# Copiotroph

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{{Short description|Organism found in carbon-rich environments}}A '''copiotroph''' is an [organism](/source/organism) found in environments rich in [nutrient](/source/nutrient)s, particularly [carbon](/source/carbon). They are the opposite to [oligotroph](/source/oligotroph)s, which survive in much lower carbon concentrations.<ref name="Panikov1995">{{cite book|author=N.S. Panikov|title=Microbial Growth Kinetics|url=https://books.google.com/books?id=6Z8X0lOjY6kC&pg=PA82|date=31 March 1995|publisher=Springer Science & Business Media|isbn=978-0-412-56630-1|page=82}}</ref>

Copiotrophic organisms tend to grow in high [organic](/source/organic_matter) substrate conditions. For example, copiotrophic organisms grow in  [Sewage lagoons](/source/Sewage_treatment). They grow in organic substrate conditions up to 100x higher than oligotrophs. Due to this substrate concentration inclination, copiotrophs are often found in nutrient rich waters near coastlines or [estuaries](/source/Estuary).<ref name=":0">{{Cite journal |last1=Lauro |first1=Federico M. |last2=McDougald |first2=Diane |last3=Thomas |first3=Torsten |last4=Williams |first4=Timothy J. |last5=Egan |first5=Suhelen |last6=Rice |first6=Scott |last7=DeMaere |first7=Matthew Z. |last8=Ting |first8=Lily |last9=Ertan |first9=Haluk |last10=Johnson |first10=Justin |last11=Ferriera |first11=Steven |last12=Lapidus |first12=Alla |last13=Anderson |first13=Iain |last14=Kyrpides |first14=Nikos |last15=Munk |first15=A. Christine |date=2009-09-15 |title=The genomic basis of trophic strategy in marine bacteria |journal=Proceedings of the National Academy of Sciences |language=en |volume=106 |issue=37 |pages=15527–15533 |doi=10.1073/pnas.0903507106 |issn=0027-8424 |pmc=2739866 |pmid=19805210 |doi-access=free }}</ref>

== Classification and Identification ==

The [bacterial phyla](/source/bacterial_phyla) can be differentiated into [copiotrophic](/source/copiotrophic) or [oligotrophic](/source/oligotrophic) categories that correspond and structure the functions of soil bacterial communities.

== Interaction with other organisms ==

Copiotrophic relation between oligotrophic bacteria depends on the amount of concentration the soil has of C compounds. If the soil has large amounts of organic C, it would then favor the copiotrophic bacteria. 

== Ecology ==

Copiotrophic bacteria are a key component in the soil C cycle. It is most important during the period of the year when vegetation is [photosynthetically active](/source/photosynthetically_active) and exudes large amounts of simple C compounds like [sugar](/source/sugar), [amino acids](/source/amino_acids), and [organic acids](/source/organic_acids). Copiotrophic bacteria are also found within marine life.

== Lifestyle ==
Copiotrophs have a higher [Michaelis-Menten constant](/source/Michaelis%E2%80%93Menten_kinetics) than [oligotrophs](/source/Oligotroph).<ref name=":1">{{Cite journal |last=Ho |first=Adrian |date=23 January 2017 |title=Revisiting life strategy concepts in environmental microbial ecology |url=https://academic.oup.com/femsec/article/93/3/fix006/2937747 |access-date=2023-04-22 |journal=FEMS Microbiology Ecology|volume=93 |issue=3 |article-number=fix006 |doi=10.1093/femsec/fix006 |pmid=28115400 |hdl=20.500.11755/97637b47-779a-413c-8397-81f77393a479 |hdl-access=free }}</ref> This constant is directly correlated to environmental substrate preference.<ref name=":1" /> In these high resource environments, copiotrophs exhibit a "feast-and-famine" lifestyle.<ref name=":72">{{Cite journal |last=Soler-Bistue |first=Alfonso |date=2023-03-01 |title=The evolving copiotrophic/oligotrophic dichotomy: From Winogradsky to physiology and genomics |journal=Environmental Microbiology|volume=25 |issue=7 |pages=1232–1237 |doi=10.1111/1462-2920.16360 |pmid=36856667 |bibcode=2023EnvMi..25.1232S |s2cid=257256291 }}</ref> They utilize the available nutrients in the environment rapidly resulting in nutrient depletion which forces them to starve.<ref name=":72" /> This is possible through increasing their growth rate with nutrient uptake.<ref name=":8">{{Cite journal |last1=Noell |first1=Stephen E. |last2=Brennan |first2=Elizabeth |last3=Washburn |first3=Quinn |last4=Davis |first4=Edward W. |last5=Hellweger |first5=Ferdi L. |last6=Giovannoni |first6=Stephen J. |date=2023-01-15 |title=Differences in the regulatory strategies of marine oligotrophs and copiotrophs reflect differences in motility |journal=Environmental Microbiology |volume=25 |issue=7 |language=en |pages=1265–1280 |doi=10.1111/1462-2920.16357|biorxiv=10.1101/2022.07.21.501054|pmid=36826469 |s2cid=251021339 |doi-access=free|bibcode=2023EnvMi..25.1265N }}</ref> However, when nutrients in the environment get depleted, copiotrophs struggle to survive for long periods of time.<ref name=":2">{{Cite journal |last=Koch |first=Arthur L. |date=2001-07-12 |title=Oligotrophs versus copiotrophs |url=https://onlinelibrary.wiley.com/doi/10.1002/bies.1091 |journal=BioEssays |language=en |volume=23 |issue=7 |pages=657–661 |doi=10.1002/bies.1091 |pmid=11462219 |s2cid=39126203 |issn=0265-9247|url-access=subscription }}</ref>  Copiotrophs do not have the ability to respond to starvation.<ref name=":2" /> It is hypothesized that this may be a lost trait.<ref name=":2" /> Another possibility is that microbes never evolved to survive these extreme conditions.<ref name=":2" /> Oligotrophs can outcompete copiotrophs in low-nutrient environments.<ref name=":2" /> This causes low-nutrient conditions to continue for extended periods of time, making it difficult for copiotrophs to sustain life.<ref name=":2" /> Copiotrophs are larger than oligotrophs and need more energy, requiring larger concentrations of substrate for survival.<ref name=":2" />

Copiotrophs are motile.<ref name=":0" /> Copiotrophs can have external organelles such as flagella that extend out of a microbe's cell to facilitate movement.<ref name=":8" /> Copiotrophs are also [chemotactic](/source/Chemotaxis), meaning they can detect nutrients in the environment.<ref name=":3">{{Cite book |last=Kirchman |first=David |title=Processes In Microbial Ecology |publisher=Oxford University Press Inc. |year=2012 |isbn=978-0-19-958693-6 |edition=1st |location=New York |pages=46–48}}</ref> These help the microbes travel quickly to nearby food sources.<ref name=":3" /> Chemotaxis also enables the organism to travel away from a restricting compound.<ref name=":3" /> There are multiple methods for chemotaxis in these organisms.<ref name=":3" /> This includes the "run and tumble" strategy in which the organism randomly picks a direction to move in.<ref name=":3" /> However, if it senses that the concentration gradient is decreasing they stop and choose another random direction to travel in.<ref name=":3" />Another strategy includes the "run and reverse" in which the organism runs towards a nutrient.<ref name=":3" /> If it notices the gradient decreasing, it moves back to where the gradient is larger and heads in another direction from this new position.<ref name=":3" />

Through their motility and chemotaxis, copiotrophic microbes respond quickly to nutrients in their environment.<ref name=":0" /> With the help of these mechanisms, copiotrophs can travel to and stay in nutrient dense areas long enough for transcriptional regulatory systems to increase gene expression.<ref name=":8" /> This in turn helps them increase metabolic processes in high nutrient areas allowing them to maximize their growth during these patches.<ref name=":8" />

== Growth characteristics ==
Copiotrophs are characterized by a high maximum growth rate.<ref name=":4">{{Cite journal |last1=Roller |first1=Benjamin R. K. |last2=Stoddard |first2=Steven F. |last3=Schmidt |first3=Thomas M. |date=2016-09-12 |title=Exploiting rRNA operon copy number to investigate bacterial reproductive strategies |journal=Nature Microbiology |language=en |volume=1 |issue=11 |page=16160 |doi=10.1038/nmicrobiol.2016.160 |pmid=27617693 |pmc=5061577 |issn=2058-5276}}</ref> This high growth rate allows for copiotrophs to have a larger genome and cell size than their oligotrophic counterparts.<ref name=":0" />

The copiotrophic genome encompasses more ribosomal RNA operons than the oligotrophic genome.<ref name=":4" /> Ribosomal RNA operons are linearly related to growth rate.<ref name=":3" /><ref name=":4" /> The ribosomal RNA operons are responsible for expression of genes in clusters.<ref name=":5">{{Cite journal |last1=Espejo |first1=Romilio T. |last2=Plaza |first2=Nicolás |date=2018 |title=Multiple Ribosomal RNA Operons in Bacteria; Their Concerted Evolution and Potential Consequences on the Rate of Evolution of Their 16S rRNA |journal=Frontiers in Microbiology |volume=9 |page=1232 |doi=10.3389/fmicb.2018.01232 |pmid=29937760 |pmc=6002687 |issn=1664-302X |doi-access=free }}</ref> The larger amount of ribosomal content allows for more rapid growth.<ref name=":4" /> Oligotrophs have one ribosomal RNA operon while copiotrophs can contain up to fifteen operons.<ref name=":5" />

Copiotrophs tend to have a lower carbon use efficiency than oligotrophs.<ref name=":6">{{Cite journal |last1=Roller |first1=Benjamin RK |last2=Schmidt |first2=Thomas M. |date=2015-01-15 |title=The physiology and ecological implications of efficient growth |journal=The ISME Journal |language=en |volume=9 |issue=7 |pages=1481–1487 |doi=10.1038/ismej.2014.235 |pmid=25575305 |pmc=4478692 |bibcode=2015ISMEJ...9.1481R |issn=1751-7370}}</ref> This is the ratio of carbon used for production of biomass per total carbon consumed by the organism.<ref name=":6" /> Carbon use efficiency can be used to understand organisms lifestyles, whether they primarily create biomass or require carbon for maintenance energy.<ref name=":6" /><ref>{{Cite journal |last1=Pold |first1=Grace |last2=Domeignoz-Horta |first2=Luiz A. |last3=Morrison |first3=Eric W. |last4=Frey |first4=Serita D. |last5=Sistla |first5=Seeta A. |last6=DeAngelis |first6=Kristen M. |date=2020-02-25 |editor-last=Giovannoni |editor-first=Stephen J. |title=Carbon Use Efficiency and Its Temperature Sensitivity Covary in Soil Bacteria |journal=mBio |language=en |volume=11 |issue=1 |pages=e02293–19 |doi=10.1128/mBio.02293-19  |doi-access=free|issn=2161-2129 |pmc=6974560 |pmid=31964725}}</ref>  Energy is necessary for the copiotrophic lifestyle which includes motility and chemotaxis.<ref name=":9">{{Cite journal |last1=Geyer |first1=Kevin M. |last2=Kyker-Snowman |first2=Emily |last3=Grandy |first3=A. Stuart |last4=Frey |first4=Serita D. |date=2016-02-01 |title=Microbial carbon use efficiency: accounting for population, community, and ecosystem-scale controls over the fate of metabolized organic matter |journal=Biogeochemistry |language=en |volume=127 |issue=2 |pages=173–188 |doi=10.1007/s10533-016-0191-y |s2cid=54772410 |issn=1573-515X|doi-access=free |bibcode=2016Biogc.127..173G }}</ref> This energy could otherwise be used for biomass production.<ref name=":9" /> This results in a lower efficiency than the oligotrophic lifestyle which primarily uses energy for the creation of biomass.<ref name=":9" />

Copiotrophs have a lower protein yield than oligotrophs.<ref name=":4" /> Protein yield is the amount of protein synthesized per O<sub>2</sub> consumed.<ref name=":4" /> This is also associated with the higher ribosomal RNA operons.<ref name=":4" /> Overall, copiotrophs create more protein than their oligotrophic peers, however due to the copiotrophs' lower carbon use efficiency, less protein is produced per gram O<sub>2</sub> consumed by the organisms.<ref name=":4" />

== References ==
{{reflist}}
Fierer, N., Bradford, M. A., & Jackson, R. B. (2007). Toward an ecological classification of soil bacteria. Ecology, 88(6), 1354-1364. 
Ivars-Martinez, E., Martin-Cuadrado, A. B., D'auria, G., Mira, A., Ferriera, S., Johnson, J., ... & Rodriguez-Valera, F. (2008). Comparative genomics of two ecotypes of the marine planktonic copiotroph Alteromonas macleodii suggests alternative lifestyles associated with different kinds of particulate organic matter. The ISME journal, 2(12), 1194-1212.
Lladó, S., & Baldrian, P. (2017). Community-level physiological profiling analyses show potential to identify the copiotrophic bacteria present in soil environments. PLoS One, 12(2), e0171638.

{{modelling ecosystems}}

Category:Organisms by adaptation
Category:Trophic ecology

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Adapted from the Wikipedia article [Copiotroph](https://en.wikipedia.org/wiki/Copiotroph) by Wikipedia contributors ([contributor history](https://en.wikipedia.org/wiki/Copiotroph?action=history)). Available under [Creative Commons Attribution-ShareAlike 4.0 International](https://creativecommons.org/licenses/by-sa/4.0/). Changes may have been made.