种子际微生物研究展望

doi:10.11983/CBB22001

摘要/Abstract

摘要： 种子际是围绕在种子表面, 受到种子萌发影响而使微生物活性增强的狭窄土壤区域。萌发期间的种子和微生物相互影响, 对种子萌发和随后的幼苗生长具有促进或抑制作用; 同时也通过垂直传播参与植物根际和叶际微生物群落构建, 影响植物后续发育及生物/非生物胁迫响应。尽管种子际对于植物生长发育影响很大, 但由于种子萌发持续时间短、种子际的空间范围小且微生物量较小, 种子际研究要比根际和叶际更为复杂。通过基因组学和多组学联合分析等方法对种子际微生物群落组成进行深入探究, 有助于理解植物与土壤生态系统的相互作用和生态过程。该文综述了种子际微生物对种子生物学的影响、种子际微生物与根际和叶际微生物的关系以及相关研究方法, 并展望了未来的研究方向。

关键词: 种子际微生物, 萌发, 生物胁迫, 非生物胁迫, 多组学联合分析

Abstract: The spermosphere is a narrow area of soil surrounding the seed surface where microbial activity is enhanced by seed germination processes. Interaction between seeds and microorganisms promotes or inhibits seed germination and subsequent seedling establishment, and also participates in plant rhizosphere and phyllosphere microbial community building through vertical dispersal, and finally influences subsequent plant development and biotic/abiotic stress response. Although the spermosphere has a great influence on plant growth and development, it is more complex to study than rhizosphere and phyllosphere due to the short duration of seed germination, the small spatial extent of the spermosphere, and the small amount of microorganisms involved. To explore the composition of spermosphere microbial community by genomic analysis and multi-omics joint analysis, will facilitate understanding the interactions and ecological processes between plants and soil ecosystems. In this paper, we review the influence of spermosphere microorganisms on the biological processes of seed germination, the comparative meta-genomics of spermosphere with rhizosphere and phyllosphere, and provide an outlook on future research.

Key words: spermosphere microorganism, germination, biotic stress, abiotic stress, combined multi-omics analysis

任晓童, 张冉冉, 魏绍巍, 罗晓峰, 徐佳慧, 舒凯. 种子际微生物研究展望. 植物学报, 2023, 58(3): 499-509.

Xiaotong Ren, Ranran Zhang, Shaowei Wei, Xiaofeng Luo, Jiahui Xu, Kai Shu. Research Progress of Spermosphere Microorganisms. Chinese Bulletin of Botany, 2023, 58(3): 499-509.

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链接本文: https://www.chinbullbotany.com/CN/10.11983/CBB22001

https://www.chinbullbotany.com/CN/Y2023/V58/I3/499

图/表 2

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寄主植物	种子际微生物	功能与作用机制	参考文献
		种子萌发相关
苜蓿	泛菌属(Pantoea)和芽孢杆菌属(Bacillus)等	盐胁迫下通过促进生长素合成、抵抗真菌以及提高渗透胁迫耐受性促进种子萌发	Dai et al., 2020
高羊茅、鸭茅和新麦草	巴西固氮螺菌(Azospirillum brasilense)	通过降低丙二醛含量和提高抗氧化酶活性修复种子老化损伤, 提高老化种子萌发率	柳旭, 2018
番茄	米甲基杆菌(Methylobacterium oryzae (CBMB20))	通过促进植物激素合成、氮固定、磷酸盐增溶和铁载体产生等途径促进种子萌发和植株生长	Chanratana et al., 2018
拟南芥	铜绿假单胞菌(Pseudomonas aeruginosa)	通过释放可触发萌发抑制的生物化合物AMB (L-2- amino-4-methoxy-trans-3-butenoic acid)抑制种子萌发, 增强胁迫耐受性	Chahtane et al., 2018
小麦和禾本科牧草	头孢霉菌属(Acremonium)、胶枝菌属(Gliocladium)和瓶霉菌属(Phialophora)	通过与镰刀菌(Fusarium culmorum)拮抗性竞争种子际生态位抑制镰刀菌生长, 降低镰刀菌枯萎病发病率	Slykhuis, 1947
小麦	假单胞菌属(Pseudomonas)	在种子萌发和幼苗早期生长阶段分解过氧化氢, 调节植物细胞外氧化还原环境	Gerna et al., 2020
番茄	哈茨曼氏菌(Trichoderma harzianum)	减少受胁迫植物中活性氧的积累, 保护植物免受氧化损伤, 增强幼苗活力	Mastouri et al., 2010
复活蕨	芽孢杆菌属(Bacillus)等	通过促进植物激素合成、氮固定、次生代谢物分泌和诱导植物系统抗性促进幼苗生长和植物发育	Jackson et al., 2006
玉米	贝莱斯芽孢杆菌(B. velezensis)	帮助植物抵抗轮枝镰刀菌(F. verticillioides), 促进玉米种子萌发和幼苗生长	Pal et al., 2022
花生	芽孢杆菌属(Bacillus)	协助植物产生特定磷酸酶, 将有机磷转化为无机磷, 提高花生种子萌发期的耐盐能力	Xu et al., 2020
豌豆和四季豆	芽孢杆菌属(Bacillus)	分泌抗真菌化合物, 帮助植物抵抗灰葡萄孢菌(Botrytis cinerea)和腐霉属(Pythium)真菌的侵染	Walker et al., 1998

寄主植物	种子际微生物	功能与作用机制	参考文献
		种子萌发相关
苜蓿	泛菌属(Pantoea)和芽孢杆菌属(Bacillus)等	盐胁迫下通过促进生长素合成、抵抗真菌以及提高渗透胁迫耐受性促进种子萌发	Dai et al., 2020
高羊茅、鸭茅和新麦草	巴西固氮螺菌(Azospirillum brasilense)	通过降低丙二醛含量和提高抗氧化酶活性修复种子老化损伤, 提高老化种子萌发率	柳旭, 2018
番茄	米甲基杆菌(Methylobacterium oryzae (CBMB20))	通过促进植物激素合成、氮固定、磷酸盐增溶和铁载体产生等途径促进种子萌发和植株生长	Chanratana et al., 2018
拟南芥	铜绿假单胞菌(Pseudomonas aeruginosa)	通过释放可触发萌发抑制的生物化合物AMB (L-2- amino-4-methoxy-trans-3-butenoic acid)抑制种子萌发, 增强胁迫耐受性	Chahtane et al., 2018
小麦和禾本科牧草	头孢霉菌属(Acremonium)、胶枝菌属(Gliocladium)和瓶霉菌属(Phialophora)	通过与镰刀菌(Fusarium culmorum)拮抗性竞争种子际生态位抑制镰刀菌生长, 降低镰刀菌枯萎病发病率	Slykhuis, 1947
小麦	假单胞菌属(Pseudomonas)	在种子萌发和幼苗早期生长阶段分解过氧化氢, 调节植物细胞外氧化还原环境	Gerna et al., 2020
番茄	哈茨曼氏菌(Trichoderma harzianum)	减少受胁迫植物中活性氧的积累, 保护植物免受氧化损伤, 增强幼苗活力	Mastouri et al., 2010
复活蕨	芽孢杆菌属(Bacillus)等	通过促进植物激素合成、氮固定、次生代谢物分泌和诱导植物系统抗性促进幼苗生长和植物发育	Jackson et al., 2006
玉米	贝莱斯芽孢杆菌(B. velezensis)	帮助植物抵抗轮枝镰刀菌(F. verticillioides), 促进玉米种子萌发和幼苗生长	Pal et al., 2022
花生	芽孢杆菌属(Bacillus)	协助植物产生特定磷酸酶, 将有机磷转化为无机磷, 提高花生种子萌发期的耐盐能力	Xu et al., 2020
豌豆和四季豆	芽孢杆菌属(Bacillus)	分泌抗真菌化合物, 帮助植物抵抗灰葡萄孢菌(Botrytis cinerea)和腐霉属(Pythium)真菌的侵染	Walker et al., 1998

方法	原理	特点	局限性	参考文献
分子生物学分析技术
G+C含量测定	不同物种所携带的鸟嘌呤(G)+胞嘧啶(C)含量不同, 通过研究DNA中G+C含量的差异, 可以评估微生物群落的多样性	原位检测; 具有特异性, 不受菌龄和外界环境的影响; 简便、快速、灵敏且成本低	单独推断物种存在或物种丰度不准确, 应与表型特征或其它分类依据结合起来分析	Griffiths et al., 1997
磷脂脂肪酸分析(PLFA)	磷脂是活性微生物生物量的重要指标; 特定的脂肪酸表示特定的生物种群	易提取	不能精确到物种水平; 严重依赖特征脂肪酸	Haack et al., 1994
变性/温度梯度凝胶电泳(DGGE/TGGE)	利用DNA分子在具有DNA变性梯度或温度梯度的聚丙烯酰胺凝胶中迁移率的差异分离DNA	可靠、重复性好、快速且成本低	只能检测出环境样本中1%-2%的代表优势种的微生物种群	Macnaughton et al., 1996; Mühling et al., 2008
末端限制性片段长度多态性(T-RFLP)	用限制性内切酶切割带有荧光标记引物的PCR扩增产物, 由于不同微生物物种存在核苷酸序列差异, 会产生不同长度的末端限制性片段	结果可靠; 可用于比较不同样本之间的关系和检测微生物群落的时空变化	只能检测到一定数量的优势物种; 受限于通用引物的选择; 受限制性内切酶种类的影响	Marsh, 1999; Rudi et al., 2007
稳定同位素探测(SIP)	追踪特定底物中重稳定同位素与微生物同化底物相关的系统发育信息生物标记物(PLFA、DNA、RNA和蛋白质)的结合	将功能与微生物群落结构联系起来, 可用于研究植物与微生物的相互作用	可用性有限、成本高、低通量	Uhlik et al., 2013
荧光原位杂交(FISH)	通过荧光标记的寡核苷酸探针与特异的互补核酸序列杂交, 利用荧光显微镜直接观察特定微生物的分布与数量	原位无损检测, 敏感度高	环境样品自发荧光; 寡核苷酸探针特异性差	Ahmad et al., 2011; Marschner et al., 2011
高通量分析技术
多组学联合分析	通过基因组学、转录组学、蛋白质组学、代谢组学和具有原位环境特征的微生物组学研究各个层次的信息	将功能、活性和相互作用联系起来; 高通量	常用的二代测序错误率高; 遗传数据库不完备	Abram, 2015

方法	原理	特点	局限性	参考文献
分子生物学分析技术
G+C含量测定	不同物种所携带的鸟嘌呤(G)+胞嘧啶(C)含量不同, 通过研究DNA中G+C含量的差异, 可以评估微生物群落的多样性	原位检测; 具有特异性, 不受菌龄和外界环境的影响; 简便、快速、灵敏且成本低	单独推断物种存在或物种丰度不准确, 应与表型特征或其它分类依据结合起来分析	Griffiths et al., 1997
磷脂脂肪酸分析(PLFA)	磷脂是活性微生物生物量的重要指标; 特定的脂肪酸表示特定的生物种群	易提取	不能精确到物种水平; 严重依赖特征脂肪酸	Haack et al., 1994
变性/温度梯度凝胶电泳(DGGE/TGGE)	利用DNA分子在具有DNA变性梯度或温度梯度的聚丙烯酰胺凝胶中迁移率的差异分离DNA	可靠、重复性好、快速且成本低	只能检测出环境样本中1%-2%的代表优势种的微生物种群	Macnaughton et al., 1996; Mühling et al., 2008
末端限制性片段长度多态性(T-RFLP)	用限制性内切酶切割带有荧光标记引物的PCR扩增产物, 由于不同微生物物种存在核苷酸序列差异, 会产生不同长度的末端限制性片段	结果可靠; 可用于比较不同样本之间的关系和检测微生物群落的时空变化	只能检测到一定数量的优势物种; 受限于通用引物的选择; 受限制性内切酶种类的影响	Marsh, 1999; Rudi et al., 2007
稳定同位素探测(SIP)	追踪特定底物中重稳定同位素与微生物同化底物相关的系统发育信息生物标记物(PLFA、DNA、RNA和蛋白质)的结合	将功能与微生物群落结构联系起来, 可用于研究植物与微生物的相互作用	可用性有限、成本高、低通量	Uhlik et al., 2013
荧光原位杂交(FISH)	通过荧光标记的寡核苷酸探针与特异的互补核酸序列杂交, 利用荧光显微镜直接观察特定微生物的分布与数量	原位无损检测, 敏感度高	环境样品自发荧光; 寡核苷酸探针特异性差	Ahmad et al., 2011; Marschner et al., 2011
高通量分析技术
多组学联合分析	通过基因组学、转录组学、蛋白质组学、代谢组学和具有原位环境特征的微生物组学研究各个层次的信息	将功能、活性和相互作用联系起来; 高通量	常用的二代测序错误率高; 遗传数据库不完备	Abram, 2015