凡纳滨对虾养殖系统中生物絮团的形成机制及外源微生物调控作用研究进展

Research progress in forming mechanism of biofloc in Pacific white shrimp (Litopenaeus vannamei) culture systems and regulatory effects of exogenous microorganisms

摘要

凡纳滨对虾(Litopenaeus vannamei)是我国水产养殖业的支柱物种，随着市场需求持续增长和养殖技术不断进步，其养殖模式正向超高密度方向发展。生物絮团(biofloc)技术在提升水产养殖效益、改善水质、防控病害等方面发挥着关键作用，已成为凡纳滨对虾超高密度养殖体系中不可或缺的核心技术。然而，关于生物絮团通过哪些特定微生物类群改善生长性能和水体质量，以及通过何种途径调控水生动物的免疫功能这两个关键机制问题仍存在不小的争议。并且外界影响因素复杂多样，难以稳定培养出良性的生物絮团系统，严重制约其推广应用。近年的研究证实，微生物群落是调控生物絮团系统的根本要素，与调节碳氮比和添加碳源等经典的调控策略相比，添加益生菌和微藻能够更稳定、有效地改善生物絮团系统，该方向研究已成为生物絮团领域的热点方向。本文综合分析了有关生物絮团作用机制的多种观点，梳理和评估了影响其稳定性的关键因素;在此基础上，重点探讨了外源微生物的种类和添加量对凡纳滨对虾生长性能、免疫状态、养殖水质以及生物絮团群落结构的综合影响，并对未来研究方向进行了展望，以期为提高生物絮团养殖系统的稳定输出、实现凡纳滨对虾的绿色、健康养殖提供科学依据。

Abstract

Pacific white shrimp (Litopenaeus vannamei), as a pillar species of China's aquaculture industry, has seen its farming model evolve towards super-intensive density due to the continuous growth in market demand and advancements in aquacultural technology. Biofloc technology plays a critical role in enhancing farming efficiency, improving water quality, and preventing diseases, making it an indispensable core technology in the super-intensive farming systems of L. vannamei. There is still controversy about which specific microbial groups in bioflocs contribute to improving growth performance and water quality, as well as the mechanisms through which they regulate the immune functions of aquatic animals. Additionally, the complexity and diversity of external influencing factors also make it difficult to stably cultivate beneficial biofloc systems, hindering their broader applications. Recent studies have demonstrated microorganisms are the fundamental element in regulating biofloc systems. Compared to traditional strategies such as adjusting the carbon-to-nitrogen ratio and carbon sources, the exogenous addition of probiotics and microalgae has proven to be a more stable and effective approach for optimizing biofloc systems, thereby becoming a hot spot in biofloc research. This review comprehensively analyzes various perspectives on the mechanisms of biofloc, systematically identifies and evaluates the key factors affecting its stability, and focuses on the impact of different types and dosages of exogenous microorganisms on the growth performance, immune status, water quality, and biofloc community structure in L. vannamei farming. Additionally, future research directions are proposed. This study aims to provide a scientific basis for stabilizing the output of biofloc farming systems and achieving sustainable and healthy farming of L. vannamei.

关键词

凡纳滨对虾 / 生物絮团 / 高密度养殖 / 外源微生物

Key words

Litopenaeus vannamei / biofloc / super-intensive culture / exogenous microorganism

参考文献

[1] FAO. The State of World Fisheries and Aquaculture 2024[M]. FAO, 2024.

[2] 农业农村部渔业渔政管理局，全国水产技术推广总站，中国水产学会. 中国渔业统计年鉴 2024[M]. 北京:中国农业出版社，2024.

[3] Fishery and Fishery Administration Bureau of the Ministry of Agriculture and Rural Affairs, National Aquaculture Technology Promotion Station, Chinese Fisheries Society. China Fisheries Statistical Yearbook 2024[M]. Beijing:China Agriculture Press, 2024(in Chinese).

[4] Villarre al H, Juarez L. Super-intensive shrimp culture:analysis and future challenges[J]. Journal of the World Aquaculture Society, 2022, 53(5): 928-932.

[5] da Silveira LGP, Krummenauer D, Poersch LH, et al. Hyperintensive stocking densities for Litopenaeus vannamei grow-out in biofloc technology culture system [J]. Journal of the World Aquaculture Society, 2020, 51(6): 1290-1300.

[6] de Almeida MS, Gimenes RMT, Furtado PS, et al. Economic analysis of intensive and super-intensive Litopenaeus vannamei shrimp production in a biofloc technology system [J]. Boletim do Instituto de Pesca, 2022, 48: e692.

[7] Emerenciano MGC, Robbens AN, Vieira FDN, et al. Intensification of penaeid shrimp culture:a applied review of advances in production systems, nutrition and breeding[J]. Animals, 2022, 12(3): 236.

[8] Foese G, Krummenauer D, Lara G, et al. Long term storage and the compensatory growth of white shrimp Litopenaeus vannamei in aquaculture ponds [J]. Latin American Journal of Aquatic Research, 2017, 44(3): 588-594.

[9] Samocha TM, Prangnell DI, Hanson TR, et al. Design and operation of high-density, biofloc-dominated production systems of Pacific white shrimp, Penaeus vannamei [M] // Vannamei Shrimp Farming. Boca Raton:CRC Press, 2020: 14-36.

[10] Ciji A, Akhtar MS. Stress management in aquaculture:a review of dietary interventions[J]. Reviews in Aquaculture, 2021, 13(4): 2190-2247.

[11] Khanjani MH, Alizadeh M, Sharifinia M. Rearing of the Pacific white shrimp, Litopenaeus vannamei in a biofloc system:the effects of different food sources and salinity levels [J]. Aquaculture Nutrition, 2020, 26(2): 328-337.

[12] Yu YB, Lee JH, Choi JH, et al. The application and future of biofloc technology (BFT) in aquaculture industry:a review[J]. Journal of Environmental Management, 2023, 342: 118237.

[13] Ramasubbura yan R, Prakash S, Immanuel G, et al. The transformative role of prebiotics, probiotics, and microbiomes in biofloc systems for sustainable aquaculture:a comprehensive review[J]. Reviews in Aquaculture, 2025, 17(1): e13000.

[14] Sun Y, Hou HC, Dong DP, et al. Comparative life cycle assessment of white leg shrimp (Penaeus vannamei) cultured in recirculating aquaculture systems(RAS), biofloc technology(BFT)and higher-place ponds(HPP)farming systems in China [J]. Aquaculture, 2023, 574: 739625.

[15] Pacheco-Vega JM, Cadena-Roa MA, Leyva-Flores JA, et al. Effect of isolated bacteria and microalgae on the biofloc characteristics in the Pacific white shrimp culture[J]. Aquaculture Reports, 2018, 11: 24-30.

[16] Ghosh AK, Hasanuzzaman AFM, Sarower MG, et al. Unveiling the biofloc culture potential:harnessing immune functions for resilience of shrimp and resistance against AHPND:causing Vibrio parahaemolyticus infection [J]. Fish & Shellfish Immunology, 2024, 151: 109710.

[17] Rego MAS, Sabbag OJ, Soares R, et al. Risk analysis of the insertion of biofloc technology in marine shrimpLitopenaeus vannamei production in a farm in Pernambuco, Brazil:a case study [J]. Aquaculture, 2017, 469: 67-71.

[18] Gutierrez SM, Dosta M, Partida AH, et al. Effect of two carbon sources in microbial abundance in a Biofloc culture system with Oreochromis niloticus(Linnaeus, 1758) [J]. International Journal of Fisheries and Aquatic Studies, 2016, 4(3): 421-427.

[19] Kasan NA, Ghazali NA, Ikhwanuddin M, et al. Isolation of potential bacteria as inoculum for biofloc formation in Pacific white leg shrimp,Litopenaeus vannamei culture ponds [J]. Pakistan Journal of Biological Sciences, 2017, 20(6): 306-313.

[20] Tepaamorndech S, Nookaew I, Higdon SM, et al. Metagenomics in bioflocs and their effects on gut microbiome and immune responses in Pacific white shrimp[J]. Fish & Shellfish Immunology, 2020, 106: 733-741.

[21] El-Sayed AM. Use of biofloc technology in shrimp aquaculture:a comprehensive review, with emphasis on the last decade[J]. Reviews in Aquaculture, 2021, 13(1): 676-705.

[22] Rios LDM, Monta gu do EB, Barrios YC, et al. Biofloc technology and immune response of penaeid shrimp:a meta-analysis and meta-regression[J]. Fish & Shellfish Immunology, 2023, 138: 108805.

[23] 周航先， 赵建华， 徐奇友. 基于文献计量学的生物絮团技术在水产中的研究热点分析[J]. 水生态学杂志， 2025， 46(3): 193-204.

[24] Zhou H X, Zhao J H, Xu Q Y. Research hotspot analysis of biofloc technology in aquaculture based on bibliometrics[J]. Journal of Hydroecology, 2025, 46(3): 193-204(in Chinese with English abstract).

[25] Li Z, Ju CW, Jiao TY, et al. Effects of biofloc on growth performance and survival of Pacific white shrimp (Litopenaeus vannamei)through C/N ratio manipulation, probiotics supplementation, and cocultivation time:a meta-analysis [J]. Aquaculture, 2024, 587: 740837.

[26] Liu G, Verdegem M, Ye ZY, et al. Advancing aquaculture sustainability:a comprehensive review of biofloc technology trends, innovative research approaches, and future prospects[J]. Reviews in Aquaculture, 2025, 17(1): e12970.

[27] Guo HP, Huang L, Hu ST, et al. Effects of carbon/nitrogen ratio on growth, intestinal microbiota and metabolome of shrimp(Litopenaeus vannamei) [J]. Frontiers in Microbiology, 2020, 11: 652.

[28] Yang SF, Li XY. Influences of extracellular polymeric substances(EPS)on the characteristics of activated sludge under non-steady-state conditions[J]. Process Biochemistry, 2009, 44(1): 91-96.

[29] Wilen BM, Lumley D, Mattsson A, et al. Dynamics in flocculation and settling properties studied at a full-scale activated sludge plant[J]. Water Environment Research, 2010, 82(2): 155-168.

[30] Schmidt I, Sliekers O, Schmid M, et al. New concepts of microbial treatment processes for the nitrogen removal in wastewater[J]. FEMS Microbiology Reviews, 2003, 27(4): 481-492.

[31] Czepiel P, Crill P, Harriss R. Nitrous oxide emissions from municipal wastewater treatment[J]. Environmental Science & Technology, 1995, 29(9): 2352-2356.

[32] Ahmed Alkhames Y, Sultana A, Tareq Arfat S, et al. The impact of biofloc technology on water quality in aquaculture:a systematic meta-analysis[J]. Aquaculture Nutrition, 2023, 2023(1): 9915874.

[33] Kumar V, Roy S, Behera BK, et al. Biofloc microbiome with bioremediation and health benefits[J]. Frontiers in Microbiology, 2021, 12: 741164.

[34] Weber SD, Wanner G, Ludwig W, et al. Microbial composition and structure of aerobic granular sewage biofilms[J]. Applied and Environmental Microbiology, 2008, 74(1): 343.

[35] Duan SP, Zhang YY, Zheng SK. Heterotrophic nitrifying bacteria in wastewater biological nitrogen removal systems:a review[J]. Critical Reviews in Environmental Science and Technology, 2022, 52(13): 2302-2338.

[36] Shi SH, He XJ, He L, et al. Overlooked pathways of endogenous simultaneous nitrification and denitrification in anaerobic/aerobic/anoxic sequencing batch reactors with organic supplementation[J]. Water Research, 2023, 230: 119493.

[37] Li CW, Li JW, Liu G, et al. Performance and microbial community analysis of combined denitrification and biofloc technology(CDBFT)system treating nitrogen-rich aquaculture wastewater[J]. Bioresource Technology, 2019, 288: 121582.

[38] Liu HJ, Guo JB, Qu JH, et al. Catalyzing denitrification of Paracoccus versutus by immobilized 1,5-dichloroanthraquinone [J]. Biodegradation, 2012, 23(3): 399-405.

[39] Raza B, Zheng ZM, Zhu JY, et al. A review:microbes and their effect on growth performance of Litopenaeus vannamei(white leg shrimp)during culture in biofloc technology system [J]. Microorganisms, 2024, 12(5): 1013.

[40] Li Z, Du QY, Jiao TY, et al. Effects of probiostics supplementary bioflocs(Rhodospirillum rubrum, Bacillus subtilis, Providencia rettgeri)on growth, immunity, and water quality in cultures of Pacific white shrimp(Litopenaeus vannamei) [J]. Aquaculture, 2024, 591: 741141.

[41] Xu WJ, Wen GL, Su HC, et al. Effect of input C/N ratio on bacterial community of water biofloc and shrimp gut in a commercial zero-exchange system with intensive production of Penaeus vannamei [J]. Microorganisms, 2022, 10(5): 1060.

[42] Cao HP, Zhang SM, An J, et al. Rhodobacter azotoformans supplementation improves defense ability of Chinese mitten crab Eriocheir sinensis against citrobacteriosis [J]. Fish & Shellfish Immunology, 2022, 131: 991-998.

[43] Murakami K, Kamimura D, Hasebe R, et al. Rhodobacter azotoformans LPS(RAP99-LPS)is a TLR4 agonist that inhibits lung metastasis and enhances TLR3-mediated chemokine expression [J]. Frontiers in Immunology, 2021, 12: 675909.

[44] de Souza Valente C, Rodiles A, Freire Marques MR, et al. White spot syndrome virus(WSSV)disturbs the intestinal microbiota of shrimp(Penaeus vannamei)reared in biofloc and clear seawater [J]. Applied Microbiology and Biotechnology, 2020, 104(18): 8007-8023.

[45] Ebeling JM, Timmons MB, Bisogni JJ. Engineering analysis of the stoichiometry of photoautotrophic, autotrophic, and heterotrophic removal of ammonia-nitrogen in aquaculture systems[J]. Aquaculture, 2006, 257(1/2/3/4): 346-358.

[46] Hargreaves JA. Photosynthetic suspended-growth systems in aquaculture[J]. Aquacultural Engineering, 2006, 34(3): 344-363.

[47] Avnimelech Y. Carbon/nitrogen ratio as a control element in aquaculture systems[J]. Aquaculture, 1999, 176(3/4): 227-235.

[48] Minaz M, Kubilay A. Operating parameters affecting biofloc technology:carbon source, carbon/nitrogen ratio, feeding regime, stocking density, salinity, aeration, and microbial community manipulation[J]. Aquaculture International, 2021, 29(3): 1121-1140.

[49] Xu WJ, Morris TC, Samocha TM. Effects of C/N ratio on biofloc development, water quality, and performance of Litopenaeus vannamei juveniles in a biofloc-based, high-density, zero-exchange, outdoor tank system [J]. Aquaculture, 2016, 453: 169-175.

[50] Zhao DH, Pan LQ, Huang F, et al. Effects of different carbon sources on bioactive compound production of biofloc, immune response, antioxidant level, and growth performance of Litopenaeus vannamei in zero-water exchange culture tanks [J]. Journal of the World Aquaculture Society, 2016, 47(4): 566-576.

[51] Mansour AT, Ashry OA, Ashour M, et al. The optimization of dietary protein level and carbon sources on biofloc nutritive values, bacterial abundance, and growth performances of white leg shrimp(Litopenaeus vannamei)juveniles [J]. Life, 2022, 12(6): 888.

[52] Ekasari J, Hanif Azhar M, Surawidjaja EH, et al. Immune response and disease resistance of shrimp fed biofloc grown on different carbon sources[J]. Fish & Shellfish Immunology, 2014, 41(2): 332-339.

[53] Panigrahi A, Sundaram M, Saranya C, et al. Carbohydrate sources differentially influence growth performances, microbial dynamics and immunomodulation in Pacific white shrimp(Litopenaeus vannamei)under biofloc system [J]. Fish & Shellfish Immunology, 2019, 86: 1207-1216.

[54] Panigrahi A, Saranya C, Sundaram M, et al. Carbon:nitrogen(C:N)ratio level variation influences microbial community of the system and growth as well as immunity of shrimp(Litopenaeus vannamei)in biofloc based culture system [J]. Fish & Shellfish Immunology, 2018, 81: 329-337.

[55] Chakrapani S, Panigrahi A, Sundaresan J, et al. Three different C:N ratios for Pacific white shrimp, Penaeus vannamei under practical conditions:evaluation of growth performance, immune and metabolic pathways [J]. Aquaculture Research, 2021, 52(3): 1255-1266.

[56] Asaduzzaman M, Rahman MM, Azim ME, et al. Effects of C/N ratio and substrate addition on natural food communities in freshwater prawn monoculture ponds[J]. Aquaculture, 2010, 306(1/2/3/4): 127-136.

[57] Xue YM, Wei FY, Jiang YL, et al. Transcriptome signatures of the Pacific white shrimp Litopenaeus vannamei hepatopancreas in response to stress in biofloc culture systems [J]. Fish & Shellfish Immunology, 2019, 91: 369-375.

[58] Huang HH, Liao HM, Lei YJ, et al. Effects of different carbon sources on growth performance of Litopenaeus vannamei and water quality in the biofloc system in low salinity [J]. Aquaculture, 2022, 546: 737239.

[59] 鲍方剑， 黄雷， 陈伟， 等. 蔗糖输入对凡纳滨对虾养殖系统真核微生物群落的影响[J]. 水生生物学报， 2021， 45(1): 172-181.

[60] Bao F J, Huang L, Chen W, et al. The effects of sucrose on microeukaryotic community in the shrimp aquaculture system[J]. Acta Hydrobiologica Sinica, 2021, 45(1): 172-181(in Chinese with English abstract).

[61] Rego MAS, Sabbag OJ, Soares RB, et al. Technical efficiency analysis of marine shrimp farming(Litopenaeus vannamei)in biofloc and conventional systems:a case study in northeastern Brazil [J]. Anais da Academia Brasileira de Ciencias, 2018, 90(4): 3705-3716.

[62] 曲寅， 章晓栋， 任岗， 等. 间歇性饥饿对南美白对虾生物絮团养殖系统中微生物群落结构、水质和生长的影响[J]. 农业生物技术学报， 2024， 32(2): 417-426.

[63] Qu Y, Zhang XD, Ren G, et al. Effects of intermittent fasting on microbial community structure, water quality and growth of Litopenaeus vannamei in a biofloc system [J]. Journal of Agricultural Biotechnology, 2024, 32(2): 417-426(in Chinese with English abstract).

[64] Esparza-Leal HM, Ponce-Palafox JT, Alvarez-Ruiz P, et al. Effect of stocking density and water exchange on performance and stress tolerance to low and high salinity by Litopenaeus vannamei postlarvae reared with biofloc in intensive nursery phase [J]. Aquaculture International, 2020, 28(4): 1473-1483.

[65] Ray AJ, Lotz JM. Comparing salinities of 10, 20, and 30‰ in intensive, commercial-scale biofloc shrimp(Litopenaeus vannamei)production systems [J]. Aquaculture, 2017, 476: 29-36.

[66] van Hoa N, Tao CT, Terahara T, et al. Influence of salinity on nursery-phase white leg shrimp(Litopenaeus vannamei)reared in a biofloc system [J]. Journal of Applied Aquaculture, 2022, 34(4): 907-921.

[67] de Souza DM, Martins AC, Jensen L, et al. Effect of temperature on antioxidant enzymatic activity in the Pacific white shrimp Litopenaeus vannamei in a BFT(Biofloc technology)system [J]. Marine and Freshwater Behaviour and Physiology, 2014, 47(1): 1-10.

[68] de Souza DM, Borges VD, Furtado P, et al. Antioxidant enzyme activities and immunological system analysis of Litopenaeus vannamei reared in biofloc technology(BFT)at different water temperatures [J]. Aquaculture, 2016, 451: 436-443.

[69] Becerra-Dorame MJ, Martinez-Porchas M, Martinez-Cordova LR, et al. Production response and digestive enzymatic activity of the Pacific white shrimp Litopenaeus vannamei(Boone, 1931)intensively pregrown in microbial heterotrophic and autotrophic-based systems [J]. The Scientific World Journal, 2012, 2012: 723654.

[70] Baloi M, Arantes R, Schweitzer R, et al. Performance of Pacific white shrimp Litopenaeus vannamei raised in biofloc systems with varying levels of light exposure [J]. Aquacultural Engineering, 2013, 52: 39-44.

[71] Lara G, Krummenauer D, Abreu PC, et al. The use of different aerators on Litopenaeus vannamei biofloc culture system:effects on water quality, shrimp growth and biofloc composition [J]. Aquaculture International, 2017, 25(1): 147-162.

[72] Lim YS, Ganesan P, Varman M, et al. Effects of microbubble aeration on water quality and growth performance of Litopenaeus vannamei in biofloc system [J]. Aquacultural Engineering, 2021, 93: 102159.

[73] de Morais APM, Abreu PC, Wasieleisky W, et al. Effect of aeration intensity on the biofilm nitrification process during the production of the white shrimp Litopenaeus vannamei(Boone, 1931)in biofloc and clear water systems [J]. Aquaculture, 2020, 514: 734516.

[74] Pierri V, Valter-Severino D, Goulart-de-Oliveira K, et al. Cultivation of marine shrimp in biofloc technology(BFT)system under different water alkalinites[J]. Revista Brasileira de Biologia, 2015, 75(3): 558-564.

[75] Furtado PS, Poersch LH, Wasieleisky W. The effect of different alkalinity levels on Litopenaeus vannamei reared with biofloc technology(BFT) [J]. Aquaculture International, 2015, 23(1): 345-358.

[76] de Moura PS, Wasieleisky W, da Paz Serra F, et al. Partial seawater inclusion to improve Litopenaeus vannamei performance in low salinity biofloc systems [J]. Aquaculture, 2021, 531: 735905.

[77] Deng YL, Xu XY, Yin XW, et al. Effect of stock density on the microbial community in biofloc water and Pacific white shrimp(Litopenaeus vannamei)gut microbiota [J]. Applied Microbiology and Biotechnology, 2019, 103(10): 4241-4252.

[78] Liu G, Zhu SM, Liu DZ, et al. Effects of stocking density of the white shrimp Litopenaeus vannamei(Boone)on immunities, antioxidant status, and resistance against Vibrio harveyi in a biofloc system [J]. Fish & Shellfish Immunology, 2017, 67: 19-26.

[79] Said MM, El-barbary YA, Ahmed OM. Assessment of performance, microbial community, bacterial food quality, and gene expression of white leg shrimp(Litopenaeus vannamei)reared under different density biofloc systems [J]. Aquaculture Nutrition, 2022, 2022(1): 3499061.

[80] Inunes RS, Branco PC, Pressinotti LN, et al. Does the heterotrophic system influence the cellular immune response of Litopenaeus vannamei shrimp? in vitro phagocytosis indices and superoxide anion production comparisons [J]. Fish and Shellfish Immunology Reports, 2021, 2: 100009.

[81] Kim SK, Song J, Rajeev M, et al. Exploring bacterioplankton communities and their temporal dynamics in the rearing water of a biofloc-based shrimp(Litopenaeus vannamei)aquaculture system [J]. Frontiers in Microbiology, 2022, 13: 995699.

[82] Situmorang ML, Uwisetwathan U, Arayamethakorn S, et al. Supplementation of ex situ produced bioflocs improves immune response against AHPND in Pacific white leg shrimp(Litopenaeus vannamei)postlarvae [J]. Applied Microbiology and Biotechnology, 2022, 106(9): 3751-3764.

[83] Waiho K, Abd Razak MS, Abdul Rahman MZ, et al. A metagenomic comparison of clear water, probiotic, and rapid BFT^TM on Pacific white leg shrimp, Litopenaeus vannamei cultures [J]. PeerJ, 2023, 11: e15758.

[84] Wu JP, Chen YF, Xu XN, et al. Screening of biofloculant and cellulase-producing bacteria strains for biofloc culture systems with fiber-rich carbon source[J]. Frontiers in Microbiology, 2022, 13: 969664.

[85] He X, Abakari G, Tan HX, et al. Effects of different probiotics(Bacillus subtilis)addition strategies on culture of Litopenaeus vannamei in biofloc technology(BFT)aquaculture system [J]. Aquaculture, 2023, 566: 739216.

[86] Hostins B, Wasieleisky W, Decamp O, et al. Managing input C/N ratio to reduce the risk of acute hepatopancreatic necrosis disease(AHPND)outbreaks in biofloc systems:a laboratory study[J]. Aquaculture, 2019, 508: 60-65.

[87] Ge HX, Li J, Chang ZQ, et al. Effect of microalgae with semicontinuous harvesting on water quality and zootechnical performance of white shrimp reared in the zero water exchange system[J]. Aquacultural Engineering, 2016, 72: 70-76.

[88] Huang C, Luo YY, Zeng GQ, et al. Effect of adding microalgae to white leg shrimp culture on water quality, shrimp development and yield[J]. Aquaculture Reports, 2022, 22: 100916.

[89] Panigrahi A, Esakkiraj P, Saranya C, et al. A biofloc-based aquaculture system bio-augmented with probiotic bacteria Bacillus tequilensis AP BFT3 improves culture environment, production performances, and proteomic changes in Penaeus vannamei [J]. Probiotics and Antimicrobial Proteins, 2022, 14(2): 277-287.

[90] 高远， 李甍， 董登攀， 等. 凡纳滨对虾养殖系统中异养和自养型生物絮团的微生物特性及其与养殖水环境的关系[J]. 中国水产科学， 2022， 29(6): 864-873.

[91] Gao Y, Li M, Dong DP, et al. Microbial characteristics of heterotrophic and autotrophic biofloc in Litopenaeus vannamei culturing systems and their relationship with water quality [J]. Journal of Fishery Sciences of China, 2022, 29(6): 864-873(in Chinese with English abstract).

[92] McCusker S, Warberg MB, Davies SJ, et al. Biofloc technology as part of a sustainable aquaculture system:a review on the status and innovations for its expansion[J]. Aquaculture, Fish and Fisheries, 2023, 3(4): 331-352.

[93] Silva VF, Pereira PKM, Martins MA, et al. Effects of microalgae addition and fish feed supplementation in the integrated rearing of Pacific white shrimp and Nile Tilapia using biofloc technology[J]. Animals, 2022, 12(12): 1527.

[94] 陈睿， 陈晓颖， 刘智明， 等. 生物絮团技术在水产养殖业中的研究进展[J]. 工业微生物， 2023， 53(1): 51-54.

[95] Chen R, Chen XY, Liu ZM, et al. On the research progress of biofloc technology in aquaculture industry[J]. Industrial Microbiology, 2023, 53(1): 51-54(in Chinese with English abstract).

[96] Nie XF, Mubashar M, Zhang S, et al. Current progress, challenges and perspectives in microalgae-based nutrient removal for aquaculture waste:a comprehensive review[J]. Journal of Cleaner Production, 2020, 277: 124209.

[97] Ma MY, Hu Q. Microalgae as feed sources and feed additives for sustainable aquaculture:prospects and challenges[J]. Reviews in Aquaculture, 2024, 16(2): 818-835.

[98] Wan Mahari WA, Waiho K, Fazhan H, et al. Emerging paradigms in sustainable shellfish aquaculture:microalgae and biofloc technologies for wastewater treatment[J]. Aquaculture, 2024, 587: 740835.

[99] Wang H, Qi BF, Jiang XM, et al. Microalgal interstrain differences in algal-bacterial biofloc formation during liquid digestate treatment[J]. Bioresource Technology, 2019, 289: 121741.

[100] Ramanan R, Kim BH, Cho DH, et al. Algae-bacteria interactions:evolution, ecology and emerging applications[J]. Biotechnology Advances, 2016, 34(1): 14-29.

[101] Abakari G, Wu X, He X, et al. Bacteria in biofloc technology aquaculture systems:roles and mediating factors[J]. Reviews in Aquaculture, 2022, 14(3): 1260-1284.

[102] Mugwanya M, Dawood MAO, Kimer a F, et al. Biofloc systems for sustainable production of economically important aquatic species:a review[J]. Sustainability, 2021, 13(13): 7255.

[103] Khoa TND, Tao CT, van Khanh L, et al. Super-intensive culture of white leg shrimp(Litopenaeus vannamei)in outdoor biofloc systems with different sunlight exposure levels:emphasis on commercial applications [J]. Aquaculture, 2020, 524: 735277.

[104] Albright MBN, Louca S, Winkler DE, et al. Solutions in microbiome engineering:prioritizing barriers to organism establishment[J]. The ISME Journal, 2021, 16(2): 331-338.

[105] Wang L, Jia XQ, Xu L, et al. Engineering microalgae for water phosphorus recovery to close the phosphorus cycle[J]. Plant Biotechnology Journal, 2023, 21(7): 1373-1382.

[106] 何梅琳， 迟巧云， 王长海. 极地微藻对极端环境的适应机制研究进展[J]. 南京农业大学学报， 2019， 42(2): 201-208.

[107] He M L, Chi Q Y, Wang C H. Research progress of the adaptive mechanisms of polar microalgae in response to extreme environment[J]. Journal of Nanjing Agricultural University, 2019, 42(2): 201-208(in Chinese with English abstract).

[108] Schwaderer AS, Yoshiyama K, de Tezanos Pinto P, et al. Eco-evolutionary differences in light utilization traits and distributions of freshwater phytoplankton[J]. Limnology and Oceanography, 2011, 56(2): 589-598.

[109] Cooper MB, Smith AG. Exploring mutualistic interactions between microalgae and bacteria in the omics age[J]. Current Opinion in Plant Biology, 2015, 26: 147-153.

[110] Natrah FMI, Bossier P, Sorgeloos P, et al. Significance of microalgal-bacterial interactions for aquaculture[J]. Reviews in Aquaculture, 2014, 6(1): 48-61.

[111] Amin SA, Parker MS, Armbrust EV. Interactions between diatoms and bacteria[J]. Microbiology and Molecular Biology Reviews, 2012, 76(3): 667-684.

基金资助

国家重点研发计划项目(2022YFD1600401)

河北省农业科技成果转化项目(南美白对虾工厂化养殖用水预处理技术应用)()

河北省自然科学青年基金项目(C2022205042)