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作物学报 ›› 2025, Vol. 51 ›› Issue (9): 2412-2432.doi: 10.3724/SP.J.1006.2025.41079

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

大麦干旱胁迫萌发生理及分子机理的差异性与相关性研究

何鹏旭1,2,**(), 姚立蓉1,2,*,**(), 陈远玲1,2, 闫妍1,2, 张宏1,2, 汪军成1,2, 李葆春1,3, 杨轲1,2, 司二静1,2, 孟亚雄1,2, 马小乐1,2, 王化俊1,2   

  1. 1省部共建干旱生境作物学国家重点实验室 / 甘肃省作物遗传改良与种质创新重点实验室, 甘肃兰州 730070
    2甘肃农业大学农学院, 甘肃兰州 730070
    3甘肃农业大学生命科学技术学院, 甘肃兰州 730070
  • 收稿日期:2024-11-12 接受日期:2025-06-01 出版日期:2025-09-12 网络出版日期:2025-06-19
  • 通讯作者: *姚立蓉, E-mail: ylr0384@163.com
  • 作者简介:何鹏旭, E-mail: hpx306080@163.com
    **同等贡献
  • 基金资助:
    本研究由东西部科技协作专项(25CXNA030);甘肃农业大学大学生创新训练项目(202501077);财政部和农业农村部国家现代农业产业技术体系建设专项(CARS-05-02A-02);国家重点实验室开放基金项目(GSCS-2021-02);甘肃省现代农业产业技术体系(麦类作物)(Triticeae Crops);甘肃农业大学公招博士科研启动项目(GAU-KYQD-2022-10);甘肃省教育厅产业支撑计划项目(2021CYZC-12);甘肃农业大学伏羲青年英才计划(GAUfx-04Y011);甘肃农业大学伏羲青年英才计划(Gaufx-03Y06);甘肃省陇原青年英才项目(2023)资助

Differences and correlations in physiological and molecular mechanisms of barley germination under drought stress

HE Peng-Xu1,2,**(), YAO Li-Rong1,2,*,**(), CHEN Yuan-Ling1,2, YAN Yan1,2, ZHANG Hong1,2, WANG Jun-Cheng1,2, LI Bao-Chun1,3, YANG Ke1,2, SI Er-Jing1,2, MENG Ya-Xiong1,2, MA Xiao-Le1,2, WANG Hua-Jun1,2   

  1. 1State key laboratory of Aridland Crop Science / The State Key Laboratory of Crop Genetic Improvement and Germplasm Innovation of Gansu Province, Lanzhou730070, Gansu, China
    2College of Agronomy, Gansu Agricultural University, Lanzhou730070, Gansu, China
    3College of Life Science and Technology, Gansu Agricultural University, Lanzhou730070, Gansu, China
  • Received:2024-11-12 Accepted:2025-06-01 Published:2025-09-12 Published online:2025-06-19
  • Contact: *E-mail: ylr0384@163.com
  • About author:**Contributed equally to this work
  • Supported by:
    East West Science and Technology Cooperation Special Project(25CXNA030);Innovative Training Program of Gansu Agricultural University(202501077);China Agriculture Research System of MOF and MARA(CARS-05-02A-02);State Key Laboratory of Aridland Crop Science Open Fund(GSCS-2021-02);Modern Agricultural Industry Technology System in Gansu Province(Triticeae Crops);Scientific Research Start-up Funds for Openly-recuited Doctors of Gansu Agricultural University(GAU-KYQD-2022-10);Industrial Support Project of Colleges and Universities in Gansu Province(2021CYZC-12);Fuxi Young Talents Fund of Gansu Agricultural University(GAUfx-04Y011);Fuxi Young Talents Fund of Gansu Agricultural University(Gaufx-03Y06);Gansu Province Longyuan Youth Talent Project

摘要: 干旱是大麦(Hordeum vulgare L.)生长发育中面临的重要逆境, 可对种子萌发期、开花期等各个生长阶段造成不可逆影响。为了探究干旱胁迫下不同大麦种质资源的萌发特性, 本试验采用蛭石控水法对54份大麦种质资源萌发特性进行研究。将54份种质资源干旱萌发特性分为好、较好、中等、较差和差5类, 干旱萌发特性中等的种质资源占比最高(37.04%)。E02703897和ZDM5430等7份干旱萌发特性好的种质资源, 种子萌发速度快, 出苗整齐且幼苗健壮; 08京134和资源136-21490058等6份萌发特性差的种质资源, 种子发芽率较低, 萌发时间分散, 幼苗整齐度差。进一步筛选出干旱萌发特性差异显著的6份种质资源E02703897、ZDM5430、7DCADA、Z1920057W、菲特36和资源69-G231M004M进行干旱胁迫下种子萌发过程中生理生化指标、DNA损伤修复分析。结果表明, 干旱萌发特性好的大麦种质资源在干旱萌发中期HVFPG基因表达量与干旱萌发前期HVOGG1基因表达量低于干旱萌发特性差的种质资源。萌发特性好的种质资源干旱萌发前期和中期POD活性受干旱影响较小; 萌发前期SOD、半胱氨酸蛋白酶活性受干旱影响较小; 干旱萌发6 h可溶性糖含量与对照相比无显著差异。萌发特性差的种质资源干旱萌发中期POD活性极显著低于对照, 菲特36干旱萌发过程中可溶性糖含量均极显著低于对照。综上结果表明, 干旱萌发特性好的种质资源在干旱萌发过程中DNA损伤程度小、抗氧化能力和物质转化能力强, 种子萌芽速度快, 幼苗生长质量好。

关键词: 大麦, 干旱胁迫, 种子萌发, DNA修复能力, 物质转化, 抗氧化能力

Abstract:

Drought is a major abiotic stress affecting the growth and development of barley (Hordeum vulgare L.), with potentially irreversible impacts across all growth stages, including seed germination and flowering. To investigate the germination responses of diverse barley germplasm under drought stress, this study employed a vermiculite-based water control method to evaluate the germination characteristics of 54 barley germplasm accessions. Based on drought tolerance during germination, the accessions were classified into five categories: excellent, good, moderate, poor, and very poor, with the highest proportion (37.04%) falling into the moderate group. Seven accessions with strong drought tolerance during germination, such as E0270389 and ZDM5430, exhibited rapid germination, uniform emergence, and vigorous seedling growth. In contrast, six accessions with poor drought tolerance, including 08 Jing 134 and Ziyuan136-21490058, showed low germination rates, asynchronous emergence, and poor seedling uniformity. Six representative accessions (E0270389, ZDM5430, 7DCADA, Z1920057W, Feite 36, and ZY69-G231M004M) with significantly different drought germination characteristics were further analyzed for physiological and biochemical parameters, as well as DNA damage repair mechanisms during germination under drought conditions. The results revealed that accessions with good drought tolerance had lower expression of HvFPG during mid-germination and lower HvOGG1 expression during early germination compared to drought-sensitive accessions. In addition, these drought-tolerant accessions maintained higher peroxidase (POD) activity during early and middle germination stages, and showed less reduction in superoxide dismutase (SOD) and cysteine protease activities before germination. Soluble sugar content at 6 hours after drought induction did not differ significantly from the control. In contrast, drought-sensitive accessions exhibited significantly reduced POD activity during mid-germination and a marked decrease in soluble sugar content, as observed in Feite 36. In summary, barley germplasm with good drought tolerance during germination demonstrated enhanced DNA repair capacity, antioxidant activity, and metabolic adaptability under drought stress, leading to faster germination and healthier seedling development.

Key words: barley, drought stress, seed germination, DNA repair capacity, material transformation, antioxidant capacity

表1

qRT-PCR引物序列"

基因号
Gene ID
基因名称
Gene name
上游引物序列
Upstream primer sequence (5'-3')
下游引物序列
Downstream primer sequence (5'-3')
EF101498.1 Actin GCAACAGATCAGCACACTTCCA GCTGACCCTGGTACTCCATTGT
LOC123424347 FPG TTGTCAGAGCCTGGCCATTT ACCTTCAGCAACCCCCAAAA
LOC123442698 OGG1 CATCGCACGGATCGAGAAGA GCCAGCAACATACTTTGCCC

表2

不同种质资源和干旱胁迫对大麦发芽指标的方差分析"

指标
Index
变异来源
Source of variation
基因型
Genotype
处理
Treatment
基因型×处理
Genotype × treatment
发芽势GE (%) 方差SS 132,851.200 225,625.000 13,399.074
自由度DF 53 1 53
均方差MS 2506.626 225,625.000 2460.360
FF-value 30.57*** 2751.662*** 30.006***
发芽率GP (%) 方差SS 144,931.859 183,469.444 146,747.222
自由度DF 53 1 53
均方差MS 2734.563 183,469.444 2768.816
FF-value 30.011*** 2013.537*** 30.387***
发芽指数GI 方差SS 10,195.863 16,337.371 1457.406
自由度DF 53 1 53
均方差MS 192.375 16,337.371 27.498
FF-value 88.527*** 7518.092*** 12.654***
活力指数VI 方差SS 147.866 223.301 17.833
自由度DF 53 1 53
均方差MS 2.790 223.301 0.336
FF-value 93.957*** 7520.215*** 11.331***
单株干重SSDW (mg) 方差SS 0.004 0.006 0.001
自由度DF 53 1 53
均方差MS 7.6510E-05 0.006 1.7580E-05
FF-value 45.705*** 3552.814*** 10.501***
根长RL (cm) 方差SS 1287.199 1441.165 245.208
自由度DF 53 1 53
均方差MS 24.287 1441.165 4.627
FF-value 71.640*** 4251.089*** 13.647***
苗长SL (cm) 方差SS 1143.616 3170.514 155.539
自由度DF 53 1 53
均方差MS 21.578 3170.514 2.935
FF-value 62.020*** 9112.909*** 8.435***
根冠比 R/S 方差SS 1258.209 468.934 1271.404
自由度DF 53 1 53
均方差MS 23.740 468.934 23.989
FF-value 4.598*** 90.816*** 4.646***

附表1

正常条件下不同大麦种质资源发芽生长指标的差异比较"

指标
Index
类别
Category
品种数量
Number of cultivars
品种名称
Cultivar name
指标范围
Index range
发芽势GE (%) 显著高
Significantly higher
10 BGLAHA, IL-34, 资源71-CA2, 21099-2316, 金山本地大麦, 71200TTN, ZDM5430, 7DCADA, ZDM5458, IL-54
BGLAHA, IL-34, ZY71-CA2, 21099-2316, JSBD barley, 71200TTN, ZDM5430, 7DCADA, ZDM5458, IL-54
> 99%
显著低
Significantly lower
7 G0587125V, DM-1300, 资源69-G231M004M, 资源144-C7067705, 菲特36, G0584001, 7极亲本抗病154
G0587125V, DM-1300, ZY69-G231M004M, ZY144-C7067705, FT 36, G0584001, 7JQBKB154
< 90%
发芽率GP (%) 显著高
Significantly higher
17 BGLA HA, 黑龙江, EDM5189, IL-36, IL-34, G0584001, 资源- 173-3289, 资源136-21490058, 资源71-CA2, 21099-2316, 金山本地大麦, 71200TTN, ZDM5430, 7DCADA, ZDM5458, EZ04V021W, IL-54
BGLA HA, HLJ, EDM5189, IL-36, IL-34, G0584001, ZY-173- 3289, ZY136-21490058, ZY71-CA2, 21099-2316, JSBD barley, 71200TTN, ZDM5430, 7DCADA, ZDM5458, EZ04V021W, IL-54
100%
显著低
Significantly lower
7 118ZDM5512, 沾益红毛大麦, IL-49, ZYM21, 7极亲本抗病154, IL-48, IL-20
118ZDM5512, ZYHM barley, IL-49, ZYM21, 7JQBKB154, IL-48, IL-20
< 95%
发芽指数GI 显著高
Significantly higher
6 资源71-CA2, Z1920057W, 21099-2316, 金山本地大麦, 71200TTN, E02703897
ZY71-CA2, Z1920057W, 21099-2316, JSBD Barley, 71200TTN, E02703897
> 30
显著低
Significantly lower
5 资源69-G231M004M, 资源144-C7067705, 菲特36, G0584001, G0401018K-1
ZY69-G231M004M, ZY144-C7067705, FT36, G0584001, G0401018K-1
< 22
活力指数VI 显著高
Significantly higher
4 Z1920057W, 21099-2316, E02703897, 7DCADA > 0.5
显著低
Significantly lower
3 资源69-G231M004M, 资源144-C7067705, 菲特36
ZY69-G231M004M, ZY144-C7067705, FT36
< 0.4
单株干重SSDW (mg) 显著高
Significantly higher
8 GR5-419网8, BM13-007413, ZYM21, Z1920057W, 21099-2316, ZDM5430, E02703897, 7DCADA
GR5-419W8, BM13-007413, ZYM21, Z1920057W, 21099-2316, ZDM5430, E02703897, 7DCADA
> 17
显著低
Significantly lower
2 资源144-C7067705, IL-48
ZY144-C7067705, IL-48
< 15
根长RL (cm) 显著高
Significantly higher
5 118ZDM5512, 2117W46V, BE-ATRJX, GR5-419网8, ZDM5430
118ZDM5512, 2117W46V, BE-ATRJX, GR5-419W8, ZDM5430
> 12.4
显著低
Significantly lower
3 资源69-G231M004M, 资源144-C7067705, 菲特36
ZY69-G231M004M, ZY144-C7067705, FT36
< 10.2
苗长SL (cm) 显著高
Significantly higher
3 金山本地大麦, ZDM5430, ZYM21
JSBD barley, ZDM5430, ZYM21
> 11.5
显著低
Significantly lower
1 08京134
08J134
< 6
根冠比R/S 显著高
Significantly higher
4 2117W46V, 08京134, IL-48, IL-54
2117W46V, 08J134, IL-48, IL-54
> 1.6
显著低
Significantly lower
3 资源69-G231M004M, 资源144-C7067705, 菲特36
ZY69-G231M004M, ZY144-C7067705, FT36
< 0.8

附表2

干旱条件下不同大麦种质资源发芽生长指标的差异比较"

指标
Index
类别
Category
品种数量
Number of cultivars
品种名称
Cultivar name
指标范围
Index range
发芽势GE (%) 显著高
Significantly higher
4 黄茫大麦92, 资源71-CA2, 21099-2316, 金山本地大麦
HM barley 92, ZY71-CA2, 21099-2316, JSBD barley
> 95%
显著低
Significantly lower
11 Z1450066W, 丹青4号, EDM5189, BM13-007413, ZW20400211W, 资源144-C7067705, 菲特36, IL-12, Z133V057W, EZ04V021W, IL-48
Z1450066W, DQ4, EDM5189, BM13-007413, ZW20400211W, ZY144- C7067705, FT36, IL-12, Z133V057W, EZ04V021W, IL-48
< 70%
发芽率GP (%) 显著高
Significantly higher
6 黄茫大麦92, 资源71-CA2, 21099-2316, 金山本地大麦, ZDM5430, E02703897
HM barley 92, ZY71-CA2, 21099-2316, JSBD barley, ZDM5430, E02703897
> 98%
显著低
Significantly lower
8 Z1450066W, BM13-007413, ZW20400211W, 资源144-C7067705, 菲特36, IL-12, Z133V057W, EZ04V021W
Z1450066W, BM13-007413, ZW20400211W, ZY144-C7067705, FT36, IL-12, Z133V057W, EZ04V021W
< 95%
发芽指数GI 显著高
Significantly higher
5 资源71-CA2, Z1920057W, 金山本地大麦, 71200TTN, E02703897
ZY71-CA2, Z1920057W, JSBD barley, 71200TTN, E02703897
> 25
显著低
Significantly lower
9 Z1450066W, EDM5189, ZW20400211W, 资源144-C7067705, 菲特36, IL-12, Z133V057W, EZ04V021W, IL-48
Z1450066W, EDM5189, ZW20400211W, ZY144-C7067705, FT36, IL-12, Z133V057W, EZ04V021W, IL-48
< 21
活力指数VI 显著高
Significantly higher
3 Z1920057W, 金山本地大麦, E02703897
Z1920057W, JSBD Barley, E02703897
> 0.4
显著低
Significantly lower
11 Z1450066W, EDM5189, 菲特36, 资源136-21490058, IL-12, 7极亲本抗病154, Z133V057W, EZ04V021W, IL-48, IL-20, IL-54
Z1450066W, EDM5189, FT36, ZY136-21490058, IL-12, 7JQBKB154, Z133V057W, EZ04V021W, IL-48, IL-20, IL-54
< 0.3
单株干重SSDW (mg) 显著高
Significantly higher
8 资源71-CA2, ZYM21, Z1920057W, 21099-2316, 金山本地大麦, ZDM5430, E02703897, 7DCADA
ZY71-CA2, ZYM21, Z1920057W, 21099-2316, JSBD barley, ZDM5430, E02703897, 7DCADA
> 15
显著低
Significantly lower
20 Z1450066W, EDM5189, BM13-007413, 资源69-G231M004M, ZY 144-C7067, 菲特36, G0584001, 2046R0805, BONC785, ZY 136-21490, G0401018K-1, ZDM5458, IL-12, 7极亲本抗病154, Z133V057W, EZ04V021W, 08京134, IL-48, IL-20, IL-54
Z1450066W, EDM5189, BM13-007413, ZY69-G231M004M, ZY 144- C7067, FT36, G0584001, 2046R0805, BONC785, ZY 136-21490, G0401018K-1, ZDM5458, IL-12, 7JQBKB154, Z133V057W, EZ04V021W, 08J134, IL-48, IL-20, IL-54
< 12
根长RL (cm) 显著高
Significantly higher
4 黄茫大麦92, ZYM21, ZDM5430, E02703897
HM barley92, ZYM21, ZDM5430, E02703897
> 10.2
显著低
Significantly lower
8 ZY 69-G231M, 菲特36, 2046R0805, G0401018K-1, IL-12, EZ04V021W, IL-20, IL-54
ZY 69-G231M, FT36, 2046R0805, G0401018K-1, IL-12, EZ04V021W, IL-20, IL-54
< 6.2
苗长SL (cm) 显著高
Significantly higher
4 金山本地大麦, 71200TTN, ZDM5430, E02703897
JSBD barley, 71200TTN, ZDM5430, E02703897
> 6.1
显著低
Significantly lower
10 EDM5189, BM13-007413, ZY 69-G231M, ZY 144-C7067, G0584001, G0401018K-1, IL-12, 7极亲本抗病154, EZ04V021W, IL-54
EDM5189, BM13-007413, ZY 69-G231M, ZY 144-C7067, G0584001, G0401018K-1, IL-12, 7JQBKB154, EZ04V021W, IL-54
< 3.2
根冠比R/S 显著高
Significantly higher
6 EDM5189, ZDM5458, EZ04V021W, 08京134, IL-20, IL-54
EDM5189, ZDM5458, EZ04V021W, 08J134, IL-20, IL-54
> 2.3
显著低
Significantly lower
1 EDM390404 < 1.5

图1

不同大麦种质资源耐旱系数聚类分析图 I: 干旱萌发特性好的种质资源; II: 干旱萌发特性较好的种质资源; III: 干旱萌发特性中等的种质资源; IV: 干旱萌发特性较差的种质资源; V: 干旱萌发特性差的种质资源。JSBD barley: 金山本地大麦; HM barley 92: 黄茫大麦92; ZY71-CA2: 资源71-CA2; 08J134: 08京134; ZY173-3289: 资源173-3289; ZY136-21490058: 资源136-21490058; DQ4: 丹青4号; FT36: 菲特36; ZY144-C7067705: 资源144-C7067705; ZY69-G231M004M: 资源69-G231M004M; 7JQBKB154:7极亲本抗病154; HLJ: 黑龙江; ZYHM barley: 沾益红毛大麦; GR5-419W8: GR5-419网8。"

附表3

干旱胁迫下发芽生长指标降幅小的大麦种质资源"

指标
Index
降幅
Decrease amplitude
品种数量
Number of cultivars
品种名称
Cultivar name
发芽势GE (%) 无影响No effect 4 黄茫大麦92, ZYM21, Z1920057W, ZY 71-CA2
HM barley 92, ZYM21, Z1920057W, ZY 71-CA2
发芽率GP (%) 无影响No effect 6 IL-49, ZY 71-CA2, 21099-2316, 金山本地大麦, ZDM5430, E02703897
IL-49, ZY 71-CA2, 21099-2316, JSBD barley, ZDM5430, E02703897
发芽指数GI < 33% 3 金山本地大麦, 71200TTN, E02703897
JSBD barley, 71200TTN, E02703897
活力指数VI < 44% 5 黄茫大麦92, 金山本地大麦, ZDM5430, E02703897, 7DCADA
HM barley 92, JSBD Barley, ZDM5430, E02703897, 7DCADA
单株干重SSDW (mg) < 13% 4 IL-48, IL-54, IL-20, EZ04V021W
根长RL (cm) < 16% 3 ZY 144-C7067, ZY 71-CA2, ZYM21
苗长SL (cm) < 38% 3 黄茫大麦92, 金山本地大麦, 71200TTN
HM barley 92, JSBD barley, 71200TTN

附表4

干旱胁迫下发芽生长指标降幅大的大麦种质资源"

指标
Index
降幅
Decrease amplitude
品种数量
Number of cultivars
品种名称
Cultivar name
发芽势GE (%) 40%-50% 4 118ZDM5512, BGLA HA, 33329, DM-1300
发芽率GP (%) 60%-80% 3 ZDM5160, TRADI70W, BONC785
发芽指数GI 60%-70% 8 BE-ATRJX, 118ZDM5512, GR5-419网8, BGLA HA, 沾益红毛大麦, IL-53, GERTR0V, 33329
BE-ATRJX, 118ZDM5512, GR5-419W8, BGLA HA, ZYHM Barley, IL-53, GERTR0V, 33329
活力指数VI 60%-75% 12 E03V005W, IL-49, IL-36, IL-3, Z145V020W, IL-34, 21099-2316, 33329, BGLA HA, GERTR0V, 黑龙江, DM-1300
E03V005W, IL-49, IL-36, IL-3, Z145V020W, IL-34, 21099-2316, 33329, BGLA HA, GERTR0V, HLJ, DM-1300
单株干重SSDW (g) 40%-60% 9 BE-ATRJX, 118ZDM5512, ZDM5160, GR5-419网8, TRADI70W, 33329, IL-3, DM-1300, ZY 144-C7067
BE-ATRJX, 118ZDM5512, ZDM5160, GR5-419W8, TRADI70W, 33329, IL-3, DM-1300, ZY 144-C7067
根长RL (cm) 30%-35% 6 GR5-419网8, TRADI70W, GERTR0V, E03V005W, IL-3, DM-1300
GR5-419W8, TRADI70W, GERTR0V, E03V005W, IL-3, DM-1300
苗长SL (cm) 52%-60% 5 沾益红毛大麦, 黑龙江, E03V005W, IL-3, Z145V020W
ZYHM barley, HLJ, E03V005W, IL-3, Z145V020W

图2

不同大麦种质资源发芽生长指标差异分析 FT36: 菲特36; ZY69-G231M004M: 资源69-G231M004M。CK: 对照; DT: 干旱胁迫。n = 3, *表示在0.05概率水平差异显著, **代表在0.01概率水平差异显著。"

图3

不同大麦种质资源种子萌发时期与幼苗形态比较 缩写同图2。"

图4

干旱胁迫下不同大麦种质资源种胚HVFPG和HVOGG1基因表达量差异 缩写同图2。*、**分别表示在0.05和0.01水平差异显著。"

图5

干旱胁迫下不同大麦种质资源种胚DNA修复基因表达量耐旱系数 缩写同图2。"

图6

干旱胁迫对不同大麦种质资源种子萌发过程中SOD、POD活性的影响 缩写同图2。*表示在0.05概率水平差异显著, **代表在0.01概率水平差异显著。"

图7

大麦种子不同萌发时间SOD、POD活性与发芽指标抗旱系数相关性分析 GE: 发芽势; GP: 发芽率; GI: 发芽指数; VI: 活力指数; SSDW: 单株干重。SOD: 超氧化物歧化酶; POD: 过氧化物酶。*表示在0.05概率水平差异显著, **表示在0.01概率水平相关性显著。"

图8

干旱胁迫对不同大麦种质资源种子萌发过程中α-淀粉酶、半胱氨酸蛋白酶活性的影响 缩写同图2。*表示在0.05概率水平差异显著, **代表在0.01概率水平差异显著。"

图9

大麦种子不同萌发时间α-淀粉酶、半胱氨酸蛋白酶活性与发芽指标抗旱系数相关性分析 RT: 根尖数; RL: 根长; SL: 苗长; TRSA: 总根表面积。α-AL: α-淀粉酶; CP:半胱氨酸蛋白酶。*表示在0.05概率水平相关性显著; **表示在0.01概率水平相关性显著。"

图10

干旱胁迫对不同大麦种质资源种子萌发过程中可溶性糖、可溶性蛋白含量的影响 缩写同图2。*表示在0.05概率水平差异显著, **代表在0.01概率水平差异显著。"

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