烯效唑对盐胁迫下糜子幼苗形态和生理特性的调控效应

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张盼盼, 杨裕然, 薛佳欣, 王涛, 刘涵, 刘翠英, 冯佰利, 张雄. 烯效唑对盐胁迫下糜子幼苗形态和生理特性的调控效应. 草业学报, 2020,29(10): 81-90
ZHANG Pan-pan, YANG Yu-ran, XUE Jia-xin, WANG Tao, LIU Han, LIU Cui-ying, FENG Bai-li, ZHANG Xiong. Effects of uniconazole on morphology and physiological characteristics of proso millet seedlings under salt stress. Acta Prataculturae Sinica,2020,29(10): 81-90 复制到剪切板

Doi:10.11686/cyxb2020140
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烯效唑对盐胁迫下糜子幼苗形态和生理特性的调控效应

张盼盼¹, 杨裕然¹, 薛佳欣¹, 王涛¹, 刘涵¹, 刘翠英¹, 冯佰利^2,^*, 张雄^1,^*

1.榆林学院生命科学学院,陕西榆林 719000

2.西北农林科技大学农学院,陕西杨凌 712100

*通信作者. E-mail: 7012766@163.com, yulinzhang2007@163.com

作者简介:张盼盼(1985-),女,陕西渭南人,博士,副教授。E-mail: zpp35@163.com

收稿日期: 2020-03-25

基金项目:国家自然科学基金项目(31860340),陕西省自然科学基础研究计划项目(2020JQ-902),陕西省产业技术体系项目,榆林市科技局科技计划项目(2018-cxy-2,2019-106-3)和榆林学院引进人才科研启动项目(17GK21)资助

摘要

为探究叶面喷施烯效唑对糜子幼苗耐盐性的调控作用,以榆糜3号为试验材料,采用砂培方式,研究了三叶一心期叶面喷施烯效唑(50 mg·L^-1)对150 mmol·L^-1 NaCl盐胁迫下糜子幼苗生长和生理特性的影响。结果表明: 喷施烯效唑提高了盐胁迫下糜子幼苗叶片净光合速率、蒸腾速率、气孔导度和胞间CO₂浓度,同时烯效唑增加了盐胁迫下糜子叶片光系统II的最大光能转换效率、实际光化学效率和光化学猝灭系数,降低了叶片PSⅡ的非光化学猝灭系数;盐胁迫下喷施烯效唑处理株高显著降低( P<0.05),叶面积和地上部干重减小,糜子幼苗总根长、根体积、根总表面积、根平均直径和地下部干重增加;盐胁迫下经烯效唑处理的幼苗叶片和根系超氧化物歧化酶(SOD)活性、过氧化物酶(POD)活性、过氧化氢酶(CAT)活性、可溶性蛋白含量均升高;而丙二醛(MDA)含量和超氧阴离子自由基($\mathop{}_{A}^{B}$)含量均下降;其中POD活性和$\mathop{}_{A}^{B}$含量在叶片中无显著差异,而在根系中差异达显著水平( P<0.05)。因此,在三叶一心期叶面喷施50 mg·L^-1烯效唑能够调控糜子幼苗形态特征,增强根系和叶片保护酶活性和可溶性蛋白含量,减缓膜脂过氧化程度,从而有效提高植株的耐盐能力。

关键词: 烯效唑; 糜子; 叶片; 根系; 形态; 生理特性

Effects of uniconazole on morphology and physiological characteristics of proso millet seedlings under salt stress

ZHANG Pan-pan¹, YANG Yu-ran¹, XUE Jia-xin¹, WANG Tao¹, LIU Han¹, LIU Cui-ying¹, FENG Bai-li^2,^*, ZHANG Xiong^1,^*

1.College of Life Science, Yulin University, Yulin 719000, China

2.College of Agronomy, Northweat A&F University, Yangling 712100, China

*Corresponding author. E-mail: 7012766@163.com, yulinzhang2007@163.com

Abstract

The aim of this research was to investigate the effect on seedling morphology and physiological characteristics of foliar uniconazole spraying of ‘Yumi 3’ proso millet seedlings under salt stress. Leaf photosynthetic characteristics, seedling growth and seedling antioxidase systems were measured, following foliar spraying with uniconazole (50 mg·L^-1), in plants grown under 150 mmol·L^-1 NaCl stress in sand culture experiments. The results showed that foliar uniconazole spraying increased net photosynthetic rate, transpiration rate, stomatal conductance and intercellular CO₂ concentration, maximum light energy conversion efficiency, actual photochemical efficiency and photochemical quenching of the seedlings, but decreased non-photochemical quenching coefficient. In seedlings exposed to salt stress and foliar spraying with uniconazole, seedling height decreased significantly ( P<0.05), and leaf area and shoot dry weight also decreased. However, the total root length, root volume, root surface area, root average diameter and root dry weight of seedlings increased. With respect to antioxidase systems, in seedlings treated with uniconazole under salt stress, superoxide dismutase activity, peroxidase (POD) activity, catalase activity and soluble protein content were all increased, while malondialdehyde content and superoxide anion content ($\mathop{}_{A}^{B}$) were decreased. Among these parameters, POD activity and $\mathop{}_{A}^{B}$ levels showed no significant differences in leaves, but significant differences in roots ( P<0.05). Therefore, foliar uniconazole (50 mg·L^-1) spraying at the three-leaf-and-one-leaflet stage of development effectively improved the salt tolerance of proso millet seedlings by regulating the seedling morphological characteristics, enhancing the protective enzyme activity and soluble protein content in the roots and leaves, and decreasing the degree of membrane lipid peroxidation.

Keyword: uniconazole; proso millet; leaf; root; morphology; physiological property

文章图片

近年来, 随着工农业的不断发展、化肥的大量使用, 灌溉和耕作的不合理, 温室效应等因素, 盐渍化程度以每年百万公顷的速度增加^{[1, 2, 3, 4]}。据统计, 目前全世界约有5%的耕地受到不同程度的盐碱化。我国盐碱地面积约1亿hm², 并呈逐渐扩大的趋势^[5]。其中陕西的土壤盐碱化形势也非常严峻, 以陕北榆林盐碱地为主^[6]。土壤盐碱化已成为当地农业发展和土地开发利用的重要限制因素。如何合理开发利用盐碱地资源, 保障该区粮食安全及农业经济发展是迫切需要解决的问题。

糜子(Panicum miliaceum), 为禾本科黍属, 原产于我国最古老的农作物, 在小杂粮作物中占有重要的地位, 栽培历史悠久^{[7, 8]}, 在长期进化过程中积累了生长发育期短, 抗逆性强、耐贮性强等特点^{[9, 10]}。糜子是新垦土地的先锋植物和理想的复种植物, 在我国农业救灾备荒中发挥着重要的作用, 同时又是陕北糜子产区重要的饲草。然而该区盐碱化严重地制约着糜子产业的发展。因此, 通过合适的栽培技术改善糜子的耐盐能力, 提高糜子生产水平, 对增加产区农民收入, 实现农业增效具有重要的意义。

烯效唑(uniconazole, S3307)是一种具有较高生物活性和使用安全的植物生长延缓剂^[11], 其具有提高作物抗逆性的重要作用, 前人关于烯效唑对作物的影响多集中于小麦(Triticum aestivum)^[12]、玉米(Zea mays)^[13]、水稻(Oryza sativa)^[14]、油菜(Brassica napus)^[15]等主要农作物生产, 而有关叶面喷施烯效唑对盐胁迫下糜子幼苗生长和生理特性方面调控效应的研究尚未见相关报道。鉴于此, 本研究以榆糜3号为试验材料, 以烯效唑为处理药剂, 进行糜子抗盐性试验, 探讨烯效唑对糜子幼苗形态和生理特性的影响, 为植物生长调节剂的应用及糜子高产栽培提供理论依据。

1 材料与方法

1.1 供试材料

供试糜子品种为榆糜3号, 由西北农林科技大学国家小宗粮豆研究中心提供, 该品种幼苗和茎秆绿色, 生育期102 d, 单株粒重7.6 g, 千粒重8.0~8.6 g。供试烯效唑药剂由榆林学院生命科学学院农业节水实验室提供, 喷施浓度为50 mg· L^-1。

1.2 试验设计

试验于2017年8-12月在榆林学院生命科学学院农业节水实验室中进行。本试验采用完全随机设计, 设置4个处理, 分别为:CK, 0 mmol· L^-1NaCl(1/2Hogland营养液配置)+叶面喷施0 mg· L^-1烯效唑溶液; S, 150 mmol· L^-1 NaCl(1/2Hogland营养液配置)+叶面喷施0 mg· L^-1烯效唑溶液; T₁, 0 mmol· L^-1NaCl(1/2Hogland营养液配置)+叶面喷施50 mg· L^-1烯效唑溶液; T₂, 150 mmol· L^-1NaCl (1/2Hogland营养液配置)+叶面喷施50 mg· L^-1烯效唑溶液, 每个处理重复10次。试验所用150 mmol· L^-1NaCl和50 mg· L^-1烯效唑是结合前人文献及经过前期试验筛选出的浓度。盆栽试验采用上口径29.5 cm、下口径20.5 cm、高26.5 cm的聚乙烯塑料盆进行, 每盆装经过过筛的沙子4 kg。用蒸馏水将大小一致、健壮饱满的糜子种子按体积比10: 1浸种, 放置于25 ℃培养箱中, 期间摇晃2~3次, 使得种子与溶液充分接触, 24 h后用自来水清洗种子3~4遍, 阴干后, 采用打孔方式播种, 播深3 cm, 每盆播种25粒, 放置于人工气候室, 昼/夜(25± 1) ℃/(18± 1) ℃, 每天光照16 h, 黑暗8 h, 光强为1000 μ mol· m^-2· s^-1, 相对湿度为60%~80%; 播种后所有盆每3 d补充适量且等量浇水以保证种子顺利出苗; 待苗齐后, 定苗, 每盆留取长势一致的幼苗15株, 并开始浇灌1/2Hogland营养液, 以后每3 d浇灌1次, 直至三叶一心期, 将40盆幼苗随机分成4组, 开始进行处理。盐胁迫时NaCl溶液浇透, 托盘中过量的溶液倒掉, 叶面喷施烯效唑处理以叶面挂满药剂但不聚合成水流为宜。以后每3 d补充缺少的1/2Hogland营养液。于盐胁迫处理后9 d, 进行相关指标测定。

1.3 测定项目与方法

1.3.1 光合参数的测定于每个处理中选取长势一致的幼苗4株, 利用Li-6400光合测定系统测定植株最上面完全展开叶的净光合速率(net photosynthetic rate, P_n)、气孔导度(stomatal conductance, G_s)、蒸腾速率(transpiration rate, T_r)、胞间CO₂浓度(intercellular CO₂ concentration, C_i)等光合参数。

1.3.2 叶绿素荧光参数的测定在测量植物叶片光合参数同一天的上午, 使用PM2500便携式调制叶绿素荧光仪测量植株同一叶片的非光化学猝灭系数(non-photochemical quenching coefficient, NPQ)、实际光化学效率(actual photochemical efficiency, Φ _PSⅡ)、光化学猝灭(photochemical quenching, qP)、最大光能转换效率(maximum light energy conversion efficiency, F_v/F_m)等叶绿素荧光参数指标, 每个处理重复4次。

1.3.3 幼苗株高和叶面积的测定从每个处理中选取长势一致的6株糜子, 用钢尺测量植株基部地面处至生长点的长度, 即为株高(cm); 同时, 测量第一片完全展开叶(由上至下)的叶长、叶宽, 根据叶面积计算公式叶长× 叶宽× 0.75求幼苗全展叶叶面积(cm²)。

1.3.4 单株干重的测定每个处理随机抽取6株, 将待测幼苗用蒸馏水冲洗干净后, 地上部分和地下部分分装, 在105 ℃下“ 杀青” 30 min后再在75 ℃下烘干至恒重, 取出纸袋和材料, 放入干燥器中冷却至室温, 用万分之一天平分别称取地上部干重和地下部干重。

1.3.5 根系形态指标的测定每个处理选取长势一致的幼苗6株, 将根从节上剪下放入装有无色透明含有去离子水的塑料水槽, 用镊子调整根的位置, 避免交叉重叠, 使用扫描仪(EPSON PERFECTIONTM4990 PHOTO型)对根样进行扫描, 扫描仪的分辨率设为300 dpi。用根系分析软件(Win RHIZO)对扫描图像进行分析, 得到总根长度、表面积、体积、根平均直径等数据。

1.3.6 幼苗生理指标的测定根系和叶片可溶性蛋白含量采用考马斯亮蓝G-250染色法测定, 超氧化物歧化酶(superoxide dismutase, SOD)活性采用NBT光化还原法测定, 过氧化氢酶(catalase, CAT)活性采用紫外吸收法测定, 超氧阴离子自由基含量采用羟胺氧化法测定^[16]。根系和叶片过氧化物酶(peroxidase, POD)活性参照杨丽涛等^[17]介绍的愈创木酚比色法测定, 并稍做修改。根系和叶片丙二醛(malondialdehyde, MDA)含量参照高俊凤^[18]的硫代巴比妥酸比色法测定, 并稍做修改。以上指标均为3次重复, 取平均值。

1.4 数据处理

数据由Microsoft Excel 2013 软件进行录入、整理, 结果用平均值± 标准差表示, 用SPSS 20.0对数据进行均值、标准差和差异显著性分析, 并采用Origin 9.1软件绘图。

2 结果与分析

2.1 S3307对盐胁迫下糜子幼苗叶片光合特性的影响

如表1所示, 当植株受到盐胁迫时(S), 叶片P_n、G_s、C_i和T_r均显著降低, 较CK分别降低了31.81%、56.22%、23.78%和57.14%; 在正常水分环境条件下叶面喷施烯效唑(T₁)对叶片G_s无明显影响, 叶片C_i和T_r呈升高的趋势, 而叶片P_n呈降低的趋势; 与S相比, T₂处理叶片P_n、G_s、C_i和T_r均呈显著升高的趋势, 分别升高了12.69%、40.20%、13.40%和53.85%。

表1 S3307对盐胁迫下糜子幼苗叶片光合特性的影响 Table 1 Effects of S3307 on photosynthetic characteristics of proso millet seedlings under salt stress

2.2 S3307对盐胁迫下糜子幼苗叶片叶绿素荧光特性的影响

如表2所示, 当植株受到盐胁迫(S)时, 叶片F_v/F_m、Φ _PSⅡ和qP较CK均显著降低, 分别降低了22.22%、25.00%和43.24%, 而叶片NPQ反而升高了27.78%; 与CK相比, 在正常水分环境条件下叶面喷施烯效唑(T₁)均可显著提高F_v/F_m、Φ _PSⅡ和qP, 而对NPQ无明显影响; 与S相比, T₂处理叶片F_v/F_m、Φ _PSⅡ和qP均呈显著升高的趋势, 分别升高了12.50%、22.22%和19.05%, 而叶片NPQ显著降低了13.04%。

表2 S3307对盐胁迫下糜子幼苗叶片叶绿素荧光特性的影响 Table 2 Effects of S3307 on chlorophyll fluorescence characteristics of proso millet seedlings under salt stress

2.3 S3307对盐胁迫下糜子幼苗形态特征的影响

由表3可知, 在盐胁迫处理(S)下, 糜子幼苗株高显著降低, 而叶面积无明显变化; 在正常水分环境条件下喷施烯效唑(T₁), 糜子幼苗株高有大幅度的降低, 而叶面积有小幅度的增加; 与S相比, 在盐胁迫下叶面喷施烯效唑(T₂), 糜子株高和叶面积均有所降低, 分别降低了25.30%和25.62%。

表3 S3307处理对盐胁迫下糜子幼苗形态特征的影响 Table 3 Effects of S3307 on morphological characteristics of proso millet seedlings under salt stress

在盐胁迫条件(S), 糜子幼苗根系总根长、根总表面积、根体积和根平均直径均呈降低的趋势, 分别降低了5.04%、32.09%、23.03%和6.40%; 与CK相比, 在正常水分条件下喷施烯效唑(T₁), 糜子幼苗根系总根长、根总表面积、根体积和根平均直径均呈上升的趋势, 分别升高了70.78%、61.19%、19.58%和6.40%; 与S相比, 在盐胁迫下喷施烯效唑(T₂), 糜子幼苗根系总根长、根总表面积、根体积和根平均直径均升高(图1), 分别升高了39.22%、118.68%、45.14%和9.40%。

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	图1 不同处理下糜子幼苗扫根图Fig.1 Root scanning of proso millet seedlings under different treatments

2.4 S3307对盐胁迫下糜子幼苗干重的影响

由图2可知, 仅盐胁迫下(S)糜子幼苗地上部和地下部分干重较正常水分条件均减小, 其中地上部干重在CK与S之间存在显著差异(P< 0.05); 与CK相比, 在正常水分环境条件下喷施烯效唑(T₁), 糜子幼苗地上部干重无明显变化, 而植株地下部干重有小幅度的增加; 与S相比, 在盐胁迫下喷施烯效唑(T₂)处理下地上部干重降低, 而地下部干重增加, 但地上部干重和地下部干重在S和T₂之间均无显著差异。

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	图2 S3307对盐胁迫下糜子幼苗地上部及地下部干重的影响不同小写字母表示不同处理差异显著(P< 0.05)。下同。Fig.2 Effect of S3307 on shoot dry weight and root dry weight of proso millet seedlings under salt stress Different letters indicate a significant difference at P< 0.05 among different treatments. The same below.

2.5 S3307对盐胁迫下糜子幼苗叶片和根系保护酶活性的影响

由图3可知, 当植株受到盐胁迫时(S), 糜子幼苗叶片和根系SOD活性有大幅度的增加; POD活性有小幅度的提高; 叶片CAT活性下降, 而根系CAT活性显著上升。在正常水分环境条件下喷施烯效唑(T₁), 可以提高糜子幼苗叶片和根系SOD活性, 分别提高了10.12%和85.26%; 叶片POD活性有小幅度的降低, 而根系POD活性显著升高; 糜子幼苗叶片和根系CAT活性均显著上升, 分别升高了33.63%和96.35%。与S相比, 在盐胁迫下喷施烯效唑(T₂)叶片和根系SOD活性无明显变化; 糜子幼苗叶片和根系POD活性均有所升高, 分别升高了1.36%和83.61%; 糜子幼苗叶片和根系CAT活性均显著升高, 分别提高了142.22%和102.33%。

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	图3 S3307对盐胁迫下糜子幼苗叶片和根系保护酶活性的影响Fig.3 Effect of S3307 on activities of protective enzymes in leaves and roots of proso millet seedlings under salt stress

2.6 S3307对盐胁迫下糜子幼苗叶片和根系MDA和

{O_{2}}^{- \cdot}

含量的影响

由图4可知, 在盐胁迫条件下, 糜子幼苗叶片和根系MDA含量均有所增加; 在正常盐分环境下喷施烯效唑(T₁), 糜子幼苗叶片和根系MDA含量均显著降低; 与S相比, 在盐胁迫下喷施烯效唑(T₂), 可有效降低叶片和根系MDA含量。

	Figure Option View Download New Window
	图4 S3307对盐胁迫下糜子幼苗丙二醛和超氧阴离子自由基含量的影响Fig.4 Effect of S3307 on malondialdehyde and superoxide anion content of proso millet seedlings under salt stress

当植株受到150 mmol· L^-1 NaCl盐胁迫(S)时, 糜子幼苗叶片和根系内 ${O_{2}}^{- \cdot}$ 含量均显著升高; 在正常水分环境条件下喷施烯效唑(T₁), 糜子幼苗叶片 ${O_{2}}^{- \cdot}$ 含量有所升高, 而根系内 ${O_{2}}^{- \cdot}$ 含量降低了28.48%; 与S相比, 在盐胁迫下喷施烯效唑(T₂), 糜子幼苗叶片和根系 ${O_{2}}^{- \cdot}$ 含量均有所降低, 且根系 ${O_{2}}^{- \cdot}$ 含量甚至接近于正常环境(CK)。

2.7 S3307对盐胁迫下糜子幼苗叶片和根系可溶性蛋白含量的影响

由图5可知, 与CK相比, 盐胁迫处理下(S), 糜子叶片和根系可溶性蛋白质累积量均增加; 在正常水分环境条件下, 喷施烯效唑(T₁)对叶片可溶性蛋白含量无明显影响, 而根系可溶性蛋白含量显著增加(P< 0.05); 与S相比, 在盐胁迫下喷施烯效唑(T₂), 叶片和根系可溶性蛋白含量均显著增加, 分别增加了3.81%和2.84%。

	Figure Option View Download New Window
	图5 S3307对盐胁迫下糜子幼苗叶片和根系可溶性蛋白含量的影响Fig.5 Effect of S3307 on soluble protein content in leaves and roots of proso millet seedlings under salt stress

3 讨论

光合作用是影响作物生长的重要方面^[19], 光合速率反映叶片合成和供应同化物的能力, 光合速率越大, 作物生产能力越强^[20]。前人研究指出, 喷施烯效唑可以有效调控作物叶片光合性能, 提高光能利用率^[21]。叶绿素荧光可以作为光合作用研究的探针, 它不仅能反映光能吸收、激发能传递和光化学反应等光合作用的原初反应过程, 而且与电子传递、质子梯度的建立及ATP合成和CO₂固定等过程有关^{[22, 23]}。孟娜等^[24]研究表明, 烯效唑能够显著提高盐胁迫下黄13大豆(Glycine max)品种叶片光合色素含量。于奇等^[25]研究表明, 叶面喷施烯效唑能够显著提高淹水胁迫下绿豆叶片的P_n、G_s、C_i、T_r。本研究表明, 盐胁迫下糜子叶片P_n、G_s、C_i、T_r、F_v/F_m、Φ _PSⅡ和qP均显著降低; 盐胁迫下喷施烯效唑处理下, 糜子叶片P_n、G_s、C_i、T_r、F_v/F_m、Φ _PSⅡ和qP等光合特性指标较盐胁迫处理均提高, 而叶面积和NPQ均降低。说明盐胁迫抑制糜子的光合作用, 而叶面喷施烯效唑能够通过调控光合参数缓解盐胁迫对糜子的伤害。

根系是吸收水分和矿质营养的主要器官, 根的生长情况和活力水平直接影响地上部的生长发育, 同时, 根系还是最先感受和响应盐胁迫的器官。根系代谢的变化不仅可以调节根系本身的功能, 而且影响植株的生理代谢。大量研究表明^{[26, 27]}, 盐胁迫对植物的生长发育影响总体表现为抑制作用, 具体表现为根系的伸长受到抑制, 鲜重的增加和干物质的积累、侧根的减少。本试验研究表明, 在盐胁迫下, 糜子幼苗根系的根总长、根平均直径、根总表面积、根体积和根干重均明显降低, 这与慈敦伟等^[28]在花生(Arachis hypogaea)幼苗上的研究结果相一致。同时, 盐胁迫下采用叶面喷施烯效唑可以明显增加糜子幼苗的根总长、根平均直径、根总表面积和根体积, 这与蒋玉香^[29]在黄瓜(Cucumis sativus)上的研究结果相一致。

植物对盐胁迫的响应机制是一个复杂的过程, 盐胁迫时, 糜子体内的活性氧代谢平衡被破坏, ${O_{2}}^{- \cdot}$ 的积累可引起膜脂过氧化, 阻碍正常的生理功能。高含量的可溶性蛋白可以维持作物细胞的低渗透势, 并抵抗盐胁迫造成的危害。SOD、POD、CAT是植物体内酶促防御系统的重要保护酶^{[30, 31]}, 在生物体中的水平高低意味着生物体的衰老与死亡。宋胜等^[32]研究表明, 在烯效唑0~50 mg· L^-1浓度范围内, 随着烯效唑浓度的升高, 大豆根系SOD活性和POD活性增强。本研究结果表明, 与CK对比, 盐胁迫下糜子根系内SOD活性、POD活性和CAT活性均有小幅度的升高, 叶片SOD活性和POD活性有小幅度的升高, 叶片和根系可溶性蛋白含量有少量增加, 但与此同时, 叶片和根系内 ${O_{2}}^{- \cdot}$ 含量有明显的增加, 这与张军等^[33]在小麦上的研究结果一致。当盐胁迫下叶面喷施50 mg· L^-1烯效唑后, 叶片中CAT活性、可溶性蛋白含量和根系中SOD活性、POD活性、CAT活性、可溶性蛋白含量均显著升高, 而叶片中MDA含量和根系中 ${O_{2}}^{- \cdot}$ 含量、MDA含量显著降低, 表明喷施烯效唑可以提高植株体内抗氧化酶的活性, 减少 ${O_{2}}^{- \cdot}$ 和MDA的积累, 降低其对细胞膜的破坏, 从而增强植株对盐胁迫的抵抗能力, 与李宁毅等^[34]在矮牵牛(Petunia hybrida)上的研究结果基本一致。其原因可能是烯效唑正向调控内源激素ABA水平增加^[35], 从而促进植物体抗氧化酶活性提高, 并进一步影响植物体内自由基的合成, 降低MDA积累量^[36], 最终增加植物抗逆能力。

4 结论

烯效唑可以提高盐胁迫下糜子幼苗叶片的光合作用, 有效保护叶片光合机构的PSⅡ , 促进根系形态发育, 增强叶片和根系保护酶活性和可溶性蛋白含量, 减缓膜脂过氧化程度, 从而提高糜子幼苗耐盐能力。因此, 在糜子盐碱地种植中, 可以在幼苗三叶一心期进行叶面喷施50 mg· L^-1烯效唑来促进糜子生长。

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2003

0.0

... 近年来,随着工农业的不断发展、化肥的大量使用,灌溉和耕作的不合理,温室效应等因素,盐渍化程度以每年百万公顷的速度增加^[1,2,3,4] ...

2005

0.0

... 近年来,随着工农业的不断发展、化肥的大量使用,灌溉和耕作的不合理,温室效应等因素,盐渍化程度以每年百万公顷的速度增加^[1,2,3,4] ...

2008

0.0

... 近年来,随着工农业的不断发展、化肥的大量使用,灌溉和耕作的不合理,温室效应等因素,盐渍化程度以每年百万公顷的速度增加^[1,2,3,4] ...

1983

0.0

... 近年来,随着工农业的不断发展、化肥的大量使用,灌溉和耕作的不合理,温室效应等因素,盐渍化程度以每年百万公顷的速度增加^[1,2,3,4] ...

2015

0.0

. 2015, 31(17):121-127

Development and experiment on protective blanket for soil salination control

土壤盐渍化治理防护毯的研发及试验

Mao H T , Huang Q H , Long S J

毛海涛, 黄庆豪, 龙顺江

土壤盐碱化问题是中国农业面临的一个巨大挑战，严重制约着农业生产的发展。为探索治理土壤盐渍化的方法，该文基于干旱区5类代表性土壤（砾砂、粉土/粉砂、粉土/粉细砂、粉土和亚黏土）的盐碱化规律，以盐碱土形成原因为切入点，结合土壤水的毛细管作用，以海绵（强力吸水层）、吸热聚乙烯编织布（蒸散加强层）为主要材料，设计出一种盐碱地治理防护毯，并对防护毯吸盐的深度和效果进行试验研究。结果表明：5类土壤表面的积盐情况各不相同，粗颗粒土（砾砂及砂土）含量较大时，在强蒸发条件下，3～4 d土样表面积盐后形成了一层厚约3～4 mm"盐痂"；细颗粒土（亚黏土和粉土）含量相对较多时，短期内表面积盐速度相对较慢且现象也不明显，10～12 d后，土样表面可观察到一层细密的晶体盐颗粒。所以，土壤岩性质地与表层积盐量关系密切。总体趋势是：土壤颗粒越粗，土壤表层在早期的积盐量越大；土壤颗粒越细，土壤表层在后期的总体积盐量越大。在干燥通风和室外温度25℃的条件下，防护毯针对上述不同类型土壤进行吸盐试验，在4 d后0～10 cm土层的含盐量（质量分数，下同）均可减少80%（P＜0.05）以上；＞10～20 cm土层至少减少50%（P＜0.05）以上；对于砂性土来说，＞20～30 cm土层含盐量减少了50%左右（P＜0.05），而黏性土和壤土可减少20%～30%以上（P＜0.05）。对于实际中的盐碱土而言，含盐量减少30%时，土壤的肥力和可耕作程度会明显改善，能有效减小盐碱化对作物生长的影响。此外，研究还发现，铺设防护毯还能改变盐碱地表层土壤的结构，增强土壤的透气性。防护毯最佳使用时间应为土壤毛细管作用最强烈的时间段。该研究提供了一种有效的盐碱地治理措施。

... 我国盐碱地面积约1亿hm²,并呈逐渐扩大的趋势^[5] ...

1992

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... 其中陕西的土壤盐碱化形势也非常严峻,以陕北榆林盐碱地为主^[6] ...

2016

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... 糜子(Panicum miliaceum),为禾本科黍属,原产于我国最古老的农作物,在小杂粮作物中占有重要的地位,栽培历史悠久^[7,8],在长期进化过程中积累了生长发育期短,抗逆性强、耐贮性强等特点^[9,10] ...

2015

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. 2015, 48(16):3121-3131 DOI:10.3864/j.issn.0578-1752.2015.16.003

Analysis of genetic diversity and structure of proso millet core germplasm

糜子骨干种质遗传多样性和遗传结构分析

Dun J L , Wang H G , Chen L

董俊丽, 王海岗, 陈凌

1 Institute of Crop Germplasm Resources of Shanxi Academy of Agricultural Sciences/Key Laboratory of Crop Gene Resources and Germplasm Enhancement on Loess Plateau, Ministry of Agriculture/Shanxi Key Laboratory of Genetic Resources and Genetic Improvement of Minor Crops, Taiyuan 030031 2 Shanxi University, Taiyuan 030006

【Objective】 The objective of this study is to understand the genetic diversity of proso millet germplasm from different regions and to provide an important theoretical foundation for crossbreeding, excavating drought genes and germplasm creation for future, because of its characters, such as short growth period, bearing drought and barren land. 【Method】Genetic diversity detection was done for proso millet resources by phenotypic identification and SSR markers. Distribution of nine phenotypic traits, including plant height, main spike length, leaf length, blade width, number of main stem section, the main stem diameter, spike weight per plant, grain weight per plant and thousand kernel weight were analyzed by subordinate function. Genetic diversity analysis, correlation analysis and principal component analysis of phenotypic traits were done by software DPS7.05; comprehensive evaluation on proso millet germplasm. Good genomic DNA was extracted from young leaves of 96 proso millet germplasm resources from different areas following the CTAB method, and then were amplified by simple sequence repeat molecular markers to analyze genetic diversity and genetic structure. Separation of the amplified fragments was performed on 8% denaturing polyacrylamide gels. The gels were stained with AgNO3 for visualizing the SSR fragments. The No. of allele (A), Major allele frequency(M), Gene diversity(He), polymorphism information content(PIC) were estimated by software PowerMaker 3.25. Neighbor-joined cluster analysis was aslo carried by software PowerMaker 3.25. Population genetic structure was analyzed by software Structure 2.3.1.【Result】Through genetic diversity analysis of the nine phenotypes of proso millet, it was known that the spike weight and grain weight per plant of genetic variation were the most abundant. Phenotypic traits were more concentrated and most of which were apparently interrelated. Genetic diversity of phenotypic traits of germplasm from Shanxi was the richest. Genetic diversity of phenotypic traits was different in different provinces. Principal component analysis and comprehensive evaluation were done for 9 phenotypic traits of 96 copies of proso millet germplasm resources. Comprehensive evaluation value of Neimi No.1 was the lowest, while that of Ningmi No.15 was the highest. Genetic diversity of 96 copies of proso millet germplasm resources was detected by 19 pairs of SSR primers and 112 allelic variation genes were found. The number of alleles per locus ranged from 3 to 9 and the average was 5.9, the average major allele frequency was 0.7045, the average gene diversity index was 0.4097, and the average percentage of polymorphism information content sites was 39.2%. Genetic diversity analysis of germplasm from different regions showed a close relationship among the regions. Genetic diversity index and polymorphic information content percentage of proso millet germplasm resources in Shanxi were 0.357 and 33.01%, respectively. Experimental materials were divided into three groups by genetic structure analysis based on modules and cluster analysis based on genetic distance. Results of these two division methods have similarity. Resources in three groups were connected with geographical environment closely.【Conclusion】 Proso millet germplasm resources were analyzed systematically from aspects of phenotype and molecule. The results showed that proso millet is rich in genetic variation and has a high diversity. What’s more, genetic diversity of proso millet germplasm resources in Shanxi is the most abundant.

【目的】糜子生育期短、耐干旱、耐瘠薄、水分利用效率高,了解糜子资源的遗传多样性和遗传结构,为今后糜子杂交育种、种质创新、挖掘抗旱基因及资源的高效利用提供理论依据。【方法】采用表型鉴定和SSR分子标记对糜子资源进行遗传多样性检测。利用模糊隶属函数法分析糜子种质的株高、主穗长、叶片长、叶片宽、主茎节数、主茎粗、单株穗重、单株粒重和千粒重9个表型性状的分布情况。利用DPS7.05软件进行表型性状的遗传多样性分析、相关性分析和主成分分析,综合评价糜子种质资源的优劣。利用CTAB法提取糜子嫩叶基因组DNA,并利用SSR分子标记技术对不同地区的96份糜子种质资源的基因组DNA进行PCR扩增,后经8%聚丙烯酰胺凝胶电泳分离,银染后显色。利用PowerMaker 3.25软件计算每对引物的等位基因数（A）、主要等位基因频率（M）、基因多样性指数（He）和多态性信息含量指数（PIC）,并进行N-J遗传距离的统计分析；利用Structure 2.3.1分析群体遗传结构。【结果】糜子表型遗传多样性分析表明：9个表型性状分布集中,且绝大部分呈极显著相关；单株粒重和单株穗重遗传变异最丰富,不同省份的资源在表型性状上表现出不同的遗传多样性,山西资源表型遗传多样性最丰富。采用主成分分析法和综合评价法表明,内糜1号的综合性状表现最差,宁糜15号的综合性状表现最好。采用19对SSR引物对96份糜子种质资源进行遗传多样性分析,共检测出112个等位变异；每个位点的等位变异数为3—9个,平均5.9个；平均主要等位基因频率为0.7045；平均基因多样性指数为0.4097；平均多态性信息含量位点百分数为39.2%。不同地理来源糜子种质资源的遗传多样性分析表明,各省份间糜子资源的亲缘关系均较近；山西省资源的基因多样性指数及多态性信息含量百分数最高,分别为0.357和33.01%。基于模型的遗传结构分析和基于遗传距离的聚类分析将试验材料划分为3个类群,两种分类结果有一定相似性,皆与生态环境密切相关。【结论】糜子遗传变异较为丰富,遗传多样性高,尤其是山西糜子资源的遗传多样性最丰富；不同地理来源的糜子种质资源亲缘关系均较近,且其遗传多样性与生态环境密切相关。

2012

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2015

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2008

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... 烯效唑(uniconazole, S3307)是一种具有较高生物活性和使用安全的植物生长延缓剂^[11],其具有提高作物抗逆性的重要作用,前人关于烯效唑对作物的影响多集中于小麦(Triticum aestivum)^[12]、玉米(Zea mays)^[13]、水稻(Oryza sativa)^[14]、油菜(Brassica napus)^[15]等主要农作物生产,而有关叶面喷施烯效唑对盐胁迫下糜子幼苗生长和生理特性方面调控效应的研究尚未见相关报道 ...

2004

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. 2004, 30(5):502-506

Effects of uniconazole waterless-dressing seed on yield of wheat

烯效唑干拌种对小麦的增产作用

Yang W Y , Yu Z W , Yu S L

杨文钰, 于振文, 余松烈

College of Agronomy, Sichuan Agricultural University, Ya’an 625014, Sichuan

The effects of uniconazole(S-3307) waterless-dressing seed on yield of wheat under different varieties,sowing dates, planting densities and nitrogen application levels were studied from 1996 to 2001. The results showed that S-3307 20 mg per kg seed was the best treatment to increase yield. The effec

研究了不同品种、不同播期、不同密度和不同施氮条件下烯效唑干拌种对小麦的产量效应。结果表明，烯效唑干拌种提高了不同品种、播期、密度和施氮量下小麦的产量，以20 mg/kg的效果最好，烯效唑的平均增产效应在品种间差异不显著，早、迟播时增产效应更大，适宜浓度的烯效唑（10～20 mg/kg）与较高密度的基本苗（180～270×10

1996

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2006

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2005

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2000

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... 6 幼苗生理指标的测定根系和叶片可溶性蛋白含量采用考马斯亮蓝G-250染色法测定,超氧化物歧化酶(superoxide dismutase,SOD)活性采用NBT光化还原法测定,过氧化氢酶(catalase,CAT)活性采用紫外吸收法测定,超氧阴离子自由基含量采用羟胺氧化法测定^[16] ...

1990

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... 根系和叶片过氧化物酶(peroxidase,POD)活性参照杨丽涛等^[17]介绍的愈创木酚比色法测定,并稍做修改 ...

2006

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... 根系和叶片丙二醛(malondialdehyde,MDA)含量参照高俊凤^[18]的硫代巴比妥酸比色法测定,并稍做修改 ...

2001

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... 3 讨论光合作用是影响作物生长的重要方面^[19],光合速率反映叶片合成和供应同化物的能力,光合速率越大,作物生产能力越强^[20] ...

2014

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... 3 讨论光合作用是影响作物生长的重要方面^[19],光合速率反映叶片合成和供应同化物的能力,光合速率越大,作物生产能力越强^[20] ...

2018

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... 前人研究指出,喷施烯效唑可以有效调控作物叶片光合性能,提高光能利用率^[21] ...

2014

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... 叶绿素荧光可以作为光合作用研究的探针,它不仅能反映光能吸收、激发能传递和光化学反应等光合作用的原初反应过程,而且与电子传递、质子梯度的建立及ATP合成和CO₂固定等过程有关^[22,23] ...

2019

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2017

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... 孟娜等^[24]研究表明,烯效唑能够显著提高盐胁迫下黄13大豆(Glycine max)品种叶片光合色素含量 ...

2019

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. 2019, 45(7):1080-1089 DOI:10.3724/SP.J.1006.2019.84160

Effects of S3307 on the photosynthesis and yield of mung bean at R1 and R5 stages under waterlogging stress

S3307 对始花期和始粒期淹水绿豆光合作用及产量的影响

Yu Q , Feng N J , Wang S Y

于奇, 冯乃杰, 王诗雅

1 College of Agronomy, Heilongjiang Bayi Agricultural University, Daqing 163319, Heilongjiang, China 2 National Coarse Cereals Engineering Research Center, Heilongjiang Bayi Agricultural University, Daqing 163319, Heilongjiang, China

Waterlogging stress is one of the main abiotic stresses during the growth and development of crops. It is of great significance to explore the mechanisms for improving the flood resistance and waterlogging resistance cultivation of mung bean under waterlogging stress. In this experiment, the effects of uniconazole (S3307) on physiology, photosynthesis and yield of mung bean leaves under waterlogging stress were investigated in pot culture with different flood resistance mung bean varieties Lufeng 2 and Lufeng 5 from 2017 to 2018. Under the stress of waterlogging at different growth stages, the chlorophyll content (SPAD) and photosynthetic characteristic parameters of mung beans leaves were significantly decreased, malondialdehyde (MDA) content was significantly increased. The yield reduction rate of mung beans under the stress of waterlogging was 24.70%-33.63% at the beginning bloom (R1 stage), and 18.07%-28.87% at the beginning seed (R5 stage). Both mung bean varieties showed that the effect of waterlogging stress at R1 stage was greater than that at R5 stage, and the flood resistance of Lufeng 2 was stronger than that of Lufeng 5. After S3307 sprayed, SPAD, net photosynthetic rate ( P n ), transpiration rate ( T r ), and stomatal conductance ( G s ) in the leaves of mung beans could be significantly increased and MDA content could be significantly decreased. The remission rate of mung beans under waterlogging stress was 28.91%-52.34% at R1 stage, and 13.77%-27.36% at R5 stage. The results showed that S3307 sprayed on the leaf surface could effectively alleviate the physiological function and photosynthetic capacity of mung bean leaves under the stress of waterlogging, thus reduce the yield reduction, but there were differences in the regulatory response of mung bean varieties to S3307 in different waterlogging periods.

淹水胁迫是作物生长发育过程中遭受的主要非生物胁迫之一, 探究提高绿豆耐淹性的机制对绿豆抗涝栽培具有重要意义。本文在2017—2018年以耐淹性不同的绿豆品种绿丰2号和绿丰5号为试验材料, 采用盆栽方式探究了烯效唑(S3307)对淹水胁迫下绿豆叶片生理、光合作用及产量的影响。结果表明, 在不同生育时期淹水胁迫下, 绿豆叶片的叶绿素含量(SPAD)及光合特性参数均显著下降, 丙二醛(MDA)含量显著增加, 始花期(R1期)淹水胁迫下绿豆的减产率为24.70%~33.63%, 始粒期(R5期)减产率为18.07%~28.87%。2个绿豆品种均表现为R1期受淹水胁迫危害程度大于R5期, 绿丰2号耐淹性强于绿丰5号。喷施S3307后显著提高淹水胁迫下绿豆叶片的SPAD、净光合速率( P n )、蒸腾速率( T r )和气孔导度( G s ), 并显著降低了MDA含量。绿豆在R1期淹水胁迫下的缓解率为28.91%~52.34%, R5期缓解率为13.77%~27.36%。表明叶面喷施S3307可有效提高淹水胁迫下绿豆叶片的生理功能及光合能力, 进而降低减产幅度, 但不同淹水时期和绿豆品种对S3307的调控响应存在差异。

... 于奇等^[25]研究表明,叶面喷施烯效唑能够显著提高淹水胁迫下绿豆叶片的P_n、G_s、C_i、T_r ...

2015

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... 大量研究表明^[26,27],盐胁迫对植物的生长发育影响总体表现为抑制作用,具体表现为根系的伸长受到抑制,鲜重的增加和干物质的积累、侧根的减少 ...

2012

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... 大量研究表明^[26,27],盐胁迫对植物的生长发育影响总体表现为抑制作用,具体表现为根系的伸长受到抑制,鲜重的增加和干物质的积累、侧根的减少 ...

2013

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. 2013, 37(11):1018-1027 DOI:10.3724/SP.J.1258.2013.00105

Genotypic differences in salt tolerance from germination to seedling stage in peanut

花生萌发至苗期耐盐胁迫的基因型差异

Ci D W , Dai L X , Song W W

慈敦伟, 戴良香, 宋文武

Shandong Peanut Research Institute, Qingdao, Shandong 266100, China

Aims In the present study, the degree of salt tolerance in different peanut varieties/lines was investigated from emergence to seedling stage. Methods The seeds of peanut varieties/lines were planted in pots with salt levels of 0, 0.15%, 0.30%, and 0.45% NaCl in soils. The degree of salinity tolerance was systematically evaluated according to ten indices reflecting emergence, morphology, and biomass accumulation, etc. Important findings Statistic analysis for the 10 indices, including the relative emergence time, relative plant height, relative stem height, relative taproot length, relative shoot fresh mass, relative root fresh mass, relative shoot dry mass, relative root dry mass, and relative plant dry mass, showed large variations among the 107 varieties/lines; each index displayed a normal distribution under different salt stresses. Using the principal component analysis, the ten indices could be categorized into several independent factors. According to the total score, the 107 varieties/lines were divided into four groups, i.e., highly salt tolerant, salt tolerant, salt susceptible, and highly salt susceptible, by cluster analysis at salt level of 0.15%, 0.30%, and 0.45%, respectively. The 107 varieties/lines accounted for 29.0%, 39.0%, 27.5%, and 4.5%, respectively under salt level of 0.15%, and 5.5%, 34.5%, 23.5%, and 29.0%, respectively under salt level of 0.30%, in the highly salt tolerant, salt tolerant, salt susceptible, and highly salt susceptible groups. The salt tolerance in some varieties/lines varied with the level of salt stresses; some varieties/lines showed tolerance under low salt stress but susceptible under high salt stress. The number of varieties/lines showed the same tolerance or susceptibility under different salt stresses was 1 (‘Fuhua 11’), 7 (‘Huayu 28’, ‘Z11’, ‘Huayu 39’, ‘Huayu 32’, ‘HLN2’, ‘Yueyou 26’, and ‘XE019’), 0 and 1 (‘D1035’), respectively, in the highly salt tolerant, salt tolerant, salt susceptible, and highly salt susceptible groups. The 10 indices reflecting emergence, morphology, and biomass accumulation, etc., differed significantly or highly significantly among he varieties/lines in the four groups. Compared to the salt tolerant groups, the salt susceptible groups had prolonged emergence time and increased inhibition in plant morphological development and biomass accumulation under high level of salt stress.

采用盆栽试验, 设置0、0.15%、0.30%和0.45%等4个浓度的盐胁迫, 通过出苗速度、植株形态和生物量等指标研究不同基因型花生( Arachis hypogaea )品种(系)萌发至幼苗期的耐盐性差异。结果表明, 各浓度盐胁迫下, 相对出苗时间、相对株高、相对主茎高、相对主根长、相对地上部鲜质量、相对地下部鲜质量、相对植株鲜质量、相对地上部干质量、相对地下部干质量和相对植株干质量等10个指标耐盐指数品种间呈正态分布, 变异较大。主成分分析将10个指标归结为不相关的独立因子, 采用综合得分将107个品种(系)聚类为高度耐盐型、耐盐型、盐敏感型和高度盐敏感型4组, 0.15%浓度下4个类型品种数分别占供试材料的45.79%、39.25%、7.48%和7.48%; 0.30%浓度下4个类型品种(系)数分别占供试材料的11.21%、47.66%、22.43%和18.69%; 0.45%浓度下4个类型品种(系)数分别占供试材料的6.54%、23.36%、50.47%和19.63%。部分品种(系)在各强度盐胁迫下均表现为耐盐或盐敏感, 部分品种(系)在低强度胁迫下表现为耐盐, 而在高强度胁迫下表现为盐敏感。3个浓度盐胁迫下高度耐盐型、耐盐型、盐敏感型和盐高度敏感型分组一致的品种(系)分别有1个(‘阜花11’)、7个(‘花育28’、‘Z11’、‘花育39’、‘花育32’、‘HLN2’、‘粤油26’、‘XE019’)、0个和1个(‘D1035’)。盐胁迫下, 不同耐盐类型花生品种(系)出苗时间、植株形态和生物量等10个指标间差异达显著或极显著水平, 盐敏感类型比耐盐类型品种(系)出苗时间延长, 植株形态建成抑制加重, 物质积累减少。

... 本试验研究表明,在盐胁迫下,糜子幼苗根系的根总长、根平均直径、根总表面积、根体积和根干重均明显降低,这与慈敦伟等^[28]在花生(Arachis hypogaea)幼苗上的研究结果相一致 ...

2014

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... 同时,盐胁迫下采用叶面喷施烯效唑可以明显增加糜子幼苗的根总长、根平均直径、根总表面积和根体积,这与蒋玉香^[29]在黄瓜(Cucumis sativus)上的研究结果相一致 ...

2012

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... SOD、POD、CAT是植物体内酶促防御系统的重要保护酶^[30,31],在生物体中的水平高低意味着生物体的衰老与死亡 ...

1997

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... SOD、POD、CAT是植物体内酶促防御系统的重要保护酶^[30,31],在生物体中的水平高低意味着生物体的衰老与死亡 ...

2008

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... 宋胜等^[32]研究表明,在烯效唑0~50 mg#cod#x000b7 ...

2016

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... 本研究结果表明,与CK对比,盐胁迫下糜子根系内SOD活性、POD活性和CAT活性均有小幅度的升高,叶片SOD活性和POD活性有小幅度的升高,叶片和根系可溶性蛋白含量有少量增加,但与此同时,叶片和根系内 O2-·含量有明显的增加,这与张军等^[33]在小麦上的研究结果一致 ...

2011

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... L^-1烯效唑后,叶片中CAT活性、可溶性蛋白含量和根系中SOD活性、POD活性、CAT活性、可溶性蛋白含量均显著升高,而叶片中MDA含量和根系中 O2-·含量、MDA含量显著降低,表明喷施烯效唑可以提高植株体内抗氧化酶的活性,减少 O2-·和MDA的积累,降低其对细胞膜的破坏,从而增强植株对盐胁迫的抵抗能力,与李宁毅等^[34]在矮牵牛(Petunia hybrida)上的研究结果基本一致 ...

2018

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... 其原因可能是烯效唑正向调控内源激素ABA水平增加^[35],从而促进植物体抗氧化酶活性提高,并进一步影响植物体内自由基的合成,降低MDA积累量^[36],最终增加植物抗逆能力 ...

2019

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