苹果砧木八棱海棠和M9T337的根系分布及水分输导组织特征
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摘要
[目的]本文旨在研究砧木乔化/矮化和抗旱能力差别的生理机制,并探索不同土层深度根水分输导组织结构和导水能力的差别,以期为生产上合理制定栽培管理技术措施提供理论依据和试验证据。[方法]本研究测定了常用乔化砧木八棱海棠和矮化砧木M9T337的地上部分生长情况、根分布,同时测定了枝条和不同土层深度的根的木质部导管特征,并根据木质部水分输导组织特征计算了枝条和不同土层深度的根的导水能力。[结果]八棱海棠的树高、基径、枝条数、枝条长度均显著高于M9T337(P<0.05);八棱海棠在不同区域的根生物量和粗根根长密度均显著高于M9T337(P<0.05),细根根长密度除个别区域外均显著高于M9T337(P<0.05),八棱海棠的根垂直分布范围为0～60 cm,M9T337为0～40 cm;八棱海棠根和茎的导管直径和导水率均显著大于M9T337(P<0.05),根和茎的导管密度显著低于M9T337(P<0.05);两个树种深层根的导管直径和导水率均高于浅层根。[结论]八棱海棠根和茎的导水能力更大,根长密度大,根系垂直分布范围广,是其乔化和抗旱能力强的重要原因。深层根有助于根系吸收深层土壤的水分,可能是提升树木抗旱能力的重要因素。
[Objective] In order to explore physiological mechanisms and drought tolerance of apple rootstock vigorous/dwarfing, the structure and water-transport ability of root system at various soil depth of two common apple rootstock species were investigated, and research was aimed to provide technical support for fruit production management. [Methods] Vigorous rootstock Malus micromalu and dwarfing rootstock M9 T337 were used as experimental materials. The growth condition, root distribution, and xylem vessel trait of both branch and root system at different soil depth were measured. Hydraulic conductivity of root and branch was determined based on the xylem vessel traits. [Results] Study results indicated that the major parameters including plant height, basal diameter, and the number and length of branch of M. micromalu were significantly higher than that of M9 T337, and both root biomass and root length density of M. micromalu with a vertical distribution of 0～60 cm were significantly greater than that of M9 T337 with vertical distribution of 0～40 cm. Vessel diameter and hydraulic conductivity of both root and branch of M. micromalu were significantly higher than that of M9 T337, while vessel density of root and branch was significantly lower in M. micromalu, and vessel diameter and hydraulic conductivity of deep roots were greater than that of shallow roots for both species. [Conclusion] M. micromalu possessed more efficient vessels, higher root length density and wider vertical root distribution, which greatly contributed to its vigorous effect and strong drought resistance. The deep root, conducting water absorbing from deep soil, may act as an important factor for improving tree's drought resistance.
[Objective] In order to explore physiological mechanisms and drought tolerance of apple rootstock vigorous/dwarfing, the structure and water-transport ability of root system at various soil depth of two common apple rootstock species were investigated, and research was aimed to provide technical support for fruit production management. [Methods] Vigorous rootstock Malus micromalu and dwarfing rootstock M9 T337 were used as experimental materials. The growth condition, root distribution, and xylem vessel trait of both branch and root system at different soil depth were measured. Hydraulic conductivity of root and branch was determined based on the xylem vessel traits. [Results] Study results indicated that the major parameters including plant height, basal diameter, and the number and length of branch of M. micromalu were significantly higher than that of M9 T337, and both root biomass and root length density of M. micromalu with a vertical distribution of 0～60 cm were significantly greater than that of M9 T337 with vertical distribution of 0～40 cm. Vessel diameter and hydraulic conductivity of both root and branch of M. micromalu were significantly higher than that of M9 T337, while vessel density of root and branch was significantly lower in M. micromalu, and vessel diameter and hydraulic conductivity of deep roots were greater than that of shallow roots for both species. [Conclusion] M. micromalu possessed more efficient vessels, higher root length density and wider vertical root distribution, which greatly contributed to its vigorous effect and strong drought resistance. The deep root, conducting water absorbing from deep soil, may act as an important factor for improving tree's drought resistance.
引文
[1]冯轶. 苹果矮化砧木M9根系IPT5b基因表达调控及其致矮机理的研究[D]. 北京: 中国农业大学, 2017.
[2]束怀瑞. 果树栽培理论与实践[M]. 北京: 中国农业出版社,2009:430-460.
[3]De Wit I, Cook N C, Keulemans J. Characterization of tree architecture in two-year-old apple seedling populations of different progenies with a common columnar gene parent//[C].XIth Eucarpia Symposium on Fruit Breeding and Genetics, 2003, 663: 363-368.
[4]Chen B, Wang C, Tian Y, et al. Anatomical characteristics of young stems and mature leaves of dwarf pear[J]. Scientia Horticulturae, 2015, 186: 172-179.
[5]Esmaeil F, Ik-Jo C, Gerry H, et al. Effects of three rootstocks on photosynthesis, leaf mineral nutrition, and vegetation growth of “BC-2 Fuji” apple trees[J]. Journal of Plant Nutrition, 2001, 24(6): 827-834.
[6]徐继忠, 史宝胜, 马宝焜, 等. 苹果不同矮砧与其对应中间砧植株POD、IOD酶活性的研究[J]. 中国农业科学, 2002, 35(4): 415-420.
[7]廖娇, 黄春辉, 辜青青, 等. 植物矮化相关基因研究进展[J]. 生物技术通讯, 2011, 22(4): 593-597.
[8]Van Hooijdonk B, Woolley D, Warrington I, et al. Rootstocks modify scion architecture, endogenous hormones, and root growth of newly grafted ‘Royal Gala’ apple trees[J]. Journal of the American Society for Horticultural Science, 2011, 136(2): 93-102.
[9]王中英, 赵玉军. 矮化中间砧苹果树14C同化物质分配和运转的研究[J]. 山西农业科学, 1998(3): 10-14.
[10]张林森, 张海亭, 胡景江, 等. 两种苹果砧木根系水力结构及其PV曲线水分参数对干旱胁迫的响应[J]. 生态学报, 2013, 33(11): 3324-3331.
[11]Rodr?guez G J, Intrigliolo D S, Primo M E, et al. Relationships between xylem anatomy, root hydraulic conductivity, leaf/root ratio and transpiration in citrus trees on different rootstocks[J]. Physiologia Plantarum, 2010, 139(2): 159-169.
[12]Bauerle T L, Centinari M, Bauerle W L. Shifts in xylem vessel diameter and embolisms in grafted apple trees of differing rootstock growth potential in response to drought[J]. Planta, 2011, 234(5): 1045-1054.
[13]罗静. 苹果矮化中间砧矮化效应的研究[D]. 杨凌: 西北农林科技大学, 2013.
[14]Kotowska M M, Hertel D, Rajab Y A, et al. Patterns in hydraulic architecture from roots to branches in six tropical tree species from cacao agroforestry and their relation to wood density and stem growth[J]. Frontiers in Plant Science, 2015, 6: 191.
[15]Tyree M T, Ewers F W. The hydraulic architecture of trees and other woody plants[J]. New Phytologist, 2010, 119(3): 345-360.
[16]Landh?usser S M, Lieffers V J. Defoliation increases risk of carbon starvation in root systems of mature aspen[J]. Trees, 2012, 26(2): 653-661.
[17]Gieger T, Thomas F M. Effects of defoliation and drought stress on biomass partitioning and water relations of Quercus robur and Quercus petraea[J]. Basic and Applied Ecology, 2002, 3(2): 171-181.
[18]Mcelrone A J, Pockman W T, Martínez-Vilalta J, et al. Variation in xylem structure and function in stems and roots of trees to 20m depth[J]. New Phytologist, 2010, 163(3): 507-517.
[19]Vadez V. Root hydraulics: The forgotten side of roots in drought adaptation[J]. Field Crops Research, 2014, 165(3): 15-24.
[20]Sellin A, Rohej?rv A, Rahi M. Distribution of vessel size, vessel density and xylem conducting efficiency within a crown of silver birch (Betula pendula)[J]. Trees, 2008, 22(2): 205-216.
[21]王中英, 解思敏, 杨佩芳, 等. 苹果矮砧解剖构造研究[J]. 果树学报, 1988, 5(1): 6-10.
[22]李春燕, 杨廷桢, 高敬东, 等. 苹果矮化砧木致矮机理研究进展[J]. 中国农学通报, 2017, 33(28): 86-92.
[23]Olson M E, Rosell J A. Vessel diameter-stem diameter scaling across woody angiosperms and the ecological causes of xylem vessel diameter variation[J]. New Phytologist, 2013, 197(4): 1204-1213.
[2]束怀瑞. 果树栽培理论与实践[M]. 北京: 中国农业出版社,2009:430-460.
[3]De Wit I, Cook N C, Keulemans J. Characterization of tree architecture in two-year-old apple seedling populations of different progenies with a common columnar gene parent//[C].XIth Eucarpia Symposium on Fruit Breeding and Genetics, 2003, 663: 363-368.
[4]Chen B, Wang C, Tian Y, et al. Anatomical characteristics of young stems and mature leaves of dwarf pear[J]. Scientia Horticulturae, 2015, 186: 172-179.
[5]Esmaeil F, Ik-Jo C, Gerry H, et al. Effects of three rootstocks on photosynthesis, leaf mineral nutrition, and vegetation growth of “BC-2 Fuji” apple trees[J]. Journal of Plant Nutrition, 2001, 24(6): 827-834.
[6]徐继忠, 史宝胜, 马宝焜, 等. 苹果不同矮砧与其对应中间砧植株POD、IOD酶活性的研究[J]. 中国农业科学, 2002, 35(4): 415-420.
[7]廖娇, 黄春辉, 辜青青, 等. 植物矮化相关基因研究进展[J]. 生物技术通讯, 2011, 22(4): 593-597.
[8]Van Hooijdonk B, Woolley D, Warrington I, et al. Rootstocks modify scion architecture, endogenous hormones, and root growth of newly grafted ‘Royal Gala’ apple trees[J]. Journal of the American Society for Horticultural Science, 2011, 136(2): 93-102.
[9]王中英, 赵玉军. 矮化中间砧苹果树14C同化物质分配和运转的研究[J]. 山西农业科学, 1998(3): 10-14.
[10]张林森, 张海亭, 胡景江, 等. 两种苹果砧木根系水力结构及其PV曲线水分参数对干旱胁迫的响应[J]. 生态学报, 2013, 33(11): 3324-3331.
[11]Rodr?guez G J, Intrigliolo D S, Primo M E, et al. Relationships between xylem anatomy, root hydraulic conductivity, leaf/root ratio and transpiration in citrus trees on different rootstocks[J]. Physiologia Plantarum, 2010, 139(2): 159-169.
[12]Bauerle T L, Centinari M, Bauerle W L. Shifts in xylem vessel diameter and embolisms in grafted apple trees of differing rootstock growth potential in response to drought[J]. Planta, 2011, 234(5): 1045-1054.
[13]罗静. 苹果矮化中间砧矮化效应的研究[D]. 杨凌: 西北农林科技大学, 2013.
[14]Kotowska M M, Hertel D, Rajab Y A, et al. Patterns in hydraulic architecture from roots to branches in six tropical tree species from cacao agroforestry and their relation to wood density and stem growth[J]. Frontiers in Plant Science, 2015, 6: 191.
[15]Tyree M T, Ewers F W. The hydraulic architecture of trees and other woody plants[J]. New Phytologist, 2010, 119(3): 345-360.
[16]Landh?usser S M, Lieffers V J. Defoliation increases risk of carbon starvation in root systems of mature aspen[J]. Trees, 2012, 26(2): 653-661.
[17]Gieger T, Thomas F M. Effects of defoliation and drought stress on biomass partitioning and water relations of Quercus robur and Quercus petraea[J]. Basic and Applied Ecology, 2002, 3(2): 171-181.
[18]Mcelrone A J, Pockman W T, Martínez-Vilalta J, et al. Variation in xylem structure and function in stems and roots of trees to 20m depth[J]. New Phytologist, 2010, 163(3): 507-517.
[19]Vadez V. Root hydraulics: The forgotten side of roots in drought adaptation[J]. Field Crops Research, 2014, 165(3): 15-24.
[20]Sellin A, Rohej?rv A, Rahi M. Distribution of vessel size, vessel density and xylem conducting efficiency within a crown of silver birch (Betula pendula)[J]. Trees, 2008, 22(2): 205-216.
[21]王中英, 解思敏, 杨佩芳, 等. 苹果矮砧解剖构造研究[J]. 果树学报, 1988, 5(1): 6-10.
[22]李春燕, 杨廷桢, 高敬东, 等. 苹果矮化砧木致矮机理研究进展[J]. 中国农学通报, 2017, 33(28): 86-92.
[23]Olson M E, Rosell J A. Vessel diameter-stem diameter scaling across woody angiosperms and the ecological causes of xylem vessel diameter variation[J]. New Phytologist, 2013, 197(4): 1204-1213.