Editing Primary nutritional groups

{{Short description|Group of organisms}}
{{more citations needed|date=September 2010}}
'''Primary nutritional groups''' are groups of [[organism]]s, divided in relation to the nutrition mode according to the sources of energy and carbon, needed for living, growth and reproduction. The sources of energy can be light or chemical compounds; the sources of carbon can be of organic or inorganic origin.<ref name="pmid17028233">{{cite journal | vauthors = Eiler A | title = Evidence for the ubiquity of mixotrophic bacteria in the upper ocean: implications and consequences | journal = Applied and Environmental Microbiology | volume = 72 | issue = 12 | pages = 7431–7 | date = December 2006 | pmid = 17028233 | pmc = 1694265 | doi = 10.1128/AEM.01559-06 | bibcode = 2006ApEnM..72.7431E | quote = Table 1: Definitions of metabolic strategies to obtain carbon and energy }}</ref>

The terms ''[[aerobic respiration]]'', ''[[anaerobic respiration]]'' and ''[[fermentation (biochemistry)|fermentation]]'' (''[[substrate-level phosphorylation]]'') do not refer to primary nutritional groups, but simply reflect the different use of possible electron acceptors in particular organisms, such as {{O2}} in aerobic respiration, or [[nitrate]] ({{chem|NO|3|−}}), [[sulfate]] ({{chem|SO|4|2−}}) or [[fumarate]] in anaerobic respiration, or various metabolic intermediates in fermentation.

== Primary sources of energy ==
[[Phototroph|''Phototrophs'']] absorb light in [[Photosynthetic pigment|photoreceptors]] and transform it into chemical energy.<br>
[[Chemotroph|''Chemotrophs'']] release chemical energy.

The freed energy is stored as [[Potential energy#Chemical potential energy|potential energy]] in [[adenosine triphosphate|ATP]], [[carbohydrate]]s, or [[protein]]s. Eventually, the energy is used for life processes such as moving, growth and reproduction.

Plants and some bacteria can alternate between phototrophy and chemotrophy, depending on the availability of light.

== Primary sources of reducing equivalents ==

[[Organotroph|''Organotrophs'']] use organic compounds as [[electron donor|electron/hydrogen donors]].<br>
[[Lithotroph|''Lithotrophs'']] use inorganic compounds as electron/hydrogen donors.

The [[electron]]s or hydrogen atoms from [[reducing equivalent]]s (electron donors) are needed by both phototrophs and chemotrophs in [[Redox|reduction-oxidation reactions]] that transfer energy in the anabolic processes of [[ATP synthesis]] (in heterotrophs) or [[biosynthesis]] (in autotrophs). The electron or hydrogen donors are taken up from the environment.

Organotrophic organisms are often also heterotrophic, using organic compounds as sources of both electrons and carbon. Similarly, lithotrophic organisms are often also autotrophic, using inorganic sources of electrons and {{CO2}} as their inorganic carbon source.

Some lithotrophic bacteria can utilize diverse sources of electrons, depending on the availability of possible donors.

The organic or inorganic substances (e.g., oxygen) used as electron acceptors needed in the catabolic processes of aerobic or anaerobic [[Cellular respiration|respiration]] and [[fermentation]] are not taken into account here.

For example, plants are lithotrophs because they use water as their electron donor for the electron transport chain across the thylakoid membrane. Animals are organotrophs because they use organic compounds as electron donors to synthesize ATP (plants also do this, but this is not taken into account). Both use oxygen in respiration as electron acceptor, but this character is not used to define them as lithotrophs.

== Primary sources of carbon ==

[[Heterotroph|''Heterotrophs'']] [[Metabolism|metabolize]] organic compounds to obtain carbon for growth and development.<br>
[[Autotroph|''Autotrophs'']] use [[carbon dioxide]] ({{CO2}}) as their source of carbon.

== Energy and carbon ==
[[Image:K 1033CR08-9 Yellow fungus on stalk.jpeg|thumb|240px|right|Yellow fungus]]
{| class="wikitable float-right" style="text-align:center" width="50%"
|+Classification of organisms based on their metabolism
|-
| rowspan=2 bgcolor="#FFFF00" |Energy source || bgcolor="#FFFF00" | Light || bgcolor="#FFFF00" | photo- || rowspan=2 colspan=2 | &nbsp; || rowspan=6 bgcolor="#7FC31C" | -troph
|-
| bgcolor="#FFFF00" | Molecules || bgcolor="#FFFF00" | chemo-
|-
| rowspan=2 bgcolor="#FFB300" | Electron donor || bgcolor="#FFB300" | [[Organic compounds]] || rowspan=2 |  &nbsp; || bgcolor="#FFB300" | organo- ||  rowspan=2 |  &nbsp;
|-
| bgcolor="#FFB300" | [[Inorganic compounds]] || bgcolor="#FFB300" | litho-
|-
| rowspan=2 bgcolor="#FB805F" | Carbon source || bgcolor="#FB805F" | [[Organic compounds]] || rowspan=2 colspan=2 | &nbsp; || bgcolor="#FB805F" | hetero-
|-
| bgcolor="#FB805F" | [[Carbon dioxide]] || bgcolor="#FB805F" | auto-
|}
A ''chemoorganoheterotrophic'' [[organism]] is one that requires organic [[substrate (biochemistry)|substrates]] to get its [[carbon]] for growth and development, and that obtains its energy from the decomposition of an organic compound. This group of organisms may be further subdivided according to what kind of organic substrate and compound they use. [[Decomposers]] are examples of chemoorganoheterotrophs which obtain carbon and electrons or hydrogen from dead organic matter. [[Herbivores]] and [[carnivores]] are examples of organisms that obtain carbon and electrons or hydrogen from living organic matter.

Chemoorganotrophs are [[organism]]s which use the chemical energy in [[organic compound]]s as their energy source and obtain electrons or hydrogen from the organic compounds, including sugars (i.e. [[glucose]]), fats and proteins.<ref name="kt">{{cite web |url=http://textbookofbacteriology.net/nutgro.html |title=Todar's Online Textbook of Bacteriology |access-date=2014-04-19 |year=2009| first = Kenneth | last = Todar | name-list-style = vanc |work=Nutrition and Growth of Bacteria }}</ref> Chemoheterotrophs also obtain the carbon atoms that they need for cellular function from these organic compounds.

All [[animal]]s are chemoheterotrophs (meaning they oxidize chemical compounds as a source of energy and carbon), as are [[fungus|fungi]], [[protozoa]], and some [[bacteria]]. The important differentiation amongst this group is that chemoorganotrophs oxidize only organic compounds while [[chemolithotroph]]s instead use oxidation of [[inorganic compound]]s as a source of energy.<ref name="dk">{{cite book | vauthors = Kelly DP, Mason J, Wood A | chapter = Energy Metabolism in Chemolithotrophs | date = 1987 | veditors = van Verseveld HW, Duine JA | title = Microbial Growth on C1 Compounds | pages=186–7 | publisher=Springer | doi = 10.1007/978-94-009-3539-6_23 | isbn = 978-94-010-8082-8 }}</ref>

==Primary metabolism table==
The following table gives some examples for each nutritional group:<ref>{{cite journal | vauthors = Lwoff A, Van Niel CB, Ryan TF, Tatum EL | title = Nomenclature of nutritional types of microorganisms. | journal = Cold Spring Harbor Symposia on Quantitative Biology | edition = 5th | date = 1946 | volume = 11 | pages = 302–3 | url = http://symposium.cshlp.org/content/11/local/back-matter.pdf }}</ref><ref>{{cite book | vauthors = Andrews JH | date = 1991 | title = Comparative Ecology of Microorganisms and Macroorganisms | publisher = Springer | pages = 68 | url = https://books.google.com/books?id=4ZDhBwAAQBAJ | isbn = 978-0-387-97439-2 }}</ref><ref name="pmid23443991">{{cite journal | vauthors = Yafremava LS, Wielgos M, Thomas S, Nasir A, Wang M, Mittenthal JE, Caetano-Anollés G | title = A general framework of persistence strategies for biological systems helps explain domains of life | journal = Frontiers in Genetics | volume = 4 | pages = 16 | date = 2013 | pmid = 23443991 | pmc = 3580334 | doi = 10.3389/fgene.2013.00016 | doi-access = free }}</ref><ref>{{cite book | veditors = [[Lynn Margulis|Margulis L]], McKhann HI, Olendzenski L | title = Illustrated Glossary of Protoctista: Vocabulary of the Algae, Apicomplexa, Ciliates, Foraminifera, Microspora, Water Molds, Slime Molds, and the Other Protoctists | publisher = Jones & Bartlett Learning | date = 1993 | pages = xxv | url = https://books.google.com/books?id=y55Efu3baksC | isbn = 978-0-86720-081-2 }}</ref>
{| class="wikitable"
|-
! Energy<br>source
! Electron/<br>H-atom<br>donor
! Carbon source
! Name
! Examples
|-
|rowspan="4" style="background: #ddffdd"| Sun Light<br>''[[phototroph|Photo-]]''
|rowspan="2" style="background: #ffdfcc"| Organic<br>''[[organotroph|-organo-]]''
|style="background: #ffbfbf"| Organic<br>''[[heterotroph|-heterotroph]]''
| <span style="color:#008800;">Photo</span><span style="color:#888800;">organo</span><span style="color:#880000;">heterotroph</span>
|Some bacteria (''[[Rhodobacter capsulatus]]'', [[Heliobacteria]]) and some archaea ([[Haloarchaea]])<ref name="Morris 2019">Morris, J. et al. (2019). "Biology: How Life Works", 3rd edition, W. H. Freeman. {{ISBN|978-1319017637}}</ref>
|-
|style="background: #bfdfff"| Carbon dioxide<br>''[[autotroph|-autotroph]]''
|  <span style="color:#008800;">Photo</span><span style="color:#888800;">organo</span><span style="color:#000088;">autotroph</span>
|Some bacteria perform anoxygenic photosynthesis and fix atmospheric carbon (''[[Chloroflexia]]'')
|-
|rowspan="2" style="background: #dfbfff"| Inorganic<br>''[[lithotroph|-litho-]]''*
|style="background: #ffbfbf"| Organic<br>''-heterotroph''
| <span style="color:#008800;">Photo</span><span style="color:#880088;">litho</span><span style="color:#880000;">heterotroph</span>
|[[Purple non-sulfur bacteria]]
|-
|style="background: #bfdfff"| Carbon dioxide<br>''-autotroph''
| <span style="color:#008800;">Photo</span><span style="color:#880088;">litho</span><span style="color:#000088;">autotroph</span>
| Some bacteria ([[cyanobacteria]]), some eukaryotes ([[algae|eukaryotic algae]], [[land plants]]). [[Photosynthesis]]. 
|-
|rowspan="4" style="background: #ffdddd"| Breaking<br> Chemical<br> Compounds<br>''[[Chemotroph|Chemo-]]''
|rowspan="2" style="background: #ffdfcc"| Organic<br>''-organo-''
|style="background: #ffbfbf"| Organic<br>''-heterotroph''
| <span style="color:#840048;">Chemo</span><span style="color:#888800;">organo</span><span style="color:#880000;">heterotroph</span>
| [[Bacteria#Predators|Predatory]], [[parasite|parasitic]], and [[saprophyte|saprophytic]] prokaryotes. Some eukaryotes (heterotrophic [[protist]]s, [[fungi]], [[animal]]s)

|-
|style="background: #bfdfff"| Carbon dioxide<br>''-autotroph''
| <span style="color:#840048;">Chemo</span><span style="color:#888800;">organo</span><span style="color:#000088;">autotroph</span>
| Some archaea ([[Anaerobic oxidation of methane|anaerobic methanotrophic archaea]]).<ref name="pmid23129626">{{cite journal | vauthors = Kellermann MY, Wegener G, Elvert M, Yoshinaga MY, Lin YS, Holler T, Mollar XP, Knittel K, Hinrichs KU | display-authors = 6 | title = Autotrophy as a predominant mode of carbon fixation in anaerobic methane-oxidizing microbial communities | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 109 | issue = 47 | pages = 19321–6 | date = November 2012 | pmid = 23129626 | pmc = 3511159 | doi = 10.1073/pnas.1208795109 | bibcode = 2012PNAS..10919321K | doi-access = free }}</ref> [[Chemosynthesis]], synthetically autotrophic ''[[Escherichia coli]]'' bacteria<ref name=Gleizer19>{{cite journal | vauthors = Gleizer S, Ben-Nissan R, Bar-On YM, Antonovsky N, Noor E, Zohar Y, Jona G, Krieger E, Shamshoum M, Bar-Even A, Milo R | display-authors = 6 | title = Conversion of Escherichia coli to Generate All Biomass Carbon from CO<sub>2</sub> | journal = Cell | volume = 179 | issue = 6 | pages = 1255–1263.e12 | date = November 2019 | pmid = 31778652 | pmc = 6904909 | doi = 10.1016/j.cell.2019.11.009 }}</ref> and ''[[Pichia pastoris]]'' yeast.<ref name=Gassler19>{{cite journal | vauthors = Gassler T, Sauer M, Gasser B, Egermeier M, Troyer C, Causon T, Hann S, Mattanovich D, Steiger MG | display-authors = 6 | title = The industrial yeast ''Pichia pastoris'' is converted from a heterotroph into an autotroph capable of growth on CO<sub>2</sub> | journal = Nature Biotechnology | volume = 38 | issue = 2 | pages = 210–6 | date = December 2019 | pmid = 31844294 | doi = 10.1038/s41587-019-0363-0 | pmc = 7008030 }}</ref>
|-
|rowspan="2" style="background: #dfbfff"| Inorganic<br>''-litho-''*
|style="background: #ffbfbf"| Organic<br>''-heterotroph''
| <span style="color:#840048;">Chemo</span><span style="color:#880088;">litho</span><span style="color:#880000;">heterotroph</span>
|Some bacteria (''[[Oceanithermus profundus]]'')<ref name="pmid12807196">{{cite journal | vauthors = Miroshnichenko ML, L'Haridon S, Jeanthon C, Antipov AN, Kostrikina NA, Tindall BJ, Schumann P, Spring S, Stackebrandt E, Bonch-Osmolovskaya EA | display-authors = 6 | title = ''Oceanithermus profundus'' gen. nov., sp. nov., a thermophilic, microaerophilic, facultatively chemolithoheterotrophic bacterium from a deep-sea hydrothermal vent | journal = International Journal of Systematic and Evolutionary Microbiology | volume = 53 | issue = Pt 3 | pages = 747–52 | date = May 2003 | pmid = 12807196 | doi = 10.1099/ijs.0.02367-0 | doi-access = free }}</ref>
|-
|style="background: #bfdfff"| Carbon dioxide<br>''-autotroph''
| <span style="color:#840048;">Chemo</span><span style="color:#880088;">litho</span><span style="color:#000088;">autotroph</span>
|Some bacteria (''[[Nitrobacter]]''), some archaea (''[[Methanobacteria]]''). [[Chemosynthesis]].
|}

<nowiki />*Some authors use ''-hydro-'' when the source is water.

{{Wiktionary|-troph}}
The common final part ''-troph'' is from Ancient Greek {{wikt-lang|grc|τροφή}} {{transliteration|grc|trophḗ}} "nutrition".

== Mixotrophs ==

Some, usually unicellular, organisms can switch between different metabolic modes, for example between photoautotrophy, photoheterotrophy, and chemoheterotrophy in ''Chroococcales''.<ref>{{cite journal | vauthors = Rippka R | title = Photoheterotrophy and chemoheterotrophy among unicellular blue-green algae | journal = Archives of Microbiology | volume = 87 | issue = 1 | pages = 93–98 | date = March 1972 | doi = 10.1007/BF00424781 | s2cid = 155161 }}</ref> ''[[Rhodopseudomonas palustris]]'' – another example – can grow with or without [[oxygen]], use either light, inorganic or organic compounds for energy.<ref>{{Cite journal |last1=Li |first1=Meijie |last2=Ning |first2=Peng |last3=Sun |first3=Yi |last4=Luo |first4=Jie |last5=Yang |first5=Jianming |date=2022 |title=Characteristics and Application of ''Rhodopseudomonas palustris'' as a Microbial Cell Factory |journal=Frontiers in Bioengineering and Biotechnology |volume=10 |page=897003 |doi=10.3389/fbioe.2022.897003 |pmid=35646843 |pmc=9133744 |issn=2296-4185|doi-access=free }}</ref> Such [[mixotroph]]ic organisms may dominate their [[habitat]], due to their capability to use more resources than either photoautotrophic or organoheterotrophic organisms.<ref name="Eiler_2006">{{cite journal | vauthors = Eiler A | title = Evidence for the ubiquity of mixotrophic bacteria in the upper ocean: implications and consequences | journal = Applied and Environmental Microbiology | volume = 72 | issue = 12 | pages = 7431–7 | date = December 2006 | pmid = 17028233 | pmc = 1694265 | doi = 10.1128/AEM.01559-06 | bibcode = 2006ApEnM..72.7431E }}</ref>

==Examples==
All sorts of combinations may exist in nature, but some are more common than others. For example, most plants are ''photolithoautotrophic'', since they use light as an energy source, water as electron donor, and {{CO2}} as a carbon source. All animals and fungi are [[chemo-organo-heterotrophic|''chemoorganoheterotrophic'']], since they use organic substances both as chemical energy sources and as electron/hydrogen donors and carbon sources. Some [[Eukaryote|eukaryotic]] microorganisms, however, are not limited to just one nutritional mode. For example, some algae live photoautotrophically in the light, but shift to chemoorganoheterotrophy in the dark. Even higher plants retained their ability to respire heterotrophically on starch at night which had been synthesised phototrophically during the day.

[[Prokaryote]]s show a great diversity of [[microbial metabolism|nutritional categories]].<ref name="TangTangBlankenship 2011">{{cite journal |vauthors=Tang KH, Tang YJ, Blankenship RE |title=Carbon metabolic pathways in phototrophic bacteria and their broader evolutionary implications |journal=Front Microbiol |volume=2 |issue= |pages=165 |date=2011 |pmid=21866228 |pmc=3149686 |doi=10.3389/fmicb.2011.00165 |doi-access=free }}</ref> For example, [[cyanobacteria]] and many [[purple sulfur bacteria]] can be ''photolithoautotrophic'', using light for energy, {{H2O}} or sulfide as electron/hydrogen donors, and {{CO2}} as carbon source, whereas [[green non-sulfur bacteria]] can be ''photoorganoheterotrophic'', using organic molecules as both electron/hydrogen donors and carbon sources.<ref name="Morris 2019" /><ref name="TangTangBlankenship 2011" /> Many bacteria are ''chemoorganoheterotrophic'', using organic molecules as energy, electron/hydrogen and carbon sources.<ref name="Morris 2019" /> Some bacteria are limited to only one nutritional group, whereas others are facultative and switch from one mode to the other, depending on the nutrient sources available.<ref name="TangTangBlankenship 2011" /> [[Lithotroph|Sulfur-oxidizing]], [[Iron-oxidizing bacteria|iron]], and [[anammox]] bacteria as well as [[methanogen]]s are [[Lithoautotroph|''chemolithoautotrophs'']], using inorganic energy, electron, and carbon sources. ''[[Lithotroph#Lithoheterotrophs versus lithoautotrophs|Chemolithoheterotrophs]]'' are rare because heterotrophy implies the availability of organic substrates, which can also serve as easy electron sources, making lithotrophy unnecessary. ''Photoorganoautotrophs'' are uncommon since their organic source of electrons/hydrogens would provide an easy carbon source, resulting in heterotrophy.

[[Synthetic biology]] efforts enabled the transformation of the trophic mode of two [[Model organism|model microorganisms]] from heterotrophy to chemoorganoautotrophy:

* ''[[Escherichia coli]]'' was [[Genetic engineering|genetically engineered]] and then [[Adaptive laboratory evolution|evolved in the laboratory]] to use {{CO2}} as the sole carbon source while using the one-carbon molecule [[formate]] as the source of electrons.<ref name=Gleizer19/>
* The [[methylotroph]]ic ''[[Pichia pastoris]]'' yeast was genetically engineered to use {{CO2}} as the carbon source instead of [[methanol]], while the latter remained the source of electrons for the cells.<ref name=Gassler19/>

== See also ==
{{refbegin|2}}
* [[Autotrophic]]
*[[Chemosynthesis]]
* [[Chemotrophic]]
* [[Heterotrophic]]
* [[Lithotrophic]]
* [[Metabolism]]
* [[Mixotrophic]]
* [[Organotrophic]]
* [[Phototroph]]ic
{{refend}}

== Notes and references ==
{{Reflist|32em}}

{{Modelling ecosystems}}
{{Metabolism}}
{{Portal bar|Biology}}

<!-- light is not the electron donor, it stimulates the release of an electron from chlorophyll. H2O is the electron donor. It becomes oxidized in the oxygen-evolving complex and donates its electrons to chlorophyll in order to replenish its lost electron -->

{{DEFAULTSORT:Primary Nutritional Groups}}
[[Category:Trophic ecology]]
[[Category:Physiology]]