Frankia

Frankia is a genus of nitrogen-fixing bacteria that live in symbiosis with actinorhizal plants, similar to the Rhizobium bacteria found in the root nodules of legumes in the family Fabaceae. Frankia also initiate the forming of root nodules.

This genus was originally named by Jørgen Brunchorst, in 1886 to honor the German biologist Albert Bernhard Frank.[1] Brunchorst considered the organism he had identified to be a filamentous fungus. Jan-Hendrik Becking (de) redefined the genus in 1970 as containing prokaryotic actinomycetes and created the family Frankiaceae within the Actinomycetales. He retained the original name of Frankia for the genus.[2]

A section through an alder root nodule

## Contents

* 1 Overview
* 2 Symbiont plants
* 3 Nodule Formation
* 4 Phylogeny
* 5 See also
* 6 References

## Overview

Most Frankia strains are specific to different plant species. The bacteria are filamentous and convert atmospheric nitrogen into ammonia via the enzyme nitrogenase, a process known as nitrogen fixation. They do this while living in root nodules on actinorhizal plants. The bacteria can supply most or all of the nitrogen requirements of the host plant. As a result, actinorhizal plants colonise and often thrive in soils that are low in plant nutrients.[3]

Several Frankia genomes are now available which may help clarify how the symbiosis between prokaryote and plant evolved, how the environmental and geographical adaptations occurred, the metabolic diversity, and the horizontal gene flow among the symbiotic prokaryotes.[3]

Frankia can resist low concentration of heavy metals such as, Cu, Co, and Zn.[4] Frankia may be an advantage for degraded soil. Degraded soil is known as soil that is heavy metal rich or nutrient depleted due to a drought. Frankia is a nitrogen-fixed organism, explaining why it is able to resist heavy metals.[5][clarification needed]

Frankia is a gram-positive Bacteria that is found on the roots of plants. The fact that Frankia is gram-positive means that the bacteria is made up of thick cell walls made out of protein called peptidologlycan. This helps with the resistance of the heavy metals that may be in the degraded soil.[6]

Frankia tolerates a narrow range of temperatures and soil pH levels. It grows best at around 30 degrees Celsius with an environment pH between 6.5 and 7.[7] These facts shows that Frankia is very sensitive to its environment. Though Frankia would not be suitable for all agriculture it does demonstrate possibilities in select areas, or in temperature controlled environments.

## Symbiont plants

Main article: Biology:Actinorhizal plant

* All species in the genus Alnus in the family Betulaceae
* Some species in all four genera in the family Casuarinaceae
* Certain species in the genus Coriaria in the family Coriariaceae
* Datisca cannabina and Datisca glomerata in the family Datiscaceae
* All species in the three genera in the family Elaeagnaceae, Elaeagnus, Shepherdia, and Hippophae
* All species in the genera Myrica, Morella, and Comptonia in the family Myricaceae.
* All species in six genera in the family Rhamnaceae, Ceanothus, Colletia, Discaria, Trevoa, and possibly Adolphia
* Some species in the family Rosaceae including all the species in the genera Cercocarpus, Cowania, Purshia, Chamaebatia, and some species of Dryas[8]

## Nodule Formation

File:Frankia Longitudinal Section Toluidine Blue- Atchison, Black.pngLongitudinal section of a Frankia nodule dyed with Toluidine Blue to highlight the vascular tissue in blue and purple.

Frankia forms nodules via two methods of root infection, intercellularly and intracellularly.[9] Intracellular infection is characterized by initial root-hair deformation which is then infected by the filamentous Frankia. The Frankia then moves within the root cells and forms a pre-nodule which is characterized by a bump on the root. This then gives rise to a Nodule primordium which feeds the bacteria via the vascular tissue of the plant allowing the nodule to mature.[9]

In contrast the intercellular infection does not have root hair deformation. Instead, the filamentous Frankia invades the roots in the space between cells on the root. After this invasion a Nodule primordium is created similarly to the intracellular mode of formation and the nodule matures.[9]

## Phylogeny

The currently accepted taxonomy is based on the List of Prokaryotic names with Standing in Nomenclature (LPSN)[10] and National Center for Biotechnology Information (NCBI).[11]

16S rRNA based LTP_10_2024[12][13][14] | 120 marker proteins based GTDB 09-RS220[15][16][17]

|

|  | Frankia |  |  |

F. coriariae Nouioui et al. 2017

|

|  |  |  |  |  |  |

F. casuarinae Nouioui et al. 2016

|

F. canadensis Normand et al. 2018[18]

|  |  |

F. umida Normand et al. 2023

|

|  |  |

F. alni (Woronin 1866) Von Tubeuf 1895

|

|  |  |

F. torreyi Nouioui et al. 2019

|

|  |  |

"F. gtarii" Nouioui et al. 2023

|

"F. tisai" Nouioui et al. 2023

|  |  |  |  |

F. inefficax Nouioui et al. 2017

|

|  |  |

F. asymbiotica Nouioui et al. 2017

|

F. saprophytica Nouioui et al. 2018

|  |  |

F. discariae Nouioui et al. 2017

|

|  |  |

F. soli Gtari et al. 2020

|

|  |  |

F. irregularis Nouioui et al. 2018

|

|  |  |

F. colletiae Nouioui et al. 2023

|

F. elaeagni (Schroeter 1886) Becking
1970 ex Nouioui et al. 2016

Protofrankia

Frankia s.s.

Pseudofrankia

Parafrankia

|  |  |  |  | Protofrankia |  |  |

"Ca. Frankia meridionalis" Nguyen et al. 2019

|

|  |  |

"Ca. Frankia californiensis" Normand et al. 2017[19]

|

P. coriariae [incl. "Ca. P. datiscae" (Persson et al. 2011) Gtari 2022]

|  |  |  | Pseudofrankia |  |  |

P. inefficax

|

|  |  |

P. asymbiotica

|

P. saprophytica

|  | Parafrankia |  |  |  |  |

P. discariae

|

P. soli

|  |  |

P. irregularis

|

|  |  |

P. colletiae

|

P. elaeagni

Frankia |  |  |

F. casuarinae

|

|  |  |  |  |

F. canadensis

|

F. umida ["Ca. F. nodulisporulans" Herrera-Belaroussi et al. 2020]

|  |  |  |  |

F. alni

|

F. torreyi

|  |  |

"Ca. F. alpina" Pozzi et al. 2020 [incl. "F. subtilis" Brunchorst 1886]

|

|  |  |

"F. gtarii"

|

"F. tisai"

Species incertae sedis:

* F. nepalensis Nouioui et al. 2023

## See also

* List of bacterial orders
* List of bacteria genera

## References

1. ↑ Pawlowski, Katharina (2009-06-17). Prokaryotic Symbionts in Plants. Springer Science & Business Media. pp. 107. ISBN 9783540754602. https://books.google.com/books?id=QQKdiHoNE_gC&q=A.B.Frank+frankia&pg=PA107.
2. ↑ "Frankia taxonomy". http://web.uconn.edu/mcbstaff/benson/Frankia/FrankiaTaxonomy.htm.
3. ↑ 3.0 3.1 Frankia and Actinorhizal Plants
4. ↑ Abdel‐lateif, Khalid Salah El dein; Mansour, Samira R.; El‐Badawy, Mohamed F.; Shohayeb, Mohamed M. (2018). "Isolation and molecular characterization of Frankia strains resistant to some heavy metals". Journal of Basic Microbiology 58 (9): 720–729. doi:10.1002/jobm.201800122. ISSN 1521-4028. PMID 29962068.
5. ↑ El dein Abdel-lateif, Khalid Salah; Mansour, Samira R.; El-Badawy, Mohamed F.; Shohayeb, Mohamed M. (September 2018). "Isolation and molecular characterization of Frankia strains resistant to some heavy metals". Journal of Basic Microbiology 58 (9): 720–729. doi:10.1002/jobm.201800122. PMID 29962068.
6. ↑ Nouioui, Imen; Ghodhbane-Gtari, Faten; del Carmen Montero-Calasanz, Maria; Rohde, Manfred; Tisa, Louis S.; Gtari, Maher; Klenk, Hans-Peter (2017-03-01). "Frankia inefficax sp. nov., an actinobacterial endophyte inducing ineffective, non nitrogen-fixing, root nodules on its actinorhizal host plants". Antonie van Leeuwenhoek 110 (3): 313–320. doi:10.1007/s10482-016-0801-7. ISSN 1572-9699. PMID 27830471.
7. ↑ Srivastava, Amrita; Singh, Anumeha; Singh, Satya S.; Mishra, Arun K. (2017-04-16). "Salt stress–induced changes in antioxidative defense system and proteome profiles of salt-tolerant and sensitive Frankia strains". Journal of Environmental Science and Health, Part A 52 (5): 420–428. doi:10.1080/10934529.2016.1270672. ISSN 1093-4529. PMID 28085556.
8. ↑ Schwintzer, C. R.; Tjepkema, J. (1990). The Biology of Frankia and Actinorhizal Plants. San Diego: Academic Press. ISBN 978-0126332100.
9. ↑ 9.0 9.1 9.2 Santi, Carole; Bogusz, Didier; Franche, Claudine (2013-03-10). "Biological nitrogen fixation in non-legume plants". Annals of Botany 111 (5): 743–767. doi:10.1093/aob/mct048. ISSN 1095-8290. PMC 3631332. http://dx.doi.org/10.1093/aob/mct048.
10. ↑ A.C. Parte. "Frankia". List of Prokaryotic names with Standing in Nomenclature (LPSN). https://lpsn.dsmz.de/genus/frankia.
11. ↑ C.L. Schoch. "Frankia". National Center for Biotechnology Information (NCBI) taxonomy database. https://www.ncbi.nlm.nih.gov/Taxonomy/Browser/wwwtax.cgi?mode=Tree&id=1854&lvl=3&lin=f&keep=1&srchmode=1&unlock.
12. ↑ "The LTP". https://imedea.uib-csic.es/mmg/ltp/#LTP.
13. ↑ "LTP_all tree in newick format". https://imedea.uib-csic.es/mmg/ltp/wp-content/uploads/ltp/LTP_all_10_2024.ntree.
14. ↑ "LTP_10_2024 Release Notes". https://imedea.uib-csic.es/mmg/ltp/wp-content/uploads/ltp/LTP_10_2024_release_notes.pdf.
15. ↑ "GTDB release 09-RS220". https://gtdb.ecogenomic.org/about#4|.
16. ↑ "bac120_r220.sp_labels". https://data.gtdb.ecogenomic.org/releases/release220/220.0/auxillary_files/bac120_r220.sp_labels.tree.
17. ↑ "Taxon History". https://gtdb.ecogenomic.org/taxon_history/.
18. ↑ Normand, P; Nouioui, I; Pujic, P; Fournier, P; Dubost, A; Schwob, G; Klenk, HP; Nguyen, A et al. (2018). "Frankia canadensis sp. nov., isolated from root nodules of Alnus incana subspecies rugosa". Int J Syst Evol Microbiol 68 (9): 3001–3011. doi:10.1099/ijsem.0.002939. PMID 30059001.
19. ↑ Normand P; Nguyen, T.V.; Battenberg, K; Berry, A.M.; Heuvel, B.V.; Fernandez, M.P.; Pawlowski, K. (2017). "Proposal of "Candidatus Frankia californiensis", the uncultured symbiont in nitrogen-fixing root nodules of a phylogenetically broad group of hosts endemic to western North America". Int J Syst Evol Microbiol 67 (10): 3706–3715. doi:10.1099/ijsem.0.002147. PMID 28884663.

* v
* t
* e

Prokaryotes: Bacteria classification (phyla and orders)

Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta

Animal
Fungi)

G-/
OM|

| Terra-/
Glidobacteria
(BV1)|

| Eobacteria|

* Deinococcus–Thermus
* Deinococcales
* Thermales
* Chloroflexi
* Anaerolineales
* Caldilineales
* Chloroflexales
* Herpetosiphonales
* Dehalococcoidales
* Ktedonobacterales
* Thermogemmatisporales
* Thermomicrobiales
* Sphaerobacterales

|

other glidobacteria|

* Thermodesulfobacteria
* thermophiles
* Aquificae
* Thermotogae
* Cyanobacteria

Proteobacteria
(BV2)|

| Alpha|

* Caulobacterales
* Kiloniellales
* Kordiimonadales
* Magnetococcales
* Parvularculales
* Rhizobiales
* Rhodobacterales
* Rhodospirillales
* Rickettsiales
* Sneathiellales
* Sphingomonadales

|

Beta|

* Burkholderiales
* Hydrogenophilales
* Methylophilales
* Neisseriales
* Nitrosomonadales
* Procabacteriales
* Rhodocyclales

Gamma|

* Acidithiobacillales
* Aeromonadales
* Alteromonadales
* Cardiobacteriales
* Chromatiales
* Enterobacterales
* Legionellales
* Methylococcales
* Oceanospirillales
* Orbales
* Pasteurellales
* Pseudomonadales
* Salinisphaerales
* Thiotrichales
* Vibrionales
* Xanthomonadales

Delta|

* Bdellovibrionales
* Desulfarculales
* Desulfobacterales
* Desulfovibrionales
* Desulfurellales
* Desulfuromonadales
* Myxococcales
* Syntrophobacterales
* Syntrophorhabdales

Epsilon|

* Campylobacterales
* Nautiliales

Zeta|

* Mariprofundales

BV4|

| Spirochaetes|

* Spirochaetes

|

Sphingobacteria
(FCB group)|

* Fibrobacteres
* Chlorobi
* Chlorobiales
* Ignavibacteriales
* Bacteroidetes
* Bacteroidales
* Cytophagales
* Flavobacteriales
* Sphingobacteriales

Planctobacteria/
(PVC group)|

* Chlamydiae
* Lentisphaerae
* Lentisphaerales
* Oligosphaerales
* Victivallales
* Planctomycetes
* Phycisphaerales
* Planctomycetales
* Verrucomicrobia
* Puniceicoccales
* Opitutales
* Chthoniobacterales
* Verrucomicrobiales
* "Poribacteria"

Other GN|

* Acidobacteria
* Acidobacteriales
* Acanthopleuribacterales
* Holophagales
* Solibacterales
* Armatimonadetes
* Armatimonadales
* Chthonomonadales
* Fimbriimonadales
* Caldiserica
* Chrysiogenetes
* Deferribacteres
* Dictyoglomi
* Elusimicrobia
* Fusobacteria
* Gemmatimonadetes
* Nitrospirae
* Synergistetes

G+/
no OM|

| Firmicutes
(BV3)|

| Bacilli|

* Bacillales
* Lactobacillales
* Aphragmobacteria
* Erysipelotrichia
* Erysipelotrichiales
* Mollicutes
* Mycoplasmatales
* Entomoplasmatales
* Anaeroplasmatales
* Acholeplasmatales
* Haloplasmatales

|

Clostridia|

* Clostridiales
* Halanaerobiales
* Thermoanaerobacteriales
* Natranaerobiales
* Negativicutes
* Selenomonadales

Thermolithobacteria|

* Thermolithobacterales

Actinobacteria
(BV5)|

| Actinobacteria|

* Actinomycetales
* Bifidobacteriales

|

Acidimicrobiia|

* Acidimicrobiales

Coriobacteriia|

* Coriobacteriales

Nitriliruptoria|

* Euzebyales
* Nitriliruptorales

Rubrobacteria|

* Gaiellales
* Rubrobacterales
* Thermoleophilales
* Solirubrobacterales

Incertae
sedis|

* †Archaeosphaeroides
* †Eobacterium
* †Leptotrichites

Source: Bergey's Manual (2001–2012). Alternative views: Wikispecies.

Wikidata ☰ Q134931 entry

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