Polyhydroxybutyrate (PHB) production from halophiles: a comprehensive review

Nadana Raja Vadivu Ganapathy , Sakshi Singh , Shivansh Ranawat , Navya Hada , Mugesh Sankaranarayanan , Punniyakotti Parthipan

Bioresources and Bioprocessing ›› 2026, Vol. 13 ›› Issue (1) : 127

PDF
Bioresources and Bioprocessing ›› 2026, Vol. 13 ›› Issue (1) :127 DOI: 10.1186/s40643-026-01119-z
Review
review-article
Polyhydroxybutyrate (PHB) production from halophiles: a comprehensive review
Author information +
History +
PDF

Abstract

The growing environmental impact of petroleum-based plastics has intensified the search for sustainable and biodegradable alternatives. Polyhydroxybutyrate (PHB), a microbial polyester from the polyhydroxyalkanoate (PHA) family, has emerged as a promising biopolymer due to its biodegradability, biocompatibility, and thermoplastic qualities that rival those of conventional polymers. Halophilic bacteria have attracted considerable attention among PHB-producing microorganisms because they thrive in hypersaline environments, enabling non-sterile cultivation, reducing contamination risks, and facilitating cost-effective downstream recovery by osmotic cell lysis. This review provides a comprehensive overview of recent improvements in PHB production by halophilic bacteria, covering physiological adaptations, metabolic pathways, substrate usage, fermentation techniques, and bioreactor optimisation. Particular emphasis is placed on the use of agro-industrial residues and waste-derived feedstocks as sustainable carbon sources to reduce production costs and increase circular bioeconomy results. Recent advances in downstream processing, such as green extraction technologies, metabolic engineering, CRISPR-based gene editing, and synthetic biology approaches to increasing PHB productivity, are critically reviewed. Additionally, developments in polymer modification, life cycle assessment, industrial scalability, regulatory frameworks, and potential applications in packaging, agriculture, and biomedical engineering are discussed. Despite significant progress, issues such as process economics, saline wastewater control, polymer brittleness, and large-scale commercialisation remain. Integrating halophilic biotechnology with waste valorisation, green recovery technologies, sophisticated metabolic engineering, and circular biorefinery concepts offers a promising strategy for developing economically and environmentally sustainable PHB production systems.

Graphical abstract

Keywords

Circular economy / Waste reduction / Sustainability / Environmental conservation

Cite this article

Download citation ▾
Nadana Raja Vadivu Ganapathy, Sakshi Singh, Shivansh Ranawat, Navya Hada, Mugesh Sankaranarayanan, Punniyakotti Parthipan. Polyhydroxybutyrate (PHB) production from halophiles: a comprehensive review. Bioresources and Bioprocessing, 2026, 13 (1) : 127 DOI:10.1186/s40643-026-01119-z

登录浏览全文

4963

注册一个新账户 忘记密码

References

[1]

Abate T, Amabile C, De Crescenzo C, Migliaccio A, Capece G, Ciampa R, et al.. A Simulation Study for Comparing Halogenated and Green Solvents during the Extraction of Poly(3-Hydroxybutyrate). Chem Eng Trans Italian Association Chem Eng - AIDIC, 2023, 98: 183-188

[2]

Abdelrahman SA, Barakat OS, Ahmed MN (2024) Genetic characterization of a novel Salinicola salarius isolate applied for the bioconversion of agro-industrial wastes into polyhydroxybutyrate. Microb Cell Fact 23. https://doi.org/10.1186/s12934-024-02326-z

[3]

Adnan M, Siddiqui AJ, Ashraf SA, Snoussi M, Badraoui R, Alreshidi M et al (2022) Polyhydroxybutyrate (PHB)-based biodegradable polymer from agromyces indicus: enhanced production, characterization, and optimization. Polymers (Basel) 14. https://doi.org/10.3390/polym14193982

[4]

Aizaz M, Lubna, Ahmad W, Khan I, Asaf S, Bilal S et al (2023) Exploring the potential of halotolerant bacteria from coastal regions to mitigate salinity stress in wheat: physiological, molecular, and biochemical insights. Front Plant Sci 14. https://doi.org/10.3389/fpls.2023.1224731

[5]

Al Rawahi AM, Zafar M, Khan TA, Al Araimi S, Mahanty B, Behera SK. Genetic algorithm-optimized artificial neural network for multi-objective optimization of biomass and exopolysaccharide production by Haloferax mediterranei. Bioprocess Biosyst Eng [Internet], 2025, 48: 785-798

[6]

Alsafadi D, Aljariri Alhesan JS, Mansoura A, Oqdeha S. Production of polyhydroxyalkanoate from sesame seed wastewater by sequencing batch reactor cultivation process of Haloferax mediterranei. Arab J Chem, 2023, 16: 104584

[7]

Andler R, González-Arancibia F, Vilos C, Sepulveda-Verdugo R, Castro R, Mamani M, et al.. Production of poly-3-hydroxybutyrate (PHB) nanoparticles using grape residues as the sole carbon source. Int J Biol Macromol [Internet], 2024, 261: 129649

[8]

Ansari S, Sami N, Yasin D, Ahmad N, Fatma T (2021) Biomedical applications of environmental friendly poly-hydroxyalkanoates. Int J Biol Macromol 549–563. https://doi.org/10.1016/j.ijbiomac.2021.04.171

[9]

Aramvash A, Gholami-Banadkuki N, Moazzeni-Zavareh F, Hajizadeh-Turchi S. An environmentally friendly and efficient method for extraction of PHB biopolymer with non-halogenated solvents. J Microbiol Biotechnol Korean Soc Microbiolog Biotechnol, 2015, 25: 1936-1943

[10]

Aransiola SA, Victor-Ekwebelem MO, Daza BX, Oladoye PO, Alli YA, Bamisaye A et al (2025) Micro- and nano-plastics pollution in the marine environment: progresses, drawbacks and future guidelines. Chemosphere. https://doi.org/10.1016/j.chemosphere.2025.144211

[11]

Arcila-Echavarría DC, Lu-Chau TA, Gómez-Vanegas NA (2025) Polyhydroxybutyrate synthesis by the halophilic bacterium, halomonas boliviensis, in oil palm empty fruit bunch hydrolysate. Biopolymers 116:e23644. https://doi.org/10.1002/bip.23644

[12]

Balan L, Mohandas SP, Priyaja P, Gopi J, Cubelio SS, Philip R et al (2025) Development of a mini-repository of marine bacteria having the potential of Polyhydroxyalkanoates production. Microbe (Netherlands) 7. https://doi.org/10.1016/j.microb.2025.100338

[13]

Bhaskar S, Steer DL, Anand R, Panjikar S (2020) Structural basis for differentiation between two classes of thiolase: degradative vs biosynthetic thiolase. J Struct Biol 4. https://doi.org/10.1016/j.yjsbx.2019.100018

[14]

Bhat GS, Deekshitha BK, Thivaharan V, Divyashree MS (2024) Physicochemical cell disruption of Bacillus sp. for recovery of polyhydroxyalkanoates: future bioplastic for sustainability. 3 Biotech 14. https://doi.org/10.1007/s13205-024-03913-y

[15]

Boy C, Lesage J, Alfenore S, Guillouet SE, Gorret N (2021) Investigation of the robustness of Cupriavidus necator engineered strains during fed-batch cultures. AMB Express 11. https://doi.org/10.1186/s13568-021-01307-4

[16]

Branca C, Fabrizi F, Mghili B, Conti-Nibali V, Gunasekaran K, Bottari T et al (2025) Plastic pollution in a special protected area for migratory birds. Sci Total Environ 958. https://doi.org/10.1016/j.scitotenv.2024.177918

[17]

Bugnicourt E, Cinelli P, Lazzeri A, Alvarez V. Polyhydroxyalkanoate (PHA): Review of synthesis, characteristics, processing and potential applications in packaging. Express Polym Lett BME-PT GTE, 2014, 8: 791-808

[18]

Carlozzi P, Di Lorenzo T, Ghanotakis DF, Touloupakis E. Effects of pH, temperature and salinity on P3HB synthesis culturing the marine Rhodovulum sulfidophilum DSM-1374. Appl Microbiol Biotechnol Springer, 2020, 104: 2007-2015

[19]

Chathalingath N, Kingsly JS, Gunasekar A (2023) Biosynthesis and biodegradation of poly(3-hydroxybutyrate) from Priestia flexa; a promising mangrove halophyte towards the development of sustainable eco-friendly bioplastics. Microbiol Res 267. https://doi.org/10.1016/j.micres.2022.127270

[20]

Chauhan V, Chinmay, Bendi AIslam S, Shahid M. Advances in Polymer Nanocomposites for Active Food Packing. Green Materials for Active Food Packaging [Internet], 2025, Singapore, Springer Nature Singapore, 229-251

[21]

Chen Q, Zhang LHApplied Mechanics and Materials [Internet]2014

[22]

Chen Q, Zhang LH (2014) Study on synthesis of PHB by moderate halophile and aqueous extraction of PHB. Appl Mech Mater 160–163. https://doi.org/10.4028/www.scientific.net/AMM.448-453.160

[23]

Chen D-D, Fang B-Z, Manzoor A, Liu Y-H, Li L, Mohamad OAA, et al.. Revealing the salinity adaptation mechanism in halotolerant bacterium Egicoccus halophilus EGI 80432T by physiological analysis and comparative transcriptomics. Appl Microbiol Biotechnol [Internet], 2021, 105: 2497-2511

[24]

Choi J, Lee SY (1999) Factors affecting the economics of polyhydroxyalkanoate production by bacterial fermentation. Appl Microbiol Biotechnol 51(1):13–21

[25]

Coimbra AAB, Prakash S, Jiménez JI, Rios-Solis L. Establishing Halomonas as a chassis for industrial biotechnology: advances in synthetic biology tool development and metabolic engineering strategies. Microb Cell Fact BioMed Cent Ltd, 2025

[26]

Cristea A, Baricz A, Leopold N, Floare CG, Borodi G, Kacso I, et al.. Polyhydroxybutyrate production by an extremely halotolerant Halomonas elongata strain isolated from the hypersaline meromictic Fără Fund Lake (Transylvanian Basin, Romania). J Appl Microbiol John Wiley Sons Inc, 2018, 125: 1343-1357

[27]

Cristea A, Pustan M, Bîrleanu C, Dudescu C, Floare CG, Tripon AM, et al.. Mechanical Evaluation of Solvent Casted Poly(3-hydroxybutyrate) Films Derived from the Storage Polyesters Produced by Halomonas elongata DSM 2581T. J Polym Environ Springer, 2022, 30: 424-430

[28]

Das SK, Eshkalak SK, Chinnappan A, Ghosh R, Jayathilaka WADM, Baskar C et al (2021) Plastic recycling of polyethylene terephthalate (PET) and polyhydroxybutyrate (PHB)—a comprehensive review. Mat Circular Econ 3. https://doi.org/10.1007/s42824-021-00025-3

[29]

De Eugenio LI, Galán B, Escapa IF, Maestro B, Sanz JM, García JL, et al.. The PhaD regulator controls the simultaneous expression of the pha genes involved in polyhydroxyalkanoate metabolism and turnover in Pseudomonas putida KT2442. Environ Microbiol, 2010, 12: 1591-1603

[30]

Deantas-Jahn C, Mendoza SN, Licona-Cassani C, Orellana C, Saa PA (2024) Metabolic modeling of Halomonas campaniensis improves polyhydroxybutyrate production under nitrogen limitation. Appl Microbiol Biotechnol 108. https://doi.org/10.1007/s00253-024-13111-8

[31]

Delamarre SC (2004) Engineering of biological systems for the production of Polyhydroxyalkanoate copolymers and the biofabrication of polymeric microstructures. Cornell University, New York. https://books.google.co.in/books?id=P9dPAAAAYAAJ

[32]

Dubey S, Mishra S (2021) Efficient production of polyhydroxyalkanoate through halophilic bacteria utilizing algal biodiesel waste residue. Front Bioeng Biotechnol 9. https://doi.org/10.3389/fbioe.2021.624859

[33]

Esposito FP, Vecchiato V, Buonocore C, Tedesco P, Noble B, Basnett P et al (2023) Enhanced production of biobased, biodegradable, Poly(3-hydroxybutyrate) using an unexplored marine bacterium Pseudohalocynthiibacter aestuariivivens, isolated from highly polluted coastal environment. Bioresour Technol 368. https://doi.org/10.1016/j.biortech.2022.128287

[34]

Faruga A, Cichoń E, Karcz R, Kryściak-Czerwenka J, Szumera M, Prajsnar J et al (2025) Integrated chemo-biotechnological process for upcycling polyesters into new PHB. Biotechnol Environ 2. https://doi.org/10.1186/s44314-025-00028-3

[35]

Faulkner M, Hoeven R, Kelly PP, Sun Y, Park H, Liu LN et al (2023) Chemoautotrophic production of gaseous hydrocarbons, bioplastics and osmolytes by a novel Halomonas species. Biotechnol Biofuels Bioproducts 16. https://doi.org/10.1186/s13068-023-02404-1

[36]

Fei T, Cazeneuve S, Wen Z, Wu L, Wang T. Effective recovery of poly-β-hydroxybutyrate (PHB) biopolymer from Cupriavidus necator using a novel and environmentally friendly solvent system. Biotechnol Prog John Wiley Sons Inc, 2016, 32: 678-685

[37]

Feranc J, Repiská M, Plavec R, Tomanová K, Ďurfina M, Vanovčanová Z et al (2025) Biodegradable PLA/PHB composites with inorganic fillers and modifiers. Polymers (Basel) 17. https://doi.org/10.3390/polym17202721

[38]

Fogašová M, Figalla S, Danišová L, Medlenová E, Hlaváčiková S, Vanovčanová Z et al (2022) PLA/PHB-based materials fully biodegradable under both industrial and home-composting conditions. Polymers (Basel) 14. https://doi.org/10.3390/polym14194113

[39]

Frehner A, Itten R, Stucki M Life cycle assessment of bio-based materials—environmental impacts of the value chain from cyanobacteria to PHB. https://www.zhaw.ch/iunr/lca

[40]

Fuentes del Carmen D, Soto-Urzua L, Martínez-Soto LJ, Martínez-Morales LJ (2024) Bioinformatic and functional analysis of a PHB polymerase (PhbC) from Azospirillum baldaniorum. J Genetic Eng Biotechnol 22. https://doi.org/10.1016/j.jgeb.2024.100403

[41]

Gasparyan KG, Tyubaeva PM, Varyan IA, Vetcher AA, Popov AA (2023) Assessing the biodegradability of PHB-based materials with different surface areas: a comparative study on soil exposure of films and electrospun materials. Polymers (Basel) 15. https://doi.org/10.3390/polym15092042

[42]

Getino L, Martín JL, Chamizo-Ampudia A (2024) A review of Polyhydroxyalkanoates: characterization, production, and application from waste. Microorganisms. https://doi.org/10.3390/microorganisms12102028

[43]

Hammami K, Souissi Y, Souii A, Ouertani A, El-Hidri D, Jabberi M et al (2022) Extremophilic bacterium halomonas desertis G11 as a cell factory for Poly-3-hydroxybutyrate-co-3-hydroxyvalerate copolymer’s production. Front Bioeng Biotechnol 10. https://doi.org/10.3389/fbioe.2022.878843

[44]

Harding KG, Dennis JS, von Blottnitz H, Harrison STL. Environmental analysis of plastic production processes: Comparing petroleum-based polypropylene and polyethylene with biologically-based poly-β-hydroxybutyric acid using life cycle analysis. J Biotechnol, 2007, 130: 57-66

[45]

Hezayen FF, Rehm BHA, Eberhardt R, Steinbüchel A. Polymer production by two newly isolated extremely halophilic archaea: application of a novel corrosion-resistant bioreactor. Appl Microbiol Biotechnol [Internet], 2000, 54: 319-325

[46]

Jia L, Kaur G, Juneja A, Majumder EL-W, Ramarao BV, Kumar D (2024) Polyhydroxybutyrate production from non-recyclable fiber rejects and acid whey as mixed substrate by recombinant Escherichia coli. Biotechnol Sustain Mat 1. https://doi.org/10.1186/s44316-024-00013-y

[47]

Jung HJ, Kim SH, Cho DH, Kim BC, Bhatia SK, Lee J, Jeon J-M, Yoon J-J, Yang Y-H. Finding of Novel Galactose Utilizing Halomonas sp. YK44 for Polyhydroxybutyrate (PHB) Production. Polymers, 2022, 14(245407

[48]

Kalia VC, Patel SKS, Lee JKMultidisciplinary Digital Publishing Institute (MDPI). Exploiting Polyhydroxyalkanoates for Biomedical Applications. Polym (Basel), 2023

[49]

Kalia VC, Patel SKS, Karthikeyan KK, Jeya M, Kim IW, Lee JK (2024) Manipulating microbial cell morphology for the sustainable production of biopolymers. Polymers (Basel). https://doi.org/10.3390/polym16030410

[50]

Khosravi-Darani K, Mokhtari ZB, Amai T, Tanaka K (2013) Microbial production of poly(hydroxybutyrate) from C1 carbon sources. Appl Microbiol Biotechnol 1407–24. https://doi.org/10.1007/s00253-012-4649-0

[51]

Koller M (2022) Advances in Polyhydroxyalkanoate (PHA) Production, Volume 3. Bioengineering. https://doi.org/10.3390/bioengineering9070328

[52]

Koller M, Mukherjee A (2022) Polyhydroxyalkanoates (PHAs)—production, properties, and biodegradation. Biodegrad Polymers Circular Plastics Econ 145–204. https://doi.org/10.1002/9783527827589.ch6

[53]

Kong Y, Koh HG, Cha H-G, Lee BW, Yu K, Park S-H, et al.. Isolation and characterization of two halophilic bacteria producing polyhydroxybutyrate from high-salt environment. Biotechnol Bioprocess Eng [Internet], 2024, 29: 1003-1013

[54]

Kong Y, Koh HG, Cha H-G, Lee BW, Yu K, Park S-H, et al.. Isolation and characterization of two halophilic bacteria producing polyhydroxybutyrate from high-salt environment. Biotechnol Bioprocess Eng [Internet], 2024, 29: 1003-1013

[55]

Kurabi A, Pak K, Chavez E, Doan J, Ryan AF (2022) A transcytotic transport mechanism across the tympanic membrane. Sci Rep 12. https://doi.org/10.1038/s41598-021-04748-w

[56]

Leandro T, Oliveira MC, da Fonseca MMR, Cesário MT (2023) Co-production of Poly(3-hydroxybutyrate) and gluconic acid from glucose by halomonas elongata. Bioeng 10. https://doi.org/10.3390/bioengineering10060643

[57]

Leandro T, Oliveira MC, da Fonseca MMR, Cesário MT (2023) Co-production of Poly(3-hydroxybutyrate) and gluconic acid from glucose by halomonas elongata. Bioeng 10. https://doi.org/10.3390/bioengineering10060643

[58]

Lee MS, Salleh KM. An overview on microalgal polyhydroxybutyrate (PHB) production and improvement of mechanical properties. Int J Biol Macromol Elsevier B V, 2025

[59]

Li D, Yang Y, Liu R, Wu Y, Guo F (2025) Review of biopolymer Polyhydroxybutyrate (PHB) and blends: modification of thermal and mechanical properties via additive manufacturing processing. Polymers (Basel). https://doi.org/10.3390/polym17223083

[60]

Lin YC, Ng IS (2025) Biofabrication of polyhydroxybutyrate (PHB) in engineered Cupriavidus necator H16 from waste molasses. J Taiwan Inst Chem Eng 167. https://doi.org/10.1016/j.jtice.2024.105843

[61]

Lin Z, Zhang Y, Yuan Q, Liu Q, Li Y, Wang Z et al (2015) Metabolic engineering of Escherichia coli for poly(3-hydroxybutyrate) production via threonine bypass. Microb Cell Fact 14. https://doi.org/10.1186/s12934-015-0369-3

[62]

Linekha R, Jeno JGA, Abirami K, Yamunadevi B, Nakkeeran E (2024) Halophiles and their adaptations: a comprehensive review on recent progress and prospects in biodesalination applications. Clean (Weinh) 52:2300260. https://doi.org/10.1002/clen.202300260

[63]

Liu C, Baffoe DK, Zhan Y, Zhang M, Li Y, Zhang G. Halophile, an essential platform for bioproduction. J Microbiol Methods Elsevier B V, 2019

[64]

Liu L, Zhou S, Deng Y (2020) The 3-ketoacyl-CoA thiolase: an engineered enzyme for carbon chain elongation of chemical compounds. Appl Microbiol Biotechnol. https://doi.org/10.1007/s00253-020-10848-w/Published

[65]

Liu R, Liang L, Freed EF, Choudhury A, Eckert CA, Gill RT (2020) Engineering regulatory networks for complex phenotypes in E. coli. Nat Commun Nat Res 11. https://doi.org/10.1038/s41467-020-17721-4

[66]

Liu C, Wang X, Yang H, Liu C, Zhang Z, Chen G. Biodegradable polyhydroxyalkanoates production from wheat straw by recombinant Halomonas elongata A1. Int J Biol Macromol [Internet], 2021, 187: 675-682

[67]

Loan TT, Trang DTQ, Huy PQ, Ninh PX, Van Thuoc D (2022) A fermentation process for the production of poly(3-hydroxybutyrate) using waste cooking oil or waste fish oil as inexpensive carbon substrate. Biotechnol Rep 33. https://doi.org/10.1016/j.btre.2022.e00700

[68]

Longo A, Fanelli F, Villano M, Montemurro M, Rizzello CG (2024) Bioplastic production from agri-food waste through the use of haloferax mediterranei: a comprehensive initial overview. Microorganisms. https://doi.org/10.3390/microorganisms12061038

[69]

Lopez-Arenas T, González-Contreras M, Anaya-Reza O, Sales-Cruz M. Analysis of the fermentation strategy and its impact on the economics of the production process of PHB (polyhydroxybutyrate). Comput Chem Eng [Internet], 2017, 107: 140-150

[70]

Loukas A, Kappas I, Abatzopoulos TJ (2018) HaloDom: a new database of halophiles across all life domains. J Biolog Res. 25. https://doi.org/10.1186/s40709-017-0072-0

[71]

Ma H, Zhao Y, Huang W, Zhang L, Wu F, Ye J et al (2020) Rational flux-tuning of Halomonas bluephagenesis for co-production of bioplastic PHB and ectoine. Nat Commun 11. https://doi.org/10.1038/s41467-020-17223-3

[72]

Mahansaria R, Bhowmik S, Dhara A, Saha A, Mandal M, Ghosh R et al (2020) Production enhancement of poly(3-hydroxybutyrate-co-3-hydroxyvalerate) in Halogeometricum borinquense, characterization of the bioplastic and desalination of the bioreactor effluent. Process Biochem 94. https://doi.org/10.1016/j.procbio.2020.04.004

[73]

Majerczak K, Wadkin-Snaith D, Magueijo V, Mulheran P, Liggat J, Johnston K (2022) Polyhydroxybutyrate: a review of experimental and simulation studies of the effect of fillers on crystallinity and mechanical properties. Polym Int 1398–408. https://doi.org/10.1002/pi.6402

[74]

Manikandan NA, Pakshirajan K, Pugazhenthi G. Preparation and characterization of environmentally safe and highly biodegradable microbial polyhydroxybutyrate (PHB) based graphene nanocomposites for potential food packaging applications. Int J Biol Macromol [Internet], 2020, 154: 866-877

[75]

Markl E (2018) PHB—bio based and biodegradable replacement for PP: a review. Novel Techniq Nutrit Food Sci 2. https://doi.org/10.31031/ntnf.2018.02.000546

[76]

Martínez-García E, Aparicio T, de Lorenzo V, Nikel PI (2014) New transposon tools tailored for metabolic engineering of Gram-negative microbial cell factories. Front Bioeng Biotechnol 2. https://doi.org/10.3389/fbioe.2014.00046

[77]

Martínez-Herrera RE, Rutiaga-Quiñones OM, Alemán-Huerta ME. Integration of Agave plants into the polyhydroxybutyrate (PHB) production: A gift of the ancient Aztecs to the current bioworld. Ind Crops Prod [Internet], 2021, 174: 114188

[78]

Matarredona L, Camacho M, Zafrilla B, Bravo-Barrales G, Esclapez J, Bonete MJ. The survival of haloferax mediterranei under stressful conditions. Microorganisms MDPI AG, 2021, 9: 1-16

[79]

McAdam B, Fournet MB, McDonald P, Mojicevic M (2020) Production of polyhydroxybutyrate (PHB) and factors impacting its chemical and mechanical characteristics. Polymers (Basel) 1–20. https://doi.org/10.3390/polym12122908

[80]

McAdam B, Fournet MB, McDonald P, Mojicevic M (2020) Production of polyhydroxybutyrate (PHB) and factors impacting its chemical and mechanical characteristics. Polymers (Basel) 1–20. https://doi.org/10.3390/polym12122908

[81]

Meereboer KW, Misra M, Mohanty AK (2020) Review of recent advances in the biodegradability of polyhydroxyalkanoate (PHA) bioplastics and their composites. Green Chem 5519–5558. https://doi.org/10.1039/d0gc01647k

[82]

Meng D, Wang S, Zhao K, Luo Y, Li X, Wang Y (2025) Improvement of acetate tolerance of Escherichia coli by introducing the PHB mobilization pathway. Appl Environ Microbiol 91. https://doi.org/10.1128/aem.02454-24

[83]

Miranda DA, Marín K, Sundman O, Hedenström M, Quillaguaman J, Gorzsás A et al (2023) Production and characterization of Poly(3-hydroxybutyrate) from halomonas boliviensis LC1 cultivated in hydrolysates of quinoa stalks. Ferment 9. https://doi.org/10.3390/fermentation9060556

[84]

Mitra R, Xu T, Xiang H, Han J (2020) Current developments on polyhydroxyalkanoates synthesis by using halophiles as a promising cell factory. Microb Cell Fact. https://doi.org/10.1186/s12934-020-01342-z

[85]

Mitra R, Xu T, Chen GQ, Xiang H, Han J (2022) An updated overview on the regulatory circuits of polyhydroxyalkanoates synthesis. Microb Biotechnol 1446–70. https://doi.org/10.1111/1751-7915.13915

[86]

Mohammadalipour M, Alihosseini F, Toloue EB, Karbasi S, Mohammadalipour Z (2025) Evaluation of PHB-chitosan/CNC scaffolds’ applicability for bone tissue engineering via MG-63 osteoblastic cell cultivation and osteogenic markers gene expression. Int J Biol Macromol 320. https://doi.org/10.1016/j.ijbiomac.2025.146041

[87]

Mohan A, Girdhar M, Kumar R, Chaturvedi HS, Vadhel A, Solanki PR et al (2021) Polyhydroxybutyrate-based nanocomposites for bone tissue engineering. Pharmaceuticals 14. https://doi.org/10.3390/ph14111163

[88]

Montemurro M, Salvatori G, Alfano S, Martinelli A, Verni M, Pontonio E et al (2022) Exploitation of wasted bread as substrate for polyhydroxyalkanoates production through the use of Haloferax mediterranei and seawater. Front Microbiol 13. https://doi.org/10.3389/fmicb.2022.1000962

[89]

Montiel-Jarillo G, Morales-Urrea DA, Contreras EM, López-Córdoba A, Gómez-Pachón EY, Carrera J et al (2022) Improvement of the Polyhydroxyalkanoates recovery from mixed microbial cultures using sodium hypochlorite pre-treatment coupled with solvent extraction. Polymers (Basel) 14. https://doi.org/10.3390/polym14193938

[90]

Moopantakath J, Imchen M, Kumavath R, Martínez-Espinosa RM (2021) Ubiquitousness of haloferax and carotenoid producing genes in Arabian sea coastal biosystems of India. Mar Drugs 19. https://doi.org/10.3390/md19080442

[91]

Motiee E-S, Karbasi S, Bidram E, Sheikholeslam M. Investigation of physical, mechanical and biological properties of polyhydroxybutyrate-chitosan/graphene oxide nanocomposite scaffolds for bone tissue engineering applications. Int J Biol Macromol [Internet], 2023, 247: 125593

[92]

Müller-Santos M, Koskimäki JJ, Alves LPS, De Souza EM, Jendrossek D, Pirttilä AM (2021) The protective role of PHB and its degradation products against stress situations in bacteria. FEMS Microbiol Rev. https://doi.org/10.1093/femsre/fuaa058

[93]

Naitam MG, Tomar GS, Pushpad U, Singh S, Kaushik R. Halophilic bacteria mediated poly-β-hydroxybutyrate production using paddy straw as a substrate. Bioresour Technol Rep [Internet], 2022, 17: 100915

[94]

Narwal S, Jaiswal S, Deswal T, Kalshan S, Yadav P, Yadav A, et al.. Bioproduction of Poly-β-hydroxybutyrate (PHB) Using Dairy Wastewater: A Sustainable and Greener Approach. Orient J Chem Orient Sci Publishing Co, 2025, 40: 1681-1687

[95]

Novackova I, Kourilova X, Mrazova K, Sedlacek P, Kalina M, Krzyzanek V et al (2022) Combination of hypotonic lysis and application of detergent for isolation of Polyhydroxyalkanoates from extremophiles. Polym (Basel) 14. https://doi.org/10.3390/polym14091761

[96]

Obruča S, Dvořák P, Sedláček P, Koller M, Sedlář K, Pernicová I, et al.. Polyhydroxyalkanoates synthesis by halophiles and thermophiles: towards sustainable production of microbial bioplastics. Biotechnol Adv [Internet], 2022, 58: 107906

[97]

Obruča S, Dvořák P, Sedláček P, Koller M, Sedlář K, Pernicová I, et al.. Polyhydroxyalkanoates synthesis by halophiles and thermophiles: towards sustainable production of microbial bioplastics. Biotechnol Adv [Internet], 2022, 58: 107906

[98]

Olavarria K, Carnet A, van Renselaar J, Quakkelaar C, Cabrera R, Guedes da Silva L, et al.. An NADH preferring acetoacetyl-CoA reductase is engaged in poly-3-hydroxybutyrate accumulation in Escherichia coli. J Biotechnol Elsevier B V, 2021, 325: 207-216

[99]

Oren A (2013) Salinibacter: an extremely halophilic bacterium with archaeal properties. FEMS Microbiol Lett 1–9. https://doi.org/10.1111/1574-6968.12094

[100]

Ouyang P, Wang H, Hajnal I, Wu Q, Guo Y, Chen GQ. Increasing oxygen availability for improving poly(3-hydroxybutyrate) production by Halomonas. Metab Eng Acad Press Inc, 2018, 45: 20-31

[101]

Paduvari R, Somashekara DM (2025) Advancements in genetic engineering for enhanced Polyhydroxyalkanoates (PHA) production: a comprehensive review of metabolic pathway manipulation and gene deletion strategies. Bioengineered. https://doi.org/10.1080/21655979.2025.2458363

[102]

Park C, Park W. Survival and energy producing strategies of Alkane degraders under extreme conditions and their biotechnological potential. Front Microbiol Front Media S A, 2018

[103]

Park YL, Bhatia SK, Gurav R, Choi TR, Kim HJ, Song HS, et al.. Fructose based hyper production of poly-3-hydroxybutyrate from Halomonas sp. YLGW01 and impact of carbon sources on bacteria morphologies. Int J Biol Macromol Elsevier B V, 2020, 154: 929-936

[104]

Park YL, Song HS, Choi TR, Lee SM, Park SL, Lee HS, et al.. Revealing of sugar utilization systems in Halomonas sp. YLGW01 and application for poly(3-hydroxybutyrate) production with low-cost medium and easy recovery. Int J Biol Macromol Elsevier B V, 2021, 167: 151-159

[105]

Parroquin-Gonzalez M, Winterburn J (2023) Continuous bioreactor production of polyhydroxyalkanoates in Haloferax mediterranei. Front Bioeng Biotechnol 11. https://doi.org/10.3389/fbioe.2023.1220271

[106]

Patience M, Singh S, Kaushik R. Characterization of poly-β-hydroxybutyrate producing halophilic bacteria and nutrient optimization for its maximum production. Indian J Agricultural Sci Indian Council Agricultural Res, 2021, 91: 350-354

[107]

Paul S, Bag SK, Das S, Harvill ET, Dutta C (2008) Molecular signature of hypersaline adaptation: insights from genome and proteome composition of halophilic prokaryotes. Genome Biol 9. https://doi.org/10.1186/gb-2008-9-4-r70

[108]

Pérez-Rivero C, López-Gómez JP, Roy I (2019) A sustainable approach for the downstream processing of bacterial polyhydroxyalkanoates: state-of-the-art and latest developments. Biochem Eng J 150. https://doi.org/10.1016/j.bej.2019.107283

[109]

Peregrina A, Martins-Lourenço J, Freitas F, Reis MAM, Arraiano CM (2021) Post-transcriptional control in the regulation of polyhydroxyalkanoates synthesis. Life. https://doi.org/10.3390/life11080853

[110]

Pillai Balakrishna A, Jaya Kumar A, Kumarapillai H (2018) Enhanced production of poly(3-hydroxybutyrate) in recombinant Escherichia coli and EDTA–microwave-assisted cell lysis for polymer recovery. AMB Express 8. https://doi.org/10.1186/s13568-018-0672-6

[111]

Pryadko AS, Botvin VV, Mukhortova YR, Pariy I, Wagner DV, Laktionov PP et al (2022) Core-shell magnetoactive phb/gelatin/magnetite composite electrospun scaffolds for biomedical applications. Polym (Basel) 14. https://doi.org/10.3390/polym14030529

[112]

Qin Q, Ling C, Zhao Y, Yang T, Yin J, Guo Y, et al.. CRISPR/Cas9 editing genome of extremophile Halomonas spp. Metab Eng [Internet], 2018, 47: 219-229

[113]

Quillaguamán J, Delgado O, Mattiasson B, Hatti-Kaul R. Poly(β-hydroxybutyrate) production by a moderate halophile, Halomonas boliviensis LC1. Enzyme Microb Technol [Internet], 2006, 38: 148-154

[114]

Quillaguamán J, Doan-Van T, Guzmán H, Guzmán D, Martín J, Everest A, et al.. Poly(3-hydroxybutyrate) production by Halomonas boliviensis in fed-batch culture. Appl Microbiol Biotechnol [Internet], 2008, 78: 227-232

[115]

Radchenkova N, Hasköylü ME, Vassilev S, Yıldız SY, Boyadzhieva I, Oner ET, et al.. Improved exopolymer production by chromohalobacter canadensis cultures for its potential cosmeceutical applications. Microorganisms MDPI AG, 2020, 8: 1-9

[116]

Rai R, Keshavarz T, Roether JA, Boccaccini AR, Roy I (2011) Medium chain length polyhydroxyalkanoates, promising new biomedical materials for the future. Mat Sci Eng R: Rep 29–47. https://doi.org/10.1016/j.mser.2010.11.002

[117]

Ramos-Valdovinos MA, Salas-Navarrete PC, Amores GR, Hernández-Orihuela AL, Martínez-Antonio A (2024) Qualitative perturbation analysis and machine learning: elucidating bacterial optimization of tryptophan production. Algorithms 17. https://doi.org/10.3390/a17070282

[118]

Rathi DN, Amir HG, Abed RMM, Kosugi A, Arai T, Sulaiman O, et al.. Polyhydroxyalkanoate biosynthesis and simplified polymer recovery by a novel moderately halophilic bacterium isolated from hypersaline microbial mats. J Appl Microbiol, 2013, 114: 384-395

[119]

Ravagnan G, Schmid J. Promising non-model microbial cell factories obtained by genome reduction. Front Bioeng Biotechnol Front Media SA, 2024

[120]

Raza ZA, Khalil S, Abid S (2020) Recent progress in development and chemical modification of poly(hydroxybutyrate)-based blends for potential medical applications. Int J Biol Macromol 77–100. https://doi.org/10.1016/j.ijbiomac.2020.05.114

[121]

Rekhi P, Goswami M, Ramakrishna S, Debnath M (2022) Polyhydroxyalkanoates biopolymers toward decarbonizing economy and sustainable future. Crit Rev Biotechnol 668–92. https://doi.org/10.1080/07388551.2021.1960265

[122]

Rivera-Terceros P, Tito-Claros E, Torrico S, Carballo S, Van-Thuoc D, Quillaguamán J (2015) Production of poly(3-hydroxybutyrate) by halomonas boliviensis in an air-lift reactor. J Biol Res-Thessaloniki 22. https://doi.org/10.1186/s40709-015-0031-6

[123]

Rivera-Terceros P, Tito-Claros E, Torrico S, Carballo S, Van-Thuoc D, Quillaguamán J (2015) Production of poly(3-hydroxybutyrate) by Halomonas boliviensis in an air-lift reactor. J Biol Res-Thessaloniki 22. https://doi.org/10.1186/s40709-015-0031-6

[124]

Rizki WOS, Ratnaningsih E, Hertadi R. Production of poly®3-hydroxybutyrate from halophilic bacterium Salinivibrio sp. utilizing palm oil mill effluent as a carbon source. Biocatal Agric Biotechnol [Internet], 2023, 47: 102558

[125]

Safaeian P, Yazdian F, Khosravi-Darani K, Rashedi H, Lackner M. P3HB from CH4 using methanotrophs: aspects of bioreactor, fermentation process and modelling for cost-effective biopolymer production. Front Bioeng Biotechnol Front Media SA, 2023

[126]

Sagar A, Rai S, Ilyas N, Sayyed RZ, Al-Turki AI, El Enshasy HA et al (2022) Halotolerant Rhizobacteria for salinity-stress mitigation: diversity, mechanisms and molecular approaches. Sustainability (Switzerland).https://doi.org/10.3390/su14010490

[127]

Salma M, Abdulla MK, Samina M (2020) Res. Res. J. Biotechnol.

[128]

Samorì C, Martinez GA, Bertin L, Pagliano G, Parodi A, Torri C, et al.. PHB into PHB: Recycling of polyhydroxybutyrate by a tandem thermolytic distillation-microbial fermentation process. Resour Conserv Recycl [Internet], 2022, 178: 106082

[129]

Sathiyanarayanan G, Saibaba G, Kiran GS, Yang YH, Selvin J (2017) Marine sponge-associated bacteria as a potential source for polyhydroxyalkanoates. Crit Rev Microbiol 294–312. https://doi.org/10.1080/1040841X.2016.1206060

[130]

Savoca MS, Abreo NA, Arias AH, Baes L, Baini M, Bergami E et al (2024) Monitoring plastic pollution using bioindicators: a global review and recommendations for marine environments. Environ Sci: Adv 10–32. https://doi.org/10.1039/d4va00174e

[131]

Schmid M, Raschbauer M, Song H, Bauer C, Neureiter M. Effects of nutrient and oxygen limitation, salinity and type of salt on the accumulation of poly(3-hydroxybutyrate) in Bacillus megaterium uyuni S29 with sucrose as a carbon source. N Biotechnol Elsevier B V, 2021, 61: 137-144

[132]

Shukla PJ, Bhatt VD, Suriya J, Mootapally C (2020) Marine extremophiles: adaptations and biotechnological applications. Encyclopedia Marine Biotechnol 1753–71. https://doi.org/10.1002/9781119143802.ch74

[133]

Singh A, Das M, Grover A. Molecular mechanism of acetoacetyl-CoA enhanced kinetics for increased bioplastic production from Cupriavidus necator 428. J Biomol Struct Dyn [Internet] Taylor Francis, 2020, 38: 827-840

[134]

Sivashankari RM, Mierzati M, Miyahara Y, Mizuno S, Nomura CT, Taguchi S et al (2023) Exploring class I polyhydroxyalkanoate synthases with broad substrate specificity for polymerization of structurally diverse monomer units. Front Bioeng Biotechnol 11. https://doi.org/10.3389/fbioe.2023.1114946

[135]

Song WS, Kim SM, Jo SH, Lee JS, Jeon HJ, Ko BJ, et al.. Multi-omics characterization of the osmotic stress resistance and protease activities of the halophilic bacterium: Pseudoalteromonas phenolica in response to salt stress. RSC Adv Royal Soc Chem, 2020, 10: 23792-23800

[136]

Stanley A, Murthy PSK, Vijayendra SVN. Characterization of Polyhydroxyalkanoate Produced by Halomonas venusta KT832796. J Polym Environ Springer, 2020, 28: 973-983

[137]

Stanley A, Mutturi S, Vijayendra SVNKuddus M, Roohi. Halophilic Microorganisms as Potential Producers of Polyhydroxyalkanoates. Bioplastics for Sustainable Development [Internet], 2021, Singapore, Springer Singapore, 277-294

[138]

Tan D, Xue YS, Aibaidula G, Chen GQ. Unsterile and continuous production of polyhydroxybutyrate by Halomonas TD01. Bioresour Technol, 2011, 102(17): 8130-8136

[139]

Tanweer S, Mishra P, Dash K, Panda BKalia VC. Two Decades Research on Cyanobacterial PHB: Challenges and Opportunities. Polyhydroxyalkanoates: Sustainable Production and Biotechnological Applications I: Microbial Biodiversity, Biowastes, and Bioprocesses [Internet], 2025, Singapore, Springer Nature Singapore, 71-102

[140]

Tao W, Lv L, Chen GQ (2017) Engineering Halomonas species TD01 for enhanced polyhydroxyalkanoates synthesis via CRISPRi. Microb Cell Fact 16. https://doi.org/10.1186/s12934-017-0655-3

[141]

Tao G-B, Tian L, Pu N, Li Z-J. Efficient production of poly-3-hydroxybutyrate from acetate and butyrate by halophilic bacteria Salinivibrio spp. TGB4 and TGB19. Int J Biol Macromol [Internet], 2022, 221: 1365-1372

[142]

Thomas T, Sudesh K, Bazire A, Elain A, Tan HT, Lim H et al (2020) PHA production and pha synthases of the halophilic bacterium halomonas sp. SF2003. Bioeng 7. https://doi.org/10.3390/bioengineering7010029

[143]

Turco R, Santagata G, Corrado I, Pezzella C, Di Serio M (2021) In vivo and post-synthesis strategies to enhance the properties of PHB-based materials: a review. Front Bioeng Biotechnol. https://doi.org/10.3389/fbioe.2020.619266

[144]

Van Thuoc D, My DN, Loan TT, Sudesh K. Utilization of waste fish oil and glycerol as carbon sources for polyhydroxyalkanoate production by Salinivibrio sp. M318. Int J Biol Macromol Elsevier B V, 2019, 141: 885-892

[145]

Van-Thuoc D, Huu-Phong T, Thi-Binh N, Thi-Tho N, Minh-Lam D, Quillaguamán J. Polyester production by halophilic and halotolerant bacterial strains obtained from mangrove soil samples located in Northern Vietnam. Microbiologyopen Blackwell Publishing Ltd, 2012, 1: 395-406

[146]

Velázquez-Sánchez C, Espín G, Peña C, Segura D. The Modification of Regulatory Circuits Involved in the Control of Polyhydroxyalkanoates Metabolism to Improve Their Production. Front Bioeng Biotechnol Front Media S A, 2020

[147]

Vlaeminck E, Uitterhaegen E, Quataert K, Delmulle T, De Winter K, Soetaert WK (2022) Industrial side streams as sustainable substrates for microbial production of poly(3-hydroxybutyrate) (PHB). World J Microbiol Biotechnol. https://doi.org/10.1007/s11274-022-03416-z

[148]

Wang J, Huang J, Liu S. The production, recovery, and valorization of polyhydroxybutyrate (PHB) based on circular bioeconomy. Biotechnol Adv [Internet], 2024

[149]

Wang S, Rao MPN, Quadri SR (2024) Assessing the metabolism, phylogenomic, and taxonomic classification of the halophilic genus Halarchaeum. FEMS Microbiol Lett 371. https://doi.org/10.1093/femsle/fnae001

[150]

Wang Z, Du C, Yan R, Li S, Zheng G, Ding D. Sustainable polyhydroxybutyrate (PHB) production from biowastes by Halomonas sp. WZQ-1 under non-sterile conditions. Int J Biol Macromol [Internet], 2025, 311: 143643

[151]

Wang Z, Du C, Yan R, Li S, Zheng G, Ding D. Sustainable polyhydroxybutyrate (PHB) production from biowastes by Halomonas sp. WZQ-1 under non-sterile conditions. Int J Biol Macromol [Internet], 2025, 311: 143643

[152]

Wen Q, Wang Z, Liu B, Liu S, Huang H, Chen Z. Enrichment performance and salt tolerance of polyhydroxyalkanoates (PHAs) producing mixed cultures under different saline environments. Environ Res [Internet], 2024, 251: 118722

[153]

Willey JM, Prescott LM, Sandman KM, Wood DH (2020) Prescott’s microbiology. McGraw-Hill Education

[154]

Wongmoon C, Napathorn SC (2022) Optimization for the efficient recovery of poly(3-hydroxybutyrate) using the green solvent 1,3-dioxolane. Front Bioeng Biotechnol 10. https://doi.org/10.3389/fbioe.2022.1086636

[155]

Xiao-Ran J, Jin Y, Xiangbin C, Guo-Qiang C (2018) Chapter eleven—Halomonas and pathway engineering for bioplastics production. In: Scrutton N, editor. Methods enzymol. Academic Press, Massachusetts; pp. 309–28. https://doi.org/10.1016/bs.mie.2018.04.008

[156]

Yao L, Zhao S, Tremblay LA, Wang W, LeBlanc GA (2025) Implications of plastic pollution on global carbon cycle. Carbon Res 4. https://doi.org/10.1007/s44246-024-00188-z

[157]

Ye J, Huang W, Wang D, Chen F, Yin J, Li TJohn Wiley & Sons, Ltd. Pilot Scale-up of Poly(3-hydroxybutyrate-co-4-hydroxybutyrate) Production by Halomonas bluephagenesis via Cell Growth Adapted Optimization Process. Biotechnol J [Internet], 2018, 13: 1800074

[158]

Ye J-W, Lin Y-N, Yi X-Q, Yu Z-X, Liu X, Chen G-Q. Synthetic biology of extremophiles: a new wave of biomanufacturing. Trends Biotechnol [Internet], 2023, 41: 342-357

[159]

Yue H, Ling C, Yang T, Chen X, Chen Y, Deng H et al (2014) A seawater-based open and continuous process for polyhydroxyalkanoates production by recombinant Halomonas campaniensis LS21 grown in mixed substrates. Biotechnol Biofuels 7. https://doi.org/10.1186/1754-6834-7-108

[160]

Zhang X, Lin Y, Chen GQ (2018) Halophiles as chassis for bioproduction. Adv Biosyst. https://doi.org/10.1002/adbi.201800088

[161]

Zhang Z, Shao M, Zhang G, Sun S, Yi X, Zhang Z, et al.. Engineering Halomonas bluephagenesis for synthesis of polyhydroxybutyrate (PHB) in the presence of high nitrogen containing media. Metab Eng Acad Press Inc, 2024, 86: 242-249

Funding

Manipal University Jaipur

Rights & permissions

The Author(s)

PDF

8

Accesses

0

Citation

Detail

Sections
Recommended

/