乌日娜,
石凤翎^,,
徐舶
内蒙古农业大学草原与资源环境学院/草地资源教育部重点实验室(内蒙古农业大学) 呼和浩特 010000
基金项目: 内蒙古自治区科技计划项目2019GG244

详细信息

作者简介:乌日娜, 主要从事牧草遗传育种研究。E-mail:caokundeyouxiang@126.com

通讯作者:石凤翎, 主要从事牧草遗传育种研究。E-mail:sfl0000@126.com

中图分类号:Q945.78;Q944.53

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出版历程

收稿日期:2020-04-24
录用日期:2020-07-04
刊出日期:2020-12-01

Medicago ruthenica (L.) Sojak. cv. Zhilixing response and adaptation strategy to drought stress and rehydration

WU Rina,
SHI Fengling^,,
XU Bo
College of Grassland, Resources and Environment, Inner Mongolia Agricultural University/Key Laboratory of Grassland Resources of Ministry of Education(Inner Mongolia Agricultural University), Huhhot 010000, China
Funds: the Science and Technology Project of Inner Mongolia Autonomous Region in China2019GG244

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Corresponding author:SHI Fengling, E-mail:sfl0000@126.com

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摘要
摘要:以直立型扁蓿豆[Medicago ruthenica(L.)Sojak.cv.Zhilixing]为材料，于苗期连续干旱处理12 d后复水4 d，研究直立型扁蓿豆幼苗形态结构特征、生理代谢及生物量分配对干旱胁迫及复水的响应，揭示直立型扁蓿豆对干旱胁迫及复水的适应策略。结果表明：随着干旱胁迫时间延长，直立型扁蓿豆叶片气孔开放率逐渐降低，处理9 d后气孔及表皮细胞密度比正常浇水处理(CK)分别增加48.5%和36.6%，形成小而密的表皮细胞和气孔。生理上，除MDA含量随胁迫时间的延长逐渐增加外，其余指标均先升高后降低，干旱胁迫9 d时达最高，SOD、POD、叶绿素、可溶性糖、可溶性蛋白及脯氨酸分别较CK提高88.9%、111.2%、86.7%、140.5%、147.8%和124.6%。同时，生物量随胁迫时间的延长先增大后减小，于干旱胁迫9 d达最大值，比CK增加16.4%，总的分配格局表现出地上生物量投资高于地下，地下生物量投资比例随胁迫时间的延长逐渐增加，而地上生物量变化与其相反。复水后各指标均能恢复至CK水平或超过CK，表现出极强的复水敏感性和潜在恢复能力。该品种扁蓿豆对干旱胁迫及复水的适应主要分为3个时期：主动适应期，其生理参数的可塑性指数为形态参数的1.33倍，主要通过抗氧化及渗透调节来减少水分散失增加水分吸收、缓解氧化伤害以适应干旱逆境；被动适应期，其形态参数的可塑性指数为生理参数的1.31倍，主要采用牺牲生物量的生存策略以及降低色素含量减少光吸收的光保护机制来提高逆境下的生存能力；复水恢复期，根冠比、气孔开放率、气孔及表皮细胞密度比CK分别增加25.9%、29.7%、24.2%和16.3%，其较高的根冠比和叶片较高的气孔开放率及小而密的气孔及表皮细胞特征，保证了直立型扁蓿豆吸水能力以及水分运输效率的迅速恢复。综上，直立型扁蓿豆抗旱能力较强，能够通过形态生理的改变以及调整不同器官的生物量分配来应对与适应干旱逆境及复水，且在不同处理阶段采取不同的适应策略以达到生存目的。
关键词:直立型扁蓿豆/
干旱胁迫/
复水/
形态/
生理/
适应策略/
叶片表皮细胞/
气孔密度
Abstract:The effect of drought stress and rehydration on Medicago ruthenica (L.) Sojak cv. Zhilixing morphological structures, physiological metabolism, and matter distribution were investigated to determine the adaptation strategy in an arid environment. Seedling-stage M. ruthenica (with 6-8 leaves) were subjected to continuous drought stress for 12 days and then re-watered for 4 days; samples were collected from the control group (CK), after 9 and 12 days of drought stress, and 4 days after rehydration. Compared with the CK, the stoma opening rate decreased, while the stoma and epidermis cell densities increased by 48.5% and 36.6%, respectively. Smaller and denser epidermal cells formed. The malondialdehyde (MDA) content increased gradually during stress period. Superoxide dismutase activity (SOD), peroxidase activity (POD), and contents of chlorophyll, soluble sugar, soluble protein, and proline first increased and then decreased with prolonged drought. Total biomass also first increased then decreased, reaching the maximum after 9 days (0.433 7 g; 16.4% increase). The distribution of the above-ground biomass was higher than the under-ground biomass, and the under-ground biomass proportion increased with drought, while the opposite effect was observed with the above-ground biomass. After re-watering, all of the physiological and biochemical indexes recovered or exceeded those of the control, indicating strong rehydration sensitivity and resilience. M. ruthenica adaptation to drought stress and rehydration was divided into three periods:active adaptation, passive adaptation, and re-watering. The physiological parameter plasticity index was 0.16, 1.33 times greater than the morphological parameter index, during the active adaptation period. Drought stress adaptation was achieved by altering the antioxidant and osmotic regulation systems to reduce water loss, improving water retention and absorption efficiency, and maintaining the water absorption and loss balance. The morphological parameter plasticity index was 0.24, 1.31 times greater than the physiological parameter index, during the passive adaptation period. M. ruthenica sacrificed biomass and reduced pigments to aid survival. The root-shoot ratio (25.9%), stomatal opening rate (29.7%), and stomatal and epidermal cell density (24.2% and 16.3%, respectively) were higher than those in the CK during the re-watering period. These characteristics promoted rapid water recovery, absorption capacity, and water transport efficiency. Morphological and physiological changes allowed M. ruthenica to adapt to drought stress and rehydration, contributing to its survival in arid conditions.
Key words:Medicago ruthenica (L.) Sojak. cv. Zhilixing/
Drought stress/
Rehydration/
Morphology/
Physiology/
Adaptive strategy/
Epidermis cell of leaf/
Stomatal density

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图1不同处理条件下直立型扁蓿豆幼苗叶片表皮细胞及气孔
CK为正常水分处理; 9d和12d为干旱胁迫9 d和12 d, Rewater为复水后4 d。不同小写字母表示CK及各处理间差异显著(P < 0.05), *表示上下表皮间差异显著(P < 0.05)。
Figure1.Epidermis cell density (A), cell length (B), cell width (C), and stoma opening rate (D), density (E), length (F) and width (G) of leaves of Medicago ruthenica (L.) Sojak. cv. Zhilixing under drought stress and rehydration
CK is the control of normal water condition. 9d and 12d are 9 and 12 days of drought stress. Rewater is rehydration for 4 days after drought stress. Different lowercase letters indicate significant differences among CK and different treatments at P < 0.05 level. * indicate significant difference between epicuticle and lower epidermis at P < 0.05 level.


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图2不同处理条件下直立型扁蓿豆幼苗叶片上表皮(A)和下表皮(B)的显微结构
Figure2.Microstructure of epidermis (A) and lower epidermis (B) of leaf of Medicago ruthenica (L.) Sojak. cv. Zhilixing under drought stress and rehydration


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表1干旱和复水条件下直立型扁蓿豆生理指标的变化
Table1.Physiological indexes of Medicago ruthenica (L.) Sojak. cv. Zhilixing under drought stress and rehydration

生理指标 Physiological index		正常水分 Control	干旱胁迫天数Drought stress days (d)		复水4 d Rehydration for 4 days
			9	12
叶绿素含量 Chlorophyll content [mg·g-1(FW)]	a	1.12±0.14b	2.28±0.12a	0.89±0.10c	1.16±0.09b
	b	0.45±0.045b	0.67±0.07a	0.39±0.05b	0.43±0.057b
	a+b	1.58±0.14c	2.95±0.09a	1.28±0.05d	1.59±0.07c
	a/b	2.47±0.47c	3.39±0.22a	2.28±0.55c	2.70±0.38b
可溶性糖含量Soluble sugar content (mg·g-1)		22.54±2.14b	54.21±1.43a	13.41±0.72c	20.43±1.29b
可溶性蛋白含量Soluble protein content (mg·g-1)		14.34±0.54b	35.53±2.99a	7.86±0.43c	13.14±1.09b
脯氨酸含量Proline content (μg·g-1)		8.74±0.45b	19.64±1.33a	5.99±0.47c	8.66±0.61b
SOD活性SOD activity (U·g-1·min-1)		107.02±5.74c	202.22±6.59a	78.55±6.32d	108.77±11.36c
POD活性POD activity (U·g-1·min-1)		10 205.18±930.74c	21 556.87±1 436.91a	5 772.46±327.54d	10 045.42±979.20c
MDA含量MDA content (mmol·g-1)		0.038±0.003 5c	0.047±0.002b	0.086±0.002 4a	0.048±0.002 0b
同行不同小写字母表示不同处理间在P < 0.05水平差异显著。Different lowercase letters indicate significant differences at 0.05 level among different treatments.

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表2干旱和复水条件下直立型扁蓿豆生物量分配
Table2.Biomass distribution of Medicago ruthenica (L.) Sojak. cv. Zhilixing under drought stress and rehydration

项目 Item	胁迫9天Drought stress 9 days			胁迫12天Drought stress 12 days			复水4天Rehydration 4 days
	正常水分 Control	干旱胁迫 Drought stress		正常水分 Control	干旱胁迫 Drought stress		正常水分 Control	干旱胁迫 Drought stress
总生物量Total biomass (g)	0.372 7*	0.433 7b*		0.375 9*	0.402 4c*		0.379 4*	0.473 6a*
地上生物量Aboveground biomass (g)	0.264 9*	0.296 4b*		0.267 1*	0.251 6c*		0.269 9*	0.313 6a*
地下生物量Root biomass (g)	0.107 8*	0.137 4b*		0.108 8*	0.150 8ab*		0.109 4*	0.160 1a*
根冠比Root/shoot	0.407 1*	0.463 6c*		0.407 4*	0.599 6a*		0.405 4*	0.510 3b*
不同小写字母表示不同胁迫天数之间差异显著，*表示不同处理与其对照间差异显著(P < 0.05)。Different lowercase letters indicate significant differences at 0.05 level among different days of stress. * indicate significant differences between control and stress treatments on the same day at P < 0.05 level.

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表3干旱条件下直立型扁蓿豆叶片形态及生理参数的可塑性指数
Table3.Plasticity index of Medicago ruthenica (L.) Sojak. cv. Zhilixing under drought stress

参数Parameter		干旱胁迫天数Drought stress days (d)
		9	12
形态可塑性 Morphological plasticity	气孔开放率 Stoma opening rate	0.13	0.09
	气孔密度Stomatal density	0.15	0.10
	表皮细胞密度 Epidermal cell density	0.12	0.23
	根冠比Root/shoot	0.10	0.50
	生物量Biomass	0.09	0.26
	均值Average	0.12	0.24
生理可塑性 Physiological plasticity	可溶性糖Soluble sugar	0.15	0.21
	可溶性蛋白Soluble protein	0.16	0.22
	脯氨酸Proline	0.19	0.24
	SOD	0.19	0.18
	POD	0.20	0.17
	MDA	0.09	0.11
	叶绿素Chlorophyll	0.12	0.13
	均值Average	0.16	0.18

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