长白山阔叶红松林3个主要树种的非结构性碳储存特征
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摘要
非结构性碳水化合物(NSC,包括可溶性糖和淀粉)作为树木生命代谢的关键物质之一,在树木器官中的储存特征受到了广泛关注,但NSC在器官内部具有不同功能的组织间(韧皮部与木质部)的分配和权衡还不清楚.本研究以长白山阔叶红松林的3个优势树种——红松、水曲柳和紫椴为研究对象,对比分析NSC在根、树干韧皮与木质部中的浓度和分配特征.结果表明:树木韧皮部和木质部间的NSC浓度差异显著,总体分配趋势为韧皮部以可溶性糖为主,而木质部以淀粉为主.树干外侧(以年轮划分,0～20年)、中段(20～40年)和内侧(>40年)的NSC浓度在不同树种间差异显著,而根中的差异不显著.红松和水曲柳树干韧皮部可溶性总糖浓度显著高于紫椴,在木质部中差异不显著.本研究结果表明,树体内部NSC在韧皮部和木质部上的分配存在明显分异,这与树种的演替阶段及组织的功能进化有关.研究结果对于深入理解温带树木的碳储存特征和分配机理具有参考意义.
Non-structural carbohydrates(NSC, including soluble sugars and starch) are key meta-bolites in tree, the storage characteristics of which in tree organs have received extensive attention. It is still unclear how NSC are allocated in the tissues(phloem and xylem) that have different function. In this study, we analyzed the concentration and allocation of NSC in the roots, and in phloem and xylem of the trunk in three dominant species of broadleaved Korean pine forest in the Changbai Mountain, Pinus koraiensis, Fraxinus mandschurica, and Tilia amurensis. The results showed that there was a significant difference in the concentration of NSC between the phloem and xylem. The soluble sugar dominated in the phloem, while starch dominated in the xylem. The concentration of NSC in trunk outside(divided by annual rings, 0-20 years), intermediate(20-40 years) and inner(>40 years) of different tree species was significantly different, but with no difference in the roots. The total soluble sugar concentration in the phloem of P. koraiensis and F. mandschurica was significantly higher than that of T. amurensis, while the difference in xylem was not significant. The results indicated that NSC allocation in the phloem and xylem of the tree had clear tissue differentiation, which might be related to the succession stage of the tree species or the functional evolution of the tissue. These findings would improve our understanding of the carbon storage characteristics and allocation mechanism in temperate trees.
Non-structural carbohydrates(NSC, including soluble sugars and starch) are key meta-bolites in tree, the storage characteristics of which in tree organs have received extensive attention. It is still unclear how NSC are allocated in the tissues(phloem and xylem) that have different function. In this study, we analyzed the concentration and allocation of NSC in the roots, and in phloem and xylem of the trunk in three dominant species of broadleaved Korean pine forest in the Changbai Mountain, Pinus koraiensis, Fraxinus mandschurica, and Tilia amurensis. The results showed that there was a significant difference in the concentration of NSC between the phloem and xylem. The soluble sugar dominated in the phloem, while starch dominated in the xylem. The concentration of NSC in trunk outside(divided by annual rings, 0-20 years), intermediate(20-40 years) and inner(>40 years) of different tree species was significantly different, but with no difference in the roots. The total soluble sugar concentration in the phloem of P. koraiensis and F. mandschurica was significantly higher than that of T. amurensis, while the difference in xylem was not significant. The results indicated that NSC allocation in the phloem and xylem of the tree had clear tissue differentiation, which might be related to the succession stage of the tree species or the functional evolution of the tissue. These findings would improve our understanding of the carbon storage characteristics and allocation mechanism in temperate trees.
引文
[1] Pan Q-M (潘庆民),Han X-G (韩兴国),Bai Y-F (白永飞),et al.Advances in physiology and ecology studies on stored non-structure carbohydrates in plants.Chinese Bulletin of Botany (植物学通报),2002,19(1):30-38 (in Chinese)
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[3] K?rner C.Carbon limitation in trees.Journal of Ecology,2003,91:4-17
[4] Barbaroux C,Bréda N.Contrasting distribution and seasonal dynamics of carbohydrate reserves in stem wood of adult ring-porous sessile oak and diffuse-porous beech trees.Tree Physiology,2002,22:1201-1210
[5] Gough CM,Flower CE,Vogel CS,et al.Whole-ecosystem labile carbon production in a north temperate deciduous forest.Agricultural and Forest Meteorology,2009,149:1531-1540
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[7] Richardson AD,Carbone MS,Keenan TF,et al.Seasonal dynamics and age of stemwood nonstructural carbohydrates in temperate forest trees.New Phytologist,2013,197:850-861
[8] Rocha AV,Goulden ML.Why is marsh productivity so high?New insights from eddy covariance and biomass measurements in a Typha marsh.Agricultural and Forest Meteorology,2009,149:159-168
[9] Sala A,Woodruff DR,Meinzer FC.Carbon dynamics in trees:Feast or famine?Tree Physiology,2012,32:764-775
[10] Palacio S,Hoch G,Sala A,et al.Does carbon storage limit tree growth?New Phytologist,2014,201:1096-1100
[11] Barbaroux C,Bréda N,Dufrêne E.Distribution of above-ground and below-ground carbohydrate reserves in adult trees of two contrasting broad-leaved species (Quercus petraea and Fagus sylvatica).New Phytologist,2003,157:605-615
[12] Hoch G,Richter A,K?rner C.Non-structural carbon compounds in temperate forest trees.Plant,Cell & Environment,2003,26:1067-1081
[13] Dietze MC,Sala A,Carbone MS,et al.Nonstructural carbon in woody plants.Annual Review of Plant Biology,2014,65:667-687
[14] R Core Team.R:A Language and Environment for Statistical Computing.Vienna,Austria:R Foundation for Statistical Computing,2018
[15] Zhang H-Y (张海燕),Wang C-K (王传宽),Wang X-C (王兴昌),et al.Spatial variation of non-structural carbohydrates in Betula platyphylla and Tilia amurensis stems.Chinese Journal of Applied Ecology (应用生态学报),2013,24(11):3050-3056 (in Chinese)
[16] Myers JA,Kitajima K.Carbohydrate storage enhances seedling shade and stress tolerance in a neotropical forest.Journal of Ecology,2007,95:383-395
[17] Kobe RK.Carbohydrate allocation to storage as a basis of interspecific variation in sapling survivorship and growth.Oikos,1997,80:226-233
[18] Klein T,Vitasse Y,Hoch G.Coordination between growth,phenology and carbon storage in three coexisting deciduous tree species in a temperate forest.Tree Physiology,2016,36:847-855
[19] Regier N,Streb S,Zeeman SC,et al.Seasonal changes in starch and sugar content of poplar (Populus deltoides × nigra cv.Dorskamp) and the impact of stem girdling on carbohydrate allocation to roots.Tree Physiology,2010,30:979-987
[20] Bazot S,Barthes L,Blanot D,et al.Distribution of non-structural nitrogen and carbohydrate compounds in mature oak trees in a temperate forest at four key phenological stages.Trees Structure and Function,2013,27:1023-1034
[21] Gruber A,Pirkebner D,Oberhuber W.Seasonal dynamics of mobile carbohydrate pools in phloem and xylem of two alpine timberline conifers.Tree Physiology,2013,33:1076-1083
[22] Scartazza A,Moscatello S,Matteucci G,et al.Seasonal and inter-annual dynamics of growth,non-structural carbohydrates and C stable isotopes in a Mediterranean beech forest.Tree Physiology,2013,33:730-742
[23] Li MH,Xiao WF,Wang SG,et al.Mobile carbohydrates in Himalayan treeline trees.Ⅰ.Evidence for carbon gain limitation but not for growth limitation.Tree Physiology,2008,28:1287-1296
[24] Ryan MG,Asao S.Phloem transport in trees.Tree Physiology,2014,34:1-4
[25] McDowell NG,Sevanto S.The mechanisms of carbon starvation:How,when,or does it even occur at all?New Phytologist,2010,186,264-266
[26] Sala A,Piper F,Hoch G.Physiological mechanisms of drought induced tree mortality are far from being resolved.New Phytologist,2010,186,274-281
[27] Yu L-M (于丽敏),Wang C-K (王传宽),Wang X-C (王兴昌).Allocation of nonstructural carbohydrates for three temperate tree species in Northeast China.Chinese Journal of Plant Ecology (植物生态学报),2011,35(12):1245-1255 (in Chinese)
[28] Newell EA,Mulkey SS,Wright SJ.Seasonal patterns of carbohydrate storage in four tropical tree species.Oecologia,2002,131:333-342
[29] Essiamah S,Eschrich W.Changes of starch content in the storage tissues of deciduous trees during winter and spring.IAWA Journal,1985,6:97-106
[30] Gérard B,Bréda N.Radial distribution of carbohydrate reserves in the trunk of declining European beech trees (Fagus sylvatica L.).Annals of Forest Science,2014,71:675-682
[2] Zheng Y-P (郑云普),Wang H-X (王贺新),Lou X (娄鑫),et al.Changes of non-structural carbohydrates and its impact factors in trees:A review.Chinese Journal of Applied Ecology (应用生态学报),2014,25(4):1188-1196 (in Chinese)
[3] K?rner C.Carbon limitation in trees.Journal of Ecology,2003,91:4-17
[4] Barbaroux C,Bréda N.Contrasting distribution and seasonal dynamics of carbohydrate reserves in stem wood of adult ring-porous sessile oak and diffuse-porous beech trees.Tree Physiology,2002,22:1201-1210
[5] Gough CM,Flower CE,Vogel CS,et al.Whole-ecosystem labile carbon production in a north temperate deciduous forest.Agricultural and Forest Meteorology,2009,149:1531-1540
[6] Würth MKR,Pelaez-Riedl S,Wright SJ,et al.Non-structural carbohydrate pools in a tropical forest.Oecologia,2005,143:11-24
[7] Richardson AD,Carbone MS,Keenan TF,et al.Seasonal dynamics and age of stemwood nonstructural carbohydrates in temperate forest trees.New Phytologist,2013,197:850-861
[8] Rocha AV,Goulden ML.Why is marsh productivity so high?New insights from eddy covariance and biomass measurements in a Typha marsh.Agricultural and Forest Meteorology,2009,149:159-168
[9] Sala A,Woodruff DR,Meinzer FC.Carbon dynamics in trees:Feast or famine?Tree Physiology,2012,32:764-775
[10] Palacio S,Hoch G,Sala A,et al.Does carbon storage limit tree growth?New Phytologist,2014,201:1096-1100
[11] Barbaroux C,Bréda N,Dufrêne E.Distribution of above-ground and below-ground carbohydrate reserves in adult trees of two contrasting broad-leaved species (Quercus petraea and Fagus sylvatica).New Phytologist,2003,157:605-615
[12] Hoch G,Richter A,K?rner C.Non-structural carbon compounds in temperate forest trees.Plant,Cell & Environment,2003,26:1067-1081
[13] Dietze MC,Sala A,Carbone MS,et al.Nonstructural carbon in woody plants.Annual Review of Plant Biology,2014,65:667-687
[14] R Core Team.R:A Language and Environment for Statistical Computing.Vienna,Austria:R Foundation for Statistical Computing,2018
[15] Zhang H-Y (张海燕),Wang C-K (王传宽),Wang X-C (王兴昌),et al.Spatial variation of non-structural carbohydrates in Betula platyphylla and Tilia amurensis stems.Chinese Journal of Applied Ecology (应用生态学报),2013,24(11):3050-3056 (in Chinese)
[16] Myers JA,Kitajima K.Carbohydrate storage enhances seedling shade and stress tolerance in a neotropical forest.Journal of Ecology,2007,95:383-395
[17] Kobe RK.Carbohydrate allocation to storage as a basis of interspecific variation in sapling survivorship and growth.Oikos,1997,80:226-233
[18] Klein T,Vitasse Y,Hoch G.Coordination between growth,phenology and carbon storage in three coexisting deciduous tree species in a temperate forest.Tree Physiology,2016,36:847-855
[19] Regier N,Streb S,Zeeman SC,et al.Seasonal changes in starch and sugar content of poplar (Populus deltoides × nigra cv.Dorskamp) and the impact of stem girdling on carbohydrate allocation to roots.Tree Physiology,2010,30:979-987
[20] Bazot S,Barthes L,Blanot D,et al.Distribution of non-structural nitrogen and carbohydrate compounds in mature oak trees in a temperate forest at four key phenological stages.Trees Structure and Function,2013,27:1023-1034
[21] Gruber A,Pirkebner D,Oberhuber W.Seasonal dynamics of mobile carbohydrate pools in phloem and xylem of two alpine timberline conifers.Tree Physiology,2013,33:1076-1083
[22] Scartazza A,Moscatello S,Matteucci G,et al.Seasonal and inter-annual dynamics of growth,non-structural carbohydrates and C stable isotopes in a Mediterranean beech forest.Tree Physiology,2013,33:730-742
[23] Li MH,Xiao WF,Wang SG,et al.Mobile carbohydrates in Himalayan treeline trees.Ⅰ.Evidence for carbon gain limitation but not for growth limitation.Tree Physiology,2008,28:1287-1296
[24] Ryan MG,Asao S.Phloem transport in trees.Tree Physiology,2014,34:1-4
[25] McDowell NG,Sevanto S.The mechanisms of carbon starvation:How,when,or does it even occur at all?New Phytologist,2010,186,264-266
[26] Sala A,Piper F,Hoch G.Physiological mechanisms of drought induced tree mortality are far from being resolved.New Phytologist,2010,186,274-281
[27] Yu L-M (于丽敏),Wang C-K (王传宽),Wang X-C (王兴昌).Allocation of nonstructural carbohydrates for three temperate tree species in Northeast China.Chinese Journal of Plant Ecology (植物生态学报),2011,35(12):1245-1255 (in Chinese)
[28] Newell EA,Mulkey SS,Wright SJ.Seasonal patterns of carbohydrate storage in four tropical tree species.Oecologia,2002,131:333-342
[29] Essiamah S,Eschrich W.Changes of starch content in the storage tissues of deciduous trees during winter and spring.IAWA Journal,1985,6:97-106
[30] Gérard B,Bréda N.Radial distribution of carbohydrate reserves in the trunk of declining European beech trees (Fagus sylvatica L.).Annals of Forest Science,2014,71:675-682
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