中国农业科学 ›› 2021, Vol. 54 ›› Issue (11): 2287-2301.doi: 10.3864/j.issn.0578-1752.2021.11.004

• 耕作栽培·生理生化·农业信息技术 • 上一篇    下一篇

非生物逆境锻炼提高作物耐逆性的生理机制研究进展

王笑(),蔡剑,周琴,戴廷波,姜东()   

  1. 南京农业大学小麦区域创新中心/农业农村部作物生理生态与生产管理重点实验室,南京 210095
  • 收稿日期:2020-08-10 接受日期:2020-10-12 出版日期:2021-06-01 发布日期:2021-06-09
  • 联系方式: 王笑,E-mail:xiaowang@njau.edu.cn。
  • 基金资助:
    重点研发计划项目(2016YFD0300107);国家自然科学基金(31771693);国家自然科学基金(U1803235);国家现代小麦产业技术体系(CARS-03);江苏省协同创新中心(JCIC-MCP)

Physiological Mechanisms of Abiotic Stress Priming Induced the Crops Stress Tolerance: A Review

WANG Xiao(),CAI Jian,ZHOU Qin,DAI TingBo,JIANG Dong()   

  1. National Technique Innovation Center for Regional Wheat Production, Nanjing Agricultural University/Key Laboratory of Crop Physiology Ecology and Production Management, Ministry of Agriculture and Rural Affairs, Nanjing 210095
  • Received:2020-08-10 Accepted:2020-10-12 Published:2021-06-01 Online:2021-06-09

摘要:

非生物逆境(如,高温、低温、干旱、渍水胁迫等)是限制作物产量提升的重要因子,并且非生物逆境发生的频率、程度以及持续时间随着全球气候变化呈显著上升趋势。因此,提高作物对非生物逆境的抗性,或采取缓解措施降低非生物逆境对作物产量和品质形成的不利影响,对于确保作物稳产及粮食安全有重要意义。逆境锻炼(priming)是指植株经过前期适度的逆境处理后,对再次发生的逆境胁迫表现出较强的抗/耐性,也称为逆境胁迫记忆。与未经过锻炼植株相比,经过锻炼植株的信号调控物质、次级代谢产物、胁迫保护性物质等可以更快、更有效地对再次发生的逆境胁迫产生响应,从而增强植株耐逆性。根据再次逆境发生的类型及时间,逆境锻炼主要包括当代同种逆境锻炼效应(锻炼阶段的逆境和再次发生的逆境是同一种)、当代交叉逆境锻炼效应(锻炼阶段的逆境和再次发生的逆境不是同一种)、跨代同种逆境锻炼效应(经过逆境锻炼的种子在子一代或子几代的同种逆境锻炼效应)、跨代交叉逆境锻炼效应(经过逆境锻炼的种子在子一代或子几代的交叉逆境锻炼效应)四大类型。本文重点围绕高温锻炼、低温锻炼、干旱锻炼及渍水锻炼介导的上述四大类型锻炼效应的生理机制进行了综述,生理机制主要包括植株光合机构响应机制、抗氧化系统在清除活性氧减轻对细胞膜脂过氧化伤害机制、胁迫诱导的信号物质(激素类物质、Ca2+、过氧化氢、一氧化氮等)在诱导下游基因表达及生理生化过程机制。此外,表观遗传修饰如DNA甲基化、组蛋白修饰为长期甚至传代胁迫记忆提供了潜在机制。对作物逆境锻炼机制的深入解析,可以找到对作物耐逆性获得起关键调控作用的基因和蛋白,这样在作物生产上,我们可以在生育前期,配合外源调控物质诱导起关键作用的基因和蛋白,可通过人为方法提前刺激这种物质在逆境来临之前表达,主动诱导作物对关键时期逆境耐性的形成,从而有效缓解在产量形成关键生育时期发生的胁迫对作物产量的不利影响,因而具有重要的实际生产意义。

关键词: 非生物逆境, 逆境锻炼, 当代锻炼, 跨代锻炼, 生理机制, 信号调控机制

Abstract:

Abiotic stress factors, including heat stress, cold stress, drought stress, waterlogging stress etc., are the limiting factors for plant growth and crop production. Moreover, the frequency, extent and duration of abiotic stresses have predicted to be increased with global climate change. Therefore, improving crops resistance to abiotic stress or finding strategy to reduce the adverse effects of abiotic stress on crop yield and quality is of great significance for ensuring stable crop production and food security. “Priming” by exposing plants to moderate stress in the early growth stage can induce plant resistance to a later severe stress episode which happened during critical stage of the plant growth. The priming is an adaptive strategy that primed plants could effectively mount a faster and/or stronger defense response and actively improve the defensive capacity of plants under stress, compared with non-primed plants. According to the reoccurred stress types, the priming can be generally separated to four types, including in-generational stress tolerance (the priming stimulus and the stress occurred at later stage are the same stress type), intra-generational cross tolerance (the priming stimulus and the stress occurred at later stage are different stress types), transgenerational stress tolerance (the priming stimulus and the stress occurred at next generations are the same stress type), and transgenerational cross tolerance (the priming stimulus and the stress occurred at next generations are different stress types). In this review, the main physiological mechanisms were discussed, including plant photosynthetic response mechanism, antioxidant mechanism, and osmotic regulation, signal transduction mechanisms (plant hormones, Ca2+, hydrogen peroxide, nitric oxide, etc.), and epigenetic modification mechanisms (DNA methylation, histone modification, etc.). Then, the perspectives for the further research on understanding the underlying mechanisms of stress priming and the application of priming effects in the crop production were suggested. After revealing the mechanisms of priming, the candidate genes and proteins which play key regulatory roles in the acquisition of crop stress tolerance are found. And then, we can stimulate the related genes and protein expression which can actively induce the formation of stress tolerance in the critical crops growth stage, thereby effectively alleviating the adverse effects of abiotic stress on crop yield during the critical stage, which is meaningful for the crop production.

Key words: abiotic stress, priming, intra-generational priming, transgenerational priming, physiological mechanisms, signal transduction mechanisms