微生物学报

时间Year	重要事件 Key events
启蒙孕育期(几千年前至20世纪中叶) Enlightenment period (from thousands of years ago to the middle 20th century)
1703	人类首次对淡水硅藻——平板藻(Tabellaria)进行细胞形态学观察和记录 The first observation and record of cell morphology of fresh-water diatom Tabellaria
1753	卡尔·林奈在《植物种志》(Speciesl Panatrum)中，将地衣归类到Lihcen属 Carl Linnaeus classified lichens in the genus Lihcen in the book titled Speciesl Panatrum
1798	埃里克·阿卡里乌斯开创地衣分类学，出版多部地衣学著作，推动其成为一门独立学科 Erik Acharius created lichen taxonomy and published several books on lichenology, and promoted it to become an independent discipline
1867	西蒙·施文德纳发现地衣是由真菌与藻共生的复合生物体，揭示了地衣的共生本质 Simon Schwendener discovered that lichens were complex organisms which were composed of fungi and algae, thus revealed the dual nature of the lichens
1872	约翰内斯·莱因克提出“consortium”一词，用来表示构成地衣的藻与真菌的关系 Johannes Reinke coined the term “consortium” to describe the relationship between algae and fungi that make up the lichens
1890	马丁努斯·贝耶林克分离获得藻际细菌 Martinus Beijerinck was the first people who isolated phycosphere bacteria
1915	查尔斯·沃尔科特从前古生代石灰岩藻沉积物化石中发现藻际细菌存在证据 Charles Walcott found the evidence for the existence of phycosphere bacteria in fossilized limestone from the prepaleozoic era
1940s	恩斯特·普林斯海姆分离保存大量藻株并进行分享；为世界主要藻株保藏中心作出了重要贡献 Ernst Prinsheim isolated, preserved, and shared a large number of algae strains, and made significant contributions to major algae collections in the world
创建成长期(20世纪中叶至21世纪10年代) Establishment and growth period (from middle 20th century to 2010s)
1943	迈耶-阿比奇提出全生物体(holobiont)概念及全共生理论(Theory of Holobiosis) Meyer-Abich coined the concept of holobiont and proposed the Theory of Holobiosis
1952	威廉·奥斯特瓦尔德等利用藻-菌共生体系开发生活污水的生物处理技术 Wilhelm Ostwald et al. developed a biological treatment technology for domestic sewage based on the bacteria-algae symbiotic system
1953	哈罗德·米尔纳发明透明容器推动规模化培养微藻细胞，为光生物反应器实际应用奠定重要基础 Harold Milner invented the transparent container to promote the large-scale cultivation of microalgae cells, and laid important foundation for the practical application of photobioreactors
1957	乔治·斯科特提出藻际生命共同体(phycobiont)概念 George Scott coined the concept of phycobiont
1963	萨弗曼发现蓝藻病毒及噬藻体(1969) Saffman discovered cyanobacterial viruses in 1963, and then cyanophages in 1969
1969	詹姆斯·斯图尔特及马尔科姆·布朗首次发现溶藻菌 James Stewart and Malcolm Brown first discovered the algicidal bacteria
1970	林恩·马古利斯提出内共生学说(Endosymbiosis Theory) Lynn Margulis proposed the Endosymbiosis Theory
1972	韦恩·贝尔及拉尔夫·米切尔提出藻际(phycosphere)概念 Wayne Bell and Ralph Mitchell coined the concept of phycosphere
1973	魏江春院士领衔编研《中国地衣志》一书 Wei Jiangchun began to compiled the book titled Chinese Lichens
1978	埃斯特拉·席尔瓦首次发现甲藻胞内细菌；提出著名的“甲藻共生菌产毒”假说 Estela Silva discovered intracellular bacteria in dinoflagellates and proposed the famous hypothesis of “toxic production by symbiotic bacteria of dinoflagellate”
1982	乔纳森·科尔提出菌藻相互作用(interactions between bacteria and algae) Jonathan Cole coined the concept of interactions between bacteria and algae
1988	儿玉正明发现来源于产毒海洋甲藻的藻际自产毒细菌莫拉氏菌PTB-1 Masaaki Kodama discovered the paralytic shellfish poisoning (PSP) toxin-producing bacterium, Moraxella sp. PTB-1 from a PSP toxin-producing marine dinoflagellate
1989	约翰·拉齐尔发现影响藻际生态位大小的因素 John Lazear studied and discovered the factors that influence the size of the phycosphere niche
1994	郑天凌等提出“以菌治藻”新思路 Zheng Tianling et al. proposed a new idea of “bacterial control the harmful algal blooms (HABs)”
1995	格雷戈里·杜塞特发现溶藻菌及藻-菌相互作用 Gregory Doucette discovered algicidal bacteria and algae-bacteria interactions
1999	伯纳德·雅克等提出互作组(interactome)概念 Bernard Jacques et al. coined the concept of interactome
2000	玛丽·莫兰等研究藻际玫瑰杆菌(Roseobacter)类细菌；发现微藻对藻际菌群结构起决定性作用 Mary Moran’s team studied the bacteria of the Roseobacter group in the the phycosphere niche, and found that microalgae play a decisive role in the community structure of the phycosphere microbiota
2003	魏江春院士团队提出“沙漠生物地毯工程” Wei Jiangchun’s team proposed the “Desert Biocarpet Project”
2008	伊拉娜·齐尔伯-罗森伯格等提出全基因组(hologenome)概念 Ilana Zilber-Rosenberg et al. proposed the concept of hologenome
2009	弗吉尼亚·阿姆布鲁斯特团队提出硅藻藻-菌关系 Virginia Armbrust’s team proposed the interactions between diatoms and algae
2009	杨桥团队开展海洋甲藻的可培养藻际微生物的分离及其生物活性功能研究 Yang Qiao’s team isolated the cultivable phycosphere bacteria from toxic marine dinoflagellates and studied their bioactive functions
快速发展期 (21世纪10年代至今) Rapid development period (from 2010s to the present)
2012	弗吉尼亚·阿姆布鲁斯特综述硅藻藻-菌关系并展望对于调节海洋生态、预测及调节气候的重要意义 Virginia Armbrust reviewed the interactions between diatoms and algae and looked forward to its importance in regulating marine ecology and predicting and regulating global climate
2014	魏江春院士团队完成首个石果衣真菌(Endocarpon pusillum Hedwig)的全基因组测序 Wei Jiangchun’s team completed the first whole genome sequencing of Endocarpon pusillum Hedwig
2014	杨桥等利用高通量测序技术揭示海洋典型赤潮甲藻的藻际微生物的多样性信息 Yang Qiao’s team revealed the biodiversity of phycosphere microbiota of some typical HAB-causing marine dinoflagellates using high-throughput sequencing technology
2015	弗吉尼亚·阿姆布鲁斯特团队阐明吲哚乙酸的藻菌互作信号新机制 Virginia Armbrust’s team elucidated new signaling mechanism of algae-bacteria interaction by indole-3-acetic acid (IAA)
2015	林森杰团队破译了重要珊瑚共生甲藻虫黄藻(Symbiodinium kawagutii)基因组 Lin Senjie’s team deciphered the genome of Symbiodinium kawagutii which is an important coral symbiotic dinoflagellate
2016	里希拉姆·拉曼等发表藻-菌关系重要综述，展望了其在推动可持续发展的巨大潜力 Rishiram Raman et al. published the review of algae-bacteria interactions, and looked forward to the great potential in promoting sustainable development
2017	贾斯汀·西摩发表重要综述，为揭示水生系统浮游植物与细菌互作机制及意义提供新视角 Justin Seymour published an important review, providing new perspectives on the mechanisms and significance of interactions between phytoplankton and bacteria in aquatic systems
2019	埃米利奥·西里及乔治·波纳特发表综述，总结展望藻菌互作对全球元素循环的重要影响 Emilio Ciri and Giorgio Bonnet published a review and summarized and looked forward to the important impact of algae-bacteria interactions on global element cycles
2019	中山卓郎等发现海洋甲藻中普遍存在共生蓝细菌，为重新认识海洋蓝细菌生态功能提供重要线索 Takuro Nakayama et al. found that symbiotic cyanobacteria were common in marine dinoflagellates, providing important clues for re-understanding the ecological functions of marine cyanobacteria.
2019	穆罕默德·多尼亚等发现藻-菌-动物复杂共生体化学防御机制，表明不同生物体共用化学防御体系 Mohammed Doniya et al. discovered the chemical defense mechanism of algae-bacteria-animal complex symbiosis, indicating that different organisms share a common chemical defense system
2021	法里巴·雷兹瓦尼等综述藻菌互作对废水营养物去除研究进展，展望了藻菌互作增效机制巨大应用潜力 Fariba Rezvani et al. reviewed the research progress on nutrient removal from wastewater by algae-bacteria interactions and looked forward to the huge application potential of the synergistic mechanism of algae-bacteria interactions
2022	《麻痹性贝类毒素生物学研究与应用》学术著作撰写出版 Yang Qiao’s team published the academic book Biological Research and Application of Paralytic Shellfish Toxins
2023	玛利亚·米塔等揭示微藻与互利菌及拮抗菌之间的互作机制，为研究藻菌互作机制及其应用奠定基础 Maria Mitta et al. revealed the interaction mechanism between microalgae and mutually beneficial and antagonistic bacteria, laying a foundation for the study of algae-bacteria interactions mechanism and application
2023	杨桥团队开始推动藻际微生物学新学科的筹建 Yang Qiao’s team began to promote to establish phycosphere microbiology as a new emerging interdiscipline
2024	《藻际微生物学导论》《藻际微生物学：百年百人百事》等藻际微生物学系列丛书陆续出版 Yang Qiao’s team published series of books on Phycosphere Microbiology, including Introduction to Phycosphere Microbiology and Phycosphere Microbiology with Hundreds of Years, Peoples and Events
2024	泰勒·科尔等发现海洋藻固氮菌内共生体进化为硝质体，将原核细胞的功能认识扩展至真核生物 Taylor Cole et al. discovered that the endosymbionts of nitrogen-fixing bacteria of marine microalgae evolved into the nitraplasts, extending the understanding of the functions of prokaryotic cells to eukaryotic organisms

时间Year	重要事件 Key events
启蒙孕育期(几千年前至20世纪中叶) Enlightenment period (from thousands of years ago to the middle 20th century)
1703	人类首次对淡水硅藻——平板藻(Tabellaria)进行细胞形态学观察和记录 The first observation and record of cell morphology of fresh-water diatom Tabellaria
1753	卡尔·林奈在《植物种志》(Speciesl Panatrum)中，将地衣归类到Lihcen属 Carl Linnaeus classified lichens in the genus Lihcen in the book titled Speciesl Panatrum
1798	埃里克·阿卡里乌斯开创地衣分类学，出版多部地衣学著作，推动其成为一门独立学科 Erik Acharius created lichen taxonomy and published several books on lichenology, and promoted it to become an independent discipline
1867	西蒙·施文德纳发现地衣是由真菌与藻共生的复合生物体，揭示了地衣的共生本质 Simon Schwendener discovered that lichens were complex organisms which were composed of fungi and algae, thus revealed the dual nature of the lichens
1872	约翰内斯·莱因克提出“consortium”一词，用来表示构成地衣的藻与真菌的关系 Johannes Reinke coined the term “consortium” to describe the relationship between algae and fungi that make up the lichens
1890	马丁努斯·贝耶林克分离获得藻际细菌 Martinus Beijerinck was the first people who isolated phycosphere bacteria
1915	查尔斯·沃尔科特从前古生代石灰岩藻沉积物化石中发现藻际细菌存在证据 Charles Walcott found the evidence for the existence of phycosphere bacteria in fossilized limestone from the prepaleozoic era
1940s	恩斯特·普林斯海姆分离保存大量藻株并进行分享；为世界主要藻株保藏中心作出了重要贡献 Ernst Prinsheim isolated, preserved, and shared a large number of algae strains, and made significant contributions to major algae collections in the world
创建成长期(20世纪中叶至21世纪10年代) Establishment and growth period (from middle 20th century to 2010s)
1943	迈耶-阿比奇提出全生物体(holobiont)概念及全共生理论(Theory of Holobiosis) Meyer-Abich coined the concept of holobiont and proposed the Theory of Holobiosis
1952	威廉·奥斯特瓦尔德等利用藻-菌共生体系开发生活污水的生物处理技术 Wilhelm Ostwald et al. developed a biological treatment technology for domestic sewage based on the bacteria-algae symbiotic system
1953	哈罗德·米尔纳发明透明容器推动规模化培养微藻细胞，为光生物反应器实际应用奠定重要基础 Harold Milner invented the transparent container to promote the large-scale cultivation of microalgae cells, and laid important foundation for the practical application of photobioreactors
1957	乔治·斯科特提出藻际生命共同体(phycobiont)概念 George Scott coined the concept of phycobiont
1963	萨弗曼发现蓝藻病毒及噬藻体(1969) Saffman discovered cyanobacterial viruses in 1963, and then cyanophages in 1969
1969	詹姆斯·斯图尔特及马尔科姆·布朗首次发现溶藻菌 James Stewart and Malcolm Brown first discovered the algicidal bacteria
1970	林恩·马古利斯提出内共生学说(Endosymbiosis Theory) Lynn Margulis proposed the Endosymbiosis Theory
1972	韦恩·贝尔及拉尔夫·米切尔提出藻际(phycosphere)概念 Wayne Bell and Ralph Mitchell coined the concept of phycosphere
1973	魏江春院士领衔编研《中国地衣志》一书 Wei Jiangchun began to compiled the book titled Chinese Lichens
1978	埃斯特拉·席尔瓦首次发现甲藻胞内细菌；提出著名的“甲藻共生菌产毒”假说 Estela Silva discovered intracellular bacteria in dinoflagellates and proposed the famous hypothesis of “toxic production by symbiotic bacteria of dinoflagellate”
1982	乔纳森·科尔提出菌藻相互作用(interactions between bacteria and algae) Jonathan Cole coined the concept of interactions between bacteria and algae
1988	儿玉正明发现来源于产毒海洋甲藻的藻际自产毒细菌莫拉氏菌PTB-1 Masaaki Kodama discovered the paralytic shellfish poisoning (PSP) toxin-producing bacterium, Moraxella sp. PTB-1 from a PSP toxin-producing marine dinoflagellate
1989	约翰·拉齐尔发现影响藻际生态位大小的因素 John Lazear studied and discovered the factors that influence the size of the phycosphere niche
1994	郑天凌等提出“以菌治藻”新思路 Zheng Tianling et al. proposed a new idea of “bacterial control the harmful algal blooms (HABs)”
1995	格雷戈里·杜塞特发现溶藻菌及藻-菌相互作用 Gregory Doucette discovered algicidal bacteria and algae-bacteria interactions
1999	伯纳德·雅克等提出互作组(interactome)概念 Bernard Jacques et al. coined the concept of interactome
2000	玛丽·莫兰等研究藻际玫瑰杆菌(Roseobacter)类细菌；发现微藻对藻际菌群结构起决定性作用 Mary Moran’s team studied the bacteria of the Roseobacter group in the the phycosphere niche, and found that microalgae play a decisive role in the community structure of the phycosphere microbiota
2003	魏江春院士团队提出“沙漠生物地毯工程” Wei Jiangchun’s team proposed the “Desert Biocarpet Project”
2008	伊拉娜·齐尔伯-罗森伯格等提出全基因组(hologenome)概念 Ilana Zilber-Rosenberg et al. proposed the concept of hologenome
2009	弗吉尼亚·阿姆布鲁斯特团队提出硅藻藻-菌关系 Virginia Armbrust’s team proposed the interactions between diatoms and algae
2009	杨桥团队开展海洋甲藻的可培养藻际微生物的分离及其生物活性功能研究 Yang Qiao’s team isolated the cultivable phycosphere bacteria from toxic marine dinoflagellates and studied their bioactive functions
快速发展期 (21世纪10年代至今) Rapid development period (from 2010s to the present)
2012	弗吉尼亚·阿姆布鲁斯特综述硅藻藻-菌关系并展望对于调节海洋生态、预测及调节气候的重要意义 Virginia Armbrust reviewed the interactions between diatoms and algae and looked forward to its importance in regulating marine ecology and predicting and regulating global climate
2014	魏江春院士团队完成首个石果衣真菌(Endocarpon pusillum Hedwig)的全基因组测序 Wei Jiangchun’s team completed the first whole genome sequencing of Endocarpon pusillum Hedwig
2014	杨桥等利用高通量测序技术揭示海洋典型赤潮甲藻的藻际微生物的多样性信息 Yang Qiao’s team revealed the biodiversity of phycosphere microbiota of some typical HAB-causing marine dinoflagellates using high-throughput sequencing technology
2015	弗吉尼亚·阿姆布鲁斯特团队阐明吲哚乙酸的藻菌互作信号新机制 Virginia Armbrust’s team elucidated new signaling mechanism of algae-bacteria interaction by indole-3-acetic acid (IAA)
2015	林森杰团队破译了重要珊瑚共生甲藻虫黄藻(Symbiodinium kawagutii)基因组 Lin Senjie’s team deciphered the genome of Symbiodinium kawagutii which is an important coral symbiotic dinoflagellate
2016	里希拉姆·拉曼等发表藻-菌关系重要综述，展望了其在推动可持续发展的巨大潜力 Rishiram Raman et al. published the review of algae-bacteria interactions, and looked forward to the great potential in promoting sustainable development
2017	贾斯汀·西摩发表重要综述，为揭示水生系统浮游植物与细菌互作机制及意义提供新视角 Justin Seymour published an important review, providing new perspectives on the mechanisms and significance of interactions between phytoplankton and bacteria in aquatic systems
2019	埃米利奥·西里及乔治·波纳特发表综述，总结展望藻菌互作对全球元素循环的重要影响 Emilio Ciri and Giorgio Bonnet published a review and summarized and looked forward to the important impact of algae-bacteria interactions on global element cycles
2019	中山卓郎等发现海洋甲藻中普遍存在共生蓝细菌，为重新认识海洋蓝细菌生态功能提供重要线索 Takuro Nakayama et al. found that symbiotic cyanobacteria were common in marine dinoflagellates, providing important clues for re-understanding the ecological functions of marine cyanobacteria.
2019	穆罕默德·多尼亚等发现藻-菌-动物复杂共生体化学防御机制，表明不同生物体共用化学防御体系 Mohammed Doniya et al. discovered the chemical defense mechanism of algae-bacteria-animal complex symbiosis, indicating that different organisms share a common chemical defense system
2021	法里巴·雷兹瓦尼等综述藻菌互作对废水营养物去除研究进展，展望了藻菌互作增效机制巨大应用潜力 Fariba Rezvani et al. reviewed the research progress on nutrient removal from wastewater by algae-bacteria interactions and looked forward to the huge application potential of the synergistic mechanism of algae-bacteria interactions
2022	《麻痹性贝类毒素生物学研究与应用》学术著作撰写出版 Yang Qiao’s team published the academic book Biological Research and Application of Paralytic Shellfish Toxins
2023	玛利亚·米塔等揭示微藻与互利菌及拮抗菌之间的互作机制，为研究藻菌互作机制及其应用奠定基础 Maria Mitta et al. revealed the interaction mechanism between microalgae and mutually beneficial and antagonistic bacteria, laying a foundation for the study of algae-bacteria interactions mechanism and application
2023	杨桥团队开始推动藻际微生物学新学科的筹建 Yang Qiao’s team began to promote to establish phycosphere microbiology as a new emerging interdiscipline
2024	《藻际微生物学导论》《藻际微生物学：百年百人百事》等藻际微生物学系列丛书陆续出版 Yang Qiao’s team published series of books on Phycosphere Microbiology, including Introduction to Phycosphere Microbiology and Phycosphere Microbiology with Hundreds of Years, Peoples and Events
2024	泰勒·科尔等发现海洋藻固氮菌内共生体进化为硝质体，将原核细胞的功能认识扩展至真核生物 Taylor Cole et al. discovered that the endosymbionts of nitrogen-fixing bacteria of marine microalgae evolved into the nitraplasts, extending the understanding of the functions of prokaryotic cells to eukaryotic organisms

藻际微生物学：全面助推可持续发展的新兴交叉学科

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张晓玲 , 杨桥

微生物学报 | 综述 2025,65(5): 1831-1848

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微生物学报 | 综述 2025, 65(5): 1831-1848

藻际微生物学：全面助推可持续发展的新兴交叉学科

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张晓玲, 杨桥

作者信息

浙江海洋大学海洋科学与技术学院，浙江舟山

Phycosphere Microbiology: an emerging interdiscipline comprehensively promoting sustainable development

Xiaoling ZHANG, Qiao YANG

Affiliations

College of Marine Science and Technology, Zhejiang Ocean University, Zhoushan, Zhejiang, China

出版时间: 2025-05-04 doi: 10.13343/j.cnki.wsxb.20240783

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摘要

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广泛分布在自然界中的藻类和细菌均是地球上最古老的生命形式，经过数十亿年的漫长自然进化，藻类与细菌演化成为以藻-菌相互作用为核心、“牵一发而动全身”的藻际生命共同体。构建这一复杂而精妙的藻菌共生系统的科学理论体系，并对其进行创新和发展，不仅对于人类探索生命起源与进化的奥秘具有重要的科学意义，而且，其在推动可持续发展的重要领域也展现出广阔的应用前景。在此背景下，作为微生物学的一个新的分支学科，藻际微生物学已经历了3个多世纪的孕育和发展。目前，推动其成为一门新兴交叉学科的时机已经到来。为此，本文深入探讨了藻际微生物学的核心概念；全面梳理了藻际生命共同体与地球环境保护、人类生命与健康、资源综合利用及绿色能源开发之间的紧密关系；系统回顾了藻际微生物学的发展历史，并归纳总结了其3个发展时期的主要研究成果。本文还对藻际微生物学的未来发展趋势进行了展望，并提出了若干潜在的交叉研究方向。

关键词

藻际微生物学 / 藻际生命共同体 / 藻际微生物 / 藻-菌相互作用 / 新兴交叉学科

Abstract

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Algae and bacteria are both the oldest forms of life on our planet, and billions of years of natural evolution have driven the algae and their microbiomes to evolve into interactive phycobionts. Through complex, flexible, intelligent, and multi-interface interactions between algae and bacteria, the functions of both sides of the phycobionts are exquisitely regulated. The creation, innovation, and development of the phycobiont theory shows vital scientific value for revealing the mystery of the origin and evolution of the life on Earth, and this theory is also being transformed into diverse practical applications in significant fields for sustainable development. After centuries of incubation, knowledge accumulation and development, currently, it is the right time to promote Phycosphere Microbiology to develop into an emerging interdiscipline. In this review, we comprehensively discussed the core concepts of Phycosphere Microbiology, sorted out its vital relationships with environment protection, human health maintenance, resource utilization, and green-oriented transition of energy, then reviewed its development history, and summarized the main research achievements during three development periods. Finally, we also proposed and discussed the future development trends and potential research directions for this emerging interdiscipline.

Key words

Phycosphere Microbiology / phycobiont / phycosphere microbiome / interactions between algae and bacteria / emerging interdiscipline

引用本文

张晓玲, 杨桥. 藻际微生物学：全面助推可持续发展的新兴交叉学科. 微生物学报, 2025 , 65 (5) : 1831 -1848 . DOI: 10.13343/j.cnki.wsxb.20240783

Xiaoling ZHANG, Qiao YANG. Phycosphere Microbiology: an emerging interdiscipline comprehensively promoting sustainable development[J]. Acta Microbiologica Sinica, 2025 , 65 (5) : 1831 -1848 . DOI: 10.13343/j.cnki.wsxb.20240783

正文

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目前，人类正面临着日益严峻的环境、健康、资源及能源等全球性难题，而坚持全面协调可持续发展是破解这些难题的“金钥匙”，也成为构建人与自然生命共同体的重要基石^[1]。然而，在实现可持续发展的过程中，如何高效协调资源与环境等关键性环节，是人类面临的巨大挑战，同时也是未来科技与行业革命的关键突破口^[2]。科学知识体系化下新兴交叉学科的创建，将为推动人类社会高质量发展注入强大的动能^[3]。

藻类与细菌均是地球上最古老的生命形式。在长达数十亿年的漫长自然进化过程中，藻及其藻际微生物组演化形成了以藻菌互作为核心、“牵一发而动全身”的藻际生命共同体(phycobiont)^[4]，在维系人类生存与发展过程中，显现出了重要的科学价值及广阔的应用前景^[5]。微生物学虽然仅有100多年的发展历史，但它作为一门极具生命力的学科，为推动人类可持续发展作出了巨大的贡献^[6]。相比之下，作为微生物学的一个新的分支学科，藻际微生物学已经历了3个多世纪的启蒙、孕育和发展，目前正呈现出生机勃勃的发展趋势。为了科学凝练藻际微生物学的学科发展规律并探究其未来发展方向，从而推动多学科交叉融合下这一新兴交叉学科的快速成长，本文系统回顾总结了藻际微生物学的发展历程，并对其未来发展趋势进行了展望。

1 藻际生命共同体

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1.1 古老藻类——自然界中的“多面手”

蓝细菌(旧称蓝绿藻或蓝藻)作为地球上早期的生命形式之一，正是因其利用自身的光合作用能力而释放氧气，才使得地球上真核生物等高等生命形式的诞生成为可能^[7]。藻类具有极丰富的物种及超强的环境适应力，遍布地球的水圈、生物圈和岩石圈，并通过O₂及CO₂等气体与大气圈紧密相连^[8]。漫长的自然进化过程使藻类成为了地球上生命起源与进化的见证者，地球上最重要的初级生产者之一，以及水生生物多样性的重要维持者^[9-10]。此外，藻类自身就具有重要的经济价值。其不仅贡献了地球上50%以上的O₂，还成为了地球碳循环与海洋碳汇的关键驱动者^[10-11]、全球的气候调控者以及重要的生物修复物种^[12]。因此，藻类被誉为自然界中的“多面手”^[13]。

1.2 藻及其微生物组

藻类广泛地分布在地球上，而围绕着藻细胞周围形成了以藻为核心、富含营养物质的微米尺度的特殊微环境，即藻际(phycosphere)^[14]。这一独特微型生态位的化学物质基础以多糖、蛋白质等胞外聚合物为主^[15]。藻细胞正是借由藻际招募并供养特定类型的微生物种群在此聚集和定殖，从而构建了具有稳定共生生物体结构的藻际微生物群落^[16-18]。藻际微生物既可生存于藻细胞的周围，也可共存于胞内的细胞质和细胞核等细胞器内^[17-19]。藻际微生物的基因数量远大于宿主藻，因此也被称为藻类的第二基因组，从而构成了藻细胞结构与功能的必需元件^[16-18]。由此，“藻-藻际-微生物”通过复杂的跨界相互作用，构建成为密不可分的藻际生命共同体^[20]。这与广泛存在于地球陆地生态系统中的根际生命共同体(rhizobiont)较为相似^[21-22]。藻际生命共同体早已渗透在自然、农业、医药及工程技术等领域中，为推动人类可持续发展发挥着重要作用^[23-25]。

藻际生命共同体理论^[20-25]的创新与发展，推动了藻际概念内涵的不断延伸与扩展，从最初的狭义概念(仅指藻细胞周围的微环境)^[14]，进一步扩展至广义的概念(同时包括藻细胞内部的微环境)^{[19-20,23-24]}。根据藻-菌在藻际生命共同体中所处地位的不同，可将其分为2大类型。一类以藻类为主体，被称作藻-菌共生系统(algae-bacteria symbiotic system)^[14-18]；另一类主要包括广泛分布在陆地的地衣(lichen)生物，它们是由地衣型真菌作为建群种、藻类作为伴生种而构建形成的稳定共生生命系统，被称作菌-藻共生系统(bacteria-algae symbiotic system)^[20]。

藻际微生物蕴含着从中发掘新颖生物合成基因及新型天然活性产物的巨大潜力，其已成为人类赖以生存与发展的生物基础的重要组成部分^[22-25]。然而，由于微生物个体微小，且大多数难以分离培养，加上受到方法学局限性等因素的制约^[26-27]，藻际微生物所蕴含的“四两拨千斤”的巨大潜能却极易被人们忽略或者轻视。对于奥妙的藻际生命共同体而言，长期以来过于倚重藻类一方，而未能充分发掘利用藻菌互作增效的巨大潜力，这极大限制了藻际生命共同体的巨大潜能的全面释放^[23,25,27]。因此，亟须系统探究藻际生命共同体的藻菌互作增效机制，发掘其系统生物学的调控新思路，并推动其在可持续发展中的广泛应用。这不仅对于破解环境、健康、资源及能源等全球性难题具有重要的现实意义，由此而产生的经济效益与社会效益也将难以估量。

1.3 藻际生命共同体与可持续发展

在漫长的自然进化过程中，藻-菌之间形成了互惠、共生、寄生或拮抗等错综复杂的藻-菌相互作用关系^[22-24]。藻际生命共同体通过藻-藻、藻-菌、菌-菌之间的多界面相互作用，精妙地调控着藻-菌的功能^[22-25]。藻际生命共同体科学理论的创新与发展，不仅对人类探索生命的起源与进化的奥秘具有重要的科学意义，而且其在环境、健康、资源与能源等重要领域也获得了广泛的应用^[23-29]，从而成为推动人类社会高质量发展的中坚力量。

1.3.1 生命起源与进化

藻类与细菌共同见证了地球生命的起源与进化，而共生现象也早已遍布在自然界中，如根瘤菌与豆科植物^[30]、地衣型真菌与蓝细菌或绿藻^[31]等。蓝细菌胞内共生现象的发现，启发了人类开始思考真核生物内重要且独特的自主细胞器——叶绿体的起源。林恩·马古利斯(Lynn Margulis)^[32]系统总结了康斯坦丁·梅列施柯夫斯基(Konstantin Merezhkovsky)^[33]、伊万·沃林(Ivan Wallin)^[34]等科学家的研究发现，进而提出了现代生物学的开创性理论之一的内共生学说(endosymbiotic theory)^[35-36]。该学说的诞生，促使人类开始思考并逐渐认识到，与人类共生及进化的微生物是人类生存与发展不可分割的一部分。相比之下，“人与自然和谐共生”、构建“人与自然生命共同体”等一系列重要科学理念的提出，正是源于人类对自然界共生现象的深刻哲学思考与探索实践^[1,21-25]。

1.3.2 地球环境保护

作为海洋碳汇的重要驱动者和全球气候的调节者，浮游藻类是海洋中最主要的初级生产者，其光合作用产物构成了海洋食物链的物质和能量基础。藻际富含多糖、蛋白质、核酸等胞外多聚物，形成了重要的海洋微型碳库^[10,27]。此外，藻际微生物是海洋碳循环的重要调节者，它们通过高效的碳转化过程实现了长达数千年的碳储存，对于维持地球水生生物多样性、调控海洋碳汇及全球气候作出了巨大贡献^[37]。

早在20世纪40年代，人类就已开始利用藻-菌共生体系进行污水生物处理^[38]。由此创立了著名的藻-菌系统(algae and bacteria system)，并在氮磷、重金属、抗生素药残、多环芳烃、石油等水体污染物的生物降解与资源化利用等重要领域获得了广泛应用^[39]。目前，国内外普遍使用的高速率藻塘(high rate algal pond, HRAP)^[40]等主流水处理技术的核心，均基于藻-菌共生体系。此外，典型藻毒素-麻痹性贝类毒素来源的“甲藻内共生菌产毒”假说的提出^[41-42]、溶藻菌(Streptomyces、Bacillus及Shewanella等)^[43]及其活性物质(酶、生物碱、羟胺、环肽及醌等)^[44]、原核藻类病毒——噬藻体(cyanophage)^[45]、真核藻类病毒(phycovirus)^[46-47]的发现等一系列重要科学研究成果的取得，不仅推动了“以菌治藻”^[48]成为极具应用潜力的赤潮治理新技术，从而为全球有害藻华的科学防控指明了新方向，同时也成为了人类智慧与自然资源融合、推动人与自然和谐共生的经典案例。

1.3.3 守护人类健康

中国历史上关于地衣的记录最早见于《诗经》中对女萝(松萝，松萝科松萝属地衣)的描绘^[49]。此外，我国古代医学名著如《名医别录》^[50]、《齐民要术》^[51]、《药性本草》^[52]及《本草纲目》^[53]中均有将地衣和藻类作为药材使用的记载。人类很早就开始利用藻际生命共同体系统生产胡萝卜素、藻蛋白、多糖、酶类、抗生素等活性物质服务于人类的生产生活^[54-55]。此外，藻-菌共生体系能够高效协同去除水体中的多种有害物质从而净化水体，同时为水产养殖生物提供丰富的饵料，实现变废为宝。基于此，人类发明了虾-贝-鱼-菌-藻、鱼-虫-虾-贝-菌-藻等绿色、立体生态循环养殖技术并进行了广泛应用，有力推动了水产养殖业的绿色可持续发展^[56]。藻际生命共同体在功能食物、保健品、农业生产、水产疾病防治及生物医药研发等重要领域，同样展现出了广阔的应用前景，逐步发展成为守护人类生命与健康的重要力量^[57-59]。

1.3.4 生物资源利用

自1890年人类首次分离获得藻际细菌以来^[60]，藻际微生物资源逐渐被人类发掘并应用到日常生产生活中。然而，目前其大部分微生物尚难以获得纯培养物，它们被称为“微生物暗物质(microbial dark matter)”^[61]。藻际生命共同体蕴含着大量的未知生物物种，其大部分资源尚且处于“沉寂”状态，亟待进行深入系统的发掘与利用。此外，作为藻类的第二基因组，藻际微生物中蕴藏的新颖合成基因及新型天然活性产物的巨大潜力同样不容小觑^[4,27]。随着培养组学、系统生物学等新技术的不断涌现，突破困扰微生物学百年的微生物低培养性的研究瓶颈，必将不断获得新的研究进展，由此也将发现数量更多的藻际微生物菌株新资源，并将其活性功能推向多领域的实际应用^[26]。合成生物学、人工智能等多组学技术的联用，也将为推动藻际微生物资源的巨大潜力全面转化为新质生产力创造新的发展契机。

1.3.5 绿色能源开发

日益严峻的全球能源危机促使全球各国竞相发展绿色新能源^[62]。其中，微藻被视为极具应用潜力的生物燃料生产原料，如柴油、甲烷、乙醇、氢气及丁醇等^[63]。藻-菌共生体自絮凝技术的出现，突破性地实现了产油藻细胞的高效收获，从而大幅降低了产油微藻的生产成本，且该培养体系不仅不易被污染，而且油脂产率和高附加值副产物产量高。此外，藻际促生菌的发现显著提高了藻细胞生长速度、脂质含量及产物化学多样性，由此催生了一系列微藻生物质能源新技术^[64]。基于藻-菌共生体系的废污水处理技术，可将污水中的多种污染物高效转化为微藻生物质柴油，同时有效降低二氧化碳排放，实现污水处理与生物资源的高效回收利用。该技术已发展成为符合“双碳”发展内在需求的绿色生物制造新技术，为保障国家能源安全发挥重要作用^[65-66]。

2 藻际微生物学的发展历史

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藻际微生物学(Phycosphere Microbiology)是研究藻际生命共同体的基本生物特征，探索并揭示其生命活动的基本规律和生命现象的本质的一门新兴交叉学科。其研究对象是广泛存在于自然界中的藻际生命共同体中的微小生物，它们栖息于藻细胞周围及其胞内的藻际生境中。其种类包括细菌、古菌、放线菌、霉菌、地衣型真菌、酵母、病毒、噬菌体、微藻及其他未知生物物种。藻际微生物学的研究内容涵盖了分子、细胞、物种、群落、生态系统等多个水平，旨在研究藻际微生物的形态、生长、生理代谢、遗传、生态、分类、进化等基本特征及其规律；揭示藻际生命共同体的相互作用过程及其本质，并推动其在环境、健康、资源及能源等可持续发展领域的广泛应用(图1)。

藻际微生物学作为微生物学的一个新的分支学科，相较于海洋微生物学、土壤微生物学等其他分支学科，其出现时间较晚。藻际微生物学具有长达3个多世纪的发展历史，可被划分为3个时期。其中包括：启蒙孕育期、创建成长期及快速发展期(图2)。藻际微生物学在研究藻际微生物的多样性及其功能、揭示藻菌互作机制及其应用方面取得了一系列重要的研究成果(表1)。

2.1 启蒙孕育期(几千年前至20世纪中叶)

2.1.1 20世纪中期之前：藻际微生物的初步认识

早在几千多年前，人类就有食用或药用藻类的古老记载，但人们对藻细胞的观察最早只能追溯到18世纪初期，当时伦敦皇家学会收到的一篇稿件中记录了附生于杂草根部或游离在水中的硅藻——平板藻(Tabellaria)的细胞形态插图^[67]。1915年，美国学者从前古生代(2.3-5.7亿年前)石灰岩藻沉积物化石中首次发现了藻际细菌的存在证据^[68]。人类成功分离获得活的藻际细菌也仅仅发生在一个多世纪之前。1890年，马丁努斯·贝耶林克(Martinus Beijerinck)在对藻细胞进行纯化时首次分离出一些藻际细菌^[60]，他也因此成为首位获得藻、动物共生藻(zoochlorellae)及地衣纯培养物的科学家。尽管早期人们在进行藻细胞纯化时分离出了藻际细菌，但当时普遍将其视为藻细胞的污染物。此外，在该发展时期内，与藻际生命共同体相关的一系列重要概念被陆续提出，如孢丝粉体(mazaedium)^[69]、共生(symbiosis)及共生体(homobium)^[70]、趋化性(chemotaxis)^[71]、菌根(mykorrhiza)^[72]、副共生(parasymbiose)^[73]、假副共生(pseudoparasymbiose)^[74]、真共生(eusymbiose)^[75]等。当时对于藻际微生物的研究主要停留在形态学描述及个体细胞观察的初级阶段，而对于藻际微生物资源在人类生产生活中的重要性尚缺乏足够的深入认识。尽管如此，这些对于藻际微生物的初步探索性研究成果的获得，标志着人类认识及利用藻际微生物资源的开始。

2.1.2 地衣——陆地藻际生命共同体的最完美典范

广泛分布于陆地表面的地衣生物，是藻际生命共同体中菌-藻共生系统的代表，被誉为地球生物共生的最完美典范^[76]。2005年，中国学者对距今约6亿年的磷块岩类地衣化石的重要发掘研究，首次证实了可能早在地球古海洋时代，真菌与蓝细菌就已发展成为相互依存的共生关系；在苔藓和蕨类植物登陆之前，地衣就已经开始改造地球的岩石圈，因此地衣极有可能是最早登陆的地球生物之一^[77]。地衣也被誉为“生命拓荒者”、人类未来移居外星球的“先锋生物”^[78]。丰富的地衣生物资源早已被广泛应用于医药、食品、化工及环保等领域^[76,79]，为推动人类可持续发展作出了重要贡献。

18世纪末至19世纪初，地衣学之父埃里克·阿卡里乌斯(Erik Acharius)所撰写的Lichenographiae Suecicae Prodromus^[80]、Methodus qua omnes Detectos Lichenes^[81]、Lichenographia Universalis^[82]以及Synopsis Methodica Lichenum^[83]等地衣学重要学术著作相继出版。这些重要研究成果的获得，不仅孕育了藻际生命共同体的科学理论萌芽，同时也奠定了现代地衣学的基础，使地衣学逐渐成长为一门独立的学科。1867年，西蒙·施文德奈尔(Simon Schwendener)^[84]发现地衣是由真菌和藻类共生而形成的生物复合体，这一发现揭开了长期悬而未决的“地衣之谜”，成为地衣科学史上的首个里程碑事件。此外，一些沿用至今的藻际生命共同体重要概念，如二元性(dual nature)^[84]、联合体(consortium)^[85]等均与地衣生物有关。中国地衣学科的开创者和主要奠基人魏江春院士，在藻际生命共同体领域创建了一系列重要研究成果^[86]。

2.2 创建成长期(20世纪中叶至21世纪10年代)

2.2.1 重要概念的创建

1957年，乔治·斯科特(George Scott)提出了藻际生命共同体的概念^[20]。1972年，韦恩·贝尔(Wayne Bell)和拉尔夫·米切尔(Ralph Mitchell)创建了藻际微生物学的核心概念藻际(phycosphere)^[14]。此外，菌-藻相互作用(interactions between bacteria and algae)^[23]、租借共生(mietsymbiose)及地衣共生(flechtensymbiose)^[87]、群体感应(quorum sensing, QS)^[88]、互作组(interactome)^[89]及全基因组(hologenome)^[90]等一系列与藻际生命共同体相关的重要概念也被陆续提出。同时，一些关键性研究技术也被发明并获得推广应用，如藻细胞规模培养光生物反应器^[39]、藻无菌化技术^[91]、藻际细菌特异性分离免疫磁珠法^[92]、高速率藻塘(HRAP)技术^[40]等。在该时期内，人们利用藻-菌共生体系进行富营养废水的生物修复^[38]。藻内共生菌^[93]、溶藻菌及其活性物质^[23]、噬藻体^[45]及藻病毒^[46-47]的陆续发现，极大地拓展了人们对藻际微生物物种丰富程度及巨大应用潜力的新认识。

2.2.2 内共生学说——藻际生命共同体的经典案例

内共生学说是林恩在继承发展内共生假说的基础上而提出的科学新理论，该理论认为生物的共生起源构成了进化论的基础。这一超前的科学理论一经问世，就引起了当时生物舆论界的广泛关注，并引发了诸多批评之声，这也直接导致林恩的最重要著作《真核细胞的起源》(Origin of Eukaryotic Cells)^[35]被推迟到1970年才获得出版。然而，该学说自诞生以来陆续获得了大量科学研究证据的支持与佐证。该学说的创建过程，也成为了人类科学发展史上关于藻际生命共同体研究的经典案例。

2.2.3 分子生物学时代的快速发展

从20世纪80年代起，随着第一代测序、基因芯片等新技术的兴起^[94]，生物学研究进入了分子生物学时代。藻际微生物这一独特的生物种群逐渐引起了人们的重视，这一新兴研究领域也陆续取得了一系列重要的研究进展。1978年，埃斯特拉·席尔瓦(Estela Silva)提出了著名的“甲藻内共生菌产毒假说”^[41-42]；藻菌互作的典型研究示例，如维生素B₁₂^[95]及铁载体(siderophore)^[96]的功能被陆续揭示；重要珊瑚共生甲藻虫黄藻(Symbiodinium kawagutii)的基因组序列被解析^[97]。同时，一系列重要的科学理论或研究思路，如“海洋微型生物碳泵”^[98]、“以菌治藻”^[48]、“沙漠生物地毯工程”^[99]等被陆续提出。藻际生命共同体在绿色能源开发、蓝碳理论与应用、地球环境保护及生物治理修复、功能食品与天然药物等重要领域的巨大应用潜力也被不断发掘^{[18,96,100-102]}。在此发展时期内，也随之相继涌现出来Susan Gallacher^[103-108]及魏江春^[109-113]等一大批杰出科学家及其研究团队。

在此发展时期内，人类对藻际微生物的生命活动及其对生产生活的影响进行了系统性研究，由此也推动了藻际微生物学研究从早期的形态描述，上升到了生理生化与分子生物学水平。这一系列藻际微生物学研究新发现的获得，不仅进一步丰富了藻际生命共同体的科学理论体系，同时，也奠定了藻际微生物学作为一门独立学科的重要基础，从而开创了藻际微生物学研究的黄金时期。

2.3 多组学助力的快速发展期(20世纪10年代至今)

21世纪初以来，随着高通量测序、质谱及计算机等新技术的迅猛发展，生物学研究从单一组学迈入了多组学(multiomics)时代^[114]。自此，多组学技术成为藻际微生物学的主流研究工具，并取得了一系列重要研究进展。例如，在藻际生命共同体中发现了一些重要的天然活性物质，如roseobacticide B^[115]等；揭示了藻菌互作信号，如二甲基巯基丙酸(dimethylsulfoniopropionate, DMSP)^[116]及二甲基亚砜丙酸内盐(dimethylsulfoxonium propionate, DMSOP)^[117]；2015年，Amin等^[118]报道了吲哚乙酸(indole-3-acetic acid, IAA)在藻菌互作中的关键信号传导功能，自此IAA被视作藻际生命共同体中跨界互作的交互性语言(cross-talking language)，为揭示藻-菌跨界互作的本质及其实际应用提供了新思路^[119]。2019年，科学家通过单细胞基因组学研究首次发现，全球海洋甲藻中广泛隐藏着共生蓝细菌，这一发现暗示人们需要重新认识藻际生命共同体在自然水体环境中的普遍性，以及其对促进人类探索自然界生命奥秘的重要意义^[120]。同年，Science发表了关于藻-菌-海洋动物共生体三方动态互动的化学防御机制的研究，发现自然界的多生物共生体可通过复杂的共生机制分享重要的化学防御分子，从而高效抵御外部猎食者；这一重大发现暗示着以防御分子为中心的细菌-食物-捕食者的共生系统可能广泛存在于海洋环境中^[121]。2024年，Science报道了关于海洋微藻的固氮内共生体硝质体(nitroplast)的重大科学发现，更是将人类对于原核细胞固氮功能的原有认识，颠覆性地扩展到了真核生物界^[122]。

目前，随着人类所面临的环境、资源等全球性挑战的日益严峻，藻际生命共同体愈发显现出其在全面助推可持续发展中的巨大潜能。藻际生命共同体能够通过生物制造生产一系列对人类生产生活至关重要的替代品，包括创新药物及功能食品等高值化产品；绿色生物质燃料和可降解塑料重要原料羟基丁酸酯等中值化学品，以及水产养殖食品及动物饲料等低值产品^[22,27,123]。此外，藻际生命共同体在废/污水生物处理、有毒有害物生物修复等重要领域将继续发挥其重要作用，为推动人与自然和谐共生贡献力量。在此发展时期内，一系列关于藻际生命共同体的重要综述陆续发表^{[12,16-17,24,27,30,44,123-134]}。在检索到的300多篇综述(截至2024年12月)中，有116篇论文引用超过百次，12篇论文引用已超过1 000次。这些重要研究成果的取得不仅表明藻际微生物学领域已成为当前多学科交叉的研究热点，而且它们进一步丰富扩展了藻际生命共同体科学理论体系的内涵。由此，也标志着藻际微生物学研究迈入了多组学技术助力、深入揭示藻际生命共同体的生命规律与本质的系统生物学研究新时代。

目前，我国藻际微生物学领域研究方兴未艾。从事该新兴研究领域的科研人员尤其是青年科研工作者人数逐年攀升，相关研究资助也呈现出稳步上升的趋势。获得的标志性研究成果及国际影响力也在持续提升，从而展现出藻际微生物学这一新兴交叉学科的蓬勃发展的良好态势。其中，藻际微生物学系列丛书，包括《麻痹性贝类毒素生物学研究与应用》^[135]、《藻际微生物学导论》^[136]、《藻际微生物学：百年百人百事》^[137]、《可培养藻际微生物的发现及其应用》^[138]陆续出版，为全面推动这一新兴学科的快速发展奠定坚实的基础^[139-142]。随着当前多学科深度交叉融合的不断深入，也必将为藻际微生物学的快速发展注入新的活力。

3 展望

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藻际微生物学作为微生物学的新分支，已发展成为融合了自然、农业、医药及工程技术科学等多个领域的新兴交叉研究领域。然而，藻际生命共同体理论的创新发展和应用仍面临诸多亟待解答的重要科学问题。藻际微生物资源作为维系人类生存发展基础的重要组成部分，其中蕴含着丰富的新物种、新颖基因及其新型活性天然产物的巨大潜力，亟待对其进行系统性发掘利用。尽管对藻际生命共同体互作增效机制有了新的认识，但其在全面助推可持续发展中的巨大应用潜力，尚未被完全发挥出来。随着合成生物学、人工智能等新技术的涌现与发展，藻际生命共同体与生物经济的融合必将不断加深，并日益加速未来科技与行业革命进程，从而为藻际微生物新资源转化为新质生产力创造新契机。因此，藻际微生物学未来的交叉创新研究可以集中在以下3个方面。

(1) 构建并完善藻际生命共同体的核心科学理论体系。藻菌互作驱动着由“藻-藻际-微生物”构成的藻际生命共同体的稳步运行。其中蕴含着复杂的物质流、信息流及能量流。这些流动之间的相互影响、动态平衡、互作功能的级联放大及其互联互通途径与机制是什么？藻与菌如何通过合成并释放跨界信号分子来调控其复杂的相互作用过程？通过多学科交叉融合，有望在阐析藻际生命共同体的生物及化学组成及其功能方面不断取得新突破。辨明其中的物质流、能量流和信息流的互通过程与本质，进而揭示藻际生命共同体的运行规律及互作机制；并突破传统研究过分偏重单方作用或单一过程而忽视或削弱整体潜能的瓶颈，从而开创藻-菌跨界互作增效的交叉创新研究领域。

(2) 贯通藻际生命共同体互作协同增效机制的关键环节。基于藻际生命共同体的科学理论体系，发掘并创立其多界面互作增效的调控新途径，对于深度释放藻际生命共同体的巨大潜力，从而全面助推人类可持续发展显得至关重要。藻际生命共同体如何协调藻-藻、藻-菌、菌-菌之间的多界面相互作用以精确调控藻-菌功能？在此过程中，哪些特定微生物种群承担了核心功能？其藻际核心功能群的群落装配与稳定机制是如何维持高效运作的？它们如何与宿主藻进行营养与能量等物质交换及跨界信号分子的传递？藻细胞又是如何感知及调控其核心功能类群的？解答这一系列重要科学问题，有望系统阐明藻际微生物核心功能群的定殖、互作信号传递与调控机制，从而将藻际生命共同体的多界面互作认知提升到以功能性应用为核心的新高度。

(3) 推动多学科的深度交叉融合研究与应用。进一步整合利用系统生物学、生物信息学、机器学习等人工智能研究工具，同时高度关注方法学的持续创新，有效提升对海量生物学数据的解读效率。构建开放、共享的藻际生命共同体互作知识数据库与生态网络信息库。同时，系统开展多组学助力下的藻际微生物可培养性关键技术攻关，力求突破微生物学的重要研究瓶颈，为阐析藻际生命共同体互作增效机制，并推动其在可持续发展的广泛应用奠定重要基础。

藻菌生命共同体与保护人类生命健康及环境等息息相关，已成为助推全面协调可持续发展的中坚力量。随着对藻际生命共同体的认识与应用的不断深入，藻际微生物学作为一门新兴交叉学科，有望为人类应对日益严峻的环境与资源等全球性重大挑战提供新的解决方案，从而为全面协调人类可持续发展注入新的强大动能。因此，藻际微生物学方兴未艾，未来发展前景广阔。

基金

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国家自然科学基金(41876114)
国家自然科学基金(41206093)
浙江省属高校基本科研业务费(创新群体项目)(2024J006)
浙江省自然科学基金(LY23D060005)
浙江省自然科学基金(LY18D060007)
浙江省高等教育学会高等教育研究课题(KT2024099)
浙江省“十四五”重点教材建设项目
浙江省产学合作协同育人项目
浙江省“十四五”本科省级教改项目(JGBA2024256)

参考文献

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文献

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2025年第65卷第5期

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doi: 10.13343/j.cnki.wsxb.20240783

接收时间：2024-12-05
首发时间：2026-02-05
出版时间：2025-05-04

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收稿日期：2024-12-05
录用日期：2025-01-19

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National Natural Science Foundation of China(41876114)

国家自然科学基金(41876114)

National Natural Science Foundation of China(41206093)

国家自然科学基金(41206093)

Fundamental Research Funds for Zhejiang Provincial Universities (Innovation Group Project)(2024J006)

浙江省属高校基本科研业务费(创新群体项目)(2024J006)

Natural Science Foundation of Zhejiang Province(LY23D060005)

浙江省自然科学基金(LY23D060005)

浙江省自然科学基金(LY18D060007)

Project of Higher Education Society of Zhejiang Province(KT2024099)

浙江省高等教育学会高等教育研究课题(KT2024099)

Key Textbook Project of Zhejiang for the 14th Five-year Plan

浙江省“十四五”重点教材建设项目

Industry-University Collaborative Education Project of Zhejiang

浙江省产学合作协同育人项目

Undergraduate Teaching Reform Project of Zhejiang(JGBA2024256)

浙江省“十四五”本科省级教改项目(JGBA2024256)

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浙江海洋大学海洋科学与技术学院，浙江舟山

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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时间Year	重要事件 Key events
启蒙孕育期(几千年前至20世纪中叶) Enlightenment period (from thousands of years ago to the middle 20th century)
1703	人类首次对淡水硅藻——平板藻(Tabellaria)进行细胞形态学观察和记录 The first observation and record of cell morphology of fresh-water diatom Tabellaria
1753	卡尔·林奈在《植物种志》(Speciesl Panatrum)中，将地衣归类到Lihcen属 Carl Linnaeus classified lichens in the genus Lihcen in the book titled Speciesl Panatrum
1798	埃里克·阿卡里乌斯开创地衣分类学，出版多部地衣学著作，推动其成为一门独立学科 Erik Acharius created lichen taxonomy and published several books on lichenology, and promoted it to become an independent discipline
1867	西蒙·施文德纳发现地衣是由真菌与藻共生的复合生物体，揭示了地衣的共生本质 Simon Schwendener discovered that lichens were complex organisms which were composed of fungi and algae, thus revealed the dual nature of the lichens
1872	约翰内斯·莱因克提出“consortium”一词，用来表示构成地衣的藻与真菌的关系 Johannes Reinke coined the term “consortium” to describe the relationship between algae and fungi that make up the lichens
1890	马丁努斯·贝耶林克分离获得藻际细菌 Martinus Beijerinck was the first people who isolated phycosphere bacteria
1915	查尔斯·沃尔科特从前古生代石灰岩藻沉积物化石中发现藻际细菌存在证据 Charles Walcott found the evidence for the existence of phycosphere bacteria in fossilized limestone from the prepaleozoic era
1940s	恩斯特·普林斯海姆分离保存大量藻株并进行分享；为世界主要藻株保藏中心作出了重要贡献 Ernst Prinsheim isolated, preserved, and shared a large number of algae strains, and made significant contributions to major algae collections in the world
创建成长期(20世纪中叶至21世纪10年代) Establishment and growth period (from middle 20th century to 2010s)
1943	迈耶-阿比奇提出全生物体(holobiont)概念及全共生理论(Theory of Holobiosis) Meyer-Abich coined the concept of holobiont and proposed the Theory of Holobiosis
1952	威廉·奥斯特瓦尔德等利用藻-菌共生体系开发生活污水的生物处理技术 Wilhelm Ostwald et al. developed a biological treatment technology for domestic sewage based on the bacteria-algae symbiotic system
1953	哈罗德·米尔纳发明透明容器推动规模化培养微藻细胞，为光生物反应器实际应用奠定重要基础 Harold Milner invented the transparent container to promote the large-scale cultivation of microalgae cells, and laid important foundation for the practical application of photobioreactors
1957	乔治·斯科特提出藻际生命共同体(phycobiont)概念 George Scott coined the concept of phycobiont
1963	萨弗曼发现蓝藻病毒及噬藻体(1969) Saffman discovered cyanobacterial viruses in 1963, and then cyanophages in 1969
1969	詹姆斯·斯图尔特及马尔科姆·布朗首次发现溶藻菌 James Stewart and Malcolm Brown first discovered the algicidal bacteria
1970	林恩·马古利斯提出内共生学说(Endosymbiosis Theory) Lynn Margulis proposed the Endosymbiosis Theory
1972	韦恩·贝尔及拉尔夫·米切尔提出藻际(phycosphere)概念 Wayne Bell and Ralph Mitchell coined the concept of phycosphere
1973	魏江春院士领衔编研《中国地衣志》一书 Wei Jiangchun began to compiled the book titled Chinese Lichens
1978	埃斯特拉·席尔瓦首次发现甲藻胞内细菌；提出著名的“甲藻共生菌产毒”假说 Estela Silva discovered intracellular bacteria in dinoflagellates and proposed the famous hypothesis of “toxic production by symbiotic bacteria of dinoflagellate”
1982	乔纳森·科尔提出菌藻相互作用(interactions between bacteria and algae) Jonathan Cole coined the concept of interactions between bacteria and algae
1988	儿玉正明发现来源于产毒海洋甲藻的藻际自产毒细菌莫拉氏菌PTB-1 Masaaki Kodama discovered the paralytic shellfish poisoning (PSP) toxin-producing bacterium, Moraxella sp. PTB-1 from a PSP toxin-producing marine dinoflagellate
1989	约翰·拉齐尔发现影响藻际生态位大小的因素 John Lazear studied and discovered the factors that influence the size of the phycosphere niche
1994	郑天凌等提出“以菌治藻”新思路 Zheng Tianling et al. proposed a new idea of “bacterial control the harmful algal blooms (HABs)”
1995	格雷戈里·杜塞特发现溶藻菌及藻-菌相互作用 Gregory Doucette discovered algicidal bacteria and algae-bacteria interactions
1999	伯纳德·雅克等提出互作组(interactome)概念 Bernard Jacques et al. coined the concept of interactome
2000	玛丽·莫兰等研究藻际玫瑰杆菌(Roseobacter)类细菌；发现微藻对藻际菌群结构起决定性作用 Mary Moran’s team studied the bacteria of the Roseobacter group in the the phycosphere niche, and found that microalgae play a decisive role in the community structure of the phycosphere microbiota
2003	魏江春院士团队提出“沙漠生物地毯工程” Wei Jiangchun’s team proposed the “Desert Biocarpet Project”
2008	伊拉娜·齐尔伯-罗森伯格等提出全基因组(hologenome)概念 Ilana Zilber-Rosenberg et al. proposed the concept of hologenome
2009	弗吉尼亚·阿姆布鲁斯特团队提出硅藻藻-菌关系 Virginia Armbrust’s team proposed the interactions between diatoms and algae
2009	杨桥团队开展海洋甲藻的可培养藻际微生物的分离及其生物活性功能研究 Yang Qiao’s team isolated the cultivable phycosphere bacteria from toxic marine dinoflagellates and studied their bioactive functions
快速发展期 (21世纪10年代至今) Rapid development period (from 2010s to the present)
2012	弗吉尼亚·阿姆布鲁斯特综述硅藻藻-菌关系并展望对于调节海洋生态、预测及调节气候的重要意义 Virginia Armbrust reviewed the interactions between diatoms and algae and looked forward to its importance in regulating marine ecology and predicting and regulating global climate
2014	魏江春院士团队完成首个石果衣真菌(Endocarpon pusillum Hedwig)的全基因组测序 Wei Jiangchun’s team completed the first whole genome sequencing of Endocarpon pusillum Hedwig
2014	杨桥等利用高通量测序技术揭示海洋典型赤潮甲藻的藻际微生物的多样性信息 Yang Qiao’s team revealed the biodiversity of phycosphere microbiota of some typical HAB-causing marine dinoflagellates using high-throughput sequencing technology
2015	弗吉尼亚·阿姆布鲁斯特团队阐明吲哚乙酸的藻菌互作信号新机制 Virginia Armbrust’s team elucidated new signaling mechanism of algae-bacteria interaction by indole-3-acetic acid (IAA)
2015	林森杰团队破译了重要珊瑚共生甲藻虫黄藻(Symbiodinium kawagutii)基因组 Lin Senjie’s team deciphered the genome of Symbiodinium kawagutii which is an important coral symbiotic dinoflagellate
2016	里希拉姆·拉曼等发表藻-菌关系重要综述，展望了其在推动可持续发展的巨大潜力 Rishiram Raman et al. published the review of algae-bacteria interactions, and looked forward to the great potential in promoting sustainable development
2017	贾斯汀·西摩发表重要综述，为揭示水生系统浮游植物与细菌互作机制及意义提供新视角 Justin Seymour published an important review, providing new perspectives on the mechanisms and significance of interactions between phytoplankton and bacteria in aquatic systems
2019	埃米利奥·西里及乔治·波纳特发表综述，总结展望藻菌互作对全球元素循环的重要影响 Emilio Ciri and Giorgio Bonnet published a review and summarized and looked forward to the important impact of algae-bacteria interactions on global element cycles
2019	中山卓郎等发现海洋甲藻中普遍存在共生蓝细菌，为重新认识海洋蓝细菌生态功能提供重要线索 Takuro Nakayama et al. found that symbiotic cyanobacteria were common in marine dinoflagellates, providing important clues for re-understanding the ecological functions of marine cyanobacteria.
2019	穆罕默德·多尼亚等发现藻-菌-动物复杂共生体化学防御机制，表明不同生物体共用化学防御体系 Mohammed Doniya et al. discovered the chemical defense mechanism of algae-bacteria-animal complex symbiosis, indicating that different organisms share a common chemical defense system
2021	法里巴·雷兹瓦尼等综述藻菌互作对废水营养物去除研究进展，展望了藻菌互作增效机制巨大应用潜力 Fariba Rezvani et al. reviewed the research progress on nutrient removal from wastewater by algae-bacteria interactions and looked forward to the huge application potential of the synergistic mechanism of algae-bacteria interactions
2022	《麻痹性贝类毒素生物学研究与应用》学术著作撰写出版 Yang Qiao’s team published the academic book Biological Research and Application of Paralytic Shellfish Toxins
2023	玛利亚·米塔等揭示微藻与互利菌及拮抗菌之间的互作机制，为研究藻菌互作机制及其应用奠定基础 Maria Mitta et al. revealed the interaction mechanism between microalgae and mutually beneficial and antagonistic bacteria, laying a foundation for the study of algae-bacteria interactions mechanism and application
2023	杨桥团队开始推动藻际微生物学新学科的筹建 Yang Qiao’s team began to promote to establish phycosphere microbiology as a new emerging interdiscipline
2024	《藻际微生物学导论》《藻际微生物学：百年百人百事》等藻际微生物学系列丛书陆续出版 Yang Qiao’s team published series of books on Phycosphere Microbiology, including Introduction to Phycosphere Microbiology and Phycosphere Microbiology with Hundreds of Years, Peoples and Events
2024	泰勒·科尔等发现海洋藻固氮菌内共生体进化为硝质体，将原核细胞的功能认识扩展至真核生物 Taylor Cole et al. discovered that the endosymbionts of nitrogen-fixing bacteria of marine microalgae evolved into the nitraplasts, extending the understanding of the functions of prokaryotic cells to eukaryotic organisms

时间Year

重要事件

Key events

启蒙孕育期(几千年前至20世纪中叶) Enlightenment period (from thousands of years ago to the middle 20th century)

1703

人类首次对淡水硅藻——平板藻(Tabellaria)进行细胞形态学观察和记录

The first observation and record of cell morphology of fresh-water diatom Tabellaria

1753

卡尔·林奈在《植物种志》(Speciesl Panatrum)中，将地衣归类到Lihcen属

Carl Linnaeus classified lichens in the genus Lihcen in the book titled Speciesl Panatrum

1798

埃里克·阿卡里乌斯开创地衣分类学，出版多部地衣学著作，推动其成为一门独立学科

Erik Acharius created lichen taxonomy and published several books on lichenology, and promoted it to become an independent discipline

1867

西蒙·施文德纳发现地衣是由真菌与藻共生的复合生物体，揭示了地衣的共生本质

Simon Schwendener discovered that lichens were complex organisms which were composed of fungi and algae, thus revealed the dual nature of the lichens

1872

约翰内斯·莱因克提出“consortium”一词，用来表示构成地衣的藻与真菌的关系

Johannes Reinke coined the term “consortium” to describe the relationship between algae and fungi that make up the lichens

1890

马丁努斯·贝耶林克分离获得藻际细菌

Martinus Beijerinck was the first people who isolated phycosphere bacteria

1915

查尔斯·沃尔科特从前古生代石灰岩藻沉积物化石中发现藻际细菌存在证据

Charles Walcott found the evidence for the existence of phycosphere bacteria in fossilized limestone from the prepaleozoic era

1940s

恩斯特·普林斯海姆分离保存大量藻株并进行分享；为世界主要藻株保藏中心作出了重要贡献

Ernst Prinsheim isolated, preserved, and shared a large number of algae strains, and made significant contributions to major algae collections in the world

创建成长期(20世纪中叶至21世纪10年代) Establishment and growth period (from middle 20th century to 2010s)

1943

迈耶-阿比奇提出全生物体(holobiont)概念及全共生理论(Theory of Holobiosis)

Meyer-Abich coined the concept of holobiont and proposed the Theory of Holobiosis

1952

威廉·奥斯特瓦尔德等利用藻-菌共生体系开发生活污水的生物处理技术

Wilhelm Ostwald et al. developed a biological treatment technology for domestic sewage based on the bacteria-algae symbiotic system

1953

哈罗德·米尔纳发明透明容器推动规模化培养微藻细胞，为光生物反应器实际应用奠定重要基础

Harold Milner invented the transparent container to promote the large-scale cultivation of microalgae cells, and laid important foundation for the practical application of photobioreactors

1957

乔治·斯科特提出藻际生命共同体(phycobiont)概念

George Scott coined the concept of phycobiont

1963

萨弗曼发现蓝藻病毒及噬藻体(1969)

Saffman discovered cyanobacterial viruses in 1963, and then cyanophages in 1969

1969

詹姆斯·斯图尔特及马尔科姆·布朗首次发现溶藻菌

James Stewart and Malcolm Brown first discovered the algicidal bacteria

1970

林恩·马古利斯提出内共生学说(Endosymbiosis Theory)

Lynn Margulis proposed the Endosymbiosis Theory

1972

韦恩·贝尔及拉尔夫·米切尔提出藻际(phycosphere)概念

Wayne Bell and Ralph Mitchell coined the concept of phycosphere

1973

魏江春院士领衔编研《中国地衣志》一书

Wei Jiangchun began to compiled the book titled Chinese Lichens

1978

埃斯特拉·席尔瓦首次发现甲藻胞内细菌；提出著名的“甲藻共生菌产毒”假说

Estela Silva discovered intracellular bacteria in dinoflagellates and proposed the famous hypothesis of “toxic production by symbiotic bacteria of dinoflagellate”

1982

乔纳森·科尔提出菌藻相互作用(interactions between bacteria and algae)

Jonathan Cole coined the concept of interactions between bacteria and algae

1988

儿玉正明发现来源于产毒海洋甲藻的藻际自产毒细菌莫拉氏菌PTB-1

Masaaki Kodama discovered the paralytic shellfish poisoning (PSP) toxin-producing bacterium, Moraxella sp. PTB-1 from a PSP toxin-producing marine dinoflagellate

1989

约翰·拉齐尔发现影响藻际生态位大小的因素

John Lazear studied and discovered the factors that influence the size of the phycosphere niche

1994

郑天凌等提出“以菌治藻”新思路

Zheng Tianling et al. proposed a new idea of “bacterial control the harmful algal blooms (HABs)”

1995

格雷戈里·杜塞特发现溶藻菌及藻-菌相互作用

Gregory Doucette discovered algicidal bacteria and algae-bacteria interactions

1999

伯纳德·雅克等提出互作组(interactome)概念

Bernard Jacques et al. coined the concept of interactome

2000

玛丽·莫兰等研究藻际玫瑰杆菌(Roseobacter)类细菌；发现微藻对藻际菌群结构起决定性作用

Mary Moran’s team studied the bacteria of the Roseobacter group in the the phycosphere niche, and found that microalgae play a decisive role in the community structure of the phycosphere microbiota

2003

魏江春院士团队提出“沙漠生物地毯工程”

Wei Jiangchun’s team proposed the “Desert Biocarpet Project”

2008

伊拉娜·齐尔伯-罗森伯格等提出全基因组(hologenome)概念

Ilana Zilber-Rosenberg et al. proposed the concept of hologenome

2009

弗吉尼亚·阿姆布鲁斯特团队提出硅藻藻-菌关系

Virginia Armbrust’s team proposed the interactions between diatoms and algae

2009

杨桥团队开展海洋甲藻的可培养藻际微生物的分离及其生物活性功能研究

Yang Qiao’s team isolated the cultivable phycosphere bacteria from toxic marine dinoflagellates and studied their bioactive functions

快速发展期 (21世纪10年代至今) Rapid development period (from 2010s to the present)

2012

弗吉尼亚·阿姆布鲁斯特综述硅藻藻-菌关系并展望对于调节海洋生态、预测及调节气候的重要意义

Virginia Armbrust reviewed the interactions between diatoms and algae and looked forward to its importance in regulating marine ecology and predicting and regulating global climate

2014

魏江春院士团队完成首个石果衣真菌(Endocarpon pusillum Hedwig)的全基因组测序

Wei Jiangchun’s team completed the first whole genome sequencing of Endocarpon pusillum Hedwig

2014

杨桥等利用高通量测序技术揭示海洋典型赤潮甲藻的藻际微生物的多样性信息

Yang Qiao’s team revealed the biodiversity of phycosphere microbiota of some typical HAB-causing marine dinoflagellates using high-throughput sequencing technology

2015

弗吉尼亚·阿姆布鲁斯特团队阐明吲哚乙酸的藻菌互作信号新机制

Virginia Armbrust’s team elucidated new signaling mechanism of algae-bacteria interaction by indole-3-acetic acid (IAA)

2015

林森杰团队破译了重要珊瑚共生甲藻虫黄藻(Symbiodinium kawagutii)基因组

Lin Senjie’s team deciphered the genome of Symbiodinium kawagutii which is an important coral symbiotic dinoflagellate

2016

里希拉姆·拉曼等发表藻-菌关系重要综述，展望了其在推动可持续发展的巨大潜力

Rishiram Raman et al. published the review of algae-bacteria interactions, and looked forward to the great potential in promoting sustainable development

2017

贾斯汀·西摩发表重要综述，为揭示水生系统浮游植物与细菌互作机制及意义提供新视角

Justin Seymour published an important review, providing new perspectives on the mechanisms and significance of interactions between phytoplankton and bacteria in aquatic systems

2019

埃米利奥·西里及乔治·波纳特发表综述，总结展望藻菌互作对全球元素循环的重要影响

Emilio Ciri and Giorgio Bonnet published a review and summarized and looked forward to the important impact of algae-bacteria interactions on global element cycles

2019

中山卓郎等发现海洋甲藻中普遍存在共生蓝细菌，为重新认识海洋蓝细菌生态功能提供重要线索

Takuro Nakayama et al. found that symbiotic cyanobacteria were common in marine dinoflagellates, providing important clues for re-understanding the ecological functions of marine cyanobacteria.

2019

穆罕默德·多尼亚等发现藻-菌-动物复杂共生体化学防御机制，表明不同生物体共用化学防御体系

Mohammed Doniya et al. discovered the chemical defense mechanism of algae-bacteria-animal complex symbiosis, indicating that different organisms share a common chemical defense system

2021

法里巴·雷兹瓦尼等综述藻菌互作对废水营养物去除研究进展，展望了藻菌互作增效机制巨大应用潜力

Fariba Rezvani et al. reviewed the research progress on nutrient removal from wastewater by algae-bacteria interactions and looked forward to the huge application potential of the synergistic mechanism of algae-bacteria interactions

2022

《麻痹性贝类毒素生物学研究与应用》学术著作撰写出版

Yang Qiao’s team published the academic book Biological Research and Application of Paralytic Shellfish Toxins

2023

玛利亚·米塔等揭示微藻与互利菌及拮抗菌之间的互作机制，为研究藻菌互作机制及其应用奠定基础

Maria Mitta et al. revealed the interaction mechanism between microalgae and mutually beneficial and antagonistic bacteria, laying a foundation for the study of algae-bacteria interactions mechanism and application

2023

杨桥团队开始推动藻际微生物学新学科的筹建

Yang Qiao’s team began to promote to establish phycosphere microbiology as a new emerging interdiscipline

2024

《藻际微生物学导论》《藻际微生物学：百年百人百事》等藻际微生物学系列丛书陆续出版

Yang Qiao’s team published series of books on Phycosphere Microbiology, including Introduction to Phycosphere Microbiology and Phycosphere Microbiology with Hundreds of Years, Peoples and Events

2024

泰勒·科尔等发现海洋藻固氮菌内共生体进化为硝质体，将原核细胞的功能认识扩展至真核生物

Taylor Cole et al. discovered that the endosymbionts of nitrogen-fixing bacteria of marine microalgae evolved into the nitraplasts, extending the understanding of the functions of prokaryotic cells to eukaryotic organisms