不同抗旱性花生品种根系形态及生理特性

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厉广辉, 万勇善, 刘风珍, 张昆. 不同抗旱性花生品种根系形态及生理特性. 作物学报, 34(3): 531-541
LI Guang-Hui, WAN Yong-Shan, LIU Feng-Zhen, ZHANG Kun. Morphological and Physiological Traits of Root in Different Drought Resistant Peanut Cultivars. 作物学报, 34(3): 531-541 复制到剪切板

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不同抗旱性花生品种根系形态及生理特性

厉广辉, 万勇善^*, 刘风珍, 张昆

作物生物学国家重点实验室 / 山东农业大学农学院, 山东泰安271018

^* 通讯作者(Corresponding author): 万勇善, E-mail:yswan@sdau.edu.cn

收稿日期:2013-12-15

基金:本研究由国家现代农业产业技术体系建设专项(CARS-14), 国家自然科学基金项目(31201167)和山东省花生良种产业化工程项目资助。;

摘要

以12个花生品种为试验材料, 在人工控水条件下, 通过苗期及结荚期干旱试验, 对比分析花生品种苗期根系性状与抗旱性的关系。结果表明, 花生苗期与结荚期抗旱性基本一致。利用产量抗旱系数可把12个花生品种的抗旱性划分为强、中、弱3级, 抗旱性强的品种为A596、山花11和如皋西洋生, 中度抗旱品种为花育20、农大818、海花1号、山花9号和79266, 抗旱性弱的品种有ICG6848、白沙1016、花17和蓬莱一窝猴。山花11可作为花生强抗旱性鉴定的标准品种, 79266可作为花生弱抗旱性鉴定的标准品种。山花9号、山花11、花育20的根系具有较大的根量及根系吸收能力, 而A596、如皋西洋生、农大818、山花11具有较强的根系抗氧化能力及膜稳定性。相关分析表明, 苗期重度干旱胁迫下的单株根系干重、体积、总吸收面积、超氧化物歧化酶(SOD)活性和丙二醛(MDA)含量与品种抗旱系数的相关性达极显著水平, 对照与重度干旱胁迫下的以上性状呈极显著正相关。因此, 在花生出苗后10 d进行40%土壤相对含水量的干旱胁迫, 持续胁迫至出苗24 d的单株根系干重、体积、总吸收面积、根尖SOD活性和MDA含量可鉴定花生品种的根系抗旱能力, 正常水分下的性状值也能反映根系性状的抗旱级别。山花11可作为花生根系形态及生理优异抗旱性状鉴定的标准品种。

关键词: 花生品种; 干旱胁迫; 根系特性; 抗旱性

Morphological and Physiological Traits of Root in Different Drought Resistant Peanut Cultivars

LI Guang-Hui, WAN Yong-Shan^*, LIU Feng-Zhen, ZHANG Kun

National Key Laboratory of Crop Biology / College of Agronomy, Shandong Agricultural University, Tai#cod#x02019;an 271018, China

Fund:

Abstract

Drought stress is a serious constraint for peanut production worldwide. It is necessary to identify the drought resistance mechanisms of different peanut cultivars in drought-resistance breeding. Under the artificial water control condition, the peanut root morphological and physiological characteristics under drought stress at seedling stage and pod-setting stage were studied using 12 different drought-resistance peanut cultivars as material. The results showed that drought resistance at seedling stage was basically identical with that at pod-setting stage. According to yield-drought resistance coefficient, 12 peanut cultivars were divided into three grades: high-resistance, including A596, Shanhua 11, and Rugaoxiyangsheng; mid-resistance, including Huayu 20, Nongda 818, Haihua 1, Shanhua 9, and 79266; and weak-resistance, including ICG6848, Baisha 1016, Hua 17, and Penglaiyiwohou. In those peanut cultivars, Shanhua11 can be used as the standard cultivar for high drought resistance identification, and 79266 as the standard cultivar for weak one. The root drought resistance mechanism of 12 peanut cultivars were different, Shanhua 9, Shanhua11 and Huayu 20 presented a larger biomass and strong absorption capacity, while A596, Nongda 818, Shanhua 11, and Rugaoxiyangsheng had strong antioxidant capacity and membrane stability under drought stress. Correlations between drought resistance and root weight, volume, total absorption area per plant, root superoxide dismutase (SOD) activity, malondialdehyde (MDA) content under serious drought stress were significant, also under control condition. Therefore, under drought stress of 40% RWC for 10 to 24 days after germination, the root weight, volume, total absorption area per plant, SOD activity and MDA content could be used for identifies the drought resistance ability of peanut roots, the resistances degree also can be reflected by the indices above under normal water condition. Shanhua 11 can be used as a suitable standard cultivar for root morphological and physiological drought resistance characteristics identification in peanut.

Keyword: Peanut cultivars; Drought stress; Root traits; Drought resistance

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我国花生( Arachis hypogaea)总产居世界第一位, 是稳定植物油市场的重要支柱, 但主要分布于干旱半干旱的丘陵地区, 生长季降雨量不均且年度间波动较大, 有70%的花生受到不同程度的干旱威胁, 干旱引起的花生减产率平均在20%以上, 干旱是限制花生产量提高的主要因素^{[ 1, 2]}。解决花生旱害最直接有效的方法就是选育抗旱品种。因此, 研究花生品种的抗旱性, 准确评价抗旱相关性状, 进而筛选不同种质所具有的优异抗旱性状, 是抗旱育种的重要前提。

多年来, 国内外学者在作物抗旱性鉴定方法和指标方面做了大量的研究工作^{[ 3, 4, 5, 6]}, 主要是利用多种综合性状, 采取隶属函数值、主成分分析及灰色关联等分析方法对品种的抗旱性评价分级^{[ 6, 7, 8]}。有关花生品种的抗旱鉴定也有初步研究。已有学者利用干旱胁迫下多种农艺性状与生理性状相对值的隶属函数值和抗旱性综合评价值(D)综合评价花生品种的抗旱性, 并对部分品种的抗旱性分级^{[ 9, 10]}; 亦有研究利用产量的抗旱系数评价花生的抗旱性, 发现部分形态性状与抗旱性显著相关^{[ 11]}; 品种的抗旱性主要是以自然干旱条件下叶片的萎蔫指数来分级^{[ 12]}。由于作物抗旱性是受多基因控制的数量性状, 与基因型、农艺性状及生理生化性状密切相关, 产量是植株整体生理生态机制对干旱响应的结果, 因此, 以生理性状及萎蔫指数评价品种的抗旱性存在片面性。产量抗旱系数是整株形态及生理生化性状对干旱综合反应的结果, 是品种抗旱性鉴定的准确评价

指标, 得到了一致认可^{[ 13, 14]}。但不同品种的抗旱机制并不完全相同, 仅用抗旱系数鉴定抗旱性不能反映品种间抗旱机制的差异。

若以产量抗旱系数评价品种抗旱性, 根据形态及生理性状与抗旱系数的相关分析筛选出抗旱性状, 对比各品种抗旱性状指标可评价其优势性状, 进而可以揭示不同品种的抗旱机制, 鉴定出具有不同优异性状的抗旱花生品种。根系不但是水分吸收的主要器官, 而且通过释放干旱信号调节气孔开度, 影响水分的散失, 与作物抗旱性关系十分密切^{[ 15]}, 不同耐旱品种根系形态特征与生理活性存在显著差异^{[ 16, 17, 18]}。本文以5大花生类型12个品种为材料, 设置苗期与结荚期干旱胁迫试验, 利用产量抗旱系数评价品种的抗旱性, 研究根系形态及生理生化性状与抗旱性的关系, 阐述鉴定不同根系抗旱性状的方法, 筛选具有的优异抗旱性状的品种, 以期为大量品种的抗旱性状鉴定及抗旱亲本选用提供试验方法和理论依据。

1 材料与方法

1.1 供试材料

选用花生栽培种5种植物学类型中12个代表性的品种(表1)。龙生型品种的抗旱性较其他植物学类型的花生强, 珍珠豆型和多粒型品种的抗旱性较弱。所选的珍珠豆型、中间型和普通型材料是山东省不同时期主推的花生品种。

表1 试验材料及其植物学类型 Table 1 Materials and their botanical types

1.2 试验设计

2011—2012年在山东农业大学农学实验站, 采用盆栽进行苗期干旱试验, 以聚酯薄膜制作的防雨棚遮雨。挖取0~30 cm耕作层土壤自然风干过筛, 风干土含水量为1.62%, 田间持水量为23.56%。盆内径30 cm、深40 cm, 每盆装风干土18 kg, 灌等量水自然蒸发失水至适宜含水量时播种, 选取饱满一致的种子2粒穴播, 每盆播3穴, 每处理种植10盆。出苗后10 d开始干旱胁迫处理, 设定土壤相对含水量分别为70% (对照)、55% (轻度干旱)和40% (重度干旱), 采用每日称重法控水, 于每天早8:00称盆重补水并记录(忽略不同盆间生物量的变化), 处理期间除盆内土壤水分差异外其他管理一致。干旱胁迫至始花期恢复供水, 持续14 d。在复水前一天取每处理长势一致的3盆, 去掉地上部, 将盆土倒入网袋, 在水桶浸泡后再冲洗去泥土, 用纸巾拭去根表面的水分后测量整株根系农艺性状, 取侧根根尖部位分别编号, 封入密封袋并迅速放入液氮速冻, 带回实验室于-40℃低温冰箱中贮存, 用于生理生化指标的测定。每处理取3盆, 植株按根茎叶分开烘干称重, 用于干物质积累量和抗旱系数的计算。

于花生栽培池内进行结荚期干旱试验, 池子上方有电动防雨棚遮雨, 栽培池规格为3 m×3 m, 深1.5 m, 下不封底, 池墙由混凝土浇铸。试验材料同苗期干旱试验。采用平种覆膜, 行距40 cm, 穴距18 cm, 密度为每公顷13.9万穴, 每穴播2粒。结荚期维持50%±5%的土壤相对含水量, 持续至饱果出现(30 d), 用CNC503DR型中子水分仪测定0~80 cm土层含水量, 每20 cm一层, 每4 d测定一次, 采用测墒补灌法保证在处理水平, 以70%±5%相对含水量作为对照, 每处理3次重复。其他生育期不遮雨, 同一般高产田管理。收获时从每小区量取4.5 m²实收测产, 计算抗旱系数。

1.3 测定项目与方法

1.3.1 抗旱性参照胡标林等^{[ 13]}方法计算抗旱系数(drought resistance coefficient, DC)和抗旱指数(drought resistance index, DI), DC = Y_d/ Y_p, DI= ( Y_d / Y_p) × ( Y_d / Y_md), 式中 Y_d为干旱胁迫下产量或生物量, Y_p为非胁迫下产量或生物量, Y_md为参试品种干旱胁迫下的平均产量。以抗旱系数评价品种的抗旱性。

1.3.2 根系农艺性状取整株根系, 以排水法测定根系体积, 用吸水纸小心吸干根系表面水分, 以甲烯蓝吸附法测定根系总吸收面积和活跃吸收面积^{[ 19]}; 以根系总吸收面积与干重的比值计算根系比吸收面积^{[ 15]}。105℃杀青半小时, 70℃烘干至恒重称取整株根系干重。

1.3.3 根系生理特性称取0.5 g根系样品用5 mL含1%聚乙烯吡咯烷酮(PVP)的50 mol L^-1磷酸缓冲液(pH 7.0)及少量石英沙, 冰浴研磨, 匀浆于10 000× g冷冻离心20 min, 上清液即为酶提取液。参照王爱国等^{[ 20]}的方法测定超氧化物歧化酶(superoxide dismutase, SOD)活性, 利用SOD抑制氮蓝四唑(NBT)在荧光下的还原作用, 酶活性以抑制NBT还原50%为一个酶活性单位。参照Wakamatsu等^{[ 21]}的方法测定过氧化氢酶(catalase, CAT)活性, 以每分钟减少0.01为1个酶活单位。参照林植芳等^{[ 22]}的方法测定丙二醛(malondialdehyde, MDA)含量。参照陈建勋等^{[ 23]}的方法测定抗坏血酸(ascorbic acid, AsA)和还原型谷胱甘肽(glutathione, GSH)含量。

1.4 数据处理

采用Microsoft Excel 2003计算试验数据平均值, DPS7.5统计分析及SigmaPlot 10.0作图。各性状相对值(relative value, RV)=水分胁迫下性状测定值/正常供水性状测定值。

2 结果与分析

2.1 不同花生品种的产量及抗旱性

由表2可知, 苗期干旱胁迫降低了花生的干物质积累量, 随干旱胁迫加剧呈逐渐下降趋势, 而不同品种对干旱胁迫的响应存在显著差异。12个品种在轻度干旱和重度干旱胁迫下的苗期生物量抗旱系数, 2011年分别为0.77~0.99和0.58~0.82, 2012年分别为0.83~0.97和0.50~0.79。结荚期干旱胁迫降低了各花生品种的产量, 不同品种的降幅不同, 反映了品种抗旱性差异(表2)。2011年12个品种产量的抗旱系数和抗旱指数分别在0.73~0.97和0.57~1.17之间; 2012年也分别在0.76~0.99和0.49~1.24之间, 两年结果基本一致。

两年中苗期轻度干旱胁迫的生物量抗旱系数与结荚期产量抗旱系数有一定的差异( r = 0.56、0.58), 重度干旱胁迫下苗期与结荚期抗旱系数大小排序基本一致( r = 0.93、0.94, P<0.01)。表明重度干旱胁迫下生物量抗旱系数鉴定的苗期抗旱性和以产量抗旱系数鉴定的结荚期抗旱性基本一致。根据抗旱系数的大小, 12个品种抗旱性为如皋西洋生>山花11>A596>花育20>农大818>山花9号>海花1号>79266>蓬莱一窝猴>花17>白沙1016>ICG6848。

2.2 干旱胁迫下不同花生品种根系形态

花生品种根系形态特性差异显著, 不同品种根系性状对干旱胁迫的响应不同, 根系生物量、总吸收面积、活跃吸收面积在轻度干旱下有所增加, 重度干旱下不同程度降低; 根系体积随干旱胁迫加重呈逐渐降低的趋势; 根系比吸收面积在不同干旱胁迫下表现较稳定(图1)。品种间2年结果表现的规律基本一致, 以2012年结果为例进行分析。

2.2.1 根系干重 12个品种根系干重在0.73~ 0.98 g之间, A596、海花1号、花育20和山花9号具有较大的根系生物量(图1)。苗期55%相对含量的干旱胁迫不同程度地增加了各花生品种的单株根系干重, 相对值为1.04~1.22, A596和如皋西洋生的增幅较大。土壤含水量降至40%, 不同品种单株根系干重较对照有不同程度的降低, 相对值为0.70~1.11, 农大818、如皋西洋生和山花11仍高于对照, 花育20、如皋西洋生和山花9号具有较大的根系生物量。

2.2.2 根系体积在对照、轻度干旱及重度干旱胁迫下, 12个花生品种根系体积变化范围分别为7.13~ 14.80 mL、7.07~11.95 mL、6.03~10.27 mL; 花育20、山花9号、山花11的根系体积在对照及干旱胁迫下均显著高于其他品种(图1)。55%相对含水量的干旱胁迫下, 各品种根系体积相对值为0.78~0.99, 其中, A596、农大818与对照接近, 其他品种的降幅明显但品种间无显著差异。40%相对含水量的干旱胁迫下, 各品种根系体积相对值为0.62~0.97, 山花9号、花育20的根系体积显著高于其他品种, A596、农大818的根系体积仍与对照接近。

2.2.3 根系吸收能力由图1可知, 山花9号、山花11、花育20的根系总吸收面积和活跃吸收面积在对照及干旱胁迫下均较高; ICG6848、白沙1016、蓬莱一窝猴则表现相反的规律。55%相对含水量干旱胁迫降低了白沙1016、花育20、花17和蓬莱一窝猴的根系总吸收面积与活跃吸收面积, 其他品种均有不同程度的增加。在40%相对含水量的干旱胁迫下农大818、山花9号和山花11的根系总吸收面积与活跃吸收面积仍高于对照, ICG6848、白沙1016与蓬莱一窝猴大幅降低且相对值显著低于其他品种。品种间的根系比吸收面积无显著差异, 相对值在0.91~1.24之间。

表2 苗期及结荚期干旱胁迫下不同花生品种的抗旱性 Table 2 Drought resistance of peanut cultivars under drought stress at seedling and pod-setting stages

品种 Cultivar	苗期生物量及抗旱性 Drought resistance and biomass at seedling stage					结荚期产量及抗旱性 Drought resistance and yield at pod-setting stage
品种 Cultivar	对照 Control (g plant^-1)	轻度干旱 MD (g plant^-1)	重度干旱 SD (g plant^-1)	轻度干旱抗旱系数 DC of MD	重度干旱抗旱系数 DC of SD	对照 Control (kg hm^-2)	干旱胁迫 Drought (kg hm^-2)	抗旱系数 DC	抗旱指数 DI
2011
79266	3.12 d	2.55 ef	1.99 e	0.82 def	0.64 def	4291 c	3452 b	0.80 efg	0.82 de
A596	3.68 c	3.33 abc	2.81 abc	0.91 bc	0.76 ab	3621 ef	3429 b	0.95 ab	0.96 c
ICG6848	3.86 bc	3.27 bcd	2.23 de	0.85 cd	0.58 f	3299 fg	2518 e	0.76 gh	0.57 g
白沙1016 Baisha 1016	3.15 d	2.44 f	1.85 e	0.77 f	0.59 f	3765 de	2751 de	0.73 h	0.59 g
海花1号 Haihua 1	3.58 c	3.04 bcd	2.56 bcd	0.85 cd	0.71 bc	4908 a	4163 a	0.85 def	1.04 bc
花17 Hua 17	4.25 a	3.70 a	2.72 abc	0.87 bcd	0.64 def	4119 cd	3023 cd	0.73 h	0.66 fg
花育20 Huayu 20	4.30 a	3.41 ab	2.90 ab	0.79 ef	0.67 cde	3878 de	3191 bc	0.82 efg	0.78 ef
农大818 Nongda 818	3.54 c	2.90 de	2.49 cd	0.82 def	0.70 bcd	4436 bc	3965 a	0.89 bcd	1.05 bc
蓬莱一窝猴 Penglaiyiwohou	3.12 d	2.92 cde	1.93 e	0.93 ab	0.62 ef	3178 g	2494 e	0.79 fgh	0.58 g
如皋西洋生 Rugaoxiyangsheng	3.71 c	3.69 a	3.05 a	0.99 a	0.82 a	3089 g	2997 cd	0.97 a	0.86 d
山花11 Shanhua 11	3.73 c	3.34 abc	3.01 a	0.89 bcd	0.81 a	4682 ab	4305 a	0.92 abc	1.17 a
山花9号 Shanhua 9	4.15 ab	3.49 ab	2.96 a	0.84 cde	0.71 bc	4979 a	4322 a	0.87 cde	1.11 ab
2012
79266	3.59 d	3.13 ef	2.25 d	0.87 bcd	0.63 d	4173 d	3620 cd	0.87 de	0.88 cd
A596	4.28 b	3.99 bc	3.16 abc	0.93 ab	0.74 b	3818 ef	3684 cd	0.97 ab	1.01 bc
ICG6848	4.13 bc	3.65 cd	2.07 d	0.88 bc	0.50 f	2979 h	2267 h	0.76 g	0.49 f
白沙1016 Baisha 1016	3.49 d	2.91 f	1.87 d	0.83 d	0.53 ef	3749 f	2978 fg	0.79 fg	0.67 e
海花1号 Haihua 1	4.23 bc	3.85 c	2.83 bc	0.91 abc	0.67 c	4767 ab	4190 ab	0.88 de	1.04 b
花17 Hua 17	5.15 a	4.44 a	3.21 ab	0.86 cd	0.62 d	4089 de	3359 de	0.82 ef	0.78 de
花育20 Huayu 20	4.96 a	4.28 ab	3.27 a	0.86 cd	0.66 cd	3822 ef	3599 cde	0.94 abc	0.95 c
农大818 Nongda 818	4.11 bc	3.62 cd	2.80 c	0.88 bc	0.68 c	4268 cd	3908 bc	0.92 bcd	1.01 bc
蓬莱一窝猴 Penglaiyiwohou	3.63 d	3.38 de	2.01 d	0.93 ab	0.55 e	3378 g	2801 g	0.83 ef	0.65 e
如皋西洋生 Rugaoxiyangsheng	3.83 cd	3.73 cd	2.91 abc	0.97 a	0.76 ab	3290 g	3245 ef	0.99 a	0.90 cd
山花11 Shanhua 11	4.14 bc	3.78 cd	3.29 a	0.91 abc	0.79 a	4548 bc	4475 a	0.98 a	1.24 a
山花9号 Shanhua 9	4.25 bc	3.80 cd	2.96 abc	0.89 bc	0.70 c	5045 a	4508 a	0.89 cd	1.13 ab

MD: 中度干旱; SD: 重度干旱; DC: 抗旱系数; DI: 抗旱指数。同一列内标有相同小写字母的值差异未达到5%显著水平。

MD: mild drought; SD: serious drought; DC: drought resistance coefficient; DI: drought resistance index. Values followed by different letters are significantly different at the 5% probability level.

表2 苗期及结荚期干旱胁迫下不同花生品种的抗旱性 Table 2 Drought resistance of peanut cultivars under drought stress at seedling and pod-setting stages

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图1 干旱胁迫下不同花生品种苗期根系形态指标品种编号1~12分别代表79266、A596、ICG6848、白沙1016、海花1号、花17、花育20、农大818、蓬莱一窝猴、如皋西洋生、山花11和山花9号。Fig. 1 Roots morphological traits of different peanut cultivars under drought stress at seedling stageThe number of cultivars from 1 to 12 represented 79266, A596, ICG6848, Baisha 1016, Haihua 1, Hua 17, Huayu 20, Nongda 818, Penglaiyiwohou, Rugaoxiyangsheng, Shanhua 11, and Shanhua 9, respectively.

2.3 干旱胁迫下不同花生品种根系的抗氧化能力

不同花生品种的根系抗氧化能力显著不同, 能表现品种间抗旱性的差异。随干旱胁迫加重, 抗旱性强的品种根系SOD及CAT活性、GSH及ASA含量呈逐渐增加的趋势, MDA含量缓慢增加; 其他品种以上抗氧化指标呈先增加后降低的趋势, MDA含量在轻度干旱时就迅速增加(图2)。两年结果的趋势基本一致, 文中分析了2012年的结果。

2.3.1 SOD、CAT活性由图2可知, 农大818、山花11、A596、如皋西洋生的两种酶活性在对照及干旱胁迫下均较高。苗期干旱胁迫增加了两种酶的活性, 随土壤含水量的下降, A596、农大818、如皋西洋生、山花11呈逐渐增加的趋势; ICG6848、白沙1016、花17、79266、山花9号呈先增加后降低的趋势; 海花1号与蓬莱一窝猴表现稳定。55%相对含水量的干旱胁迫下, 海花1号与花育20的SOD活性增幅最大, ICG6848、白沙1016与山花9号的CAT活性显著升高, 农大818与蓬莱一窝猴的两种酶活性均未明显升高。40%相对含水量的干旱胁迫下, A596、农大818、如皋西洋生、山花11的2种酶活性及其增幅显著高于其他品种。

2.3.2 GSH、ASA含量随干旱胁迫加重A596、农大818、如皋西洋生、山花11的根系GSH、ASA含量逐渐增加, 且在对照与干旱胁迫下均有较高的含量; 其他品种均呈先增加后降低的趋势(图2)。55%相对含水量的干旱胁迫下, 12个品种的根系GSH、ASA含量相对值分别在1.06~1.64和1.21~1.75之间, 总趋势是抗旱性强的品种其相对值较小, 其含量增加幅度较小, 抗旱性弱的品种, 增加幅度较大。40%相对含水量的干旱胁迫下, GSH、ASA含量相对值分别在0.76~1.50和0.94~1.68之间, 总趋势是抗旱性强的品种其含量增加幅度较大。

2.3.3 MDA含量 55%土壤相对含水量的干旱胁迫下, 12个品种根系MDA含量相对值在1.06~ 1.75之间, 其中, ICG6848、79266、白沙1016、花17快速增加, A596与如皋西洋生基本维持在对照水平(图2)。40%相对含水量的干旱胁迫下, 根系MDA含量相对值在1.35~2.06之间, 山花11、农大818、A596、如皋西洋生的相对值显著低于其他品种, 膜伤害程度较轻。两种干旱胁迫下均表现为抗旱性强的品种的增加幅度明显低于抗旱性弱的品种。

2.4 花生根系特性与品种抗旱性的关系

正常供水时花生根系干重、体积、吸收面积、抗氧化酶活性与品种的抗旱系数的相关性未达显著水平(表3)。55%相对含水量干旱胁迫下, 2年间仅SOD活性和MDA含量与品种的抗旱性呈极显著相关( P<0.01)。在40%相对含水量胁迫下, 2年间根系干重、体积、吸收面积、SOD、CAT活性及GSH、AsA含量与品种抗旱系数的相关性分别达显著( P<0.05)或极显著水平( P<0.01)。MDA含量在正常供水及干旱胁迫下均与抗旱性极显著负相关。轻度及重度干旱胁迫下, 根系干重、根系体积、根系总吸收面积、根系活跃吸收面积、SOD活性、AsA和MDA含量与对照水平下对应的各性状的相关性显著或极显著(表4), 抗旱性较强的花生品种在正常生长及干旱胁迫下均具有较大的根系生物量及较强的抗氧化能力。

3 讨论

不同花生品种在抗旱性方面存在遗传上的差异, 这是许多研究者的一致结论^{[ 2, 9, 10, 11]}。本研究分别以花生苗期干旱处理的生物量抗旱系数和结荚期干旱处理的产量抗旱系数评价12个花生品种的抗旱性, 发现苗期重度干旱胁迫下品种的抗旱性与结荚期抗旱性基本一致, 相关极显著( P<0.01)。12个花生品种两年的结荚期干旱处理的产量抗旱系数分别在0.73~0.97和0.76~0.99之间, 结果基本一致, 说明品种间抗旱性存在明显的遗传差异。根据封海胜等^{[ 24]}的标准, 把12个花生品种的抗旱性划分为强、中、弱3级。有研究表明龙生型品种A596是强抗旱性花生品种, 79266的抗旱性较弱^{[ 9, 11]}。张智猛等^{[ 6, 10]}研究认为抗旱系数大于0.98为抗旱性强的品种, 小于0.86为弱抗旱品种。本文发现, A596和79266的抗旱系数分别为0.97和0.87, 本试验条件下, 产量抗旱系数高于0.97为抗旱性强的品种, 0.87~0.97为中度抗旱品种, 小于0.87为抗旱性弱的品种, 与前人研究结果基本一致^{[ 6, 9, 10, 11]}。但由于干旱时间、胁迫强度等不同, 同一品种的抗旱系数存有差异。12个品种的产量抗旱指数在0.49~1.24之间, 反映了在干旱胁迫下品种的产量差异。通过对抗旱系数与抗旱指数的综合评价, 山花11是抗旱高产品种, 如皋西洋生是抗旱低产品种, ICG6848与白沙1016属弱抗低产品种, 海花1号与山花9号属中抗高产品种。

作物抗旱机制的鉴定和评价, 不仅与所用品种抗旱性差异有关, 选用的评价性状也非常重要, 抗旱相关性状缺乏适当的鉴定方法及评价标准已成为

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图2 干旱胁迫下不同花生品种苗期根系抗氧化保护相关生理性状品种编号1~12分别代表79266、A596、ICG6848、白沙1016、海花1号、花17、花育20、农大818、蓬莱一窝猴、如皋西洋生、山花11和山花9号。Fig. 2 Roots physiological traits related to antioxidant system of peanut cultivars under drought stress at seedling stageThe number of cultivars from 1 to 12 represented 79266, A596, ICG6848, Baisha 1016, Haihua 1, Hua 17, Huayu 20, Nongda 818, Penglaiyiwohou, Rugaoxiyangsheng, Shanhua 11, and Shanhua 9, respectively.

表3 苗期花生根系形态及生理性状与品种抗旱性的相关系数 Table 3 Correlation coefficients of root traits of peanut cultivars with drought resistance

TAA: 总吸收面积; TAA/DR: 比吸收面积; AAA: 活跃吸收面积。^*表示 P< 0.05的显著水平,^**表示 P< 0.01的显著水平。

TAA: total absorption area; TAA/DR: ratio of total absorption area to root dry weight; AAA: active absorption area.^* Significant at P< 0.05;^** significant at P< 0.01.

表4 苗期不同干旱处理下根系性状间的相关系数 Table 4 Correlation coefficients of root traits under drought stress and control

TAA: 总吸收面积; TAA/DR: 比吸收面积; AAA: 活跃吸收面积。^*表示 P< 0.05的显著水平;^**表示 P< 0.01的显著水平。

TAA: total absorption area; TAA/DR: ratio of total absorption area to root dry weight; AAA: active absorption area.^* Significant at P< 0.05;^** significant at P< 0.01.

作物抗旱机制研究的关键问题。本研究选择的12个花生品种, 分属五大植物学类型, 抗旱性差异较大, 利于研究花生的抗旱机制。作物首先感受土壤干旱的器官是根系, 根系特性与抗旱性有密切的关系, 已成为研究和改良作物抗旱性的一个重要组成部分^{[ 13, 14, 15, 16]}。大量的研究认为根系干重、根系体积、根系吸收面积等形态指标与品种抗旱性密切相关^{[ 16, 25, 26]}。根系总吸收面积与活跃吸收面积是衡量根系吸收功能和活力的重要指标, 干旱胁迫下较大的根系吸收面积能提高作物吸水效率, 有利于吸收尽可能多的水分, 提高抗旱性^{[ 27, 28]}。前人只在定性水平上研究, 无法定量描述各性状的抗旱级别, 亦没有研究具体品种的根系特性。本研究发现, 抗旱性越强的花生品种根系干重、根系体积、根系总吸收面积、根系活跃吸收面积越大, 说明较大的根量及较强的根系吸收能力可以有效抵御干旱胁迫对植株所造成的伤害。山花9号的根系干重和根系体积最大, 花育20的根系总吸收面积和活跃吸收面积显著高于其他品种, 山花11的根系体积和总吸收面积较大, 这是它们抗旱的根系机制; A596与如皋西洋生有较强的抗旱性, 但根系生物量及吸收能力未有显著优势, 可能是其他抗旱机制起主导作用。杨守萍等^{[ 26]}研究认为应以干旱胁迫下根系性状的相对值鉴定品种的抗旱性, 刘莹等^{[ 29]}的研究表明干旱胁迫下根系性状占整株的比重可作为衡量大豆抗旱性的指标。本研究认为, 根系形态变化是作物抗旱的机制之一, 并不是整株对干旱胁迫的综合响应, 不能全面反映品种的抗旱性。相关分析表明, 两年间重度干旱胁迫下单株根系总吸收面积与品种抗旱系数均呈极显著正相关( P<0.01), 根系干重和体积与抗旱性的相关系数也达显著( P<0.05)或极显著水平。虽然对照处理的根系性状与品种抗旱性的相关性未达显著水平, 但对照与轻度和重度干旱处理各根系性状的相关性分别达显著和极显著水平, 正常生长时具有较大根系及较强吸收能力的品种, 在干旱胁迫下仍具有显著优势。因此, 可用40%相对含水量胁迫下的单株根系干重、体积和总吸收面积鉴定花生种质的根系抗旱能力。如果鉴定的种质规模较大, 难以实现控水条件, 则正常水分下的根系性状也能反映品种间抗旱性的差异, 这减少了大量的测定工作, 便于多次重复与大规模的品种鉴定。

干旱胁迫导致植物体内活性氧水平提高时, SOD、CAT等酶活性和GSH、AsA等非酶类抗氧化物质增加, 以保持体内活性氧积累与清除系统的平衡^{[ 2, 30]}。齐伟等^{[ 31]}研究认为, 根系SOD、CAT、POD活性在水分胁迫下与品种的抗旱性密切相关, 抗旱性强的品种能维持较高的SOD、CAT活性; 尚晓颍等^{[ 32]}研究认为根系SOD、CAT活性可以用来鉴别干旱胁迫下品种的抗旱性。本研究表明, 干旱胁迫下, 抗旱性越强的品种根系SOD、CAT活性及GSH、AsA含量越高, 说明相对较高的抗氧化酶活性及抗氧化剂含量能减轻活性氧对细胞的伤害, 维持细胞膜的稳定性, 保证根系生长及吸收能力正常。山花11、A596、农大818、如皋西洋生在对照和干旱条件下均保持较高的SOD、CAT活性及GSH、AsA含量, MDA含量显著低于其他品种, 这是它们抗旱的根系生理机制; ICG6848、白沙1016、花17在对照和干旱条件下均有较高的MDA含量, 膜伤害程度较严重, 这可能是它们抗旱性弱的主要原因之一。相关分析表明, 两年间重度干旱胁迫下的根系SOD活性和MDA含量均与品种抗旱性呈极显著相关, 正常灌水下根系MDA含量与品种抗旱性的相关系数也达显著水平, 它们是花生苗期抗旱的根系生理性状。本研究还发现, 对照与重度干旱处理的根系SOD活性和MDA含量呈极显著正相关, 不同品种间正常水分时根系的抗氧化能力能反映干旱胁迫下的趋势, 王思思等^{[ 33]}在玉米研究中也得出相似结论。由此可见, 抗旱花生品种的选育应高度重视正常灌水条件下根系的生理特性。

干旱的持续时间及胁迫强度影响各抗旱性状的测定值, 直接以测定数值的大小划分抗旱性状的等级难免出现误差, 盖钧镒等^{[ 34]}提出了大豆品种生育期组鉴定的标准品种, 依据标准品种作为参考来划分品种的生育期可以排除环境因素的影响。陈加敏^{[ 35]}在研究大豆品种抗旱性时, 根据抗旱性强弱分别筛选出了抗旱标准品种与干旱敏感标准品种。本研究抗旱性强的3个品种中, 山花11是当前生产上使用的品种(中间型, 连续开花), 如皋西洋生、A596为龙生型, 交替开花, 匍匐生长, 生育期特长, 产量非常低, 植物学性状和生物学特性与当前推广品种差异最大, 代表性差。因此, 确定山花11为抗旱性强的标准品种。两年结果发现, 79266均是中抗级别中抗旱系数最低的, 处于中抗和弱抗的临界值。另外, 考虑到79266曾是我国北方花生产区大面积推广的品种, 目前仍有一定种植面积, 也是许多育成品种的亲本之一。属于连续开花型, 与当前推广品种的植物学性状、生物学特性、产量水平差异较小, 在生产中有广泛的代表性。而处于弱抗级别的前两位品种蓬莱一窝猴和花17植物学类型均普通型, 交替开花, 生育期较长, 产量水平较低, 植物学性状和生物学特性与当前推广品种差异较大, 代表性差。因此确定79266为花生弱抗旱性鉴定的标准品种。山花11不仅具有稳定的强抗旱性, 其根系形态和抗氧化能力在对照及干旱胁迫下均表现突出, 可作为花生根系性状鉴定的标准品种, 这样可消除因试验环境不同而产生的性状差异, 使品种抗旱性状得以量化, 鉴定结果更科学、准确。若根系性状优于山花11, 即可认为是品种的优异抗旱性状, 由此可以评价不同花生品种抗旱的主导机制。

4 结论

根据抗旱系数筛选出A596、如皋西洋生和山花11为强抗旱性花生品种。山花11可作为花生强抗旱性鉴定的标准品种, 79266可作为弱抗旱性鉴定的标准品种。在花生出苗后10 d进行40%土壤相对含水量持续14 d (出苗后24 d)的干旱胁迫下, 以单株根系干重、体积、总吸收面积、根尖SOD活性和MDA含量可鉴定花生种质的根系抗旱能力, 出苗后维持70%相对含水量至始花期的上述性状值也能反映种质根系抗旱能力的差异。山花11可作为花生根系优异抗旱性状鉴定的标准品种。

The authors have declared that no competing interests exist.

作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.

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[31]	齐伟, 张吉旺, 王空军, 刘鹏, 董树亭. 干旱胁迫对不同耐旱性玉米杂交种产量和根系生理特性的影响. 应用生态学报, 2010, 21: 48-52 Qi W, Zhang J W, Wang K J, Liu P, Dong S T. Effects of drought stress on the grain yield and root physiological traits of maize varieties with different drought tolerance. Chin J Appl Ecol, 2010, 21: 48-52 (in Chinese with English abstract) [本文引用:1] [CJCR: 1.742]
[32]	尚晓颍, 刘化冰, 张小全, 林娟, 段旺军, 杨铁钊. 干旱胁迫对不同烤烟品种根系生长和生理特性的影响. 西北植物学报, 2010, 30: 357-361 Shang X Y, Liu H B, Zhang X Q, Lin J, Duan W J, Yang T Z. Growth and physiological characteristics of roots in different flue-cured tobacco varieties under drought stress. Acta Bot Boreal-Occident Sin, 2010, 30: 357-361 (in Chinese with English abstract) [本文引用:1]
[33]	王思思, 张吉旺, 刘鹏, 董树亭, 王空军. 干旱对不同玉米品种苗期根系生理生化特性的影响. 山东农业科学, 2009, (6): 36-38 Wang S S, Zhang J W, Liu P, Dong S T, Wang K J. Effect of drought on root physiological and biochemical characters of different maize cultivars during seedling stage. Shand ong Agric Sci, 2009, (6): 36-38 (in Chinese with English abstract) [本文引用:1]
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[35]	陈加敏. 大豆苗期耐旱性的鉴定及苗期耐旱性和根系性状的遗传研究. 南京农业大学硕士学位论文, 2004 Chen J M. Studies on Identification of Drought Tolerance and Genetic Mechanism of Drought Tolerance and Root Traits of Soybean Seedling. MS Thesis of Nanjing Agricultural University, Nanjing, China, 2004 (in Chinese with English abstract) [本文引用:1]

2004

0.0

1.667

. 2004, 30:169-174

The effect on SOD activity and protein content in groundnut leaves by drought stress

(干旱胁迫对花生叶片SOD活性和蛋白质的影响) . (作物学报) , 2004 , 30 : 169 - 174 Jiang H F , Ren X P . The effect on SOD activity and protein content in groundnut leaves by drought stress

Jiang H F

(姜慧芳) , (任小平) . (干旱胁迫对花生叶片SOD活性和蛋白质的影响) . (作物学报) , 2004 , 30 : 169 - 174 Jiang H F

Oil Crops Research Institute, CAAS, Wuhan 430062, Hubei

Drought stress is a serious constraint for groundnut production worldwide. Developing genotypes with drought tolerance is one economic approach to abate the damage of stress. Study on the tolerance mechanism of groundnut is necessary to its breeding. The SOD activity, protein content and water poten

以不同类型的抗旱花生种质为材料，于花针期干旱胁迫处理43 d，调查和分析了水分胁迫对叶片SOD活性、蛋白质和水势影响的动态变化以及干旱胁迫对花生主要经济性状的影响。分析结果表明在干旱胁迫初期，花生叶片的SOD活性下降，蛋白质含量增加，此时品种之间的SOD活性变化差异不明显，但蛋白质含量差异显著。在严重干旱胁迫时

... 我国花生(Arachis hypogaea)总产居世界第一位, 是稳定植物油市场的重要支柱, 但主要分布于干旱半干旱的丘陵地区, 生长季降雨量不均且年度间波动较大, 有70%的花生受到不同程度的干旱威胁, 干旱引起的花生减产率平均在20%以上, 干旱是限制花生产量提高的主要因素^[1,2] ...

2007

0.0

1.667

. 2007, 33:113-119

Effects of drought during seedling stage on physiological traits, yield and quality of different peanut cultivars

(苗期干旱胁迫对不同抗旱花生品种生理特性、产量和品质的影响) . (作物学报) , 2007 , 33 : 113 - 119 Yan M L , Li X D , Lin Y J , Wang L L , Zhou L Y . Effects of drought during seedling stage on physiological traits, yield and quality of different peanut cultivars

Yan M L

(严美玲) , (李向东) , (林英杰) , (王丽丽) , (周录英) . (苗期干旱胁迫对不同抗旱花生品种生理特性、产量和品质的影响) . (作物学报) , 2007 , 33 : 113 - 119 Yan M L

The Key Laboratory of Crop Biology of Shandong/ College of Agronomy, Shandong Agricultural University, Tai’an 271018, Shandong, China

Peanut is one of the ideal crops in dry farming. Drought is the main limit factor of peanut to increase yield and improve quality. Seedling stage is constantly hit by drought for summer peanut. In order to understand the suitable watering amount at peanut seedling stage, provide theoretical basis and technique for dry farming peanut producing the experiment was conducted with drought light-resistant peanut cv. Luhua 11 and drought resistant peanut cv. Nongda 818 from 2003 to 2004 in the experimental station of Shandong Agricultural University. Five treatments with watering 20 mm(serious drought stress), 40 mm(middle drought stress), 60 mm(slight-middle drought stress), 80 mm(slight drought stress) and 100 mm(CK) were made at 10 days after seedling(27 June) with three replicates in a randomized block design. The drought stress stopped at full-bloom stage(20 July) and then all treatments were irrigated 100 mm water at full-bloom stage, pod setting stage and pod filling stage separately. The results showed that the photosynthesis rate(P_n) decreased, and malondialdeyde (MDA) content increased remarkably with the drought degree raising. After rewatering, the MDA content decreased and Pn returned to the level of check rapidly, the recovery ability of Nongda 818 was bigger than that of Luhua 11. Suitable drought treatment could enhance the activities of superoxide dismutase(SOD), peroxidase (POD) and catalase (CAT), and increase the soluble protein content. The highest activities of SOD, POD, CAT and soluble protein content presented at the slight-middle drought treatment(watering 60–80 mm ) for Luhua 11, but at the middle drought treatment(watering 40–60 mm ) for Nongda 818. At the same drought treatment, especially at serious drought treatment(watering 20 mm), the activities of SOD, POD, CAT of Nongda 818 were obviously higher than that of Luhua 11. After rewatering, the activities of SOD, POD, CAT and soluble protein content reduced significantly, but the activities of SOD, POD, CAT of Nongda 818 were still higher than that of Luhua 11. With the drought degree raising, pod and kernel yields substantially decreased and the yield losses of Nongda 818 were less than that of Luhua 11. The irrigating amount at peanut seedling stage is no less than 60–80 mm to avoid the yield loss. Middle and slight drought stress at seedling stage could increase the protein content of peanut kernel, and have a little effect on fat content. Serious drought stress at seedling stage remarkably decreased oleic content and the ratio of oleic/linoleic(O/L), increased linoleic content of peanut kernel, and have a little effect on protein content. The optimal irrigating amount at seedling stage for the best kernel quality of Luhua 11 is 60–80 mm and that of Nongda 818 is 40–60 mm.

在人工控水条件下，研究了两个不同抗旱花生品种苗期干旱下的生理特性及其产量和品质。结果表明，随着干旱程度的增加，两品种叶片光合速率(P_n)逐渐下降，丙二醛(MDA)含量增加；解除干旱后，叶内MDA含量降低，Pn很快恢复到或超过对照水平，农大818的恢复能力强于鲁花11。适当干旱处理，可增加叶片中超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)的活性，提高叶片可溶性蛋白质(P_r)含量，鲁花11以中轻度干旱（灌水60~80 mm），而农大818以中度干旱（灌水40~60 mm）时酶活性最强、Pr含量最高，同一干旱处理、特别重度干旱（灌水20 mm）处理的酶活性，农大818明显高于鲁花11；解除干旱后，两品种叶中SOD、POD、CAT活性和Pr含量显著降低，但农大818的酶活性仍高于鲁花11。随着苗期干旱程度的增加，花生荚果和籽仁产量降低，但农大818的降幅小于鲁花11；若苗期灌水量低于60~80 mm则影响产量。花生苗期中、轻度干旱胁迫可增加籽仁蛋白质含量，而对脂肪含量的影响不大；重度干旱明显降低籽仁脂肪含量、脂肪中油酸组分和O/L比值，增加亚油酸组分，但对蛋白质的影响较小。鲁花11以60~80 mm水分处理及农大818以40~60 mm水分处理时籽仁品质最佳。

... 55%相对含水量干旱胁迫下, 2年间仅SOD活性和MDA含量与品种的抗旱性呈极显著相关(PPP3 讨论不同花生品种在抗旱性方面存在遗传上的差异, 这是许多研究者的一致结论^[2,9,10,11] ...

... 相关分析表明, 两年间重度干旱胁迫下单株根系总吸收面积与品种抗旱系数均呈极显著正相关(PP干旱胁迫导致植物体内活性氧水平提高时, SOD、CAT等酶活性和GSH、AsA等非酶类抗氧化物质增加, 以保持体内活性氧积累与清除系统的平衡^[2,30] ...

1984

0.657

0.0

... 多年来, 国内外学者在作物抗旱性鉴定方法和指标方面做了大量的研究工作^[3,4,5,6], 主要是利用多种综合性状, 采取隶属函数值、主成分分析及灰色关联等分析方法对品种的抗旱性评价分级^[6,7,8] ...

2005

1.513

0.0

2008

2.474

0.0

2012

0.0

1.667

. 2012, 38:495-504 DOI:10.3724/SP.J.1006.2012.00471

Identification and evaluation of drought resistance in different peanut varieties widely grown in northern China

(中国北方主栽花生品种抗旱性鉴定与评价) . (作物学报) , 2012 , 38 : 495 - 504 Zhang Z M , Dai L X , Ding H , Chen D X , Yang W Q , Song W W , Wan S B . Identification and evaluation of drought resistance in different peanut varieties widely grown in northern China

Zhang Z M

(张智猛) , (戴良香) , (丁红) , (陈殿绪) , (杨伟强) , (宋文武) , (万书波) . (中国北方主栽花生品种抗旱性鉴定与评价) . (作物学报) , 2012 , 38 : 495 - 504 Zhang Z M

1 Peanut Research Institute of Shandong Province, Qingdao 266100, China; 2 Shandong Academy of Agricultural Sciences, Ji’nan, 250100, China

It is necessary to systematically identify and evaluate drought-resistant varieties for stable development of peanut production in arid and semi-arid regions in China. To establish drought resistance index of peanut cultivars, we determined 13 morphological and biochemical indices including plant height, number of branch, biomass, leaf water content and photosynthetic pigment content in the pot experiment at pod setting and pod filling stages with 29 peanut cultivars (strains) widely grown in north China under artificial water stress condition. Drought resistances of varieties were scored with drought coefficient method and subordinate function value under water stresses. The results showed that the same index of drought resistance characterized in different peanut varieties achieved different results, and 29 peanut varieties were divided into four grads: high tolerant, moderate tolerant, weaker tolerant, and drought susceptible. In the 29 cultivars (strains), we screened out the high drought tolerant varieties consisting of Tangke 8, Jihua 2, Jihua 4, Huayu 25, Huayu 17, Huayu 22, Datangyou, and Huayu 21. The cultivars (strains) of Huayu 20, Huayu 24, Weihua 6, Huayu 27, Luhua 11, Fuhua 11, Fuhua 13, Tangyou 4, Fenghua 1, and G2 had moderate drought resistance. The cultivars (strains) with weaker drought resistance were 16-8, Fuhua 10, M5, Huayu 16, and Luhua 14. At pod setting and pod filling stages Huayu 19, Huayu 23, Fenghua 6, Weihua 8, M7, and TE-2 were all drought susceptible. From seedling to pod setting stages under water stress, all morphological and biochemical indices but the number of branch and carotenoid content and their comprehensive D values could be taken as evaluation criteria. From seedling to pod filling stages under water stress, there only existed significanct correlation between leaf area and drought resistance. Either at pod setting or pod filling stages under water stress, comprehensive D value of the indices had significant correlation with drought resistance, which could be used to identify peanuts drought resistance.

人工控水条件下，通过盆栽试验，在花生植株的结荚期和饱果期, 测定29个品种(系)的株高、分枝数、生物累积量、叶片含水量和光合色素含量等与抗旱性有关的13个表型性状和生理生化指标，采用抗旱系数法和隶属函数值法，鉴定各性状指标水分胁迫下的抗旱性。结果表明,相同指标评价不同花生品种抗旱性时得出的结果不同,依此可将供试品种(系)划分为抗旱性强、中、弱和不抗旱4类,即供试品种中唐科8号、冀花2号、冀花4号、花育25、花育17、花育22、大唐油、花育21具有较强的抗旱能力；花育20、花育24、潍花6号、花育27、鲁花11、阜花11、阜花13、唐油4号、丰花1号和G2具有中等抗旱能力；具有较弱抗旱能力的品种(系)有16-8、阜花10号、M5、花育16和鲁花14；而花育19、花育23、丰花6号、潍花8号、M7和TE-2在结荚期和饱果期均无抗旱能力。水分胁迫至结荚期，除分枝数和类胡萝卜素外，其余各形态型状和生理生化指标及其综合D值均可作为鉴定品种(系)抗旱性的依据；水分胁迫至饱果期，只有叶面积与抗旱系数间的相关极显著。无论在结荚期或饱果期，综合D值可作为鉴定品种抗旱性的指标，单一指标均不能准确判定某品种(系)的抗旱性。

... 张智猛等^[6,10]研究认为抗旱系数大于0 ...

... 87为抗旱性弱的品种, 与前人研究结果基本一致^[6,9,10,11] ...

2007

0.0

1.667

. 2007, 33:1523-1529

Screening indexes of drought resistance during seedling stage in rice

(水稻苗期抗旱性指标的筛选) . (作物学报) , 2007 , 33 : 1523 - 1529 Wang H Z , Li Y , Ma J , Zhang R P , Li X Y , Wang R Q . Screening indexes of drought resistance during seedling stage in rice

Wang H Z

(王贺正) , (李艳) , (马均) , (张荣萍) , (李旭毅) , (汪仁全) . (水稻苗期抗旱性指标的筛选) . (作物学报) , 2007 , 33 : 1523 - 1529 Wang H Z

Rice Research Institute, Sichuan Agricultural University, Wenjiang 611130, Sichuan, China

Rice plants may suffer serious water stress during its growth period, so that change their morphological characteristics and the process of physiological metabolic activity. Screening indexes of drought resistance has been focused in the concerned researched. To explore the morphological and physiological mechanisms and screen indexes of drought resistance during seedling stage in rice, in this paper, morphological and physiological characteristics correlating with drought resistance were monitored under both normal and drought conditions during seedling stage using 15 paddy rice cultivars. The results showed that relative leaf age, relative area of first leaf under the central leaf, relative leaf fresh weight, relative root length were significantly and very significantly correlated with survival percentage under repeated drought condition and could be used as the morphological indexes to identify the drought resistance. Relative activities of superoxide dismutase(SOD), catalase(CAT), peroxidase(POD) and relative contents of amino acid(AA), ascorbic acid(AsA), reduced glutathione (GSH), soluble protein and malondialdehyde(MDA) were significantly and very significantly correlated with the survival percentage under repeated drought condition and could be used as the physiological indexes to identify the drought resistance. The integrated drought resistance indexes including the relative activity of SOD, relative GSH content, leaf fresh weight, and leaf age were established by principal components analysis. The drought resistance of 15 rice cultivars was evaluated by stepwise regression equation and was basically consistent with the results from paddy field, proving it is effective to predict the drought resistance during seedling stage in rice using the integrated identification index screened out in this study.

为探索水稻苗期抗旱性形态和生理机制，筛选苗期抗旱性指标，以15个水稻品种为试验材料，在干旱棚内系统研究了水、旱条件下苗期形态和生理指标的变化。结果表明，叶龄、心叶下倒数第1叶叶面积、叶鲜重、根长4个指标的相对值与反复干旱存活率显著或极显著相关，可作为苗期抗旱性的形态指标；超氧化物歧化酶（SOD）、过氧化物酶（POD）、过氧化氢酶（CAT）活性，氨基酸（AA）、抗坏血酸（AsA）、还原性谷甘肽（GSH）、可溶性蛋白质、丙二醛（MDA）含量8个指标的相对值与反复干旱存活率显著或极显著相关，可作为苗期抗旱性的生理指标。通过主成分分析确立了SOD酶活性、GSH含量、叶鲜重和叶龄4个指标相对值为水稻苗期抗旱性的综合指标，以反复干旱存活率为因变量，4个综合指标为自变量建立的回归方程对供试品种的抗旱性进行预测效果较好。

2007

0.0

1.889

. 2007, 40:2452-2459

Gray relational grade analysis of agronomical and physi-biochemical traits related to drought tolerance in wheat

Wang S Q

(王士强) , (胡银岗) , (佘奎军) , (周琳璘) , (孟凡磊) . (小麦抗旱相关农艺性状和生理生化性状的灰色关联度分析) . (中国农业科学) , 2007 , 40 : 2452 - 2459 Wang S Q

西北农林科技大学农学院

【Objective】This paper aims to probe the relationship between agronomical characters and physi-biochemical indexes of wheat cultivar with its drought tolerance. 【Method】 12 indexes, including 5 agronomical characters and 7 physi-biochemical indexes that measured in 65 wheat cultivars grown at drought shelf and no-irrigation conditions, were analyzed using gray relational grade analysis method. 【Result】 Gray correlation degrees facilitating to identify the drought tolerance of wheat were calculated between each index and its drought resistance index, they were ranked as follows by their contribution to drought tolerance: stomata conductance (γ=0.7995), photosynthetic rate (γ=0.7909), transpiration rate (γ=0.7556), soluble sugar content (γ=0.7467), MDA content (γ=0.7336), dissociative proline content (γ=0.7267), spike length (γ=0.7102), number of kernel per spike (γ=0.7095), 1000-kernel weight (γ=0.7005), water potential (γ=0.6959), tiller (γ=0.6951), plant height (γ=0.6866). And the clustering analysis based on the weighted drought index of cultivars could finely represent their breeding and application areas. Species that have the best drought resistance among the 65 species are: Luohan 6, Xiaoyan 6, Jinmai 47, Shanmai 168 etc. 【Conclusion】 The result indicates that the physi-biochemical indexes related to leaf stomata have the closest relationship with the drought resistance of wheat, and these indexes are significantly influenced under water stress, thus have the greatest relation degree with drought resistance index, so the physi-biochemical indexes including stomata conductance, photosynthetic rate and transpiration rate should be used for the assistant selection of drought tolerance to improve the selection efficiency.

【目的】探讨小麦主要农艺性状和部分生理生化性状与抗旱性的关系。【方法】采用灰色关联分析（gray correlation analysis），对分别种植于干旱棚和大田自然降水两个条件下的65个小麦品种的主要农艺性状和部分生理生化性状与其抗旱指数的关联度进行分析和评价。【结果】通过灰色关联度分析确定了各性状与抗旱指数的关联度。各农艺性状和生理生化性状与抗旱指数的关联程度大小依次为：气孔导度（0.7995）、光合速率（0.7909）、蒸腾速率（0.7556）、可溶性糖含量（0.7467）、丙二醛（MDA）含量（0.7336）、游离脯氨酸含量（0.7267）、穗长（0.7102）、穗粒数（0.7095）、千粒重（0.7005）、水势（0.6959）、有效分蘖数（0.6951）和株高（0.6866）。并依据各品种的加权抗旱指数对参试品种进行了聚类分析，聚类结果可以较好的反映品种的选育及适应地区。参试品种中较抗旱的品种有洛旱6号、晋麦47、陕麦168、小偃6号等。【结论】与抗旱指数关联度最大的是和叶片气孔相关的生理生化性状，它们在作物受到干旱胁迫时所受的影响最大，与作物抗旱性关系最为密切。因而可以选用气孔导度、光合速率和蒸腾速率等生理生化性状进行小麦抗旱性的以性状为基础的选择，以提高选择效率。

2004

0.0

. 2004, 33:8-12

Identification of drought resistance in different peanut varieties

(不同花生品种的抗旱性比较鉴定) . (花生学报) , 2004 , 33 : 8 - 12 Yan M L , Li X D , Jiao Y L , Wang L L . Identification of drought resistance in different peanut varieties

Yan M L

(严美玲) , (李向东) , (矫岩林) , (王丽丽) . (不同花生品种的抗旱性比较鉴定) . (花生学报) , 2004 , 33 : 8 - 12 Yan M L

通过研究花生种子在盛花期与抗旱性密切相关的形态指标与生理指标,对9个不同的花生品种(系)进行了抗旱性鉴定,并用隶属函数法对花生品种的抗旱性进行了综合评价.结果表明,同一花生品种在干旱与正常供水条件下相比较,其株高、单株叶面积、根冠比、比叶重、根茎叶干重及叶绿素含量差异显著,因此,它们可以作为鉴定花生抗旱性的简易指标.不同花生品种的抗旱性有明显差异,并鉴定出高抗旱品种丰花1号,抗旱品种海花1号、农大142、鲁花14号,不抗旱品种农大818、鲁花5号与79266.

... 已有学者利用干旱胁迫下多种农艺性状与生理性状相对值的隶属函数值和抗旱性综合评价值(D)综合评价花生品种的抗旱性, 并对部分品种的抗旱性分级^[9,10] ...

... 有研究表明龙生型品种A596是强抗旱性花生品种, 79266的抗旱性较弱^[9,11] ...

... 87为抗旱性弱的品种, 与前人研究结果基本一致^[6,9,10,11] ...

2011

0.0

1.989

. 2011, 35:100-109 DOI:10.3724/SP.J.1258.2011.00100

Estimating and screening of drought resistance indexes of peanut

(花生抗旱性鉴定指标的筛选与评价) . (植物生态学报) , 2011 , 35 : 100 - 109 Zhang Z M , Wan S B , Dai L X , Song W W , Chen J , Shi Y Q . Estimating and screening of drought resistance indexes of peanut

Zhang Z M

(张智猛) , (万书波) , (戴良香) , (宋文武) , (陈静) , (石运庆) . (花生抗旱性鉴定指标的筛选与评价) . (植物生态学报) , 2011 , 35 : 100 - 109 Zhang Z M

1Peanut Research Institute of Shandong Province, Qingdao 266100, China; 2Shandong Academy of Agricultural Sciences, Jinan 250100, China

Aims Our objective was to define an index of peanut (Arachis hypogaea) drought resistance and comprehensively evaluate drought resistance of peanut varieties under controlled water conditions. Methods The experiment was conducted on the farm of Qingdao Academy of Agricultural Science with 29 peanut varieties. Two levels of water content (80%–85% and 45%–50% of field moisture capacity) were used in a randomized complete block design with four replications. We measured 13 morphological and physiological characters such as plant height, branch numbers, biomass, leaf water content and photosynthetic pigment content in seedling and pod-pin stages. Important findings The 29 peanut cultivars can divided into four classes: high-, mid- and low-tolerant cultivars and intolerant cultivars. The high-tolerant cultivars included ‘Tangke 8’, ‘Jihua 2’, ‘Datangyou’, ‘Huayu 25’, ‘Huayu 17’, ‘Luhua 14’ and ‘Fenghua 1’. Subordinate function values of the morphological indexes including main stem height, branch number and biomass, and photosynthetic pigments contents of the same cultivars at seedling stage were different. In addition, the subordinate function value of the seedling stage was unrelated to drought resistance, so a single physiological index cannot be used to identify the drought resistance of peanut. But the values drought resistance comprehensive evaluation (D) of physiological indexes were significantly related to drought resistance and can be used to identify the drought resistance of peanut. At the pod-pin stage, the D values of morphological indexes and physiological indexes were both significantly related to drought resistance, so the subordinate function value and D value of morphological indexes and physiological indexes at this stage can serve as indexes for identifying peanut drought resistance.

为确定鉴定花生(Arachis hypogaea)品种(系)抗旱性指标体系, 综合评价花生品种(系)的抗旱性, 在人工控水条件下, 通过盆栽试验, 测定了29个花生品种(系)苗期和花针期的株高、分枝数、生物累积量、叶片含水量和光合色素含量等与抗旱性有关的13个表观形态性状和生理性状的指标, 采用抗旱系数法和隶属函数值法, 对各指标性状进行了水分胁迫下的抗性评价和鉴定。结果表明, 29个花生品种(系)可划分为抗旱性较强、中等、较弱和不抗旱4类, 其中‘唐科8号’、‘冀花2号’、‘大唐油’、‘花育25号’、‘花育17号’、‘鲁花14号’、‘丰花1号’ 7个品种(系)具有较强的抗旱能力; 苗期同一品种(系)的主茎高、分枝数和生物累积量等形态指标和光合色素等生理指标的隶属函数值均有较大差别, 苗期各指标隶属函数值与品种(系)抗旱性无显著相关关系, 苗期单一形态指标不能作为鉴定品种(系)抗旱性的指标; 但苗期抗旱性综合评价值(D)与抗旱系数间存在显著相关关系, D的大小可作为抗旱性的鉴定指标。花针期形态指标和生理指标D值间, 以及各类指标D值与抗旱系数间均存在显著或极显著的相关关系, 此期植株形态指标、生理指标隶属函数值以及综合D值均可作为鉴定品种(系)抗旱性的指标。

... 张智猛等^[6,10]研究认为抗旱系数大于0 ...

... 87为抗旱性弱的品种, 与前人研究结果基本一致^[6,9,10,11] ...

1997

0.0

... 亦有研究利用产量的抗旱系数评价花生的抗旱性, 发现部分形态性状与抗旱性显著相关^[11] ...

... 有研究表明龙生型品种A596是强抗旱性花生品种, 79266的抗旱性较弱^[9,11] ...

... 87为抗旱性弱的品种, 与前人研究结果基本一致^[6,9,10,11] ...

1999

0.0

1.889

... 品种的抗旱性主要是以自然干旱条件下叶片的萎蔫指数来分级^[12] ...

2007

0.0

1.667

. 2007, 33:425-432

Drought-resistance identification of Dongxiang common wild rice (Oryza rufipogon) in whole growth period

(东乡普通野生稻全生育期抗旱性鉴定) . (作物学报) , 2007 , 33 : 425 - 432 Hu B L , Yu S W , Wan Y , Zhang Z , Qiu B Y , Xie J K . Drought-resistance identification of Dongxiang common wild rice (Oryza rufipogon) in whole growth period

Hu B L

(胡标林) , (余守武) , (万勇) , (张铮) , (邱兵余) , (谢建坤) . (东乡普通野生稻全生育期抗旱性鉴定) . (作物学报) , 2007 , 33 : 425 - 432 Hu B L

Jiangxi Provincial Key Laboratory for Rice Physiology and Genetics/ Institute of Rice Research, Jiangxi Academy of Agricultural Sciences, Nanchang 330200, Jiangxi, China

The drought and scarcity of water resource are more and more serious, and restrict agricultural development, specially rice production in China. Dongxiang common wild rice distributed in the most north region all over the world contains plentiful genetic diversity and excellent genes encoding drought resistance, cytoplasm male sterility (CMS), cold tolerance, fertility restoration, and so on. Screening drought resistance of dongxiang common wild rice is important for breeding rice with drought resistance and creating new drought resistance resource in current cultivated rice.
With the water stress and non-stress treatments designed, twelve morphological and physiological traits correlating with drought resistance were monitored in whole growth period for the back-cross inbred line population (R974//DongYe/R974) BC₁F₅, which were plant height (PH), tillers number per plant (TNP), effective tillers number per plant (ETNP), spikelets per panicle (SP), filled spikelets per panicle (FSP), length of panicle (LP), panicle density (PD), 1000-grain weight (TGW), seed setting rate (SR), grain weight per plant(GWP), leaf relative water content (LRWC), and level for rolling leaf (LRL). GWP is the most sensitive trait that is influenced by drought. Single variable stepwise regression, path analysis, gray correlative analysis, and multiple partial correlations analysis were conducted between the relative value of various traits and drought resistance coefficient. The results showed that six traits including LRWC, TNP, ETNP, TGW, PH, and ETNP significantly influenced the drought resistance and could be the indices to identify the drought resistance in rice. The D value of six traits elected was comprehensively used to evaluate the drought resistance by subordinate function values analysis, the results showed six index traits were feasible to predict the drought resistance in rice for whole growth stage. Four different drought resistance evaluated methods were compared, which indicated the drought resistance index is the best one for drought resistance direct evaluating.

利用组合R974//东野/R974 BC₁F₅ BIL群体，设水分胁迫和非胁迫两种处理，在全生育期测定了株高、单株分蘖数、单株有效穗数、穗长、穗实粒数、穗总粒数、着粒密度、千粒重、结实率、单株产量、叶片相对含水量、卷叶级别等12个与抗旱相关的形态及生理性状，其中单株产量对水分胁迫影响最敏感。通过各性状旱、水相对值对抗旱系数分别进行单因素逐步回归、通径、灰色关联分析，结果表明，叶片相对含水量、单株分蘖数、穗实粒数、千粒重、株高、单株有效穗数等6个性状与水稻抗旱性相关显著，可作为全生育期抗旱鉴定指标。利用模糊隶属函数法对入选性状的抗旱D值进行抗旱性综合评价，更表明上述性状作为水稻全生育期的抗旱鉴定指标是可行的。同时对5种抗旱性评价方法进行比较指出，抗旱指数是最合适的抗旱性直接评价方法。

... 指标, 得到了一致认可^[13,14] ...

... 1 抗旱性参照胡标林等^[13]方法计算抗旱系数(drought resistance coefficient, DC)和抗旱指数(drought resistance index, DI), DC = Y_d/Y_p, DI= (Y_d / Y_p) #cod#x000d7 ...

... 作物首先感受土壤干旱的器官是根系, 根系特性与抗旱性有密切的关系, 已成为研究和改良作物抗旱性的一个重要组成部分^{[13,14,15,16]} ...

1995

0.0

1.889

. 1995, 28(5):65-72

Drought resistance and its physiological characteristics in rice varieties

(水稻品种的抗旱性及其生理特性的研究) . (中国农业科学) , 1995 , 28 ( 5 ): 65 - 72 Yang J C , Wang Z Q , Zhu Q S . Drought resistance and its physiological characteristics in rice varieties

Yang J C

(杨建昌) , (王志琴) , (朱庆森) . (水稻品种的抗旱性及其生理特性的研究) . (中国农业科学) , 1995 , 28 ( 5 ): 65 - 72 Yang J C

江苏农学院，扬州 225009

研究了10个中熟水、陆稻品种（含杂交组合）的抗旱性以及对水分胁迫的生理反应。结果表明：不同品种的抗旱系数为：旱-A03＞旱-501＞珍珠矮＞扬稻4号＞汕优63＞六优1号＞亚优2号＞南京11＞盐粳2号＞盐粳235，而抗旱指数和在水分胁迫下的产量则以3个杂交组合汕优63、亚优2号和六优1号最高；在水分胁迫下，抗旱性较好（抗旱系数较高）的品种SOD（超氧化物歧化酶）活性明显增加的叶水势阈值较高，叶片质膜透性明显增加的叶水势阈值较低，抗旱性较差（抗旱系数较低）的品种则相反；水分胁迫下库源比、茎鞘物质的输出率和转换率的相对比率（胁迫处理/对照×100%），抗旱性较好的品种较高，抗旱性较差的品种较低；在水分胁迫下剑叶的气孔密度增加，但抗旱性较好的品种增加较少，抗旱性较差的品种增加较多。

... 指标, 得到了一致认可^[13,14] ...

... 作物首先感受土壤干旱的器官是根系, 根系特性与抗旱性有密切的关系, 已成为研究和改良作物抗旱性的一个重要组成部分^{[13,14,15,16]} ...

2005

0.0

1.667

. 2005, 31:203-208

Relationship between maize root hydraulic conductivity and drought resistance

(玉米整株根系水导与其表型抗旱性的关系研究) . (作物学报) , 2005 , 31 : 203 - 208 Mu Z X , Zhang S Q , Liang A H , Liang Z S . Relationship between maize root hydraulic conductivity and drought resistance

Mu Z X

(慕自新) , (张岁岐) , (梁爱华) , (梁宗锁) . (玉米整株根系水导与其表型抗旱性的关系研究) . (作物学报) , 2005 , 31 : 203 - 208 Mu Z X

Northwest Sci-Tech University of Agriculture and Forestry，Yangling 712100, Shaanxi

Solution culture method was used to study the relationship between maize whole root hydraulic conductivity（Lpwr） and drought resistance. The results indicated that for all the three maize genotypes, there was a significant positive correlation between Lpwr and leafψs, Pn, Tr and Cs, which meant w

以一组遗传背景清楚，表型性状已知的玉米遗传材料杂交种户单4号（F1代）及其双亲（♂，♀）为供试材料，用PEG-6000模拟干旱胁迫（ψs =－0.2 MPa），在室内溶液培养条件下研究了玉米整株根系水导（Lpwr）与其表型抗旱性间的关系，结果表明玉米整株根系水导与其品种抗旱性间具有显著的正相关（P<0.05）。在干旱胁迫下，玉米

... 根系不但是水分吸收的主要器官, 而且通过释放干旱信号调节气孔开度, 影响水分的散失, 与作物抗旱性关系十分密切^[15], 不同耐旱品种根系形态特征与生理活性存在显著差异^[16,17,18] ...

... 以根系总吸收面积与干重的比值计算根系比吸收面积^[15] ...

... 作物首先感受土壤干旱的器官是根系, 根系特性与抗旱性有密切的关系, 已成为研究和改良作物抗旱性的一个重要组成部分^{[13,14,15,16]} ...

1997

0.0

1.742

. 1997, 8:134-138

Relationship between root system characteristics and drought resistance of wheat populations in semiarid region

(半干旱地区小麦群体的根系特征与抗旱性的关系) . (应用生态学报) , 1997 , 8 : 134 - 138 Duan S S , Gu W X , Zhang D Y , Li F M . Relationship between root system characteristics and drought resistance of wheat populations in semiarid region

Duan S S

(段舜山) , (谷文祥) , (张大勇) , (李凤民) . (半干旱地区小麦群体的根系特征与抗旱性的关系) . (应用生态学报) , 1997 , 8 : 134 - 138 Duan S S

1. South China Agricultural University, Guangzhou 510642; 2. The State Key Laboratory of Arid Agroecology, Lazhou University, Lanzhou 730000

Afield experiment with five spring wheat (Triticum aestivum) varieties 8275(A),79157(B),8832(C),8624(D) and DX24(E) which have different drought resistance shows that the average root density in 0～120 cm soil layer is 66 μg·cm^-3 for A,104 μg·cm^-3 for B,137 μg·cm^-3 for C,102 μg·cm^-3 for Dand 99 μg·cm^-3 E, and their total root weight is 78.7,124.3,164.3,121.8 and 118.7 g·m^-2 ,respectively.The grain yield of these five varieties are respectively 201,207,141,218 and 190 g·m^-2 .When water is a primary limiting factor, the drought resistance of wheat is worse for those with large root systems,and an intense competition for water is existed among individuals;but is better for those with small ones, and no competition for water is existed.

对抗旱性能不同的5个春小麦品种(分别为A、B、C、D、E)的试验表明,在0～120cm土层中,其根密度分别为66、104、137、102和99μg·cm^-3;总根量分别为78.7、124.3、164.3、121.8和118.7g·m^-2;产量分别为201、207、141、218和190g·m^-2.在水分为主导因子的旱作条件下,根系过大的品种,个体间竞争激烈,其群体的抗旱性较差;根系较小的品种,个体间竞争较弱或无竞争,故群体抗旱性较强.

... 作物首先感受土壤干旱的器官是根系, 根系特性与抗旱性有密切的关系, 已成为研究和改良作物抗旱性的一个重要组成部分^{[13,14,15,16]} ...

... 大量的研究认为根系干重、根系体积、根系吸收面积等形态指标与品种抗旱性密切相关^[16,25,26] ...

2006

2.474

0.0

2011

2.474

0.0

2002

0.0

... 2 根系农艺性状取整株根系, 以排水法测定根系体积, 用吸水纸小心吸干根系表面水分, 以甲烯蓝吸附法测定根系总吸收面积和活跃吸收面积^[19] ...

1983

0.0

... 参照王爱国等^[20]的方法测定超氧化物歧化酶(superoxide dismutase, SOD)活性, 利用SOD抑制氮蓝四唑(NBT)在荧光下的还原作用, 酶活性以抑制NBT还原50%为一个酶活性单位 ...

1993

6.555

0.0

... 参照Wakamatsu等^[21]的方法测定过氧化氢酶(catalase, CAT)活性, 以每分钟减少0 ...

1984

0.599

0.0

... 参照林植芳等^[22]的方法测定丙二醛(malondialdehyde, MDA)含量 ...

2006

0.0

... 参照陈建勋等^[23]的方法测定抗坏血酸(ascorbic acid, AsA)和还原型谷胱甘肽(glutathione, GSH)含量 ...

1987

0.0

... 本研究分别以花生苗期干旱处理的生物量抗旱系数和结荚期干旱处理的产量抗旱系数评价12个花生品种的抗旱性, 发现苗期重度干旱胁迫下品种的抗旱性与结荚期抗旱性基本一致, 相关极显著(P[24]的标准, 把12个花生品种的抗旱性划分为强、中、弱3级 ...

2006

0.0

1.667

. 2006, 32:811-816

Effects of manure on root and shoot growth of winter wheat under water stress

(水分胁迫条件下有机肥对小麦根苗生长的影响) . (作物学报) , 2006 , 32 : 811 - 816 Zhang Y Q , Miao G Y . Effects of manure on root and shoot growth of winter wheat under water stress

Zhang Y Q

(张永清) , (苗果园) . (水分胁迫条件下有机肥对小麦根苗生长的影响) . (作物学报) , 2006 , 32 : 811 - 816 Zhang Y Q

(1 Collage of Life Science, Shanxi Teachers University, Linfen 041004, Shanxi; 2 College of Agriculture, Shanxi Agricultural University, Taigu 030801,Shanxi, China)

Corp root, an important organ for absorbing water and nutrients and synthesizing growth regulator, has a direct influence on the crop growth and yield. Roots play a remarkable role in dryland agriculture under serious soil water restriction. However, the research on crop root system is not so thorough as that on aerial parts of crop for lack of advanced root treating techniques, suggesting that the study on roots is very important for the further improvement of the yield. The pot culture experiment was conducted on the Experimental Farm of Shanxi Agricultural University to determine if supplemental irrigation and/or application of soil amendments rich in organic matter could improve root growth of winter wheat (Triticum aestivum L.). Root length, root weight, root activity, adventitious root number, root total and active absorbing areas, root SOD and POD activities, root MDA content and root-to-shoot ratio of winter wheat were studied under three levels of soil moisture (13%, 16%, 20%) and three levels of fertilization (0,333,and 666 g cattle manure per pot were measured) during 2003–2004. The results indicated that the content of MDA and the ratio of root to canopy were increased, but the dry weight of root, total root length, adventitious root number, root activity, the SOD and POD activities of root, the total and active absorbing areas decreased significantly with drought treatment regardless of fertilizing or not; root growth, root physiological characteristics, seedling growth, the water use efficiency and the yield as well as drought resistance were improved (Table 3); the highest yield of wheat was found under F₃(666 g/kg) and W3(soil water capacity 20%) treatment;. the activity of soil enzymes (Fig.2), especially in the rhizosphere could be increased by manure application, resulting in the improvement of soil nutrients, especially in the rhizosphere and enhancing the nutrients absorption and growth in wheat.

水分胁迫条件下施用有机肥对小麦根苗生长的影响研究表明：随着水分胁迫程度的加剧，无论施肥与否，小麦根系均表现出根重下降、根长缩短、根活力降低、次生根数与根系吸收面积减小、根系SOD、POD活性明显降低、MDA含量和根冠比明显增高的趋势。但施有机肥促进小麦根系的生长，改善根系生理特性，增加根系吸收面积和活力，具以肥促根、以肥调水、延缓根系衰老的作用。有机肥和土壤水分对小麦产量的增加具有明显的正交互作用，特别是在小麦生长中心发生转换的关键时期，即拔节期，水肥的显著互作促进根系的生长，在一定程度上缓解干旱胁迫的影响，最终导致土壤水分利用率的提高和产量的增加。

... 大量的研究认为根系干重、根系体积、根系吸收面积等形态指标与品种抗旱性密切相关^[16,25,26] ...

2005

0.0

. 2005, 24:176-182

Identification and analysis of drought tolerance and root traits of seedlings in soybeans

(大豆苗期耐旱性与根系性状的鉴定和分析) . (大豆科学) , 2005 , 24 : 176 - 182 Yang S P , Chen J M , Liu Y , Yu D Y , Gai J Y . Identification and analysis of drought tolerance and root traits of seedlings in soybeans

Yang S P

(杨守萍) , (陈加敏) , (刘莹) , (喻德跃) , (盖钧镒) . (大豆苗期耐旱性与根系性状的鉴定和分析) . (大豆科学) , 2005 , 24 : 176 - 182 Yang S P

2003年对63份大豆品种在盆栽自然干旱和盆栽水分控制两种条件下进行苗期耐旱性鉴定,两者鉴定结果间的关联分析表明两者呈极显著相关,两者间的鉴定分级结果比较一致,筛选出一批苗期耐旱型品种.综合分析2001～2003年的大豆苗期耐旱性鉴定结果,筛选出稳定耐旱型(1级)品种4份(晋豆14、汉中八月黄、科丰1号和因黑豆)和稳定干旱敏感型(5级)品种1份(宁海晚黄豆).苗期耐旱隶属函数值与根系性状间的相关分析表明,比根长、比根干重、比根表面积和比根体积等根系性状相对值与耐旱隶属函数值呈极显著相关,根总长、侧根总长、侧根数等根系性状绝对值与耐旱隶属函数值间的相关不显著,说明利用根系性状绝对值进行苗期耐旱性的评价较为牵强,而利用根系性状相对值进行耐旱品种的鉴定筛选比较可靠.

... 大量的研究认为根系干重、根系体积、根系吸收面积等形态指标与品种抗旱性密切相关^[16,25,26] ...

... 杨守萍等^[26]研究认为应以干旱胁迫下根系性状的相对值鉴定品种的抗旱性, 刘莹等^[29]的研究表明干旱胁迫下根系性状占整株的比重可作为衡量大豆抗旱性的指标 ...

1995

0.0

... 根系总吸收面积与活跃吸收面积是衡量根系吸收功能和活力的重要指标, 干旱胁迫下较大的根系吸收面积能提高作物吸水效率, 有利于吸收尽可能多的水分, 提高抗旱性^[27,28] ...

2005

0.0

2005

0.0

1.667

. 2005, 31:1132-1137

Identification of rhizosphere abiotic stress tolerance and related root traits in soybean [Glycine max (L.) Merr.]

(大豆根区逆境耐性的种质鉴定及其与根系性状的关系) . (作物学报) , 2005 , 31 : 1132 - 1137 Liu Y , Gai J Y , L#cod#x000fc; H N . Identification of rhizosphere abiotic stress tolerance and related root traits in soybean [Glycine max (L.) Merr.]

Liu Y

(刘莹) , (盖钧镒) , (吕慧能) . (大豆根区逆境耐性的种质鉴定及其与根系性状的关系) . (作物学报) , 2005 , 31 : 1132 - 1137 Liu Y

Soybean Research Institute of Nanjing Agricultural University, National Center for Soybean Improvement/ National Key Laboratory of Crop Genetics and Germplasm Enhancement, Nanjing 210095, Jiangsu

Sixty two accessions of soybean [Glycine max (L.) Merr.] selected from 301 representative entries from Huang-Huai-Hai and Middle-Lower Changjiang Valleys according to their root performance were used to identify their tolerance to rhizosphere-abiotic stresses, including drought, aluminum toxin and l

依根系类型从黄淮海和长江中下游地区301份代表性材料中选取62份，以株高、叶龄、地上部干物重、地下部干物重为指标，采用平均隶属函数值方法鉴定了苗期耐旱性、苗期耐铝毒性，加上主茎节数、分枝数、单株荚数、单株粒数、百粒重等性状鉴定了后期耐旱性，并通过钒钼黄比色法测定植株P含量鉴定了苗期耐低磷性。筛选出1级苗期

2004

6.555

0.0

2010

0.0

1.742

. 2010, 21:48-52

Effects of drought stress on the grain yield and root physiological traits of maize varieties with different drought tolerance

(干旱胁迫对不同耐旱性玉米杂交种产量和根系生理特性的影响) . (应用生态学报) , 2010 , 21 : 48 - 52 Qi W , Zhang J W , Wang K J , Liu P , Dong S T . Effects of drought stress on the grain yield and root physiological traits of maize varieties with different drought tolerance

Qi W

(齐伟) , (张吉旺) , (王空军) , (刘鹏) , (董树亭) . (干旱胁迫对不同耐旱性玉米杂交种产量和根系生理特性的影响) . (应用生态学报) , 2010 , 21 : 48 - 52 Qi W

State Key Laboratory of Crop Biology, College of Agronomy, Shandong Agricultural University,Tai’an 271018, Shandong, China

A pot experiment was conducted to study the effects of simulated moderate drought in whole growth period on the grain yield and root physiological traits of drought-tolerant maize variety JK 628 and drought-sensitive maize variety ND-95. Compared with the control, the grain yield of JK-628 and ND-95 under the stress of moderate drought was decreased by 33.7% and 62.3%, respectively. Under drought stress, the root biomass of the two varieties decreased, with the maximum value appeared ahead. The root/shoot ratio of JK-628 increased at early growth stage but decreased at late growth stage, while that of ND-95 increased all along. ND-95 had a larger decrement of root activity than JK-628. The root SOD activity in drought stress plants was higher in prophase but lower in anaphase, compared with that in the control, and the time of SOD activity begun to lower was later for JK-628 than for ND-95. The root MDA content in drought-stress plants was increased, and the increment was much higher for ND-95 than for JK-628. In addition, the root soluble protein content in drought-stress plants was decreased, and ND-95 had a larger decrement than JK-628. In a word, under drought stress, the root activity, SOD activity, and soluble protein content of JK-628 were higher than those of ND-95, which could slow down the root aging process and prolong the root functional period, possibly being one of the important factors of getting high yield of drought-tolerant maize under drought stress.

通过盆栽人工模拟干旱试验，研究了全生育期中度干旱胁迫对不同耐旱性玉米杂交种（耐旱：京科628；不耐旱：农大95）产量及根系生理特性的影响.结果表明：干旱胁迫下，耐旱玉米产量比对照减少33.7%，不耐旱玉米则比对照减少62.3%.干旱胁迫下，玉米根系生物量降低且最大值出现时间提前，与对照相比，不耐旱玉米根冠比升高，耐旱玉米根冠比前期升高后期降低；根系活力降低，不耐旱玉米根系活力降低幅度大于耐旱玉米；根系超氧化物歧化酶（SOD）活性前期高于对照后期低于对照，耐旱玉米根系SOD活性开始低于对照的时间比不耐旱玉米晚；根系丙二醛（MDA）含量升高，随干旱胁迫处理时间的延长，不耐旱玉米MDA含量比对照升高的幅度大于耐旱玉米；根系可溶性蛋白含量降低，不耐旱玉米的降低幅度大于耐旱玉米.干旱胁迫下耐旱玉米杂交种根系活力﹑根系SOD活性及可溶性蛋白含量较高，减缓了根系的衰老进程，延长了根系功能期，这可能是耐旱玉米杂交种在干旱胁迫下仍能获得较高产量的重要原因之一.

... 齐伟等^[31]研究认为, 根系SOD、CAT、POD活性在水分胁迫下与品种的抗旱性密切相关, 抗旱性强的品种能维持较高的SOD、CAT活性 ...

2010

0.0

. 2010, 30:357-361

Growth and physiological characteristics of roots in different flue-cured tobacco varieties under drought stress

(干旱胁迫对不同烤烟品种根系生长和生理特性的影响) . (西北植物学报) , 2010 , 30 : 357 - 361 Shang X Y , Liu H B , Zhang X Q , Lin J , Duan W J , Yang T Z . Growth and physiological characteristics of roots in different flue-cured tobacco varieties under drought stress

Shang X Y

(尚晓颍) , (刘化冰) , (张小全) , (林娟) , (段旺军) , (杨铁钊) . (干旱胁迫对不同烤烟品种根系生长和生理特性的影响) . (西北植物学报) , 2010 , 30 : 357 - 361 Shang X Y

以抗旱型烤烟品种‘农大202'及一般型烤烟品种‘NC89'和‘K326'为材料,采用盆栽试验研究了干旱胁迫对根系生长和生理特性的影响,以明确各烟草品种的抗旱能力及其与根系生长和生理特性的关系.结果显示:严重干旱胁迫之后各烤烟品种根系鲜重、干重上升,而根系活力、根系吸收面积以及根系SOD和POD活性等根系生理指标则呈下降趋势.在干旱胁迫条件下,‘农大202'的根系总吸收面积、活跃吸收面积、根系活力、SOD活性和POD活性均显著高于两对照品种‘NC89'和‘K326',但其根系鲜重和干重等生物量并不具有太大的优势.研究表明,烟草的根系生长和生理特性对环境水分条件的响应存在明显的基因型差异;在干旱胁迫条件下, ‘农大202'根系能保持较高总吸收面积、活跃吸收面积、根系活力和保护酶活性,是其具有较强抗旱性的生理基础.

Abstract：

The experiment were carried out to study the effects of drought stress on growth and physiological characteristics of roots among three stress,the root biomass including root fresh weight and root dry weight increased,whereas,the root activity,root absorbing area,SOD activity and appeared significantly greater root total absorbing area,active absorbing area,SOD activity and POD activity,but the root biomass including fresh weight and dry weight did not have significant difference.In conclusion,there are obvious differences among flue-cured tobacco genotypes in root growth and physiological characteristics,and their activity and POD activity,more root total and active absorbing area under seriously drought stress,which were the physiological basis of stronger drought resistance.

... 尚晓颍等^[32]研究认为根系SOD、CAT活性可以用来鉴别干旱胁迫下品种的抗旱性 ...

2009

0.0

. 2009, (6):36-38

Effect of drought on root physiological and biochemical characters of different maize cultivars during seedling stage

(干旱对不同玉米品种苗期根系生理生化特性的影响) . (山东农业科学) , 2009 , ( 6 ): 36 - 38 Wang S S , Zhang J W , Liu P , Dong S T , Wang K J . Effect of drought on root physiological and biochemical characters of different maize cultivars during seedling stage

Wang S S

(王思思) , (张吉旺) , (刘鹏) , (董树亭) , (王空军) . (干旱对不同玉米品种苗期根系生理生化特性的影响) . (山东农业科学) , 2009 , ( 6 ): 36 - 38 Wang S S

以抗旱性不同的23个玉米品种为试材,用20%PEG溶液对幼苗进行模拟水分胁迫,研究了干旱胁迫下苗期根系生理生化特性的变化及其与品种抗旱性的关系.结果表明:苗期干旱导致根冠比和根系活力呈升高趋势,耐旱性品种升高幅度最大.苗期干旱,根系超氧化物歧化酶(SOD)活性升高,且耐旱性品种升高幅度大于中等耐旱和不耐旱品种;根系中MDA含量均增加,且耐旱玉米品种丙二醛(MDA)含量增量最小.本研究认为耐旱型玉米品种苗期根系在干旱胁迫下优势的内在机制是根系活力和清除活性氧的能力较强.

... 本研究还发现, 对照与重度干旱处理的根系SOD活性和MDA含量呈极显著正相关, 不同品种间正常水分时根系的抗氧化能力能反映干旱胁迫下的趋势, 王思思等^[33]在玉米研究中也得出相似结论 ...

2001

0.0

1.667

. 2001, 27:286-292

Studies on the classification of maturity groups of soybeans in China

(中国大豆品种熟期组划分的研究) . (作物学报) , 2001 , 27 : 286 - 292 Gai J Y , Wang Y S , Zhang M C , Wang J A , Chang R Z . Studies on the classification of maturity groups of soybeans in China

Gai J Y

(盖钧镒) , (汪越胜) , (张孟臣) , (王继安) , (常汝镇) . (中国大豆品种熟期组划分的研究) . (作物学报) , 2001 , 27 : 286 - 292 Gai J Y

根据北美13个熟期组大豆代表品种48份及我国各地地方品种256份在南京春播自然条件结合18小时长光照条件的试验，部分品种在石家庄、哈尔滨的春播试验，获得以下主要结果：(1)将我国大豆品种归属为相应的000～Ⅸ共12个熟期组，未发现熟期组Ⅹ品种；(2)按同一熟期组品种生育前期变异的地理分布，0～Ⅲ熟期组内各划分为秦岭淮河

... 干旱的持续时间及胁迫强度影响各抗旱性状的测定值, 直接以测定数值的大小划分抗旱性状的等级难免出现误差, 盖钧镒等^[34]提出了大豆品种生育期组鉴定的标准品种, 依据标准品种作为参考来划分品种的生育期可以排除环境因素的影响 ...

2004

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... 陈加敏^[35]在研究大豆品种抗旱性时, 根据抗旱性强弱分别筛选出了抗旱标准品种与干旱敏感标准品种 ...