摘要
土壤盐渍化严重影响作物生长发育,制约着作物产量和品质的提高。在设施栽培中由于特殊的小气候环境,加之复种指数高、不科学的肥水管理加重了土壤盐渍化,解决土壤盐渍化是当前设施蔬菜生产中急需解决的难题。脯氨酸是植物体内重要的渗透调节物质,许多植物受到逆境胁迫时会积累较高水平的脯氨酸,它在增强植物抗逆性上具有重要的作用。甜瓜因其果实香甜,富含营养,深为人们喜爱,由于甜瓜对土壤盐渍化所造成盐害胁迫比较敏感,限制了其种植的范围和生长效果。本研究以耐盐性较强的甜瓜品种‘玉皇’和耐盐性较弱的品种‘雪美’为试材,研究外源脯氨酸对盐(100mmol·L-1NaCl)胁迫下甜瓜幼苗生长、光合代谢、活性氧代谢和矿质元素吸收与分布等的影响；在此基础上,以‘雪美’为试材,研究了外源脯氨酸对盐胁迫下甜瓜幼苗体内硝酸还原作用与脯氨酸代谢的影响,旨在揭示外源脯氨酸提高甜瓜幼苗耐盐性的生理调节功能,主要结果如下：
采用隶属函数分析法对12个甜瓜品种进行了耐盐性分析,筛选出耐盐性强的品种3个,耐盐性中等的品种6个,耐盐性弱的品种3个。
适宜浓度脯氨酸处理可以缓解盐胁迫对‘雪美’甜瓜幼苗的伤害,0.1-0.4mmol·L-1脯氨酸能显著提高盐胁迫下甜瓜幼苗的鲜重和干重,增加甜瓜幼苗叶片的叶绿素含量,降低根系超氧阴离子(O2·-)产生速率、过氧化氢(H2O2)含量、丙二醛(MDA)含量和质膜相对透性,其中以添加0.2mmol·L-1脯氨酸效果最好,而较高浓度的脯氨酸(1.6-3.2mmol·L-1)加重了盐胁迫下甜瓜幼苗的伤害。
盐胁迫降低了甜瓜幼苗根系的SOD、CAT、APX、DHAR和GR抗氧化酶活性,耐盐性较弱的‘雪美’甜瓜变化幅度大于耐盐性较强的‘玉皇’,施用外源脯氨酸可提高盐胁迫下2个甜瓜品种体内的抗氧化酶活性,降低甜瓜体内O2·-产生速率和H2O2、 MDA含量及质膜相对透性,增强甜瓜的耐盐性,其中对‘雪美’效果好于‘玉皇’。外源脯氨酸还提高了盐胁迫和正常营养液培养下非酶促抗氧化物质ASA和GSH的含量。这些结果表明,外源脯氨酸可通过增强盐胁迫下甜瓜幼苗体内抗氧化酶活性和抗氧化剂来清除盐胁迫产生的活性氧,保护膜的完整性,缓解盐胁迫伤害。
盐胁迫导致甜瓜幼苗叶片叶绿素含量下降,降低甜瓜幼苗叶片的净光合速率(Pn)和PSⅡ实际光化学效率(ΦPSⅡ),抑制了甜瓜幼苗的生长,对‘雪美’甜瓜抑制作用大于‘玉皇’。盐胁迫显著地降低了‘玉皇’甜瓜气孔导度(Gs)和胞间CO2浓度(Ci),提高了‘雪美’甜瓜幼苗的Ci,说明‘玉皇’Pn下降是由气孔因素引起的,‘雪美'Pn下降是由非气孔因素引起的。外源脯氨酸通过提高‘雪美’甜瓜叶片的叶绿素含量、气孔导度(Gs)、胞间CO2浓度(Ci)、PS Ⅱ实际光化学效率(OPS Ⅱ)和光化学淬灭(qP),提高‘玉皇'Pn、Chlb和ΦPS Ⅱ,增强甜瓜植株光合速率,提高PsⅡ的运转效率。表明,外源施加脯氨酸可以通过增强光合电子传递效率,增加了光化学反应分配的光能份额,从而提高了实际光化学效率,增强甜瓜的耐盐性,对耐盐性较弱的‘雪美’甜瓜使用效果较好。
盐胁迫降低了植株体内Ca2+、K+、Mg2+等大量元素和Fe、Mo、Mn、Zn、Cu等微量元素含量,增加了Na+和Cl-的含量,破坏了甜瓜体内矿质营养元素平衡,并且耐盐性较弱的‘雪美’甜瓜变化的幅度大于耐盐性较强的‘玉皇’甜瓜；盐胁迫提高了甜瓜幼苗根系N和P的含量,降低了叶片中N和P含量。盐胁迫下外源施用脯氨酸对甜瓜幼苗体内的矿质营养元素的吸收具有明显地调节作用,进一步提高了N和P含量和Ca2+、K+、Mg2+、Fe、Mo、Mn、Zn、Cu等含量,增加了组织中的K+/Na+、Mg2+/Na+和Ca2+/Na+的比值,提高对Ca2+、K+、Mg2+的选择性运输能力。表明,脯氨酸可以促进甜瓜对各种必需营养元素的吸收,增强离子区域化分布能力,调节植物体内的矿质营养元素平衡,缓解盐胁迫对植物的伤害。
盐胁迫提高了‘雪美’甜瓜幼苗叶片和根系内铵态氮(NH4+-N)含量,降低了硝态氮(N03--N)含量以及硝酸还原酶(NR)和谷氨酰胺合成酶(GS)活性,抑制了甜瓜对氮素的吸收利用；盐胁迫下,甜瓜幼苗可溶性蛋白和脯氨酸含量提高。外源施用脯氨酸增强了盐胁迫下甜瓜幼苗体内NR和GS活性,提高了盐胁迫下甜瓜幼苗叶片和根系内N03--N含量、降低了盐胁迫下甜瓜幼苗叶片和根系内NH4+-N含量,促进了甜瓜对氮素的吸收利用。施用外源脯氨酸进一步提高了盐胁迫下甜瓜幼苗体内可溶性蛋白和脯氨酸含量,抑制了吡咯啉-5-羧酸合成酶(P5CS)活性,提高了鸟氨酸转氨酶(OAT)和脯氨酸脱氢酶(ProDH)活性,改变了植物体内脯氨酸代谢途径。这些结果表明,脯氨酸可以调节甜瓜幼苗体内硝酸还原作用,促进植株对氮素的吸收,减轻盐胁迫伤害；同时也通过调节甜瓜体内脯氨酸代谢,增加脯氨酸含量,提高甜瓜耐盐性。
Soil salinity severely affects the growth and development of crops and restricts their yields and quality. However, this phenomenon is becoming more and more serious because of special microclimate environment, high multiple crop index and unscientific management of water and fertilizer and needs to be solved urgently in greenhouse cultivation. Proline is a crucial osmotic adjustment in plants and will be accumulated to a higher lever when plants are in environmental stress, which plays an important role in enhancing resisting adversity of plants. In this paper, melon cultivars salt tolerance were identified and filtered. Based on the effects of exogenous proline on growth, photosynthesis, ROS metabolism, the absorption and distribution of mineral elements in melon cultivars (Cucumis melo L. cv. Yuhuang and cv. Xuemei) under100mmol·L-1salt stress, had studied the effects of exogenous proline on the nitrate reducation and proline metabolism in Xuemei, Main results were as follows:
Salt tolerance of the12melon cultivars melon were compared by membership function method, and selected three high salt tolerant, six moderate salt tolerant and three sensitivity salt tolerant melon cultivars.
Suitable concentration proline could alleviate damage caused by salt stress,0.1-0.4mmol·L-1proline could improve the fresh and dry weight of the melon seedlings under the salt stress remarkably and increase their chlorophyll contents as well as decreased H2O2content, O2-production rate, MDA content and membrane permeability. Among these results, the0.2mmol·L-1proline treatment showed the best effect. But the higher concentration of proline treatment1.6and3.2mmol·L-1made the damage on melon seedlings worse.
Salt stress decreased SOD, CAT, DHAR, APX and GR activities of both melon cultivars, and the variation range of Xuemei was larger than Yuhuang. After using proline, the two cultivars'antioxidant activities and salt tolerance was increased while their O2·-production rate was decreased with H2O2content, MDA content and membrane permeability decreased as well under salt stress,and its effects on Xuemei was better than Yuhuang. exogenous proline also increased the content of non-enzymatic antioxidants such as ASA and GSH under salt stress and normal nutrient solution culture. Therefore, it showed that the exogenous proline can clear the ROS caused by salt stress by increasing the activities of antioxidant enzymed and non-enzymatic antioxidants content so that it can protect the membrane integrity to lessen the damage caused by the salt stress.
Salt stress decreased the chlorophyll content, Pn and ΦPSⅡ, inhibited the growth, the effect was more markedly Xuemei than Yuhuang. Salt stress decreased Gs and Ci in cv. Yuhuang melon, increased Ci in cv. Xuemei. Exogenous proline increased chlorophyll content, Gs, Ci, ΦPS Ⅱ and qP in cv. Xuemei, increased Pn, Chlb and ΦPS Ⅱ in Yuhuang, Therefore, these results showed that exogenous proline can increase the actual photoche-mical efficiency to strengthen melons'salt tolerance by enhancing the transport rate of photo synthetic electron and light sharing distributed by photochemical reaction and its effects on Xuemei was better.
Salt stress decreased ion contents, in creased the Na+and Cl-contents in melon, destroy the balance of mineral nutrient, the effect was more markedly Xuemei than Yuhuang; salt stress increased the N and P contents in roots, decreased in leaves. Exogenous proline markedly regulate the absorption to mineral nutrient under salt stress, increased N and P content in melon; and also Ca、K、Mg、Fe、Mo、Mn、Zn and Cu contents, enhanced K+/Na+、Mg2+/Na+and Ca2+/Na+ratio. These results showed that exogenous proline could alleviate the damage by accelerating melons'absorption of necessary nutrients, enhancing ion's regional distribution ability and regulating the absorption balance of mineral nutrients.
Salt stress increased NH4+-N, decreased NO3--N content, NR and GS activities in leaves and roots of cv. Xuemei melon, inhibit the absorption of nitrogen in melon; melon increased the salt torelent via to increase soluble protein and proline contents. Exogenous proline increased NR and GS activities, increased NO3--N, decreased NH4+-N content, promoting the absorption of N. exogenous proline improved the soluble protein and proline contents, feedback inhibite P5CS activity, increase OAT and ProDH activities. it could be concluded that the proline could regulate Nitrate reduction and foster their absorption to nitrogen to alleviate the damage caused by salt stress.melons salt tolerance could be improved by regulating their proline metabolism to increase the proline content.
引文
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