Effects of canopy density on the growth of broadleaved tree species under artificial regeneration of Pinus sylvestris forest
-
摘要:
目的 研究在樟子松林不同郁闭度条件下,水曲柳、胡桃楸和紫椴人工种植幼苗的生长差异,为樟子松林不同郁闭度下适合更新的阔叶树种选择提供理论依据以及技术支撑。 方法 以水曲柳、胡桃楸和紫椴2年生裸根苗为试验材料;在樟子松林下选择4种光环境(全光、郁闭度0.4、郁闭度0.6、郁闭度0.8),1个生长季后观测人工更新的3个树种苗高增长量、地径增长量、光合参数、光合色素含量以及叶片养分含量变化。 结果 胡桃楸在郁闭度0.6 ~ 0.8下的苗高生长量和地径生长量显著高于全光和郁闭度0.4处理下的苗木,在郁闭度0.8处理下的苗高增长量和地径增长量最大,水曲柳和紫椴在郁闭度0.4 ~ 0.6处理下的苗高增长量和地径增长量显著高于全光和郁闭度0.8处理下的苗木,整体上胡桃楸的苗高增长量和地径增长量在4个处理下都优于水曲柳和紫椴。随着郁闭度的增大,3个树种的叶绿素含量表现出显著的上升趋势,光合速率(Pn)的变化趋势和叶绿素总量含量(Ct)变化趋势保持一致。胡桃楸的气孔导度(Gs)以及胞间CO2浓度(Ci)随郁闭度的增大呈上升趋势;水曲柳和紫椴的Gs呈上升趋势,Ci整体呈先升高后降低的趋势;蒸腾速率(Tr)随着郁闭度的增大都呈下降趋势。3个树种的叶片全N含量随着郁闭度的增加呈上升趋势;胡桃楸的叶片全C含量各处理间差异不显著,水曲柳和紫椴的叶片全C含量随郁闭度的增大呈下降趋势,其中水曲柳苗木叶片的全C含量在全光、郁闭度0.4和郁闭度0.6处理下差异不显著,但都高于郁闭度0.8处理下的叶片的全C含量;紫椴的叶片P含量呈先上升后下降的趋势,胡桃楸则呈逐渐下降的趋势,而水曲柳的叶片全P含量在各处理间的差异不显著;3个树种的叶片全K含量整体呈现先下降后上升的趋势。在樟子松不同郁闭度下胡桃楸的生长情况均优于水曲柳和紫椴;其中紫椴在各个郁闭度下光合速率变化不显著,光照强度变化对其影响不大。 结论 本研究针对樟子松林不同郁闭度下3个阔叶树种幼苗栽植1年后生长情况进行了对比分析,研究发现水曲柳、紫椴和胡桃楸3个树种幼苗在栽植当年均可在樟子松林土壤环境下生长,且均具有一定的耐阴性。水曲柳和紫椴在郁闭度0.4 ~ 0.6的生长优于全光和郁闭度0.8,胡桃楸在郁闭度0.8下生长最好,且不同郁闭度下胡桃楸的生长状况均优于水曲柳和紫椴。随着胡桃楸的生长,其与周围植被的关系,以及对光照资源的需求也可能会发生变化,后续仍需跟踪调查,为樟子松人工林更新提供扎实的理论基础。 Abstract:Objective The growth differences of Fraxinus mandshurica, Juglans mandshurica and Tilia amurensis artificially planted seedlings under different canopy densities of Pinus sylvestris var. mongolica forest were investigated in this study to provide theoretical basis and technical support for the selection of broadleaved trees suitable for regeneration under different canopy densities of P. sylvestris var. mongolica forest. Method A artificial regeneration experiment was conducted with two-year-old bare root seedlings of F. mandshurica, J. mandshurica, T. amurensis under the P. sylvestris var. mongolica forest. Furthermore, four light environments of P. sylvestris var. mongolica forest were applied, i.e. full light, canopy density of 0.4, 0.6 and 0.8. At the end of the growing season, the changes of seedling height, base diameter, photosynthetic parameters, photosynthetic pigment content and leaf nutrient content of the three seedlings were measured. Result By the end of the growing season, the height growth and ground diameter growth of J. mandshurica seedlings under canopy density of 0.6−0.8 were significantly higher than those under full light and canopy density of 0.4, and those were the largest under canopy density of 0.8. The height growth and ground diameter growth of F. mandshurica and T. amurensis seedlings under canopy density of 0.4−0.6 were significantly higher than those under full light and canopy density of 0.8. The height growth and ground diameter growth of J. mandshurica seedlings were better than F. mandshurica and T. amurensis seedlings under the four treatments. With the increase of canopy density, chlorophyll content of the three tree species showed a significant upward trend, and the trend of photosynthetic rate (Pn) was consistent with the trend of total chlorophyll content (Ct). The stomatal conductance (Gs) and intercellular CO2 concentration (Ci) of J. mandshurica increased with the increase of canopy density; the Gs of F. mandshurica and T. amurensis showed an upward trend, and the Ci showed an overall trend of first increasing and then decreasing; transpiration rate (Tr) showed a downward trend with the increase of canopy density. The leaf N content of the three tree species increased with the increase of canopy density; there was no significant difference in the total C content of J. mandshurica leaves among the treatments. The total C content of F. mandshurica and T. amurensis leaves showed a downward trend with the increase of canopy density, among which, the total C content of F. mandshurica seedling leaves under full light, canopy density of 0.4 and 0.6 treatments had no significant difference, but they were all higher than that under canopy density of 0.8 treatment; total P content in the leaves of T. amurensis increased first and then decreased, J. mandshurica decreased gradually, while total P content in the leaves of F. mandshurica was not significantly different among the treatments, and total K content decreased first and then increased as a whole. The growth of J. mandshurica was better than F. mandshurica and T. amurensis under different canopy densities of P. sylvestris; the growth of J. mandshurica was better than F. mandshurica and T. amurensis under different canopy densities of P. sylvestris; among them, the photosynthetic rate of T. amurensis under each canopy density did not change significantly, the change of light intensity had little effect on it. Conclusion This study conducted a comparative analysis of the growth of three broadleaved tree species under different canopy densities of P. sylvestris var. mongolica forest in one year of pianting. It was found that F. mandshurica, T. amurensis and J. mandshurica can grow in the soil environment of P. sylvestris var. mongolica forest in the year of planting, and all of them have certain negative tolerance. F. mandshurica and T. amurensis grow better in canopy density of 0.4−0.6 than in full light and canopy density of 0.8, J. mandshurica grows best in canopy density of 0.8, and the growth status of J. mandshurica under different canopy densities is better than F. mandshurica and T. amurensis. With the growth of seedlings, the relationship between J. mandshurica and its surrounding vegetation and its demand for light resources may change. Thus, to provide a solid theoretical basis for the regeneration of P. sylvestris var. mongolica plantation, followed investigations are still needed. -
图 5 不同郁闭度下各树种生长指标的Spearman相关性
红色表示正相关关系,蓝色表示负相关关系,圆圈面积越大相关性越强,右侧数值表示相关性系数值;*表示显著相关(P < 0.05);Sd.不同郁闭度;Sh.苗高增长量;Gd.地径增长量;Ct.叶绿素总量;TC.全碳;TN.全氮;TK.全钾;TP.全磷。Red indicates positive correlation, blue indicates negative correlation, and the larger the circle area is, the stronger the correlation is, the value on the right indicates correlation coefficient value; * indicates significant correlation (P < 0.05); Sd, different canopy densities; Sh, seedling height growth; Gd, ground diameter growth; Ct, total chlorophyll content; TC, total carbon; TN, total nitrogen; TK, total potassium; TP, total phosphorus.
Figure 5. Spearman correlation of growth indexes of various tree species under different canopy densities
-
[1] 崔海鸥, 刘珉. 我国第九次森林资源清查中的资源动态研究[J]. 西部林业科学, 2020, 49(5): 90−95. doi: 10.16473/j.cnki.xblykx1972.2020.05.014Cui H O, Liu M. Analysis on the results of the 9th National Forest Inventory[J]. Journal of West China Forestry Science, 2020, 49(5): 90−95. doi: 10.16473/j.cnki.xblykx1972.2020.05.014 [2] Nguyen H, Firn J, Lamb D, et al. Wood density: a tool to find complementary species for the design of mixed species plantations[J]. Forest Ecology and Management, 2014, 334: 106−113. doi: 10.1016/j.foreco.2014.08.022 [3] Li Y Z, Chen X S, Xie Y H, et al. Effects of young poplar plantations on understory plant diversity in the Dongting Lake Wetlands, China[J]. Scientific Reports, 2014, 4(1): 6339. doi: 10.1038/srep06339 [4] Chaudhary A, Burivalova Z, Koh L P, et al. Impact of forest management on species richness: global meta-analysis and economic trade-offs[J]. Scientific Reports, 2016, 6: 23954. doi: 10.1038/srep23954 [5] 赵庆喜. 红松皆伐迹地更新红松的效果分析[J]. 防护林科技, 2014(4): 28−29, 32. doi: 10.13601/j.issn.1005-5215.2014.04.012Zhao Q X. Effects of regeneration of Pinus koraiensis in clear-cutting site[J]. Protection Forest Science and Technology, 2014(4): 28−29, 32. doi: 10.13601/j.issn.1005-5215.2014.04.012 [6] 林文树, 穆丹, 王丽平, 等. 针阔混交林不同演替阶段表层土壤理化性质与优势林木生长的相关性[J]. 林业科学, 2016, 52(5): 17−25.Lin W S, Mu D, Wang L P, et al. Correlation between the growth of dominant trees and surface soil physiochemical of conifer and broad-leaved mixed forest at different succession stages[J]. Scientia Silvae Sinicae, 2016, 52(5): 17−25. [7] Zhang X Z, Guan D X, Li W B, et al. The effects of forest thinning on soil carbon stocks and dynamics: a meta-analysis[J]. Forest Ecology and Management, 2018, 429: 36−43. doi: 10.1016/j.foreco.2018.06.027 [8] 周磊, 王树力. 树种混交对红皮云杉人工林土壤养分的影响[J]. 东北林业大学学报, 2019, 47(2): 37−41. doi: 10.3969/j.issn.1000-5382.2019.02.009Zhou L, Wang S L. Effects of mixed tree species on soil nutrients in Picea koraiensis plantations[J]. Journal of Northeast Forestry University, 2019, 47(2): 37−41. doi: 10.3969/j.issn.1000-5382.2019.02.009 [9] Manson D G, Schmidt S, Bristow M, et al. Species-site matching in mixed species plantations of native trees in tropical Australia[J]. Agroforestry Systems, 2013, 87(1): 233−250. doi: 10.1007/s10457-012-9538-0 [10] 张东来, 毛子军, 朱胜英, 等. 黑龙江省帽儿山林区6种主要林分类型凋落物研究[J]. 植物研究, 2008, 28(1): 104−108. doi: 10.7525/j.issn.1673-5102.2008.01.022Zhang D L, Mao Z J, Zhu S Y, et al. Litter falls of 6 major stands in Maoershan Mountain of Heilongjiang Province[J]. Bulletin of Botanical Research, 2008, 28(1): 104−108. doi: 10.7525/j.issn.1673-5102.2008.01.022 [11] 韦庆钰, 黄海龙, 吴纯泽, 等. 3种倍性青杨扦插苗对覆膜滴肥的生长响应[J]. 南京林业大学学报(自然科学版), 2021, 45(5): 93−101.Wei Q Y, Huang H L, Wu C Z, et al. Response of Populus cathayana cutting seedlings of three ploidy types to fertilizer[J]. Journal of Nanjing Forestry University (Natural Sciences Edition), 2021, 45(5): 93−101. [12] 马常钦, 李敏, 卫星. 菌渣还田比例及粒级对土壤特性及水曲柳苗木生长的影响[J]. 西南林业大学学报(自然科学), 2022, 42(6): 80−87.Ma C Q, Li M, Wei X. Effect of ratio and size added spent mushroom substrate on soil properties and growth of Fraxinus mandshurica seedlings[J]. Journal of Southwest Forestry University (Natural Sciences), 2022, 42(6): 80−87 [13] 陈璐, 王庆贵, 闫国永, 等. 不同施氮水平对兴安落叶松化学计量特征的影响[J]. 森林工程, 2019, 35(2): 1−10. doi: 10.3969/j.issn.1006-8023.2019.02.001Chen L, Wang Q G, Yan G Y, et al. Effect of long-term nitrogen deposition on the stoichiometric characteristics of Larix gmelinii[J]. Forest Engineering, 2019, 35(2): 1−10. doi: 10.3969/j.issn.1006-8023.2019.02.001 [14] 李红丽, 徐立清, 胡建文, 等. 落叶松和红松纯林土壤对1年生胡桃楸和水曲柳苗木生长的影响[J]. 东北林业大学学报, 2021, 49(11): 1−7. doi: 10.3969/j.issn.1000-5382.2021.11.001Li H L, Xu L Q, Hu J W, et al. Effect of pure forest soil of Larix olgensis and Pinus koraiensis on growth of one-year Juglans mandshurica and Fraxinus mandshurica seedlings[J]. Journal of Northeast Forestry University, 2021, 49(11): 1−7. doi: 10.3969/j.issn.1000-5382.2021.11.001 [15] Cheng X, Yu M, Wang G G, et al. Growth, morphology and biomass allocation in response to light gradient in five subtropical evergreen broadleaved tree seedlings[J]. Journal of Tropical Forest Science, 2013, 25(4): 537−546. [16] Nakagawa M, Isogimi T, Inanaga M, et al. Inter-specific and sexual differences in architectural traits of two dioecious Lindera species (Lauraceae)[J]. Plant Ecology, 2015, 216(1): 99−109. doi: 10.1007/s11258-014-0419-7 [17] Spetea C, Rintamaki E, Schoefs B. Changing the light environment: chloroplast signalling and response mechanisms[J]. Philosophical transactions of the Royal Society B-Biological Sciences, 2014, 369: 20130220. doi: 10.1098/rstb.2013.0220 [18] 孙欣欣. 遮荫对胡桃楸和紫椴苗木形态和生理的影响[D]. 哈尔滨: 东北林业大学, 2013.Sun X X. Effect of shading on morphology and physiology in Juglans mandshurica and Tilia amurensis seedlings[D]. Harbin: Northeast Forestry University, 2013. [19] 张东来, 张玲, 葛文志. 不同光环境对胡桃楸幼苗生物量及光合生理的影响[J]. 森林工程, 2017, 33(5): 8−11.Zhang D L, Zhang L, Ge W Z. Effect of different light environment on biomass and photosynthetic physiology of Juglan mandshurica seedling[J]. Forest Engineering, 2017, 33(5): 8−11. [20] Liu B, Liu Q Q, Daryanto S, et al. Responses of Chinese fir and Schima superba seedlings to light gradients: implications for the restoration of mixed broadleaf-conifer forests from Chinese fir monocultures[J]. Forest Ecology and Management, 2018, 419: 51−57. [21] Way D A, Pearcy R W. Sunflecks in trees and forests: from photosynthetic physiology to global change biology[J]. Tree Physiology, 2012, 32(9): 1066−1081. doi: 10.1093/treephys/tps064 [22] 刘柿良, 马明东, 潘远智, 等. 不同光环境对桤木幼苗生长和光合特性的影响[J]. 应用生态学报, 2013, 24(2): 351−358.Liu S L, Ma M D, Pan Y Z, et al. Effects of light regime on the growth and photosynthetic characteristics of Alnus formosana and A. cremastogyne seedlings[J]. Chinese Journal of Applied Ecology, 2013, 24(2): 351−358. [23] 马莉薇, 张文辉, 周建云, 等. 秦岭北坡林窗大小对栓皮栎实生幼苗生长发育的影响[J]. 林业科学, 2013, 49(12): 43−50.Ma L W, Zhang W H, Zhou J Y, et al. Effects of forest gap size on the growth of Quercus variabilis seedlings on north slopes of the Qinling Mountains[J]. Scientia Silvae Sinicae, 2013, 49(12): 43−50. [24] 平晓帆, 遇文婧, 宋小双. 遮荫对引种玉簪生长及光合特性的影响[J]. 森林工程, 2018, 34(5): 20−26. doi: 10.3969/j.issn.1006-8023.2018.05.004Ping X F, Yu W J, Song X S. Effects of shading on the growth and leaf photosynthetic characteristics of introduced Hosta[J]. Forest Engineering, 2018, 34(5): 20−26. doi: 10.3969/j.issn.1006-8023.2018.05.004 [25] 智西民, 王梦颖, 牛畔青, 等. 遮荫对青桐幼苗生长性状与化学计量特征的影响[J]. 生态学杂志, 2021, 40(3): 664−671.Zhi X M, Wang M Y, Niu P Q, et al. Effects of shading on the growth indices and stoichiometric characteristics of Firmiana platanifolia seedlings[J]. Chinese Journal of Ecology, 2021, 40(3): 664−671. [26] Chun J H, Kim S, Arasu M V, et al. Combined effect of nitrogen, phosphorus and potassium fertilizers on the contents of glucosinolates in rocket salad (Eruca sativa Mill.)[J]. Saudi Journal of Biological Sciences, 2017, 24(2): 436−443. doi: 10.1016/j.sjbs.2015.08.012 [27] 刘青青, 黄智军, 马祥庆, 等. 遮阴条件下杉木幼苗生长和C、N、P化学计量特征的变化[J]. 南京林业大学学报(自然科学版), 2022, 46(3): 74−82.Liu Q Q, Huang Z J, Ma X Q, et al. Changes of seedling growth and C, N, P stoichiometric characteristics in Chinese fir under shading[J]. Journal of Nanjing Forestry University (Natural Science Edition), 2022, 46(3): 74−82. [28] 柳凤娟, 向双, 阳小成, 等. 两种光照生境下4种常绿阔叶树的单位叶面积干重、光合能力与化学防御物质含量比较[J]. 应用与环境生物学报, 2010, 16(4): 462−467.Liu F J, Xiang S, Yang X C, et al. Comparison of leaf mass per area, photosynthetic capacity and chemical defense traits of four evergreen broad-leaved tree species under contrasting light conditions[J]. Chinese Journal of Applied and Environmental, 2010, 16(4): 462−467. [29] Quiles M J, Lopez N I. Photoinhibition of photosystems Ⅰ and Ⅱ induced by exposure to high light intensity during oat plant growth: effects on the chloroplast NADH dehydrogenase complex[J]. Plant Science, 2004, 166(3): 815−823. doi: 10.1016/j.plantsci.2003.11.025 [30] Baig M J, Anand A, Mandal P K, et al. Irradiance influences contents of photosynthetic pigments and proteins in tropical grasses and legumes[J]. Photosynthetica, 2005, 43(1): 47−53. doi: 10.1007/s11099-005-7053-5 [31] 乔新荣. 光照强度对烤烟生长发育、光合特性及品质的影响[D]. 郑州: 河南农业大学, 2007.Qiao X R. Effects of light intensity on the growth, photosynthetic characteristics and quality of flue-cured tobacco[D]. Zhengzhou: Henan Agricultural University, 2007. [32] 吕强. 绥化市兰西县玉米氮磷钾最佳施肥配比研究[D]. 哈尔滨: 东北农业大学, 2018.Lü Q. Study on optimum fertilization ratio of maize nitrogen, phosphorus and potassium in Lanxi County of Suihua City[D]. Harbin: Northeast Agricultural University, 2018. -