大豆响应高温胁迫的生理和分子遗传机理研究现状与展望

doi:10.3864/j.issn.0578-1752.2017.14.004

摘要/Abstract

摘要： 大豆是重要的经济作物，是植物油脂和蛋白质的重要来源。近年来，因全球气候变化引起的高温胁迫频发，危及到大豆生长的各个时期，成为制约大豆产量和品质的重要因素之一。为了揭示大豆耐高温性的遗传机理，建立综合高效大豆耐高温评价体系，促进大豆耐高温特性的遗传改良，现对大豆响应高温胁迫的生理生化基础和分子调控机制进行综述。相较于适温条件，高温胁迫可使大豆植株发生叶片增厚、气孔导度下降、细胞膜透性增加、细胞微观组织结构受损以及渗透调节物质（脯氨酸、可溶性糖和可溶性蛋白）含量变化和抗氧化防御系统关键酶活性丧失等生理异常反应，导致植株光合、蒸腾和呼吸作用及物质含量等一系列生理生化过程紊乱。高温胁迫还造成大豆花粉形态异常，绒毡层细胞结构松散、空泡化和自溶化，花粉活力及其萌发率明显下降，花粉败育率增高，致使大豆结荚率和结实率显著降低，进而影响大豆籽粒的正常发育、蛋白的积累和产量形成等，最终导致减产；高温胁迫对大豆籽粒外观品质性状也能造成一定损伤，进而对其经济价值带来不利影响。高温胁迫从转录、翻译和代谢水平影响大豆正常的生理代谢调控。目前，已通过高通量测序等方法鉴定出多个与大豆高温胁迫响应相关的转录因子、蛋白及代谢产物，但与水稻、拟南芥的研究相比仍存在较大差距；初步建立了以花粉活力和多项生理指标为基础的大豆耐高温鉴定方法，但在生产上尚缺乏系统高效耐高温评价体系，耐高温大豆育种工作进展缓慢。国内在大豆耐高温预防措施、耐高温综合评鉴体系建立、优异耐高温大豆种质资源筛选及耐高温关键功能基因挖掘等方面仍然存在挑战。为应对极端生态环境给大豆生产造成的不利影响，未来应通过建立综合高效大豆耐高温评价技术体系，提高大豆优异种质资源鉴定水平，重点解析大豆耐高温的分子遗传机理、挖掘耐高温关键基因，结合常规育种和分子育种技术培育综合性状优异的耐高温大豆新品种，以实现大豆品种耐高温性与高产、优质的统一。

关键词: 大豆, 高温胁迫, 耐热性, 生理生化特性, 分子机理

Abstract: Soybean is a key economic crop, and is an important source of plant protein and oil. In recent years, high temperature (HT) stress caused by global climate change has threatened the key growth period of soybean, and has become one of the main environmental factors which limited the yield and quality of soybean. In order to reveal the molecular mechanism, establish a comprehensive efficiency evaluation system and promoting genetic improvement of soybean resistance to HT stress, the physiological and biochemical basis and molecular regulation mechanism responding to HT stress of soybean were reviewed in this paper. Compared to optimum temperature, the thickness of leaves increased, the stomatal conductance decreased, the membrane permeability increased, cell microstructure and its integrity damaged, the osmoregulation substances (proline, soluble sugar and protein) changed and the key enzyme activities of the antioxidant defense system in soybean have been lost under HT stress, these results will lead to disorders of a series of physiological processes in soybean, which include the photosynthesis, transpiration, respiration and substance content, etc. Besides, HT stress also caused the abnormalities of pollen morphology, the tapetum cytoplasm is not dense, showing vacuolated and autolysis, the pollen vigour and germination rate are significantly decreased, and pollen abortion rate increased, thus resulting in the pod-set rate and seed-set percentage remarkable decrease, and affecting the normal development and protein accumulation of seed and yield formation of soybean, and ultimately leading to lower yields of soybean. Meanwhile, HT stress can also cause some damage to quality traits of seed, and then has an adverse effect on its economic value of soybean. Moreover, normal physiological metabolism is widely affected by HT stress at the level of transcription, translation and metabolism. At present, many transcription factors, proteins and metabolites response to HT stress in soybean have been identified by the method of high-throughput sequencing, however, there is still a wide gap compared with research on rice and Arabidopsis. Currently, the comprehensive stress response indexes based on pollen viability and several physiological parameters have been used to screen HT resistance soybean varieties, but, the method of system identification is still imperfect in soybean production, and thus leading to the slow progress in selection of HT resistant soybean. Up to date, challenges are still existed in preventive measures, the establishment of a comprehensive evaluation system, screening for excellent soybean germplasm related to resistance to HT, and identification of key functional genes resistance to HT stress in soybean. The adverse effects of extreme ecological environment on soybean production will be further aggravated with the uncertainty of global climate change. In the future, the main focuses should be put on the establishment of a comprehensive evaluation technology system, improvement of the identification level of soybean germplasm resources, elucidation of molecular genetic mechanism and excavation of key genes resistance to HT in soybean. Selecting new soybean varieties with resistance to HT stress by both of the conventional breeding and molecular breeding technologies, thus will provide a theoretical basis and technical support for achieving the unification of HT resistance as well as high yield and quality in soybean.

Key words: soybean, high temperature stress, heat tolerance, physiological and biochemical characteristics, molecular mechanism

李佳佳，郑双雨，孙根楼，张文明，王晓波，邱丽娟. 大豆响应高温胁迫的生理和分子遗传机理研究现状与展望[J]. 中国农业科学, 2017, 50(14): 2670-2682.

LI JiaJia, ZHENG ShuangYu, SUN GenLou, ZHANG WenMing, WANG XiaoBo, QIU LiJuan. Advances and Perspectives in Research of Physiological and Molecular Mechanism of Soybean Response to High Temperature Stress[J]. Scientia Agricultura Sinica, 2017, 50(14): 2670-2682.

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