缺硼对脐橙幼苗硼分配及叶片细胞壁组分硼含量的影响
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摘要
【目的】硼在维持细胞壁正常结构方面具有重要的作用,前期结果证实缺硼严重的脐橙叶片细胞壁结构改变程度也更大,但这种变化与细胞壁组分中硼的含量变化是否有关尚不清楚。本研究通过分析缺硼对脐橙幼苗各部分硼分配及叶片细胞壁组分硼含量的影响,明确缺硼症状表现及细胞壁结构变化程度与细胞壁各组分中硼含量变化之间的关系。【方法】以纽荷尔脐橙幼苗为试材,利用营养液培养方法进行缺硼处理,测定根、砧木茎、接穗茎、上部叶、下部叶、叶片细胞壁以及细胞壁各组分硼含量的变化情况。【结果】缺硼处理9周后上部叶出现叶片卷曲及叶片失绿等症状,而下部叶没有出现任何可见的症状。缺硼处理的脐橙幼苗各部位硼含量和硼吸收量均显著降低,缺硼降低了硼向地上部的相对分配比例且上部叶受到的影响程度更大。在硼正常供应条件下,上部叶和下部叶游离态硼、原生质体硼和细胞壁硼的含量和相对分配比例没有显著差异,说明硼在不同类型脐橙叶片细胞各组分中的分配是相对稳定的。缺硼后水溶性硼(包括游离态硼和原生质体硼)在脐橙上部叶和下部叶中都降到极低的水平,尤其是原生质体硼百分含量下部叶甚至是低于上部叶的。缺硼后细胞壁硼占总硼的比例则由22%左右增加到80%以上。与叶片中硼含量的变化趋势一致,缺硼以后虽然上部叶和下部叶细胞壁硼含量都显著降低,但上部叶降低的程度远大于下部叶。进一步分析细胞壁组分硼含量变化,发现缺硼显著降低了上部叶细胞壁中离子结合态果胶硼含量而对下部叶的无明显影响,其他组分硼含量的变化趋势下部叶和上部叶一致。【结论】原生质体硼含量的高低并不是决定缺硼症状的主要因素,离子结合态果胶与硼的结合能力对缺硼条件下细胞壁的结构及缺硼症状表现起着至关重要的作用。
【Objectives】Boron(B) plays an important role in maintaining the normal structure of cell walls. The previous results showed that the cell wall structure of leaves with severe B deficiency had a lot of change, but it was not clear whether this change was related to the change of B concentration in the cell wall components. This paper aims to clarify the relationship between B deficiency symptoms and changes of cell wall structure and B content in cell wall component.【Methods】In this study, the changes of B distribution in different plant parts and B concentrations in leaf cell wall and cell wall components of navel orange plants were measured using hydroponics method.【Results】The symptoms of leaf chlorosis and curling of the leaves were observed only in the upper leaves of B-deficient plants. The lower leaves did not show any visible symptoms. The B concentration and B accumulation in different parts of the boron-deficient navel orange plants were significantly decreased.Boron deficiency reduced the relative distribution ratio of B in the shoots and the greater impact on the upper leaves. There was no evident difference in concentrations of free B, protoplast B and cell wall B in leaves between the two types of leaves under normal B condition. However, the water-soluble B(including free B and protoplast B) decreased to extremely low levels in the upper and lower leaves of navel orange under B deficiency, especially the protoplast B in the lower leaves was even lower than that in the upper leaves. In control plants, the upper and lower leaves contained only about 22% of cellular B in the cell wall, whereas in B-deficient plants the cell walls of the upper and lower leaves contained >80% of total cell B. Furthermore, the concentrations of B in lower leaves and their cell walls were much higher than those in the upper leaves and their cell walls under B-deficient conditions. Further analysis of the changes of B content in cell wall components showed that B deficiency significantly reduced the boron content in the ion-bound pectin in the upper leaf cell wall and had no significant effect on that in the lower leaf cell wall. In contrast, no significant difference was found in changes of B content in other wall components due to B deficiency between the upper leaves and lower leaves. In addition, the structural changes of the upper leaf cell wall were much larger than those of the lower leaf cell wall under B deficiency.【Conclusions】Boron concentration in the protoplast was not the main factor determining the symptoms of B deficiency, while the binding capacity of ion-bound pectin to B was crucial to the structure of cell wall and the occurrence of B deficiency symptoms in navel orange leaves.
【Objectives】Boron(B) plays an important role in maintaining the normal structure of cell walls. The previous results showed that the cell wall structure of leaves with severe B deficiency had a lot of change, but it was not clear whether this change was related to the change of B concentration in the cell wall components. This paper aims to clarify the relationship between B deficiency symptoms and changes of cell wall structure and B content in cell wall component.【Methods】In this study, the changes of B distribution in different plant parts and B concentrations in leaf cell wall and cell wall components of navel orange plants were measured using hydroponics method.【Results】The symptoms of leaf chlorosis and curling of the leaves were observed only in the upper leaves of B-deficient plants. The lower leaves did not show any visible symptoms. The B concentration and B accumulation in different parts of the boron-deficient navel orange plants were significantly decreased.Boron deficiency reduced the relative distribution ratio of B in the shoots and the greater impact on the upper leaves. There was no evident difference in concentrations of free B, protoplast B and cell wall B in leaves between the two types of leaves under normal B condition. However, the water-soluble B(including free B and protoplast B) decreased to extremely low levels in the upper and lower leaves of navel orange under B deficiency, especially the protoplast B in the lower leaves was even lower than that in the upper leaves. In control plants, the upper and lower leaves contained only about 22% of cellular B in the cell wall, whereas in B-deficient plants the cell walls of the upper and lower leaves contained >80% of total cell B. Furthermore, the concentrations of B in lower leaves and their cell walls were much higher than those in the upper leaves and their cell walls under B-deficient conditions. Further analysis of the changes of B content in cell wall components showed that B deficiency significantly reduced the boron content in the ion-bound pectin in the upper leaf cell wall and had no significant effect on that in the lower leaf cell wall. In contrast, no significant difference was found in changes of B content in other wall components due to B deficiency between the upper leaves and lower leaves. In addition, the structural changes of the upper leaf cell wall were much larger than those of the lower leaf cell wall under B deficiency.【Conclusions】Boron concentration in the protoplast was not the main factor determining the symptoms of B deficiency, while the binding capacity of ion-bound pectin to B was crucial to the structure of cell wall and the occurrence of B deficiency symptoms in navel orange leaves.
引文
[1]Voxeur A,Fry S C.Glycosylinositol phosphorylceramides from Rosa cell cultures are boron-bridged in the plasma membrane and form complexes with rhamnogalacturonan II[J].The Plant Journal,2014,79:139-149.
[2]Brown P H,Bellaloui N,Wimmer M A,et al.Boron in plant biology[J].Plant Biology,2002,4:205-223.
[3]O’Neill M A,Eberhard S,Albersheim P,et al.Requirement of borate cross-linking of cell wall rhamnogalacturonan II for Arabidopsis growth[J].Science,2001,294:846-849.
[4]姜存仓,王运华,刘桂东,等.赣南脐橙叶片黄化及施硼效应研究[J].植物营养与肥料学报,2009,15(3):656-661.Jiang C C,Wang Y H,Liu G D,et al.Effect of boron on the leaves etiolation and fruit fallen of Newhall navel orange[J].Plant Nutrition and Fertilizer Science,2009,15(3):656-661.
[5]Liu G D,Jiang C C,Wang Y H.Distribution of boron and its forms in young‘Newhall’navel orange(Citrus sinensis Osb.)plants grafted on two rootstocks in response to deficient and excessive boron[J].Soil Science and Plant Nutrition,2011,57:93-104.
[6]Matoh T,Ishigaki K,Ohno K,et al.Isolation and characterization of a boron-polysaccharide complex from radish roots[J].Plant and Cell Physiology,1993,34:639-642.
[7]Kobayashi M,Matoh T,Azuma J.Two chains of rhamnogalacturonan II are cross-linked by borate-diol ester bonds in higher plant cell walls[J].Plant Physiology,1996,110:1017-1020.
[8]O’Neill M A,Warrenfeltz D,Kates K,et al.Rhamnogalacturonan-II,a pectic polysaccharide in the walls of growing plant cell,forms a dimer that is covalently cross-linked by a borate ester[J].The Journal of Biological Chemistry,1996,271:22923-22930.
[9]Ishii T,Matsunaga T,Pellerin P,et al.The plant cell wall polysaccharide rhamnogalacturonan II self-assembles into a covalently cross-linked dimer[J].The Journal of Biological Chemistry,1999,274:13098-13104.
[10]Chormova D,Messenger D J,Fry S C.Boron bridging of rhamnogalacturonan-II,monitored by gel electrophoresis,occurs during polysaccharide synthesis and secretion but not postsecretion[J].The Plant Journal,2014,77:534-546.
[11]Kaku T,Tabuchi A,Wakabayashi K,et al.Action of xyloglucan hydrolase within the native cell wall architecture and its effect on cell wall extensibility in azuki bean epicotyls[J].Plant and Cell Physiology,2002,43:21-26.
[12]Sakurai N.Cell wall functions in growth and development,a physical and a chemical point of view[J].Curtis's Botanical Magazine,1991,104:235-251.
[13]Hoson T.Apoplast as the site of response to environmental signals[J].Journal of Plant Research,1998,111:167-177.
[14]Liu G D,Dong X C,Liu L C,et al.Boron deficiency is correlated with changes in cell wall structure that lead to growth defects in the leaves of navel orange plants[J].Scientia Horticulturae,2014,176:54-62.
[15]Goldbach H E,Wimmer M A,Findeklee P.Discussion paper:BoronHow can the critical level be defined?[J].Journal of Plant Nutrition and Soil Science,2000,163:115-121.
[16]Dannel F,Pfeffer H,R?mheld V.Compartmentation of boron in roots and leaves of sunflower as affected by boron supply[J].Journal of Plant Physiology,1998,153:615-622.
[17]Pan Y,Wang Z H,Yang L,et al.Differences in cell wall components and allocation of boron to cell walls confer variations in sensitivities of Brassica napus cultivars to boron deficiency[J].Plant and Soil,2012,354:383-394.
[18]Hu H,Brown P H.Localization of boron in cell walls of squash and tobacco and its association with pectin(Evidence for a structural role of boron in the cell wall)[J].Plant Physiology,1994,105:681-689.
[19]Liu G D,Wang R D,Liu L C,et al.Cellular boron allocation and pectin composition in two citrus rootstock seedlings differing in boron-deficiency response[J].Plant and Soil,2013,370:555-565.
[20]Hoagland D R,Arnon D I.The water-culture method for growing plants without soil[J].California Agricultural Experiment Station Circular,1950,347:1-32.
[21]Redgwell R J,Selvendran R R.Structural features of cell-wall polysaccharides of onion Allium cepa[J].Carbohydrate Research,1986,157:183-199.
[22]M?tt?nen M,Lehto T,Aphalo P J.Growth dynamics and mycorrhizas of Norway spruce(Picea abies)seedlings in relation to boron supply[J].Trees,2001,15:319-326.
[23]Asad A,Blamey F P C,Edwards D G.Dry matter production and boron concentration of vegetative and reproductive tissues of canola and sunflower plants grown in nutrition solution[J].Plant and Soil,2002,243(2):243-252.
[24]Dong X C,Liu G D,Wu X W,et al.Different metabolite profile and metabolic pathway with leaves and roots in response to boron deficiency at the initial stage of citrus rootstock growth[J].Plant Physiology and Biochemistry,2016,108:121-131.
[25]Liu G D,Wang R D,Wu L S,et al.Boron distribution and mobility in navel orange grafted on citrange and trifoliate orange[J].Plant and Soil,2012,360:123-133.
[26]杨玉华,王运华,杜昌文,等.硼对不同硼效率甘蓝型油菜品种细胞壁组成的影响[J].植物营养与肥料学报,2002,8(3):340-343.Yang Y H,Wang Y H,Du C W,et al.Study on the influence of B on cell wall components in different B efficiency rape cultivars[J].Plant Nutrition and Fertilizer Science,2002,8(3):340-343.
[2]Brown P H,Bellaloui N,Wimmer M A,et al.Boron in plant biology[J].Plant Biology,2002,4:205-223.
[3]O’Neill M A,Eberhard S,Albersheim P,et al.Requirement of borate cross-linking of cell wall rhamnogalacturonan II for Arabidopsis growth[J].Science,2001,294:846-849.
[4]姜存仓,王运华,刘桂东,等.赣南脐橙叶片黄化及施硼效应研究[J].植物营养与肥料学报,2009,15(3):656-661.Jiang C C,Wang Y H,Liu G D,et al.Effect of boron on the leaves etiolation and fruit fallen of Newhall navel orange[J].Plant Nutrition and Fertilizer Science,2009,15(3):656-661.
[5]Liu G D,Jiang C C,Wang Y H.Distribution of boron and its forms in young‘Newhall’navel orange(Citrus sinensis Osb.)plants grafted on two rootstocks in response to deficient and excessive boron[J].Soil Science and Plant Nutrition,2011,57:93-104.
[6]Matoh T,Ishigaki K,Ohno K,et al.Isolation and characterization of a boron-polysaccharide complex from radish roots[J].Plant and Cell Physiology,1993,34:639-642.
[7]Kobayashi M,Matoh T,Azuma J.Two chains of rhamnogalacturonan II are cross-linked by borate-diol ester bonds in higher plant cell walls[J].Plant Physiology,1996,110:1017-1020.
[8]O’Neill M A,Warrenfeltz D,Kates K,et al.Rhamnogalacturonan-II,a pectic polysaccharide in the walls of growing plant cell,forms a dimer that is covalently cross-linked by a borate ester[J].The Journal of Biological Chemistry,1996,271:22923-22930.
[9]Ishii T,Matsunaga T,Pellerin P,et al.The plant cell wall polysaccharide rhamnogalacturonan II self-assembles into a covalently cross-linked dimer[J].The Journal of Biological Chemistry,1999,274:13098-13104.
[10]Chormova D,Messenger D J,Fry S C.Boron bridging of rhamnogalacturonan-II,monitored by gel electrophoresis,occurs during polysaccharide synthesis and secretion but not postsecretion[J].The Plant Journal,2014,77:534-546.
[11]Kaku T,Tabuchi A,Wakabayashi K,et al.Action of xyloglucan hydrolase within the native cell wall architecture and its effect on cell wall extensibility in azuki bean epicotyls[J].Plant and Cell Physiology,2002,43:21-26.
[12]Sakurai N.Cell wall functions in growth and development,a physical and a chemical point of view[J].Curtis's Botanical Magazine,1991,104:235-251.
[13]Hoson T.Apoplast as the site of response to environmental signals[J].Journal of Plant Research,1998,111:167-177.
[14]Liu G D,Dong X C,Liu L C,et al.Boron deficiency is correlated with changes in cell wall structure that lead to growth defects in the leaves of navel orange plants[J].Scientia Horticulturae,2014,176:54-62.
[15]Goldbach H E,Wimmer M A,Findeklee P.Discussion paper:BoronHow can the critical level be defined?[J].Journal of Plant Nutrition and Soil Science,2000,163:115-121.
[16]Dannel F,Pfeffer H,R?mheld V.Compartmentation of boron in roots and leaves of sunflower as affected by boron supply[J].Journal of Plant Physiology,1998,153:615-622.
[17]Pan Y,Wang Z H,Yang L,et al.Differences in cell wall components and allocation of boron to cell walls confer variations in sensitivities of Brassica napus cultivars to boron deficiency[J].Plant and Soil,2012,354:383-394.
[18]Hu H,Brown P H.Localization of boron in cell walls of squash and tobacco and its association with pectin(Evidence for a structural role of boron in the cell wall)[J].Plant Physiology,1994,105:681-689.
[19]Liu G D,Wang R D,Liu L C,et al.Cellular boron allocation and pectin composition in two citrus rootstock seedlings differing in boron-deficiency response[J].Plant and Soil,2013,370:555-565.
[20]Hoagland D R,Arnon D I.The water-culture method for growing plants without soil[J].California Agricultural Experiment Station Circular,1950,347:1-32.
[21]Redgwell R J,Selvendran R R.Structural features of cell-wall polysaccharides of onion Allium cepa[J].Carbohydrate Research,1986,157:183-199.
[22]M?tt?nen M,Lehto T,Aphalo P J.Growth dynamics and mycorrhizas of Norway spruce(Picea abies)seedlings in relation to boron supply[J].Trees,2001,15:319-326.
[23]Asad A,Blamey F P C,Edwards D G.Dry matter production and boron concentration of vegetative and reproductive tissues of canola and sunflower plants grown in nutrition solution[J].Plant and Soil,2002,243(2):243-252.
[24]Dong X C,Liu G D,Wu X W,et al.Different metabolite profile and metabolic pathway with leaves and roots in response to boron deficiency at the initial stage of citrus rootstock growth[J].Plant Physiology and Biochemistry,2016,108:121-131.
[25]Liu G D,Wang R D,Wu L S,et al.Boron distribution and mobility in navel orange grafted on citrange and trifoliate orange[J].Plant and Soil,2012,360:123-133.
[26]杨玉华,王运华,杜昌文,等.硼对不同硼效率甘蓝型油菜品种细胞壁组成的影响[J].植物营养与肥料学报,2002,8(3):340-343.Yang Y H,Wang Y H,Du C W,et al.Study on the influence of B on cell wall components in different B efficiency rape cultivars[J].Plant Nutrition and Fertilizer Science,2002,8(3):340-343.