不同高粱品种萌发期抗旱性筛选与鉴定

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吴奇, 周宇飞, 高悦, 张姣, 陈冰嬬, 许文娟, 黄瑞冬. . 不同高粱品种萌发期抗旱性筛选与鉴定. 作物学报, 2016, 42(8): 1233-1246
WU Qi, ZHOU Yu-Fei, GAO Yue, ZHANG Jiao, CHEN Bing-Ru, XU Wen-Juan, HUANG Rui-Dong. . Screening and Identification for Drought Resistance during Germination in Sorghum Cultivars. ACTA AGRONOMICA SINICA, 2016, 42(8): 1233-1246 复制到剪切板

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不同高粱品种萌发期抗旱性筛选与鉴定

吴奇¹, 周宇飞^1,^*, 高悦¹, 张姣¹, 陈冰嬬^1,², 许文娟¹, 黄瑞冬^1,^*

¹沈阳农业大学农学院, 辽宁沈阳110866

²吉林省农业科学院作物资源研究所, 吉林公主岭130124

^*通讯作者(Corresponding authors): 周宇飞, E-mail: zhouyufei2002@aliyun.com; 黄瑞冬, E-mail: r_huang@126.com, Tel: 024-88487135

第一作者联系方式：E-mail:597912452@qq.com, Tel: 18842343657

收稿日期:2015-12-10 接受日期:2016-05-09 网络出版日期:2016-05-13

基金:本研究由国家现代农业产业技术体系建设专项(CARS-06-02-02),辽宁省教育厅一般项目(L2014271)和沈阳农业大学博士后基金项目(139173)资助

摘要

萌发期是高粱表现抗旱性的关键阶段之一。选用54份高粱品种(组合)为供试材料, 采用人工气候箱培养, 以水势为-0.4 MPa的PEG-6000水溶液模拟干旱胁迫环境, 萌发10 d后测定发芽势、发芽率、萌发抗旱指数等8个萌发性状。基于主成分分析的综合因子得分函数能够对萌发期高粱品种的抗旱性综合评定, 各品种(组合)综合因子得分( Y值)经聚类分析后, 将54份高粱品种(组合)按萌发期抗旱性的强弱分为4大类, 13218A×20982R是一个抗旱性极强的杂交组合, 吉杂133等13个品种(组合)为抗旱品种(组合), 苏马克等23个品种为干旱敏感品种, 辽杂35等17个品种为高度干旱敏感品种。萌发抗旱指数、发芽率和根长等性状可作为高粱品种萌发期抗旱性鉴定的主要指标。

关键词: 高粱; 萌发期; 抗旱性; 主成分分析; 聚类分析

Screening and Identification for Drought Resistance during Germination in Sorghum Cultivars

WU Qi¹, ZHOU Yu-Fei^1,^*, GAO Yue¹, ZHANG Jiao¹, CHEN Bing-Ru^1,², XU Wen-Juan¹, HUANG Rui-Dong^1,^*

¹College of Agronomy, Shenyang Agricultural University, Shenyang 110866, China

² Institute of Crop Resources, Jilin Academy of Agricultural Sciences, Gongzhuling 130124, China

Fund:This study was supported by the China Agriculture Research System (CARS-06-02-02), the General Fund of Department of Education of Liaoning Province (L2014271), and the Postdoctoral Fund of Shenyang Agricultural University (139173)

Abstract

Germination is a key stage showing drought resistance in sorghum. The present study was conducted to investigate the responses of 54 different sorghum cultivars (hybrid combinations) to drought stress during germination and screen sorghum cultivars suitable for growing in arid environment. PEG-6000 (-0.4 MPa) was used to simulate the typical drought stress environment in artificial climate chamber. Eight parameters including germination potential, germination rate, germination drought resistance index were measured. The results indicated that the comprehensive factor score functions were suitable for the comprehensive assessment of drought resistance during germination of sorghum cultivars based on principal component analysis (PCA). Based on comprehensive factor scores ( Y-value), 54 sorghum cultivars (hybridized combinations) were clustered into four groups, in which 13218A×20982R was the highest drought resistant hybrid, 13 cultivars (hybridized combinations) were drought resistant, e.g. Jiza 133, 23 cultivars were drought sensitive, e.g. Sumac, and 17 cultivars were highly drought sensitive, e.g. Liaoza 35. Results of PCA showed that germination drought resistance index, germination rate and root length can be used to screen a large number of sorghum cultivars with drought resistance.

Keyword: Sorghum; Germination; Drought resistance; Principal component analysis; Cluster analysis

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近年来, 随着全球气候的变化水资源供需矛盾加剧, 干旱已经成为农业中常见的问题。其对粮食产量、品质和效益的影响在将来亦有增加的趋势^[1]。因此, 提高作物的抗旱性是保证粮食高产、稳产的必要措施。高粱[Sorghum bicolor (L.) Moench]是我国最早栽培的禾谷类作物之一, 在干旱和半干旱地区分布较广, 具有食用、酿造和饲用等多种使用价值^{[2, 3]}。与其他作物相比, 虽然高粱具有较强的抗旱能力^{[4, 5, 6]}, 但其生长受干旱影响较大。在干旱环境中, 选择抗旱性更高的作物品种是创建优良栽培群体的重要途径^{[7, 8]}, 鉴于此, 抗旱性已成为高粱逆境生理研究的重要内容之一。

干旱条件下, 作物出苗情况的优劣依赖于种子的发芽速度及整齐度, 这与种子在低水分条件下的发芽能力密切相关^{[9, 10]}。由于北方地区春季多风少雨, 种子在萌发阶段极易受到干旱的干扰, 为此, 许多学者都很注重萌发期抗旱性的研究。裴帅帅等^[11]认为用单一的性状或生理指标不能对萌发期的抗旱机制进行综合评价。陈新等^[12]研究表明, 利用主成分分析的方法能消除各鉴定指标间信息重叠的干扰, 可有效评价裸燕麦萌发期的抗旱性。这种综合分析抗逆性的方法在高粱的耐盐碱研究中已得到应用^{[13, 14]}。近年来, 已广泛使用PEG溶液模拟干旱环境进行萌发试验^{[15, 16]}。种子的发芽率、发芽势、萌发抗旱指数是玉米^[17]、小麦^[18]、水稻^[19]等作物萌发期抗旱性评定的常用指标, 具有一定的可靠性, 也可以运用到高粱萌发期抗旱性的鉴定中, 但是不同作物萌发期抗旱性的主要鉴定指标仍存在一定差异^{[12, 17, 20, 21, 22]}。萌发期是作物生长的关键时期, 对作物群体数量和结构具有重要的决定作用。然而, 到目前为止, 基于不同基因型试材、综合鉴定高粱萌发期抗旱性的研究并不多见。对于高粱萌发期抗旱能力有针对性和系统性的研究, 必将有助于促进高粱抗旱机制的了解。本试验在人工气候条件下模拟干旱处理, 考察具有不同遗传背景的高粱品种(组合)在萌发期对干旱胁迫的响应, 以期筛选和鉴定出抗旱能力更强的种质资源, 并为高粱抗旱新品种选育提供理论依据。

1 材料与方法

1.1 供试材料

选用高粱材料共54份, 来自全国9家高粱育种单位。具体品种名称、保存单位及父母本等信息见附表1。

1.2 试验设计

2014年在沈阳农业大学高粱生理实验室进行试验。预备试验, 随机选取每个高粱品种(组合) 25粒饱满、大小一致的种子, 经7%的NaClO溶液(有效氯含量≥ 5.2%, 国药集团化学试剂有限公司生产)消毒10 min, 蒸馏水冲洗5次, 用滤纸吸干, 放入直径15 cm培养皿(铺有双层滤纸)。于RXZ-1000B型人工气候箱培养, 箱内光照/黑暗为12 h/12 h, 温度为28℃/25℃, 湿度为60%, 光照强度为134 μ mol m^-2 s^-1, 连续培养10 d, 各处理3次重复。以PEG-6000水溶液模拟干旱环境, 设置4个干旱胁迫水平, PEG-6000质量浓度分别为80、120、150和175 g L^-1(对应水势分别为-0.1、-0.2、-0.3和-0.4 MPa), 以蒸馏水作为对照, 每个培养皿中加入8 mL胁迫液或蒸馏水。从种子培养之日起作为起始期(0 d), 每隔2 d补充适量蒸馏水, 且在2、4、6、8和10 d调查种子发芽数。通过预备试验观察到在水势为-0.4 MPa时, 不同高粱品种(组合)间及处理与对照间的发芽性状存在较大差异, 所以选择175 g L^-1的PEG-6000水溶液作为正式试验的干旱胁迫浓度。

正式试验的人工培养条件和调查过程与预备试验相同, 每品种(组合)各处理条件下3次重复。在试验结束时, 从各处理随机取高粱幼苗5株, 测定芽长、根长、芽干重、根干重、籽粒干重, 计算干物质转运率^[23]。为消除试验材料间的差异, 采用各项测定指标的相对值来反映对干旱的响应。

发芽势(GP) = 4 d发芽种子数/供试种子数× 100%, 相对发芽势 = (处理发芽势/对照发芽势)× 100%; 发芽率(GR) = 10 d发芽种子数/供试种子数× 100%。

相对发芽率 = (处理发芽率/对照发芽率)× 100%; 种子萌发抗旱指数(DGRI) = 干旱种子萌发指数/对照种子萌发指数; 萌发指数 = 1.00nd₂+ 0.75nd₄+ 0.50nd₆+ 0.25nd₈, 其中, nd₂、nd₄、nd₆、nd₈分别为2、4、6、8 d的种子发芽率。

干物质转运率 = (芽干重+根干重)/ (芽干重+根干重+籽粒干重)× 100%, 相对干物质转运率 = (处理干物质转运率/对照干物质转运率)× 100%。

1.3 数据处理与分析

应用Microsoft Excel 2007整理数据, SPSS18.0进行方差分析、相关分析及聚类分析。

2 结果与分析

2.1 干旱胁迫对不同高粱品种萌发的影响

在水势为-0.4 MPa的干旱胁迫下, 不同高粱品种的萌发指标表现出不同程度的下降(表1)。发芽势相对值变化范围为0.33~1.35, 发芽率相对值在0.69~1.58之间, 萌发指数的相对值在0.47~1.22之间, 芽长的相对值在0.39~1.01之间, 根长的相对值在0.42~1.42之间, 芽干重的相对值在0.44~1.18之间, 根干重的相对值在0.49~2.16之间, 干物质转运率的相对值为0.40~1.34。

表1 高粱品种萌发性状相对值 Table 1 Relative value of germination traits of sorghum under drought stress

编号 Code	X₁	X₂	X₃	X₄	X₅	X₆	X₇	X₈
1	1.06 ad	1.02 bc	1.05 af	0.74 cm	0.68 dj	0.89 bh	1.47 bd	1.01 bc
2	1.00 ad	1.01 bc	1.00 ag	0.73 cm	0.73 dj	1.05 ab	1.58 ac	1.02 b
3	0.96 ad	0.98 bc	0.99 ag	0.72 dm	0.81 ch	0.99 ae	1.35 bf	0.97 bd
4	0.87 ae	0.92 bc	0.92 ah	0.80 bl	0.91 bf	0.87 bi	1.12 bl	0.88 bf
5	1.02 ad	1.03 bc	1.05 af	0.84 ah	1.20 ab	0.78 bl	1.31 bh	0.91 be
6	1.04 ad	1.07 bc	1.07 ae	0.86 ae	0.77 di	0.86 bi	1.28 bi	0.93 bd
7	0.98 ad	1.10 bc	1.01 ag	0.89 ad	1.01 bd	0.80 bl	1.10 bl	0.83 dh
8	0.92 ad	0.98 bc	0.95 ag	0.72 dm	0.81 ch	0.79 bl	1.11 bl	1.34 a
9	0.95 ad	1.01 bc	0.96 ag	0.84 ah	0.74 dj	0.87 bi	1.06 bl	0.92 bd
10	0.97 ad	0.88 bc	0.90 ah	0.83 ai	0.81 ch	0.93 af	1.19 bk	0.96 bd
11	1.04 ad	1.04 bc	1.07 ae	0.78 bm	0.57 gj	0.86 bj	0.95 bl	0.80 dj
12	1.10 ad	1.03 bc	1.08 ae	0.76 cm	0.96 be	0.89 bh	1.30 bh	0.93 bd
13	0.99 ad	0.72 bc	1.04 ag	0.94 ac	0.69 dj	1.02 ac	1.03 bl	0.93 bd
14	0.91 ad	0.95 bc	0.93 ah	0.86 af	0.71 dj	0.79 bl	0.81 dl	0.80 dj
15	1.27 ac	1.00 bc	1.15 ac	0.86 ag	0.74 dj	0.88 bi	1.22 bk	0.94 bd
16	0.97 ad	0.92 bc	0.96 ag	0.85 ag	0.96 be	0.80 bl	1.38 be	0.85 dj
17	1.08 ad	0.90 bc	0.99 ag	1.01 a	1.42 a	1.18 a	2.12 a	0.97 bd
18	1.34 ab	1.20 abc	1.14 ad	0.87 ad	0.79 ch	0.85 bj	2.06 a	0.96 bd
19	1.35 a	1.58 a	1.22 a	0.88 ad	1.11 bc	0.93 af	2.16 a	0.95 bd
20	1.05 ad	1.05 bc	0.99 ag	0.80 bl	0.80 ch	0.84 bk	1.32 bg	0.90 be
21	1.00 ad	0.94 bc	0.96 ag	0.80 bl	0.82 ch	0.99 ae	1.46 bd	0.88 bf
22	1.06 ad	1.00 bc	0.92 ah	0.75 cm	0.67 ej	0.75 cl	0.99 bl	0.80 di
23	0.93 ad	1.05 bc	0.94 ah	0.66 eo	0.71 dj	0.66 fm	1.32 bh	0.83 dh
24	0.79 ae	0.92 bc	0.94 ah	0.85 ag	0.61 fj	1.00 ad	1.35 bf	0.88 bf
25	0.88 ae	0.96 bc	1.17 ab	0.98 ab	0.79 ch	0.93 af	1.60 ab	0.84 ch
26	1.06 ad	0.91 bc	0.97 ag	0.71 dn	0.44 ij	0.74 cm	1.09 bl	0.67 hm
27	0.95 ad	1.04 bc	0.99 ag	0.69 do	0.42 j	0.90 ag	1.26 bj	0.85 cg
28	0.94 ad	1.03 bc	0.96 ag	0.80 bl	0.50 hj	0.88 bi	1.32 bh	0.69 gl
29	0.89 ae	0.94 bc	0.91 ah	0.85 ag	0.57 gj	0.93 af	1.32 bh	0.89 be
30	0.88 ae	0.96 bc	0.90 ah	0.65 go	0.50 hj	0.73 cm	1.08 bl	0.85 bg
31	1.06 ad	1.00 bc	0.97 ag	0.82 aj	0.52 gj	1.01 ad	1.09 bl	0.84 ch
32	0.98 ad	0.97 bc	0.99 ag	0.77 cm	0.56 gj	0.79 bl	1.16 bk	0.71 fl
33	1.16 ad	1.23 ab	1.14 ad	0.81 bk	0.77 di	0.72 cm	0.98 bl	0.75 ek
34	1.02 ad	0.97 bc	0.97 ag	0.76 cm	0.85 cg	0.78 bl	1.35 bf	0.75 ek
35	0.94 ad	0.89 bc	0.75 ah	0.69 dn	0.54 gj	0.60 gm	0.58 kl	0.55 lp
36	0.72 ce	0.85 bc	0.57 fh	0.52 np	0.49 hj	0.54 km	0.66 gl	0.48 np
37	0.87 ae	0.79 bc	0.71 bh	0.69 do	0.50 hj	0.68 em	0.77 el	0.59 ko
38	0.71 ce	0.81 bc	0.56 gh	0.57 mo	0.53 gj	0.62 gm	0.75 el	0.60 ko
39	0.76 ae	0.84 bc	0.63 eh	0.60 lo	0.56 gj	0.59 hm	0.65 hl	0.50 np
40	0.89 ae	0.90 cb	0.69 bh	0.68 do	0.59 fj	0.66 fm	0.74 el	0.58 lo
41	0.76 be	0.78 bc	0.72 bh	0.58 mo	0.58 fj	0.58 im	0.75 el	0.48 np
42	0.70 ce	0.82 bc	0.71 bh	0.39 p	0.61 fj	0.44 m	0.86 dl	0.44 op
43	1.25 ac	1.20 abc	1.11 ae	0.69 do	0.63 ej	0.61 gm	0.77 el	0.52 mp
44	1.02 ad	1.09 bc	0.95 ag	0.49 op	0.63 ej	0.53 lm	0.49 l	0.44 op
45	0.83 ae	0.83 bc	0.67 dh	0.63 io	0.83 ch	0.77 bl	0.88 dl	0.64 jn
46	0.70 ce	0.87 bc	0.67 eh	0.61 ko	0.70 dj	0.63 fm	0.92 cl	0.50 np
47	0.90 ae	0.93 bc	0.83 ah	0.62 io	0.61 fj	0.56 jm	0.70 fl	0.40 p
48	0.93 ad	0.93 bc	0.88 ah	0.60 lo	0.67 ej	0.70 dm	0.88 dl	0.62 kn
49	0.65 de	0.91 bc	0.79 ah	0.65 fo	0.63 ej	0.75 cl	0.76 el	0.59 ko
50	0.74 ce	0.79 bc	0.69 bh	0.66 eo	0.85 cg	0.73 cm	0.60 jl	0.58 ko
51	0.33 e	0.69 c	0.47 h	0.64 ho	0.77 di	0.73 cm	0.85 dl	0.72 fl
52	0.80 ae	0.85 bc	0.70 bh	0.62 jo	0.80 ch	0.69 em	0.80 dl	0.59 ko
53	0.71 ce	0.89 bc	0.79 ah	0.63 io	0.67 ej	0.62 gm	0.73 el	0.65 in
54	0.62 de	0.94 bc	0.69 bh	0.60 lo	0.80 ch	0.58 im	0.63 il	0.48 np

X₁: relative germination potential; X₂: relative germination rate; X_3: relative promptness index; X₄: relative bud length; X₅: relative root length; X₆: relative bud dry weight; X₇: relative root dry weight; X₈: relative dry matter translocation rate.

X₁: 相对发芽势; X₂: 相对发芽率; X₃: 相对萌发抗旱指数; X₄: 相对芽长; X₅: 相对根长; X₆: 相对芽干重; X₇: 相对根干重; X₈: 对干物质转运率。

表1 高粱品种萌发性状相对值 Table 1 Relative value of germination traits of sorghum under drought stress

部分品种的发芽率、发芽势和萌发抗旱指数大于对照, 如杂交组合1335A× TY3560R、13218A× 20982R和吉杂99、吉杂124等11个品种。其中杂交组合13218A× 20982R的发芽率、发芽势和萌发抗旱指数的相对值显著高于其他品种, 对干旱条件不敏感, 而龙杂11的发芽率、发芽势和萌发抗旱指数的观测值下降幅度最大, 分别为0.33、0.69和0.47, 说明受干旱胁迫的影响较大。L470A× LZ1050R和13218A× 20982R在芽长、根长、芽干重几个指标上的相对值都比较大, 其余大部分品种这3个指标的相对值大多都处于0.65~0.85之间, 说明干旱胁迫使发芽迟缓、根系生长缓慢; 吉杂90、辽杂12和龙杂9号在这3个指标上的相对值都很低, 说明受到干旱胁迫的影响较大, 对干旱环境极为敏感; 大部分高粱品种根干重的相对值都大于1, 可见高粱根生物积累量受到干旱条件的激发反而呈上升趋势。种子萌发过程中的营养转化会使种子的重量逐渐减小, 干旱胁迫会降低营养物质转化的速率, 干物质转运率是一个能够反映种子营养物质转化速率与效率的指标, 白杂11的干物质转运率的相对值最高, 为1.34, 辽杂11的干物质转运率最低, 为0.40。

2.2 干旱胁迫下高粱萌发期各性状间的相关性

由表2可知, 大部分萌发性状间的相关性都已达到显著或极显著的水平。相对发芽势与相对萌发抗旱指数的相关系数为0.853, 相对芽长与相对芽干重的相关系数为0.815。相对芽干重和相对干物质转运率之间与相对根干重之间的关系也达到极显著水平, 分别为0.794和0.751。相对发芽率和相对发芽势之间与相对萌发抗旱指数之间的相关性也较高, 分别为0.746和0.732。

表2 干旱条件下高粱萌发期幼苗各生长指标的相关系数 Table 2 Correlation coefficients among germination traits of sorghum cultivars under drought stress

2.3 对高粱品种萌发期抗旱性的主成分分析

表3表明前3个主成分贡献率分别为62.04%、16.07%、9.20%, 累计贡献率达87.31%, 考虑到累计贡献率大于80%的原则, 说明这3个主成分已足以反映该数据的信息, 符合主成分分析的要求。

表3 3个主成分的特征根及贡献率 Table 3 Eigen value of three principal components and their contributions and cumulative contribution rates

第I主成分的特征根λ ₁= 4.96, 贡献率为62.04%, 在得分函数Y₁中第I主成分上X₃、X₇和X₄的系数较大, 说明在第I主成分上有较大的载荷, 反映了相对萌发抗旱指数、相对根干重和相对芽长的综合指标。第II主成分的特征根λ ₂= 1.29, 贡献率为16.07%, 在得分函数Y₂上X₂和X₁的系数较大, 分别为0.511和0.403, 说明在第II主成分上发芽率和发芽势具有较高的载荷, 反映了在干旱胁迫下各品种的萌发状况。第III主成分的贡献率为9.20%, 相对根长在第III主成分的载荷较大, 反映了干旱胁迫下根部的生长情况。

表4 各因子的载荷矩阵 Table 4 Loading matrix of each component

表5 因子得分系数矩阵 Table 5 Coefficient matrix of component score

根据主成分分析得出2个矩阵(表4和表5), 由此可得出因子得分函数公式。

Y₁= 0.157X₁+0.131X₂+0.178X₃+0.173X₄+0.112X₅+0.164X₆+0.176X₇+0.166X₈

Y₂= 0.403X₁+0.511X₂+0.261X₃-0.200X₄-0.253X₅-0.339X₆-0.105X₇-0.239X₈

Y₃= -0.071X₁+0.257X₂-0.165X₃-0.158X₄+1.020 X₅-0.355X₆+0.102X₇+0.240X₈

单一的Y₁、Y₂或Y₃并不能对高粱品种的抗旱性作出综合评价, 而由Y₁、Y₂和Y₃可以得到综合因子得分函数, 即Y = 0.71Y₁+0.18Y₂+0.11Y₃。

综合因子得分(Y)的结果能够对不同品种高粱抗旱能力作出综合性评价, Y值越大, 代表着该品种抗旱能力越强。由表6可知, 13218A× 20982R的Y值最大, 说明它在众多品种(组合)中抗旱性最好, 1335A× TY3560R和L407A× LZ1050R分别位居第2和第3, 抗旱能力较高; 锦杂93、辽杂12、龙杂11分别居后3位, 属于干旱敏感品种。

2.4 对不同高粱品种萌发期抗旱性的聚类分析

根据表6得出的Y值进行聚类分析, 利用系统分类的组内联接法, 根据欧式距离> 5的标准, 将54个高粱品种分为4大类(图1)。

第I类群只包含13218A× 20982R一个组合, 该组合的抗旱指标大于其他品种(组合), 位于抗旱能力排序的第一位, 是具有高抗旱能力的杂交组合。

第II类群包含吉杂133、晋杂102、川糯粱15、泸糯3号、吉杂305、白杂8号、意大利、HL8、L470A× 1181R、吉杂99、吉杂124、L470A× LZ1050R和1335A× TY3560R这13个品种(组合), 这些品种抗旱指标的相对值都比较大, 说明这些品种在萌发期对干旱胁迫有不敏感的特性, 属于抗旱品种。

表6 各品种综合因子得分Y值及抗旱性的排序 Table 6 Y-value of comprehensive component score and the ranks of drought-resistance of sorghum cultivars

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	图1 54个高粱品种聚类分析图Fig. 1 Dendrogram of 54 sorghum cultivars based on Y-value

第III类群包含苏马克、沈杂5号、特雷利、龙杂9号、赤杂24、敖杂1号、白杂11、吉杂123、晋杂18、川糯粱2号、泸糯12、罗马、丽欧、赤杂28、非洲白、吉杂127、贝利、凯勒、晋杂101、泸糯13、戴尔、泸州红1号、考利共23个品种, 这些品种的一些抗旱指标容易受到干旱条件的影响, 所以属于干旱敏感品种。

第IV类群的品种包括辽杂35、葫梁1号、龙杂12、辽杂21、锦杂105、龙杂13、辽杂10号、锦杂106、辽杂19、辽杂12、锦杂93、锦杂103、四杂29、吉杂90和龙杂11, 这些品种抗旱指标受干旱条件影响较大, 说明对干旱条件非常敏感, 属于高度敏感品种。

3 讨论

植物的抗旱性是一个复杂的综合性状, 在植物生长发育的各个阶段表现出抗旱机制也有所不同。萌发期是作物群体数量建成的关键时期, 作物在萌发期极易受到干旱环境的影响, 所以在萌发期筛选出具有抗旱能力的品种对于干旱地区农业生产具有重要意义^[24]。PEG作为一种理想的渗透调节剂, 在栽培基质中可以降低水势且能保持稳定的渗透压, 经常被用来模拟干旱环境进行发芽试验^[15]。

近年来对高粱抗旱性的研究多集中于生长和生理特性方面^{[25, 26, 27]}, 而对高粱萌发期抗旱性的系统研究并不多见。本研究的试验材料, 涉及全国主要高粱育种单位选育的高粱品种、杂交组合及部分甜高粱品种, 遗传背景较为丰富, 对不同材料的应用具有一定的借鉴作用。虽然试验材料来自不同生产地区, 受遗传背景影响在萌发期的表现可能会存在一定差异, 但本试验以人工气候箱控制相同的培养条件, 应用PEG在一致的低渗透压下(-0.4 MPa)对不同来源高粱品种萌发特性的研究, 减少或避免了由于品种栽培环境差异对观测结果所产生的偏差。同时, 正式试验前还进行了预备试验, 对干旱水平的控制及试验材料的观测均不少于6次重复, 保证了试验结果的可靠性。

本研究发现, 萌发期干旱胁迫下, 除根干重外, 大部分品种(组合)的萌发指标都有不同程度的降低, 说明干旱胁迫对高粱的萌发生长造成了消极的影响。但也有一些品种(组合)对于干旱的敏感度较低, 甚至干旱激发了高粱的某些萌发状态, 表现为相对值大于1, 如吉杂99、吉杂124等品种和组合1335A× TY3560R、13218A× 20982R的萌发指标都大于对照, 这与前人研究的一定程度的干旱可以提高种子发芽率和幼苗活力的观点较为相似^{[17, 18, 19]}。干旱条件下, 作物生长需要更加庞大的根系以便吸收和利用土壤中的水分^[28]; 一定程度的干旱胁迫可以促进作物根系的生长, 这是作物在一定程度上适应干旱环境的表现^{[29, 30]}, 本试验中大部分高粱品种(组合)根部生物积累量高于对照就证明了此观点。

作物的抗旱性是一个与环境因素密切相关的复杂的数量性状。不同作物萌发期的抗旱性研究都已表明, 种子的发芽率和萌发抗旱指数等指标在鉴定作物抗旱性方面已经得到了广泛的应用, 并能够充分反映作物种子的抗旱能力^{[29, 31]}。但任何单项指标都不能有效、准确地评价其抗旱性。所以, 必须同时观测多个指标, 在综合分析的基础上对试验材料抗旱能力进行有效、准确的评定。陈新等^[12]研究表明, 萌发期裸燕麦的种子活力指数与抗旱性关系密切, 可作为其抗旱性鉴定的主要指标。赫福霞等^[17]和徐蕊等^[22]认为种子萌发抗旱指数可作为鉴定玉米萌发期抗旱性的可靠指标, 发芽势、发芽率等指标与其显著相关。刘桂红等^[20]在谷子萌发期的抗旱性研究中发现, 发芽势、发芽率、幼芽长度、侧根数和根长均可作为抗旱鉴定的主要指标; 安永平等^[21]研究认为, 干旱胁迫下水稻种子的幼苗成活率和萌发胁迫指数可作为鉴定抗旱性的有效评价指标。前人对不同作物萌发期抗旱性的主要鉴定指标提出了不同的观点, 可见这些指标在不同作物抗旱性鉴定中的作用存在一定差异, 对不同作物抗旱性的鉴定需要不同的指标。本试验中, 通过主成分分析将8个抗旱指标归类成三大成分因子, 相对萌发抗旱指数、相对发芽率和相对根长又分别为各成分因子上最大的载荷, 可以作为高粱品种抗旱性鉴定的主要指标, 基于主成分分析的结果得出每个品种(组合)的因子得分, 对高粱萌发期的抗旱性综合评价。从排序和聚类分析的结果上来看, 与试验过程中的表现(表1)也较为一致, 说明能够对不同高粱品种萌发期的抗旱性作出有效、准确的评定。

作物抗旱性受自身遗传因素和环境条件的影响, 有些品种全生育期的抗旱性是一致的, 有些品种则存在着所处生育时期的差异, 即萌发期抗旱不等于中后期也抗旱, 但对于旱粮作物高粱来说, 萌发期抗旱性是非常重要的, 可以保证出苗率和合理群体的建立。我们对部分试验品种生育中后期的抗旱性进行了鉴定, 发现吉杂305在不同生育时期均属于抗旱性较高的品种, 而辽杂35、吉杂127、锦杂103在不同生育时期均属于抗旱性较弱的品种, 同时发现锦杂106生育后期的抗旱能力高于萌发期。本试验仅研究了对不同高粱品种(组合)萌发期的抗旱性, 而对不同干旱胁迫水平以及不同生育时期部分材料抗旱性的差异仍有待从形态、生理和分子水平深入研究。

4 结论

54个高粱品种(组合)按照抗旱能力的强弱分为4大类, 13218A× 20982R是一个抗旱性极强的杂交组合, 吉杂133等13个品种(组合)为抗旱品种(组合), 苏马克等23个品种为干旱敏感品种, 辽杂35等17个品种为干旱高敏感型品种。高粱品种的萌发抗旱指数、发芽率和根长等指标可用作抗旱性鉴定的筛选依据。

附表1 供养试高梁品种及特征 Supplementary table 1 Tested sorghum cultivars and their characteristics

The authors have declared that no competing interests exist.

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[25]	周宇飞, 王德权, 陆樟镳, 王娜, 王艺陶, 李丰先, 许文娟, 黄瑞冬. 干旱胁迫对持绿性高粱光合特性和内源激素ABA, CTK含量的影响. 中国农业科学, 2013, 47: 655-663 Zhou Y F, Wang D Q, Lu Z B, Wang N, Wang Y T, Li F X, Xu W J, Huang R D. Effects of drought stress on photosynthetic characteristics and endogenous hormone ABA and CTK contents in green-stayed sorghum. Sci Agric Sin, 2013, 47: 655-663 (in Chinese with English abstract) [本文引用:1]
[26]	周宇飞, 王德权, 陆樟镳, 王娜, 王艺陶, 李丰先, 许文娟, 黄瑞冬. 干旱胁迫对持绿性高粱叶片渗透调节及叶绿体超微结构的影响. 应用生态学报, 2013, 24: 2545-2550 Zhou Y F, Wang D Q, Lu Z B, Wang N, Wang Y T, Li F X, Xu W J, Huang R D. Impacts of drought stress on leaf osmotic adjustment and chloroplast ultrastructure of stay-green sorghum. Chin Appl Ecol, 2013, 24: 2545-2550 (in Chinese with English abstract) [本文引用:1]
[27]	邵艳军, 山仑, 李广敏. 干旱胁迫与复水条件下高粱, 玉米苗期渗透调节及抗氧化比较研究. 中国生态农业学报, 2006, 14(1): 68-70 Shao Y J, Shan L, Li G M. Comparison of osmotic regulation and antioxidation between sorghum and maize seedlings under soil drought stress and water recovering conditions. Chin J Eco-Agric, 2006, 14(1): 68-70 (in Chinese with English abstract) [本文引用:1]
[28]	张美俊, 杨武德, 乔治军, 冯美臣, 王冠, 段云, 陈凌. 不同糜子品种萌发期对干旱胁迫的响应及抗旱性评价. 草地学报, 2013, 21: 302-307 Zhang M J, Yang W D, Qiao Z J, Feng M C, Wang G, Duan Y, Chen L. Resistance evaluation and response of 16 millet varieties at germination stage to drought stress. Acta Agres Sin, 2013, 21: 302-307 (in Chinese with English abstract) [本文引用:1]
[29]	伏兵哲, 兰剑, 李小伟, 高雪芹. PEG-6000 干旱胁迫对16个苜蓿品种种子萌发的影响. 种子, 2012, 31(4): 10-14 Fu B Z, Lan J, Li X W, Gao X Q. Effects of PEG-6000 drought stress on seed germination of 16 varieties of alfalfa. Seed, 2012, 31(4): 10-14 (in Chinese with English abstract) [本文引用:2]
[30]	罗音, 王玉军, 谢胜利, 赵新西, 杨兴洪, 王玮. 等渗水分与盐分胁迫对烟草种子萌发的影响及外源甜菜碱的保护作用. 作物学报, 2005, 31: 1029-1034 Luo Y, Wang Y J, Xie S L, Zhao X X, Yang X H, Wang W. Effects of isotonic water and salt stress on seed germination of tobacco and protective function of exogenous glycine betaine. Acta Agron Sin, 2005, 31: 1029-1034 (in Chinese with English abstract) [本文引用:1]
[31]	张玉梅, 林琪, 姜雯, 张延胜, 李忠. 渗透胁迫条件下不同抗旱性小麦品种萌发期生理生化指标的变化. 麦类作物学报, 2006, 26(6): 125-130 Zhang Y M, Lin Q, Jiang W, Zhang Y S, Li Z. Effect of osmotic stress on physiological and biochemical indices at germination stage of different drought-resistant wheat. J Triticeae Crops, 2006, 26(6): 125-130 (in Chinese with English abstract) [本文引用:1]

2002

0.0

... 其对粮食产量、品质和效益的影响在将来亦有增加的趋势^[1] ...

2009

0.0

... ) Moench]是我国最早栽培的禾谷类作物之一, 在干旱和半干旱地区分布较广, 具有食用、酿造和饲用等多种使用价值^[2,3] ...

1984

0.0

... ) Moench]是我国最早栽培的禾谷类作物之一, 在干旱和半干旱地区分布较广, 具有食用、酿造和饲用等多种使用价值^[2,3] ...

1976

0.0

... 与其他作物相比, 虽然高粱具有较强的抗旱能力^[4,5,6], 但其生长受干旱影响较大 ...

1987

0.0

... 与其他作物相比, 虽然高粱具有较强的抗旱能力^[4,5,6], 但其生长受干旱影响较大 ...

2009

0.0

山仑, 徐炳成. 论高粱的抗旱性及在旱区农业中的地位. 中国农业科学, 2009, 42: 2342-2348

Shan

, Xu B

Discussion on drought resistance of sorghum and its status in agriculture in arid and semiarid regions.

Sci Agric Sin, 2009, 42: 2342-2348 (in Chinese with English abstract)

Based on authors’ research results and taking maize as the main reference crop, the paper discussed the yield performance, water utilization and drought resistance of sorghum under drought stress environment. Sorghum is regarded as a crop having strong and comprehensive ability of coping with drought, especially low water consumption and high water use-efficiency characteristics. The drought tolerance of sorghum is significantly higher than maize, millet and alfalfa. Sorghum is a typical drought-resistant model crop, which has important values for production and research. The paper also discussed the reasons for the sharp declining in total planting areas of sorghum, and pointed out that as a multifunctional crop possessing high stress resistibility, sorghum can be used for food, fodder, the production of beverages, as well as biofuels, which indicates its favorable developmental prospects. To bring more land under cultivation for sorghum at the regions where the rainfall is lower than 450mm and heat is satisfiable for growth, and strengthen the investigations about drought-resistant mechanism in whole and drought-resistant genome were proposed.

（西北农林科技大学/中国科学院/水利部水土保持研究所）

本文以笔者研究组多年研究结果为基础，以玉米为主要参比作物，就干旱逆境下高粱的产量表现、水分利用及抗旱特性进行了论述。认为高粱是一种综合抗旱能力很强的作物，尤其具有低耗水、高水分利用效率特性，其耐旱性显著高于玉米，也高于谷子和苜蓿，属于一种典型的模式抗旱作物，具有重要的生产和研究价值。指出高粱作为抗逆性很强的粮饲酿兼用作物，特别是作为一种能源植物，仍具良好的发展前景。建议今后在降水量低于450 mm、热量可满足生长的地区扩大高粱种植面积，同时加强对其整体抗旱性机理及抗旱基因组的研究。

... 与其他作物相比, 虽然高粱具有较强的抗旱能力^[4,5,6], 但其生长受干旱影响较大 ...

2010

0.0

... 在干旱环境中, 选择抗旱性更高的作物品种是创建优良栽培群体的重要途径^[7,8], 鉴于此, 抗旱性已成为高粱逆境生理研究的重要内容之一 ...

2006

0.0

... 在干旱环境中, 选择抗旱性更高的作物品种是创建优良栽培群体的重要途径^[7,8], 鉴于此, 抗旱性已成为高粱逆境生理研究的重要内容之一 ...

2011

0.0

... 干旱条件下, 作物出苗情况的优劣依赖于种子的发芽速度及整齐度, 这与种子在低水分条件下的发芽能力密切相关^[9,10] ...

2013

0.0

刘自刚, 沈冰, 张雁. 桔梗种子萌发对低温、干旱及互作胁迫的响应. 生态学报, 2013, 33: 2615-2622

Liu Z

, Shen

, Zhang

Morphological responses to temperature, drought stress and their interaction during seed germination of Platycodon grand iflorum.

Acta Ecol Sin, 2013, 33: 2615-2622 (in Chinese with English abstract)

Controlled experiments were conducted to investigate the effect of temperature, drought and their interaction on seeds germination and seedling growth, aiming to understand how they adapt to the seasonally drought and cold in spring at seed germination stage, for giving base to population resume and cultivation of Platycodon . The results of a two-way ANOVA showed that temperature, drought and their interaction significantly affected the seed germination and seedling growth. Seed germination was optimal at the fluctuating night/day temperature regime of 10/20℃ and constant temperature range from 15℃ to 25℃ in the concentration of 0-10% PEG (except for the concentration of 10% PEG at 20℃), with more than 62.7% of seeds germinated, while seed germination percentage was significantly decreased by the concentration of 15% PEG. Seed germination was significantly inhibited and their initiatory time was delayed at 10℃, germination percentage was increased and then decreased with the increasing of drought stress, while both germination percentage and vigor index all declined at other temperatures. Root length, shoot length and fresh weight were declined with the decreasing of temperature, and root was more sensitive than shoot to changes of temperature.

以药用植物桔梗种子为材料,研究了低温、干旱及其互作对种子萌发和幼苗生长的影响,旨在探索在春寒干旱条件下其种子萌发对策及生态适应性,为桔梗野生种群恢复及人工栽培群体构建提供依据。结果表明,温度、干旱及其互作对桔梗种子萌发和幼苗生长均有显著影响。0-10% PEG水分条件下,在变温10/20℃及恒温15-25℃下桔梗种子能够良好萌发(10%PEG、20℃除外),而15% PEG 可显著抑制萌发。低温(10℃)下种子萌发始时间延迟,发芽率降低。随干旱胁迫程度的增加,10℃发芽率呈先升后降趋势,而其它温度下,发芽率和发芽指数均呈下降趋势。随温度的降低,幼苗根长、茎长及鲜重均呈下降趋势,且幼根生长对温度变化更为敏感。

... 干旱条件下, 作物出苗情况的优劣依赖于种子的发芽速度及整齐度, 这与种子在低水分条件下的发芽能力密切相关^[9,10] ...

2014

0.0

裴帅帅, 尹美强, 温银元, 黄明镜, 张彬, 郭平毅, 王玉国, 原向阳. 不同品种谷子种子萌发期对干旱胁迫的生理响应及其抗旱性评价. 核农学报, 2014, 28: 1897-1904

Pei S

, Yin M

, Wen Y

, Huang M

, Zhang

, Guo P

, Wang Y

, Yuan X

Physiological response of foxtail millet to drought stress during seed germination and drought resistance evaluation.

J Nucl Agric Sci, 2014, 28: 1897-1904 (in Chinese with English abstract)

The drought resistance of four cultivars foxtail millet which was ‘Jingu 28', ‘Jingu 45', ‘Jingu 42' and ‘Jingu 77-322' was studied in seed germination stage. The suitable concentration of PEG was selected firstly for simulating drought stress, then simulated drought stress using the suitable concentration of PEG. Seed Germination and seedling growth characteristics, physiological and biochemical indexes were measured in normal water supply and drought stress conditions. It was shown that: the suitable PEG6000 concentration for drought resistance identification was 18%. The relative germination potential,relative germination rate,germination index and drought index were decreased under drought stress. The content of soluble protein and proline,the activity of SOD and POD,the content of MDA were increased.Comparison with the CK,the content of proline and protein and the activity of SOD and POD of ‘Jingu 45' was raised by 462.34%, 34.17%, 34.17% and 61.22%,respectively. After drought stress, the activity of SOD of ‘Jingu 28' was increased by 266.35%.The content of POD of ‘Jingu 42' was reduced by 9.9%, 7.5% and 19.5% respectively than ‘Jingu 28', ‘Jingu 45' and ‘Jingu 77-322'.The largest increase of MDA content was ‘Jingu 42',it was increased 98.7% higher than CK;the smallest increase was Jingu28.The comprehensive evaluation of the relative seed germination rates, relative plumule:radicle ratio and some of physiological and biochemical parameters of four cultivars foxtail millet by membership function method indicated that the drought resistance of the four cultivars decreased in the order of : ‘Jingu 45'>‘Jingu 28'>‘Jingu 77-322'>‘Jingu 42'. These results provides the basis for millet germplasm resources apprasal and the new drought-resistant variety's seed selection.

1. College of Agriculture, Shanxi Agricultural University, Taigu, Shanxi 030801; 2. Arid Farming Research Center, Shanxi Academy of Agricultural Sciences, Taiyuan Shanxi 030031

对4个品种的谷子晋谷28、晋谷45、晋谷42和晋谷77-322进行种子萌发期抗旱性研究.试验首先筛选出合适的PEG浓度模拟干旱胁迫条件;在正常供水和模拟干旱条件下测定了种子发芽特性及生长、生理生化等指标.结果表明:18%PEG浓度为合适的干旱胁迫条件;随着干旱胁迫的加强,谷子的相对发芽势、相对发芽率、发芽指数和抗旱指数都呈下降趋势.在18%PEG的胁迫下,谷子的可溶性蛋白、脯氨酸含量、SOD和POD 活性、MDA含量都有所升高,差异较显著.其中,晋谷45的脯氨酸含量、蛋白质含量、SOD、POD活性比对照分别提高 462.34%、34.17%、113.84%、61,22%,晋谷28的SOD活性提高266.35%.晋谷42的POD活性比28、45、77-322三个品种分别降低了9.9%、7.5%、19.5%,MDA含量28增幅最小,42增幅最大比CK增加98.7%.经过隶属函数法分析谷子萌芽期抗旱性的强弱顺序为:晋谷45>晋谷28>晋谷77-322>晋谷42.研究为谷子抗旱种质资源鉴定评价及品种选育提供理论依据.

... 裴帅帅等^[11]认为用单一的性状或生理指标不能对萌发期的抗旱机制进行综合评价 ...

2014

0.0

... 陈新等^[12]研究表明, 利用主成分分析的方法能消除各鉴定指标间信息重叠的干扰, 可有效评价裸燕麦萌发期的抗旱性 ...

... 种子的发芽率、发芽势、萌发抗旱指数是玉米^[17]、小麦^[18]、水稻^[19]等作物萌发期抗旱性评定的常用指标, 具有一定的可靠性, 也可以运用到高粱萌发期抗旱性的鉴定中, 但是不同作物萌发期抗旱性的主要鉴定指标仍存在一定差异^{[12,17,20,21,22]} ...

... 陈新等^[12]研究表明, 萌发期裸燕麦的种子活力指数与抗旱性关系密切, 可作为其抗旱性鉴定的主要指标 ...

2012

0.0

孙璐, 周宇飞, 汪澈, 肖木辑, 陶冶, 许文娟, 黄瑞冬. 高粱品种萌发期耐盐性筛选与鉴定. 中国农业科学, 2012, 45: 1714-1722

Sun

, Zhou Y F. Wang C, Xiao M

, Tao

, Xu W

, Huang R

Screening and identification of sorghum cultivars for salinity tolerance during germination.

Sci Agric Sin, 2012, 45: 1714-1722 (in Chinese with English abstract)

【Objective】The objectives of the current study were to explore the response of different sorghum cultivars to salt stress, screen sorghum cultivars suitable for growing in saline soils, and provide a foundation for management of salt stress.【Method】 Thirty seeds of each of 42 sorghum cultivars were treated with 150 mmol•L-1NaCl or distilled water and incubated in Petri dishes for 10 days at humidity of 60%, light/dark of 12h/12h, illumination of 34 μmol•m-2•s-1 and 28℃/25℃ of day/night. Germination and growth were assessed by counting the number of radicals and coleoptiles and leaves that had appeared on a daily basis from the fourth day and measuring their lengths and dry weights on the tenth day using destructive sampling. The cultivars were classified on the basis of salinity tolerance by means of principal component analysis and cluster analysis considering the range of data on germination and growth traits.【Result】The correlation analysis on the relative values of most germination and growth parameters showed positive relationships among several variables. The cultivars were ranked based on the principal components analysis and five groups on salinity tolerance were sorted for the 42 sorghum cultivars according to the cluster analysis.【Conclusion】Principal component analysis results showed that radical length, coleoptiles and leaf weight and germination rate were the most significant factors and are recommended as the main indexes to identify salinity tolerance of sorghum at germination. The analysis showed five of the 42 cultivars were highly salt tolerant, e.g. Liaoza 15, fourteen cultivars were salt tolerant, e.g. Shenshi 104, twelve cultivars were medium salt sensitive, e.g. Aoza 1, eight cultivars were salt sensitive, e.g Tieza 17, and three were highly salt sensitive, e.g. Longza 10. As this research only relates to ranking sensitivity during germination, further work is needed to determine the mechanism of tolerance and its persistence through the plant life cycle.

1.沈阳农业大学农学院，沈阳 110866 2.沈阳农业大学生物科学技术学院，沈阳 110866

【目的】根据不同高粱品种萌发期对盐胁迫的响应，筛选出适合在盐渍土壤上种植的高粱品种及为耐盐胁迫人工调控提供依据。【方法】采用人工气候箱内培养皿培养，用150 mmol•L-1NaCl对42个高粱品种进行胁迫处理，蒸馏水处理为对照，每培养皿放30粒种子。人工气候箱内湿度60%，光照/黑暗时间为12h/12h，光照强度为134 μmol•m-2•s-1，昼/夜温度为28℃/25℃。培养第4天测定发芽势，第10天测定发芽率、根长、叶长、根重、叶重。通过主成分分析、聚类分析和对各品种的表现综合评定，对高粱品种进行耐盐性强弱的分类。【结果】多项萌发和生长参数的相对值之间存在着极显著或显著的正相关。主成分分析结果表明，根长、叶重和发芽率分别在根部、叶部和萌发因子中的负荷量最大，可作为萌发期高粱耐盐性筛选的主要鉴定指标。42个高粱品种的耐盐性存在很大差异，其中辽杂15等5个品种为高度耐盐品种，沈试104等14个品种为耐盐品种，敖杂1等12个品种为中等耐盐品种，铁杂17等8个品种为盐敏感品种，龙杂10等3个品种为高度盐敏感品种。【结论】150 mmol•L-1NaCl可作为高粱萌发期耐盐性鉴定的适宜盐浓度，根长、叶重和发芽率等指标可用于大批量高粱品种耐盐性的评价。

... 这种综合分析抗逆性的方法在高粱的耐盐碱研究中已得到应用^[13,14] ...

2013

0.0

李丰先, 周宇飞, 王艺陶, 孙璐, 白薇, 闫彤, 许文娟, 黄瑞冬. 高粱品种萌发期耐碱性筛选与综合鉴定. 中国农业科学, 2013, 46: 1762-1771

Li F

, Zhou Y

, Wang Y

, Sun

, Bai

, Yan

, Xu W

, Huang R

Screening and identification of sorghum cultivars for alkali tolerance during germination.

Sci Agric Sin, 2013, 46: 1762-1771 (in Chinese with English abstract)

【Objective】The objective of the current study was to explore the response of 35 different sorghum cultivars to alkali stress, and screen them for suitability for growing in alkaline soil and provide some guidance for their management under alkali stress.【Method】Mixed alkali were used (NaHCO3﹕Na2CO3 with the mole ratio of 9﹕1) to simulate the typical alkali stress environment. The 35 sorghum cultivars were treated with 25 mmol•L-1 solution or distilled water and incubated in Petri dishes for 10 days at humidity of 60%, light/dark of 12 h/12 h, illumination of 34 μmol•m-2•s-1 and 28℃/25℃ of day/night during germination. The germination force was measured on the fourth day and the germination percentage, leaf length, root length, leaf fresh weight, root fresh weight, leaf dry weight, root dry weight and remaining seed dry weight were measured on the tenth day. Through the fuzzy mathematics method of membership function, principal component analysis (PCA) and clustering analysis (CA), the alkali resistance of different sorghum cultivars were evaluated.【Result】The PCA results showed that root length, germination index and leaf dry weight were the most significant factors and are recommended as the main indexes for identifing alkali tolerance of sorghum at germination. The cultivars were ranked based on the membership function, and three groups on alkali tolerance were sorted from the 35 sorghum cultivars according to the CA. The analysis showed that seven of the 35 cultivars were strongly tolerant of alkaline conditions, e. g. Siza25, that twenty-two were had medium tolerance, e. g. Jiza 319 and Chiza 28, and that six were sensitive, e. g. Longza 9.【Conclusion】Principal component analysis showed that root length, germination index and leaf dry weight were the most significant factors and these are recommended as the main indexes to identify alkali tolerance of sorghum at germination, and in plant breeding and evaluation projects should be supported by CA and membership function to provide rigorous assessment of cultivars tolerance of alkalinity.

College of Agronomy, Shenyang Agricultural University, Shenyang 110866

【目的】根据不同高粱品种萌发期对碱胁迫的响应，筛选出适合在盐渍土壤上种植的高粱品种，从而为减轻混合碱胁迫提供调控依据。【方法】采用人工气候箱内培养皿培养，以混合碱（NaHCO3﹕Na2CO3的摩尔比为9﹕1）模拟典型碱胁迫环境，在萌发期以25 mmol•L-1溶液处理35个高粱品种，蒸馏水处理为对照，每培养皿放30粒种子。人工气候箱内湿度为60%，光照/黑暗时间为12h/12h，光照强度为134 μmol•m-2•s-1，昼/夜温度为28℃/25℃。培养第4天测定发芽势，第10天测定发芽率、叶长、根长、叶鲜重、根鲜重、叶干重、根干重、子粒残余干重等性状，通过模糊数学隶属函数法和主成分分析法对各品种进行耐碱性综合评定，并进行聚类分析。【结果】主成分分析结果表明，根长、发芽指数和叶干重分别在根部、萌发因子和叶部中的负荷量最大，可作为萌发期高粱耐碱性筛选的主要鉴定指标。通过模糊数学隶属函数法可以对不同高粱品种进行耐碱性排序，不同高粱品种间耐碱性表现出明显的差异，最后通过聚类分析把35个高粱品种按耐碱性强弱分为三大类，其中四杂25号等7个品种为耐碱品种，吉杂319、赤杂28等22个品种为中等耐碱品种，龙杂9号等6个品种为碱敏感品种。【结论】根长、发芽指数、叶干重等指标可用于大量高粱品种萌发期耐碱性的筛选，隶属函数和聚类分析也可为高粱品种耐碱性在植物育种和鉴定方面提供依据。

... 这种综合分析抗逆性的方法在高粱的耐盐碱研究中已得到应用^[13,14] ...

1982

0.0

... 近年来, 已广泛使用PEG溶液模拟干旱环境进行萌发试验^[15,16] ...

... PEG作为一种理想的渗透调节剂, 在栽培基质中可以降低水势且能保持稳定的渗透压, 经常被用来模拟干旱环境进行发芽试验^[15] ...

2004

0.0

... 近年来, 已广泛使用PEG溶液模拟干旱环境进行萌发试验^[15,16] ...

2014

0.0

... 20982R的萌发指标都大于对照, 这与前人研究的一定程度的干旱可以提高种子发芽率和幼苗活力的观点较为相似^[17,18,19] ...

... 赫福霞等^[17]和徐蕊等^[22]认为种子萌发抗旱指数可作为鉴定玉米萌发期抗旱性的可靠指标, 发芽势、发芽率等指标与其显著相关 ...

2008

0.0

... 20982R的萌发指标都大于对照, 这与前人研究的一定程度的干旱可以提高种子发芽率和幼苗活力的观点较为相似^[17,18,19] ...

2014

0.0

... 20982R的萌发指标都大于对照, 这与前人研究的一定程度的干旱可以提高种子发芽率和幼苗活力的观点较为相似^[17,18,19] ...

2013

0.0

... 刘桂红等^[20]在谷子萌发期的抗旱性研究中发现, 发芽势、发芽率、幼芽长度、侧根数和根长均可作为抗旱鉴定的主要指标 ...

2006

0.0

安永平, 强爱玲, 张媛媛, 张文银, 曹桂兰, 韩龙植. 渗透胁迫下水稻种子萌发特性及抗旱性鉴定指标研究. 植物遗传资源学报, 2006, 4: 421-426

An Y

, Qiang A

, Zhang Y

, Zhang W

, Cao G

, Han L

Study on characteristics of germination and drought-resistance index by osmotic stress in rice.

J Plant Genet Resourc, 2006, 4: 421-426 (in Chinese with English abstract)

用聚乙二醇(PEG)-6000作为渗透剂模拟干旱胁迫,探讨在4种不同浓度的渗透胁迫下水稻种子的萌发特性,并确立间接评价水稻苗期抗旱性的萌发期鉴定指标.结果表明,高浓度渗透液对种子萌发的抑制作用明显大于低浓度,萌发率的干旱胁迫反应指数随渗透液浓度的降低和萌发进程而增加.在不同浓度的渗透胁迫处理中,20%浓度下萌发6 d后的萌发率表现较大的变异幅度和变异系数以及较均衡的品种频率分布,较好地反映了水稻种质对渗透胁迫的不同反应,认为20%/6 d条件较适合作为萌发率鉴定的渗透液模拟干旱胁迫;经反复干旱后的幼苗存活率与萌发胁迫指数和20%胁迫浓度下萌发8 d后芽鞘长的干旱胁迫反应指数(DRI)表现极显著或显著正相关,认为萌发胁迫指数和芽鞘长的DRI可作为评价水稻苗期抗旱性的间接鉴定指标.

... 安永平等^[21]研究认为, 干旱胁迫下水稻种子的幼苗成活率和萌发胁迫指数可作为鉴定抗旱性的有效评价指标 ...

2009

0.0

徐蕊, 王启柏, 张春庆, 吴承来. 玉米自交系抗旱性评价指标体系的建立. 中国农业科学, 2009, 42: 72-84

, Wang Q

, Zhang C

, Wu C

Drought-resistance evaluation system of maize inbred.

Sci Agric Sin, 2009, 42: 72-84 (in Chinese with English abstract)

【Objective】 The present study was to choose drought-resistance indexes of maize in different growing period and establish a resistance evaluation system. 【Method】 Seven indexes were tested at seeding stage treated with different concentrations of solution PEG-6000, and twenty-four indexes were tested at growth stage treated with drought-stress and non-stress treatments in the PVC shed and drought shed.【Result】According to the results of simple relative analyses, nine indexes selected including germination drought resistance index of seed and relative germination energy under the condition of 20% PEG-6000 drought stress, GDRI and relative germination energy under the condition of 15% PEG-6000 drought stress, relative value of leaf water potential at seeding stage, leaf water conservation, kernel per row, axis weight per plant and 100-grain weight, they were correlated with drought resistance coefficient or drought resistance index at P≤0.05 and P≤0.01 level. Stepwise regression analysis was conducted between the relative value of nine indexes and drought resistance coefficient and drought resistance index. The result showed that relative germination energy under the condition of 20% PEG-6000 drought stress, relative value of leaf water potential at seeding stage, leaf water conservation, kernel per row and 100-grain weight significantly influenced the drought resistance. According to these indexes and partial relative coefficient, drought-resistance evaluation equation (D value) and evaluation system was established. 【Conclusion】 The five indexes could be selected as identification indexes for drought resistance in maize. Using D value equation to evaluate the drought resistance of maize is considered scientifically reliable.

山东农业大学农学院/作物生物学国家重点实验室

【目的】筛选玉米不同时期抗旱鉴定指标，建立玉米抗旱性评价体系。【方法】室内运用PEG（6000）胁迫处理，研究了种子吸水速率等7个指标与玉米抗旱性的关系;大棚内通过水分胁迫，研究了叶片相对含水量等24个指标与玉米抗旱性的关系。【结果】通过简单相关分析筛选出15% PEG溶液处理GDRI、相对发芽势，20% PEG溶液处理GDRI、相对发芽势及胁迫后苗期叶片水势、抽雄吐丝期叶片保水力、行粒数、单轴重、百粒重和对照相对值9个与抗旱系数或抗旱指数相关显著的指标。通过逐步回归分析筛选出5个指标：20% PEG溶液处理相对发芽势和胁迫后苗期叶片水势、抽雄吐丝期叶片保水力、行粒数、百粒重和对照相对值。根据逐步回归筛选出的指标和偏相关系数，建立了不同时期的抗旱评价（D值）方程和评价体系。【结论】20% PEG溶液处理相对发芽势和胁迫后苗期叶片水势、抽雄吐丝期叶片保水力、行粒数和百粒重和对照相对值可作为玉米抗旱性鉴定指标，利用这些指标建立的评价（D值）方程可对玉米抗旱性进行较好的评价。

... 赫福霞等^[17]和徐蕊等^[22]认为种子萌发抗旱指数可作为鉴定玉米萌发期抗旱性的可靠指标, 发芽势、发芽率等指标与其显著相关 ...

2009

0.0

齐华, 许晶, 孟显华, 王晓波, 刘明, 管宏军. 水分胁迫下燕麦萌芽期抗旱指标的研究. 种子, 2009, (7): 7-10

, Xu

, Meng X

, Wang X

, Liu

, Guan H

Studies on the drought resistance index of oat at germination stage under water stress.

Seed, 2009, (7): 7-10 (in Chinese with English abstract)

利用高渗溶液模拟干旱胁迫条件,以种子萌发抗旱指数为依据进行燕麦品种萌芽期抗旱性筛选,研究结果表明:蒙2034和9342-4-1-3抗旱性最好,蒙麦H 8814抗旱性较好,坝莜五号、冀张莜二号、Ly03-08等抗旱性较差,草莜一号抗旱性最差.相关分析结果表明:胚根长、胚根干重、贮藏物质运转率与种子萌发抗旱指数相关系数达到了显著水平,这些指标可以作为燕麦萌芽期抗旱性鉴定指标.

... 在试验结束时, 从各处理随机取高粱幼苗5株, 测定芽长、根长、芽干重、根干重、籽粒干重, 计算干物质转运率^[23] ...

2010

0.0

石永红, 万里强, 刘建宁, 王运琦, 郭锐, 吴欣明, 李向林. 多年生黑麦草抗旱性主成分及隶属函数分析. 草地学报, 2010, 18: 669-672

Shi Y

, Wan L

, Liu J

, Wang Y

, Guo

, Wu X

, Li X

Analysis of the principal components and the subordinate function of Lolium perenne drought resistance.

Acta Agrest Sin, 2010, 18: 669-672 (in Chinese with English abstract)

The drought resistance of six Lolium perenne fodder varieties at the seedling stage was studied by using PEG-6000 solution to simulate drought stress in the green house.The physiological and biochemical indexes such as relative water content(RWC),exosmosis rate of cytoplasm(RPP),proline content(Pro),chlorophyll contents(Chl),content of malondialdehyde(MDA),activities of superoxide dismutase(SOD)and CAT were measured.Drought resistance was evaluated by principal components analysis and subordinate function analysis.Results show that the seven significantly different drought-resistance indexes could be classified into 3 component parts.The capacity of drought resistance of the six Lolium perenne fodder varieties is gauged as follows:Oupec>Tove>Nul>Eminent>Polim>Yatsyn.

1. Animal Husbandry and Veterinary Institute, Shanxi Academy of Agricultural Sciences, Taiyuan, Shanxi Province 030032, China; 2. Institute of animal Science, CAAS, Beijing 100193, China

在日光温室PEG-6000模拟干旱胁迫条件下,通过测定6个多年生黑麦草( Lolium perenne L.)品种的相对含水量(RWS)、细胞质外渗率(RPP)、游离脯氨酸(Pro)、叶绿素(Chl)、超氧化物歧化酶(SOD)活性、过氧化氢酶(CAT)活性和丙二醛(MDA)含量等7个理化指标,利用主成分及隶属函数分析法对其抗旱性进行综合评价。结果表明:7个差异显著的抗旱指标应用主成分分析归纳成3个主成分,通过综合隶属函数值评定的抗旱性强弱顺序为:欧必克>多福>努依>卓越>凤凰>雅晴。

... 萌发期是作物群体数量建成的关键时期, 作物在萌发期极易受到干旱环境的影响, 所以在萌发期筛选出具有抗旱能力的品种对于干旱地区农业生产具有重要意义^[24] ...

2013

0.0

... 近年来对高粱抗旱性的研究多集中于生长和生理特性方面^[25,26,27], 而对高粱萌发期抗旱性的系统研究并不多见 ...

2013

0.0

周宇飞, 王德权, 陆樟镳, 王娜, 王艺陶, 李丰先, 许文娟, 黄瑞冬. 干旱胁迫对持绿性高粱叶片渗透调节及叶绿体超微结构的影响. 应用生态学报, 2013, 24: 2545-2550

Zhou Y

, Wang D

, Lu Z

, Wang

, Wang Y

, Li F

, Xu W

, Huang R

Impacts of drought stress on leaf osmotic adjustment and chloroplast ultrastructure of stay-green sorghum.

Chin Appl Ecol, 2013, 24: 2545-2550 (in Chinese with English abstract)

Taking stay-green sorghum (B35) and non-stay green sorghum (Sanchisan) as test materials, a pot experiment was conducted to study their leaf osmotic adjustment and chloroplast ultrastructure at flowering and filling stages under impacts of drought stress (45%-50% of maximum field capacity). For the two sorghum lines, drought stress caused the reduction of their leaf free water content and relative water content, and increased the leaf bound water content, water saturation deficit, and electrical conductivity, with the increment or dement being larger for Sanchisan than for B35. Drought stress increased the leaf soluble sugar content and proline content, with the increment of the soluble sugar content being larger for Sanchisan and the increment of the proline content being larger for B35, while decreased the leaf soluble protein content, with the decrement being larger for Sanchisan than for B35. The chloroplast ultrastructure of both B35 and Sanchisan under drought stress was damaged to some extent, but the damaged degree was obviously lower for B35 than for Sanchisan. The findings indicated that stay-green sorghum had a greater adaptation to drought stress through stronger osmotic adjustment.

(College of Agronomy, Shenyang Agricultural University, Shenyang 110866, China)

在盆栽条件下,研究了开花期和灌浆期干旱胁迫(土壤含水量为田间最大持水量的45%~50%)对持绿性高粱(B35)和非持绿性高粱(三尺三)叶片水分、渗透调节物质以及叶绿体超微结构的影响.结果表明: 干旱胁迫下，两高粱品系叶片自由水含量下降,束缚水含量增加,相对含水量降低,水分饱和亏缺增加,相对电导率增大,但三尺三各指标的变化幅度均大于B35.对于渗透调节物质,干旱胁迫下，三尺三可溶性糖含量的增幅大于B35,脯氨酸含量的增幅小于B35,可溶性蛋白含量的降幅大于B35.干旱胁迫下，B35与三尺三的叶绿体超微结构均受到一定程度的破坏,但B35叶绿体结构保持相对完好,受损程度明显小于三尺三.在干旱胁迫下，持绿性高粱通过较强的渗透调节表现出更好的干旱适应能力.

... 近年来对高粱抗旱性的研究多集中于生长和生理特性方面^[25,26,27], 而对高粱萌发期抗旱性的系统研究并不多见 ...

2006

0.0

邵艳军, 山仑, 李广敏. 干旱胁迫与复水条件下高粱, 玉米苗期渗透调节及抗氧化比较研究. 中国生态农业学报, 2006, 14(1): 68-70

Shao Y

, Shan

, Li G

Comparison of osmotic regulation and antioxidation between sorghum and maize seedlings under soil drought stress and water recovering conditions.

Chin J Eco-Agric, 2006, 14(1): 68-70 (in Chinese with English abstract)

The results from pot experiment showed that the drought tolerance mechanism of osmotic regulation and antioxidation was all found in sorghum and maize seedlings. The drought tolerance of sorghum is stronger than that of maize, its leaf relative water content and water potential are higher under drought condition. The main osmotic regulation substances in sorghum are soluble sugar and protein, catalase(CAT) is used as the main antioxidant enzyme. K + and proline are the main osmotic regulation substances and the main antioxidant enzymes are superoxide dismutase(SOD) and peroxidase(POD) in maize.

盆栽试验结果表明，高粱和玉米2种作物均存在渗透调节和抗氧化耐旱途径。高粱耐旱性较强，干旱条件下叶片相对含水量较高，水势亦较大。高粱以可溶性糖和蛋白质为主要渗透调节物质，以过氧化氢酶(CAT)为主要抗氧化酶。玉米以K + 和脯氨酸为主要渗透调节物质，以超氧化物歧化酶(SOD)和过氧化物酶(POD)为主要抗氧化酶。

... 近年来对高粱抗旱性的研究多集中于生长和生理特性方面^[25,26,27], 而对高粱萌发期抗旱性的系统研究并不多见 ...

2013

0.0

张美俊, 杨武德, 乔治军, 冯美臣, 王冠, 段云, 陈凌. 不同糜子品种萌发期对干旱胁迫的响应及抗旱性评价. 草地学报, 2013, 21: 302-307

Zhang M

, Yang W

, Qiao Z

, Feng M

, Wang

, Duan

, Chen

Resistance evaluation and response of 16 millet varieties at germination stage to drought stress.

Acta Agres Sin, 2013, 21: 302-307 (in Chinese with English abstract)

Related indicators to drought resistance were measured for 16 millet ( Panicum miliaceum L.) varieties at germination stage under simulated drought stress using polyethylence glycol (PEG)-6000 solution. Drought resistance was comprehensively evaluated with subordinative function at this stage to investigate the response of varieties to drought stress. Results showed significant drought resistance among 16 millet varieties. Seed germination speed and rate were decreased by restraining seed germination vigor under drought stress. The inhibitory effect of drought stress on radicle was stronger than on embryo. Therefore, germination drought resistance index, relative germination rate, relative germination energy, relative germ and radicle length, relative germination index and relative vigor index were able to be used as effective indicators for drought resistance identification at the germination stage of millet. Hequhong millet, Yihong millet and Nianfeng millet 5 having stronger drought resistance were tested.

1. College of Agronomy, Shanxi Agricultural University, Taigu, Shanxi Province 030801, China; 2. Institute of Germplasm Resources, Shanxi Academy of Agricultural Sciences, Taiyuan, Shanxi Province 030031, China

利用PEG-6000溶液模拟干旱胁迫,对16个糜子( Panicum miliaceum L.)品种萌发期与抗旱性相关的指标进行测定,比较其抗旱性,并采用隶属函数法对其抗旱性进行综合评价。结果表明:不同糜子品种抗旱性有显著性差异。干旱胁迫抑制糜子种子萌发活力,降低了萌发速度和发芽率;干旱胁迫对糜子胚根的抑制要强于胚芽;萌发抗旱指数、相对发芽率、相对发芽势、相对胚芽长、相对胚根长、相对萌发指数和相对活力指数均可作为糜子萌发期抗旱性鉴定的参考指标。河曲红黍子、伊选红糜、粘丰5号这3个品种抗旱性较强。

... 干旱条件下, 作物生长需要更加庞大的根系以便吸收和利用土壤中的水分^[28] ...

2012

0.0

... 一定程度的干旱胁迫可以促进作物根系的生长, 这是作物在一定程度上适应干旱环境的表现^[29,30], 本试验中大部分高粱品种(组合)根部生物积累量高于对照就证明了此观点 ...

... 不同作物萌发期的抗旱性研究都已表明, 种子的发芽率和萌发抗旱指数等指标在鉴定作物抗旱性方面已经得到了广泛的应用, 并能够充分反映作物种子的抗旱能力^[29,31] ...

2005

0.0

罗音, 王玉军, 谢胜利, 赵新西, 杨兴洪, 王玮. 等渗水分与盐分胁迫对烟草种子萌发的影响及外源甜菜碱的保护作用. 作物学报, 2005, 31: 1029-1034

Luo

, Wang Y

, Xie S

, Zhao X

, Yang X

, Wang

Effects of isotonic water and salt stress on seed germination of tobacco and protective function of exogenous glycine betaine.

Acta Agron Sin, 2005, 31: 1029-1034 (in Chinese with English abstract)

Drought and salt are the two most serious abiotic stresses affecting the yield of crops including tobacco throughout the world. As a good osmolyte and macromolecular-protective substance, there are lots of researches on glycinebetaine (GB). Tobacco can’t synthesize glycinebetaine itself and transge

College of Life Sciences; 2 College of Plant Protective Science, Shandong Agricultural University, Tai’an 271018, Shandong, China

在烟草（Nicotiana tabacum L.）种子萌发期，首先用0.5 mmol/L甘氨酸甜菜碱浸种，再用等渗的25％ PEG-6000和1.6％ NaCl溶液进行胁迫处理，研究了两种胁迫对烟草种子萌发的抑制以及外源甜菜碱对该抑制的缓解作用，并对其生理机制进行了探讨。结果表明，两种胁迫处理对烟草种子的萌发均产生抑制作用，均降低其发芽率、发芽势

2006

0.0

张玉梅, 林琪, 姜雯, 张延胜, 李忠. 渗透胁迫条件下不同抗旱性小麦品种萌发期生理生化指标的变化. 麦类作物学报, 2006, 26(6): 125-130

Zhang Y

, Lin

, Jiang

, Zhang Y

, Li

Effect of osmotic stress on physiological and biochemical indices at germination stage of different drought-resistant wheat.

J Triticeae Crops, 2006, 26(6): 125-130 (in Chinese with English abstract)

In this research,the effect of osmotic stress(-0.975 MPa) on the coleoptile and radicle growth was studied with 17 drought resistant wheat genotypes at germination stage,and several physiological and biochemical indices were analyzed.The results showed th

为给小麦苗期抗旱鉴定提供依据。以17个抗旱性不同的小麦品种为材料，在渗透胁迫（-0．975MPa）条件下。研究了小麦萌发后期部分生理生化指标的变化，并在旱棚池栽控水条件下对品种的抗旱性进行了比较鉴定。结果表明，在渗透胁迫条件下，不同小麦品种萌发期胚芽鞘的脯氨酸含量、胚芽鞘和胚根的MDA含量升高，而胚芽鞘和胚根的SOD、CAT活性以及可溶性蛋白质含量下降。其中胚芽鞘脯氨酸含量、CAT活性、SOD活性均与抗旱指数、抗旱系数相关极显著，反映抗旱性强弱趋势基本一致，可作为小麦早期的抗旱性鉴定指标。渗透胁迫条件下胚芽鞘脯氨酸、CAT和SOD的T／CK值（胁迫测定值／对照值）分别大于2．2、0．8和0．8为强抗旱性品种，而抗旱性弱的品种则分别小于1.5、0．65和0．4。