Understanding the Genetic Diversity of Bacteria Isolated from Across the Atacama Desert

Abstract

Despite being one of the driest and harshest deserts on Earth, the Atacama Desert is home to a variety of bacterial life. Microorganisms that reside here may have developed adaptations to help them survive this unique environment. In this study, we used bioinformatic and genetic methods to assess the abundance of phyla that are present in this environment and focus on the types of adaptations individual bacteria have obtained. To assess bacterial diversity, we used 16S rRNA sequencing on soil samples and determined the relative composition of different phyla and archaea at sixteen locations. The whole genome sequence genome of eight selected pigmented bacteria was also performed. We found that all strains we sequenced are predicted to produce bioactive compounds. We focused on stress-tolerance capabilities, including pigment production pathways, biofilm-related genes, antibiotic production, and genome stability. We also found that the pigments that these bacteria produce have antioxidant, iron, and ion chelating, and/or antibiotic properties. This characterization allows us to assess adaptive strategies of bacteria, which is important in the fields of agriculture, biotechnology, and health.

Keywords

Extremophiles / Microbiology / Atacama Desert / Environmental microbiology / Genetic diversity

Supplementary Materials

The following supporting information can be found at: https://www.sciepublish.com/article/pii/795. Figure S1: An example of a plate from which bacteria were isolated for WGS; Figure S2: A phylogenetic tree generated by autoMLST2; Table S1: Locations, coordinates, and elevations of sampling locations for 16S sequencing; Table S2: Locations, coordinates, and elevations of sampling locations where bacteria where isolated from for WGS.

Acknowledgments

We would like to thank all Northeastern University (NU) students who participated in Dialogue of Civilizations (DOC) trips to Chile (2018, 2022, 2024). These students helped perform sample collection and bacterial isolation. Thank you to Joey Lehman Morris for co-teaching these DOC courses and participating in sample collection. Thank you also to our OneSeed team, including Sofia Mardones and Guillermo Maluenda, for guiding us to sampling locations. A special thanks also to the Riquelme Lab for supporting our team while in Chile.

Author Contributions

Conceptualization: V.G.-C., C.R., Y.C., A.R.P. and N.T.C.; Methodology: V.G.-C., N.T.C., A.R.P. and E.I.; Formal Analysis: N.T.C., A.R.P., E.I., E.A., A.H. and M.C.F.; Investigation: N.T.C., A.R.P., E.I., E.A., A.H., N.T., M.C.F. and M.T.; Writing—Original Draft: N.T.C. and A.R.P.; Writing—Review & Editing: N.T.C., A.R.P. and V.G.-C.; Supervision: Y.C., C.R. and V.G.-C.; Project Administration: V.G.-C. and Y.C.; Funding Acquisition: V.G.-C. and Y.C.

Ethics Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

WGS projects have been deposited into NCBI. Each sample is deposited under the following BioSample numbers: 0102A is under SAMN39706723, 0209A is under SAMN39706724, 0909A is under SAMN39706725, 0516A is under SAMN50557742, 1020B is under SAMN50557744, 0819A is under SAMN50557743, Iso1_2024 is under SAMN50557745, and Iso2_2024 is under SAMN50557746. Additionally, we assessed the G4 and GC content of three strains which are publicly available on NCBI. We assessed Bacillus subtilis with an NCBI ID of CP020102.1, Pseudomonas aeruginosa with an NCBI ID of NZ_CP027174.1, and Kocuria rhizophila with an NCBI ID of CP124833.1.

Funding

This work was funded by the Northeastern University (NU) Global Experience Office. E.I., E.A., A.H., N.T., M.C.F. and M.T. were funded by NU’s PEAK Undergraduate Research Fellowships. A.R.P. was funded by the NU Provost Dissertation Completion Fellowship. N.T.C. was funded by the National Science Foundation (NSF) Graduate Research Fellowship Program (1938052). V.G.-C. was funded by NuSci, a grant from HHMI. Y.C. was supported by NSF grant MCB1651732.

Declaration of Competing Interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

References

[1] Teske A, de Beer D, McKay LJ, Tivey MK, Biddle JF, Hoer D, et al. The Guaymas Basin Hiking Guide to Hydrothermal Mounds, Chimneys, and Microbial Mats: Complex Seafloor Expressions of Subsurface Hydrothermal Circulation. Front. Microbiol. 2016, 7, 75. doi:10.3389/fmicb.2016.00075.

[2] Buongiorno J, Herbert LC, Wehrmann LM, Michaud AB, Laufer K, Røy H, et al. Complex Microbial Communities Drive Iron and Sulfur Cycling in Arctic Fjord Sediments. Appl. Environ. Microbiol. 2019, 85, e00949-19. doi:10.1128/AEM.00949-19.

[3] Hadland N, Hamilton CW, Duhamel S. Young volcanic terrains are windows into early microbial colonization. Commun. Earth Environ. 2024, 5, 114. doi:10.1038/s43247-024-01280-3.

[4] Reed CJ, Lewis H, Trejo E, Winston V, Evilia C. Protein Adaptations in Archaeal Extremophiles. Archaea 2013, 2013, 373275. doi:10.1155/2013/373275.

[5] Azua-Bustos A, Urrejola C, Vicuña R. Life at the dry edge: Microorganisms of the Atacama Desert. FEBS Lett. 2012, 586, 2939-2945. doi:10.1016/j.febslet.2012.07.025.

[6] Bull AT, Asenjo JA. Microbiology of hyper-arid environments: Recent insights from the Atacama Desert, Chile. Antonie Van Leeuwenhoek 2013, 103, 1173-1179. doi:10.1007/s10482-013-9911-7.

[7] Pavan ME, López NI, Pettinari MJ. Melanin biosynthesis in bacteria, regulation and production perspectives. Appl. Microbiol. Biotechnol. 2019, 104, 1357-1370. doi:10.1007/s00253-019-10245-y.

[8] Stein T. Bacillus subtilis antibiotics: Structures, syntheses and specific functions. Mol. Microbiol. 2005, 56, 845-857. doi:10.1111/j.1365-2958.2005.04587.x.

[9] Sengupta S, Chattopadhyay MK, Grossart HP. The multifaceted roles of antibiotics and antibiotic resistance in nature. Front. Microbiol. 2013, 4, 47. doi:10.3389/fmicb.2013.00047.

[10] Hall-Stoodley L, Costerton JW, Stoodley P. Bacterial biofilms: From the Natural environment to infectious diseases. Nat. Rev. Microbiol. 2004, 2, 95-108. doi:10.1038/nrmicro821.

[11] Pandit A, Adholeya A, Cahill D, Brau L, Kochar M. Microbial biofilms in nature: Unlocking their potential for agricultural applications. J. Appl. Microbiol. 2020, 129, 199-211. doi:10.1111/jam.14609.

[12] Vlamakis H, Chai Y, Beauregard P, Losick R, Kolter R. Sticking together: Building a biofilm the Bacillus subtilis way. Nat. Rev. Microbiol. 2013, 11, 157-168. doi:10.1038/nrmicro2960.

[13] de Carvalho CCCR. Biofilms:Microbial Strategies for Surviving UV Exposure. In Ultraviolet Light in Human Health, Diseases and Environment; Ahmad SI, Ed.; Springer International Publishing: Cham, Switzerland, 2017; pp. 233-239. doi:10.1007/978-3-319-56017-5_19.

[14] Hall CW, Mah TF. Molecular mechanisms of biofilm-based antibiotic resistance and tolerance in pathogenic bacteria. FEMS Microbiol. Rev. 2017, 41, 276-301. doi:10.1093/femsre/fux010.

[15] Gerardin Y, Springer M, Kishony R. A competitive trade-off limits the selective advantage of increased antibiotic production. Nat. Microbiol. 2016, 1, 16175. doi:10.1038/nmicrobiol.2016.175.

[16] Ferri M, Ranucci E, Romagnoli P, Giaccone V. Antimicrobial resistance: A global emerging threat to public health systems. Crit. Rev. Food Sci. Nutr. 2015, 57, 2857-2876. doi:10.1080/10408398.2015.1077192.

[17] Lewis K, Caboni M. The Making of a Pathogen. Cell Host Microbe 2017, 21, 653-654. doi:10.1016/j.chom.2017.06.001.

[18] Kool ET. Hydrogen Bonding, Base Stacking, and Steric Effects in DNA Replication. Annu. Rev. Biophys. Biomol. Struct. 2001, 30, 1-22. doi:10.1146/annurev.biophys.30.1.1.

[19] Yakovchuk P, Protozanova E, Frank-Kamenetskii MD. Base-stacking and base-pairing contributions into thermal stability of the DNA double helix. Nucleic Acids Res. 2006, 34, 564-574. doi:10.1093/nar/gkj454.

[20] Šmarda P, Bureš P, Horová L, Leitch IJ, Mucina L, Pacini E, et al. Ecological and evolutionary significance of genomic GC content diversity in monocots. Proc. Natl. Acad. Sci. USA 2014, 111, E4096-E4102. doi:10.1073/pnas.1321152111.

[21] Beaume N, Pathak R, Yadav VK, Kota S, Misra HS, Gautam HK, et al. Genome-wide study predicts promoter-G4 DNA motifs regulate selective functions in bacteria: Radioresistance of D. radiodurans involves G 4 DNA-mediated regulation. Nucleic Acids Res. 2013, 41, 76-89. doi:10.1093/nar/gks1071.

[22] Yadav P, Kim N, Kumari M, Verma S, Sharma TK, Yadav V, et al. G-Quadruplex Structures in Bacteria: Biological Relevance and Potential as an Antimicrobial Target. J. Bacteriol. 2021, 203, 10-1128. doi:10.1128/JB.00577-20.

[23] Bull AT, Andrews BA, Dorador C, Goodfellow M. Introducing the Atacama Desert. Antonie Van Leeuwenhoek 2018, 111, 1269-1272. doi:10.1007/s10482-018-1100-2.

[24] Veblen TT, Young KR, Orme AR. The Physical Geography of South America; Oxford University Press: Oxford, UK, 2015; p. 751.

[25] Cordero RR, Seckmeyer G, Damiani A, Riechelmann S, Rayas J, Labbe F, et al. The world’s highest levels of surface UV. Photochem. Photobiol. Sci. 2013, 13, 70-81. doi:10.1039/c3pp50221j.

[26] Cordero RR, Damiani A, Jorquera J, Sepúlveda E, Caballero M, Fernandez S, et al. Ultraviolet radiation in the Atacama Desert. Antonie Van Leeuwenhoek 2018, 111, 1301-1313. doi:10.1007/s10482-018-1075-z.

[27] Stoker CR, Glass BJ, Stucky TR, Dave AI, Kobayashi LT, Quinn RC, et al. A Mission Simulating the Search for Life on Mars with Automated Drilling, Sample Handling, and Life Detection Instruments Performed in the Hyperarid Core of the Atacama Desert, Chile. Astrobiology 2023, 23, 1284-1302. doi:10.1089/ast.2022.0055.

[28] Reverdy A, Hathaway D, Jha J, Michaels G, Sullivan J, McAdoo DD, et al. Insights into the diversity and survival strategies of soil bacterial isolates from the Atacama Desert. Front. Microbiol. 2024, 15, 1335989. doi:10.3389/fmicb.2024.1335989.

[29] Angelini LL, dos Santos RAC, Fox G, Paruthiyil S, Gozzi K, Shemesh M, et al. Pulcherrimin protects Bacillus subtilis against oxidative stress during biofilm development. NPJ Biofilms Microbiomes 2023, 9, 50. doi:10.1038/s41522-023-00418-z.

[30] Schöner TA, Gassel S, Osawa A, Tobias NJ, Okuno Y, Sakakibara Y, et al. Aryl Polyenes, a Highly Abundant Class of Bacterial Natural Products, Are Functionally Related to Antioxidative Carotenoids. ChemBioChem 2016, 17, 247-253. doi:10.1002/cbic.201500474.

[31] Callahan BJ, McMurdie PJ, Rosen MJ, Han AW, Johnson AJA, Holmes SP. DADA2: High-resolution sample inference from Illumina amplicon data. Nat. Methods 2016, 13, 581-583. doi:10.1038/nmeth.3869.

[32] Huse SM, Mark Welch DB, Voorhis A, Shipunova A, Morrison HG, Eren AM, et al. VAMPS: A website for visualization and analysis of microbial population structures. BMC Bioinform. 2014, 15, 41. doi:10.1186/1471-2105-15-41.

[33] Shemesh M, Chai Y. A Combination of Glycerol and Manganese Promotes Biofilm Formation in Bacillus subtilis via Histidine Kinase KinD Signaling. J. Bacteriol. 2013, 195, 2747-2754. doi:10.1128/JB.00028-13.

[34] Cavanaugh NT, Kumar G, Couto Frignani M, Thewedros N, Twahirwa M, Riquelme C, et al. Whole-genome sequencing of three extremophile Bacillus sp. strains isolated from the Atacama Desert. Microbiol. Resour. Announc. 2025, 14, e00679-24. doi:10.1128/mra.00679-24.

[35] Wick R. rrwick/Filtlong. 2025. Available online: https://github.com/rrwick/Filtlong (accessed on 14 August 2025).

[36] Li H. Minimap and miniasm: Fast mapping and de novo assembly for noisy long sequences. Bioinformatics 2016, 32, 2103-2110. doi:10.1093/bioinformatics/btw152.

[37] Kolmogorov M, Yuan J, Lin Y, Pevzner PA. Assembly of long, error-prone reads using repeat graphs. Nat. Biotechnol. 2019, 37, 540-546. doi:10.1038/s41587-019-0072-8.

[38] Pourmohsenin B, Wiese A, Ziemert N. AutoMLST2: A web server for phylogeny and microbial taxonomy. Nucleic Acids Res. 2025, 53, W45-W50. doi:10.1093/nar/gkaf397.

[39] Parks DH, Chuvochina M, Rinke C, Mussig AJ, Chaumeil PA, Hugenholtz P. GTDB: An ongoing census of bacterial and archaeal diversity through a phylogenetically consistent, rank normalized and complete genome-based taxonomy. Nucleic Acids Res. 2022, 50, D785-D794. doi:10.1093/nar/gkab776.

[40] Blin K, Shaw S, Vader L, Szenei J, Reitz ZL, Augustijn HE, et al. antiSMASH 8.0: Extended gene cluster detection capabilities and analyses of chemistry, enzymology, and regulation. Nucleic Acids Res. 2025, 53, W32-W38. doi:10.1093/nar/gkaf334.

[41] Buchfink B, Xie C, Huson DH. Fast and sensitive protein alignment using DIAMOND. Nat. Methods 2014, 12, 59-60. doi:10.1038/nmeth.3176.

[42] Brázda V, Kolomazník J, Lýsek J, Bartas M, Fojta M, Šťastný J, et al. G4Hunter web application: A web server for G-quadruplex prediction. Bioinformatics 2019, 35, 3493-3495. doi:10.1093/bioinformatics/btz087.

[43] Schulze-Makuch D, Wagner D, Kounaves SP, Mangelsdorf K, Devine KG, de Vera JP, et al. Transitory microbial habitat in the hyperarid Atacama Desert. Proc. Natl. Acad. Sci. USA 2018, 115, 2670-2675. doi:10.1073/pnas.1714341115.

[44] Demergasso C, Escudero L, Casamayor EO, Chong G, Balagué V, Pedrós-Alió C. Novelty and spatio-temporal heterogeneity in the bacterial diversity of hypersaline Lake Tebenquiche (Salar de Atacama). Extremophiles 2008, 12, 491-504. doi:10.1007/s00792-008-0153-y.

[45] Aguayo P, González P, Campos V, Maugeri TL, Papale M, Gugliandolo C, et al. Comparison of Prokaryotic Diversity in Cold, Oligotrophic Remote Lakes of Chilean Patagonia. Curr. Microbiol. 2017, 74, 598-613. doi:10.1007/s00284-017-1209-y.

[46] Demergasso C, Dorador C, Meneses D, Blamey J, Cabrol N, Escudero L, et al. Prokaryotic diversity pattern in high-altitude ecosystems of the Chilean Altiplano. J. Geophys. Res. Biogeosci. 2010, 115. doi:10.1029/2008JG000836.

[47] Mandakovic D, Maldonado J, Pulgar R, Cabrera P, Gaete A, Urtuvia V, et al. Microbiome analysis and bacterial isolation from Lejía Lake soil in Atacama Desert. Extremophiles 2018, 22, 665-673. doi:10.1007/s00792-018-1027-6.

[48] Dorador C, Fink P, Hengst M, Icaza G, Villalobos AS, Vejar D, et al. Microbial community composition and trophic role along a marked salinity gradient in Laguna Puilar, Salar de Atacama, Chile. Antonie Van Leeuwenhoek 2018, 111, 1361-1374. doi:10.1007/s10482-018-1091-z.

[49] Bates ST, Berg-Lyons D, Caporaso JG, Walters WA, Knight R, Fierer N. Examining the global distribution of dominant archaeal populations in soil. ISME J. 2011, 5, 908-917. doi:10.1038/ismej.2010.171.

[50] Dorador C, Vila I, Remonsellez F, Imhoff JF, Witzel KP. Unique clusters of Archaea in Salar de Huasco, an athalassohaline evaporitic basin of the Chilean Altiplano. FEMS Microbiol. Ecol. 2010, 73, 291-302. doi:10.1111/j.1574-6941.2010.00891.x.

[51] Ondov BD, Treangen TJ, Melsted P, Mallonee AB, Bergman NH, Koren S, et al. Mash: Fast genome and metagenome distance estimation using MinHash. Genome Biol. 2016, 17, 132. doi:10.1186/s13059-016-0997-x.

[52] Lutgring JD, Machado MJ, Benahmed FH, Conville P, Shawar RM, Patel J, et al. FDA-CDC Antimicrobial Resistance Isolate Bank: A Publicly Available Resource To Support Research, Development, and Regulatory Requirements. J. Clin. Microbiol. 2018, 56, 10-1128. doi:10.1128/JCM.01415-17.

[53] Nye TM, Schroeder JW, Kearns DB, Simmons LA. Complete Genome Sequence of Undomesticated Bacillus subtilis Strain NCIB 3610. Genome Announc. 2017, 5, 10-1128. doi:10.1128/genomeA.00364-17.

[54] Connon SA, Lester ED, Shafaat HS, Obenhuber DC, Ponce A. Bacterial diversity in hyperarid Atacama Desert soils. J. Geophys. Res. Biogeosci. 2007, 112. doi:10.1029/2006JG000311.

[55] Kerrigan Z, Kirkpatrick JB, D’Hondt S. Influence of 16S rRNA Hypervariable Region on Estimates of Bacterial Diversity and Community Composition in Seawater and Marine Sediment. Front. Microbiol. 2019, 10, 1640. doi:10.3389/fmicb.2019.01640.

[56] George PBL, Lallias D, Creer S, Seaton FM, Kenny JG, Eccles RM, et al. Divergent national-scale trends of microbial and animal biodiversity revealed across diverse temperate soil ecosystems. Nat. Commun. 2019, 10, 1107. doi:10.1038/s41467-019-09031-1.

[57] Wagner R, Zona D, Oechel W, Lipson D. Microbial community structure and soil pH correspond to methane production in Arctic Alaska soils. Environ. Microbiol. 2017, 19, 3398-3410. doi:10.1111/1462-2920.13854.

[58] Alanjary M, Steinke K, Ziemert N. AutoMLST: An automated web server for generating multi-locus species trees highlighting natural product potential. Nucleic Acids Res. 2019, 47, W276-W282. doi:10.1093/nar/gkz282.

[59] Rooney AP, Price NPJ, Ehrhardt C, Swezey JL, Bannan JD. Phylogeny and molecular taxonomy of the Bacillus subtilis species complex and description of Bacillus subtilis subsp. inaquosorum subsp. nov. Int. J. Syst. Evol. Microbiol. 2009, 59, 2429-2436. doi:10.1099/ijs.0.009126-0.

[60] Roberts MS,Nakamura LK, Cohan FM. Bacillus vallismortis sp. nov., a Close Relative of Bacillus subtilis, Isolated from Soil in Death Valley, California. Int. J. Syst. Bacteriol. 1996, 46, 470-475. doi:10.1099/00207713-46-2-470.

[61] Hartley AJ, Chong G, Houston J, Mather AE. 150 million years of climatic stability: Evidence from the Atacama Desert, northern Chile. J. Geol. Soc. 2022, 162, 421-424. doi:10.1144/0016-764904-071.

[62] Houston J, Hartley AJ. The central Andean west-slope rainshadow and its potential contribution to the origin of hyper-aridity in the Atacama Desert. Int. J. Climatol. 2003, 23, 1453-1464. doi:10.1002/joc.938.

[63] Navarro-González R, Rainey FA, Molina P, Bagaley DR, Hollen BJ, de la Rosa J, et al. Mars-Like Soils in the Atacama Desert, Chile, and the Dry Limit of Microbial Life. Science 2003, 302, 1018-1021. doi:10.1126/science.1089143.

[64] Li CJ, Jiang ZM, Zhi XY, Chen HH, Yu LY, Li GF, et al. Genomic insights into Kocuria: Taxonomic revision and identification of five IAA-producing extremophiles. Front. Microbiol. 2025, 16, 1547983. doi:10.3389/fmicb.2025.1547983.

[65] Azua-Bustos A, González-Silva C, Fernández-Martínez MÁ, Arenas-Fajardo C, Fonseca R, Martín-Torres FJ, et al. Aeolian transport of viable microbial life across the Atacama Desert, Chile: Implications for Mars. Sci. Rep. 2019, 9, 11024. doi:10.1038/s41598-019-47394-z.

[66] Ghssein G, Ezzeddine Z. A Review of Pseudomonas aeruginosa Metallophores: Pyoverdine, Pyochelin and Pseudopaline. Biology 2022, 11, 1711. doi:10.3390/biology11121711.

[67] Hassan HM, Fridovich I. Mechanism of the antibiotic action pyocyanine. J. Bacteriol. 1980, 141, 156-163. doi:10.1128/jb.141.1.156-163.1980.

[68] Jayaseelan S, Ramaswamy D, Dharmaraj S. Pyocyanin: Production, applications, challenges and new insights. World J. Microbiol. Biotechnol. 2013, 30, 1159-1168. doi:10.1007/s11274-013-1552-5.

[69] Dumas Z, Ross-Gillespie A, Kümmerli R. Switching between apparently redundant iron-uptake mechanisms benefits bacteria in changeable environments. Proc. R. Soc. B Biol. Sci. 2013, 280, 20131055. doi:10.1098/rspb.2013.1055.

[70] Visca P, Imperi F, Lamont IL. Pyoverdine siderophores: From biogenesis to biosignificance. Trends Microbiol. 2007, 15, 22-30. doi:10.1016/j.tim.2006.11.004.

[71] Ernst S, Volkov AN, Stark M, Hölscher L, Steinert K, Fetzner S, et al. Azetidomonamide and Diazetidomonapyridone Metabolites Control Biofilm Formation and Pigment Synthesis in Pseudomonas aeruginosa. J. Am. Chem. Soc. 2022, 144, 7676-7685. doi:10.1021/jacs.1c13653.

[72] Tamehiro N, Okamoto-Hosoya Y, Okamoto S, Ubukata M, Hamada M, Naganawa H, et al. Bacilysocin, a Novel Phospholipid Antibiotic Produced by Bacillus subtilis 168. Antimicrob. Agents Chemother. 2002, 46, 315-320. doi:10.1128/AAC.46.2.315-320.2002.

[73] Chakraborty K, Kizhakkekalam VK, Joy M, Chakraborty RD. Bacillibactin class of siderophore antibiotics from a marine symbiotic Bacillus as promising antibacterial agents. Appl. Microbiol. Biotechnol. 2021, 106, 329-340. doi:10.1007/s00253-021-11632-0.

[74] Geissler M, Heravi KM, Henkel M, Hausmann R. Lipopeptide Biosurfactants from Bacillus Species. In Biobased Surfactants, 2nd ed.; Hayes DG, Solaiman DKY, Ashby RD,Eds.; AOCS Press:Champaign, IL, USA, 2019; pp. 205-240. Available online: https://www.sciencedirect.com/science/article/pii/B978012812705600006X. (accessed on 14 August 2025).

[75] Butcher RA, Schroeder FC, Fischbach MA, Straight PD, Kolter R, Walsh CT, et al. The identification of bacillaene, the product of the PksX megacomplex in Bacillus subtilis. Proc. Natl. Acad. Sci. USA 2007, 104, 1506-1509. doi:10.1073/pnas.0610503104.

[76] Babasaki K, Takao T, Shimonishi Y, Kurahashi K. Subtilosin A, a New Antibiotic Peptide Produced by Bacillus subtilis 168: Isolation, Structural Analysis, and Biogenesis1. J. Biochem. 1985, 98, 585-603. doi:10.1093/oxfordjournals.jbchem.a135315.

[77] Stannius RO, Kovács ÁT. Plipastatin is a shared good by Bacillus subtilis during combating Fusarium spp. FEMS Microbiol. Ecol. 2025, 101, fiaf020. doi:10.1093/femsec/fiaf020.

[78] O’Connor EB, Cotter PD, O’Connor P, O’Sullivan O, Tagg JR, Ross RP, et al. Relatedness between the two-component lantibiotics lacticin 3147 and staphylococcin C55 based on structure, genetics and biological activity. BMC Microbiol. 2007, 7, 24. doi:10.1186/1471-2180-7-24.

[79] Schalk IJ. Bacterial siderophores: Diversity, uptake pathways and applications. Nat. Rev. Microbiol. 2024, 23, 24-40. doi:10.1038/s41579-024-01090-6.

[80] Rojas Murcia N, Lee X, Waridel P, Maspoli A, Imker HJ, Chai T, et al. The Pseudomonas aeruginosa antimetabolite L -2-amino-4-methoxy-trans-3-butenoic acid (AMB) is made from glutamate and two alanine residues via a thiotemplate-linked tripeptide precursor. Front. Microbiol. 2015, 6, 170. doi:10.3389/fmicb.2015.00170.

[81] Blumer C, Haas D. Mechanism, regulation, and ecological role of bacterial cyanide biosynthesis. Arch. Microbiol. 2000, 173, 170-177. doi:10.1007/s002039900127.

[82] Tymiak AA, Culver CA, Malley MF, Gougoutas JZ. Structure of obafluorin: An antibacterial.beta.-lactone from Pseudomonas fluorescens. J. Org. Chem. 2002, 50, 5491-5495. doi:10.1021/jo00350a010.

[83] Schimming O, Fleischhacker F, Nollmann FI, Bode HB. Yeast Homologous Recombination Cloning Leading to the Novel Peptides Ambactin and Xenolindicin. ChemBioChem 2014, 15, 1290-1294. doi:10.1002/cbic.201402065.

[84] Asnani A, Amaliyah R, Yuniaty A. Screening Anti-MRSA Activities of Indigenous Microbes and Prediction of The Biosynthetic Gene Clusters. J. Phys. Conf. Ser. 2020, 1665, 012001. doi:10.1088/1742-6596/1665/1/012001.