal of Theoretical Biology. 470: 101–107. Bibcode:2019JThBi.470..101P. doi:10.1016/j.jtbi.2019.03.009. PMID 30876803. S2CID 80625250.
- Bateson, Patrick (February 2005). "The return of the whole organism". Journal of Biosciences. 30 (1): 31–39. doi:10.1007/BF02705148. PMID 15824439. S2CID 26656790.
- Clarke, E. (2010). "The problem of biological individuality". Biological Theory. 5 (4): 312–325. doi:10.1162/BIOT_a_00068. S2CID 28501709.
- Pepper, J.W.; Herron, M.D. (November 2008). "Does biology need an organism concept?". Biological Reviews of the Cambridge Philosophical Society. 83 (4): 621–627. doi:10.1111/j.1469-185X.2008.00057.x. PMID 18947335. S2CID 4942890.
- Wilson, R. (2007). "The biological notion of individual". Stanford Encyclopedia of Philosophy.
- Folse, H.J., III; Roughgarden, J. (December 2010). "What is an individual organism? A multilevel selection perspective". The Quarterly Review of Biology. 85 (4): 447–472. doi:10.1086/656905. PMID 21243964. S2CID 19816447.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - Queller, David C.; Strassmann, Joan E. (November 2009). "Beyond society: the evolution of organismality". Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences. 364 (1533): 3143–3155. doi:10.1098/rstb.2009.0095. PMC 2781869. PMID 19805423.
- Gardner, A.; Grafen, A. (April 2009). "Capturing the superorganism: a formal theory of group adaptation". Journal of Evolutionary Biology. 22 (4): 659–671. doi:10.1111/j.1420-9101.2008.01681.x. PMID 19210588. S2CID 8413751.
- Santelices, B. (April 1999). "How many kinds of individual are there?". Trends in Ecology & Evolution. 14 (4): 152–155. doi:10.1016/s0169-5347(98)01519-5. PMID 10322523.
- Lücking, Robert; Leavitt, Steven D.; Hawksworth, David L. (2021). "Species in lichen-forming fungi: balancing between conceptual and practical considerations, and between phenotype and phylogenomics". Fungal Diversity. 109 (1): 99–154. doi:10.1007/s13225-021-00477-7.
- Hine, R.S. (2008). A Dictionary of Biology (6th ed.). Oxford: Oxford University Press. p. 461. ISBN 978-0-19-920462-5.
- Kelly, Kevin (1994). Out of control: the new biology of machines, social systems and the economic world. Boston: Addison-Wesley. pp. 98. ISBN 978-0-201-48340-6.
- Díaz-Muñoz, Samuel L.; Boddy, Amy M.; Dantas, Gautam; Waters, Christopher M.; Bronstein, Judith L. (2016). "Contextual organismality: Beyond pattern to process in the emergence of organisms". Evolution. 70 (12): 2669–2677. doi:10.1111/evo.13078. ISSN 0014-3820. PMC 5132100. PMID 27704542.
- Forterre, Patrick (4 October 2012). "The virocell concept and environmental microbiology". The ISME Journal. 7 (2): 233–236. doi:10.1038/ismej.2012.110. ISSN 1751-7362. PMC 3554396. PMID 23038175.
- Bandea, Claudiu I. (1983). "A new theory on the origin and the nature of viruses". Journal of Theoretical Biology. 105 (4): 591–602. Bibcode:1983JThBi.105..591B. doi:10.1016/0022-5193(83)90221-7. PMID 6672474.
- Clawson, Wesley P.; Levin, Michael (1 January 2023). "Endless forms most beautiful 2.0: teleonomy and the bioengineering of chimaeric and synthetic organisms". Biological Journal of the Linnean Society. 138 (1): 141. doi:10.1093/biolinnean/blac116. ISSN 0024-4066.
External links
- "The Tree of Life". Tree of Life Web Project.
- "Indexing the world's known species". Species 2000.
An organism is any living thing that functions as an individual Such a definition raises more problems than it solves not least because the concept of an individual is also difficult Many criteria few of them widely accepted have been proposed to define what an organism is Among the most common is that an organism has autonomous reproduction growth and metabolism This would exclude viruses despite the fact that they evolve like organisms Other problematic cases include colonial organisms a colony of eusocial insects is organised adaptively and has germ soma specialisation with some insects reproducing others not like cells in an animal s body The body of a siphonophore a jelly like marine animal is composed of organism like zooids but the whole structure looks and functions much like an animal such as a jellyfish the parts collaborating to provide the functions of the colonial organism The evolutionary biologists David Queller and Joan Strassmann state that organismality the qualities or attributes that define an entity as an organism has evolved socially as groups of simpler units from cells upwards came to cooperate without conflicts They propose that cooperation should be used as the defining trait of an organism This would treat many types of collaboration including the fungus alga partnership of different species in a lichen or the permanent sexual partnership of an anglerfish as an organism EtymologyThe term organism from the Ancient Greek ὀrganismos derived from organon meaning instrument implement tool organ of sense or apprehension first appeared in the English language in the 1660s with the now obsolete meaning of an organic structure or organization It is related to the verb organize In his 1790 Critique of Judgment Immanuel Kant defined an organism as both an organized and a self organizing being Whether criteria exist or are neededOne criterion proposes that an organism cannot be divided without losing functionality This basil plant cutting is however developing new adventitious roots from a small bit of stem forming a new plant Among the criteria that have been proposed for being an organism are autonomous reproduction growth and metabolism noncompartmentability structure cannot be divided without losing functionality Richard Dawkins stated this as the quality of being sufficiently heterogeneous in form to be rendered non functional if cut in half However many organisms can be cut into pieces which then grow into whole organisms individuality the entity has simultaneous holdings of genetic uniqueness genetic homogeneity and autonomy an immune response separating self from foreign anti entropy the ability to maintain order a concept first proposed by Erwin Schrodinger or in another form that Claude Shannon s information theory can be used to identify organisms as capable of self maintaining their information content Other scientists think that the concept of the organism is inadequate in biology that the concept of individuality is problematic and from a philosophical point of view question whether such a definition is necessary Problematic cases include colonial organisms for instance a colony of eusocial insects fulfills criteria such as adaptive organisation and germ soma specialisation If so the same argument or a criterion of high co operation and low conflict would include some mutualistic e g lichens and sexual partnerships e g anglerfish as organisms If group selection occurs then a group could be viewed as a superorganism optimized by group adaptation Another view is that attributes like autonomy genetic homogeneity and genetic uniqueness should be examined separately rather than requiring that an organism possess all of them On this view there are multiple dimensions to biological individuality resulting in several types of organism Organisms at differing levels of biological organisationA lichen consists of a body formed mainly by fungi with unicellular algae or cyanobacteria green interspersed within the structure and a bacterial microbiome The species are mutually interdependent like cells within a multicellular organism Differing levels of biological organisation give rise to potentially different understandings of the nature of organisms A unicellular organism is a microorganism such as a protist bacterium or archaean composed of a single cell which may contain functional structures called organelles A multicellular organism such as an animal plant fungus or alga is composed of many cells often specialised A colonial organism such as a siphonophore is a being which functions as an individual but is composed of communicating individuals A superorganism is a colony such as of ants consisting of many individuals working together as a single functional or social unit A mutualism is a partnership of two or more species which each provide some of the needs of the other A lichen consists of fungi and algae or cyanobacteria with a bacterial microbiome together they are able to flourish as a kind of organism the components having different functions in habitats such as dry rocks where neither could grow alone The evolutionary biologists David Queller and Joan Strassmann state that organismality has evolved socially as groups of simpler units from cells upwards came to cooperate without conflicts They propose that cooperation should be used as the defining trait of an organism Queller and Strassmann s view of organisms as cooperating entities at differing levels of biological organisation Level Example Composition Metabolism growth reproduction Co operationVirus Tobacco mosaic virus Nucleic acid protein No No metabolism so not living not an organism say many biologists but they evolve their genes collaborating to manipulate the hostUnicellular organism Paramecium One cell with organelles e g cilia for specific functions Yes Inter cellular inter organismal signallingSwarming protistan Dictyostelium cellular slime mould Unicellular amoebae Yes Free living unicellular amoebae for most of lifetime swarm and aggregate to a multicellular slug cells specialising to form a dead stalk and a fruiting bodyMulticellular organism Mushroom forming fungus Cells grouped into organs for specific functions e g reproduction Yes Cell specialisation communicationPermanent sexual partnership Anglerfish Male and female permanently fastened together Yes Male provides male gametes female provides all other functionsMutualism Lichen Organisms of different species Yes Fungus provides structure absorbs water and minerals alga photosynthesisesJoined colony Siphonophore Zooids joined together Yes Organism specialisation inter organism signallingSuperorganism Ant colony Individuals living together Yes Organism specialisation many ants do not reproduce inter organism signalling Samuel Diaz Munoz and colleagues 2016 accept Queller and Strassmann s view that organismality can be measured wholly by degrees of cooperation and of conflict They state that this situates organisms in evolutionary time so that organismality is context dependent They suggest that highly integrated life forms which are not context dependent may evolve through context dependent stages towards complete unification Boundary casesViruses A virus such as tobacco mosaic virus is not a cell it contains only its genetic material and a protein coat Viruses are not typically considered to be organisms because they are incapable of autonomous reproduction growth metabolism or homeostasis Although viruses have a few enzymes and molecules like those in living organisms they have no metabolism of their own they cannot synthesize the organic compounds from which they are formed In this sense they are similar to inanimate matter Viruses have their own genes and they evolve Thus an argument that viruses should be classed as living organisms is their ability to undergo evolution and replicate through self assembly However some scientists argue that viruses neither evolve nor self reproduce Instead viruses are evolved by their host cells meaning that there was co evolution of viruses and host cells If host cells did not exist viral evolution would be impossible As for reproduction viruses rely on hosts machinery to replicate The discovery of viruses with genes coding for energy metabolism and protein synthesis fuelled the debate about whether viruses are living organisms but the genes have a cellular origin Most likely they were acquired through horizontal gene transfer from viral hosts Comparison of cellular organisms and viruses Capability Cellular organism VirusMetabolism Yes No rely entirely on host cellGrowth Yes No just self assemblyReproduction Yes No rely entirely on host cellStore genetic information about themselves DNA DNA or RNAAble to evolve Yes mutation recombination natural selection Yes high mutation rate natural selection There is an argument for viewing viruses as cellular organisms Some researchers perceive viruses not as virions alone which they believe are just spores of an organism but as a virocell an ontologically mature viral organism that has cellular structure Such virus is a result of infection of a cell and shows all major physiological properties of other organisms metabolism growth and reproduction therefore life in its effective presence Organism like colonies Apolemia a colonial siphonophore that functions as a single individual The philosopher Jack A Wilson examines some boundary cases to demonstrate that the concept of organism is not sharply defined In his view sponges lichens siphonophores slime moulds and eusocial colonies such as those of ants or naked molerats all lie in the boundary zone between being definite colonies and definite organisms or superorganisms Jack A Wilson s analysis of the similar organism like nature of siphonophores and jellyfish Function Colonial siphonophore JellyfishBuoyancy Top of colony is gas filled JellyPropulsion Nectophores co ordinate to pump water Body pulsates to pump waterFeeding Palpons and gastrozooids ingest prey feed other zooids Tentacles trap prey pass it to mouthFunctional structure Single functional individual Single functional individualComposition Many zooids possibly individuals Many cells Synthetic organisms Insect cyborg Scientists and bio engineers are experimenting with different types of synthetic organism from chimaeras composed of cells from two or more species cyborgs including electromechanical limbs hybrots containing both electronic and biological elements and other combinations of systems that have variously evolved and been designed An evolved organism takes its form by the partially understood mechanisms of evolutionary developmental biology in which the genome directs an elaborated series of interactions to produce successively more elaborate structures The existence of chimaeras and hybrids demonstrates that these mechanisms are intelligently robust in the face of radically altered circumstances at all levels from molecular to organismal Synthetic organisms already take diverse forms and their diversity will increase What they all have in common is a teleonomic or goal seeking behaviour that enables them to correct errors of many kinds so as to achieve whatever result they are designed for Such behaviour is reminiscent of intelligent action by organisms intelligence is seen as an embodied form of cognition ReferencesMosby s Dictionary of Medicine Nursing and Health Professions 10th ed St Louis Missouri Elsevier 2017 p 1281 ISBN 978 0 3232 2205 1 ὄrganon Liddell Henry George Scott Robert A Greek English Lexicon at the Perseus Project organism n Online Etymology Dictionary Retrieved 11 April 2024 Kant Immanuel 1790 Critique of Judgment Lagarde und Friederich 65 5 374 Huneman Philippe 2017 Kant s Concept of Organism Revisited A Framework for a Possible Synthesis between Developmentalism and Adaptationism The Monist 100 3 373 390 doi 10 1093 monist onx016 JSTOR 26370801 Rosen Robert September 1958 A relational theory of biological systems The Bulletin of Mathematical Biophysics 20 3 245 260 doi 10 1007 BF02478302 ISSN 0007 4985 Moreira D Lopez Garcia P N April 2009 Ten reasons to exclude viruses from the tree of life Nature Reviews Microbiology 7 4 306 311 doi 10 1038 nrmicro2108 PMID 19270719 S2CID 3907750 Wilson Jack A 2000 Ontological butchery organism concepts and biological generalizations Philosophy of Science 67 301 311 doi 10 1086 392827 JSTOR 188676 S2CID 84168536 Santelices Bernabe April 1999 How many kinds of individual are there Trends in Ecology amp Evolution 14 4 152 155 doi 10 1016 S0169 5347 98 01519 5 PMID 10322523 Pradeu T 2010 What is an organism An immunological answer History and Philosophy of the Life Sciences 32 2 3 247 267 PMID 21162370 Bailly Francis Longo Giuseppe 2009 Biological Organization and Anti entropy Journal of Biological Systems 17 1 63 96 doi 10 1142 S0218339009002715 ISSN 0218 3390 Piast Radoslaw W June 2019 Shannon s information Bernal s biopoiesis and Bernoulli distribution as pillars for building a definition of life Journal of Theoretical Biology 470 101 107 Bibcode 2019JThBi 470 101P doi 10 1016 j jtbi 2019 03 009 PMID 30876803 S2CID 80625250 Bateson Patrick February 2005 The return of the whole organism Journal of Biosciences 30 1 31 39 doi 10 1007 BF02705148 PMID 15824439 S2CID 26656790 Clarke E 2010 The problem of biological individuality Biological Theory 5 4 312 325 doi 10 1162 BIOT a 00068 S2CID 28501709 Pepper J W Herron M D November 2008 Does biology need an organism concept Biological Reviews of the Cambridge Philosophical Society 83 4 621 627 doi 10 1111 j 1469 185X 2008 00057 x PMID 18947335 S2CID 4942890 Wilson R 2007 The biological notion of individual Stanford Encyclopedia of Philosophy Folse H J III Roughgarden J December 2010 What is an individual organism A multilevel selection perspective The Quarterly Review of Biology 85 4 447 472 doi 10 1086 656905 PMID 21243964 S2CID 19816447 a href wiki Template Cite journal title Template Cite journal cite journal a CS1 maint multiple names authors list link Queller David C Strassmann Joan E November 2009 Beyond society the evolution of organismality Philosophical Transactions of the Royal Society of London Series B Biological Sciences 364 1533 3143 3155 doi 10 1098 rstb 2009 0095 PMC 2781869 PMID 19805423 Gardner A Grafen A April 2009 Capturing the superorganism a formal theory of group adaptation Journal of Evolutionary Biology 22 4 659 671 doi 10 1111 j 1420 9101 2008 01681 x PMID 19210588 S2CID 8413751 Santelices B April 1999 How many kinds of individual are there Trends in Ecology amp Evolution 14 4 152 155 doi 10 1016 s0169 5347 98 01519 5 PMID 10322523 Lucking Robert Leavitt Steven D Hawksworth David L 2021 Species in lichen forming fungi balancing between conceptual and practical considerations and between phenotype and phylogenomics Fungal Diversity 109 1 99 154 doi 10 1007 s13225 021 00477 7 Hine R S 2008 A Dictionary of Biology 6th ed Oxford Oxford University Press p 461 ISBN 978 0 19 920462 5 Kelly Kevin 1994 Out of control the new biology of machines social systems and the economic world Boston Addison Wesley pp 98 ISBN 978 0 201 48340 6 Diaz Munoz Samuel L Boddy Amy M Dantas Gautam Waters Christopher M Bronstein Judith L 2016 Contextual organismality Beyond pattern to process in the emergence of organisms Evolution 70 12 2669 2677 doi 10 1111 evo 13078 ISSN 0014 3820 PMC 5132100 PMID 27704542 Forterre Patrick 4 October 2012 The virocell concept and environmental microbiology The ISME Journal 7 2 233 236 doi 10 1038 ismej 2012 110 ISSN 1751 7362 PMC 3554396 PMID 23038175 Bandea Claudiu I 1983 A new theory on the origin and the nature of viruses Journal of Theoretical Biology 105 4 591 602 Bibcode 1983JThBi 105 591B doi 10 1016 0022 5193 83 90221 7 PMID 6672474 Clawson Wesley P Levin Michael 1 January 2023 Endless forms most beautiful 2 0 teleonomy and the bioengineering of chimaeric and synthetic organisms Biological Journal of the Linnean Society 138 1 141 doi 10 1093 biolinnean blac116 ISSN 0024 4066 External links The Tree of Life Tree of Life Web Project Indexing the world s known species Species 2000
