东北地区3种桦木木质部导管特征对气候变化响应的趋同与差异

doi:10.17521/cjpe.2022.0300

植物生态学报 ›› 2023, Vol. 47 ›› Issue (8): 1144-1158.DOI: 10.17521/cjpe.2022.0300 cstr: 32100.14.cjpe.2022.0300

东北地区3种桦木木质部导管特征对气候变化响应的趋同与差异

白雨鑫¹, 苑丹阳¹, 王兴昌¹, 刘玉龙²^,³, 王晓春¹^,^*()

¹东北林业大学生态研究中心, 森林生态系统可持续经营教育部重点实验室, 哈尔滨 150040
²黑龙江省生态研究所, 哈尔滨 150081
³黑龙江牡丹江森林生态系统国家定位观测研究站, 黑龙江牡丹江 157500

收稿日期:2022-07-20 接受日期:2022-11-16 出版日期:2023-08-20 发布日期:2022-11-16
通讯作者: *ORCID: 王晓春: 0000-0002-8897-5077,(wangx@nefu.edu.cn)
基金资助:
国家自然科学基金(41877426);国家自然科学基金(42177421);中央高校基本科研业务费专项资金(2572022AW04)

Comparison of characteristics of tree trunk xylem vessels among three species of Betula in northeast China and their relationships with climate

BAI Yu-Xin¹, YUAN Dan-Yang¹, WANG Xing-Chang¹, LIU Yu-Long²^,³, WANG Xiao-Chun¹^,^*()

¹Center for Ecological Research, Key Laboratory of Sustainable Forestry Ecosystem Management-Ministry of Education, Northeast Forestry University, Harbin 150040, China
²Heilongjiang Ecology Institute, Harbin 150081, China
³Mudanjiang Forest Ecosystem Research Station, Mudanjiang, Heilongjiang 157500, China

Received:2022-07-20 Accepted:2022-11-16 Online:2023-08-20 Published:2022-11-16
Contact: *WANG Xiao-Chun(wangx@nefu.edu.cn)
Supported by:
National Natural Science Foundation of China(41877426);National Natural Science Foundation of China(42177421);Fundamental Research Funds for the Central Universities(2572022AW04)

摘要/Abstract

摘要：

桦木属(Betula)树种作为北方温带森林的先锋树种, 在次生林恢复中起重要作用。在当前的气候变化背景下, 桦木属不同种类树干木质部解剖特征对气候变化的响应与适应策略还知之甚少。该研究以黑龙江省穆棱市东北红豆杉国家级自然保护区的3种天然桦木: 白桦(B. platyphylla)、黑桦(B. dahurica)和硕桦(B. costata)为研究对象, 运用树轮年代学和树轮解剖学方法, 比较了3种桦木木质部导管特征, 分析了导管特征与季节气候因子关系、时间稳定性及生长对极端气候的抵抗力与恢复力。结果表明: 3种桦木导管数量和密度都与轮宽显著正相关。白桦在3个树种中平均轮宽最宽, 导管小且多。黑桦和硕桦平均轮宽较小, 导管明显大而少, 可能使黑桦和硕桦更容易产生栓塞。3种桦木的径向生长主要受水分因素的限制, 温度限制作用不明显。3种桦木导管数量与各季节降水量正相关, 其中硕桦的正相关关系最强。春季气温上升, 促进白桦导管变多, 非生长季(11月至次年4月)气温升高使黑桦导管变多而硕桦导管数量变少。随着气候变暖, 黑桦导管更趋向于小而多, 硕桦导管更倾向于小而少。3种桦木径向生长对生长季干旱和非生长季高温的抵抗力和恢复力趋势在种间大致相同, 且对非生长季高温的抵抗力和恢复力均较低。黑桦对非生长季高温的响应在个体间变异较大。该研究发现不同桦木木质部导管应对气候变暖的策略不同, 白桦采取较为保守的策略(产生较多且较小导管)应对气候变化; 硕桦则采用大导管提高水分运输效率的策略, 这可能导致其最先衰退, 甚至死亡; 黑桦介于白桦和硕桦之间, 导管数量和面积适中。

关键词: 树干, 木质部解剖, 导管, 气候变暖, 极端气候, 桦属

Abstract:

Aims As pioneer tree species in temperate and boreal forests, birch species (Betula spp.) play an important role in the restoration of secondary forests. Under the current climate change, little is known about the anatomical characteristics of the xylem of different birch species in response to climate change and their adaptation strategies. Therefore, we aim to study the relationship between the characteristics of their xylem vessels and climate, to reveal the response and adaptation strategies of Betula spp. to climate change, and to provide theoretical basis for accurately assessing the impact of climate change on different Betula spp. populations.

Methods In this paper, three natural birch species (B. platyphylla, B. dahurica and B. costata) from Taxus cuspidata National Nature Reserve in Muling, Heilongjiang, China were selected as the research objects. By means of dendrochronology and tree-ring anatomy, we compared these characteristics of xylem vessels of the three birch species, and analyzed the relationship between the characteristics of xylem vessels and seasonal climatic factors, temporal stability, as well as the resistance and recovery of growth to extreme climate.

Important findings (1) The vessel number and vessel density were significantly positively correlated with ring width for all three birch species. The average ring width of B. platyphylla was the widest among the three species, and the vessels were small and numerous. In B. dahurica and B. costata, the average ring width was smaller, and the vessel was significantly larger and less, which made B. dahurica more prone to embolism. (2) The growth of the three species of birch was mainly limited by moisture factors and less limited by temperature. The vessel number of the three birch species was positively correlated with precipitation in each season, and the strongest positive correlation was found in B. costata. The increase of temperature in spring promoted the increase of number of vessels, while the number of birch vessels decreased with the increase of temperature in non-growing seasons. As the climate warmed, B. dahurica tended to have smaller and more vessels, while B. costata tended to have smaller and fewer vessels. (3) The trends of resistance and recovery to drought in the growing season and heat in the non-growing season were similar among the three species, and the resistance and recovery to heat in the non-growing season were lower among the three species. There was great variation among individuals of B. dahurica in response to high temperature in non-growing season. We found that different birch species had different strategies of xylem vessel characteristics to cope with climate warming. B. platyphylla adopted a more conservative strategy (producing more and smaller vessels) to cope with climate change, while B. costata initially adopted a strategy to improve water transport efficiency through large vessels, which may lead to the first decline and even death. B. dahurica’s strategy was between B. platyphylla and B. costata, with moderate number and size of vessels.

Key words: trunk, xylem anatomy, vessel, climate warming, extreme climate, Betula

白雨鑫, 苑丹阳, 王兴昌, 刘玉龙, 王晓春. 东北地区3种桦木木质部导管特征对气候变化响应的趋同与差异. 植物生态学报, 2023, 47(8): 1144-1158. DOI: 10.17521/cjpe.2022.0300

BAI Yu-Xin, YUAN Dan-Yang, WANG Xing-Chang, LIU Yu-Long, WANG Xiao-Chun. Comparison of characteristics of tree trunk xylem vessels among three species of Betula in northeast China and their relationships with climate. Chinese Journal of Plant Ecology, 2023, 47(8): 1144-1158. DOI: 10.17521/cjpe.2022.0300

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2022.0300

https://www.plant-ecology.com/CN/Y2023/V47/I8/1144

图/表 11

表1 穆棱东北红豆杉国家级自然保护区3种桦木树轮采样的基本信息

Table 1 Basic information of tree-ring sampling of three birch species in Taxus cuspidata National Nature Reserve in Muling, Heilongjiang, China

树种(代码) Tree species (code)	纬度 Latitude (° N)	经度 Longitude (° E)	海拔 Altitude (m)	切片样本数 Core number	时间跨度 Time span	与气象站距离 Distance from weather station (km)
硕桦(BC) Betula costata (BC)	44.00	130.10	735	6	1971-2019	77
黑桦(BD) Betula dahurica (BD)	44.03	130.14	690	6	1928-2019	76
白桦(BP) Betula platyphylla (BP)	44.00	130.14	590	6	1976-2019	81

图1 穆棱东北红豆杉国家级自然保护区3种桦木木质部解剖图片。BC, 硕桦; BD, 黑桦; BP, 白桦。

Fig. 1 Anatomical pictures of xylem of three birch species in Taxus cuspidata National Nature Reserve in Muling, Heilongjiang, China. BC, Betula costata; BD, Betula dahurica; BP, Betula platyphylla.

图2 穆棱1951-2019年月平均气温和月降水量(A)、年降水量(B)、年平均气温(C)和年相对湿度(D)变化。P, 月降水量; T, 月平均气温; Tmax, 月平均最高气温; Tmin, 月平均最低气温。k, 趋势线的斜率。

Fig. 2 Variation in monthly mean air temperature and monthly precipitation (A), annual precipitation (B), mean annual temperature (C) and annual relative humidity (D) in Muling 1951-2019. P, monthly precipitation; T, mean monthly air temperature; Tmax, mean monthly maximum air temperature; Tmin, mean monthly minimum air temperature. k, slope of the trend line.

图3 3种桦木年轮宽度(RW)与导管主要特征的差异(平均值±标准差)。Kh, 理论导水率; Ks, 特异性导水率; MVA, 平均导管面积; RCTA, 导管占比; TVA, 导管总面积; VD, 导管密度; VN, 导管数量。树种代码同表1。ns, p ≥ 0.05; *, p < 0.05; **, p < 0.01; ***, p < 0.001; ****, p < 0.000 1。

Fig. 3 Difference in tree ring width (RW) and main vessel traits for three birch species (mean ± SD). Kh, theoretical hydraulic conductivity; Ks, specific hydraulic conductivity; MVA, mean vessel area; RCTA, mean percentage of conductive area within xylem; TVA, total vessel area; VD, vessel density; VN, number of vessels. See Table 1 for the codes of tree species. ns, p ≥ 0.05; *, p < 0.05; **, p < 0.01; ***, p < 0.001; ****, p < 0.000 1.

图4 3种桦木年轮宽度(RW)与主要导管特征随形成层年龄的变化趋势。Kh, 理论导水率; Ks, 特异性导水率; MVA, 平均导管面积; RCTA, 导管占比; TVA, 导管总面积; VD, 导管密度; VN, 导管数量。树种代码同表1。

Fig. 4 Trends of tree ring width (RW) and the main vessel traits for three birch species. Kh, theoretical hydraulic conductivity; Ks, specific hydraulic conductivity; MVA, mean vessel area; RCTA, mean percentage of conductive area within xylem; TVA, total vessel area; VD, vessel density; VN, number of vessels. See Table 1 for the codes of tree species.

图5 3种桦木年轮宽度(RW)与主要导管特征之间的关系。Kh, 理论导水率; Ks, 特异性导水率; MVA, 平均导管面积; RCTA, 导管占比; TVA, 导管总面积; VD, 导管密度; VN, 导管数量。Corr, 相关系数。树种代码同表1。×, p ≥ 0.05。

Fig. 5 Relationship between ring width (RW) and main vessel characteristics of three birch species. Kh, theoretical hydraulic conductivity; Ks, specific hydraulic conductivity; MVA, mean vessel area; RCTA, mean percentage of conductive area within xylem; TVA, total vessel area; VD, vessel density; VN, number of vessels. Corr, correlation coefficient. See Table 1 for the codes of tree species. ×, p ≥ 0.05.

图6 3种桦木树轮解剖参数主成分(PC)分析。Kh, 理论导水率; Ks, 特异性导水率; MVA, 平均导管面积; RCTA, 导管占比; TVA, 导管总面积; VD, 导管密度; VN, 导管数量; RW, 年轮宽度。cos2, 变量在因子图上的表示质量。树种代码同表1。

Fig. 6 Principal component (PC) analysis of anatomical parameters of the annual rings of three birch species. Kh, theoretical hydraulic conductivity; Ks, specific hydraulic conductivity; RCTA, mean percentage of conductive area within xylem; MVA, mean vessel area; TVA, total vessel area; VD, vessel density; VN, number of vessels; RW, ring width. cos2, the quality of representation for variables on the factor map. See Table 1 for the codes of tree species.

图7 3个桦木属树种树轮解剖参数主成分(PC)与各季节气候因素的相关关系。AN, 全年; AUT, 秋季; GS, 当年生长季; PGS, 上一年生长季; PNG, 上一年非生长季; pWin, 上一年冬季; SPR, 春季; SUM, 夏季。P, 月降水量; Rh, 年相对湿度; SPEI, 标准化降水-蒸散指数; T, 月平均气温; Tmax, 月平均最高气温; Tmin, 月平均最低气温。树种代码同表1。*, p < 0.05。

Fig. 7 Relationship between the principal components (PC) of the anatomical parameters of the tree whorls of three birch species and climatic factors by season. AN, annual; AUT, autumn; GS, growing season; PGS, previous growing season; PNG, previous non-growing season; pWin, previous winter; SPR, spring; SUM, summer. P, monthly precipitation; Rh, annual relative humidity; SPEI, standardized precipitation evapotranspiration index; T, mean monthly air temperature; Tmax, mean monthly maximum air temperature; Tmin, mean monthly minimum air temperature. See Table 1 for the codes of tree species. *, p < 0.05.

图8 3种桦木年轮解剖参数主成分(PC)与主要季节气候因素的滑动相关。GS, 当年生长季; PGS, 上一年生长季; PNG, 上一年非生长季。P, 月降水量; T, 月平均气温; Tmax, 月平均最高气温; Tmin, 月平均最低气温; Rh, 年相对湿度; SPEI, 标准化降水-蒸散指数。r, 相关系数。树种代码同表1。*, p < 0.05。

Fig. 8 Sliding correlations between the principal components (PC) of the anatomical parameters of the annual rings of three birch species and the main seasonal climatic factors. GS, growing season; PGS, previous growing season; PNG, previous non-growing season. P, monthly precipitation; Rh, annual relative humidity; SPEI, standardized precipitation evapotranspiration index; T, mean monthly air temperature; Tmax, mean monthly maximum air temperature; Tmin, mean monthly minimum air temperature. r, correlation coefficient. See Table 1 for the codes of tree species. *, p < 0.05.

图9 3种桦木对2个生长季干旱年平均抵抗力(Rt)和恢复力(Rc)的比较。Kh, 理论导水率; Ks, 特异性导水率; MVA, 平均导管面积; RCTA, 导管占比; RW, 年轮宽度; TVA, 导管总面积; VD, 导管密度; VN, 导管数量。树种代码同表1。ns, p ≥ 0.05。

Fig. 9 Comparison of average resistance (Rt) and recovery (Rc) of three birch species to extreme drought between the two-growing seasons. Kh, theoretical hydraulic conductivity; Ks, specific hydraulic conductivity; RCTA, mean percentage of conductive area within xylem; RW, ring width; MVA, mean vessel area; TVA, total vessel area; VD, vessel density; VN, number of vessels. See Table 1 for the codes of tree species. ns, p ≥ 0.05.

图10 3种桦木对5个非生长季高温年的平均抵抗力(Rt)和恢复力(Rc)的比较。Kh, 理论导水率; Ks, 特异性导水率; MVA, 平均导管面积; RCTA, 导管占比; RW, 年轮宽度; TVA, 导管总面积; VD, 导管密度; VN, 导管数量。树种代码同表1。ns, p ≥ 0.05。

Fig. 10 Comparison of average resistance (Rt) and recovery (Rc) of three birch species to five non-growing season heat years. Kh, theoretical hydraulic conductivity; Ks, specific hydraulic conductivity; RCTA, mean percentage of conductive area within xylem; RW, ring width; MVA, mean vessel area; TVA, total vessel area; VD, vessel density; VN, number of vessels. See Table 1 for the codes of tree species. ns, p ≥ 0.05.

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东北地区3种桦木木质部导管特征对气候变化响应的趋同与差异

Comparison of characteristics of tree trunk xylem vessels among three species of Betula in northeast China and their relationships with climate

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