鉬在草莓中的吸收轉(zhuǎn)運(yùn)及其對(duì)氮代謝調(diào)控機(jī)制的研究
發(fā)布時(shí)間：2018-08-05 14:32
【摘要】：鉬(molybdenum,Mo)是植物正常生長(zhǎng)必不可少的微量元素之一,參與到植物的氮代謝過(guò)程中。缺鉬的條件下植物容易積累較多的硝態(tài)氮從而表現(xiàn)出較低的氮素利用率(nitrogen utilization efficiency,NUE)。提高氮素利用率可以減少化學(xué)肥料過(guò)多施用帶來(lái)的負(fù)面效應(yīng),但是關(guān)于鉬提高氮素利用率的機(jī)理尚不清楚。本研究分別在溫室無(wú)土栽培和組培條件下進(jìn)行。以‘章姬’(Fragaria×ananassa Duch.cv.‘Akihime’)草莓為試驗(yàn)材料,在溫室無(wú)土栽培條件下噴施不同濃度的鉬酸鈉(sodium molybdate,Na_2MoO_4),試驗(yàn)設(shè)置T1(葉面噴施清水)、T2(0.02%Na_2MoO_4)、T3(0.04%Na_2MoO_4)、T4(0.05%Na_2MoO_4)和T5(0.06%Na_2MoO_4)5個(gè)處理。研究了其對(duì)植株生物量、果實(shí)營(yíng)養(yǎng)元素含量和果實(shí)品質(zhì)(可溶性固形物(total soluble solids,TSS)、抗壞血酸(ascorbic acid,AsA)、可滴定酸(titratable acidity,TA)、芳香物質(zhì))、光合作用、鉬轉(zhuǎn)運(yùn)蛋白基因MOT1表達(dá)量、植株體內(nèi)Mo濃度、氮代謝關(guān)鍵酶活性(硝酸還原酶,nitrate reductase,NR;亞硝酸還原酶,NiR;谷氨酰胺合成酶,glutamine synthetase,GS;NADH-谷氨酸合酶,NADH-glutamate synthase,NADH-GOGAT)、氮代謝關(guān)鍵基因(NRT1.1,NRT2.1,NIA,NiR,GS-1,NADH-GOGAT)表達(dá)量、15N吸收、分配及利用的影響。在組培條件下,MS固態(tài)培養(yǎng)基中加Na_2MoO_4或者鎢酸鈉(sodium tungstate,Na2WO4),設(shè)置CK(-Na_2MoO_4-Na2WO4)、Mo處理(+Na_2MoO_4)、W處理(+Na2WO4)、Mo+W處理(+Na_2MoO_4+Na2WO4)4個(gè)處理。研究其對(duì)植株表型、根系顯微結(jié)構(gòu)及根系活力、葉片顯微結(jié)構(gòu)與超顯微結(jié)構(gòu)及葉綠素含量、MOT1表達(dá)量、鉬輔因子合成通路關(guān)鍵基因(CNX2,CNX3,CNX5,CNX6,CNX7,CNX1)表達(dá)量、植株體內(nèi)Mo濃度、氮代謝關(guān)鍵酶活性(NR,NiR,GS,NADH-GOGAT)以及15N利用的影響。主要結(jié)果如下:1.適宜濃度的Mo供應(yīng)能顯著提高幼苗的生物量。葉面噴施Mo肥對(duì)草莓果實(shí)的營(yíng)養(yǎng)元素(N,Mo,Fe和Cu)含量,TSS,TA和芳香物質(zhì)含量有一定的提升作用。但是施Mo對(duì)果實(shí)中的Se和部分芳香物質(zhì)沒(méi)有顯著影響。0.04%Na_2MoO_4處理在所有處理中表現(xiàn)出最高的果實(shí)TSS,N和Fe濃度。0.02%Na_2MoO_4處理的AsA含量顯著高于對(duì)照。利用頂空固相微萃取法和氣質(zhì)聯(lián)用技術(shù)一共鑒別出97中芳香物質(zhì)。0.04%Na_2MoO_4處理的6種特征酯類(lèi)(丁酸甲酯,γ-癸內(nèi)酯,丁酸乙酯,己酸甲酯,γ-十二內(nèi)酯,己酸乙酯)含量最高。2.草莓幼苗根部、莖部和葉片的Mo含量隨著施Mo量的增加而呈現(xiàn)上升的趨勢(shì)。0.04%Na_2MoO_4處理的根部的MOT1的表達(dá)水平高于其它處理,當(dāng)Na_2MoO_4的濃度從0.05%增加到0.06%時(shí),根部的MOT1的mRNA水平呈現(xiàn)下降的趨勢(shì),培養(yǎng)5天后,葉片的MOT1的表達(dá)水平隨著噴施Na_2MoO_4濃度的提高而提高;培養(yǎng)10天和15天后,當(dāng)Na_2MoO_4的濃度從0.00%增加到0.04%時(shí),葉片的MOT1的表達(dá)水平呈現(xiàn)升高的趨勢(shì),而后隨著噴施Na_2MoO_4的提高而呈現(xiàn)下降的趨勢(shì)。施Mo影響幼苗根部Fe、Cu和Se濃度。0.04%Na_2MoO_4處理的幼苗表現(xiàn)出最高的15N地上部/地下部比例和15N利用率。0.04%Na_2MoO_4處理的氮代謝關(guān)鍵酶活性較高,且硝態(tài)氮吸收基因(NRT1.1和NRT2.1)以及硝態(tài)氮響應(yīng)基因的表達(dá)水平上調(diào)。0.04%Na_2MoO_4處理的根部和葉片的NO3--N濃度低于其它處理,葉片的NH4+-N的濃度較高,Gln/Glu的濃度也較高。而T5處理表現(xiàn)出相反的趨勢(shì)。因此,葉面噴施0.04%Na_2MoO_4通過(guò)調(diào)控氮代謝關(guān)鍵酶活性和NO3-吸收同化相關(guān)基因的表達(dá)提高了根部NO3-向地上部的轉(zhuǎn)運(yùn),提高了氮素利用率。3.MS固態(tài)培養(yǎng)基中正常加入Na_2MoO_4的草莓幼苗長(zhǎng)勢(shì)較好,葉片為正常綠色,葉肉細(xì)胞顯微結(jié)構(gòu)及超微結(jié)構(gòu)完整,葉綠體較多,葉綠素含量較高。根系發(fā)育正常,根系活力較高,根尖細(xì)胞排列整齊緊密,中央髓部細(xì)胞體積較大,排列緊密。而缺Mo培養(yǎng)基上生長(zhǎng)的幼苗葉片下表皮細(xì)胞排列不整齊,葉肉細(xì)胞超微結(jié)構(gòu)異常,細(xì)胞器及內(nèi)膜系統(tǒng)受損,葉綠素含量較低,葉片發(fā)黃。根系較長(zhǎng),但是長(zhǎng)勢(shì)較弱,根尖細(xì)胞微變形,細(xì)胞排列疏松,細(xì)胞間隙較大。只加Na2WO4和同時(shí)加入Na_2MoO_4和Na2WO4的培養(yǎng)基上的幼苗株型變小,長(zhǎng)勢(shì)減弱,細(xì)胞器受損,膜結(jié)構(gòu)不清晰,部分葉片表現(xiàn)為黃色,葉綠素含量降低,出現(xiàn)與氮素缺乏類(lèi)似的癥狀。施入Na_2MoO_4可以顯著誘導(dǎo)草莓地上部和根部FaMOT1的表達(dá),加入抑制劑Na2WO4的FaMOT1表達(dá)量則表現(xiàn)出相對(duì)下降,且施入Na_2MoO_4幼苗的Mo濃度顯著高于其它處理。培養(yǎng)基中加入Na_2MoO_4的組培苗的CNX2和CNX3的表達(dá)量較高,培養(yǎng)基加入Na_2MoO_4的組培苗中CNX7和CNX1的表達(dá)量高于加入抑制劑的組培苗。加入抑制劑Na2WO4使得NR活性顯著降低,加入Na_2MoO_4組培苗的NR活性最高,其它氮代謝關(guān)鍵酶(NiR,GS和NADH-GOGAT)表現(xiàn)出類(lèi)似的趨勢(shì)。培養(yǎng)基加入Na_2MoO_4的組培苗的NO3--N濃度低于其它處理,NO2--N和NH4+-N濃度顯著高于其它處理。同位素15N-Ca(NO3)2標(biāo)記進(jìn)一步表明培養(yǎng)基加入Na_2MoO_4的組培苗吸收利用了較多的15N,轉(zhuǎn)運(yùn)到地上部的15N也較多,植株的15N利用率最高,而加入抑制劑的組培苗的15N利用率較低。
[Abstract]:Molybdenum (Mo) is one of the essential trace elements essential for the normal growth of plants. It participates in the process of nitrogen metabolism in plants. Under the condition of lack of molybdenum, plants accumulate more nitrate nitrogen and thus show low nitrogen use efficiency (nitrogen utilization efficiency, NUE). Increasing nitrogen use rate can reduce the excessive application of chemical fertilizer. The negative effect is not clear, but the mechanism of increasing nitrogen utilization rate of molybdenum is not clear. This study was carried out under the conditions of soil free cultivation and tissue culture in greenhouse. The experimental materials of "Fragaria x ananassa Duch.cv. 'Akihime") strawberry were used to spray different concentrations of sodium molybdate (sodium molybdate, N) under the condition of soil free cultivation in greenhouse. A_2MoO_4), the experiment set up T1 (foliar spraying water), T2 (0.02%Na_2MoO_4), T3 (0.04%Na_2MoO_4), T4 (0.05%Na_2MoO_4) and T5 (0.06%Na_2MoO_4). The plant biomass, fruit nutrition element content and fruit quality (total soluble), ascorbic acid, titratable acid were studied. Table acidity, TA), photosynthesis, MOT1 expression of molybdenum transporter gene, Mo concentration in the plant, activity of key enzymes in nitrogen metabolism (nitrate reductase, nitrate reductase, NR; nitrite reductase, NiR; glutamine synthetase, glutamine synthetase, GS; glutamate synthase, depletion), nitrogen metabolism The expression of key genes (NRT1.1, NRT2.1, NIA, NiR, GS-1, NADH-GOGAT), the effect of 15N absorption, distribution and utilization. Under the condition of tissue culture, Na_2MoO_4 or sodium tungstate (sodium tungstate, Na2WO4) in the solid culture medium of MS (sodium tungstate, Na2WO4), 4 treatments. The plant phenotype, root microstructure and root activity, leaf microstructure and ultrastructure and chlorophyll content, MOT1 expression, the expression of key genes (CNX2, CNX3, CNX5, CNX6, CNX7, CNX1), the concentration of Mo in the plant, the activity of the key enzymes of nitrogen metabolites (NR, NiR, GS, NADH-GOGAT), and the effects of the utilization of the plants. The results are as follows: 1. the suitable concentration of Mo supply can significantly increase the biomass of the seedlings. Spraying Mo fertilizer on the leaf surface can improve the content of nutrient elements (N, Mo, Fe and Cu), TSS, TA and aromatic substance in strawberry fruit. However, the application of Mo on Se and some aromatic substances in the fruit has no significant effect on the treatment of.0.04%Na_2MoO_4 in all treatments. The highest TSS, N and Fe concentrations of.0.02%Na_2MoO_4 were significantly higher than those of the control. A total of 6 kinds of esters (methyl butyrate, gamma decanide, ethyl butyrate, methyl Hexanate, gamma twelve lactone, gamma twelve lactone, ethyl caproate, ethyl hexanoate) were identified by headspace solid phase microextraction and GC-MS. The content of Mo in the roots of the strawberry seedlings with the highest content of.2., the content of the stems and leaves increased with the increase of Mo, and the expression level of MOT1 in the root of the.0.04%Na_2MoO_4 treatment was higher than that of the other treatments. When the concentration of Na_2MoO_4 increased from 0.05% to 0.06%, the mRNA level of MOT1 in the root showed a decline trend, and the leaves was MO after 5 days. The expression level of T1 increased with the increase of the concentration of Na_2MoO_4. When the concentration of Na_2MoO_4 increased from 0% to 0.04% after 10 and 15 days, the expression level of MOT1 in leaves showed a trend of increase, and then decreased with the increase of spraying Na_2MoO_4. Mo influenced Fe, Cu and Se.0.04%Na_2MoO_4 in the roots of seedlings. The seedlings of the treated seedlings showed the highest 15N upper / underground ratio and the 15N utilization rate.0.04%Na_2MoO_4 treated with higher nitrogen metabolism key enzyme activity, and the expression level of nitrate nitrogen absorption gene (NRT1.1 and NRT2.1) and nitrate nitrogen response gene up regulated the NO3--N concentration of the root and leaf of.0.04%Na_2MoO_4 treatment was lower than that of other treatments. The concentration of NH4+-N was higher and the concentration of Gln/Glu was higher, while T5 treatment showed the opposite trend. Therefore, the foliar spraying 0.04%Na_2MoO_4 enhanced the transport of NO3- to the top of the root by regulating the activity of key enzyme activities of nitrogen metabolism and the expression of NO3- assimilation related genes, and increased the normal addition of Na_ in the solid-state medium of nitrogen utilization ratio.3.MS. The growth trend of strawberry seedlings in 2MoO_4 is better, the leaf is normal green, the microstructure and ultrastructure of the mesophyll cells are complete, the chloroplasts are more, the chlorophyll content is higher, the root development is normal, the root activity is high, the root tip cells are arranged neatly and closely, the central marrow cells are large and arranged closely. The leaves are short under the leaves of the seedlings grown on the Mo medium. The epidermal cells were not arranged regularly, the ultrastructure of the mesophyll cells was abnormal, the organelles and the intima system were damaged, the chlorophyll content was low, the leaves were yellow and the roots were long, but the growth potential was weak, the root tip cells were slightly deformed, the cells were loosely arranged and the cell gaps were large. The plant types on the medium with Na2WO4 and Na_2MoO_4 and Na2WO4 were smaller. The cell organ is impaired, the organelle is damaged, the membrane structure is not clear, some leaves are yellow, the content of chlorophyll is reduced, and there is a similar symptom with nitrogen deficiency. Na_2MoO_4 can significantly induce the expression of FaMOT1 in the upper and root parts of the strawberry, and the amount of the FaMOT1 surface of the inhibitor Na2WO4 shows a relative decline and is applied to the Na_2MoO_4 seedlings. The concentration of Mo was significantly higher than that of other treatments. The expression of CNX2 and CNX3 in tissue culture seedlings with Na_2MoO_4 was higher. The expression of CNX7 and CNX1 in the tissue culture medium with Na_2MoO_4 was higher than that of the tissue culture seedlings which were added to the culture medium. The addition of inhibitor Na2WO4 made the NR activity significantly lower, and the NR activity of the group of Na_2MoO_4 tissue cultured seedlings was the highest. The key enzyme of nitrogen metabolism (NiR, GS and NADH-GOGAT) showed a similar trend. The concentration of NO3--N in the culture medium with Na_2MoO_4 was lower than that of other treatments, and the concentration of NO2--N and NH4+-N was significantly higher than that of other treatments. The isotopic 15N-Ca (NO3) 2 marker further indicated that the tissue culture seedlings of the culture medium added to Na_2MoO_4 were absorbed and utilized more 15N and transported to the ground. There were more 15N in the upper part, and the highest utilization rate of 15N in plants. The 15N utilization rate of tissue culture seedlings with inhibitors was lower.
【學(xué)位授予單位】：山東農(nóng)業(yè)大學(xué)
【學(xué)位級(jí)別】：博士
【學(xué)位授予年份】：2017
【分類(lèi)號(hào)】：S668.4
本文編號(hào)：2166079
[Abstract]:Molybdenum (Mo) is one of the essential trace elements essential for the normal growth of plants. It participates in the process of nitrogen metabolism in plants. Under the condition of lack of molybdenum, plants accumulate more nitrate nitrogen and thus show low nitrogen use efficiency (nitrogen utilization efficiency, NUE). Increasing nitrogen use rate can reduce the excessive application of chemical fertilizer. The negative effect is not clear, but the mechanism of increasing nitrogen utilization rate of molybdenum is not clear. This study was carried out under the conditions of soil free cultivation and tissue culture in greenhouse. The experimental materials of "Fragaria x ananassa Duch.cv. 'Akihime") strawberry were used to spray different concentrations of sodium molybdate (sodium molybdate, N) under the condition of soil free cultivation in greenhouse. A_2MoO_4), the experiment set up T1 (foliar spraying water), T2 (0.02%Na_2MoO_4), T3 (0.04%Na_2MoO_4), T4 (0.05%Na_2MoO_4) and T5 (0.06%Na_2MoO_4). The plant biomass, fruit nutrition element content and fruit quality (total soluble), ascorbic acid, titratable acid were studied. Table acidity, TA), photosynthesis, MOT1 expression of molybdenum transporter gene, Mo concentration in the plant, activity of key enzymes in nitrogen metabolism (nitrate reductase, nitrate reductase, NR; nitrite reductase, NiR; glutamine synthetase, glutamine synthetase, GS; glutamate synthase, depletion), nitrogen metabolism The expression of key genes (NRT1.1, NRT2.1, NIA, NiR, GS-1, NADH-GOGAT), the effect of 15N absorption, distribution and utilization. Under the condition of tissue culture, Na_2MoO_4 or sodium tungstate (sodium tungstate, Na2WO4) in the solid culture medium of MS (sodium tungstate, Na2WO4), 4 treatments. The plant phenotype, root microstructure and root activity, leaf microstructure and ultrastructure and chlorophyll content, MOT1 expression, the expression of key genes (CNX2, CNX3, CNX5, CNX6, CNX7, CNX1), the concentration of Mo in the plant, the activity of the key enzymes of nitrogen metabolites (NR, NiR, GS, NADH-GOGAT), and the effects of the utilization of the plants. The results are as follows: 1. the suitable concentration of Mo supply can significantly increase the biomass of the seedlings. Spraying Mo fertilizer on the leaf surface can improve the content of nutrient elements (N, Mo, Fe and Cu), TSS, TA and aromatic substance in strawberry fruit. However, the application of Mo on Se and some aromatic substances in the fruit has no significant effect on the treatment of.0.04%Na_2MoO_4 in all treatments. The highest TSS, N and Fe concentrations of.0.02%Na_2MoO_4 were significantly higher than those of the control. A total of 6 kinds of esters (methyl butyrate, gamma decanide, ethyl butyrate, methyl Hexanate, gamma twelve lactone, gamma twelve lactone, ethyl caproate, ethyl hexanoate) were identified by headspace solid phase microextraction and GC-MS. The content of Mo in the roots of the strawberry seedlings with the highest content of.2., the content of the stems and leaves increased with the increase of Mo, and the expression level of MOT1 in the root of the.0.04%Na_2MoO_4 treatment was higher than that of the other treatments. When the concentration of Na_2MoO_4 increased from 0.05% to 0.06%, the mRNA level of MOT1 in the root showed a decline trend, and the leaves was MO after 5 days. The expression level of T1 increased with the increase of the concentration of Na_2MoO_4. When the concentration of Na_2MoO_4 increased from 0% to 0.04% after 10 and 15 days, the expression level of MOT1 in leaves showed a trend of increase, and then decreased with the increase of spraying Na_2MoO_4. Mo influenced Fe, Cu and Se.0.04%Na_2MoO_4 in the roots of seedlings. The seedlings of the treated seedlings showed the highest 15N upper / underground ratio and the 15N utilization rate.0.04%Na_2MoO_4 treated with higher nitrogen metabolism key enzyme activity, and the expression level of nitrate nitrogen absorption gene (NRT1.1 and NRT2.1) and nitrate nitrogen response gene up regulated the NO3--N concentration of the root and leaf of.0.04%Na_2MoO_4 treatment was lower than that of other treatments. The concentration of NH4+-N was higher and the concentration of Gln/Glu was higher, while T5 treatment showed the opposite trend. Therefore, the foliar spraying 0.04%Na_2MoO_4 enhanced the transport of NO3- to the top of the root by regulating the activity of key enzyme activities of nitrogen metabolism and the expression of NO3- assimilation related genes, and increased the normal addition of Na_ in the solid-state medium of nitrogen utilization ratio.3.MS. The growth trend of strawberry seedlings in 2MoO_4 is better, the leaf is normal green, the microstructure and ultrastructure of the mesophyll cells are complete, the chloroplasts are more, the chlorophyll content is higher, the root development is normal, the root activity is high, the root tip cells are arranged neatly and closely, the central marrow cells are large and arranged closely. The leaves are short under the leaves of the seedlings grown on the Mo medium. The epidermal cells were not arranged regularly, the ultrastructure of the mesophyll cells was abnormal, the organelles and the intima system were damaged, the chlorophyll content was low, the leaves were yellow and the roots were long, but the growth potential was weak, the root tip cells were slightly deformed, the cells were loosely arranged and the cell gaps were large. The plant types on the medium with Na2WO4 and Na_2MoO_4 and Na2WO4 were smaller. The cell organ is impaired, the organelle is damaged, the membrane structure is not clear, some leaves are yellow, the content of chlorophyll is reduced, and there is a similar symptom with nitrogen deficiency. Na_2MoO_4 can significantly induce the expression of FaMOT1 in the upper and root parts of the strawberry, and the amount of the FaMOT1 surface of the inhibitor Na2WO4 shows a relative decline and is applied to the Na_2MoO_4 seedlings. The concentration of Mo was significantly higher than that of other treatments. The expression of CNX2 and CNX3 in tissue culture seedlings with Na_2MoO_4 was higher. The expression of CNX7 and CNX1 in the tissue culture medium with Na_2MoO_4 was higher than that of the tissue culture seedlings which were added to the culture medium. The addition of inhibitor Na2WO4 made the NR activity significantly lower, and the NR activity of the group of Na_2MoO_4 tissue cultured seedlings was the highest. The key enzyme of nitrogen metabolism (NiR, GS and NADH-GOGAT) showed a similar trend. The concentration of NO3--N in the culture medium with Na_2MoO_4 was lower than that of other treatments, and the concentration of NO2--N and NH4+-N was significantly higher than that of other treatments. The isotopic 15N-Ca (NO3) 2 marker further indicated that the tissue culture seedlings of the culture medium added to Na_2MoO_4 were absorbed and utilized more 15N and transported to the ground. There were more 15N in the upper part, and the highest utilization rate of 15N in plants. The 15N utilization rate of tissue culture seedlings with inhibitors was lower.
【學(xué)位授予單位】：山東農(nóng)業(yè)大學(xué)
【學(xué)位級(jí)別】：博士
【學(xué)位授予年份】：2017
【分類(lèi)號(hào)】：S668.4
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