安徽大别山木本植物幼树小枝薄壁组织组成特征初探
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摘要
木本植物次生木质部轴向薄壁组织和射线薄壁组织在物质存储和转运等功能中起着重要的作用,剖析次生木质部薄壁组织组成有助于深入探究其功能,而小枝木质部薄壁组织组成特征还缺乏研究。该研究以天马国家级自然保护区内的18种木本植物幼树为研究对象,测算了各物种小枝木质部薄壁组织的组成含量并检测了其谱系信号,结合有关树干薄壁组织含量数据集,初步探讨木本植物小枝木质部薄壁组织含量特征。研究结果表明:(1) 18种木本植物幼树小枝的薄壁组织总含量为9.96%–18.56%,平均为14.80%;其中射线薄壁组织含量为7.74%–15.45%,高于轴向薄壁组织的含量(1.13%–7.49%);(2)小枝中薄壁组织总含量呈低于树干的趋势,其中小枝射线薄壁组织含量低于树干,而轴向薄壁组织含量高于树干,这可能是由器官差异和生活史阶段差异造成的;(3)轴向薄壁组织的含量具有显著的谱系信号,即物种亲缘关系越近其含量越相近。该研究初步验证了木本植物次生木质部薄壁组织的谱系信号,同时暗示了器官和生活史阶段差异对薄壁组织含量具有重要影响。
Aims Axial parenchyma(AP) and ray parenchyma(RP) in secondary xylem have many important ecological functions, such as storage and translocation. Quantifying the compositions of parenchyma in secondary xylem will benefit the further research about their functions. However, the understanding of parenchyma compositions in current-year branches is still lacking.Methods Eighteen woody saplings in the Tianma National Nature Reserve were selected for sampling. The proportions of cross-sectional area occupied by AP and RP were measured. The variations in parenchyma among species and their phylogenetic signals were analyzed. Compared with the related dataset, the differences between twigs and trunks were tested.Important findings(1) The proportions of total parenchyma in twigs of the 18 woody saplings were9.96%–18.56%, with the average of 14.80%; the proportions of RP(7.74%–15.45%) were higher than that of AP(1.13%–7.49%).(2) The total parenchyma(RAP) in twigs was lower than that in trunks, of which, RP in twigs was lower than that in trunks while AP showed an opposite pattern. The differences between twigs and trunks may be caused by the differences between different organs and different life history periods.(3) Significant phylogenetic signal was detected in the amount of AP in secondary xylem of twigs. This study primitively verified the phylogenetic signal of the secondary xylem parenchyma, and suggested that the difference between organs, and the difference between life history periods, had important effects on the variations of parenchyma.
Aims Axial parenchyma(AP) and ray parenchyma(RP) in secondary xylem have many important ecological functions, such as storage and translocation. Quantifying the compositions of parenchyma in secondary xylem will benefit the further research about their functions. However, the understanding of parenchyma compositions in current-year branches is still lacking.Methods Eighteen woody saplings in the Tianma National Nature Reserve were selected for sampling. The proportions of cross-sectional area occupied by AP and RP were measured. The variations in parenchyma among species and their phylogenetic signals were analyzed. Compared with the related dataset, the differences between twigs and trunks were tested.Important findings(1) The proportions of total parenchyma in twigs of the 18 woody saplings were9.96%–18.56%, with the average of 14.80%; the proportions of RP(7.74%–15.45%) were higher than that of AP(1.13%–7.49%).(2) The total parenchyma(RAP) in twigs was lower than that in trunks, of which, RP in twigs was lower than that in trunks while AP showed an opposite pattern. The differences between twigs and trunks may be caused by the differences between different organs and different life history periods.(3) Significant phylogenetic signal was detected in the amount of AP in secondary xylem of twigs. This study primitively verified the phylogenetic signal of the secondary xylem parenchyma, and suggested that the difference between organs, and the difference between life history periods, had important effects on the variations of parenchyma.
引文
Anhui Tianma National Nature Reserve Administration,Anhui Forestry Survey and Planning Institute(2014).Comprehensive Scientific Investigation Report of Tianma National Nature Reserve in Anhui.Anhui Science&Technology Publishing House,Hefei.[安徽天马国家级自然保护区管理局,安徽省林业调查规划院(2014).安徽天马国家级自然保护区综合科学考察报告.安徽科学技术出版社,合肥.]
Bel AJEV(1990).Xylem-phloem exchange via the rays:The undervalued route of transport.Journal of Experimental Botany,41,631-644.
Bhat KM,Bhat KV,Dhamodaran TK(1985).Wood and ba rk properties of branches of selected tree species growi ng in Kerala.KFRI Research Report,Kerala Forest Re search Institute.https://pdfs.semanticscholar.org/6d17/0853fce456bb543f7e060b3030b435dc9a9f.pdf.Cited:2018-12-17.
Blomberg SP,Garland T,Ives AR(2003).Testing for phylogenetic signal in comparative data:Behavioral traits are more labile.Evolution,57,717-745.
Burgert I,Eckstein D(2001).The tensile strength of isolated wood rays of beech(Fagus sylvatica L.)and its significance for the biomechanics of living trees.Trees,15,168-170.
Cao K,Rao MD,Yu JZ,Liu XJ,Mi XC,Chen JH(2013).The phylogenetic signal of functional traits and their effects on community structure in an evergreen broad-leaved forest.Biodiversity Science,21,564-571.[曹科,饶米德,余建中,刘晓娟,米湘成,陈建华(2013).古田山木本植物功能性状的系统发育信号及其对群落结构的影响.生物多样性,21,564-571.]
Du Y,Mao L,Queenborough SA,Freckleton RP,Chen B,Ma K(2015).Phylogenetic constraints and trait correlates of flowering phenology in the angiosperm flora of China.Global Ecology and Biogeography,24,928-938.
Hearn DJ(2009).Developmental patterns in anatomy are shared among separate evolutionary origins of stem succulent and storage root-bearing growth habits in Adenia(Passifloraceae).American Journal of Botany,96,1941-1956.
Hu Y,Yan L,Li H(2006).Studies on the anatomical characteristics of the stems of 14 desert plants.Journal of Arid Land Resources and Environment,20,204-210.[胡云,燕玲,李红(2006).14种荒漠植物茎的解剖结构特征分析.干旱区资源与环境,20,204-210.]
Martínez-Cabrera HI,Zheng J,Estrada-Ruiz E(2017).Wood functional disparity lags behind taxonomic diversification in angiosperms.Review of Palaeobotany&Palynology,246,251-257.
Morris H,PlavcováL,Cvecko P,Fichtler E,Gillingham MAF,Martínez-Cabrera HI,Mcglinn DJ,Wheeler E,Zheng JM,Ziemińska K,Jansen S(2016).A global analysis of parenchyma tissue fractions in secondary xylem of seed plants.New Phytologist,209,1553-1565.
Olano JM,Arzac A,García-Cervigón AI,von Arx G,Rozas V(2013).New star on the stage:Amount of ray parenchyma in tree rings shows a link to climate.New Phytologist,198,486-495.
PlavcováL,Hoch G,Morris H,Ghiasi S,Jansen S(2016).The amount of parenchyma and living fibers affects storage of nonstructural carbohydrates in young stems and roots of temperate trees.American Journal of Botany,103,1-10.
PlavcováL,Jansen S(2015).The role of xylem parenchyma in the storage and utilization of Nonstructural Carbohydrates.In:Hacke U ed.Functional and Ecological Xylem Anatomy.Springer,Cham.209-234.
Poorter L,Mcdonald I,Alarcón A,Fichtler E,Licona JC,Pe?a-Claros M,Sterck F,Villegas Z,Sass-Klaassen U(2010).The importance of wood traits and hydraulic conductance for the performance and life history strategies of42 rainforest species.New Phytologist,185,481-492.
Rungwattana K,Hietz P(2018).Radial variation of wood functional traits reflect size-related adaptations of tree mechanics and hydraulics.Functional Ecology,32,260-272.
Spicer R(2014).Symplasmic networks in secondary vascular tissues:parenchyma distribution and activity supporting long-distance transport.Journal of Experimental Botany,65,1829-1848.
Tan HY,Hao BZ,Wu JL(2000).Seasonal ultrastructural changes of secondary phloem parenchyma cells in a tropical tree,Dalbergia odorifera.Acta Botanica Yunnanica,22,461-466.[谭海燕,郝秉中,吴继林(2000).热带落叶树降香黄檀次生韧皮部薄壁组织细胞超微结构的季节变化.云南植物研究,22,461-466.]
Wei X,Liu Y,Chen HB(2008).Anatomical and functional heterogeneity among different root orders of Phellodendron amurense.Journal of Plant Ecology(Chinese Version),32,1238-1247.[卫星,刘颖,陈海波(2008).黄波罗不同根序的解剖结构及其功能异质性.植物生态学报,32,1238-1247.]
Wheeler EA,Baas P,Rodgers S(2007).Variations in dicot wood anatomy:A global analysis based on the insidewood database.Iawa Journal,28,229-258.
Yang XD,Ji PP,Re YS,Li HX(2018).Relationship between flowering phenology and phylogeny in 31 woody plants of Urumqi,Xinjiang.Acta Ecologica Sinica,38,1003-1015.
[杨晓东,姬盼盼,热依沙,李宏侠(2018).31种木本植物开花物候与系统发育的关系.生态学报,38,1003-1015.]
Zhang HY,Wang CK,Wang XC(2013).Comparison of concentrations of non-structural carbohydrates between new twigs and old branches for 12 temperate species.Acta Ecologica Sinica,33,5675-5685.[张海燕,王传宽,王兴昌(2013).温带12个树种新老树枝非结构性碳水化合物浓度比较.生态学报,33,5675-5685.]
Zhang HY,Wang CK,Wang XC(2015).Within-crown variation in concentrations of non-structural carbohydrates of five temperate tree species.Acta Ecologica Sinica,35,6496-6506.[张海燕,王传宽,王兴昌(2015).5个温带树种冠层枝叶非结构性碳水化合物浓度的空间变异.生态学报,35,6496-6506.]
Zheng GQ,Bao H,Yang J,Su XL,Hu ZH(2015).Ultrastructure of phloem and the flesh sink-cells during fruit development of Lucium barbarum.Acta Botanica BorealiOccidentalia Sinica,35,2211-2218.[郑国琦,包晗,杨涓,苏雪玲,胡正海(2015).宁夏枸杞果实韧皮部及其周围细胞超微结构研究.西北植物学报,35,2211-2218.]
Zheng GQ,Zhao M,Zhang L,Zheng GB,Hu ZH(2010).Structures and compositions of root and stem secondary xylem anatomy of Lycium barbarum with different irrigation amounts.Acta Botanica Boreali-Occidentalia Sinica,30,2170-2176.[郑国琦,赵猛,张磊,郑国保,胡正海(2010).灌水量对枸杞根茎次生木质部结构和组成的影响.西北植物学报,30,2170-2176.]
Zheng JM,Martínez-Cabrera HI(2013).Wood anatomical correlates with theoretical conductivity and wood density across China:Evolutionary evidence of the functional differentiation of axial and radial parenchyma.Annals of Botany,112,927-935.
Ziemińska K,Butler DW,Gleason SM,Wright IJ,Westoby M(2013).Fibre wall and lumen fractions drive wood density variation across 24 Australian angiosperms.AoB Plants,5,plt046.DOI:101093/aobpla/plt046.
Bel AJEV(1990).Xylem-phloem exchange via the rays:The undervalued route of transport.Journal of Experimental Botany,41,631-644.
Bhat KM,Bhat KV,Dhamodaran TK(1985).Wood and ba rk properties of branches of selected tree species growi ng in Kerala.KFRI Research Report,Kerala Forest Re search Institute.https://pdfs.semanticscholar.org/6d17/0853fce456bb543f7e060b3030b435dc9a9f.pdf.Cited:2018-12-17.
Blomberg SP,Garland T,Ives AR(2003).Testing for phylogenetic signal in comparative data:Behavioral traits are more labile.Evolution,57,717-745.
Burgert I,Eckstein D(2001).The tensile strength of isolated wood rays of beech(Fagus sylvatica L.)and its significance for the biomechanics of living trees.Trees,15,168-170.
Cao K,Rao MD,Yu JZ,Liu XJ,Mi XC,Chen JH(2013).The phylogenetic signal of functional traits and their effects on community structure in an evergreen broad-leaved forest.Biodiversity Science,21,564-571.[曹科,饶米德,余建中,刘晓娟,米湘成,陈建华(2013).古田山木本植物功能性状的系统发育信号及其对群落结构的影响.生物多样性,21,564-571.]
Du Y,Mao L,Queenborough SA,Freckleton RP,Chen B,Ma K(2015).Phylogenetic constraints and trait correlates of flowering phenology in the angiosperm flora of China.Global Ecology and Biogeography,24,928-938.
Hearn DJ(2009).Developmental patterns in anatomy are shared among separate evolutionary origins of stem succulent and storage root-bearing growth habits in Adenia(Passifloraceae).American Journal of Botany,96,1941-1956.
Hu Y,Yan L,Li H(2006).Studies on the anatomical characteristics of the stems of 14 desert plants.Journal of Arid Land Resources and Environment,20,204-210.[胡云,燕玲,李红(2006).14种荒漠植物茎的解剖结构特征分析.干旱区资源与环境,20,204-210.]
Martínez-Cabrera HI,Zheng J,Estrada-Ruiz E(2017).Wood functional disparity lags behind taxonomic diversification in angiosperms.Review of Palaeobotany&Palynology,246,251-257.
Morris H,PlavcováL,Cvecko P,Fichtler E,Gillingham MAF,Martínez-Cabrera HI,Mcglinn DJ,Wheeler E,Zheng JM,Ziemińska K,Jansen S(2016).A global analysis of parenchyma tissue fractions in secondary xylem of seed plants.New Phytologist,209,1553-1565.
Olano JM,Arzac A,García-Cervigón AI,von Arx G,Rozas V(2013).New star on the stage:Amount of ray parenchyma in tree rings shows a link to climate.New Phytologist,198,486-495.
PlavcováL,Hoch G,Morris H,Ghiasi S,Jansen S(2016).The amount of parenchyma and living fibers affects storage of nonstructural carbohydrates in young stems and roots of temperate trees.American Journal of Botany,103,1-10.
PlavcováL,Jansen S(2015).The role of xylem parenchyma in the storage and utilization of Nonstructural Carbohydrates.In:Hacke U ed.Functional and Ecological Xylem Anatomy.Springer,Cham.209-234.
Poorter L,Mcdonald I,Alarcón A,Fichtler E,Licona JC,Pe?a-Claros M,Sterck F,Villegas Z,Sass-Klaassen U(2010).The importance of wood traits and hydraulic conductance for the performance and life history strategies of42 rainforest species.New Phytologist,185,481-492.
Rungwattana K,Hietz P(2018).Radial variation of wood functional traits reflect size-related adaptations of tree mechanics and hydraulics.Functional Ecology,32,260-272.
Spicer R(2014).Symplasmic networks in secondary vascular tissues:parenchyma distribution and activity supporting long-distance transport.Journal of Experimental Botany,65,1829-1848.
Tan HY,Hao BZ,Wu JL(2000).Seasonal ultrastructural changes of secondary phloem parenchyma cells in a tropical tree,Dalbergia odorifera.Acta Botanica Yunnanica,22,461-466.[谭海燕,郝秉中,吴继林(2000).热带落叶树降香黄檀次生韧皮部薄壁组织细胞超微结构的季节变化.云南植物研究,22,461-466.]
Wei X,Liu Y,Chen HB(2008).Anatomical and functional heterogeneity among different root orders of Phellodendron amurense.Journal of Plant Ecology(Chinese Version),32,1238-1247.[卫星,刘颖,陈海波(2008).黄波罗不同根序的解剖结构及其功能异质性.植物生态学报,32,1238-1247.]
Wheeler EA,Baas P,Rodgers S(2007).Variations in dicot wood anatomy:A global analysis based on the insidewood database.Iawa Journal,28,229-258.
Yang XD,Ji PP,Re YS,Li HX(2018).Relationship between flowering phenology and phylogeny in 31 woody plants of Urumqi,Xinjiang.Acta Ecologica Sinica,38,1003-1015.
[杨晓东,姬盼盼,热依沙,李宏侠(2018).31种木本植物开花物候与系统发育的关系.生态学报,38,1003-1015.]
Zhang HY,Wang CK,Wang XC(2013).Comparison of concentrations of non-structural carbohydrates between new twigs and old branches for 12 temperate species.Acta Ecologica Sinica,33,5675-5685.[张海燕,王传宽,王兴昌(2013).温带12个树种新老树枝非结构性碳水化合物浓度比较.生态学报,33,5675-5685.]
Zhang HY,Wang CK,Wang XC(2015).Within-crown variation in concentrations of non-structural carbohydrates of five temperate tree species.Acta Ecologica Sinica,35,6496-6506.[张海燕,王传宽,王兴昌(2015).5个温带树种冠层枝叶非结构性碳水化合物浓度的空间变异.生态学报,35,6496-6506.]
Zheng GQ,Bao H,Yang J,Su XL,Hu ZH(2015).Ultrastructure of phloem and the flesh sink-cells during fruit development of Lucium barbarum.Acta Botanica BorealiOccidentalia Sinica,35,2211-2218.[郑国琦,包晗,杨涓,苏雪玲,胡正海(2015).宁夏枸杞果实韧皮部及其周围细胞超微结构研究.西北植物学报,35,2211-2218.]
Zheng GQ,Zhao M,Zhang L,Zheng GB,Hu ZH(2010).Structures and compositions of root and stem secondary xylem anatomy of Lycium barbarum with different irrigation amounts.Acta Botanica Boreali-Occidentalia Sinica,30,2170-2176.[郑国琦,赵猛,张磊,郑国保,胡正海(2010).灌水量对枸杞根茎次生木质部结构和组成的影响.西北植物学报,30,2170-2176.]
Zheng JM,Martínez-Cabrera HI(2013).Wood anatomical correlates with theoretical conductivity and wood density across China:Evolutionary evidence of the functional differentiation of axial and radial parenchyma.Annals of Botany,112,927-935.
Ziemińska K,Butler DW,Gleason SM,Wright IJ,Westoby M(2013).Fibre wall and lumen fractions drive wood density variation across 24 Australian angiosperms.AoB Plants,5,plt046.DOI:101093/aobpla/plt046.