树木木质部生长动态及其调节机制研究进展
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摘要
全球变化对树木木质部生长产生了深远影响,进而影响了森林生态系统的固碳功能以及全球生态系统能量和物质的循环过程。树木木质部生长动态主要包括形成层活动开始和结束的时间、生长季长度以及分裂速率等,其受到多种因素的共同调节,如植物激素、碳水化合物、氮素和气象因子等。通过在精细的时间尺度上对比研究树木木质部生长动态,揭示木质部形成的决定因子,可以加深对树木生长生理机制的理解,从而提高其对气候变化响应的预测精度。对近年来在木质部的形成动态及其调节机制方面取得的研究进展进行了综述,并对未来的研究方向进行了展望。
Global changes impose a profound impact on the xylem formation, which in turn affects the carbon sequestration of forest ecosystems and fundamental services of global ecosystems. The xylem formation dynamic of tree is mainly characterized by the timing of the onset and the end of cambial activity, the length of the growing season, and the growth rate, etc., which are jointly regulated by various factors, such as phytohormone, carbohydrate, nitrogen and meteorological factors. By investigating the formation dynamics of xylem over a fine time scale, the determinants of xylem formation could be revealed, the understanding of physiological mechanism of tree growth would be deepen, and the prediction accuracy of the tree growth response to climate changes would further improve. The recent research progresses in the xylem formation dynamic and its regulation mechanism were reviewed, and the prospects for the future research were provided.
Global changes impose a profound impact on the xylem formation, which in turn affects the carbon sequestration of forest ecosystems and fundamental services of global ecosystems. The xylem formation dynamic of tree is mainly characterized by the timing of the onset and the end of cambial activity, the length of the growing season, and the growth rate, etc., which are jointly regulated by various factors, such as phytohormone, carbohydrate, nitrogen and meteorological factors. By investigating the formation dynamics of xylem over a fine time scale, the determinants of xylem formation could be revealed, the understanding of physiological mechanism of tree growth would be deepen, and the prediction accuracy of the tree growth response to climate changes would further improve. The recent research progresses in the xylem formation dynamic and its regulation mechanism were reviewed, and the prospects for the future research were provided.
引文
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[13] REN P, ROSSI S, GRICAR J, et al. Is precipitation a trigger for the onset of xylogenesis in Juniperus przewalskii on the north-eastern Tibetan Plateau?[J]. Ann Bot, 2015, 115(4):629–639. doi:10.1093/aob/mcu259.
[14] HE M H, YANG B, SHISHOV V, et al. Relationships between wood formation and Cambium phenology on the Tibetan Plateau during1960–2014[J]. Forests, 2018, 9(2):86. doi:10.3390/f9020086.
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[18] ZHANG J Z, GOU X H, MANZANEDO R D, et al. Cambial phenology and xylogenesis of Juniperus przewalskii over a climatic gradient is influenced by both temperature and drought[J]. Agric For Meteor, 2018, 260–261:165–175. doi:10.1016/j.agrformet.2018.06.011.
[19] ZHANG J Z, GOU X H, PEDERSON N, et al. Cambial phenology in Juniperus przewalskii along different altitudinal gradients in a cold and arid region[J]. Tree Physiol, 2018, 38(6):840–852. doi:10.1093/tree phys/tpx160.
[20] REN P, ZIACO E, ROSSI S, et al. Growth rate rather than growing season length determines wood biomass in dry environments[J]. Agric For Meteor, 2019, 271:46–53. doi:10.1016/j.agrformet.2019.02.031.
[21] WEIJERS D, NEMHAUSER J, YANG Z B. Auxin:Small molecule, big impact[J]. J Exp Bot, 2018, 69(2):133–136. doi:10.1093/jxb/erx463.
[22] ALONI R. Role of hormones in controlling vascular differentiation and the mechanism of lateral root initiation[J]. Planta, 2013, 238(5):819–830. doi:10.1007/s00425-013-1927-8.
[23] COSGROVE D J. Loosening of plant cell walls by expansins[J].Nature, 2000, 407(6802):321–326. doi:10.1038/35030000.
[24] PERROT-RECHENMANN C. Cellular responses to auxin:Division versus expansion[J]. Cold Spring Harb Perspect Biol, 2010, 2(5):a001446. doi:10.1101/cshperspect.a001446.
[25] BHALERAO R P, FISCHER U. Auxin gradients across woodinstructive or incidental?[J]. Physiol Plant, 2014, 151(1):43–51. doi:10.1111/ppl.12134.
[26] FAJSTAVR M, PASCHOVáZ, GIAGLI K, et al. Auxin(IAA)and soluble carbohydrate seasonal dynamics monitored during xylogenesis and phloemogenesis in Scots pine[J]. iForest-Biogeosci Forestry, 2018,11:553–562. doi:10.3832/ifor2734-011.
[27] KIJIDANI Y, NAGAI T, SUWASHITA T, et al. Seasonal variations of tracheid formation and amount of auxin(IAA)and gibberellin A4(GA4)in cambial-region tissues of mature sugi(Cryptomeria japonica)cultivar grown in a Nelder plot with different tree densities[J]. J Wood Sci, 2017, 63(4):315–321. doi:10.1007/s10086-017-1626-3.
[28] LARSON P R. Wood formation and the concept of wood quality[J].Yale Univ Sch For Bull, 1969, 74:1–54.
[29] UGGLA C, MAGEL E, MORITZ T, et al. Function and dynamics of auxin and carbohydrates during earlywood/latewood transition in Scots pine[J]. Plant Physiol, 2001, 125(4):2029–2039. doi:10.1104/pp.125.4.2029.
[30] IMMANEN J, NIEMINEN K, SMOLANDER O P, et al. Cytokinin and auxin display distinct but interconnected distribution and signaling profiles to stimulate cambial activity[J]. Curr Biol, 2016, 26(15):1990–1997. doi:10.1016/j.cub.2016.05.053.
[31] ISRAELSSON M, SUNDBERG B, MORITZ T. Tissue-specific localization of gibberellins and expression of gibberellin-biosynthetic and signaling genes in wood-forming tissues in aspen[J]. Plant J, 2005, 44(3):494–504. doi:10.1111/j.1365-313X.2005.02547.x.
[32] NIEMINEN K, IMMANEN J, LAXELL M, et al. Cytokinin signaling regulates cambial development in poplar[J]. Proc Natl Acad Sci USA,2008, 105(50):20032–20037. doi:10.1073/pnas.0805617106.
[33] ALONI R. Ecophysiological implications of vascular differentiation and plant evolution[J]. Trees, 2015, 29(1):1–16. doi:10.1007/s00468-014-1070-6.
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