云南松、思茅松和卡西亚松天然种群间的针叶表型变异

doi:10.17521/cjpe.2022.0263

植物生态学报 ›› 2023, Vol. 47 ›› Issue (6): 833-846.DOI: 10.17521/cjpe.2022.0263

云南松、思茅松和卡西亚松天然种群间的针叶表型变异

李卫英¹^,², 章正仁², 辛雅萱¹, 王飞¹, 辛培尧¹^,^*(), 高洁²^,^*()

¹西南林业大学国家林业和草原局西南风景园林工程技术研究中心, 西南林业大学西南地区生物多样性保育国家林业和草原局重点实验室, 昆明 650224
²中国科学院西双版纳热带植物园热带森林生态学重点实验室, 云南勐腊 666303

收稿日期:2022-06-22 接受日期:2022-10-31 出版日期:2023-06-20 发布日期:2022-10-31
通讯作者: * (Xin PY, xpytgyx@163.com;Gao J, gaojie@xtbg.org.cn)
作者简介:ORCID:李卫英: 0000-0003-1055-2522
基金资助:
国家自然科学基金(31770701);云南省基础研究专项(202201AT070217)

Needle phenotype variation among natural populations of Pinus yunnanensis, P. kesiya var. langbianensis and P. kesiya

LI Wei-Ying¹^,², ZHANG Zheng-Ren², XIN Ya-Xuan¹, WANG Fei¹, XIN Pei-Yao¹^,^*(), GAO Jie²^,^*()

¹Southwest Research Center for Engineering Technology of Landscape Architecture National Forestry and Grassland Administration, Key Laboratory of National Forestry and Grassland Administration on Biodiversity Conservation in Southwest China, Southwest Forestry University, Kunming 650224, China
²CAS Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Garden, Chinese Academy of Sciences, Mengla, Yunnan 666303, China

Received:2022-06-22 Accepted:2022-10-31 Online:2023-06-20 Published:2022-10-31
Contact: * (Xin PY, xpytgyx@163.com;Gao J, gaojie@xtbg.org.cn)
Supported by:
National Natural Science Foundation of China(31770701);Basic Research Program of Yunnan Province(202201AT070217)

摘要/Abstract

摘要：

研究近缘种复合体的针叶表型变异及其地理分布格局, 有助于从生理生态机制上理解针叶树种的地理变异、种群动态以及对区域气候的响应。云南松(Pinus yunnanensis)、思茅松(P. kesiya var. langbianensis)和卡西亚松(P. kesiya)是一个跨越了东南亚热带和亚热带地区的近缘种复合体, 具有典型的地理替代分布特征, 拥有丰富的表型变异和遗传变异。该研究在3种松树分布区采集了31个代表种群, 每个种群选择10个样木, 测量了针叶长度、气孔密度、气孔保卫细胞长度和宽度、木质增厚层长度和宽度、气孔腔长度和宽度等共8个性状。采用巢式方差分析计算性状在树种间和种群间的差异, 通过主成分分析和聚类分析揭示针叶性状的种群变异结构; 分析针叶性状随纬度的变化规律, 运用多元线性回归模型来确定影响针叶性状变异的主要环境因子。结果表明: (1)针叶性状在种群间的变异系数为12.01%-34.08%, 气孔保卫细胞长度、木质增厚层长度和宽度、针叶长度和气孔密度的表型分化系数较高; (2)云南松和卡西亚松在大多数性状上存在显著差异, 思茅松介于二者之间, 聚类分析结果表明3种松树针叶性状的种群变异结构与其地理区域分布一致; (3)针叶长度和气孔密度与纬度正相关, 气孔腔长度和宽度与纬度负相关; (4)影响针叶性状变化的关键环境因子是最干季平均气温、降水量季节性变异系数、平均气温日较差、最湿月降水量和最干月降水量。3种松树针叶性状具有丰富的种间和种群间变异, 呈现显著的纬度梯度变异趋势, 反映了其对环境长期适应的结果。热带亚热带松树针叶性状变异及其对环境因子的响应可为造林育种的地理种源选择提供科学依据。

关键词: 云南松, 思茅松, 卡西亚松, 地理变异, 针叶, 解剖性状, 温度, 降水

Abstract:

Aims Investigating the needle phenotypic variation and its geographical distribution pattern of related species complexes will aid in understanding conifer geographic variation, population dynamics, and physiological and ecological responses to geographical climates. Here we focus on the Pinus kesiya, P. yunnanensis, and the hybrid species P. kesiyavar. langbianensis, which are primarily distributed across tropical and subtropical Southeast Asia, exhibiting abundant climate and genetic variation.

Methods We selected thirty-one representative populations covering the distribution areas of three pine species, and sampled ten individuals from each population. Eight traits were measured, including needle length, stomatal density, stomatal guard cell length, stomatal guard cell width, woody thickening layer length, woody thickening layer width, stomatal cavity length, and stomatal cavity width. Variation between species and populations was assessed using a nested variance analysis. Principal components analysis was applied to evaluate the underlying dimensionality of the needle variation. Clustering analysis among populations was performed using the Ward method to infer the population structure. Pearson’s correlation coefficient between latitude and needle characters was assessed. Finally, a multiple linear regression model was used to identify the main environmental factors influencing needle trait variation.

Important findings The variation coefficients of the eight needle traits ranged from 12.01% to 34.08% across populations. The phenotypic differentiation coefficient was higher for stomata guard cell length, woody thickening layer length and width, needle length, and stomatal density. Most morphological needle characteristics were significantly different between P. yunnanensis and P. kesiya,while P. kesiyavar. langbianensis showed intermediate values between the two parental pines. Hierarchical clustering analysis showed that the variation pattern of needle traits was related to geographical region. Needle length and stomatal density were positively correlated with latitude, while the stomatal cavity length and width were negatively correlated with latitude. The key environmental factors affecting the needle traits of the three pine species were mean temperature during the driest quarter, precipitation seasonality, mean diurnal range, precipitation during wettest month, and precipitation during the driest month. The needle traits of the three pine species displayed high interspecific and interpopulation variation. The significant latitudinal gradient trend in needle trait variation indicates long-term evolutionary adaptation to the environment. The response of needle character variation to environmental factors can provide essential insights for geographical provenance selection in afforestation breeding

Key words: Pinus yunnanensis, Pinus kesiya var. langbianensis, Pinus kesiya, geographical variation, needle, anatomical characters, temperature, precipitation

李卫英, 章正仁, 辛雅萱, 王飞, 辛培尧, 高洁. 云南松、思茅松和卡西亚松天然种群间的针叶表型变异. 植物生态学报, 2023, 47(6): 833-846. DOI: 10.17521/cjpe.2022.0263

LI Wei-Ying, ZHANG Zheng-Ren, XIN Ya-Xuan, WANG Fei, XIN Pei-Yao, GAO Jie. Needle phenotype variation among natural populations of Pinus yunnanensis, P. kesiya var. langbianensis and P. kesiya. Chinese Journal of Plant Ecology, 2023, 47(6): 833-846. DOI: 10.17521/cjpe.2022.0263

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https://www.plant-ecology.com/CN/Y2023/V47/I6/833

图/表 9

表1 云南松、卡西亚松及思茅松采集信息表

Table 1 Sample informations of Pinus yunnanensis, P. kesiya and P. kesiya var. langbianensis

物种 Species	种群地点 Population location	种群编号 Population No.	经度 Longitude (° E)	纬度 Latitude (° N)	海拔 Altitude (m)
卡西亚松 P. kesiya	老挝阿速坡 Attapu, Laos	ASPPk	107.19	15.22	1 061
	缅甸掸邦, 东枝 Dongzhi, Shan State, Myanmar	DZPk	97.06	20.79	1 583
	老挝川圹, 丰沙湾 Phonsavan, Xieng Khouang, Laos	FSWPk	103.12	19.50	1 070
	缅甸掸邦, 格劳 GeLao, Shan State, Myanmar	KLPk	96.55	20.65	1 022
	菲律宾 Philippines	PHP1	120.85	16.19	1 201
	菲律宾 Philippines	PHP4	120.89	17.14	751
	菲律宾 Philippines	PHP8	120.89	18.10	50
	老挝华潘, 桑怒 Sam Neua, Houaphanh, Laos	SNPk	103.96	20.22	1 399
	越南安沛, 拉潘滩 La Pan Tan, Yen Bai, Vietnam	V04	104.13	21.75	1 410
	越南得农 Dak Nong Vietnam	V08	107.43	12.15	847
思茅松 P. kesiya var. langbianensis	中国云南 Yunnan, China
	景谷 Jinggu	JGPk	100.50	23.49	1 707
	景洪贺建 Hejian, Jinghong	JHPk02	100.51	22.25	1 244
	勐腊 Mengla	MLPk	101.67	21.25	921
	宁洱 Ning’er	NEPk	100.97	23.19	1 004
	思茅 Simao	SMPk	100.95	22.71	1 340
	景洪大渡岗 Dadugang, Jinghong	JHPk01	100.99	22.42	1 113
	勐海布朗山 Bulangshan, Menghai	MHPk01	100.24	21.66	885
	墨江 Mojiang	MJPk	101.61	23.40	1 690
	镇沅 Zhengyuan	ZYPk	101.14	23.88	1 109
云南松 P. yunnanensis	中国云南 Yunnan, China
	宾川 Binchuan	BCPy	100.35	25.97	1 809
	大理云龙 Yunlong, Dali	DLPy	99.29	25.87	2 572
	景东 Jingdong	JDPy	100.67	24.70	1 713
	昆明 Kunming	KMPy	102.62	24.91	2 041
	丽江 Lijiang	LJPy	100.22	26.88	2 651
	腾冲 Tengchong	TCPy	98.58	24.91	1 776
	文山 Wenshan	WSPy02	104.63	23.97	1 573
	宜良 Yiliang	YLPy	103.16	24.71	1 938
	玉溪 Yuxi	YXPy	102.15	24.24	1 861
	中国广西 Guangxi, China
	乐业 Leye	LYPy	106.37	24.80	1 336
	西林 Xilin	XLPy	104.62	24.59	857
	隆林 Longlin	LLPy	104.96	24.69	1 294

表1 云南松、卡西亚松及思茅松采集信息表

Table 1 Sample informations of Pinus yunnanensis, P. kesiya and P. kesiya var. langbianensis

物种 Species	种群地点 Population location	种群编号 Population No.	经度 Longitude (° E)	纬度 Latitude (° N)	海拔 Altitude (m)
卡西亚松 P. kesiya	老挝阿速坡 Attapu, Laos	ASPPk	107.19	15.22	1 061
	缅甸掸邦, 东枝 Dongzhi, Shan State, Myanmar	DZPk	97.06	20.79	1 583
	老挝川圹, 丰沙湾 Phonsavan, Xieng Khouang, Laos	FSWPk	103.12	19.50	1 070
	缅甸掸邦, 格劳 GeLao, Shan State, Myanmar	KLPk	96.55	20.65	1 022
	菲律宾 Philippines	PHP1	120.85	16.19	1 201
	菲律宾 Philippines	PHP4	120.89	17.14	751
	菲律宾 Philippines	PHP8	120.89	18.10	50
	老挝华潘, 桑怒 Sam Neua, Houaphanh, Laos	SNPk	103.96	20.22	1 399
	越南安沛, 拉潘滩 La Pan Tan, Yen Bai, Vietnam	V04	104.13	21.75	1 410
	越南得农 Dak Nong Vietnam	V08	107.43	12.15	847
思茅松 P. kesiya var. langbianensis	中国云南 Yunnan, China
	景谷 Jinggu	JGPk	100.50	23.49	1 707
	景洪贺建 Hejian, Jinghong	JHPk02	100.51	22.25	1 244
	勐腊 Mengla	MLPk	101.67	21.25	921
	宁洱 Ning’er	NEPk	100.97	23.19	1 004
	思茅 Simao	SMPk	100.95	22.71	1 340
	景洪大渡岗 Dadugang, Jinghong	JHPk01	100.99	22.42	1 113
	勐海布朗山 Bulangshan, Menghai	MHPk01	100.24	21.66	885
	墨江 Mojiang	MJPk	101.61	23.40	1 690
	镇沅 Zhengyuan	ZYPk	101.14	23.88	1 109
云南松 P. yunnanensis	中国云南 Yunnan, China
	宾川 Binchuan	BCPy	100.35	25.97	1 809
	大理云龙 Yunlong, Dali	DLPy	99.29	25.87	2 572
	景东 Jingdong	JDPy	100.67	24.70	1 713
	昆明 Kunming	KMPy	102.62	24.91	2 041
	丽江 Lijiang	LJPy	100.22	26.88	2 651
	腾冲 Tengchong	TCPy	98.58	24.91	1 776
	文山 Wenshan	WSPy02	104.63	23.97	1 573
	宜良 Yiliang	YLPy	103.16	24.71	1 938
	玉溪 Yuxi	YXPy	102.15	24.24	1 861
	中国广西 Guangxi, China
	乐业 Leye	LYPy	106.37	24.80	1 336
	西林 Xilin	XLPy	104.62	24.59	857
	隆林 Longlin	LLPy	104.96	24.69	1 294

图1 云南松、卡西亚松和思茅松31个种群的地理分布。

Fig. 1 Geographic locations of 31 populations from Pinus yunnanensis, P. kesiya and P. kesiya var. langbianensis.

表2 云南松、思茅松和卡西亚松天然种群针叶的表型性状、变异系数和种间Tukey’s HSD多重比较

Table 2 Measurement values, variation coefficients and Tukey’s HSD test of the needle morphological traits among the natural populations of Pinus yunnanensis, P. kesiya var. langbianensis and P. kesiya

性状 Trait	云南松 Pinus yunnanensis		卡西亚松 P. kesiya		思茅松 P. kesiya var. langbianensis		种群间 Among populations
性状 Trait	平均值±标准差 Mean ± SD	变异系数 CV (%)	平均值±标准差 Mean ± SD	变异系数 CV (%)	平均值±标准差 Mean ± SD	变异系数 CV (%)	平均值±标准差 Mean ± SD	变异系数 CV (%)
LSGC (μm)	30.20 ± 3.41^b	11.30	35.45 ± 3.83^a	10.80	31.41 ± 3.86^b	12.28	32.28 ± 4.32	13.39
WSGC (μm)	24.70 ± 2.78^b	11.26	27.32 ± 3.17^a	11.62	24.94 ± 2.53^ab	10.13	25.63 ± 3.08	12.01
WTLL (μm)	16.94 ± 2.62^b	15.50	22.66 ± 4.64^a	20.49	18.31 ± 2.69^b	14.68	19.21 ± 4.23	22.01
WTLW (μm)	13.84 ± 2.20^c	15.77	17.65 ± 2.00^a	11.35	15.58 ± 2.18^b	14.01	15.63 ± 2.64	16.87
LSC (μm)	6.99 ± 2.17^a	31.01	8.07 ± 2.96^a	36.70	6.57 ± 1.90^a	28.84	7.23 ± 2.46	34.08
WSC (μm)	5.88 ± 1.45^b	24.56	6.95 ± 1.82^a	26.12	5.67 ± 1.74^b	30.70	6.17 ± 1.74	28.26
NL (cm)	21.34 ± 3.57^a	16.72	18.43 ± 2.87^ab	15.59	14.93 ± 5.45^b	36.53	18.53 ± 4.78	25.78
SD (600 μm^-2)	98.43 ± 25.28^a	25.68	68.92 ± 16.11^b	23.38	66.87 ± 17.41^b	26.04	79.62 ± 25.21	31.66

表3 云南松、思茅松和卡西亚松天然种群针叶表型性状变异方差分析

Table 3 Variance portions and differentiation coefficients of the needle morphological traits among the natural populations of Pinus yunnanensis, P. kesiya var. langbianensis and P. kesiya

性状 Trait	均方 Mean square			F		方差分量 Variance components (%)			V_ST (%)
性状 Trait	种间 Interspecies	种内群体间 Populations within species	残差 Residuals	种间 Interspecies	种内群体间 Populations within species	种间 Interspecies	种内群体间 Populations within species	残差 Residuals	V_ST (%)
LSGC (μm)	770.50	43.80	10.40	17.59^***	4.21^***	34.66	16.40	48.94	67.88
WSGC (μm)	209.53	31.04	5.71	6.75^***	5.44^***	17.78	25.97	56.25	40.64
WTLL (μm)	914.30	50.20	7.80	18.21^***	6.44^***	41.77	21.06	37.17	66.48
WTLW (μm)	363.60	9.30	4.00	39.10^***	2.33^***	43.89	6.67	49.44	86.80
LSC (μm)	56.49	21.60	4.06	2.62^***	5.32^***	5.63	29.28	65.08	16.14
WSC (μm)	45.31	6.77	2.33	6.69^***	2.91^***	12.24	14.50	73.26	45.79
NL (cm)	1 005.90	61.00	11.50	16.49^***	5.30^***	36.54	19.68	43.78	64.99
SD (600 μm^-2)	32 767.00	1 112.00	345.00	29.47^***	3.22^***	43.03	10.72	46.25	80.06
平均值 Mean	4 516.58	166.96	48.85	-	-	29.44	18.04	52.52	58.60

图2 云南松、思茅松和卡西亚松针叶性状指标的主成分(PC)分析(A)和31个天然群体的针叶性状聚类图(B)。LSC, 气孔腔长度; LSGC, 气孔保卫细胞长度; NL, 针叶长度; SD, 气孔密度; WSC, 气孔腔宽度; WSGC, 气孔保卫细胞宽度; WTLL, 木质增厚层长度; WTLW, 木质增厚层宽度。群体编号信息见表1。

Fig. 2 Scatter diagram of the first two principal coordinates from principal component (PC) analysis (A) and ward cluster (B) for the 31 populations based on needle traits of Pinus yunnanensis, P. kesiya var. langbianensis and P. kesiya. LSC, length of the stomata cavity; LSGC, length of stomata guard cell; NL, needle length; SD, stomatal density; WSC, width of the stomata cavity; WSGC, width of the stomata guard cell; WTLL, length of the woody thickening layer; WTLW, width of the woody thickening layer. See Table 1 for abbreviations of population ID.

图3 云南松、思茅松和卡西亚松针叶性状与纬度梯度相关性分析。LSC, 气孔腔长度; LSGC, 气孔保卫细胞长度; NL, 针叶长度; SD, 气孔密度; WSC, 气孔腔宽度; WSGC, 气孔保卫细胞宽度; WTLL, 木质增厚层长度; WTLW, 木质增厚层宽度。

Fig. 3 Latitudinal gradients of the needle traits of Pinus yunnanensis, P. kesiya var. langbianensis and P. kesiya. LSC, length of the stomata cavity; LSGC, length of stomata guard cell; NL, needle length; SD, stomatal density; WSC, width of the stomata cavity; WSGC, width of the stomata guard cell; WTLL, length of the woody thickening layer; WTLW, width of the woody thickening layer.

图4 多元线性模型筛选出的重要气候变量与云南松、思茅松和卡西亚松针叶性状的相关性。LSC, 气孔腔长度; LSGC, 气孔保卫细胞长度; NL, 针叶长度; SD, 气孔密度; WSC, 气孔腔宽度; WSGC, 气孔保卫细胞宽度; WTLL, 木质增厚层长度; WTLW, 木质增厚层宽度。BIO2, 平均气温日较差; BIO9, 最干季平均气温; BIO13, 最湿月降水量; BIO14, 最干月降水量; BIO15, 降水量季节变异系数。r, 相关系数。所有值为基于lg转换。

Fig. 4 Significant correlation between the needle traits and climatic variables (selected by the multiple linear regression model) of Pinus yunnanensis, P. kesiya var. langbianensis and P. kesiya. LSC, length of the stomata cavity; LSGC, length of stomata guard cell; NL, needle length; SD, stomatal density; WSC, width of the stomata cavity; WSGC, width of the stomata guard cell; WTLL, length of the woody thickening layer; WTLW, width of the woody thickening layer. BIO2, mean temperature diurnal range; BIO9, mean temperature of driest quarter; BIO13, precipitation of wettest month; BIO14, precipitation of driest month; BIO15, precipitation seasonality. r, correlation coefficient. The values was lg transformed.

图5 云南松、思茅松和卡西亚松的针叶表型性状主成分分析中第一主成分的群体分值(PC1)随重要气候变量(多元线性模型筛选)的地理变异趋势。BIO2, 平均气温日较差; BIO9, 最干季平均气温; BIO13, 最湿月降水量; BIO14, 最干月降水量; BIO15, 降水量季节变异系数。所有值为基于lg转换后扩大100倍。

Fig. 5 Distribution patterns of the population score of the first principal component value (PC1) based on the analysis of conifer phenotypic traits of Pinus yunnanensis, P. kesiya var. langbianensis and P. kesiya was the geographical variation trend of important climatic variables (multivariate linear model screening). BIO2, mean temperature diurnal range; BIO9, mean temperature of driest quarter; BIO13, precipitation of wettest month; BIO14, precipitation of driest month; BIO15, precipitation seasonality. The values was lg transformed and multiplied by 100.

表4 多元线性回归模型筛选的影响云南松、思茅松和卡西亚松种群针叶表型性状变异的气候变量

Table 4 Multiple linear regression models selected the climatic variables that predict variation in measured traits across all populations of Pinus yunnanensis, P. kesiya var. langbianensis and P. kesiya.

性状 Trait	截距 Intercept	平均气温日较差 BIO2	最湿季平均气温 BIO8	最干季平均气温 BIO9	最湿月降水量 BIO13	最干月降水量 BIO14	降水量季节变异系数 BIO15
LSGC (μm)	-0.000 8	0.1 759			0.7 516		-0.3 924
WSGC (μm)	-0.054 6		-0.5 432		0.6 846	0.3 786
WTLL (μm)	0.007 7			0.2 861	0.6 523		-0.4 449
WTLW (μm)	-0.008 3		-0.3 561	0.9 913		0.1 731
LSC (μm)	-0.031 4	-0.4 884			0.7 315	0.7 785	-1.3 658
WSC (μm)	-0.027 5				0.4 550		-0.3 918
NL (cm)	-0.007 0	-0.3 137		-0.5 250
SD ( 600 μm^-2)	-0.018 1			-0.8 011		0.1 918

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云南松、思茅松和卡西亚松天然种群间的针叶表型变异

Needle phenotype variation among natural populations of Pinus yunnanensis, P. kesiya var. langbianensis and P. kesiya

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