基于叶片显微及亚显微结构的新疆扁桃10个主栽品种抗旱性综合评价
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摘要
【目的】从叶片形态解剖角度综合评价新疆10个扁桃(Amygdalus communis L.)主栽品种的抗旱性,为扁桃抗旱性评价指标体系的建立提供理论依据。【方法】通过光学和扫描电子显微镜观察叶片显微及亚显微结构相关20项指标并进行生物统计学分析。【结果】所有品种叶片均形成不同程度的旱生结构特征,角质层、栅栏细胞和维管系统均较发达,有含晶细胞和粘液细胞分布、气孔只分布于下表皮等。通过主成分分析,选取主脉厚度、叶比面积、叶肉细胞紧实度和气孔密度4项代表性指标。隶属函数评价结果表明,其抗旱能力由大到小排序为S14> S3> M7> M8>M1> M2> S12> S1>S9> S11。【结论】初步确定S14和S3的抗旱能力较强,而S9和S11的抗旱能力较弱,美国品种均属于中度抗旱类型。
【Objective】The almond(Amygdalus communis L.)is one of the four most famous nuts in the world.It has a long history of cultivation, and has important nutritional and ecological value. As one of the origin of Almond in the world, Southern Xinjiang is rich in almond resistant resources. In this study, the drought resistant capacity of 10 different almond cultivars including 6 main local cultivars and 4 main American introduced cultivars collected from the Southern Xinjiang were evaluated by means of leaf microstructure and ultrastructure characteristics in order to provide a theoretical basis for the establishment of evaluation index system for drought resistance of almond,and to help understanding the drought resistant mechanism of almond from the anatomy aspects.【Methods】Fully-expanded matured leaves on one–year old twigs were sampled from 10 different almond cultivars on local peach rootstock from the southern Xinjiang. Leaf samples were fixed in FAA and 2.5% glutaraldehyde separately. Paraffin sections were made. At least 4 different leaves were used for cultivar, and the mesophyll cells structure and main vein structure were observed under the light microscopy; the scanning electronic microscopy was used for observing the ultrastructure of epidermis(4 different leaves for each cultivar, 2 of them used for observing upper epidermis, other 2 for observing lower epidermis). Besides of the anatomical structures, the SLA(specific leaf area) value of each cultivar was also calculated at least from 20 representative leaves. Except for the regular leaf anatomical indicators, crystal cell density(number of crystal cells per um2 mesophyll tissue), mucous cell distributed thickness(measured at least from 8 different points) and cell porosity ratio(ratio of intercellular space at mesophyll cells of the paraxial surface to the total area of mesophyll cells of the paraxial surface) etc., xerophytic plant related several leaf structural features were quantified. Thickness of cuticle, epidermis, palisade cell and total leaf were measured under 10×20 magnifications, and the thickness of main vain, vascular bundle, phloem and xylem vessel diameter etc. were measured at 10×4 magnifications by using Image-J software.Comparisons of total 20 leaf anatomical indicators between cultivars were made with a one-way ANOVA. The principal component analysis was used for further screening the main representative indicators.The drought resistant degrees of different cultivars were evaluated by subordinate function values analysis. Pearson correlation analysis was carried out to further understand the interrelations between anatomical indicators.【Results】Under the same drought environmental conditions, the leaves of the 10 different almond cultivars exhibited different degrees of drought resistant anatomical structure. The cuticle,palisade tissue cells, main veins and lateral vascular bundles were well developed. There were crystal cells in the mesophyll tissue, and mucous cells around the vascular bundles. The stomata were distributed only in the lower epidermis. Among the 20 anatomical structure indexes, 18 indexes were differnt significantly between cultivars(p < 0.05), and the CV was 2.17% to 27.44%. There were significant differences in some anatomical features between local cultivars and American introduced cultivars, such as SLA, LT, CTR%, thickness of epidermis and cuticle, etc. Total 13 drought resistant-related anatomical indexes were selected among 18 indexes based on the published research papers related to leaf anatomy and drought resistance, and also the leaf structural features of xerophytic plants. The main component analysis was carried out to further screen the representative indexes from 13 drought resistant related indexes. The cumulative contribution rate of the first four principal components was about 83%, the main vein thickness(MVT), specific leaf area(SLA), cell tightness ratio(CTR%) and stomata density(SD) were selected as the four principal component for drought resistant indicators. The results of subordinate function values analysis showed that the drought resistance capacity of 10 almond cultivars was S14>S3> M7 > M8>M1> M2> S12 > S1 >S9> S11. According to the Pearson correlation analysis,there were negative significant correlations between SLA and TL, LCT, indicating that the almond with lower SLA had thicker leaf lamina and cuticles(p < 0.05). There were close positive correlations between cuticle thickness, lateral vein density and mucous cell distributed thickness(p < 0.05), and this was one of the drought adaptive strategies of almond for surviving in the extremely drought environment.【Conclusion】The leaf tissue structures of American introduced cultivars were very similar to each other and there were large differences in the leaf tissue structures among Xinjiang local cultivars.Among 10 almond cultivars, Xinjiang local cultivars of S14 and S3 showed higherger drought resistance than other cultivars; the drought resistance capacity of American introduced cultivars was moderate and local cultivars of S11, S9 showed weaker drought resistance.
【Objective】The almond(Amygdalus communis L.)is one of the four most famous nuts in the world.It has a long history of cultivation, and has important nutritional and ecological value. As one of the origin of Almond in the world, Southern Xinjiang is rich in almond resistant resources. In this study, the drought resistant capacity of 10 different almond cultivars including 6 main local cultivars and 4 main American introduced cultivars collected from the Southern Xinjiang were evaluated by means of leaf microstructure and ultrastructure characteristics in order to provide a theoretical basis for the establishment of evaluation index system for drought resistance of almond,and to help understanding the drought resistant mechanism of almond from the anatomy aspects.【Methods】Fully-expanded matured leaves on one–year old twigs were sampled from 10 different almond cultivars on local peach rootstock from the southern Xinjiang. Leaf samples were fixed in FAA and 2.5% glutaraldehyde separately. Paraffin sections were made. At least 4 different leaves were used for cultivar, and the mesophyll cells structure and main vein structure were observed under the light microscopy; the scanning electronic microscopy was used for observing the ultrastructure of epidermis(4 different leaves for each cultivar, 2 of them used for observing upper epidermis, other 2 for observing lower epidermis). Besides of the anatomical structures, the SLA(specific leaf area) value of each cultivar was also calculated at least from 20 representative leaves. Except for the regular leaf anatomical indicators, crystal cell density(number of crystal cells per um2 mesophyll tissue), mucous cell distributed thickness(measured at least from 8 different points) and cell porosity ratio(ratio of intercellular space at mesophyll cells of the paraxial surface to the total area of mesophyll cells of the paraxial surface) etc., xerophytic plant related several leaf structural features were quantified. Thickness of cuticle, epidermis, palisade cell and total leaf were measured under 10×20 magnifications, and the thickness of main vain, vascular bundle, phloem and xylem vessel diameter etc. were measured at 10×4 magnifications by using Image-J software.Comparisons of total 20 leaf anatomical indicators between cultivars were made with a one-way ANOVA. The principal component analysis was used for further screening the main representative indicators.The drought resistant degrees of different cultivars were evaluated by subordinate function values analysis. Pearson correlation analysis was carried out to further understand the interrelations between anatomical indicators.【Results】Under the same drought environmental conditions, the leaves of the 10 different almond cultivars exhibited different degrees of drought resistant anatomical structure. The cuticle,palisade tissue cells, main veins and lateral vascular bundles were well developed. There were crystal cells in the mesophyll tissue, and mucous cells around the vascular bundles. The stomata were distributed only in the lower epidermis. Among the 20 anatomical structure indexes, 18 indexes were differnt significantly between cultivars(p < 0.05), and the CV was 2.17% to 27.44%. There were significant differences in some anatomical features between local cultivars and American introduced cultivars, such as SLA, LT, CTR%, thickness of epidermis and cuticle, etc. Total 13 drought resistant-related anatomical indexes were selected among 18 indexes based on the published research papers related to leaf anatomy and drought resistance, and also the leaf structural features of xerophytic plants. The main component analysis was carried out to further screen the representative indexes from 13 drought resistant related indexes. The cumulative contribution rate of the first four principal components was about 83%, the main vein thickness(MVT), specific leaf area(SLA), cell tightness ratio(CTR%) and stomata density(SD) were selected as the four principal component for drought resistant indicators. The results of subordinate function values analysis showed that the drought resistance capacity of 10 almond cultivars was S14>S3> M7 > M8>M1> M2> S12 > S1 >S9> S11. According to the Pearson correlation analysis,there were negative significant correlations between SLA and TL, LCT, indicating that the almond with lower SLA had thicker leaf lamina and cuticles(p < 0.05). There were close positive correlations between cuticle thickness, lateral vein density and mucous cell distributed thickness(p < 0.05), and this was one of the drought adaptive strategies of almond for surviving in the extremely drought environment.【Conclusion】The leaf tissue structures of American introduced cultivars were very similar to each other and there were large differences in the leaf tissue structures among Xinjiang local cultivars.Among 10 almond cultivars, Xinjiang local cultivars of S14 and S3 showed higherger drought resistance than other cultivars; the drought resistance capacity of American introduced cultivars was moderate and local cultivars of S11, S9 showed weaker drought resistance.
引文
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[3]杨波,车玉红,郭春苗,龚鹏,徐叶挺,孙涛,刘河疆.扁桃生理落果期不同组织激素浓度的动态变化及其对落果的影响[J].西北植物学报,2015,35(1):0118-0124.YANG Bo,CHE Yuhong,GUO Chunmiao,GONG Peng,XUYeting,SUN Tao,LIU Hejiang.Dynamic changes of hormones in the different tissues of Almond during the physiological fruit drop and its effect on fruit drop[J].Acta Botanica Boreali-Occidentalia Sinica,2015,35(1):0118-0124.
[4]TORRECILLAS A,ALARCON J J,DOMINGO R,PLANES J,SANCHEZ-BLANCO M J.Strategies for drought resistance in leaves of two almond cultivars[J].Plant Science,1996,118(2):135-143.
[5]YADOLLAHI A,ARZANI K,EBADI A,WIRTHENSOHN M,KARIMI S.The response of different almond genotypes to moderate and severe water stress in order to screen for drought tolerance[J].Scientia Horticulturae,2011,129(3):403-413.
[6]EHSAN A,ALI L,SHAHIN FY.Physiological and Morphological Responses of Almond Cultivars under In Vitro Drought Stress[J].Journal of Nuts,2017,8(1):61-72
[7]郭改改,封斌,麻保林,张应龙,郭春会,井赵斌.不同区域长柄扁桃叶片解剖结构及其抗旱性分析[J].西北植物学报,2013,33(4):720-728.GUO Gaigai,FENG Bin,MA Baolin,ZHANG Yinglong,GUOChunhui,JING Zhaobin.Leaf anatomical structures of different regional Amygdalus pedunculata Pall.and their drought resistance analysis[J].Acta Botanica Boreali-Occidentalia Sinica,2013,33(4):720-728.
[8]WANG J,ZHENG R,BAI S,GAO X,LIU M,YAN W.Mongolian Almond(Prunus mongolica Maxim):The Morpho-Physiological,Biochemical and Transcriptomic Response to Drought Stress[J].Plos One,2015,10(4):e0124442.
[9]王丽娜,克热木·伊力,侯江涛.水分胁迫对扁桃砧木叶片脯氨酸、可溶性蛋白质、质膜透性、相对含水量的影响[J].新疆农业大学学报,2006,29(3):53-58.WANG Lina,YILI·Keremu,HOU Jiangtao.Effects of water stress on proline,soluble protein,membrane permeability,relative water content of Almond rootstock[J].Journal of Xinjiang Agricultural University,2006,29(3):53-58.
[10]克热木·伊力,王丽娜,侯江涛.水分胁迫对扁桃砧木干物质和叶绿素含量的影响[J].经济林研究,2007,25(4):1-5.YILI·Keremu,WANG Lina,HOU Jiangtao.Effects of Water Stress on Contents of Dry Matters and Chlorophyll in Almond Rootstock[J].Nonwood Forest Research,2007,25(4):1-5.
[11]李芳兰,包维楷.植物叶片形态解剖结构对环境变化的响应与适应[J].植物学通报,2005,22(增刊):118-127LI Fanglan,BAO Werkai.Responses of the morphological and anatomical structure of the plant leaf to environmental change[J].Chinese Bulletin of Botany,2005,22(Suppl.):118-127
[12]王耀芝,王勋陵,李蔚.荒漠化草原常见植物叶内部结构的观察[J].兰州大学学报,1983(3):90-99.WANG Yaozhi,WANG Xunling,LI Yu.Observation on leaf structure of some species in desert steppe[J].Journal of Lanzhou University(Natural Science Eddition),1983(3):90-99.
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