Comparison of tree-ring xylem anatomical parameters of three tree species under different moisture conditions in the Muling area
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摘要:
目的 明确不同生境下树木木质部的解剖特征差异和暖干化气候条件下树木木质部的生长变化。 方法 在黑龙江省穆棱市针阔叶混交林中设置了水分偏多和偏少2个生境采样点，在2个样点内分别对红松、蒙古栎、紫椴进行取样，使用滑走切片机对所取样芯进行切片，获取木质部解剖参数。 结果 偏干生境下蒙古栎的平均导管面积（MVA）、木质部脆弱性指数（Vx）显著增大，但蒙古栎的水力效率（Ks）在2个生境间差异不显著。从偏湿生境到偏干生境，紫椴的MVA、Vx、Ks显著降低。在3个树种中，只有红松的平均管胞面积（MTA）和Vx在2个生境下差异不显著，同时在偏干生境下Ks显著提高。偏干生境下红松、蒙古栎、紫椴的年轮宽度（RW）均与生长季的帕默尔干旱指数（PDSI）呈显著正相关，且其敏感性随土壤含水量的增加而降低。偏干生境下蒙古栎的Ks与前一年11月―当年10月的PDSI显著负相关，紫椴的RW、MVA与前一年非生长季最低温显著正相关。 结论 生长季初期干旱是制约穆棱地区红松管胞面积和水力效率的重要因素，在未来红松可能会更强烈地感受到东北地区的气候变化。紫椴在偏干生境下降低水力效率以提高水力安全。偏干生境下的蒙古栎生长量和平均导管面积大于偏湿生境，所以未来的暖干化气候条件可能对蒙古栎的生长更有益，但存在空穴化风险。明确全球变暖带来的干旱胁迫是否会改变树种生长状况，对调整当前的造林措施以使针阔混交林更好地应对未来的暖干化气候条件至关重要。 Abstract:Objective To clarify the differences in the anatomical characteristics of the xylem of trees in different habitats and the growth changes of the xylem of trees under warm and dry climate conditions. Method The relatively dry and relatively wet habitat sampling sites were set up in mixed coniferous forests in Muling City, Heilongjiang Province, and Pinus koraiensis, Quercus mongolica, and Tilia amurensis were sampled. Micro-sections were cut with a rotary microtome to obtain the anatomical parameters of the xylem. Result The mean vessel area (MVA) and vulnerability index (Vx) of Quercus mongolica in the relatively dry habitat increased significantly, but the theoretical xylem-specific hydraulic conductivity (Ks) of Quercus mongolica did not differ significantly between the two habitats. From relatively wet habitat to relatively dry habitat, MVA, Vx, and Ks of Tilia amurensis were significantly reduced. Among the three tree species, only the mean tracheid area (MTA) and Vx of Pinus koraiensis did not differ significantly in the two habitats, and at the same time, the Ks was significantly increased in the relatively dry habitat. The tree-ring widths (RW) of Pinus koraiensis, Quercus mongolica, and Tilia amurensis in relatively dry habitat were all significantly positively correlated with the Palmer drought severity index (PDSI) of the growing season, and the sensitivity decreased when the water content of soil rose. The Ks of Quercus mongolica in the relatively dry habitat was significantly negatively correlated with the PDSI from November of the previous year to October of the current year. The RW and MVA of Tilia amurensis in the relatively dry habitat was significantly positively correlated with the lowest temperature in the previous non-growing season (PNG). Conclusion At the beginning of the growing season, drought was an important factor that restricted the tracheid area and hydraulic efficiency of Pinus koraiensis in the Muling area. In the future, Pinus koraiensis may feel the change of climatic conditions in northeast China more strongly. Tilia amurensis reduces hydraulic efficiency in relatively dry habitats to improve hydraulic safety. The growth and MVA of Quercus mongolica in the relatively dry habitat were greater than that in the relatively wet habitat, so future warm and dry climate conditions may be more beneficial to the growth of Quercus mongolica but there was also a risk of cavitation to it. Clarifying whether drought stress due to global warming will alter tree growth is critical to adjusting current silvicultural practices, which is better for mixed coniferous forests to cope with future warming and drying climatic conditions. -
图 1 红松、紫椴、蒙古栎的木质部显微切片图
空心箭头表示生长方向；T. 管胞；V. 导管；红松/紫椴木质部显微切片图的比例尺为100 μm，蒙古栎木质部显微切片图的比例尺为1 mm。Hollow arrows indicate growth direction; T, tracheid; V, vessel; Scale bars are 100 μm in the microsection of xylem of Pinus koraiensis/Tilia amurensis and 1 mm in the microsection of xylem of Quercus mongolica.
Figure 1. Microsection of xylem of Pinus koraiensis, Tilia amurensis, and Quercus mongolica among sites
图 2 不同生境下红松、蒙古栎、紫椴树轮宽度和木质部解剖参数的主成分分析
RW. 年轮宽度；MVA/MTA. 平均导管/管胞面积；VAmax/TAmax. 最大导管/管胞面积；Vx. 木质部脆弱性指数；Ks. 水力效率；RCTA. 导水面积百分比；CWT. 细胞壁厚度；Dh. 平均水力直径。下同。 RW, ring width; MVA/MTA, mean vessel area/mean tracheid area; VAmax/TAmax, annual ring with largest mean vessel area/annual ring with largest mean tracheid area; Vx, vulnerability index; Ks, theoretical xylem-specific hydraulic conductivity; RCTA, mean percentage of the conductive area within xylem; CWT, overall mean thickness of all cell walls; Dh, overall mean hydraulic diameter. The same below.
Figure 2. Principal component analysis （PCA） of ring width and xylem anatomical parameters of Pinus koraiensis, Quercus mongolica, and Tilia amurensis among sites
图 3 不同生境下红松、蒙古栎、紫椴树轮宽度和木质部解剖特征的箱线图对比
“*”、“NS”分别表示差异显著（P < 0.05）和无显著差异。*, NS indicate significant difference at P < 0.05 and P > 0.05 levels, respectively.
Figure 3. Boxplots for characteristics of tree-ring width and xylem anatomy features of Pinus koraiensis, Quercus mongolica, and Tilia amurensis among sites
图 4 不同生境下的红松、蒙古栎、紫椴的木质部解剖特征年表/树轮宽度年表与1951—2019年期间月PDSI数据的相关性
小写字母代表上一年的月份。 “实心”、“空心”分别表示在P < 0.05水平上的显著相关和不显著相。Lower case letters represent months of the previous year. “solid”, “hollow” indicate significant correlations at P < 0.05 and P > 0.05 levels, respectively.
Figure 4. Correlations between xylem anatomy features/ring width chronologies of Pinus koraiensis, Quercus mongolica, and Tilia amurensis among sites and monthly climate variables, PDSI for the period 1951–2019
图 6 不同生境下的红松、蒙古栎、紫椴的木质部解剖特征年表/树轮宽度年表与1951—2019年期间季节PDSI和季节最低温数据的相关性
PNG. 前一年非生长季（前一年11—当年3月）；BGS. 生长季初期（当年4—5月）；MGS. 生长季中期（当年6—8月）；EGS. 生长季末期（当年9—10）。PNG, Previous non-growing season (previous November to current March); BGS, Beginning of the growing season (current April to May); MGS, Middle of the growing season (current June to August); EGS, End of the growing season (current September to October).
Figure 6. Correlations between xylem anatomy features/ring width chronologies of Pinus koraiensis, Quercus mongolica, and Tilia amurensis among sites and the season climate variables, PDSI and minimum temperature for the period 1951–2019
图 5 不同生境下的红松、蒙古栎、紫椴的木质部解剖特征年表/树轮宽度年表与1951—2019年期间月最低温数据的相关性
Figure 5. Correlations between xylem anatomy features/ring width chronologies of Pinus koraiensis, Quercus mongolica, and Tilia amurensis among sites and the monthly climate variables, minimum temperature for the period 1951–2019
表 1 不同生境下红松、蒙古栎、紫椴不同土层土壤的含水率
Table 1. Soil moisture content in different soil layers of Pinus koraiensis, Quercus mongolica, and Tilia amurensis among sites
树种 Tree species 生境 Habitat 土壤含水率 Soil moisture content/% 0 ~ 20 cm 20 ~ 40 cm 40 ~ 60 cm 60 ~ 80 cm 80 ~ 100 cm 红松 Pinus koraiensis 偏干 Relatively dry 10.15a 5.34a 3.54a 3.24a 2.62a 偏湿 Relatively wet 15.78b 12.53b 9.2b 7.62b 4.68a 蒙古栎 Quercus mongolica 偏干 Relatively dry 30.49a 10.55a 5.37a 5.18a 5.16a 偏湿 Relatively wet 42.69b 14.12b 12.84b 12.66b 11.08b 紫椴 Tilia amurensis 偏干 Relatively dry 22.89a 12.19a 5.14a 4.85a 4.54a 偏湿 Relatively wet 30.85b 19.71b 12.31b 11.68b 10.64b 注：同一树种同一土层不同小写字母代表P < 0.05水平的差异显著性。Notes: Different lowercase letters for the same tree species and the same soil layer represent significant differences at the P < 0.05 level. 
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表 2 不同生境下红松、蒙古栎、紫椴的采样点信息及所取样芯数量
Table 2. The information of the sampling sites and the number of sampled cores of Pinus koraiensis, Quercus mongolica, and Tilia amurensis among sites
树种
Tree species生境
Habitat坡向
Aspect海拔
Altitude/m土壤有机质
Soil organic matter/（g·kg−1）纬度
Latitude经度
Longitude样芯数量
Core number红松
Pinus koraiensis偏干
Relatively dry南 South 694 109.33a 44°02′N 130°09′E 40 偏湿
Relatively wet西 West 654 162.85b 43°58′N 130°05′E 36 蒙古栎
Quercus mongolica偏干
Relatively dry南 South 708 89.88a 44°02′N 130°08′E 42 偏湿
Relatively wet东北 Northeast 559 120.46b 44°00′N 130°10′E 40 紫椴
Tilia amurensis偏干
Relatively dry东南 Southeast 713 118.32a 43°57′N 130°04′E 36 偏湿
Relatively wet西 West 707 244.5b 44°11′N 130°12′E 40 注：同一树种不同小写字母代表P < 0.05水平的差异显著性。Notes: Different lowercase letters for the same tree species represent significant differences at the P < 0.05 level.
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表 3 不同生境下红松、蒙古栎、紫椴标准年表的主要统计特征
Table 3. Major statistic characteristics of the standard chronologies of Pinus koraiensis, Quercus mongolica, and Tilia amurensis among sites
树种
Tree species生境
Habitat时间跨度
Time span标准年表特征值
Eigenvalue of standard chronology公共区间统计量
Common interval analysisMS SD MC AC SNR EPS VF/% 红松
Pinus koraiensis偏干
Relatively dry1890—2019 0.30 0.29 0.69 0.56 26.1 0.96 52.64 偏湿
Relatively wet1922—2019 0.18 0.38 0.52 0.88 14.1 0.93 35.8 蒙古栎
Quercus mongolica偏干
Relatively dry1802—2019 0.24 0.21 0.62 0.42 10.0 0.91 34.1 偏湿
Relatively wet1809—2019 0.28 0.27 0.67 0.47 19.8 0.95 41.8 紫椴
Tilia amurensis偏干
Relatively dry1843—2019 0.31 0.28 0.59 0.53 13.3 0.93 39.2 偏湿
Relatively wet1825—2019 0.30 0.32 0.56 0.70 17.4 0.95 43.2 注：MS. 平均敏感度；SD. 标准偏差；MC. 平均相关系数；AC. 一阶自相关；SNR. 信噪比；EPS. 样本量的总体解释信号；VF. 第一特征根解释量。Notes: MS, mean sensitivity; SD, standard deviation; MC, mean correlation; AC, first order autocorrelation; SNR, signal-to-noise ratio; EPS, expressed population signal; VF, variance in first eigenvector.
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