细叶云南松针叶形态和显微性状地理变异及其环境解释

doi:10.17521/cjpe.2023.0041

植物生态学报 ›› 2023, Vol. 47 ›› Issue (8): 1116-1130.DOI: 10.17521/cjpe.2023.0041

所属专题：植物功能性状

细叶云南松针叶形态和显微性状地理变异及其环境解释

冯珊珊, 黄春晖, 唐梦云, 蒋维昕, 白天道^*()

广西大学林学院, 广西森林生态与保育重点实验室, 广西高校亚热带人工林培育与利用重点实验室, 南宁 530004

收稿日期:2023-02-14 接受日期:2023-04-26 出版日期:2023-08-20 发布日期:2023-05-04
通讯作者: *白天道(btd@gxu.edu.cn)
基金资助:
国家自然科学基金(32160381);广西自然科学基金(2018GXNSFBA281110)

Geographical variation of needles phenotypic and anatomic traits between populations of Pinus yunnanensis var. tenuifolia and its environmental interpretation

FENG Shan-Shan, HUANG Chun-Hui, TANG Meng-Yun, JIANG Wei-Xin, BAI Tian-Dao^*()

Guangxi Key Laboratory of Forest Ecology and Conservation, Guangxi Colleges and Universities Key Laboratory for Cultivation and Utilization of Subtropical Forest Plantation, College of Forestry, Guangxi University, Nanning 530004, China

Received:2023-02-14 Accepted:2023-04-26 Online:2023-08-20 Published:2023-05-04
Contact: *BAI Tian-Dao(btd@gxu.edu.cn)
Supported by:
National Natural Science Foundation of China(32160381);Natural Science Foundation of Guangxi(2018GXNSFBA281110)

摘要/Abstract

摘要：

叶片是林木获取和利用生存资源的重要器官, 其形态结构一定程度上反映了林木的生境适应性。探究南盘江—红水河流域的地理气候对该区域重要树种细叶云南松(Pinus yunnanensis var. tenuifolia)针叶形态及显微结构的塑造作用, 对理解该树种的生态适应性及资源保育具有重要参考价值。该研究以分布于贵州和广西的细叶云南松9个自然种群的18个针叶形态及显微性状为研究对象, 结合种群所在地7个地理和气候因子, 采用巢氏方差分析、相关分析、多元统计分析(主成分分析、冗余分析和系统聚类分析)方法解析其种群差异及环境关联。结果表明, 除叶截面面积与中柱截面面积之比(V1)外, 其他指标在种群间均存在不同程度的分化(表型分化系数(V_ST) = 22.32%-51.42%), 即种群间生境异质性对大部分指标有显著影响。Pearson相关分析和多元统计分析表明: 针叶树脂道相关指标(树脂道数目、周长、面积等)随纬度、海拔、降水量增加而增大, 随年平均气温升高而减小; 气孔相关指标(气孔密度、气孔密度与V1之比)随经纬度增加而增大, 随相对湿度增加而减小; 针叶横截面大小相关指标(叶宽、叶厚、叶截面面积、中柱面积等)则主要与采样地至南盘江—红水河谷的距离显著相关, 距离越近, 指标值越小。综上, 细叶云南松在与云南松(P. yunnanensis)截然不同的干热生境驱动下, 表现出所有树脂道性状小型化趋势, 细叶云南松较高的气孔密度(以及下陷的气孔)有利于其在干旱环境下平衡呼吸和蒸腾失水作用, 而其相对细柔的针叶则可能主要由该区域特殊的河谷地形造成的焚风及峡谷强风的胁迫作用以及季节性暖干气候对针叶的生长限制作用所塑造。

关键词: 生态适应性, 干热生境, 地理因子, 气候因子, 地理变异, 资源保育

Abstract:

Aims Leaf is an important organ for forest trees to acquire and utilize survival resources, and its morphological structure reflects the habitat adaptability of trees to a certain extent. Exploring the role of geography and climate of the Nanpan-Hongshui River basin in shaping the needles morphology and microstructure of an important tree species in the region, Pinus yunnanensis var. tenuifolia, has important reference value for understanding the ecological adaptability and resource conservation of this tree species.

Methods Eighteen morphological and microscopic characters of nine wild populations which distributed in Guizhou and Guangxi were measured, and seven geographical and climatic factors of the nine population locations were recorded. The population divergence and environmental associations were analyzed by nested ANOVA, correlation analysis, multivariate statistical analysis (principal component analysis, redundancy analysis, and hierarchical clustering analysis).

Important findings The results showed that, except for the ratio of needle cross-sectional area to central cylinder area (V1), all indicators had different degrees of differentiation among populations (phenotypic differentiation coefficient (V_ST) = 22.32%-51.42%). It implied that the habitat heterogeneity among populations had a significant impact on most indicators. Pearson correlation analysis and multivariate statistical analysis showed that the indicators which related to needle resin canals (resin canals number, resin canals perimeter, resin canals area, etc.) increased with the increase of latitude, altitude, and mean annual precipitation, but decreased with the increase of mean annual temperature; the stomatal indicators (stomatal density, ratio of stomatal density to V1) increased with increasing latitude and longitude, and decreased with increasing relative humidity; the indicators which related to the needle cross-sectional size (needle width, needle thickness, needle cross-sectional area, central cylinder area, etc.) are mainly affected by the distance from the sampling site to the Nanpan-Hongshui River. The closer the distance, the smaller the indicator value is. To sum up, P. yunnanensis var. tenuifolia exhibits a trend of miniaturization of resin canal, which was driven by dry-hot habitat selection that is different from that of the original species Pinus yunnanensis. Its higher stomatal density (and sunken stomata) is conducive to balancing respiration and transpiration dehydration in arid environments. The relatively slender needles may be mainly shaped by the stress effects of foehn and strong canyon winds which caused by the special valley terrain in the region, as well as the growth limiting effects of seasonal warm and dry climate on needles.

Key words: ecological adaptivity, dry-hot habitat, geographical factor, climatic factor, geographical variation, resource conservation

冯珊珊, 黄春晖, 唐梦云, 蒋维昕, 白天道. 细叶云南松针叶形态和显微性状地理变异及其环境解释. 植物生态学报, 2023, 47(8): 1116-1130. DOI: 10.17521/cjpe.2023.0041

FENG Shan-Shan, HUANG Chun-Hui, TANG Meng-Yun, JIANG Wei-Xin, BAI Tian-Dao. Geographical variation of needles phenotypic and anatomic traits between populations of Pinus yunnanensis var. tenuifolia and its environmental interpretation. Chinese Journal of Plant Ecology, 2023, 47(8): 1116-1130. DOI: 10.17521/cjpe.2023.0041

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2023.0041

https://www.plant-ecology.com/CN/Y2023/V47/I8/1116

图/表 10

图1 细叶云南松采样点分布图。

Fig. 1 Sampling sites of Pinus yunnanensis var. tenuifolia.

表1 细叶云南松针叶参试性状指标名称、缩写及单位

Table 1 Names, abbreviations, and units of tested needle characters for Pinus yunnanensis var. tenuifolia

性状缩写 Abbreviation of character	单位 Unit	性状名称 Name of character
α_A	°	针叶截面夹角 Angle of needle cross section
NW	mm	叶宽 Needle width
NT	mm	叶厚 Needle thickness
NSP	mm	叶截面周长 Needle cross-section perimeter
NSA	mm²	叶截面面积 Needle cross-section area
CCW	mm	中柱宽 Central cylinder width
CCT	mm	中柱厚 Central cylinder thickness
CCP	mm	中柱截面周长 Central cylinder perimeter
CCA	mm2	中柱截面面积 Central cylinder area
RCN	No.	树脂道数 Number of resin canals
RCP	mm	树脂道总周长 Resin canal perimeter
RCA	mm2	树脂道总面积 Resin canal area
MA	mm2	叶肉面积 Mesophyll area
MSD	No.·mm^-2	气孔密度 Mean stomatal density
V1		叶截面面积与中柱截面面积之比 NSA/CCA
V2		树脂道数与气孔密度之比 RCN/MSD
V3		树脂道数与V1之比 RCN/V1
V4		气孔密度与V1之比 MSD/V1

图2 细叶云南松针叶显微形态。A, 针叶表面(纵向)气孔分布状态。B, 针叶横截面图。C, 针叶横截面外皮层、树脂道及气孔放大。D, 针叶横截面模式图。αA, 针叶截面夹角; c, 中柱; e, 表皮; h, 皮下层; m, 叶肉; p, 韧皮部; r, 树脂道; s, 气孔; x, 木质部。CCT, 中柱厚; CCW, 中柱宽; NT, 针叶厚; NW, 针叶宽; RCA, 树脂道总面积; RCP, 树脂道总周长。

Fig. 2 Needle microscopic structures of Pinus yunnanensis var. tenuifolia. A, Stomata and stomatal line distribution. B, Cross section of needle. C, Enlarged exodermis, resin canal and stomata. D, Model picture of cross section of needle. αA, angle of needle cross section; c, central cylinder; e, epidermis; h, subcutaneous layer; m, mesophyll; p, phloem; r, resin canal; s, stomata; x, xylem. CCT, central cylinder thickness; CCW, central cylinder width; NT, needle thickness; NW, needle width; RCA, resin canal area; RCP, resin canal perimeter.

表2 细叶云南松针叶性状种群间和种群内F值、方差分量百分比及表型分化系数

Table 2 F value, percentage of variance component among and within populations of needle characters and phenotypic differentiation coefficient of Pinus yunnanensis var. tenuifolia

性状 Trait	F		方差分量百分比 Percentage of variance component (%)			表型分化系数 Phenotypic differentiation coefficient (V_ST, %)
性状 Trait	种群间 Among populations	种群内 Within population	种群间 Among populations	种群内 Within population	随机误差 Random error	表型分化系数 Phenotypic differentiation coefficient (V_ST, %)
α_A	2.49 ^*	1.59^*	2.83	9.85	87.32	22.32
NW	16.88^***	3.52^***	17.99	27.13	54.88	39.87
NT	14.83^***	2.92^***	17.64	22.57	59.79	43.86
NSP	12.40^***	3.41^***	13.59	27.62	58.79	32.98
NSA	13.74^***	3.20^***	15.54	25.43	59.04	37.93
CCW	11.04^***	2.38^***	14.36	18.21	67.43	44.09
CCT	3.68^***	1.46^*	5.68	7.74	86.58	42.31
CCP	7.62^***	2.36^***	10.27	18.87	70.86	35.24
CCA	6.96^***	2.17^***	9.37	16.71	73.92	35.94
RCN	55.63^***	11.90^***	26.79	50.01	23.20	34.88
RCP	39.13^***	9.31^***	22.44	49.66	27.90	31.13
RCA	20.74^***	7.02^***	15.23	45.99	38.77	24.88
MA	17.64^***	4.80^***	15.95	35.87	48.18	30.78
MSD	12.04^***	2.33^***	18.08	17.08	64.85	51.42
V1	1.70	3.90^***	0.00	34.08	65.92	0.00
V2	22.60^***	5.68^***	18.68	38.97	42.35	32.40
V3	44.45^***	10.77^***	23.25	50.53	26.22	31.51
V4	10.02^***	2.95^***	12.78	24.17	63.04	34.59
平均值 Mean			14.47	28.92	56.61	33.67

表3 细叶云南松种群针叶性状多重比较结果(平均值±标准差)

Table 3 Tukey HSD test on the needle traits between populations of Pinus yunnanensis var. tenuifolia (mean ± SD)

图3 细叶云南松针叶性状与地理和气候因子的Pearson相关分析。***, p < 0.001; **, p < 0.01;*, p < 0.05。αA, 针叶截面夹角; CCA, 中柱截面积; CCP, 中柱截面周长; CCT, 中柱厚; CCW, 中柱宽; MA, 叶肉面积; MSD, 平均气孔密度; NSA, 针叶截面面积; NSP, 针叶截面周长; NT, 针叶厚; NW, 针叶宽; RCA, 树脂道总面积; RCN, 树脂道数; RCP, 树脂道总周长; V1, NSA/CCA; V2, RCN/MSD; V3, RCN/V1; V4, MSD/V1。ALT, 海拔; DFSSR, 样点至河流距离; LAT, 纬度; LON, 经度; MAP, 年降水量; MAT, 年平均气温; MRH, 相对湿度。

Fig. 3 Pearson correlations between needle characters of Pinus yunnanensis var. tenuifolia and geographical and climatic factors. ***, p < 0.001; **, p < 0.01; *, p < 0.05. αA, angle of needle cross section; CCA, central cylinder area; CCP, central cylinder perimeter; CCT, central cylinder thickness; CCW, central cylinder width; MA, mesophyll area; MSD, mean stomatal density; NSP, needle cross-section perimeter; NSA, needle cross-section area; NT, needle thickness; NW, needle width; RCA, resin canal area; RCN, number of resin canals; RCP, resin canal perimeter; V1, NSA/CCA; V2, RCN/MSD; V3, RCN/V1; V4, MSD/V1. ALT, altitude; DFSSR, distance from sampling site to river; LAT, latitude; LON, longitude; MAP, annual precipitation; MAT, mean annual air temperature; MRH, mean relative humidity.

表4 细叶云南松种群针叶性状的地理和气候因子主成分(PC)分析

Table 4 Principal component (PC) analysis of geographical and climatic factors for needle characters of Pinus yunnanensis var. tenuifolia populations

地理和气候因子 Geographical and climatic factor	主成分载荷 Principal component loading
地理和气候因子 Geographical and climatic factor	PC1	PC2	PC3	PC4
纬度 Latitude	0.213	-0.877	0.380	0.152
经度 Longitude	-0.850	-0.360	0.369	-0.010
年平均气温 Mean annual air temperature	-0.869	0.441	-0.100	0.044
相对湿度 Mean relative humidity	0.205	0.902	0.242	-0.238
年降水量 Mean annual precipitation	0.867	0.306	-0.259	0.274
海拔 Altitude	0.838	-0.288	0.321	-0.252
采样点至河流距离 Distance from sampling site to river	0.079	0.704	0.660	0.241
特征值 Eigenvalue	3.02	2.58	0.95	0.28
贡献率 Contribution rate (%)	43.21	36.85	13.64	3.98

图4 细叶云南松种群地理和气候因子主成分(PC)与针叶性状的回归分析(平均值±标准误)。A-E为与PC1回归显著的指标。F-G为与PC2回归显著的指标。MSD, 平均气孔密度; RCA, 树脂道总面积; RCN, 树脂道数; RCP, 树脂道总周长; V1, 叶截面面积与中柱截面面积之比(NSA/CCA); V2, RCN/MSD; V3, RCN/V1; V4, MSD/V1。

Fig. 4 Regression analysis between principal components (PC) of geographical and climatic factors and needle characters of Pinus yunnanensis var. tenuifolia populations (mean ± SE). A-E are characters that have significant regression with PC1. F-G are characters that have significant regression with PC2. MSD, mean stomatal density; RCA, resin canal area; RCN, number of resin canals; RCP, resin canal perimeter; V1, needle cross-section area divided by central cylinder area (NSA/CCA); V2, RCN/MSD; V3, RCN/V1; V4, MSD/V1.

图5 细叶云南松种群针叶性状与地理和气候因子的冗余分析(RDA)。αA, 针叶截面夹角; CCA, 中柱截面面积; CCP, 中柱截面周长; CCT, 中柱厚; CCW, 中柱宽; MA, 叶肉面积; MSD, 平均气孔密度; NSA, 针叶截面面积; NSP, 针叶截面周长; NT, 针叶厚; NW, 针叶宽; RCA, 树脂道总面积; RCN, 树脂道数; RCP, 树脂道总周长; V1, NSA/CCA; V2, RCN/MSD; V3, RCN/V1; V4, MSD/V1。

Fig. 5 Redundancy analysis (RDA) between needle characters and geo-climatic factors of Pinus yunnanensis var. tenuifolia populations. αA, angle of needle cross section; CCA, central cylinder area; CCP, central cylinder perimeter; CCT, central cylinder thickness; CCW, central cylinder width; MA, mesophyll area; MSD, mean stomatal density; NSA, needle cross-section area; NSP, needle cross-section perimeter; NT, needle thickness; NW, needle width; RCA, resin canal area; RCN, number of resin canals; RCP, resin canal perimeter; V1, NSA/CCA; V2, RCN/MSD; V3, RCN/V1; V4, MSD/V1.

图6 参试细叶云南松种群地理和气候及针叶性状最佳分组数(A-C)和层次聚类树(D)。A, 基于针叶性状的种群最佳分组。B, 地理和气候因子最佳分组。C, 针叶性状最佳分组。D, 参试种群针叶性状及地理和气候因子聚类结果。αA, 针叶截面夹角; CCA, 中柱截面面积; CCP, 中柱截面周长; CCT, 中柱厚; CCW, 中柱宽; MA, 叶肉面积; MSD, 平均气孔密度; NSA, 针叶截面面积; NSP, 针叶截面周长; NT, 针叶厚; NW, 针叶宽; RCA, 树脂道总面积; RCN, 树脂道数; RCP, 树脂道总周长; V1, NSA/CCA; V2, RCN/MSD; V3, RCN/V1; V4, MSD/V1。ALT, 海拔; DFSSR, 样点至河流距离; LAT, 纬度; LON, 经度; MAP, 年降水量; MAT, 年平均气温; MRH, 相对湿度。图D中呈现的针叶性状值、地理和气候因子为分别标准化后的数值。

Fig. 6 Optimal number of clusters (A-C) and hierarchical cluster tree (D) of geographical and climatic factors and needle traits of Pinus yunnanensis var. tenuifolia populations. A, Optimal number of clusters of populations based on needle traits. B, Optimal number of clusters of geographical and climatic factors. C, Optimal number of clusters of needle traits. D, Cluster results of needle traits and geographical and climatic factors in the tested populations. αA, angle of needle cross section; CCA, central cylinder area; CCP, central cylinder perimeter; CCT, central cylinder thickness; CCW, central cylinder width; MA, mesophyll area; MSD, mean stomatal density; NSA, needle cross-section area; NT, needle thickness; NW, needle width; NSP, needle cross-section perimeter; RCA, resin canal area; RCN, number of resin canals; RCP, resin canal perimeter; V1, NSA/CCA; V2, RCN/MSD; V3, RCN/V1; V4, MSD/V1. ALT, altitude; DFSSR, distance from sampling site to river; LAT, latitude; LON, longitude; MAP, annual precipitation; MAT, mean annual air temperature; MRH, mean relative humidity. The values of needle character and geographical and climatic factors in plot D were standardized before visualization.

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细叶云南松针叶形态和显微性状地理变异及其环境解释

Geographical variation of needles phenotypic and anatomic traits between populations of Pinus yunnanensis var. tenuifolia and its environmental interpretation

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