Non-structural carbohydrate storage strategy in trunk tissues of different wood porosity species in warm temperate zone

doi:10.13287/j.1001-9332.202508.009

Abstract

Abstract: Wood porosity types (non-porous, diffuse-porous, and ring-porous) reflect evolutionary gradients cha-racteristics of xylem anatomy of temperate tree species. The mechanisms linking porosity type to non-structural carbohydrate (NSC) storage strategy in stem tissues remain unclear. We conducted an experiment with 77 warm-tempe-rate tree species in the Baotianman National Nature Reserve, Henan Province. Among the examined species, there were 3 non-porous wood species, 45 diffuse-porous wood species, and 29 ring-porous wood species (including semi-ring-porous wood). We measured soluble sugars, starch, total NSC concentrations and sugars/starch ratio in bark, sapwood, and heartwood at breast height to explore the influence of wood porosity on NSC storage strategy across stem tissues (bark, sapwood and heartwood). The results showed that the types of trunk tissue and wood porosity had significant effects on the concentration of NSC and its components (soluble sugars, and starch). As for the trunk cross-section, NSC and its components exhibited an inward decline trend, with total NSC concentration in bark (6.4%) being notably higher than that in sapwood (3.2%) and heartwood (2.5%). The total NSC in various trunk tissues was dominant in the form of soluble sugars. Across the three wood porosity types, the concentrations of soluble sugars, total NSC, and sugars/starch ratio in tree bark displayed a counter-evolutionary sequence pattern (non-porous > diffuse-porous > ring-porous), while that in sapwood and heartwood increased along the evolutionary gradient. The concentrations of NSC and its components in sapwood and that in heartwood were significantly correlated for both diffuse-porous and ring-porous tree species. Soluble sugars and starch exhibited significant positive correlations in bark of non-porous and diffuse-porous trees and the three trunk tissues of ring-porous tree species. Along the evolutionary gradient of wood porosity, warm-temperate trees tended to optimize resource allocation by reducing the NSC concentration from bark to sapwood and enhancing the functional differentiation between sapwood and heartwood, reflecting the coordination between xylem anatomical structure and storage function.

Key words: bark, sapwood, heartwood, wood porosity, non-structural carbohydrate

DUAN Yichen, ZHAO Huabin, HAN Yongjie, LIU Xiaojing, ZHANG Yi, YAN Hailei, CHEN Zhicheng, WANG Xingchang. Non-structural carbohydrate storage strategy in trunk tissues of different wood porosity species in warm temperate zone[J]. Chinese Journal of Applied Ecology, 2025, 36(8): 2335-2343.

References

[1] Hartmann H, Trumbore S. Understanding the roles of nonstructural carbohydrates in forest trees-from what we can measure to what we want to know. New Phytologist, 2016, 211: 386-403
[2] Landháusser S, Adams H. Getting to the root of carbon reserve dynamics in woody plants: Progress, challenges, and goals. Tree Physiology, 2024, 44: 1-10
[3] Blumstein M, Sala A, Weston DJ, et al. Plant carbohydrate storage: Intra- and inter-specific trade-offs reveal a major life history trait. New Phytologist, 2022, 235: 2211-2222
[4] Blumstein MJ, Furze ME. Interannual dynamics of stemwood nonstructural carbohydrates in temperate forest trees surrounding drought. Journal of Forestry Research, 2023, 34: 77-86
[5] Wang ZG, Wang CK. Dynamics of nonstructural carbohydrates during drought and subsequent recovery: A global meta-analysis. Agricultural and Forest Meteoro-logy, 2025, 363: 110429
[6] Zhang HY, Wang CK, Wang XC. Spatial variations in non-structural carbohydrates in stems of twelve tempe-rate tree species. Trees-Structure and Function, 2014, 28: 77-89
[7] Hoch G, Richter A, Körner C. Non-structural carbon compounds in temperate forest trees. Plant, Cell & Environment, 2003, 26: 1067-1081
[8] 董涵君, 王兴昌, 苑丹阳, 等. 温带不同材性树种树干非结构性碳水化合物的径向分配差异. 植物生态学报, 2022, 46(6): 722-734
[9] Taylor A, Gartner B, Morrell J. Heartwood formation and natural durability: A review. Wood and Fiber Science, 2002, 34: 587-611
[10] Würth MKR, Peláez-Riedl S, Wright SJ, et al. Non-structural carbohydrate pools in a tropical forest. Oecologia, 2005, 143: 11-24
[11] Spicer R. 22-Senescence in Secondary Xylem: Heartwood Formation as an Active Developmental Program. Burlington, USA: Academic Press, 2005: 457-475
[12] Tyree MT, Zimmermann MH. Xylem Structure and the Ascent of Sap. New York: Springer-Verlag, 2002
[13] Chen ZC, Zhu SD, Zhang YT, et al. Tradeoff between storage capacity and embolism resistance in the xylem of temperate broadleaf tree species. Tree Physiology, 2020, 40: 1029-1042
[14] Morris H, Plavcová L, Cvecko P, et al. A global analysis of parenchyma tissue fractions in secondary xylem of seed plants. New Phytologist, 2016, 209: 1553-1565
[15] Plavcová L, Jansen S. The role of xylem parenchyma in the storage and utilization of nonstructural carbohydrates. Cham, Switzerland: Springer, 2015: 209-234
[16] 于丽敏, 王传宽, 王兴昌. 三种温带树种非结构性碳水化合物的分配. 植物生态学报, 2011, 35(12): 1245-1255
[17] Rosell J, Gleason S, Méndez-Alonzo R, et al. Bark functional ecology: Evidence for tradeoffs, functional coordination, and environment producing bark diversity. New Phytologist, 2013, 201: 486-497
[18] Rosell JA. Bark in woody plants: Understanding the diversity of a multifunctional structure. Integrative and Comparative Biology, 2019, 59: 535-547
[19] Zhang GQ, Mao Z, Fortunel C, et al. Parenchyma fractions drive the storage capacity of nonstructural carbohydrates across a broad range of tree species. American Journal of Botany, 2022, 109: 535-549
[20] 代永欣, 王林, 王延书, 等. 摘叶造成的碳限制对刺槐碳素分配和水力学特性的影响. 植物科学学报, 2017, 35(5): 750-758
[21] Herrera-Ramírez D, Hartmann H, Römermann C, et al. Anatomical distribution of starch in the stemwood influences carbon dynamics and suggests storage-growth trade-offs in some tropical trees. Journal of Ecology, 2023, 111: 2532-2548
[22] 王凯, 石亮. 科尔沁沙地3种主要造林树种树干非结构性碳水化合物的分布规律. 生态学杂志, 2023, 42(8): 1841-1850
[23] Wang XC, Wang CK, Zhang QZ, et al. Heartwood and sapwood allometry of seven Chinese temperate tree species. Annals of Forest Science, 2010, 67: 410
[24] Gartner BL, Baker DC, Spicer R. Distribution and vita-lity of xylem rays in relation to tree leaf area in Douglas-fir. IAWA Journal, 2000, 21: 389-401
[25] Gleason SM, Westoby M, Jansen S, et al. Weak tradeoff between xylem safety and xylem-specific hydraulic efficiency across the world’s woody plant species. New Phytologist, 2016, 209: 123-136
[26] Zhang SB, Wen GJ, Qu YY, et al. Trade-offs between xylem hydraulic efficiency and mechanical strength in Chinese evergreen and deciduous savanna species. Tree Physiology, 2022, 42: 1337-1349
[27] Bush SE, Pataki DE, Hultine KR, et al. Wood anatomy constrains stomatal responses to atmospheric vapor pressure deficit in irrigated, urban trees. Oecologia, 2008, 156: 13-20
[28] Ziemińska K, Rosa E, Gleason SM, et al. Wood day capacitance is related to water content, wood density, and anatomy across 30 temperate tree species. Plant, Cell & Environment, 2020, 43: 3048-3067
[29] 殷笑寒, 郝广友. 长白山阔叶树种木质部环孔和散孔结构特征的分化导致其水力学性状的显著差异. 应用生态学报, 2018, 29(2): 352-360
[30] Hacke UG, Sperry JS, Wheeler JK, et al. Scaling of angiosperm xylem structure with safety and efficiency. Tree Physiology, 2006, 26: 689-701
[31] Nardini A, Pedà G, Rocca N. Trade-offs between leaf hydraulic capacity and drought vulnerability: Morpho-anatomical bases, carbon costs and ecological consequences. New Phytologist, 2012, 196: 788-798
[32] Bittencourt PRL, Pereira L, Oliveira RS. On xylem hydraulic efficiencies, wood space-use and the safety-efficiency tradeoff. New Phytologist, 2016, 211: 1152-1155
[33] Godfrey JM, Riggio J, Orozco J, et al. Ray fractions and carbohydrate dynamics of tree species along a 2750 m elevation gradient indicate climate response, not spatial storage limitation. New Phytologist, 2020, 225: 2314-2330
[34] Peltier DMP, Carbone MS, Ogle K, et al. Decades-old carbon reserves are widespread among tree species, constrained only by sapwood longevity. New Phytologist, 2025, 245: 1468-1480
[35] 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