尾叶樱桃天然种群叶表型性状变异研究

doi:10.17521/cjpe.2018.0196

摘要/Abstract

摘要：

为揭示中国特有植物尾叶樱桃(Cerasus dielsiana)在现代核心分布区天然种群的叶表型地理变异规律及其生态适应性特征, 该研究通过多重比较、巢氏方差分析、相关性分析、主成分分析(PCA)、主坐标分析(PCoA)、非加权配对算术平均法(UPGMA)聚类分析等数理方法, 对来自四川、湖北、湖南、江西、台湾5省8个尾叶樱桃天然种群的11个叶表型性状进行了比较分析, 研究其不同地理单元间叶表型多样性和地理变异规律及对地理气候的响应。结果显示: 1)尾叶樱桃主要叶表型性状变异在种群内和种群间均存在显著差异, 平均变异系数为22.44%, 其中变异系数最大和最小的分别为叶面积(50.83%)与一级侧脉数(7.96%); 平均叶表型性状的分化系数为30.78%, 种群内的变异(51.55%)大于种群间的变异(22.55%)。2) PCA表明对尾叶樱桃叶表型性状变异起主要贡献作用的前三大主成分累计贡献率达到92.400%, 可以综合概括和排序为“大小性状” (73.242%)与“形状性状” (19.158%)。3)叶宽(r = -0.641)、叶面积(r = -0.658)和一级侧脉数(r = 0.659)性状均与经度呈显著负相关或正相关关系, 气温季节变化和最湿季降水量对叶表型性状变异影响较大。4)基于PCoA和UPGMA聚类分析可将8个天然种群划分为4类。尾叶樱桃天然种群叶表型性状变异丰富, 在数量上表现出一定的连续性, “大小性状”是性状间变异的主要来源, 平均表型分化处于中等程度水平, 种群内是叶表型性状变异的主要来源; 各种群间表型分化划分结果与地理位置基本一致, 在地理空间上呈现以经度为主的梯度变异模式, “气候变异性”与“展叶期降水量”是驱动叶表型性状变异的主要气候因子, 推测这是尾叶樱桃在长期进化中与亚热带季风气候环境相适应的结果。

关键词: 尾叶樱桃, 天然种群, 地理-气候因子, 叶表型分化

Abstract:

Aims Cerasus dielsianais a wild cherry species endemic to the subtropical forest of China, and was regarded as a promising ornamental resource. Our objectives were to determine the leaf phenotypic variation, adaptation and patterns in eight natural C. dielsiana populations.

Methods We analyzed eleven leaf phenotypic traits from five provinces of China in eight natural populations of C. dielsianaby using multiple comparisons, nested analysis of variance, correlation analysis, principal component analysis (PCA), principal coordinate analysis (PCoA) and unweighted pair-group method with arithmetic mean (UPGMA) cluster analysis.

Important findings Results showed that 1) Rich leaf phenotypic variation existed among and within populations, and the average coefficient of variation (CV) was 22.44%, the maximum and the minimum were leaf area (CV = 50.83%) and primary lateral veins (CV = 7.96%), respectively. The mean differentiation coefficient (V_st) for all traits was 30.78%, and the variation within populations (51.55%) was higher than that among populations (22.55%). 2) The principal component analysis showed that the cumulative contribution rate of the first three main components of variation from leaf phenotypic traits of C. dielsiana made a major contribution reached to 92.400%, and can be comprehensively summarized and sorted as “size traits” (73.242%) and “shape traits” (19.158%). 3) Leaf width (r = -0.641), leaf area (r = -0.658) and primary lateral veins (r = 0.659) showed significant negative or positive correlation with longitude, and the temperature seasonality and precipitation of wettest quarter were showed more influence on leaf phenotype variation. 4) The eight natural populations of C. dielsiana could be divided into four groups according to principal coordinate analysis (PCA) and UPGMA cluster analysis. To sum up, leaf phenotypic variation in C. dielsiana is abundant, with a certain of continuity in quantity, and “size trait” is the main source of inter-trait variation. The mean differentiation coefficient at a moderate level, the phenotypic variation within populations was the main source of leaf traits variation. The results of phenotypic differentiation among populations were found to be consistent with the geographical location, and presented a gradient variation pattern dominated by longitude geographically. Meanwhile, the “climate variability” and “leaf-expansion period” are the main climatic factors that drive leaf phenotypic variation. We speculate the phenomena results from a long evolutionary adaptation of C. dielsiana to the subtropical monsoon climate.

Key words: Cerasus dielsiana, natural population, geography and climatic factors, leaf phenotypic differentiation

朱弘, 朱淑霞, 李涌福, 伊贤贵, 段一凡, 王贤荣. 尾叶樱桃天然种群叶表型性状变异研究. 植物生态学报, 2018, 42(12): 1168-1178. DOI: 10.17521/cjpe.2018.0196

ZHU Hong, ZHU Shu-Xia, LI Yong-Fu, YI Xian-Gui, DUAN Yi-Fan, WANG Xian-Rong. Leaf phenotypic variation in natural populations of Cerasus dielsiana. Chinese Journal of Plant Ecology, 2018, 42(12): 1168-1178. DOI: 10.17521/cjpe.2018.0196

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https://www.plant-ecology.com/CN/Y2018/V42/I12/1168

图/表 8

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代码 Code	气候变量 Climatic variables	特征值 Eigenvalue	作为第1主成分的贡献率 Variance contribution rate as first principal component (%)
bio4	气温季节变化 Temperature seasonality (SD × 100)	-0.151	97.459%
bio12	年降水量 Annual precipitation (mm)	0.784
bio16	最湿季降水量 Precipitation of wettest quarter (mm)	0.420
bio18	最暖季降水量 Precipitation of warmest quarter (mm)	0.403

代码 Code	气候变量 Climatic variables	特征值 Eigenvalue	作为第1主成分的贡献率 Variance contribution rate as first principal component (%)
bio4	气温季节变化 Temperature seasonality (SD × 100)	-0.151	97.459%
bio12	年降水量 Annual precipitation (mm)	0.784
bio16	最湿季降水量 Precipitation of wettest quarter (mm)	0.420
bio18	最暖季降水量 Precipitation of warmest quarter (mm)	0.403

种群 Population	纬度 Latitude (N)	经度 Longitude (E)	平均海拔 Mean altitude (m)	气温季节变化 Temperature Seasonality bio4 (SD × 100)	年降水量 Annual precipitation bio12 (mm)	最湿季降水量 Precipitation of wettest quarter bio16 (mm)	最暖季降水量 Precipitation of warmest quarter bio18 (mm)
四川峨眉山 Mt. Emei, Sichuan (EMS)	29.57°	103.37°	1 365	7.53	1 264.00	92.13	710.00
湖北神农架 Shennongjia, Hubei (SNJ)	31.45°	110.25°	860	8.15	1 696.00	51.61	557.00
湖北星斗山 Mt. Xingdou, Hubei (XDS)	30.02°	109.12°	839	8.19	1 772.00	60.22	568.00
湖南莽山 Mt. Mangshan, Hunan (MS)	24.97°	112.88°	1 228	7.66	1 492.00	56.44	601.00
湖南张家界 Zhangjiajie, Hunan (ZJJ)	29.32°	110.42°	1 118	7.91	1 591.00	62.14	556.00
江西金竹飞瀑 Jinzhufeipu, Jiangxi (JZFP)	27.00°	115.92°	966	7.20	1 508.00	60.26	609.00
江西仙姑坛 Xiangutan, Jiangxi (XGT)	28.45°	114.38°	1 364	6.86	1 405.00	62.72	591.00
台湾阿里山 Mt. Ali, Taiwan (ALS)	23.50°	120.80°	1 182	8.20	3 747.00	80.66	1 780.00

种群 Population	纬度 Latitude (N)	经度 Longitude (E)	平均海拔 Mean altitude (m)	气温季节变化 Temperature Seasonality bio4 (SD × 100)	年降水量 Annual precipitation bio12 (mm)	最湿季降水量 Precipitation of wettest quarter bio16 (mm)	最暖季降水量 Precipitation of warmest quarter bio18 (mm)
四川峨眉山 Mt. Emei, Sichuan (EMS)	29.57°	103.37°	1 365	7.53	1 264.00	92.13	710.00
湖北神农架 Shennongjia, Hubei (SNJ)	31.45°	110.25°	860	8.15	1 696.00	51.61	557.00
湖北星斗山 Mt. Xingdou, Hubei (XDS)	30.02°	109.12°	839	8.19	1 772.00	60.22	568.00
湖南莽山 Mt. Mangshan, Hunan (MS)	24.97°	112.88°	1 228	7.66	1 492.00	56.44	601.00
湖南张家界 Zhangjiajie, Hunan (ZJJ)	29.32°	110.42°	1 118	7.91	1 591.00	62.14	556.00
江西金竹飞瀑 Jinzhufeipu, Jiangxi (JZFP)	27.00°	115.92°	966	7.20	1 508.00	60.26	609.00
江西仙姑坛 Xiangutan, Jiangxi (XGT)	28.45°	114.38°	1 364	6.86	1 405.00	62.72	591.00
台湾阿里山 Mt. Ali, Taiwan (ALS)	23.50°	120.80°	1 182	8.20	3 747.00	80.66	1 780.00

叶表型性状 Leaf phenotypic trait	种群 Population
叶表型性状 Leaf phenotypic trait	EMS	SNJ	XDS	MS	ZJJ	JZFP	XGT	ALS	平均值 Mean	变异系数 CV (%)
叶长 LL (mm)	88.80 ± 22.85^a	79.08 ± 8.62^b	83.40 ± 17.18^ab	87.48 ± 11.27^a	89.63 ± 12.56^a	54.57 ± 12.41^d	62.87 ± 8.19^c	69.17 ± 9.14^c	76.94	23.61
叶宽 LW (mm)	43.95 ± 8.73^a	37.15 ± 5.44^bc	33.90 ± 9.02^c	34.93 ± 4.96^bc	38.92 ± 8.12^b	22.64 ± 6.34^e	26.68 ± 4.44^d	37.43 ± 3.93^bc	34.45	26.39
叶长宽比 LWR	2.01 ± 0.21^c	2.15 ± 0.19^c	2.50 ± 0.27^ab	2.53 ± 0.35^a	2.36 ± 0.38^b	2.44 ± 0.27^ab	2.37 ± 0.19^b	1.86 ± 0.14^d	2.28	14.93
叶柄长 PL (mm)	11.84 ± 3.86^a	8.86 ± 1.49^b	7.59 ± 2.46^cd	11.10 ± 2.11^a	8.25 ± 1.60^bc	6.63 ± 1.35^d	8.53 ± 0.89^bc	7.54 ± 0.89^cd	8.80	29.92
叶面积 LA (mm²)	2 746.99 ± 1 337.93^a	1 870.77 ± 476.40^bc	2 045.84 ± 1 040.24^bc	2 020.51 ± 516.34^bc	2 306.43 ± 829.25^a	890.91 ± 463.20^f	1 095.62 ± 295.07^e	1 826.66 ± 482.46^d	1 846.13	50.83
叶周长 LP (mm)	213.94 ± 50.90^a	183.08 ± 21.64^cd	191.98 ± 44.07^bc	198.99 ± 24.00^abc	207.05 ± 32.44^ab	127.23 ± 29.26^e	146.15 ± 18.91^d	171.85 ± 22.61^c	179.81	23.77
叶形指数 LI	0.73 ± 0.04^b	0.69 ± 0.05^c	0.65 ± 0.05^d	0.63 ± 0.06^d	0.65 ± 0.07^d	0.64 ± 0.06^d	0.63 ± 0.03^d	0.77 ± 0.03^a	0.67	8.90
一级侧脉数 PLV	11.00 ± 0.46^c	12.00 ± 0.71^b	12.00 ± 1.32^b	12.00 ± 0.44^b	12.00 ± 0.70^b	12.00 ± 0.77^b	13.00 ± 0.46^a	13.00 ± 0.34^a	11.88	7.96
叶脉密度 VD	0.06 ± 0.01^d	0.07 ± 0.01^c	0.07 ± 0.01^c	0.07 ± 0.01^cd	0.07 ± 0.03^cd	0.11 ± 0.01^a	0.10 ± 0.01^b	0.10 ± 0.01^b	0.08	24.42
叶尖夹角 AA (°)	27.57 ± 3.42^cd	28.99 ± 3.5^bc	28.54 ± 4.36^bc	24.56 ± 2.79^e	31.02 ± 5.67^b	25.22 ± 7.02^de	22.99 ± 2.64^e	40.24 ± 4.46^a	28.64	22.87
叶基夹角 BA (°)	114.47 ± 14.05^b	114.88 ± 11.04^b	113.87 ± 14.78^b	128.34 ± 8.24^a	103.60 ± 9.34^c	121.53 ± 17.54^b	116.68 ± 15.03^b	130.67 ± 13.33^a	116.72	13.31
平均值 Mean	-	-	-	-	-	-	-	-	-	22.44