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大豆出苗期耐盐性鉴定方法建立及耐盐种质筛选

本站小编 Free考研考试/2021-12-26

刘谢香, 常汝镇, 关荣霞,*, 邱丽娟,*中国农业科学院作物科学研究所/国家农作物基因资源与遗传改良重大科学工程/农业部种质资源利用重点实验室, 北京 100081

Establishment of screening method for salt tolerant soybean at emergence stage and screening of tolerant germplasm

LIU Xie-Xiang, CHANG Ru-Zhen, GUAN Rong-Xia,*, QIU Li-Juan,*Institute of Crop Sciences, Chinese Academy of Agricultural Sciences/National Key Facility for Crop Gene Resources and Genetic Improvement (NFCRI)/Key Laboratory of Germplasm Utilization, Ministry of Agriculture, Beijing 100081, China

通讯作者: *邱丽娟, E-mail: qiulijuan@caas.cn 关荣霞, E-mail: guanrongxia@caas.cn

收稿日期:2019-04-17接受日期:2019-08-9网络出版日期:2019-09-03
基金资助:本研究由国家自然科学基金项目资助.31830066


Received:2019-04-17Accepted:2019-08-9Online:2019-09-03
Fund supported: This study was supported by the National Natural Science Foundation of China.31830066

作者简介 About authors
E-mail:15311442897@163.com。






摘要
土壤盐渍化是影响农业生产的重要问题, 筛选耐盐大豆资源对于大豆主产区盐渍化土壤的利用具有重要意义。以中黄35、中黄39、Williams 82、铁丰8号、Peking和NY27-38为供试材料, 以蛭石为培养基质, 设0、100和150 mmol L -1NaCl 3个处理, 进行出苗期耐盐性鉴定, 分析与生长相关的6个指标, 旨在明确大豆出苗期耐盐性鉴定指标和评价方法。结果表明, 150 mmol L -1 NaCl处理显著降低大豆的成苗率、株高、地上部鲜重、根鲜重、地上部干重和根干重, 并且不同材料间差异显著。基于幼苗生长发育状况的耐盐指数方法与耐盐系数方法对6份种质耐盐性评价结果显著相关。耐盐指数法对植株无损坏、可省略种植对照, 节约人力和物力, 提高种质鉴定的效率。因此, 以150 mmol L -1 NaCl作为出苗期耐盐鉴定浓度, 以耐盐指数作为大豆出苗期耐盐鉴定评价指标, 鉴定27份大豆资源, 获得出苗期高度耐盐大豆(1级) 3份、耐盐大豆(2级) 7份, 其中4份苗期也高度耐盐(1级), 分别为运豆101、郑1311、皖宿1015和铁丰8号。本研究建立了一种以蛭石为基质, 利用150 mmol L -1 NaCl处理, 以耐盐指数作为评价指标的大豆出苗期耐盐性鉴定评价的简便方法, 并筛选出4份出苗期和苗期均耐盐的大豆, 对耐盐大豆种质资源的高效鉴定和耐盐大豆新品种培育具有重要意义。
关键词: 大豆;出苗期;耐盐性;鉴定方法

Abstract
Salinity is an important factor affecting crop production. Screening salt tolerant soybean germplasm is of great significance for the utilization of salinized soil in major soybean production regions. In order to select salt tolerant soybean, a screening method was developed by using six soybean accessions, including Zhonghuang 35, Zhonghuang 39, Williams 82, Tiefeng 8, Peking, and NY27-38. Seeds were grown in vermiculite and treated with 0, 100, and 150 mmol L -1 NaCl solution. Seedling rate (SR), plant height (H), fresh weight of shoot and root (FWS and FWR), dry weight of shoot and root (DWS and DWR) were decreased significantly under 150 mmol L -1 NaCl treatment, with significant difference among varieties. Therefore, 150 mmol L -1 NaCl was suitable to identify salt tolerant soybean at emergence stage. The salt tolerance index (SI) based on the growth and development of seedlings and the salt tolerance coefficient (ST) were significantly correlated with the salt tolerance. The method using salt tolerance index is non-destructive and does not require planting control, which could save time and labor in salt tolerant germplasm identification. Twenty-seven soybean resources were screened, in which three were highly tolerant (grade 1) and seven tolerant (grade 2) at emergence stage. Among them, Yundou 101, Zheng 1311, Wansu 1015, and Tiefeng 8 also showed salt tolerance (grade 1) at seedling stage. In summary, an effective method for screening salt tolerant soybean at emergence stage was developed, with vermiculite as the substrate, 150 mmol L -1 NaCl as suitable treatment solution, and salt tolerance index as the indicator. Four soybean accessions were found to be salt tolerant at both emergence and seedling stages. This screening method will be useful for identification of salt tolerant soybean germplasm.
Keywords:soybean;emergence stage;salt tolerance;screening method


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本文引用格式
刘谢香, 常汝镇, 关荣霞, 邱丽娟. 大豆出苗期耐盐性鉴定方法建立及耐盐种质筛选[J]. 作物学报, 2020, 46(1): 1-8. doi:10.3724/SP.J.1006.2020.94062
LIU Xie-Xiang, CHANG Ru-Zhen, GUAN Rong-Xia, QIU Li-Juan. Establishment of screening method for salt tolerant soybean at emergence stage and screening of tolerant germplasm[J]. Acta Agronomica Sinica, 2020, 46(1): 1-8. doi:10.3724/SP.J.1006.2020.94062


土壤盐渍化影响世界上20%的可用耕地, 是导致作物减产的重要因素之一[1,2,3]。据全国第二次土壤普查, 中国盐渍土总面积约3600万公顷, 占可利用土地面积的4.88%[4]。通过对不同作物盐度临界值(不导致产量降低的最大土壤盐度)的研究发现, 大豆属于中度耐盐作物, 盐胁迫可抑制大豆的萌发、生长发育和根瘤形成[5,6,7]。当土壤盐度超过5 dS m-1时, 大豆的产量开始降低; 当土壤盐度为15 dS m-1时, 大豆产量显著降低甚至绝收[8,9]。耐盐大豆品种的培育是有效利用盐渍化土壤、促进大豆可持续发展的重要途径[2]

大豆萌发期、出苗期、营养生长期和生殖生长期等不同生育阶段的耐盐性没有明显相关性, 说明不同时期可能存在不同的耐盐机制[5,9-11]。大豆苗期耐盐性的研究较为深入, 利用分子标记技术已经检测到分布在N、D2、G等连锁群上的多个苗期耐盐性相关的数量性状位点(Quantitative Trait Locus, QTL), 特别是位于N连锁群的主效QTL, 是在野生、栽培大豆不同耐盐资源中保守的重要位点[1-2,8,10,12-19]。本实验室前期在耐盐大豆铁丰8号中图位克隆了位于N连锁群的苗期耐盐基因GmSALT3[17], 在此基础上利用分子标记构建了分别携带GmSALT3Gmsalt3等位基因的近等基因系, 研究发现GmSALT3控制大豆苗期耐盐性, 但与大豆出苗期的耐盐性无关, 进一步证明出苗期耐盐性可能存在不同的遗传调控机制[20]。有研究发现, 作物萌发期的耐盐性远高于出苗期的耐盐性, 因此出苗期耐盐性比萌发期耐盐性更具有研究和利用价值[21,22,23,24]

不同研究者对大豆萌发期和出苗期耐盐性的鉴定方法、鉴定指标和评价方法也存在一些差异。鉴定方法主要有田间鉴定和室内鉴定2类; 鉴定指标有形态指标如受害叶面积、株高和生物量, 生长发育指标如发芽率和出苗率, 以及生理生化指标如丙二醛(MDA)含量和超氧化物歧化酶(SOD)活性等; 评价方法有盐害指数和耐盐系数法等[5-6,24-29]

本研究分析6份大豆种质盐胁迫条件下植株成苗率、株高、地上部鲜重、根鲜重、地上部干重和根干重, 建立了一种简便高效的大豆出苗期耐盐性鉴定评价方法, 并利用该方法进行大豆耐盐资源的筛选, 为大豆耐盐优异资源发掘和新品种培育提供了技术支撑。

1 材料与方法

1.1 试验材料

共27份种质, 其中用于方法建立的中黄35、中黄39、Williams 82、铁丰8号、Peking和NY27-38来自国家大豆种质资源库, 其余21份大豆新品种(系)来自2018年国家黄淮海北片品种区域试验(表3)。

1.2 耐盐性鉴定

1.2.1 出苗期耐盐鉴定与性状调查 耐盐性鉴定于2018年5月至6月在中国农业科学院作物科学研究所网室遮雨棚下进行。挑选每份大豆材料90β粒饱满的种子, 种于装有蛭石的8 cm × 8 cm × 8 cm的小花盆中, 每盆10粒, 播种深度为2 cm, 每24个小花盆置于一个大蓝盒(46 cm × 32 cm × 10 cm)内, 用RO水(对照)或盐溶液(100 mmol L-1、150 mmol L-1 NaCl)处理, 3次重复。对照每个蓝盒中浇5 L水, 盐处理每个蓝盒中分别浇100 mmol L-1、150 mmol L-1 NaCl溶液5 L, 使蛭石达到最大持水量, 此后每3天浇2 L水。从第一个大豆出苗(子叶突出蛭石表面)开始, 每天调查出苗数, 第15天调查成苗数(子叶展开、具有叶片的植株)。采用单株分类记载法调查处理条件下所有材料的成苗情况(图1表1)。测量成苗植株的株高, 称量地上部鲜重、根鲜重, 随后于70°C烘箱中烘干(3 d), 分别称量地上部和根干重, 计算相对成苗率(ST_SR)、相对株高(ST_H)、相对地上部鲜重(ST_FWS)、相对根鲜重(ST_FWR)、相对地上部干重(ST_DWS)和相对根干重(ST_DWR)。各指标的相对值, 即耐盐系数(salt tolerance coefficient, ST), 为NaCl处理指标值/对照指标值。耐盐指数(salt tolerance index, SI) = ∑(类别数值×该类别株数)/播种粒数×5 (最高类别数值)。

图1

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图1出苗期耐盐性单株分类记载法的标准

Fig. 1Standard of individual classification for salt tolerance at germination stage



Table 1
表1
表1盐胁迫后大豆出苗期和苗期的盐害症状
Table 1Symptoms of soybean at emergence and seedling stages under salt stress
出苗期 Emergence stage苗期 Seedling stage
类别
Category
表型特征
Symptom
级别
Grade
表型特征
Symptom
I植株凋亡, 子叶干枯(类别数值为1)
Plant dead, cotyledons were dry (category value is 1)
1健康的绿叶, 没有观察到损伤
Healthy green leaves, no damage observed
II植株生长受到严重抑制, 子叶未展开(类别数值为2)
Plant growth was severely inhibited, cotyledon was not unfolded (category value is 2)
2轻度坏死, 真叶轻微发黄
Slight chlorosis, light yellowish color observed in true leaves
III植株生长受到抑制, 具有生长点, 但真叶未展开(类别数值为3)
Plant growth was inhibited with shoot apical meristem, but true leaves were not unfolded (category value is 3)
3中度坏死, 三出复叶发黄
Moderate chlorosis, chlorosis observed in trifoliate leaves
IV植株生长基本正常, 真叶未完全展开(类别数值为4)
Plant growth was basically normal, true leaves were not fully expanded (category value is 4)
4严重坏死, 超过75%的叶面发黄
Severe chlorosis, more than 75% of the leaf area showed chlorosis
V植株生长正常, 真叶完全展开(类别数值为5)
Plant growth was normal, true leaves were fully expanded (category value is 5)
5凋亡, 植物完全枯萎
Dead, plants were completely withered

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1.2.2 苗期耐盐鉴定与性状调查 参照本实验室前期建立的方法鉴定评价苗期耐盐性[17]。每个品种各10粒播于小花盆(8 cm × 8 cm × 8 cm)的蛭石中, 每24个小花盆置于一个大蓝盒内(46 cm × 32 cm × 10 cm), 每个蓝盒浇5 L RO水。此后每3 d浇2 L水。待子叶完全展开后间苗, 每盆留4~6株。待真叶完全展开后(10 d), 每个蓝盒浇2 L 200 mmol L-1 NaCl, 于第13天和第16天再各浇2 L 200 mmol L-1 NaCl, 设置3次重复。在最后一次盐处理5 d后对叶片坏死程度进行评估(表1)。其中1级和2级为耐盐, 3~5级为盐敏感。

1.3 统计分析

应用Microsoft Excel 2010处理数据, SAS9.4软件进行单因素方差分析, 最小显著极差法(Least Significant Differences, LSD)进行多重比较(P<0.05)。

2 结果与分析

2.1 6份大豆种质出苗期耐盐性鉴定

在100 mmol L-1 NaCl处理下, 与对照相比, 6份种质出苗正常且生长发育良好(图2-A, B), 成苗率、地上部干重均与对照差异不显著(图2-D, I)。中黄35 (ZH35)的根鲜重和株高、NY27-38的根和地上部鲜重较对照显著降低, 中黄39 (ZH39)、Williams 82 (W82)、铁丰8号(TF8)和Peking (Pek)的根鲜重、株高和地上部鲜重均与对照差异不显著(图2-E~H)。

图2

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图2不同浓度NaCl处理15 d后6份材料的表型

A、B、C分别表示0、100和150 mmol L-1 NaCl处理15 d的表型特征, 比例尺为1 cm。D: 成苗率。E: 根鲜重。F: 地上部鲜重。G: 株高。H: 根干重; I: 地上部干重。数据结果为3次生物学重复, 误差线为标准误(n = 3); 标以不同小写字母的柱值在同一品种的不同浓度处理间在0.05水平上差异显著。
Fig. 2Phenotypes of six materials after treatment with NaCl solution for 15 days

A, B, and C refer to the phenotype after treatment with 0, 100, and 150 mmol L-1 NaCl for 15 days, respectively. Bar = 1 cm. D: seedling rate. E: fresh weight of root. F: fresh weight of shoot. G: shoot height. H: dry weight of root. I: dry weight of shoot. Data are mean ± SE with three biological replicates (n = 3). Bars superscripted by different letters are significantly different at P < 0.05 between different treatments of the same variety.


当NaCl浓度为150 mmol L-1时, 6份种质出苗率下降、株高降低、生长发育迟缓, 部分种子出苗后子叶未展开, 不能正常成苗(图1图2-C)。除ZH39和NY27-38的成苗率显著下降外, 其余材料的成苗率与对照差异不显著; 6份种质的根鲜重、地上部鲜重、株高、根干重和地上部干重均显著低于对照(图2-D~I)。不同种质间相对成苗率、相对地上部鲜重、相对根鲜重、相对株高、相对地上部干重和相对根干重差异显著(图3); 盐胁迫对不同的材料的影响存在差异, W82、TF8、Pek和ZH39的成苗率、地上部鲜重、根鲜重、株高、地上部干重和根重下降程度较低, 耐盐性较强; 而ZH35和NY27-38的耐盐性较差。因此, 确定150 mmol L-1 NaCl为大豆出苗期耐盐性鉴定适宜盐浓度。根据6个性状的耐盐系数大小, 得出6个材料的耐盐性为W82 > Pek > TF8 > ZH39 > ZH35 > NY27-38。

图3

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图3150 mmol L-1 NaCl处理下6份材料的耐盐性

ST_SR: 相对成苗率; ST_H: 相对株高; ST_FWR: 相对根鲜重; ST_FWS: 相对地上部鲜重; ST_DWR: 相对根干重; ST_DWS: 相对地上部干重; 数据结果为3次生物学重复, 误差线为标准误(n = 3); 标以不同小写字母的柱值在同一指标的不同品种间在0.05水平上差异显著。
Fig. 3Salt tolerance of six materials under 150 mmol L-1 NaCl stress

ST_SR: relative seedling rate; ST_H: relative height; ST_FWR: relative fresh weight of root; ST_FWS: relative fresh weight of shoot; ST_DWR: relative dry weight of root; ST_DWS: relative dry weight of shoot. Data are mean ± SE with three biological replicates (n = 3). Bars superscripted by different letters are significantly different at P < 0.05 between different varieties of the same indicator.


2.2 大豆出苗期耐盐性鉴定的评价指标分析

根据单株分类记载法计算耐盐指数, W82、Pek、TF8、ZH39、ZH35和NY27-38的耐盐指数分别为0.94、0.74、0.72、0.68、0.52和0.34, 耐盐性以W82最强, Pek、TF8和ZH39次之, ZH35和NY27-38最弱。相对成苗率、相对地上部鲜重、相对根鲜重、相对株高、相对地上部干重、相对根干重和耐盐指数7个指标的相关分析表明, 除相对根干重与相对成苗率、相对地上部鲜重相关不显著外, 其余性状间均呈显著或极显著正相关; 相对地上部鲜重、相对株高和相对地上部干重与相对成苗率高度相关; 相对鲜重与相对干重相关系数达0.98; 耐盐指数与6个性状的耐盐系数均显著或极显著正相关(表2)。

Table 2
表2
表2大豆出苗期不同耐盐评价指标间的相关系数
Table 2Correlation coefficients between salt tolerance indexes at emergence stage in soybean
指标
Index
相对成苗率
ST_SR
相对地上部鲜重
ST_FWS
相对根鲜重
ST_FWR
相对株高
ST_H
相对根干重
ST_DWR
相对地上部干重
ST_DWS
相对地上部鲜重 ST_FWS0.97***
相对根鲜重 ST_FWR0.81*0.81*
相对株高 ST_H0.97***0.96**0.85*
相对根干重 ST_DWR0.800.770.98***0.85*
相对地上部干重 ST_DWS0.97***0.98***0.89*0.99***0.87*
耐盐指数 SI0.88*0.83*0.91**0.87*0.88*0.90**
* represent significance level at P < 0.05, ** represent significance level at P < 0.01, and *** represent significance level at P < 0.001. ST_SR: relative seedling rate. ST_H: relative height. ST_FWR: relative fresh weight of root. ST_FWS: relative fresh weight of shoot. ST_DWR: relative dry weight of root. ST_DWS: relative dry weight of shoot.
*表示显著( P < 0.05), **表示极显著 ( P < 0.01 ), ***表示极显著 ( P < 0.001 )。

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耐盐系数法是经典的出苗期耐盐性评价方法, 通过盐处理与对照的比值反映盐胁迫使大豆成苗数、生长量下降的程度。耐盐指数法同时考虑了盐胁迫后大豆的出苗数量及幼苗生长发育状况; 在实际操作方面, 省略了种植对照, 实现了耐盐性状无损调查, 可极大节约人力和物力, 为大批量优异种质的快速鉴定提供了可能。

2.3 大豆耐盐种质筛选

分别以Williams 82和中黄35为耐盐和盐敏感对照, 用150 mmol L-1 NaCl对包括上述6份在内的27份大豆种质进行耐盐性鉴定(图4), 15 d后根据单株分类记载法计算耐盐指数, 可将27份种质的出苗期耐盐性分为5级, 1级为高度耐盐型(0.80≤SI<1.00), 2级为耐盐型(0.60≤SI<0.80), 3级为中度耐盐型(0.40≤SI<0.60), 4级为敏感型(0.20≤SI<0.40), 5级为高度敏感型(0.00≤SI<0.20), 其中1级3份, 2级7份(表3)。同时, 用200 mmol L-1 NaCl对27份大豆品种(系)进行苗期耐盐鉴定, 发现12份材料苗期高度耐盐(1级)。其中, 运豆101、郑1311、皖宿1015和铁丰8号是出苗期和苗期均耐盐的大豆种质(表3)。

Table 3
表3
表327份大豆种质苗期和出苗期的耐盐等级
Table 3Salt tolerance of 27 accessions at seedling and emergence stages
品种
Variety
耐盐性 Salt tolerance品种
Variety
耐盐性 Salt tolerance
出苗期
Emergence stage
苗期
Seedling stage
出苗期
Emergence stage
苗期
Seedling stage
中黄74 Zhonghuang 7435运豆101 Yundou 10121
冀1507 Ji 150731郑1311 Zheng 131121
冀豆29 Jidou 2935冀1503 Ji 150341
冀豆23 Jidou 2341皖宿1015 Wansu 101521
安豆1498 Andou 149815齐黄39 Qihuang 3914
石豆17 Shidou 1741中黄207 Zhonghuang 20733
科豆13 Kedou 1344邯豆11 Handou 1135
中黄206 Zhonghuang 20643Williams 8214
中黄605 Zhonghuang 60525中黄35 Zhonghuang 3535
中黄80 Zhonghuang 8033中黄39 Zhonghuang 3925
中黄203 Zhonghuang 20331铁丰8号 Tiefeng 821
中黄204 Zhonghuang 20431Peking25
中黄70 Zhonghuang 7035NY27-3841
圣豆10号 Shengdou 1031

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图4

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图44份大豆种质苗期和出苗期耐盐性

A: 对照(0 mmol L-1 NaCl处理15 d)。B: 出苗期耐盐性(播种时150 mmol L-1 NaCl处理15 d)。C: 苗期耐盐性(真叶展开时200 mmol L-1 NaCl处理15 d)。比例尺 = 5 cm。
Fig. 4Salt tolerance in both seedling and emergence stages for four accessions

A: control (0 mmol L-1 NaCl treatment for 15 days). B: salt tolerance at emergence stage (150 mmol L-1 NaCl treatment for 15 days after sowing). C: salt tolerance at seedling stage (200 mmol L-1 NaCl treatment for 15 days when the unifoliate leaves of plants were fully expanded). Bars = 5 cm.


3 讨论

3.1 大豆出苗期耐盐性鉴定方法

在盐碱地上,“拿住苗”是大豆生产的关键问题, 因此, 出苗期的耐盐性十分重要[24]。大豆为中度耐盐作物, 耐盐性不同的大豆种质资源盐害症状存在明显差异[9]。国内外****采用了多种不同的方法评价大豆出苗期的耐盐性。邵桂花等[11]通过抽提地下咸水和淡水配制成一定浓度的咸水对2000多个大豆品种进行处理, 根据盐害指数法筛选出出苗期耐盐品种242个。田间鉴定方便快捷, 适用于大批量品种的鉴定。但田间漫灌会导致盐分分布不均匀, 如地头土壤盐浓度仅为3 dS m-1, 而地尾的土壤盐浓度则高达11 dS m-1 [6]。田间鉴定法还易受地力、光照、气温、降水和风力的影响, 可重复性较差, 因此, 需要严格控制鉴定条件[24,30]。罗庆云等[25]利用营养液水培法鉴定大豆出苗期的耐盐性, 发现NaCl胁迫降低种子出苗速率, 影响幼苗正常生长, 但需多次更换营养液, 费时费力。张海波等[26]鉴定发现耐盐性较强的大豆品种较对照MDA含量降低, SOD活性升高, 但生理生化指标易受胁迫方式、胁迫时间、测定部位、测定方式和所用仪器设备等因素影响。为模拟大田环境, 本研究以蛭石为基质, 在遮雨棚下利用自然光温条件进行大豆种质资源的耐盐性鉴定。与田间鉴定法相比, 该方法不受地力、降水等影响, 可重复性高。同时, 蛭石吸水迅速, 有利于保持土壤盐分的相对稳定, 且无需更换溶液或配制混合基质, 具有快速、准确、经济等特点。

邵桂花等[11]在评价大豆出苗期耐盐性时采用的盐水浓度为10~15 mS cm-1。Zhang等[27]鉴定所利用浓度为100 mmol L-1 NaCl。罗庆云等[25]研究发现, 在50 mmol L-1和100 mmol L-1 NaCl营养液胁迫下, 所有参试大豆均能形成部分正常植株, 而在150 mmol L-1 NaCl处理下则不能正常发育成苗。邵桂花等[24]认为NaCl溶液浓度可根据试验目的进行调整, 若为评价耐盐性, 则鉴定浓度可低些, 若筛选高度耐盐种质, 则可适当提高浓度。本研究中用100 mmol L-1 NaCl处理, 所有种质生长发育良好, 相对盐敏感的种质ZH35和NY27-38的成苗率、地上部鲜重、株高和地上部干重与对照差异不显著; 而150 mmol L-1 NaCl处理下, 所有种质的性状均与对照差异显著, 且不同品种受盐胁迫影响的程度差异明显。因此, 以150 mmol L-1 NaCl作为大豆出苗期耐盐性评价鉴定浓度。

邵桂花等[24]通过受害叶面积为分类标准计算盐害指数评价大豆出苗期的耐盐性, 但盐害症状的分辨会因研究人员的经验不同而产生差异[25]。本研究观察到子叶出土后, 盐胁迫对其发育影响较大, 有的子叶无法展开最终干枯凋亡, 也有的能正常生长。因此, 根据幼苗发育情况, 将其分为5级(图1), 在评价时将出苗数与生长发育状况级别相结合, 计算耐盐指数, 可准确地评价大豆种质出苗期的耐盐性。此外, 单株分类记载法只统计处理条件下品种出苗情况的差异性, 不用种植对照也不损伤植株, 从而节约大量的劳动成本, 适用于大批量种质资源的高效鉴定, 也为其他作物耐盐性鉴定提供了借鉴。

3.2 大豆不同发育阶段的耐盐性研究

作物在不同生长发育阶段对盐分的敏感性存在差异[31]。例如, 水稻的芽期耐盐性高于生殖生长期[22]。番茄萌发期与幼苗生长发育阶段的耐盐性不同[32]。同一大豆种质不同发育阶段耐盐性也存在差异, 如大豆栽培品种Lee、Coiquitt和Clark 36受盐胁迫后发芽率降低的程度相似, 而Lee苗期耐盐性高于Coiquitt和Clark 36[15]。研究发现, 大豆萌发期的耐盐性强于出苗期和苗期, 以后随生育期的增进耐盐性增强[11,23]。当NaCl浓度为220 mmol L-1时, 种子的萌发率几乎不受影响, 而此浓度下幼苗生长率较对照下降95%; NaCl提高到330 mmol L-1时, 幼苗的生长完全被抑制, 而种子的萌发率仍可达81%; 当胚轴中的Na+含量为9.3 mg g-1 FW时, 部分种子仍能萌发(40%), 而幼苗的生长则在Na+含量为6.1 mg g-1 FW时受到完全抑制[23]。这些结果表明, 与后期生长相比, 大豆芽期能耐受更高的盐浓度, 种子萌发并不意味着能够正常生长成幼苗。此外, 大田播种通常会在雨后进行, 此时田间土壤的盐度会较低。因此, 出苗期的耐盐性研究对于大豆生产更具有指导意义[21]

目前, 已克隆了大豆苗期耐盐关键基因GmSALT3/ GmCHX1/GmNcl[17-18,33], 发现在盐碱地条件下, 耐盐基因GmSALT3主要通过增加粒重, 使耐盐大豆增产30%~50%[20]。而出苗期耐盐基因挖掘的研究较少。Zhang等[27]利用257个大豆品种与135个SSR标记进行出苗期耐盐性的关联分析, 检测到83个与环境互作的微效QTL (解释的表型贡献率: 1%~5%)。因此, 本研究建立的大豆出苗期耐盐鉴定方法对于筛选耐盐种质资源、发掘耐盐基因、培育耐盐品种具有十分重要的意义。

4 结论

建立了一种以蛭石为基质, 利用150 mmol L-1 NaCl处理15 d的简便快速鉴定方法, 不需要种植对照, 只需统计处理条件下出苗数及每株苗的耐盐等级, 计算耐盐指数作为评价大豆出苗期耐盐性指标。利用该方法筛选到出苗期耐盐大豆种质10份, 为耐盐大豆品种的培育提供基础材料和鉴定方法。

参考文献 原文顺序
文献年度倒序
文中引用次数倒序
被引期刊影响因子

Hamwieh A, Xu D . Conserved salt tolerance quantitative trait loci (QTL) in wild and cultivated soybean
Breed Sci, 2008,58:355-359.

DOI:10.1270/jsbbs.58.355URL [本文引用: 2]

Hamwieh A, Tuyen D D, Cong H, Benitez E R, Takahashi R, Xu D H . Identification and validation of a major QTL for salt tolerance in soybean
Euphytica, 2011,179:451-459.

DOI:10.1007/s10681-011-0347-8URL [本文引用: 3]
To identify quantitative trait loci (QTLs) conditioning salt tolerance in soybean (Glycine max (L.) Merr.), two recombinant inbred line (RIL) populations derived from crosses of FT-Abyara x C01 and Jin dou No. 6 x 0197 were used in this study. The FT-Abyara x C01 population consisted of 96 F-7 RILs, and the Jin dou No. 6 x 0197 population included 81 F-6 RILs. The salt tolerant parents FT-Abyara and Jin dou No. 6 were originally from Brazil and China, respectively. The QTL analysis identified a major salt-tolerant QTL in molecular linkage group N, which accounted for 44.0 and 47.1% of the total variation for salt tolerance, in the two populations. In the FT-Abyara x C01 population, three RILs were found to be heterozygous around the detected QTL region. By selfing the three residual heterozygous lines, three sets of near isogenic lines (NILs) for salt tolerance were developed. An evaluation of salt tolerance of the NILs revealed that all the lines with FT-Abyara chromosome segment at the QTL region showed significantly higher salt tolerance than the lines without the FT-Abyara chromosome segment. Results of the NILs validated the salt tolerance QTL detected in the RIL populations.

Wang Z, Wang J, Bao Y, Wu Y, Zhang H . Quantitative trait loci controlling rice seed germination under salt stress
Euphytica, 2011,178:297-307.

DOI:10.1007/s10681-010-0287-8URL [本文引用: 1]
Salt tolerance of rice (Oryza sativa L.) at the seed germination stage is one of the major determinants for the stable stand establishment in salinity soil. One population of recombinant inbred lines (RILs, F(2:9)), derived from a cross between a japonica rice landrace tolerant to salt stress and a sensitive indica rice variety, was used to determine the germination traits including imbibition rate and germination percentage under control (water) and salt stress (100 mM NaCl) for 10 days at 30 A degrees C. The multiple interval mapping (MIM) were applied to conduct QTL for the traits. The results showed that seed germination was a quantitative trait controlled by several genes, and strongly affected by salt stress. A total of 16 QTLs were detected in this study, and each QTL could explain 4.6-43.7% of the total phenotypic variance. The expression of these QTLs might be developmentally regulated and growth stage-specific. In addition, only one digenic interaction was detected under salt stress, showing small effect on germination percentage with R(2) 2.7%. Among sixteen QTLs detected in this study, four were major QTLs with R(2) > 30%, and some novel alleles of salt tolerance genes in rice. The results demonstrated that the japonica rice Jiucaiqing is a good source of gene(s) for salt tolerance and the major or minor QTLs identified could be used to improve the salt tolerance by marker-assisted selection (MAS) in rice.

王佳丽, 黄贤金, 钟太洋, 陈志刚 . 盐碱地可持续利用研究综述
地理学报, 2011,66:673-684.

DOI:10.11821/xb201105010URL [本文引用: 1]
盐碱地资源的开发利用对粮食生产具有重大贡献。但水资源的缺乏、气候变化使盐碱地开发利用工作更具挑战性。文章系统总结了近年来盐碱地可持续利用研究取得的重要进展:(1)盐碱地可持续利用的技术研发、排盐水环境安全处理及技术配套管理研究;(2) 盐碱地可持续利用的农户技术选择行为研究;(3) 盐碱地可持续利用的科学研究方法。提出盐碱地可持续利用进一步研究的建议:加强多学科交叉融合的研究;强调农户技术选择行为研究;增强研究方法的科学性。
Wang J L, Huang X J, Zhong T Y, Chen Z G . Review on sustainable utilization of salt-affected land
Acta Geogr Sin, 2011,66:673-684 (in Chinese with English abstract).

DOI:10.11821/xb201105010URL [本文引用: 1]
盐碱地资源的开发利用对粮食生产具有重大贡献。但水资源的缺乏、气候变化使盐碱地开发利用工作更具挑战性。文章系统总结了近年来盐碱地可持续利用研究取得的重要进展:(1)盐碱地可持续利用的技术研发、排盐水环境安全处理及技术配套管理研究;(2) 盐碱地可持续利用的农户技术选择行为研究;(3) 盐碱地可持续利用的科学研究方法。提出盐碱地可持续利用进一步研究的建议:加强多学科交叉融合的研究;强调农户技术选择行为研究;增强研究方法的科学性。

Abel G H, Mackenzie A J . Salt tolerance of soybean varieties (Glycine max L. Merrill) during germination and later growth
Crop Sci, 1964,4:14-28.

[本文引用: 3]

Wang D, Shannon M C . Emergence and seedling growth of soybean cultivars and maturity groups under salinity
Plant Soil, 1999,214:117-124.

DOI:10.1023/A:1004719420806URL [本文引用: 3]

Singleton P W, Bohlool B B . Effect of salinity on nodule formation by soybean
Plant Physiol, 1984,74:72-76.

DOI:10.1104/pp.74.1.72URLPMID:16663389 [本文引用: 1]
A split-root growth system was employed to evaluate the effect of NaCl on nodule formation by soybean (Glycine max L. Merr. cv Davis). By applying the salinity stress and rhizobial inoculum to only one-half the root system, the effects of salinity on shoot growth were eliminated in the nodulation process. Rhizobium colonization of inoculated root surfaces was not affected by the salt treatments (0.0, 26.6, 53.2, and 79.9 millimolar NaCl). While shoot dry weight remained unaffected by the treatments, total shoot N declined from 1.26 grams N per pot at 0.0 millimolar NaCl to 0.44 grams N per pot at 79.9 millimolar NaCl. The concentration of N in the shoot decreased from 3.75% N (0.0 millimolar NaCl) to 1.26% N at 79.9 millimolar NaCl. The decrease in shoot N was attributed to a sharp reduction in nodule number and dry weight. Nodule number and weight were reduced by approximately 50% at 26.6 millimolar NaCl, and by more than 90% at 53.2 and 79.9 millimolar NaCl. Nodule development, as evidenced by the average weight of a nodule, was not as greatly affected by salt as was nodule number. Total nitrogenase activity (C(2)H(2) reduction) decreased proportionally in relation to nodule number and dry weight. Specific nitrogenase activity, however, was less affected by salinity and was not depressed significantly until 79.9 millimolar NaCl. In a second experiment, isolates of Rhizobium japonicum from nodules formed at 79.9 millimolar NaCl did not increase nodulation of roots under salt stress compared to nodule isolates from normal media (0.0 millimolar NaCl). Salt was applied (53.2 millimolar NaCl) to half root systems at 0, 4, 12, and 96 hours from inoculation in a third experiment. By delaying the application of salt for 12 hours, an increase in nodule number, nodule weight, and shoot N was observed. Nodule formation in the 12- and 96-hour treatments was, however, lower than the control. The early steps in nodule initiation are, therefore, extremely sensitive to even low concentrations of NaCl. The sensitivity is not related to rhizobial survival and is probably due to the salt sensitivity of root infection sites.

Chen H, Cui S, Fu S, Gai J, Yu D . Identification of quantitative trait loci associated with salt tolerance during seedling growth in soybean (Glycine max L.)
Aust J Agric Res, 2008,59:1086-1091.

DOI:10.1186/1471-2229-13-161URLPMID:24134188 [本文引用: 2]
Field pea (Pisum sativum L.) is a self-pollinating, diploid, cool-season food legume. Crop production is constrained by multiple biotic and abiotic stress factors, including salinity, that cause reduced growth and yield. Recent advances in genomics have permitted the development of low-cost high-throughput genotyping systems, allowing the construction of saturated genetic linkage maps for identification of quantitative trait loci (QTLs) associated with traits of interest. Genetic markers in close linkage with the relevant genomic regions may then be implemented in varietal improvement programs.

Phang T H, Lam H M . Salt tolerance in soybean
J Integr Plant Biol, 2008,50:1196-1212.

DOI:10.1111/j.1744-7909.2008.00760.xURL [本文引用: 3]
Soybean is an important cash crop and its productivity is significantly hampered by salt stress. High salt imposes negative impacts on growth, nodulation, agronomy traits, seed quality and quantity, and thus reduces the yield of soybean. To cope with salt stress, soybean has developed several tolerance mechanisms, including: (i) maintenance of ion homeostasis; (ii) adjustment in response to osmotic stress; (iii) restoration of osmotic balance; and (iv) other metabolic and structural adaptations. The regulatory network for abiotic stress responses in higher plants has been studied extensively in model plants such as Arabidopsis thaliana. Some homologous components involved in salt stress responses have been identified in soybean. In this review, we tried to integrate the relevant works on soybean and proposes a working model to describe its salt stress responses at the molecular level.

Pathan M S, Lee J D, Shannon J G, Nguyen H T. Recent advances in breeding for drought and salt stress tolerance in soybean
In: Jenks M A, Hasegawa P M, Jain S M, eds. Advances in Molecular-breeding Toward Drought and Salt Tolerant Crops. USA: Springer, 2007. pp 739-773.

[本文引用: 1]

邵桂花, 宋景芝, 刘惠令 . 大豆种质资源耐盐性鉴定初报
中国农业科学, 1986,19(6):30-35.

URL [本文引用: 4]

Shao G H, Song J Z, Liu H L . Preliminary studies on the evaluation of salt tolerance in soybean varieties
Sci Agric Sin, 1986,19(6):30-35 (in Chinese with English abstract).

URL [本文引用: 4]

郭蓓, 邱丽娟, 邵桂花, 许占友 . 大豆耐盐性种质的分子标记辅助鉴定及其利用研究
大豆科学, 2002,21:56-61.

[本文引用: 1]

Guo B, Qiu L J, Shao G H, Xu Z Y . Markers-assisted identification of the salt tolerant accessions in soybean
Soybean Sci, 2002,21:56-61 (in Chinese with English abstract).

[本文引用: 1]

Ha B K, Vuong T D, Velusamy V, Nguyen H T, Shannon J G, Lee J D . Genetic mapping of quantitative trait loci conditioning salt tolerance in wild soybean (Glycine soja) PI 483463
Euphytica, 2013,193:79-88.

DOI:10.1007/s10681-013-0944-9URL
Salt tolerance in soybean [Glycine max (L.) Merr.] is controlled by major quantitative trait loci (QTL) or single gene(s). Among soybean germplasm, wild soybean plant introduction PI 483463 was reported to have a single dominant gene for salt tolerance. The objective of this study was to genetically map the QTL in a recombinant inbred line (RIL) population derived from a cross between PI 483463 and Hutcheson. Simple sequence repeat (SSR) markers and universal soybean single nucleotide polymorphism (SNP) panel (the USLP 1.0) were utilized for molecular genotyping. The RILs were phenotyped in two independent tests in a greenhouse using a 1-5 scale visual rating method. The results showed that the salt tolerant QTL in PI 483463 was mapped to chromosome 3 in a genomic region between the Satt255 and BARC-038333-10036 markers. The favorable allele inherited from PI 483463 conferred tolerance to salinity and had an additive effect on reducing leaf scorch. A subset of 66 iso-lines was developed from the F-3 families of the same cross and was used for genetic confirmation of the QTL. The integration of recombination events and the salt reaction data indicate that the QTL is located in the region of approximately a 658 kb segment between SSR03_1335 at nucleotide 40,505,992 and SSR03_1359 at nucleotide 41,164,735 on chromosome 3. This narrow region can facilitate further genomic research for salt tolerance in soybean including cloning salt tolerance genes.

Tuyen D D, Lal S K, Xu D H . Identification of a major QTL allele from wild soybean (Glycine soja Sieb. & Zucc.) for increasing alkaline salt tolerance in soybean
Theor Appl Genet, 2010,121:229-236.

DOI:10.1007/s00122-010-1304-yURL
Salt-affected soils are generally classified into two main categories, sodic (alkaline) and saline. Our previous studies showed that the wild soybean accession JWS156-1 (Glycine soja) from the Kinki area of Japan was tolerant to NaCl salt, and the quantitative trait locus (QTL) for NaCl salt tolerance was located on soybean linkage group N (chromosome 3). Further investigation revealed that the wild soybean accession JWS156-1 also had a higher tolerance to alkaline salt stress. In the present study, an F6 recombinant inbred line mapping population (n=112) and an F2 population (n=149) derived from crosses between a cultivated soybean cultivar Jackson and JWS156-1 were used to identify QTL for alkaline salt tolerance in soybean. Evaluation of soybean alkaline salt tolerance was carried out based on salt tolerance rating (STR) and leaf chlorophyll content (SPAD value) after treatment with 180mM NaHCO3 for about 3weeks under greenhouse conditions. In both populations, a significant QTL for alkaline salt tolerance was detected on the molecular linkage group D2 (chromosome 17), which accounted for 50.2 and 13.0% of the total variation for STR in the F6 and the F2 populations, respectively. The wild soybean contributed to the tolerance allele in the progenies. Our results suggest that QTL for alkaline salt tolerance is different from the QTL for NaCl salt tolerance found previously in this wild soybean genotype. The DNA markers closely associated with the QTLs might be useful for marker-assisted selection to pyramid tolerance genes in soybean for both alkaline and saline stresses.

Essa T A . Effect of salinity stress on growth and nutrient composition of three soybean (Glycine max L. Merrill) cultivars
J Agron Crop Sci, 2002,188:86-93.

DOI:10.1046/j.1439-037X.2002.00537.xURL [本文引用: 1]

Tuyen D D, Zhang H M, Xu D H . Validation and high-resolution mapping of a major quantitative trait locus for alkaline salt tolerance in soybean using residual heterozygous line
Mol Breed, 2013,31:79-86.

DOI:10.1007/s11032-012-9771-2URL

Guan R X, Qu Y, Guo Y, Yu L L, Liu Y, Jiang J H, Chen J G, Ren Y L, Liu G Y, Tian L, Jin L G, Liu Z X, Hong H L, Chang R Z, Gilliham M, Qiu L J . Salinity tolerance in soybean is modulated by natural variation in GmSALT3
Plant J, 2015,80:937-950.

DOI:10.1111/tpj.12695URLPMID:25292417 [本文引用: 3]
The identification of genes that improve the salt tolerance of crops is essential for the effective utilization of saline soils for agriculture. Here, we use fine mapping in a soybean (Glycine max (L.) Merr.) population derived from the commercial cultivars Tiefeng 8 and 85-140 to identify GmSALT3 (salt tolerance-associated gene on chromosome 3), a dominant gene associated with limiting the accumulation of sodium ions (Na+) in shoots and a substantial enhancement in salt tolerance in soybean. GmSALT3 encodes a protein from the cation/H+ exchanger family that we localized to the endoplasmic reticulum and which is preferentially expressed in the salt-tolerant parent Tiefeng 8 within root cells associated with phloem and xylem. We identified in the salt-sensitive parent, 85-140, a 3.78-kb copia retrotransposon insertion in exon 3 of Gmsalt3 that truncates the transcript. By sequencing 31 soybean landraces and 22 wild soybean (Glycine soja) a total of nine haplotypes including two salt-tolerant haplotypes and seven salt-sensitive haplotypes were identified. By analysing the distribution of haplotypes among 172 Chinese soybean landraces and 57 wild soybean we found that haplotype 1 (H1, found in Tiefeng 8) was strongly associated with salt tolerance and is likely to be the ancestral allele. Alleles H2-H6, H8 and H9, which do not confer salinity tolerance, were acquired more recently. H1, unlike other alleles, has a wide geographical range including saline areas, which indicates it is maintained when required but its potent stress tolerance can be lost during natural selection and domestication. GmSALT3 is a gene associated with salt tolerance with great potential for soybean improvement.

Qi X, Li M W, Xie M, Liu X, Ni M, Shao G, Song C, Yim K Y, Tao Y, Wong F L, Isobe S, Wong C F, Wong K S, Xu C, Li C, Wang Y, Guan R, Sun F, Fan G, Xiao Z, Zhou F, Phang T H, Liu X, Tong S W, Chan T F, Yiu S M, Tabata S, Wang J, Xu X, Lam H M . Identification of a novel salt tolerance gene in wild soybean by whole-genome sequencing
Nat Commun, 2014,5:4340.

DOI:10.1038/ncomms5340URLPMID:25004933 [本文引用: 1]
Using a whole-genome-sequencing approach to explore germplasm resources can serve as an important strategy for crop improvement, especially in investigating wild accessions that may contain useful genetic resources that have been lost during the domestication process. Here we sequence and assemble a draft genome of wild soybean and construct a recombinant inbred population for genotyping-by-sequencing and phenotypic analyses to identify multiple QTLs relevant to traits of interest in agriculture. We use a combination of de novo sequencing data from this work and our previous germplasm re-sequencing data to identify a novel ion transporter gene, GmCHX1, and relate its sequence alterations to salt tolerance. Rapid gain-of-function tests show the protective effects of GmCHX1 towards salt stress. This combination of whole-genome de novo sequencing, high-density-marker QTL mapping by re-sequencing and functional analyses can serve as an effective strategy to unveil novel genomic information in wild soybean to facilitate crop improvement.

Do T D, Vuong T D, Dunn D, Smothers S, Patil G, Yungbluth D C, Chen P, Scaboo A, Xu D, Carter T E . Mapping and confirmation of loci for salt tolerance in a novel soybean germplasm, Fiskeby III
Theor Appl Genet, 2018,131:513-524.

DOI:10.1007/s00122-017-3015-0URLPMID:29151146 [本文引用: 1]
The confirmation of a major locus associated with salt tolerance and mapping of a new locus, which could be beneficial for improving salt tolerance in soybean. Breeding soybean for tolerance to high salt conditions is important in some regions of the USA and world. Soybean cultivar Fiskeby III (PI 438471) in maturity group 000 has been reported to be highly tolerant to multiple abiotic stress conditions, including salinity. In this study, a mapping population of 132 F2 families derived from a cross of cultivar Williams 82 (PI 518671, moderately salt sensitive) and Fiskeby III (salt tolerant) was analyzed to map salt tolerance genes. The evaluation for salt tolerance was performed by analyzing leaf scorch score (LSS), chlorophyll content ratio (CCR), leaf sodium content (LSC), and leaf chloride content (LCC) after treatment with 120?mM NaCl under greenhouse conditions. Genotypic data for the F2 population were obtained using the SoySNP6K Illumina Infinium BeadChip assay. A major allele from Fiskeby III was significantly associated with LSS, CCR, LSC, and LCC on chromosome (Chr.) 03 with LOD scores of 19.1, 11.0, 7.7 and 25.6, respectively. In addition, a second locus associated with salt tolerance for LSC was detected and mapped on Chr. 13 with an LOD score of 4.6 and an R 2 of 0.115. Three gene-based polymorphic molecular markers (Salt-20, Salt14056 and Salt11655) on Chr.03 showed a strong predictive association with phenotypic salt tolerance in the present mapping population. These molecular markers will be useful for marker-assisted selection to improve salt tolerance in soybean.

Liu Y, Yu L L, Qu Y, Chen J J, Liu X X, Hong H L, Liu Z X, Chang R Z, Gilliham M, Qiu L J, Guan R X . GmSALT3, which confers improved soybean salt tolerance in the field, increases leaf Cl- exclusion prior to Na+ exclusion but does not improve early vigor under salinity
Front Plant Sci, 2016,7:1485.

DOI:10.3389/fpls.2016.01485URLPMID:27746805 [本文引用: 2]
Soil salinity reduces soybean growth and yield. The recently identified GmSALT3 (Glycine max salt Tolerance-associated gene on chromosome 3) has the potential to improve soybean yields in salinized conditions. Here we evaluate the impact of GmSALT3 on soybean performance under saline or non-saline conditions. Three sets of near isogenic lines (NILs), with genetic similarity of 95.6-99.3% between each pair of NIL-T and NIL-S, were generated from a cross between two varieties 85-140 (salt-sensitive, S) and Tiefeng 8 (salt-tolerant, T) by using marker-assisted selection. Each NIL-T; 782-T, 820-T and 860-T, contained a common ~1000 kb fragment on chromosome 3 where GmSALT3 was located. We show that GmSALT3 does not contribute to an improvement in seedling emergence rate or early vigor under salt stress. However, when 12-day-old seedlings were exposed to NaCl stress, the NIL-T lines accumulated significantly less leaf Na+ compared with their corresponding NIL-S, while no significant difference of K+ concentration was observed between NIL-T and NIL-S; the magnitude of Na+ accumulation within each NIL-T set was influenced by the different genetic backgrounds. In addition, NIL-T lines accumulated less Cl- in the leaf and more in the root prior to any difference in Na+; in the field they accumulated less pod wall Cl- than the corresponding NIL-S lines. Under non-saline field conditions, no significant differences were observed for yield related traits within each pair of NIL-T and NIL-S lines, indicating there was no yield penalty for having the GmSALT3 gene. In contrast, under saline field conditions the NIL-T lines had significantly greater plant seed weight and 100-seed weight than the corresponding NIL-S lines, meaning GmSALT3 conferred a yield advantage to soybean plants in salinized fields. Our results indicated that GmSALT3 mediated regulation of both Na+ and Cl- accumulation in soybean, and contributes to improved soybean yield through maintaining a higher seed weight under saline stress.

Munns R, James R A . Screening methods for salinity tolerance: a case study with tetraploid wheat
Plant Soil, 2003,253:201-218.

DOI:10.1023/A:1024553303144URL [本文引用: 2]
Fast and effective glasshouse screening techniques that could identify genetic variation in salinity tolerance were tested. The objective was to produce screening techniques for selecting salt-tolerant progeny in breeding programs in which genes for salinity tolerance have been introduced by either conventional breeding or genetic engineering. A set of previously unexplored tetraploid wheat genotypes, from five subspecies of Triticum turgidum, were used in a case study for developing and validating glasshouse screening techniques for selecting for physiologically based traits that confer salinity tolerance. Salinity tolerance was defined as genotypic differences in biomass production in saline versus non-saline conditions over prolonged periods, of 3–4 weeks. Short-term experiments (1 week) measuring either biomass or leaf elongation rates revealed large decreases in growth rate due to the osmotic effect of the salt, but little genotypic differences, although there were genotypic differences in long-term experiments. Specific traits were assessed. Na+ exclusion correlated well with salinity tolerance in the durum subspecies, and K+/Na+ discrimination correlated to a lesser degree. Both traits were environmentally robust, being independent of root temperature and factors that might influence transpiration rates such as light level. In the other four T. turgidum subspecies there was no correlation between salinity tolerance and Na+ accumulation or K+/Na+ discrimination, so other traits were examined. The trait of tolerance of high internal Na+ was assessed indirectly, by measuring chlorophyll retention. Five landraces were selected as maintaining green healthy leaves despite high levels of Na+ accumulation. Factors affecting field performance of genotypes selected by trait-based techniques are discussed.

Khatun S, Flowers T J . Effects of salinity on seed set in rice
Plant Cell Environ, 1995,18:61-67.

DOI:10.1007/s10142-010-0203-2URLPMID:21213008 [本文引用: 2]
Glyoxalase pathway, ubiquitously found in all organisms from prokaryotes to eukaryotes, consists of glyoxalase I (GLY I) and glyoxalase II (GLY II) enzymes, which detoxify a cytotoxic molecule, methylglyoxal (MG). Increase in MG has been correlated with various diseases in humans and different abiotic stresses in plants. We have previously shown that overproduction of GLY I and/or GLY II enzymes in transgenic plants provide tolerance towards salinity and heavy metal stresses. We have identified nineteen potential GLY I and four GLY II proteins in rice and twenty two GLY I and nine GLY II proteins in Arabidopsis. An analysis of complete set of genes coding for the glyoxalase proteins in these two genomes is presented, including classification and chromosomal distribution. Expression profiling of these genes has been performed in response to multiple abiotic stresses, in different tissues and during various stages of vegetative and reproductive development using publicly available databases (massively parallel signature sequencing and microarray). AtGLYI8, OsGLYI3, and OsGLYI10 expresses constitutively high in seeds while AtGLYI4, AtGLYI7, OsGLYI6, and OsGLYI11 are highly stress inducible. To complement this analyses, qRT-PCR is performed in two contrasting rice genotypes, i.e., IR64 and Pokkali where OsGLYI6 and OsGLYI11 are found to be highly stress inducible.

Hosseini M, Powell A, Bingham I . Comparison of the seed germination and early seedling growth of soybean in saline conditions
Seed Sci Res, 2002,12:165-172.

DOI:10.1079/SSR2002108URL [本文引用: 3]

邵桂花 . 大豆种质资源耐盐性田间鉴定方法
作物杂志, 1986, (3):36-37.

[本文引用: 6]

Shao G H . Field identification method for salt tolerance of soybean germplasm resources
Crops, 1986, (3):36-37 (in Chinese).

[本文引用: 6]

罗庆云 . 野生大豆和栽培大豆耐盐机理及遗传研究
南京农业大学博士学位论文, 江苏南京, 2003.

[本文引用: 3]

Luo Q Y . Study on Mechanism and Inheritance of Salt Tolerance in Wild Soybean (Glycine soja) and Cultivated Soybean (G. max)
PhD Dissertation of Nanjing Agriculture University, Nanjing, China, 2003 (in Chinese with English abstract).

[本文引用: 3]

张海波, 崔继哲, 曹甜甜, 张佳彤, 刘千千, 刘欢 . 大豆出苗期和苗期对盐胁迫的响应及耐盐指标评价
生态学报, 2011,31:2805-2812.

URL [本文引用: 1]
比较了4个大豆品种出苗期和苗期的耐盐性,测定150 mmol/L NaCl胁迫下的株高、下胚轴长、侧根数、地上干/鲜重、根干/鲜重、MDA含量、SOD活性、游离Pro含量,并将幼苗移栽到田间生长至成熟。结果表明:出苗期和苗期盐胁迫下4个品种的株高都显著降低、地上干/鲜重和根干/鲜重降低;出苗期胁迫侧根数减少,下胚轴长降低;而苗期胁迫侧根数增加,下胚轴长升高。未胁迫条件下,出苗期和苗期耐盐性强的品种22021-1的MDA含量和SOD活性高于耐盐性弱的品种22293-1。胁迫后,22021-1的MDA含量降低、SOD活性升高,其MDA含量分别比对照低51.03%和21.45%,SOD活性比对照高5.85%和45.77%;22293-1的MDA含量出苗期比对照高58.97%,苗期基本无变化,SOD活性出苗期和苗期升高都不显著。MDA和SOD可以作为大豆耐盐性筛选指标。早期的短时胁迫对不同耐盐性大豆品种的经济产量影响不同。
Zhang H B, Cui J Z, Cao T T, Zhang J T, Liu Q Q, Liu H . Response to salt stresses and assessment of salt tolerability of soybean varieties in emergence and seedling stages
Acta Ecol Sin, 2011,31:2805-2812 (in Chinese with English abstract).

URL [本文引用: 1]
比较了4个大豆品种出苗期和苗期的耐盐性,测定150 mmol/L NaCl胁迫下的株高、下胚轴长、侧根数、地上干/鲜重、根干/鲜重、MDA含量、SOD活性、游离Pro含量,并将幼苗移栽到田间生长至成熟。结果表明:出苗期和苗期盐胁迫下4个品种的株高都显著降低、地上干/鲜重和根干/鲜重降低;出苗期胁迫侧根数减少,下胚轴长降低;而苗期胁迫侧根数增加,下胚轴长升高。未胁迫条件下,出苗期和苗期耐盐性强的品种22021-1的MDA含量和SOD活性高于耐盐性弱的品种22293-1。胁迫后,22021-1的MDA含量降低、SOD活性升高,其MDA含量分别比对照低51.03%和21.45%,SOD活性比对照高5.85%和45.77%;22293-1的MDA含量出苗期比对照高58.97%,苗期基本无变化,SOD活性出苗期和苗期升高都不显著。MDA和SOD可以作为大豆耐盐性筛选指标。早期的短时胁迫对不同耐盐性大豆品种的经济产量影响不同。

Zhang W J, Niu Y, Bu S H, Li M, Feng J Y, Zhang J, Yang S X, Odinga M M, Wei S P, Liu X F, Zhang Y M . Epistatic association mapping for alkaline and salinity tolerance traits in the soybean germination stage
PLoS One, 2014,9:e84750.

DOI:10.1371/journal.pone.0084750URLPMID:24416275 [本文引用: 2]
Soil salinity and alkalinity are important abiotic components that frequently have critical effects on crop growth, productivity and quality. Developing soybean cultivars with high salt tolerance is recognized as an efficient way to maintain sustainable soybean production in a salt stress environment. However, the genetic mechanism of the tolerance must first be elucidated. In this study, 257 soybean cultivars with 135 SSR markers were used to perform epistatic association mapping for salt tolerance. Tolerance was evaluated by assessing the main root length (RL), the fresh and dry weights of roots (FWR and DWR), the biomass of seedlings (BS) and the length of hypocotyls (LH) of healthy seedlings after treatments with control, 100 mM NaCl or 10 mM Na2CO3 solutions for approximately one week under greenhouse conditions. A total of 83 QTL-by-environment (QE) interactions for salt tolerance index were detected: 24 for LR, 12 for FWR, 11 for DWR, 15 for LH and 21 for BS, as well as one epistatic QTL for FWR. Furthermore, 86 QE interactions for alkaline tolerance index were found: 17 for LR, 16 for FWR, 17 for DWR, 18 for LH and 18 for BS. A total of 77 QE interactions for the original trait indicator were detected: 17 for LR, 14 for FWR, 4 for DWR, 21 for LH and 21 for BS, as well as 3 epistatic QTL for BS. Small-effect QTL were frequently observed. Several soybean genes with homology to Arabidopsis thaliana and soybean salt tolerance genes were found in close proximity to the above QTL. Using the novel alleles of the QTL detected above, some elite parental combinations were designed, although these QTL need to be further confirmed. The above results provide a valuable foundation for fine mapping, cloning and molecular breeding by design for soybean alkaline and salt tolerance.

Kan G, Zhang W, Yang W, Ma D, Zhang D, Hao D, Hu Z, Yu D . Association mapping of soybean seed germination under salt stress
Mol Genet Genomics, 2015,290:2147-2162.

DOI:10.1007/s00438-015-1066-yURLPMID:26001372
Soil salinity is a serious threat to agriculture sustainability worldwide. Seed germination is a critical phase that ensures the successful establishment and productivity of soybeans in saline soils. However, little information is available regarding soybean salt tolerance at the germination stage. The objective of this study was to identify the genetic mechanisms of soybean seed germination under salt stress. One natural population consisting of 191 soybean landraces was used in this study. Soybean seeds produced in four environments were used to evaluate the salt tolerance at their germination stage. Using 1142 single-nucleotide polymorphisms (SNPs), the molecular markers associated with salt tolerance were detected by genome-wide association analysis. Eight SNP-trait associations and 13 suggestive SNP-trait associations were identified using a mixed linear model and the TASSEL 4.0 software. Eight SNPs or suggestive SNPs were co-associated with two salt tolerance indices, namely (1) the ratio of the germination index under salt conditions to the germination index under no-salt conditions (ST-GI) and (2) the ratio of the germination rate under salt conditions to the germination rate under no-salt conditions (ST-GR). One SNP (BARC-021347-04042) was significantly associated with these two traits (ST-GI and ST-GR). In addition, nine possible candidate genes were located in or near the genetic region where the above markers were mapped. Of these, five genes, Glyma08g12400.1, Glyma08g09730.1, Glyma18g47140.1, Glyma09g00460.1, and Glyma09g00490.3, were verified in response to salt stress at the germination stage. The SNPs detected could facilitate a better understanding of the genetic basis of soybean salt tolerance at the germination stage, and the marker BARC-021347-04042 could contribute to future breeding for soybean salt tolerance by marker-assisted selection.

姬丹丹, 刘畅, 曹其聪, 向凤宁 . 不同大豆品种芽期和苗期耐盐性比较研究
山东轻工业学院学报(自然科学版), 2011,25(2):4-7.

[本文引用: 1]

Ji D D, Liu C, Cao Q C, Xiang F N . Comparison of the salt tolerance at sprout and seedling in different soybean
J Shandong Polytechnic Univ (Nat Sci Edn), 2011,25(2):4-7 (in Chinese with English abstract).

[本文引用: 1]

郭蓓, 邵桂花 . 大豆耐盐基因的PCR标记
中国农业科学, 2000,33(1):10-16.

URL [本文引用: 1]
以大豆耐盐品种和盐敏感品种及“耐盐品种×盐敏感品种”组合的F2群体为试验材料,筛选和鉴定与大豆耐盐性基因紧密连锁的PCR标记,旨在建立快速准确的大豆耐盐性鉴定方法。利用BSA法,对耐(敏)盐品种池和一个组合F2的耐(敏)盐池进行了鉴定,获得一个共显性标记。经F2分析,在盐敏感个体中仅扩增出约600bp的特异片段;在耐盐性个体中扩增出约700bp的特异片段或2个特异片段(700bp/600bp),经过连锁值测定,表明该标记与大豆耐盐基因位点紧密连锁。此外,该标记在其它2个组合F2群体及12个耐盐品种和13个盐敏感品种中得到验证,表明此标记可用于大豆耐盐种质鉴定及大豆耐盐遗传育种的分子标记辅助选择,使大豆耐盐性室内鉴定成为可能。为此,大豆耐盐性基因的分子标记及其获得方法和应用已申请了中华人民共和国发明专利。
Guo B, Shao G H . Tagging salt tolerant gene using PCR marker in soybean
Sci Agric Sin, 2000,33(1):10-16 (in Chinese with English abstract).

URL [本文引用: 1]
以大豆耐盐品种和盐敏感品种及“耐盐品种×盐敏感品种”组合的F2群体为试验材料,筛选和鉴定与大豆耐盐性基因紧密连锁的PCR标记,旨在建立快速准确的大豆耐盐性鉴定方法。利用BSA法,对耐(敏)盐品种池和一个组合F2的耐(敏)盐池进行了鉴定,获得一个共显性标记。经F2分析,在盐敏感个体中仅扩增出约600bp的特异片段;在耐盐性个体中扩增出约700bp的特异片段或2个特异片段(700bp/600bp),经过连锁值测定,表明该标记与大豆耐盐基因位点紧密连锁。此外,该标记在其它2个组合F2群体及12个耐盐品种和13个盐敏感品种中得到验证,表明此标记可用于大豆耐盐种质鉴定及大豆耐盐遗传育种的分子标记辅助选择,使大豆耐盐性室内鉴定成为可能。为此,大豆耐盐性基因的分子标记及其获得方法和应用已申请了中华人民共和国发明专利。

Flowers T J . Improving crop salt tolerance
J Exp Bot, 2004,55:307-319.

DOI:10.1093/jxb/erh003URLPMID:14718494 [本文引用: 1]
Salinity is an ever-present threat to crop yields, especially in countries where irrigation is an essential aid to agriculture. Although the tolerance of saline conditions by plants is variable, crop species are generally intolerant of one-third of the concentration of salts found in seawater. Attempts to improve the salt tolerance of crops through conventional breeding programmes have met with very limited success, due to the complexity of the trait: salt tolerance is complex genetically and physiologically. Tolerance often shows the characteristics of a multigenic trait, with quantitative trait loci (QTLs) associated with tolerance identified in barley, citrus, rice, and tomato and with ion transport under saline conditions in barley, citrus and rice. Physiologically salt tolerance is also complex, with halophytes and less tolerant plants showing a wide range of adaptations. Attempts to enhance tolerance have involved conventional breeding programmes, the use of in vitro selection, pooling physiological traits, interspecific hybridization, using halophytes as alternative crops, the use of marker-aided selection, and the use of transgenic plants. It is surprising that, in spite of the complexity of salt tolerance, there are commonly claims in the literature that the transfer of a single or a few genes can increase the tolerance of plants to saline conditions. Evaluation of such claims reveals that, of the 68 papers produced between 1993 and early 2003, only 19 report quantitative estimates of plant growth. Of these, four papers contain quantitative data on the response of transformants and wild-type of six species without and with salinity applied in an appropriate manner. About half of all the papers report data on experiments conducted under conditions where there is little or no transpiration: such experiments may provide insights into components of tolerance, but are not grounds for claims of enhanced tolerance at the whole plant level. Whether enhanced tolerance, where properly established, is due to the chance alteration of a factor that is limiting in a complex chain or an effect on signalling remains to be elucidated. After ten years of research using transgenic plants to alter salt tolerance, the value of this approach has yet to be established in the field.

Foolad M R, Lin G Y . Absence of genetic relationship between salt tolerance during seed germination and vegetative growth in tomato
Plant Breed, 1997,116:363-367.

DOI:10.1111/pbr.1997.116.issue-4URL [本文引用: 1]

Do T D, Chen H, Hien V T T, Hamwieh A, Yamada T, Sato T, Yan Y, Cong H, Shono M, Suenaga K . Ncl synchronously regulates Na+, K+, and Cl- in soybean and greatly increases the grain yield in saline field conditions
Sci Rep, 2016,6:19147.

DOI:10.1038/srep19147URLPMID:26744076 [本文引用: 1]
Salt stress inhibits soybean growth and reduces gain yield. Genetic improvement of salt tolerance is essential for sustainable soybean production in saline areas. In this study, we isolated a gene (Ncl) that could synchronously regulate the transport and accumulation of Na(+), K(+), and Cl(-) from a Brazilian soybean cultivar FT-Abyara using map-based cloning strategy. Higher expression of the salt tolerance gene Ncl in the root resulted in lower accumulations of Na(+), K(+), and Cl(-) in the shoot under salt stress. Transfer of Ncl with the Agrobacterium-mediated transformation method into a soybean cultivar Kariyutaka significantly enhanced its salt tolerance. Introgression of the tolerance allele into soybean cultivar Jackson, using DNA marker-assisted selection (MAS), produced an improved salt tolerance line. Ncl could increase soybean grain yield by 3.6-5.5 times in saline field conditions. Using Ncl in soybean breeding through gene transfer or MAS would contribute to sustainable soybean production in saline-prone areas.
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