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分根盐处理对玉米产量的影响及根系响应
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摘要
本试验于2009-2011年在黄淮海区域玉米技术创新中心和作物生物学国家重点实验室进行。本研究采用盆栽试验和室内生理生化分析相结合的技术路线,选用郑单958和登海9为试验材料,以盆栽的方式进行分根盐处理,系统研究了断根和不均匀盐处理对玉米产量、品质、渗透调节、Na+、Cl-分布、抗氧化系统,根际土壤特性,根系形态及生理生化变化的影响,探讨分根对盐处理玉米产量形成的机理及根系响应。主要研究结果如下:
     1分根盐处理对玉米产量及品质的影响
     玉米生物产量和籽粒产量在盐处理下有类似规律:低盐度时,生物产量和籽粒产量有不同程度的升高;中高盐度条件下产量显著下降,当盐浓度达到102mMNaCl时,玉米的生长受到严重影响,干物质积累量极低,不能开花结实,直至死亡。断根玉米在无盐和低盐时促进了玉米的生长,产量显著高于全根玉米,中盐度时二者差异不大或断根玉米产量略小,高盐度时无产量。无盐和低盐时断根玉米籽粒粗蛋白含量显著超过对照,淀粉和粗脂肪含量与对照差异不显著。
     不均匀盐处理改善了玉米生长,产量显著高于对照。不均匀盐处理玉米籽粒淀粉含量显著高于对照,而对粗蛋白和粗脂肪的影响在不同品种和盐度间规律不一致。
     2分根盐处理对玉米光合特性的影响
     断根显著影响盐处理下玉米的光合特性,具体表现为:无盐和低盐时,断根玉米花后光合色素含量,穗位叶净光合速率、蒸腾速率、气孔导度,叶绿素荧光参数ФPSⅡ,qP值显著高于全根玉米,干物质积累量增加;中高盐度时,断根玉米的光合性能受抑制:叶绿素含量减少,玉米叶面积显著降低,穗位叶光合速率、蒸腾速率、气孔导度下降,断根玉米干物质量与全根玉米相比差异不大或较小。
     不均匀盐处理显著增加了玉米的光合色素含量,玉米净光合速率、蒸腾速率、气孔导度超过对照,ФPSⅡ,qP值提高,胞间CO_2浓度和NPQ降低,大口期后玉米叶面积显著高于均匀盐处理,干物质积累较多。
     3分根盐处理对玉米渗透调节、抗氧化系统的影响
     3.1分根盐处理对玉米渗透调节的影响
     无盐和低盐时断根玉米叶水势、K~+、Ca~(2+)含量、脯氨酸、可溶性糖、抗坏血酸含量均显著高于全根玉米,而叶中Na~+、Cl-含量显著低于全根玉米。由此可见无盐和低盐时断根促进了玉米的渗透调节能力,而中高盐度时断根玉米小于全根玉米。耐盐品种DH9渗调能力超过ZD958。
     不均匀盐处理改善了盐处理玉米的水分状况,不均匀盐处理玉米叶中高浓度的K~+、Ca~(2+)、甜菜碱和脯氨酸含量显著高于同盐度时的均匀盐处理玉米,并促进了根叶的生长。不均匀盐处理中无机离子在长期盐处理时起主要作用,其中K~+和Ca~(2+)是最主要的无机离子,在渗透调节中起非常重要作用,特别是维持较高的K/Na比和Ca/Na比尤为重要。有机物质在渗透调节中作用相对较小。
     3.2分根盐处理对玉米抗氧化系统的影响
     断根玉米叶片MDA含量在低盐时低于全根玉米,无盐时二者差异不显著,中高盐时断根玉米叶片MDA含量显著高于全根玉米。断根玉米大口期后叶片SOD、POD酶活性在无盐和低盐时显著高于全根玉米,APX、GPX含量在不同时期呈现不同规律。
     不均匀盐处理玉米叶片MDA含量在3个时期几乎都小于对照。不均匀处理玉米大口期时叶片POD、APX活性显著高于对照,开花期时POD、GPX、APX活性高于对照,乳熟期时APX活性小于对照,POD、GPX活性高于对照。
     4分根盐处理对玉米根际土壤生理生化指标的影响
     4.1分根盐处理对玉米根际土壤电导率及离子积累的影响
     不同时期除无盐处理外,三种盐度下断根玉米根际土壤电导率,Na~+、Cl-含量均高于全根玉米。不均匀盐处理时水侧根际土壤电导率随盐侧所浇盐水浓度增加而增加,这说明玉米根系有将一侧矿质元素运至另一侧的能力。不均匀盐处理根系只有一侧浇盐水,两侧的Na~+、Cl-含量不同。两侧土钠含量除高盐度两侧Na~+含量差异显著外,其余两个不均匀盐处理差别不大。说明钠可有一侧土壤运至另一侧。而氯含量在不均匀盐处理两侧根际土壤中相差较大。
     4.2分根盐处理对玉米土壤生理生化特性的影响
     两玉米品种根际土壤脲酶活性变化规律略有不同:断根ZD958在拔节和大口期根际土壤脲酶活性小于对照,自开花期后均高于全根盐处理;断根盐处理DH9拔节期时脲酶活性小于对照,自大口期后均高于对照,玉米根际土壤过氧化氢酶活性随盐度增加先增加后减少。两品种玉米根际土壤过氧化氢酶活性变化趋势基本一致:断根玉米拔节期根际土壤过氧化氢酶活性小于全根玉米过氧化氢酶活性,断根玉米大口期后根际土壤过氧化氢酶活性均高于对照。无盐和低盐时断根玉米根际土壤蔗糖酶活性在花后高于对照,两品种表现一致。
     不均匀盐处理两侧土壤脲酶活性不同,盐度差别越大差异越显著。水侧脲酶活性显著高于盐侧土壤。不均匀盐处理根系两侧土壤脲酶活性开花后显著高于对照。蔗糖酶、过氧化氢酶活性变化规律与脲酶基本一致。
     5玉米根系对分根盐处理的响应
     无盐和低盐时断根玉米根系出现补偿生长,根条数,根总表面积、根尖数、根体积和根总长增加,根活性变大,根系抗氧化酶活性增加,植株生物量和籽粒产量提高,品质改善,植株Na~+、Cl-积累量降低,叶片光合能力提高,植株对NaCl的耐性提高。断根对中高盐度胁迫下玉米的补偿作用较小。
     不均匀盐处理促进了玉米根系的生长,根体积、根条数、根尖数、根总表面积和根总长,根活性显著高于对照,根系抗氧化酶活性升高,MDA含量相对较小,根毛区细胞受盐害程度小于对照。不均匀盐处理玉米清除活性氧能力增强可以减轻高盐对玉米的毒害作用,激素调节促进玉米的根系生长,延缓衰老,促进产量增加。
The experiment was carried out in the Maize Technological Innovation Center in the Yellow River Basin and the Huaihe River Basin and in the States Key Laboratory of Crop Biology from2010to2011. This study used the technology line of integration of pot experiment in the field and physiological and biochemical experiments indoor. We studied the effects of split-root salt treatments on maize (ZD958and DH9) under several concentrations of NaCl. The contents of study included yield, grain quality, phytosynthesic performance, osmotic adjustment, antioxidative system, soil enzymes, root morphology and physilogical characteristics. The results summaries were as follows:
     1Effects of split-root salt treatements on yield, grain quality in maize
     Biomass and grain yield had similar changing rule with salt treatments:At the lowest salt concentration (W1or N1), it enhanced these yields. When it came to higher NaCl concentrations, biomass and yield decreased significantly. When NaCl concentration reached to102mM, the growth of maize had been seriously inhibited and dry matter accumulation was very low, and plant could not flowering, even to death.
     Excision partial root significantly increased maize yield and growth when irrigated with tap water or34mM NaCl solution. When irrigated with moderate NaCl solutiion, the yield of maize with partial root pruned was not different from whole-root maize or lower than that. And high NaCl treatments had no yield. Excision partial root siginificantly improved the protein contents of maize grain when irrigated with tap water or lower concentration NaCl solution.
     Unequal salt stress siginificantly improved maize growth than equal salt stress, and got higher yield than the control. The contents of seed starch were increased in unequal salt treatments, and the contents of protein and fat were changed with the cultivar and the salt solution.
     2Effects of split-root salt treatments on phytosynthetic characteristics in maize
     Excision partial roots significantly affected phytosynthesic characteristics in maize, specifically as follows:When irrigated with tap water or lower concentration of NaCl solution, excision partial root increased chlorophyll contents, improved the net photosynthetic rate, somatal conductivity, transpirition rate, ΦPSⅡ, qP and increased dry matter accumulation in maize postanthesis; when irrigated with moderate and high NaCl solution, the photosynthetic performance of maize with partial root pruned was inhibited; chlorophyll contents and leaf area both decreased, photosynthetic rate, transpiration rate, stomatal conductivity reduced, dry matter weights had little difference or slightly smaller than whole-root maize.
     Unequal salt treatments significantly increased chlorophyll contents, photosynthetic rate, transpiration rate, stomatal conductivity,ΦPSⅡ, qP, and intercelluar CO2concentration, while NPQ were decreased. After big flare phase, leaf area significantly higher than control and dry matter accumulation were more.
     3Effects of split-root salt treatments on osmotic adjustment and antioxidative system of maize
     3.1Effects of split-root salt treatments on osmotic adjustment of maize
     when irrigated with tap water or low NaCl solution, WP, contents of K+, Ca2+, proline, soluble sugar, ascorbic acid of leave in maize with partial root pruned significantly increased than the control; and Na+and Cl-contents in leave significantly lower than the control. So excision partial root induced the ability of osmotic adjustment in maize when irrigated with tap water and low NaCl solution. Cultivar DH9has the more osmotic adjustment ability than ZD958.
     Unequal salt treatments improved water uptake and the concentration of K+, Ca2+, GB, and Pro, which induced shoot and roots growth. Inorganic ions were the main contributors in unequal salt treatments for the long-term tolerance to the salt stress, and K+and Ca2+were the primary inorganic ions, which were very important in osmotic adjustment, especially maintaining the high K/Na and Ca/Na ratios. Organics played a relatively minor role in osmotic adjustment, whereas in those GB and Pro played larger roles, followed by SS.
     3.2Effects of split-root salt treatments on antioxidant system of maize
     MDA contents in maize with partial root pruned were lower than whole-root maize when irrigated with low NaCl solution, and were undifferent when irrigated with tap water. When it came to moderate and high NaCl, the contents of MDA in pruned-root maize were significantly higher than the control. After big flare stage, SOD and POD activity of pruned-root maize leaves significantly higher than whole-root maize when irrigated with tap water and low NaCl solution. APX and GPX activity presents different rules at different stages.
     MDA contents in unequal salt treatments maize leaves were all lower than control at three stages, only at milk-ripe stage, MDA contents of U1and U2in ZD958slightly larger than the control.
     Three enzymes which responsible for the removal H2O2play different role at different stages:at big flare stage, POD and APX played the main role in unequal salt treatments, at anthesis stage, POD, GPX and APX together played the role; at milk-ripe, APX activity was lower than control, POD and GPX played the main role.
     4Effects of split-root salt treatments on rhizosphere soil of physiological and biochemical indictors of maize
     4.1Effects of split-root salt treatments on rhizosphere soil conductivity and ion accumulation in maize
     Except for tap water, at different stages, soil conductivity, contents of Na+and Cl-in pruned-root maize irrigated with three different salt concentration were significantly higher than control. In unequal salt treatments, the soil conductivity of root part irrigated with tap water were increased with the increase of the salt concentration that irrigating the other root part. This reflected a translocation of mineral elements from one root part to the other. Because irrigated with different concentration salt solution, Na+and Cl-contents in the two root parts were different. Except for high salt solution, Na+contents in two soil parts were little different.
     4.2Effects of split-root salt treatments on soil enzymes activities in maize
     The rule of soil urease activity was little different in two maize cultivars:at jionting and big flare stage, urease activity of ZD958rhizosphere soil were lower than the control, and after anthesis, the condition was the reverse; at jionting stage, urease activity of DH9rhizosphere soil were lower than the control, but after big flare stage, the condition was the reverse. With salt solution increased, soil catalase activity firstly increased and then decreased. The soil catalase activity change of two cultivars presents the same trend:at jionting stage, catalase activity in pruned-root maize rhizosphere soil was lower than the control, after big flare stage, the condition was the reverse. After anthesis, sucrase activity of pruned-root rhizosphere soil was higher than control. And the change rule of two cultivars was the same.
     Urease activity in two soil parts irrigated with different salt solution were significantly different, and the differnet was more when salt concentration was biggest in two soil parts. The urease activity of soil irrgated with tap water was significantly higher than the other soil part irrigated with salt solution. The change rule of sucrase and catalse activity was the same as urease activity.
     5Responses of root to split-root salt treatments in maize
     5.1Responses of root to pruned-root salt treatments in maize
     The roots of pruned-root maize appeared compenstory growth, the root number, root area, root tips, root volume, whole root length, root activities and antioxidative enzymes activities increased, maize biomass and yield induced, grain quality improved, Na+and Cl-contents in plants decreased, photosynthesis performance improved, and the tolerance of NaCl also impoved when maize irrigated with tap water and low salt concentration solution.
     5.2Responses of root to unequal salt treatments in maize
     Unequal salt treatments induced growth of maize root:root volume, root number, root tips, root area, root length and root activities increased, root antioxidative enzymes activities improved, MDA contents decreased, the salt damage in cell of root hair zone were less than the control. So unequal salt treatments alleviated high salt stress, induced maize growth and got higher yield.
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