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[[File:Cluster of PAOs.jpg|thumb|Cluster of PAOs]]
[[File:Cluster of PAOs.jpg|thumb|Cluster of PAOs]]


'''Polyphosphate-accumulating organisms''' (PAOs) are a group of [[microorganism]]s that, under certain conditions, facilitate the removal of large amounts of [[phosphorus]] from their environments. The most studied example of this phenomenon is in bacteria found in a form of wastewater processing known as [[enhanced biological phosphorus removal]] (EBPR), however phosphate hyperaccumulation has been found to occur in other conditions such as soil and marine environments, as well as in non-bacterial organisms such as fungi and algae.<ref>{{Cite journal |last=Akbari |first=Ali |last2=Wang |first2=ZiJian |last3=He |first3=Peisheng |last4=Wang |first4=Dongqi |last5=Lee |first5=Jangho |last6=Han |first6=Il |last7=Li |first7=Guangyu |last8=Gu |first8=April Z. |date=January 2021 |title=Unrevealed roles of polyphosphate‐accumulating microorganisms |url=https://onlinelibrary.wiley.com/doi/10.1111/1751-7915.13730 |journal=Microbial Biotechnology |language=en |volume=14 |issue=1 |pages=82–87 |doi=10.1111/1751-7915.13730 |issn=1751-7915 |pmc=7888455 |pmid=33404187}}</ref> PAOs accomplish this removal of phosphate by accumulating it within their cells as [[polyphosphate]]. PAOs are by no means the only microbes that can accumulate phosphate within their cells and in fact, the production of polyphosphate is a widespread ability among microbes. However, PAOs have many characteristics that other organisms that accumulate polyphosphate do not have that make them amenable to use in [[wastewater treatment]]. Specifically, in the case of classical PAOs, is the ability to consume simple carbon compounds (energy source) without the presence of an external electron acceptor (such as nitrate or oxygen) by generating energy from internally stored polyphosphate and glycogen. Most other bacteria cannot consume under these conditions and therefore PAOs gain a selective advantage within the mixed microbial community present in the activated sludge. Therefore, wastewater treatment plants that operate for enhanced biological phosphorus removal have an anaerobic tank (where there is no nitrate or oxygen present as external electron acceptor) prior to the other tanks to give PAOs preferential access to the simple carbon compounds in the wastewater that is influent to the plant.
'''Polyphosphate-accumulating organisms''' (PAOs) are a group of [[microorganism]]s that, under certain conditions, facilitate the removal of large amounts of [[phosphorus]] from their environments. The most studied example of this phenomenon is in bacteria found in a form of wastewater processing known as [[enhanced biological phosphorus removal]] (EBPR), however phosphate hyperaccumulation has been found to occur in other conditions such as soil and marine environments, as well as in non-bacterial organisms such as fungi and algae.<ref>{{Cite journal |last1=Akbari |first1=Ali |last2=Wang |first2=ZiJian |last3=He |first3=Peisheng |last4=Wang |first4=Dongqi |last5=Lee |first5=Jangho |last6=Han |first6=Il |last7=Li |first7=Guangyu |last8=Gu |first8=April Z. |date=January 2021 |title=Unrevealed roles of polyphosphate‐accumulating microorganisms |journal=Microbial Biotechnology |language=en |volume=14 |issue=1 |pages=82–87 |doi=10.1111/1751-7915.13730 |issn=1751-7915 |pmc=7888455 |pmid=33404187}}</ref> PAOs accomplish this removal of phosphate by accumulating it within their cells as [[polyphosphate]]. PAOs are by no means the only microbes that can accumulate phosphate within their cells and in fact, the production of polyphosphate is a widespread ability among microbes. However, PAOs have many characteristics that other organisms that accumulate polyphosphate do not have that make them amenable to use in [[wastewater treatment]]. Specifically, in the case of classical PAOs, is the ability to consume simple carbon compounds (energy source) without the presence of an external electron acceptor (such as nitrate or oxygen) by generating energy from internally stored polyphosphate and glycogen. Most other bacteria cannot consume under these conditions and therefore PAOs gain a selective advantage within the mixed microbial community present in the activated sludge. Therefore, wastewater treatment plants that operate for enhanced biological phosphorus removal have an anaerobic tank (where there is no nitrate or oxygen present as external electron acceptor) prior to the other tanks to give PAOs preferential access to the simple carbon compounds in the wastewater that is influent to the plant.


== Metabolisms ==
== Metabolisms ==


=== Classical (Canonical) PAO Metabolism ===
=== Classical (Canonical) PAO Metabolism ===
The classical or "canonical" behavior of PAOs is considered to be the release of phosphate (as [[orthophosphate]]) to the environment and transformation of intracellular polyphosphate reserves into [[polyhydroxyalkanoates]] (PHA) from [[volatile fatty acids]] (VFAs) and glycogen during anoxic conditions.<ref name=":0">{{Citation |last=Akram |first=Fatima |title=Chapter 8 - Role of polyphosphate accumulating organisms in enhanced biological phosphorous removal |date=2022-01-01 |url=https://www.sciencedirect.com/science/article/pii/B9780323918930000031 |work=Microbial Consortium and Biotransformation for Pollution Decontamination |pages=151–179 |editor-last=Dar |editor-first=Gowhar Hamid |access-date=2023-07-07 |series=Advances in Environmental Pollution Research |publisher=Elsevier |language=en |isbn=978-0-323-91893-0 |last2=Aqeel |first2=Amna |last3=Ahmed |first3=Zeeshan |last4=Zafar |first4=Javeria |last5=Haq |first5=Ikram ul |editor2-last=Bhat |editor2-first=Rouf Ahmad |editor3-last=Qadri |editor3-first=Humaira |editor4-last=Hakeem |editor4-first=Khalid Rehman}}</ref> This is followed by the consumption of the PHA/VFAs and uptake of environmental orthophosphate during oxic conditions to regenerate polyphosphate reserves within the cell.<ref name=":0" />
The classical or "canonical" behavior of PAOs is considered to be the release of phosphate (as [[orthophosphate]]) to the environment and transformation of intracellular polyphosphate reserves into [[polyhydroxyalkanoates]] (PHA) from [[volatile fatty acids]] (VFAs) and glycogen during anoxic conditions.<ref name=":0">{{Citation |last1=Akram |first1=Fatima |title=Chapter 8 - Role of polyphosphate accumulating organisms in enhanced biological phosphorous removal |date=2022-01-01 |url=https://www.sciencedirect.com/science/article/pii/B9780323918930000031 |work=Microbial Consortium and Biotransformation for Pollution Decontamination |pages=151–179 |editor-last=Dar |editor-first=Gowhar Hamid |access-date=2023-07-07 |series=Advances in Environmental Pollution Research |publisher=Elsevier |language=en |isbn=978-0-323-91893-0 |last2=Aqeel |first2=Amna |last3=Ahmed |first3=Zeeshan |last4=Zafar |first4=Javeria |last5=Haq |first5=Ikram ul |editor2-last=Bhat |editor2-first=Rouf Ahmad |editor3-last=Qadri |editor3-first=Humaira |editor4-last=Hakeem |editor4-first=Khalid Rehman}}</ref> This is followed by the consumption of the PHA/VFAs and uptake of environmental orthophosphate during oxic conditions to regenerate polyphosphate reserves within the cell.<ref name=":0" />


=== Non-Canonical (or "Fermentative) PAO Metabolism ===
=== Non-Canonical (or "Fermentative) PAO Metabolism ===
Some PAOs have been found to have alternative methods to accumulating polyphosphate, particularly to do with not storing PHA or glycogen.<ref name=":12">{{Cite journal |last=Singleton |first=C. M. |last2=Petriglieri |first2=F. |last3=Wasmund |first3=K. |last4=Nierychlo |first4=M. |last5=Kondrotaite |first5=Z. |last6=Petersen |first6=J. F. |last7=Peces |first7=M. |last8=Dueholm |first8=M. S. |last9=Wagner |first9=M. |last10=Nielsen |first10=P. H. |date=June 2022 |title=The novel genus, 'Candidatus Phosphoribacter', previously identified as Tetrasphaera, is the dominant polyphosphate accumulating lineage in EBPR wastewater treatment plants worldwide |url=https://pubmed.ncbi.nlm.nih.gov/35217776/ |journal=The ISME journal |volume=16 |issue=6 |pages=1605–1616 |doi=10.1038/s41396-022-01212-z |issn=1751-7370 |pmc=9123174 |pmid=35217776}}</ref><ref name=":23">{{Cite journal |last=Nielsen |first=Per Halkjær |last2=McIlroy |first2=Simon J. |last3=Albertsen |first3=Mads |last4=Nierychlo |first4=Marta |date=June 2019 |title=Re-evaluating the microbiology of the enhanced biological phosphorus removal process |url=https://pubmed.ncbi.nlm.nih.gov/30959426/ |journal=Current Opinion in Biotechnology |volume=57 |pages=111–118 |doi=10.1016/j.copbio.2019.03.008 |issn=1879-0429 |pmid=30959426}}</ref> This is generally believed to be seen more often in extracellular environments high in organic compounds, thus containing fermentable substrates like amino acids and sugars.<ref>{{Cite journal |last=Nguyen |first=Hien Thi Thu |last2=Kristiansen |first2=Rikke |last3=Vestergaard |first3=Mette |last4=Wimmer |first4=Reinhard |last5=Nielsen |first5=Per Halkjær |date=2015-07-15 |title=Intracellular Accumulation of Glycine in Polyphosphate-Accumulating Organisms in Activated Sludge, a Novel Storage Mechanism under Dynamic Anaerobic-Aerobic Conditions |url=http://dx.doi.org/10.1128/aem.01012-15 |journal=Applied and Environmental Microbiology |volume=81 |issue=14 |pages=4809–4818 |doi=10.1128/aem.01012-15 |issn=0099-2240}}</ref> However, the exact mechanisms of these microbes to accumulate and use polyphosphate are not well understood.<ref name=":23"/>
Some PAOs have been found to have alternative methods to accumulating polyphosphate, particularly to do with not storing PHA or glycogen.<ref name=":12">{{Cite journal |last1=Singleton |first1=C. M. |last2=Petriglieri |first2=F. |last3=Wasmund |first3=K. |last4=Nierychlo |first4=M. |last5=Kondrotaite |first5=Z. |last6=Petersen |first6=J. F. |last7=Peces |first7=M. |last8=Dueholm |first8=M. S. |last9=Wagner |first9=M. |last10=Nielsen |first10=P. H. |date=June 2022 |title=The novel genus, 'Candidatus Phosphoribacter', previously identified as Tetrasphaera, is the dominant polyphosphate accumulating lineage in EBPR wastewater treatment plants worldwide |journal=The ISME Journal |volume=16 |issue=6 |pages=1605–1616 |doi=10.1038/s41396-022-01212-z |issn=1751-7370 |pmc=9123174 |pmid=35217776}}</ref><ref name=":23">{{Cite journal |last1=Nielsen |first1=Per Halkjær |last2=McIlroy |first2=Simon J. |last3=Albertsen |first3=Mads |last4=Nierychlo |first4=Marta |date=June 2019 |title=Re-evaluating the microbiology of the enhanced biological phosphorus removal process |url=https://pubmed.ncbi.nlm.nih.gov/30959426/ |journal=Current Opinion in Biotechnology |volume=57 |pages=111–118 |doi=10.1016/j.copbio.2019.03.008 |issn=1879-0429 |pmid=30959426|s2cid=104294644 }}</ref> This is generally believed to be seen more often in extracellular environments high in organic compounds, thus containing fermentable substrates like amino acids and sugars.<ref>{{Cite journal |last1=Nguyen |first1=Hien Thi Thu |last2=Kristiansen |first2=Rikke |last3=Vestergaard |first3=Mette |last4=Wimmer |first4=Reinhard |last5=Nielsen |first5=Per Halkjær |date=2015-07-15 |title=Intracellular Accumulation of Glycine in Polyphosphate-Accumulating Organisms in Activated Sludge, a Novel Storage Mechanism under Dynamic Anaerobic-Aerobic Conditions |url=http://dx.doi.org/10.1128/aem.01012-15 |journal=Applied and Environmental Microbiology |volume=81 |issue=14 |pages=4809–4818 |doi=10.1128/aem.01012-15 |pmid=25956769 |pmc=4551194 |issn=0099-2240}}</ref> However, the exact mechanisms of these microbes to accumulate and use polyphosphate are not well understood.<ref name=":23"/>


== Known Bacterial PAOs ==
== Known Bacterial PAOs ==


=== ''Candidatus'' ''Phosphoribacter'' (previously referred to as ''[[Tetrasphaera]]'' prior to 2022) ===
=== ''Candidatus'' ''Phosphoribacter'' (previously referred to as ''[[Tetrasphaera]]'' prior to 2022) ===
''Candidatus'' ''Phosphoribacter'' is a bacterial genus that has been found to be the dominant PAO associated with wastewater treatment worldwide, and has been found to often participate more in the biological removal of phosphorus than ''Candidatus'' Accumulibacter, contrary to previous understandings.<ref name=":12"/><ref name=":4"/><ref name=":23"/> This bacteria has been found to be a non-canonical (or fermentative/"fPAO") PAO, and universally lack the genetic potential to store PHA.<ref name=":12" /><ref name=":3">{{Cite journal |last=Otieno |first=Jeremiah |last2=Kowal |first2=Przemysław |last3=Mąkinia |first3=Jacek |date=2022-10-28 |title=The Occurrence and Role of Tetrasphaera in Enhanced Biological Phosphorus Removal Systems |url=http://dx.doi.org/10.3390/w14213428 |journal=Water |volume=14 |issue=21 |pages=3428 |doi=10.3390/w14213428 |issn=2073-4441}}</ref> This genus was largely found to be capable of producing the fermentation products acetate, lactate, alanine, and succinate.<ref name=":3" /><ref>{{Cite journal |last=Kristiansen |first=Rikke |last2=Nguyen |first2=Hien Thi Thu |last3=Saunders |first3=Aaron Marc |last4=Nielsen |first4=Jeppe Lund |last5=Wimmer |first5=Reinhard |last6=Le |first6=Vang Quy |last7=McIlroy |first7=Simon Jon |last8=Petrovski |first8=Steve |last9=Seviour |first9=Robert J. |last10=Calteau |first10=Alexandra |last11=Nielsen |first11=Kåre Lehmann |last12=Nielsen |first12=Per Halkjær |date=March 2013 |title=A metabolic model for members of the genus Tetrasphaera involved in enhanced biological phosphorus removal |url=https://pubmed.ncbi.nlm.nih.gov/23178666/ |journal=The ISME journal |volume=7 |issue=3 |pages=543–554 |doi=10.1038/ismej.2012.136 |issn=1751-7370 |pmc=3578573 |pmid=23178666}}</ref> Additionally, it is suggested that the [[amino acid]]s lysine, arginine, histidine, leucine, isoleucine, valine and phenylalanine may replace the canonical purpose of PHA as an energy substrate during oxic conditions, based on genomic potential and similarity to behavior of other microbial metabolisms.<ref name=":12" /> Alternatively, the compound [[cyanophycin]] may used as an energy substrate due to the ubiquity of cyanophycin-metabolizing enzymes encoded in the species.<ref name=":12" />
''Candidatus'' ''Phosphoribacter'' is a bacterial genus that has been found to be the dominant PAO associated with wastewater treatment worldwide, and has been found to often participate more in the biological removal of phosphorus than ''Candidatus'' Accumulibacter, contrary to previous understandings.<ref name=":12"/><ref name=":4"/><ref name=":23"/> This bacteria has been found to be a non-canonical (or fermentative/"fPAO") PAO, and universally lack the genetic potential to store PHA.<ref name=":12" /><ref name=":3">{{Cite journal |last1=Otieno |first1=Jeremiah |last2=Kowal |first2=Przemysław |last3=Mąkinia |first3=Jacek |date=2022-10-28 |title=The Occurrence and Role of Tetrasphaera in Enhanced Biological Phosphorus Removal Systems |journal=Water |volume=14 |issue=21 |pages=3428 |doi=10.3390/w14213428 |issn=2073-4441 |doi-access=free }}</ref> This genus was largely found to be capable of producing the fermentation products acetate, lactate, alanine, and succinate.<ref name=":3" /><ref>{{Cite journal |last1=Kristiansen |first1=Rikke |last2=Nguyen |first2=Hien Thi Thu |last3=Saunders |first3=Aaron Marc |last4=Nielsen |first4=Jeppe Lund |last5=Wimmer |first5=Reinhard |last6=Le |first6=Vang Quy |last7=McIlroy |first7=Simon Jon |last8=Petrovski |first8=Steve |last9=Seviour |first9=Robert J. |last10=Calteau |first10=Alexandra |last11=Nielsen |first11=Kåre Lehmann |last12=Nielsen |first12=Per Halkjær |date=March 2013 |title=A metabolic model for members of the genus Tetrasphaera involved in enhanced biological phosphorus removal |journal=The ISME Journal |volume=7 |issue=3 |pages=543–554 |doi=10.1038/ismej.2012.136 |issn=1751-7370 |pmc=3578573 |pmid=23178666}}</ref> Additionally, it is suggested that the [[amino acid]]s lysine, arginine, histidine, leucine, isoleucine, valine and phenylalanine may replace the canonical purpose of PHA as an energy substrate during oxic conditions, based on genomic potential and similarity to behavior of other microbial metabolisms.<ref name=":12" /> Alternatively, the compound [[cyanophycin]] may used as an energy substrate due to the ubiquity of cyanophycin-metabolizing enzymes encoded in the species.<ref name=":12" />


=== ''Candidatus'' ''Accumlibacter phosphatis'' ===
=== ''Candidatus'' ''Accumlibacter phosphatis'' ===
[[Candidatus Accumulibacter phosphatis|''Candidatus'' ''Accumulibacter phosphatis'']] is one of the most well-studied PAOs, and is responsible for the development of the classical PAO metabolic model which ''Ca.'' ''Phosphoribacter'' later contradicted.<ref>{{Cite journal |last=He |first=Shaomei |last2=McMahon |first2=Katherine D. |date=2011-02-21 |title=Microbiology of ‘''Candidatus''Accumulibacter’ in activated sludge |url=http://dx.doi.org/10.1111/j.1751-7915.2011.00248.x |journal=Microbial Biotechnology |volume=4 |issue=5 |pages=603–619 |doi=10.1111/j.1751-7915.2011.00248.x |issn=1751-7915}}</ref> Formerly considered the most important PAO in waste treatment, the bacteria is highly abundant in wastewater treatment plants globally.<ref name=":23"/><ref>{{Cite journal |last=Onnis‐Hayden |first=Annalisa |last2=Srinivasan |first2=Varun |last3=Tooker |first3=Nicholas B. |last4=Li |first4=Guangyu |last5=Wang |first5=Dongqi |last6=Barnard |first6=James L. |last7=Bott |first7=Charles |last8=Dombrowski |first8=Paul |last9=Schauer |first9=Peter |last10=Menniti |first10=Adrienne |last11=Shaw |first11=Andrew |last12=Stinson |first12=Beverly |last13=Stevens |first13=Gerry |last14=Dunlap |first14=Patrick |last15=Takács |first15=Imre |date=March 2020 |title=Survey of full‐scale sidestream enhanced biological phosphorus removal (S2EBPR) systems and comparison with conventional EBPRs in North America: Process stability, kinetics, and microbial populations |url=https://onlinelibrary.wiley.com/doi/10.1002/wer.1198 |journal=Water Environment Research |language=en |volume=92 |issue=3 |pages=403–417 |doi=10.1002/wer.1198 |issn=1061-4303}}</ref> It can consume a range of carbon compounds, such as acetate and propionate, under anaerobic conditions and store these compounds as [[polyhydroxyalkanoates]] (PHA) which it consumes as a carbon and energy source for growth using oxygen or nitrate as electron acceptor. Historically, the hyperaccumulation of phosphate by ''Ca''. ''Accumulibacter'' was seen as a stress response, but currently it is suggested that this behavior may play an ecological role.<ref>{{Cite web |last=da Silva |first=Leonor Guedes |last2=Gamez |first2=Karel Olavarria |last3=Gomes |first3=Joana Castro |last4=Akkermans |first4=Kasper |last5=Welles |first5=Laurens |last6=Abbas |first6=Ben |last7=van Loosdrecht |first7=Mark C.M. |last8=Wahl |first8=Sebastian Aljoscha |date=2018-11-01 |title=Revealing metabolic flexibility of''Candidatus''Accumulibacter phosphatis through redox cofactor analysis and metabolic network modeling |url=http://dx.doi.org/10.1101/458331 |access-date=2023-07-07 |website=dx.doi.org}}</ref> In combination with Ca. ''Phosphoribacter'', these two PAOs are considered to account for 24-70% of phosphorus removed from wastewater during treatment processing.<ref name=":4">{{Cite journal |last=Fernando |first=Eustace Y. |last2=McIlroy |first2=Simon Jon |last3=Nierychlo |first3=Marta |last4=Herbst |first4=Florian-Alexander |last5=Petriglieri |first5=Francesca |last6=Schmid |first6=Markus C. |last7=Wagner |first7=Michael |last8=Nielsen |first8=Jeppe Lund |last9=Nielsen |first9=Per Halkjær |date=August 2019 |title=Resolving the individual contribution of key microbial populations to enhanced biological phosphorus removal with Raman-FISH |url=https://pubmed.ncbi.nlm.nih.gov/30894691/ |journal=The ISME journal |volume=13 |issue=8 |pages=1933–1946 |doi=10.1038/s41396-019-0399-7 |issn=1751-7370 |pmc=6776032 |pmid=30894691}}</ref>
[[Candidatus Accumulibacter phosphatis|''Candidatus'' ''Accumulibacter phosphatis'']] is one of the most well-studied PAOs, and is responsible for the development of the classical PAO metabolic model which ''Ca.'' ''Phosphoribacter'' later contradicted.<ref>{{Cite journal |last1=He |first1=Shaomei |last2=McMahon |first2=Katherine D. |date=2011-02-21 |title=Microbiology of '''Candidatus''Accumulibacter' in activated sludge |url=http://dx.doi.org/10.1111/j.1751-7915.2011.00248.x |journal=Microbial Biotechnology |volume=4 |issue=5 |pages=603–619 |doi=10.1111/j.1751-7915.2011.00248.x |pmid=21338476 |pmc=3819010 |issn=1751-7915}}</ref> Formerly considered the most important PAO in waste treatment, the bacteria is highly abundant in wastewater treatment plants globally.<ref name=":23"/><ref>{{Cite journal |last1=Onnis‐Hayden |first1=Annalisa |last2=Srinivasan |first2=Varun |last3=Tooker |first3=Nicholas B. |last4=Li |first4=Guangyu |last5=Wang |first5=Dongqi |last6=Barnard |first6=James L. |last7=Bott |first7=Charles |last8=Dombrowski |first8=Paul |last9=Schauer |first9=Peter |last10=Menniti |first10=Adrienne |last11=Shaw |first11=Andrew |last12=Stinson |first12=Beverly |last13=Stevens |first13=Gerry |last14=Dunlap |first14=Patrick |last15=Takács |first15=Imre |date=March 2020 |title=Survey of full‐scale sidestream enhanced biological phosphorus removal (S2EBPR) systems and comparison with conventional EBPRs in North America: Process stability, kinetics, and microbial populations |url=https://onlinelibrary.wiley.com/doi/10.1002/wer.1198 |journal=Water Environment Research |language=en |volume=92 |issue=3 |pages=403–417 |doi=10.1002/wer.1198 |pmid=31402530 |s2cid=199539909 |issn=1061-4303}}</ref> It can consume a range of carbon compounds, such as acetate and propionate, under anaerobic conditions and store these compounds as [[polyhydroxyalkanoates]] (PHA) which it consumes as a carbon and energy source for growth using oxygen or nitrate as electron acceptor. Historically, the hyperaccumulation of phosphate by ''Ca''. ''Accumulibacter'' was seen as a stress response, but currently it is suggested that this behavior may play an ecological role.<ref>{{Cite journal |last1=da Silva |first1=Leonor Guedes |last2=Gamez |first2=Karel Olavarria |last3=Gomes |first3=Joana Castro |last4=Akkermans |first4=Kasper |last5=Welles |first5=Laurens |last6=Abbas |first6=Ben |last7=van Loosdrecht |first7=Mark C.M. |last8=Wahl |first8=Sebastian Aljoscha |date=2018-11-01 |title=Revealing metabolic flexibility of''Candidatus''Accumulibacter phosphatis through redox cofactor analysis and metabolic network modeling |url=http://dx.doi.org/10.1101/458331 |access-date=2023-07-07 |website=dx.doi.org|doi=10.1101/458331 |s2cid=91862227 }}</ref> In combination with Ca. ''Phosphoribacter'', these two PAOs are considered to account for 24-70% of phosphorus removed from wastewater during treatment processing.<ref name=":4">{{Cite journal |last1=Fernando |first1=Eustace Y. |last2=McIlroy |first2=Simon Jon |last3=Nierychlo |first3=Marta |last4=Herbst |first4=Florian-Alexander |last5=Petriglieri |first5=Francesca |last6=Schmid |first6=Markus C. |last7=Wagner |first7=Michael |last8=Nielsen |first8=Jeppe Lund |last9=Nielsen |first9=Per Halkjær |date=August 2019 |title=Resolving the individual contribution of key microbial populations to enhanced biological phosphorus removal with Raman-FISH |journal=The ISME Journal |volume=13 |issue=8 |pages=1933–1946 |doi=10.1038/s41396-019-0399-7 |issn=1751-7370 |pmc=6776032 |pmid=30894691}}</ref>


=== ''Candidatus'' ''dechloromonas'' ===
=== ''Candidatus'' ''dechloromonas'' ===
[[Dechloromonas|''Candidatus'' ''Dechloromonas'']] species phosphoritropha and phosphorivorans are PAOs with classical metabolism genotype.<ref>{{Cite web |title=Midas Field Guide |url=https://www.midasfieldguide.org/guide/fieldguide/genus/dechloromonas |access-date=2023-07-07 |website=www.midasfieldguide.org}}</ref> ''Dechloromonas'' has been found in high abundances in wastewater treatment plants across the world.<ref>{{Cite journal |last=Terashima |first=Mia |last2=Yama |first2=Ayano |last3=Sato |first3=Megumi |last4=Yumoto |first4=Isao |last5=Kamagata |first5=Yoichi |last6=Kato |first6=Souichiro |date=2016 |title=Culture-Dependent and -Independent Identification of Polyphosphate-Accumulating &lt;i&gt;Dechloromonas&lt;/i&gt; spp. Predominating in a Full-Scale Oxidation Ditch Wastewater Treatment Plant |url=http://dx.doi.org/10.1264/jsme2.me16097 |journal=Microbes and Environments |volume=31 |issue=4 |pages=449–455 |doi=10.1264/jsme2.me16097 |issn=1342-6311}}</ref><ref>{{Cite journal |last=Wang |first=Baogui |last2=Jiao |first2=Erlong |last3=Guo |first3=Yu |last4=Zhang |first4=Lifang |last5=Meng |first5=Qingan |last6=Zeng |first6=Wei |last7=Peng |first7=Yongzhen |date=2020-07-02 |title=Investigation of the polyphosphate-accumulating organism population in the full-scale simultaneous chemical phosphorus removal system |url=http://dx.doi.org/10.1007/s11356-020-09912-9 |journal=Environmental Science and Pollution Research |volume=27 |issue=30 |pages=37877–37886 |doi=10.1007/s11356-020-09912-9 |issn=0944-1344}}</ref><ref>{{Cite journal |last=Stokholm-Bjerregaard |first=Mikkel |last2=McIlroy |first2=Simon J. |last3=Nierychlo |first3=Marta |last4=Karst |first4=Søren M. |last5=Albertsen |first5=Mads |last6=Nielsen |first6=Per H. |date=2017-04-27 |title=A Critical Assessment of the Microorganisms Proposed to be Important to Enhanced Biological Phosphorus Removal in Full-Scale Wastewater Treatment Systems |url=http://dx.doi.org/10.3389/fmicb.2017.00718 |journal=Frontiers in Microbiology |volume=8 |doi=10.3389/fmicb.2017.00718 |issn=1664-302X}}</ref><ref name=":23" /> The two species described here, ''Dechloromonas'' phosphoritropha and phosphorivans, are the two most abundant species in waste treatment within the genus.<ref>{{Cite journal |last=Petriglieri |first=Francesca |last2=Singleton |first2=Caitlin |last3=Peces |first3=Miriam |last4=Petersen |first4=Jette F. |last5=Nierychlo |first5=Marta |last6=Nielsen |first6=Per H. |date=December 2021 |title="Candidatus Dechloromonas phosphoritropha" and "Ca. D. phosphorivorans", novel polyphosphate accumulating organisms abundant in wastewater treatment systems |url=https://www.nature.com/articles/s41396-021-01029-2 |journal=The ISME Journal |language=en |volume=15 |issue=12 |pages=3605–3614 |doi=10.1038/s41396-021-01029-2 |issn=1751-7370}}</ref>
[[Dechloromonas|''Candidatus'' ''Dechloromonas'']] species phosphoritropha and phosphorivorans are PAOs with classical metabolism genotype.<ref>{{Cite web |title=Midas Field Guide |url=https://www.midasfieldguide.org/guide/fieldguide/genus/dechloromonas |access-date=2023-07-07 |website=www.midasfieldguide.org}}</ref> ''Dechloromonas'' has been found in high abundances in wastewater treatment plants across the world.<ref>{{Cite journal |last1=Terashima |first1=Mia |last2=Yama |first2=Ayano |last3=Sato |first3=Megumi |last4=Yumoto |first4=Isao |last5=Kamagata |first5=Yoichi |last6=Kato |first6=Souichiro |date=2016 |title=Culture-Dependent and -Independent Identification of Polyphosphate-Accumulating &lt;i&gt;Dechloromonas&lt;/i&gt; spp. Predominating in a Full-Scale Oxidation Ditch Wastewater Treatment Plant |url=http://dx.doi.org/10.1264/jsme2.me16097 |journal=Microbes and Environments |volume=31 |issue=4 |pages=449–455 |doi=10.1264/jsme2.me16097 |pmid=27867159 |pmc=5158118 |issn=1342-6311}}</ref><ref>{{Cite journal |last1=Wang |first1=Baogui |last2=Jiao |first2=Erlong |last3=Guo |first3=Yu |last4=Zhang |first4=Lifang |last5=Meng |first5=Qingan |last6=Zeng |first6=Wei |last7=Peng |first7=Yongzhen |date=2020-07-02 |title=Investigation of the polyphosphate-accumulating organism population in the full-scale simultaneous chemical phosphorus removal system |url=http://dx.doi.org/10.1007/s11356-020-09912-9 |journal=Environmental Science and Pollution Research |volume=27 |issue=30 |pages=37877–37886 |doi=10.1007/s11356-020-09912-9 |pmid=32617817 |s2cid=220305344 |issn=0944-1344}}</ref><ref>{{Cite journal |last1=Stokholm-Bjerregaard |first1=Mikkel |last2=McIlroy |first2=Simon J. |last3=Nierychlo |first3=Marta |last4=Karst |first4=Søren M. |last5=Albertsen |first5=Mads |last6=Nielsen |first6=Per H. |date=2017-04-27 |title=A Critical Assessment of the Microorganisms Proposed to be Important to Enhanced Biological Phosphorus Removal in Full-Scale Wastewater Treatment Systems |journal=Frontiers in Microbiology |volume=8 |page=718 |doi=10.3389/fmicb.2017.00718 |pmid=28496434 |pmc=5406452 |issn=1664-302X |doi-access=free }}</ref><ref name=":23" /> The two species described here, ''Dechloromonas'' phosphoritropha and phosphorivans, are the two most abundant species in waste treatment within the genus.<ref>{{Cite journal |last1=Petriglieri |first1=Francesca |last2=Singleton |first2=Caitlin |last3=Peces |first3=Miriam |last4=Petersen |first4=Jette F. |last5=Nierychlo |first5=Marta |last6=Nielsen |first6=Per H. |date=December 2021 |title="Candidatus Dechloromonas phosphoritropha" and "Ca. D. phosphorivorans", novel polyphosphate accumulating organisms abundant in wastewater treatment systems |journal=The ISME Journal |language=en |volume=15 |issue=12 |pages=3605–3614 |doi=10.1038/s41396-021-01029-2 |pmid=34155336 |pmc=8630035 |issn=1751-7370}}</ref>


=== ''Candidatus'' ''accumulimonas'' (previously referred to as '''''Candidatus''''' Halomonas phosphatis) ===
=== ''Candidatus'' ''accumulimonas'' (previously referred to as '''''Candidatus''''' Halomonas phosphatis) ===
''Candidatus'' ''accumulimonas'' is a species of PAO with classical metabolism phenotype.<ref>{{Cite journal |last=Nguyen |first=Hien Thi Thu |last2=Nielsen |first2=Jeppe Lund |last3=Nielsen |first3=Per Halkjaer |date=October 2012 |title='Candidatus Halomonas phosphatis', a novel polyphosphate-accumulating organism in full-scale enhanced biological phosphorus removal plants: Polyphosphate-accumulating uncultured Halomonas |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1462-2920.2012.02826.x |journal=Environmental Microbiology |language=en |volume=14 |issue=10 |pages=2826–2837 |doi=10.1111/j.1462-2920.2012.02826.x}}</ref><ref>{{Cite web |title=Midas Field Guide |url=https://www.midasfieldguide.org/guide/fieldguide/genus/halomonas |access-date=2023-07-07 |website=www.midasfieldguide.org}}</ref>
''Candidatus'' ''accumulimonas'' is a species of PAO with classical metabolism phenotype.<ref>{{Cite journal |last1=Nguyen |first1=Hien Thi Thu |last2=Nielsen |first2=Jeppe Lund |last3=Nielsen |first3=Per Halkjaer |date=October 2012 |title='Candidatus Halomonas phosphatis', a novel polyphosphate-accumulating organism in full-scale enhanced biological phosphorus removal plants: Polyphosphate-accumulating uncultured Halomonas |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1462-2920.2012.02826.x |journal=Environmental Microbiology |language=en |volume=14 |issue=10 |pages=2826–2837 |doi=10.1111/j.1462-2920.2012.02826.x|pmid=22827168 }}</ref><ref>{{Cite web |title=Midas Field Guide |url=https://www.midasfieldguide.org/guide/fieldguide/genus/halomonas |access-date=2023-07-07 |website=www.midasfieldguide.org}}</ref>


=== ''Microlunatis'' phosphovorus ===
=== ''Microlunatis'' phosphovorus ===
[[Microlunatus phosphovorus|''Microlunatis'' ''phosphovorus'']] is a species of PAO with likely non-canonical PAO metabolism, however exact mechanisms have not been determined.<ref>{{Cite journal |last=Nakamura |first=K. |last2=Hiraishi |first2=A. |last3=Yoshimi |first3=Y. |last4=Kawaharasaki |first4=M. |last5=Masuda |first5=K. |last6=Kamagata |first6=Y. |date=January 1995 |title=Microlunatus phosphovorus gen. nov., sp. nov., a new gram-positive polyphosphate-accumulating bacterium isolated from activated sludge |url=https://pubmed.ncbi.nlm.nih.gov/7857797/ |journal=International Journal of Systematic Bacteriology |volume=45 |issue=1 |pages=17–22 |doi=10.1099/00207713-45-1-17 |issn=0020-7713 |pmid=7857797}}</ref><ref name=":5">{{Cite journal |last=Kawakoshi |first=A. |last2=Nakazawa |first2=H. |last3=Fukada |first3=J. |last4=Sasagawa |first4=M. |last5=Katano |first5=Y. |last6=Nakamura |first6=S. |last7=Hosoyama |first7=A. |last8=Sasaki |first8=H. |last9=Ichikawa |first9=N. |last10=Hanada |first10=S. |last11=Kamagata |first11=Y. |last12=Nakamura |first12=K. |last13=Yamazaki |first13=S. |last14=Fujita |first14=N. |date=2012-08-23 |title=Deciphering the Genome of Polyphosphate Accumulating Actinobacterium Microlunatus phosphovorus |url=http://dx.doi.org/10.1093/dnares/dss020 |journal=DNA Research |volume=19 |issue=5 |pages=383–394 |doi=10.1093/dnares/dss020 |issn=1340-2838}}</ref><ref>{{Cite journal |last=Zhong |first=Chuanqing |last2=Zhang |first2=Peipei |last3=Liu |first3=Cheng |last4=Liu |first4=Meng |last5=Chen |first5=Wenbing |last6=Fu |first6=Jiafang |last7=Qi |first7=Xiaoyu |last8=Cao |first8=Guangxiang |date=2019 |title=The PolS-PolR Two-Component System Regulates Genes Involved in Poly-P Metabolism and Phosphate Transport in Microlunatus phosphovorus |url=https://www.frontiersin.org/articles/10.3389/fmicb.2019.02127 |journal=Frontiers in Microbiology |volume=10 |doi=10.3389/fmicb.2019.02127/full |issn=1664-302X}}</ref> Belonging to the same phylum as ''Ca.'' phosphoribacter, these two actinobacterial organisms exhibit similar metabolisms, however ''M.'' ''phosphovorus'' has been suggested to hyperaccumulate over ten times the amount of polyphosphate per cell mass dry weight compared to ''Ca.'' ''phosphoribacter'' or proteobacterial PAOs.<ref name=":5" />
[[Microlunatus phosphovorus|''Microlunatis'' ''phosphovorus'']] is a species of PAO with likely non-canonical PAO metabolism, however exact mechanisms have not been determined.<ref>{{Cite journal |last1=Nakamura |first1=K. |last2=Hiraishi |first2=A. |last3=Yoshimi |first3=Y. |last4=Kawaharasaki |first4=M. |last5=Masuda |first5=K. |last6=Kamagata |first6=Y. |date=January 1995 |title=Microlunatus phosphovorus gen. nov., sp. nov., a new gram-positive polyphosphate-accumulating bacterium isolated from activated sludge |url=https://pubmed.ncbi.nlm.nih.gov/7857797/ |journal=International Journal of Systematic Bacteriology |volume=45 |issue=1 |pages=17–22 |doi=10.1099/00207713-45-1-17 |issn=0020-7713 |pmid=7857797}}</ref><ref name=":5">{{Cite journal |last1=Kawakoshi |first1=A. |last2=Nakazawa |first2=H. |last3=Fukada |first3=J. |last4=Sasagawa |first4=M. |last5=Katano |first5=Y. |last6=Nakamura |first6=S. |last7=Hosoyama |first7=A. |last8=Sasaki |first8=H. |last9=Ichikawa |first9=N. |last10=Hanada |first10=S. |last11=Kamagata |first11=Y. |last12=Nakamura |first12=K. |last13=Yamazaki |first13=S. |last14=Fujita |first14=N. |date=2012-08-23 |title=Deciphering the Genome of Polyphosphate Accumulating Actinobacterium Microlunatus phosphovorus |url=http://dx.doi.org/10.1093/dnares/dss020 |journal=DNA Research |volume=19 |issue=5 |pages=383–394 |doi=10.1093/dnares/dss020 |pmid=22923697 |pmc=3473371 |issn=1340-2838}}</ref><ref>{{Cite journal |last1=Zhong |first1=Chuanqing |last2=Zhang |first2=Peipei |last3=Liu |first3=Cheng |last4=Liu |first4=Meng |last5=Chen |first5=Wenbing |last6=Fu |first6=Jiafang |last7=Qi |first7=Xiaoyu |last8=Cao |first8=Guangxiang |date=2019 |title=The PolS-PolR Two-Component System Regulates Genes Involved in Poly-P Metabolism and Phosphate Transport in Microlunatus phosphovorus |journal=Frontiers in Microbiology |volume=10 |page=2127 |doi=10.3389/fmicb.2019.02127 |pmid=31572333 |pmc=6754071 |issn=1664-302X |doi-access=free }}</ref> Belonging to the same phylum as ''Ca.'' phosphoribacter, these two actinobacterial organisms exhibit similar metabolisms, however ''M.'' ''phosphovorus'' has been suggested to hyperaccumulate over ten times the amount of polyphosphate per cell mass dry weight compared to ''Ca.'' ''phosphoribacter'' or proteobacterial PAOs.<ref name=":5" />


=== ''Pseudomonas'' spp. ===
=== ''Pseudomonas'' spp. ===
Some unnamed species of the ''[[Pseudomonas]]'' genus have been known to exhibit PAO phenotypes.<ref name=":1">{{Cite journal |last=Günther |first=S. |last2=Trutnau |first2=M. |last3=Kleinsteuber |first3=S. |last4=Hause |first4=G. |last5=Bley |first5=T. |last6=Röske |first6=I. |last7=Harms |first7=H. |last8=Müller |first8=S. |date=April 2009 |title=Dynamics of Polyphosphate-Accumulating Bacteria in Wastewater Treatment Plant Microbial Communities Detected via DAPI (4′,6′-Diamidino-2-Phenylindole) and Tetracycline Labeling |url=http://dx.doi.org/10.1128/aem.01540-08 |journal=Applied and Environmental Microbiology |volume=75 |issue=7 |pages=2111–2121 |doi=10.1128/aem.01540-08 |issn=0099-2240}}</ref>
Some unnamed species of the ''[[Pseudomonas]]'' genus have been known to exhibit PAO phenotypes.<ref name=":1">{{Cite journal |last1=Günther |first1=S. |last2=Trutnau |first2=M. |last3=Kleinsteuber |first3=S. |last4=Hause |first4=G. |last5=Bley |first5=T. |last6=Röske |first6=I. |last7=Harms |first7=H. |last8=Müller |first8=S. |date=April 2009 |title=Dynamics of Polyphosphate-Accumulating Bacteria in Wastewater Treatment Plant Microbial Communities Detected via DAPI (4′,6′-Diamidino-2-Phenylindole) and Tetracycline Labeling |url=http://dx.doi.org/10.1128/aem.01540-08 |journal=Applied and Environmental Microbiology |volume=75 |issue=7 |pages=2111–2121 |doi=10.1128/aem.01540-08 |pmid=19181836 |pmc=2663203 |issn=0099-2240}}</ref>


=== ''Paracoccus'' ''denitrificans'' ===
=== ''Paracoccus'' ''denitrificans'' ===
[[Paracoccus denitrificans|''Paracoccus'' ''denitrificans'']] has been known to exhibit a non-canonical PAO phenotype.<ref name=":1" /><ref>{{Cite journal |last=Barak |first=Yoram |last2=van Rijn |first2=Jaap |date=March 2000 |title=Atypical Polyphosphate Accumulation by the Denitrifying Bacterium ''Paracoccus denitrificans'' |url=http://dx.doi.org/10.1128/aem.66.3.1209-1212.2000 |journal=Applied and Environmental Microbiology |volume=66 |issue=3 |pages=1209–1212 |doi=10.1128/aem.66.3.1209-1212.2000 |issn=0099-2240}}</ref>
[[Paracoccus denitrificans|''Paracoccus'' ''denitrificans'']] has been known to exhibit a non-canonical PAO phenotype.<ref name=":1" /><ref>{{Cite journal |last1=Barak |first1=Yoram |last2=van Rijn |first2=Jaap |date=March 2000 |title=Atypical Polyphosphate Accumulation by the Denitrifying Bacterium ''Paracoccus denitrificans'' |url=http://dx.doi.org/10.1128/aem.66.3.1209-1212.2000 |journal=Applied and Environmental Microbiology |volume=66 |issue=3 |pages=1209–1212 |doi=10.1128/aem.66.3.1209-1212.2000 |pmid=10698794 |pmc=91965 |issn=0099-2240}}</ref>


==References==
==References==

Revision as of 02:47, 10 July 2023

Cluster of PAOs

Polyphosphate-accumulating organisms (PAOs) are a group of microorganisms that, under certain conditions, facilitate the removal of large amounts of phosphorus from their environments. The most studied example of this phenomenon is in bacteria found in a form of wastewater processing known as enhanced biological phosphorus removal (EBPR), however phosphate hyperaccumulation has been found to occur in other conditions such as soil and marine environments, as well as in non-bacterial organisms such as fungi and algae.[1] PAOs accomplish this removal of phosphate by accumulating it within their cells as polyphosphate. PAOs are by no means the only microbes that can accumulate phosphate within their cells and in fact, the production of polyphosphate is a widespread ability among microbes. However, PAOs have many characteristics that other organisms that accumulate polyphosphate do not have that make them amenable to use in wastewater treatment. Specifically, in the case of classical PAOs, is the ability to consume simple carbon compounds (energy source) without the presence of an external electron acceptor (such as nitrate or oxygen) by generating energy from internally stored polyphosphate and glycogen. Most other bacteria cannot consume under these conditions and therefore PAOs gain a selective advantage within the mixed microbial community present in the activated sludge. Therefore, wastewater treatment plants that operate for enhanced biological phosphorus removal have an anaerobic tank (where there is no nitrate or oxygen present as external electron acceptor) prior to the other tanks to give PAOs preferential access to the simple carbon compounds in the wastewater that is influent to the plant.

Metabolisms

Classical (Canonical) PAO Metabolism

The classical or "canonical" behavior of PAOs is considered to be the release of phosphate (as orthophosphate) to the environment and transformation of intracellular polyphosphate reserves into polyhydroxyalkanoates (PHA) from volatile fatty acids (VFAs) and glycogen during anoxic conditions.[2] This is followed by the consumption of the PHA/VFAs and uptake of environmental orthophosphate during oxic conditions to regenerate polyphosphate reserves within the cell.[2]

Non-Canonical (or "Fermentative) PAO Metabolism

Some PAOs have been found to have alternative methods to accumulating polyphosphate, particularly to do with not storing PHA or glycogen.[3][4] This is generally believed to be seen more often in extracellular environments high in organic compounds, thus containing fermentable substrates like amino acids and sugars.[5] However, the exact mechanisms of these microbes to accumulate and use polyphosphate are not well understood.[4]

Known Bacterial PAOs

Candidatus Phosphoribacter (previously referred to as Tetrasphaera prior to 2022)

Candidatus Phosphoribacter is a bacterial genus that has been found to be the dominant PAO associated with wastewater treatment worldwide, and has been found to often participate more in the biological removal of phosphorus than Candidatus Accumulibacter, contrary to previous understandings.[3][6][4] This bacteria has been found to be a non-canonical (or fermentative/"fPAO") PAO, and universally lack the genetic potential to store PHA.[3][7] This genus was largely found to be capable of producing the fermentation products acetate, lactate, alanine, and succinate.[7][8] Additionally, it is suggested that the amino acids lysine, arginine, histidine, leucine, isoleucine, valine and phenylalanine may replace the canonical purpose of PHA as an energy substrate during oxic conditions, based on genomic potential and similarity to behavior of other microbial metabolisms.[3] Alternatively, the compound cyanophycin may used as an energy substrate due to the ubiquity of cyanophycin-metabolizing enzymes encoded in the species.[3]

Candidatus Accumlibacter phosphatis

Candidatus Accumulibacter phosphatis is one of the most well-studied PAOs, and is responsible for the development of the classical PAO metabolic model which Ca. Phosphoribacter later contradicted.[9] Formerly considered the most important PAO in waste treatment, the bacteria is highly abundant in wastewater treatment plants globally.[4][10] It can consume a range of carbon compounds, such as acetate and propionate, under anaerobic conditions and store these compounds as polyhydroxyalkanoates (PHA) which it consumes as a carbon and energy source for growth using oxygen or nitrate as electron acceptor. Historically, the hyperaccumulation of phosphate by Ca. Accumulibacter was seen as a stress response, but currently it is suggested that this behavior may play an ecological role.[11] In combination with Ca. Phosphoribacter, these two PAOs are considered to account for 24-70% of phosphorus removed from wastewater during treatment processing.[6]

Candidatus dechloromonas

Candidatus Dechloromonas species phosphoritropha and phosphorivorans are PAOs with classical metabolism genotype.[12] Dechloromonas has been found in high abundances in wastewater treatment plants across the world.[13][14][15][4] The two species described here, Dechloromonas phosphoritropha and phosphorivans, are the two most abundant species in waste treatment within the genus.[16]

Candidatus accumulimonas (previously referred to as Candidatus Halomonas phosphatis)

Candidatus accumulimonas is a species of PAO with classical metabolism phenotype.[17][18]

Microlunatis phosphovorus

Microlunatis phosphovorus is a species of PAO with likely non-canonical PAO metabolism, however exact mechanisms have not been determined.[19][20][21] Belonging to the same phylum as Ca. phosphoribacter, these two actinobacterial organisms exhibit similar metabolisms, however M. phosphovorus has been suggested to hyperaccumulate over ten times the amount of polyphosphate per cell mass dry weight compared to Ca. phosphoribacter or proteobacterial PAOs.[20]

Pseudomonas spp.

Some unnamed species of the Pseudomonas genus have been known to exhibit PAO phenotypes.[22]

Paracoccus denitrificans

Paracoccus denitrificans has been known to exhibit a non-canonical PAO phenotype.[22][23]

References

  1. ^ Akbari, Ali; Wang, ZiJian; He, Peisheng; Wang, Dongqi; Lee, Jangho; Han, Il; Li, Guangyu; Gu, April Z. (January 2021). "Unrevealed roles of polyphosphate‐accumulating microorganisms". Microbial Biotechnology. 14 (1): 82–87. doi:10.1111/1751-7915.13730. ISSN 1751-7915. PMC 7888455. PMID 33404187.
  2. ^ a b Akram, Fatima; Aqeel, Amna; Ahmed, Zeeshan; Zafar, Javeria; Haq, Ikram ul (2022-01-01), Dar, Gowhar Hamid; Bhat, Rouf Ahmad; Qadri, Humaira; Hakeem, Khalid Rehman (eds.), "Chapter 8 - Role of polyphosphate accumulating organisms in enhanced biological phosphorous removal", Microbial Consortium and Biotransformation for Pollution Decontamination, Advances in Environmental Pollution Research, Elsevier, pp. 151–179, ISBN 978-0-323-91893-0, retrieved 2023-07-07
  3. ^ a b c d e Singleton, C. M.; Petriglieri, F.; Wasmund, K.; Nierychlo, M.; Kondrotaite, Z.; Petersen, J. F.; Peces, M.; Dueholm, M. S.; Wagner, M.; Nielsen, P. H. (June 2022). "The novel genus, 'Candidatus Phosphoribacter', previously identified as Tetrasphaera, is the dominant polyphosphate accumulating lineage in EBPR wastewater treatment plants worldwide". The ISME Journal. 16 (6): 1605–1616. doi:10.1038/s41396-022-01212-z. ISSN 1751-7370. PMC 9123174. PMID 35217776.
  4. ^ a b c d e Nielsen, Per Halkjær; McIlroy, Simon J.; Albertsen, Mads; Nierychlo, Marta (June 2019). "Re-evaluating the microbiology of the enhanced biological phosphorus removal process". Current Opinion in Biotechnology. 57: 111–118. doi:10.1016/j.copbio.2019.03.008. ISSN 1879-0429. PMID 30959426. S2CID 104294644.
  5. ^ Nguyen, Hien Thi Thu; Kristiansen, Rikke; Vestergaard, Mette; Wimmer, Reinhard; Nielsen, Per Halkjær (2015-07-15). "Intracellular Accumulation of Glycine in Polyphosphate-Accumulating Organisms in Activated Sludge, a Novel Storage Mechanism under Dynamic Anaerobic-Aerobic Conditions". Applied and Environmental Microbiology. 81 (14): 4809–4818. doi:10.1128/aem.01012-15. ISSN 0099-2240. PMC 4551194. PMID 25956769.
  6. ^ a b Fernando, Eustace Y.; McIlroy, Simon Jon; Nierychlo, Marta; Herbst, Florian-Alexander; Petriglieri, Francesca; Schmid, Markus C.; Wagner, Michael; Nielsen, Jeppe Lund; Nielsen, Per Halkjær (August 2019). "Resolving the individual contribution of key microbial populations to enhanced biological phosphorus removal with Raman-FISH". The ISME Journal. 13 (8): 1933–1946. doi:10.1038/s41396-019-0399-7. ISSN 1751-7370. PMC 6776032. PMID 30894691.
  7. ^ a b Otieno, Jeremiah; Kowal, Przemysław; Mąkinia, Jacek (2022-10-28). "The Occurrence and Role of Tetrasphaera in Enhanced Biological Phosphorus Removal Systems". Water. 14 (21): 3428. doi:10.3390/w14213428. ISSN 2073-4441.
  8. ^ Kristiansen, Rikke; Nguyen, Hien Thi Thu; Saunders, Aaron Marc; Nielsen, Jeppe Lund; Wimmer, Reinhard; Le, Vang Quy; McIlroy, Simon Jon; Petrovski, Steve; Seviour, Robert J.; Calteau, Alexandra; Nielsen, Kåre Lehmann; Nielsen, Per Halkjær (March 2013). "A metabolic model for members of the genus Tetrasphaera involved in enhanced biological phosphorus removal". The ISME Journal. 7 (3): 543–554. doi:10.1038/ismej.2012.136. ISSN 1751-7370. PMC 3578573. PMID 23178666.
  9. '^ He, Shaomei; McMahon, Katherine D. (2011-02-21). "Microbiology of CandidatusAccumulibacter' in activated sludge". Microbial Biotechnology. 4 (5): 603–619. doi:10.1111/j.1751-7915.2011.00248.x. ISSN 1751-7915. PMC 3819010. PMID 21338476.
  10. ^ Onnis‐Hayden, Annalisa; Srinivasan, Varun; Tooker, Nicholas B.; Li, Guangyu; Wang, Dongqi; Barnard, James L.; Bott, Charles; Dombrowski, Paul; Schauer, Peter; Menniti, Adrienne; Shaw, Andrew; Stinson, Beverly; Stevens, Gerry; Dunlap, Patrick; Takács, Imre (March 2020). "Survey of full‐scale sidestream enhanced biological phosphorus removal (S2EBPR) systems and comparison with conventional EBPRs in North America: Process stability, kinetics, and microbial populations". Water Environment Research. 92 (3): 403–417. doi:10.1002/wer.1198. ISSN 1061-4303. PMID 31402530. S2CID 199539909.
  11. ^ da Silva, Leonor Guedes; Gamez, Karel Olavarria; Gomes, Joana Castro; Akkermans, Kasper; Welles, Laurens; Abbas, Ben; van Loosdrecht, Mark C.M.; Wahl, Sebastian Aljoscha (2018-11-01). "Revealing metabolic flexibility ofCandidatusAccumulibacter phosphatis through redox cofactor analysis and metabolic network modeling". dx.doi.org. doi:10.1101/458331. S2CID 91862227. Retrieved 2023-07-07.
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