东亚地区栓皮栎（Quercus variabilis）叶片性状的变异格局及其对环境变化的响应
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摘要
植物叶片的性状（例如，叶形态、叶脉密度、气孔大小和密度、表皮毛类型和密度等）是植物对环境因子长期适应的结果，与植物叶片的蒸腾及光合作用两大生理过程密切相关。对不同植物叶片性状及其特点，已有大量研究报导。但是，在区域尺度上，对植物叶片性状沿气候梯度的变异格局和主要控制因子以及叶片性状对气候变化的响应等，还缺乏系统的研究和综合分析。这些方面研究是植物生理生态学的重要内容，特别是对了解气候变化条件下植物响应、群落构成变化、植物迁移等具有重要的理论意义，研究结果对在气候变化形势下合理地开展森林经营具有指导作用。
栓皮栎（Quercus variabilis）是东亚地区分布最广泛的落叶阔叶树种之一（19–42°N、97–140°E，海拔50–2000m）。在中国、韩国和日本，栓皮栎林分均都具有重要经济价值（木材、栓皮、果实、叶片等），发挥着生物多样性和环境保护作用，形成了重要森林景观。由于栓皮栎具有较强的抗旱特性，在我国华北和西北的干旱半干旱地区，是重要的荒山造林树种。栓皮栎分布区域较大，分布区内环境复杂，不同地区的种群经过长期的自然选择和种群内的遗传分化，形成了遗传结构不同的地理种群。因而，栓皮栎是在种的水平上，研究植物叶片性状变异格局及其与环境因子关系的理想树种。
在本项研究中，我们采集了44个野外种群林分样地（包括中国大陆、台湾、舟山、韩国和日本等岛屿）的叶片样品，并获得了各样点气候数据。应用光学显微镜和扫描电镜等技术和方法，观察测量了叶片大小、比叶面积、叶柄长度、各级叶脉密度、气孔大小和密度、表皮毛密度等指标，分析了这些叶片性状的地理变异格局及其与环境因子间的关系。同时，我们在2008年采集了野外15个栓皮栎种群种子，在栓皮栎分布区中部的上海地区建立了栓皮栎苗木同质园（common garden），通过野外和同质园样品数据对比，研究了叶片性状对环境因子变化的响应。主要研究结果如下：
1、在东亚地区分布范围内，栓皮栎不同种群间的叶形态性状（叶片长度、叶片宽度及叶片长宽比值等）存在显著差异（P <0.001），变异格局与环境变量显著相关。在野外条件下，栓皮栎叶片的长宽比值随纬度升高显著降低（r~2=0.134, P=0.015），叶片长度（r~2=0.132, P=0.016）、叶片宽度（r~2=0.109, P=0.028）、叶柄长（r~2=0.289, P <0.001）随样地的经度升高而显著降低。野外栓皮栎种群的叶片的长宽比值与年平均温度（MAT）呈显著正相关关系（r~2=0.135,P=0.014），叶片长度（r~2=0.207, P=0.002）及叶片的长宽比值（r~2=0.278, P <0.001）与MDSH极显著负相关。栓皮栎的叶长（r~2=0.131, P=0.033）及叶片的长宽比值（r~2=0.235, P=0.003）与土壤中的K含量显著负相关。多元回归分析结果表明，环境因子中影响叶片形状的首要因子是MDSH，其次为MAT。在同质园中生长两年后，除了叶柄长度外，不同地理种群间栓皮栎的幼苗其余叶形态性状的差异缩小，其中叶长（P=0.160）与叶面积（P=0.105）在不同地理种群间无显著差异。同质园中叶形态特性与原产地经度、纬度均不相关（P=0.081–0.949）。栓皮栎叶形态性状对环境因子敏感，野外种群间的栓皮栎叶形态变异主要是由于各样地间的环境差异，由环境引起的栓皮栎叶形态的变异的可遗传性较低。
2、在分布区范围内，野外栓皮栎种群的叶片细脉密度随原产地的纬度升高而极显著减小（r~2=0.63, P <0.001）；而且，在同质园生长条件下，不同种群的栓皮栎叶脉密度与原产地纬度仍然呈现与野外一致的变异格局（r~2=0.63, P <0.001）。也就是说，栓皮栎叶片的细脉密度是一种遗传性状（genotypic trait）。野外与同质园样地的栓皮栎叶片细脉密度的年平均降水量（MAP）显著正相关（野外r~2=0.30, P <0.001；同质园r~2=0.29, P=0.04），与年平均温度（MAT）显著正相关（野外r~2=0.53, P <0.001；同质园r~2=0.35, P=0.02）。野外叶片细脉密度与样地平均日照时数（MDSH）极显著负相关（r~2=0.24, P=0.001），而同质园叶片细脉密度与MDSH不具显著相关性（r~2=0.23, P=0.07）。野外栓皮栎种群的叶片细脉密度与叶长（r~2=0.12, P=0.02）和叶片长宽比（r~2=0.36, P <0.001）显著正相关，而同质园样地的栓皮栎细脉密度与叶长表现为显著负相关（r~2=0.44, P=0.007）。此外，野外栓皮栎种群的细脉密度与土壤中K、Ca含量显著负相关（r~2=0.16, P=0.02和r~2=0.21, P=0.007）。通径分析结果表明，影响栓皮栎叶片细脉密度的主要因子为MAT及叶片形状，而MAP与MDSH对栓皮栎细脉密度具有较大的间接影响。栓皮栎叶片的叶脉密度是对当地气候因子长期适应的结果，叶脉密度是基因型性状对于短期生长环境条件的改变不是非常敏感。
3、在栓皮栎分布区尺度上，栓皮栎叶片气孔密度（SD）及叶片气孔面积指数（SOI）随样地经度升高而显著变小（气孔密度, r~2=0.098, P=0.039; SOI, r~2=0.120, P=0.021），而对于同质园中15个不同地理种群栓皮栎来说，气孔密度及SOI都不存在显著差异，并与种群地的纬度不相关（P=0.278–0.540）。野外种群的叶片气孔密度与样地的年平均降水量（MAP）呈现极显著负相关（r~2=0.157,P=0.008），而与其它气候因子不具显著相关性（P=0.321–0.740）。与气孔密度相比，栓皮栎叶片的SOI对环境因子更为敏感。野外种群的SOI与年平均温度（MAT）（r~2=0.12, P=0.02）和MAP显著负相关（r~2=0.39, P <0.001），而与平均月太阳辐射量（MMSR）显著正相关（r~2=0.16, P=0.011）。同质园中的栓皮栎种群的气孔密度和SOI与种群地的气候因子都不相关（P=0.101–0.775）。野外栓皮栎种群的叶片的气孔密度与气孔长度（r~2=0.236, P=0.001）及气孔长宽比值（r~2=0.266, P <0.001）负相关。野外栓皮栎种群的叶片气孔密度和SOI都与单位叶面积的叶片干重（LMA）及叶柄长度显著正相关（r~2=0.199–0.299, P=<0.001–0.002），而同质园样地的气孔密度和SOI与叶片LMA及叶柄长度不相关（P=0.056–0.568）。野外种群的SOI与土壤中Ca（r~2=0.18, P=0.012）及K（r~2=0.15, P=0.019）元素含量显著正相关。野外栓皮栎种群的气孔Rp值与纬度极显著正相关（r~2=0.210, P=0.002），同质园中栓皮栎气孔Rp值与种群地纬度不相关（r~2=0.006, P=0.781）。野外种群的气孔Rp值与样地的年平均降水量（MAP）极显著负相关（r~2=0.160, P=0.007），与气孔长度及宽度显著正相关（r~2=0.231–0.235, P=0.001）。多元回归分析结果表明，栓皮栎叶片的气孔密度与SOI的主要影响因子为气孔大小、LMA及MAP，叶片气孔Rp值的主要影响因子为气孔长度与MAP。本研究结果表明栓皮栎叶片的气孔密度及SOI对于自身生长的生态环境具有长期的基因上的适应性，并对于适应短期的环境的改变具有较高的可塑性，而栓皮栎叶片的气孔Rp值相对较稳定，环境因子对栓皮栎气孔R值的影响并未改变栓皮栎气孔的非随机分布格局。
4、在栓皮栎分布区尺度上，44个样地的栓皮栎叶片表皮毛密度间差异极显著（P <0.001），而在同质园中，15个不同地理种群间的栓皮栎叶片的表皮毛密度无显著差异（P=0.521）。野外及同质园栓皮栎叶片的表皮毛密度不存在明显的变异格局，与样地纬度、经度及海拔均不相关（P=0.107–0.650）。野外栓皮栎种群的叶片表皮毛密度与样地MAP极显著负相关（r~2=0.159, P=0.007），与MMSR在临界水平上显著正相关（r~2=0.095, P=0.053），与其他气候因子不相关（P=0.074–0.613）。同质园不同地理种群的栓皮栎叶片表皮毛密度与原产地气候因子均不相关（P=0.404–0.985）。野外栓皮栎种群的叶片的表皮毛密度与叶片的叶柄长度（r~2=0.145, P=0.011）及LMA（r~2=0.235, P=0.001）均显著正相关，与叶片的气孔密度呈极显著正相关关系（r~2=0.510, P <0.001）。对栓皮栎叶片表皮毛密度而言，主要影响因子为叶片LMA与MAP。本文研究结果表明，栓皮栎叶片的表皮毛与叶片的气孔可能存在着协同演化关系。栓皮栎叶片的表皮毛密度对环境变化高度敏感，具有较强的表现型可塑性。
For plants, the leaf anatomical traits (leaf morphology traits, vein density, stomata sizeand density, trichomes types and density) were closely related with transpiration andphotosynthesis. They are the results of adaption for environmental factors during long-term evolution. There were many reports about leaf traits and characters. In regionalscales, yet there is no general conclusion about the relationships between leaf traits, thevariation pattern along a climate gradient, the main factors that drive the variations andhow do leaf traits respond to climate changes. They were important in ecology researchfield, and had important theoretical significance in understanding how plant respondedto climatic changes, community composition changes as well as plant migration. Theywould guide the resonable forest management under climatic change conditions.
Orienatal oak (Quercus variabilis) was the most wide distributioned tree species ineastern Asia (19–42°N、97–140°E，50–2000m a.s.l). Oriental oak had importanteconomic value (timber, cork, fruits and so on), biological diversity, environmentalprotection, urban and rural landscape function et al. The natural environment oforiental oak distributed sites were complex due to the large distribution area, and theoriental oak growing in different sites formed geographical populations with differentgenetic composition. This made oriental oak be an ideal plant species for investigatingthe relationship between leaf traits and environmental factors.
We examined both the variation in leaf size, LMA, leaf petiole length, leaf veindensity, stomatal size and density, leaf trichome density and other leaf traits of orientaloak (Quercus variabilis) from44in situ Populations include the mainland of China,Taiwan Island, Zhoushan Archipelago, Korea Peninsula and Japan Archipelago across temperate-subtropical biomes. We investigated the variation patterns of leaf traits andinvestigated the relationships between leaf traits and environmental factors. Theresponse of leaf traits to environmental changes were also examined by commongarden experiment with seedlings of15populations from different provenances grownin a common garden. The main results were as follows:
1) Leaf morphogy traits of oriental oak significantly differed among different insitu populations (P <0.001), and the differences among common garden populationsreduced after two years growing in common garden except for leaf petiole length.Particularly, leaf length（P=0.160） and leaf area （P=0.105）had no significantdifference among common garden populations. Leaf length to width was negativelycorrelated with latitude for in situ populations (r~2=0.134, P=0.015). Leaf length,width and petiole length showed negative correlations with longitude for in situpopulations (r~2=0.132, P=0.016for leaf length; r~2=0.109, P=0.028for leaf width;r~2=0.289, P <0.001for leaf petiole length). While leaf morphology traits wereindependent of latitude and longitude of origins (P=0.081–0.949). Leaf morphologytraits were invariant with MAP (P=0.070–0.511). Leaf length to width rationegatively correlated with MAT (r~2=0.135, P=0.014). Leaf length and leaf length towidth ratio were significantly negatively correlated with MDSH (r~2=0.207, P=0.002for leaf length and r~2=0.278, P <0.001for leaf length to width ratio). Additionally,leaf length and leaf length to width were negatively correlated with soil Kconcentrations (r~2=0.131, P=0.033for leaf length and r~2=0.235, P=0.003for leaflength to width). We concluded that the main factor that affected leaf morphology wasMDSH, secondly was MAT by multiple regression analysis. Leaf morphology traits oforiental oak were sensitive to climatic factors. The variances in field observed in thisstudy were mainly due to environmental differences among different sampling sites.The heritability of leaf morphology traits variations caused by environment was low.
2) Minor vein density (≥3rdorder) of oriental oak decreased significantly with theincreasing latitude (r~2=0.63and P <0.001) for the in situ populations, and this patternremained unchanged for the garden-Populations (r~2=0.63and P <0.001). Minor veindensity both positively correlated with mean annual precipitation (MAP) and mean annual temperature (MAT) of the origins for both the garden-Populations (r~2=0.29, P=0.04and r~2=0.35, P=0.02, respectively) and the in situ Populations (r~2=0.30, P <0.001and r~2=0.53, P <0.001, respectively). Leaf vein density significantly positivelycorrelated with mean daily sunshine hours (MDSH) of the origins for the in situpopulations (r~2=0.24, P=0.001), but were invariant for the common gardenpopulations (r~2=0.23, P=0.07). Minor vein density significantly increased with leaflength and leaf length to width for in situ populations (r~2=0.12, P=0.02and r~2=0.36,P <0.001, respectively). Leaf minor vein density were negatively correlated with leaflength (r~2=0.44, P=0.007). Additionally, leaf minor vein density showed a negativelyrelationships with soil K and Ca concentrations (r~2=0.16, P=0.02and r~2=0.21, P=0.007, respectively). MAT and leaf shape were the main factors that had directed effecton leaf minor vein density of oriental oak through Path Coefficient analysis, whileMAP and MDSH had strong in-directed effects. These results implied that leaf veindensity was a genotypic trait for adapting for local climatic factors, and this trait did notsensitively respond to temporal change in growing conditions.
3) Stomatal density and SOI of in situ populations decreased with longitude (r~2=0.098, P=0.039of stomata density; r~2=0.120, P=0.021of SOI), while stomataldensity and SOI were independent of longitude, and have no significant differencesamong15garden populations (P=0.278–0.540). Stomatal density was significantlynegatively correlated with MAP of the origins (r~2=0.157, P=0.008), but wereinvariant with other climatic factors for the in situ populations (P=0.321–0.740).Compared with stomatal density, leaf SOI was more sensitive to climatic factors. SOIshowed a negative relationship with MAT (r~2=0.12, P=0.02) and MAP (r~2=0.39, P <0.001), and a positive correlation with mean monthly solar radiation (MMSR) of theorigins for the in situ populations (r~2=0.16, P=0.011). For the common gardenpopulations, both stomatal density and SOI were invariant with climatic factors oforigins (P=0.101–0.775). Stomatal density was negatively correlated with stomatal length (r~2=0.236, P=0.001) and stomatal width-to-length (r~2=0.266, P <0.001)across in situ populations. Both of stomatal density and SOI were positively correlatedwith leaf dry mass per area (LMA) and leaf petiole length (r~2=0.199–0.299, P=<0.001–0.002) for the in situ populations, while no significant correlation for thegarden populations (P=0.056–0.568). SOI was positively correlated with soil Ca (r~2=0.18, P=0.012) and K concentrations for the in situ populations (r~2=0.15, P=0.019). Stomatal Rpvalue was significantly positively correlated with latitude for the insitu populations (r~2=0.210, P=0.002), while showed no correlations for the commongarden populations (r~2=0.006, P=0.781). Stomatal Rpvalue were negativelycorrelated with MAP of the origins (r~2=0.160, P=0.007), positively correlated withstomatal length and width for the in situ populations (r~2=0.231–0.235, P=0.001).The main factors that affected stomatal density and SOI of oriental oak were stomatalsize, LMA and MAP by multiple regression analysis. The main factors influencingstomatal Rpvalue were stomatal length and MAP. The present results implied thatstomatal density of oriental oak were genetically adaptive to long-term ecologicalenvironments and had high flexibilities in response to temporary climatic changes. Thestomatal Rpvalue was relatively stable, environmental factors influenced the stomatalRpvalue in some extent, but did not change the non-random distribution pattern ofstomata.
4) Leaf trichome density of oriental oak had significant differences among in situpopulations (P <0.001), while had no significant differences among15gardenpopulations (P=0.521). Trichome density was invariant with latitude, longitude andaltitude of the origins for both in situ and common garden populations (P=0.107– 0.650). Leaf trichome density was significantly negatively correlated with MAP (r~2=0.159, P=0.007), positively correlated with MMSR (r~2=0.095, P=0.053) and wasinvariant with other climatic factors for in situ populations (P=0.074–0.613).Trichome density was independent of climatic factors of origins for common gardenpopulations (P=0.404–0.985). Leaf trichome density was significantly positivelycorrelated with leaf petiole length (r~2=0.145, P=0.0110) and LMA (r~2=0.235, P=0.001), and significantly positively correlated with stomatal density (r~2=0.510, P <0.001) for in situ populations. Leaf LMA and MAP were the main factors that affectedleaf trichome density of oriental oak. The present study suggested that the leaf trichomeand stomata may be two leaf traits that have coevolution relationships during long-termenvironmental adaptation. Leaf trichome density of oriental oak was very sensitive toenvironmental changes and had strong phenotypic plasticity.
栓皮栎（Quercus variabilis）是东亚地区分布最广泛的落叶阔叶树种之一（19–42°N、97–140°E，海拔50–2000m）。在中国、韩国和日本，栓皮栎林分均都具有重要经济价值（木材、栓皮、果实、叶片等），发挥着生物多样性和环境保护作用，形成了重要森林景观。由于栓皮栎具有较强的抗旱特性，在我国华北和西北的干旱半干旱地区，是重要的荒山造林树种。栓皮栎分布区域较大，分布区内环境复杂，不同地区的种群经过长期的自然选择和种群内的遗传分化，形成了遗传结构不同的地理种群。因而，栓皮栎是在种的水平上，研究植物叶片性状变异格局及其与环境因子关系的理想树种。
在本项研究中，我们采集了44个野外种群林分样地（包括中国大陆、台湾、舟山、韩国和日本等岛屿）的叶片样品，并获得了各样点气候数据。应用光学显微镜和扫描电镜等技术和方法，观察测量了叶片大小、比叶面积、叶柄长度、各级叶脉密度、气孔大小和密度、表皮毛密度等指标，分析了这些叶片性状的地理变异格局及其与环境因子间的关系。同时，我们在2008年采集了野外15个栓皮栎种群种子，在栓皮栎分布区中部的上海地区建立了栓皮栎苗木同质园（common garden），通过野外和同质园样品数据对比，研究了叶片性状对环境因子变化的响应。主要研究结果如下：
1、在东亚地区分布范围内，栓皮栎不同种群间的叶形态性状（叶片长度、叶片宽度及叶片长宽比值等）存在显著差异（P <0.001），变异格局与环境变量显著相关。在野外条件下，栓皮栎叶片的长宽比值随纬度升高显著降低（r~2=0.134, P=0.015），叶片长度（r~2=0.132, P=0.016）、叶片宽度（r~2=0.109, P=0.028）、叶柄长（r~2=0.289, P <0.001）随样地的经度升高而显著降低。野外栓皮栎种群的叶片的长宽比值与年平均温度（MAT）呈显著正相关关系（r~2=0.135,P=0.014），叶片长度（r~2=0.207, P=0.002）及叶片的长宽比值（r~2=0.278, P <0.001）与MDSH极显著负相关。栓皮栎的叶长（r~2=0.131, P=0.033）及叶片的长宽比值（r~2=0.235, P=0.003）与土壤中的K含量显著负相关。多元回归分析结果表明，环境因子中影响叶片形状的首要因子是MDSH，其次为MAT。在同质园中生长两年后，除了叶柄长度外，不同地理种群间栓皮栎的幼苗其余叶形态性状的差异缩小，其中叶长（P=0.160）与叶面积（P=0.105）在不同地理种群间无显著差异。同质园中叶形态特性与原产地经度、纬度均不相关（P=0.081–0.949）。栓皮栎叶形态性状对环境因子敏感，野外种群间的栓皮栎叶形态变异主要是由于各样地间的环境差异，由环境引起的栓皮栎叶形态的变异的可遗传性较低。
2、在分布区范围内，野外栓皮栎种群的叶片细脉密度随原产地的纬度升高而极显著减小（r~2=0.63, P <0.001）；而且，在同质园生长条件下，不同种群的栓皮栎叶脉密度与原产地纬度仍然呈现与野外一致的变异格局（r~2=0.63, P <0.001）。也就是说，栓皮栎叶片的细脉密度是一种遗传性状（genotypic trait）。野外与同质园样地的栓皮栎叶片细脉密度的年平均降水量（MAP）显著正相关（野外r~2=0.30, P <0.001；同质园r~2=0.29, P=0.04），与年平均温度（MAT）显著正相关（野外r~2=0.53, P <0.001；同质园r~2=0.35, P=0.02）。野外叶片细脉密度与样地平均日照时数（MDSH）极显著负相关（r~2=0.24, P=0.001），而同质园叶片细脉密度与MDSH不具显著相关性（r~2=0.23, P=0.07）。野外栓皮栎种群的叶片细脉密度与叶长（r~2=0.12, P=0.02）和叶片长宽比（r~2=0.36, P <0.001）显著正相关，而同质园样地的栓皮栎细脉密度与叶长表现为显著负相关（r~2=0.44, P=0.007）。此外，野外栓皮栎种群的细脉密度与土壤中K、Ca含量显著负相关（r~2=0.16, P=0.02和r~2=0.21, P=0.007）。通径分析结果表明，影响栓皮栎叶片细脉密度的主要因子为MAT及叶片形状，而MAP与MDSH对栓皮栎细脉密度具有较大的间接影响。栓皮栎叶片的叶脉密度是对当地气候因子长期适应的结果，叶脉密度是基因型性状对于短期生长环境条件的改变不是非常敏感。
3、在栓皮栎分布区尺度上，栓皮栎叶片气孔密度（SD）及叶片气孔面积指数（SOI）随样地经度升高而显著变小（气孔密度, r~2=0.098, P=0.039; SOI, r~2=0.120, P=0.021），而对于同质园中15个不同地理种群栓皮栎来说，气孔密度及SOI都不存在显著差异，并与种群地的纬度不相关（P=0.278–0.540）。野外种群的叶片气孔密度与样地的年平均降水量（MAP）呈现极显著负相关（r~2=0.157,P=0.008），而与其它气候因子不具显著相关性（P=0.321–0.740）。与气孔密度相比，栓皮栎叶片的SOI对环境因子更为敏感。野外种群的SOI与年平均温度（MAT）（r~2=0.12, P=0.02）和MAP显著负相关（r~2=0.39, P <0.001），而与平均月太阳辐射量（MMSR）显著正相关（r~2=0.16, P=0.011）。同质园中的栓皮栎种群的气孔密度和SOI与种群地的气候因子都不相关（P=0.101–0.775）。野外栓皮栎种群的叶片的气孔密度与气孔长度（r~2=0.236, P=0.001）及气孔长宽比值（r~2=0.266, P <0.001）负相关。野外栓皮栎种群的叶片气孔密度和SOI都与单位叶面积的叶片干重（LMA）及叶柄长度显著正相关（r~2=0.199–0.299, P=<0.001–0.002），而同质园样地的气孔密度和SOI与叶片LMA及叶柄长度不相关（P=0.056–0.568）。野外种群的SOI与土壤中Ca（r~2=0.18, P=0.012）及K（r~2=0.15, P=0.019）元素含量显著正相关。野外栓皮栎种群的气孔Rp值与纬度极显著正相关（r~2=0.210, P=0.002），同质园中栓皮栎气孔Rp值与种群地纬度不相关（r~2=0.006, P=0.781）。野外种群的气孔Rp值与样地的年平均降水量（MAP）极显著负相关（r~2=0.160, P=0.007），与气孔长度及宽度显著正相关（r~2=0.231–0.235, P=0.001）。多元回归分析结果表明，栓皮栎叶片的气孔密度与SOI的主要影响因子为气孔大小、LMA及MAP，叶片气孔Rp值的主要影响因子为气孔长度与MAP。本研究结果表明栓皮栎叶片的气孔密度及SOI对于自身生长的生态环境具有长期的基因上的适应性，并对于适应短期的环境的改变具有较高的可塑性，而栓皮栎叶片的气孔Rp值相对较稳定，环境因子对栓皮栎气孔R值的影响并未改变栓皮栎气孔的非随机分布格局。
4、在栓皮栎分布区尺度上，44个样地的栓皮栎叶片表皮毛密度间差异极显著（P <0.001），而在同质园中，15个不同地理种群间的栓皮栎叶片的表皮毛密度无显著差异（P=0.521）。野外及同质园栓皮栎叶片的表皮毛密度不存在明显的变异格局，与样地纬度、经度及海拔均不相关（P=0.107–0.650）。野外栓皮栎种群的叶片表皮毛密度与样地MAP极显著负相关（r~2=0.159, P=0.007），与MMSR在临界水平上显著正相关（r~2=0.095, P=0.053），与其他气候因子不相关（P=0.074–0.613）。同质园不同地理种群的栓皮栎叶片表皮毛密度与原产地气候因子均不相关（P=0.404–0.985）。野外栓皮栎种群的叶片的表皮毛密度与叶片的叶柄长度（r~2=0.145, P=0.011）及LMA（r~2=0.235, P=0.001）均显著正相关，与叶片的气孔密度呈极显著正相关关系（r~2=0.510, P <0.001）。对栓皮栎叶片表皮毛密度而言，主要影响因子为叶片LMA与MAP。本文研究结果表明，栓皮栎叶片的表皮毛与叶片的气孔可能存在着协同演化关系。栓皮栎叶片的表皮毛密度对环境变化高度敏感，具有较强的表现型可塑性。
For plants, the leaf anatomical traits (leaf morphology traits, vein density, stomata sizeand density, trichomes types and density) were closely related with transpiration andphotosynthesis. They are the results of adaption for environmental factors during long-term evolution. There were many reports about leaf traits and characters. In regionalscales, yet there is no general conclusion about the relationships between leaf traits, thevariation pattern along a climate gradient, the main factors that drive the variations andhow do leaf traits respond to climate changes. They were important in ecology researchfield, and had important theoretical significance in understanding how plant respondedto climatic changes, community composition changes as well as plant migration. Theywould guide the resonable forest management under climatic change conditions.
Orienatal oak (Quercus variabilis) was the most wide distributioned tree species ineastern Asia (19–42°N、97–140°E，50–2000m a.s.l). Oriental oak had importanteconomic value (timber, cork, fruits and so on), biological diversity, environmentalprotection, urban and rural landscape function et al. The natural environment oforiental oak distributed sites were complex due to the large distribution area, and theoriental oak growing in different sites formed geographical populations with differentgenetic composition. This made oriental oak be an ideal plant species for investigatingthe relationship between leaf traits and environmental factors.
We examined both the variation in leaf size, LMA, leaf petiole length, leaf veindensity, stomatal size and density, leaf trichome density and other leaf traits of orientaloak (Quercus variabilis) from44in situ Populations include the mainland of China,Taiwan Island, Zhoushan Archipelago, Korea Peninsula and Japan Archipelago across temperate-subtropical biomes. We investigated the variation patterns of leaf traits andinvestigated the relationships between leaf traits and environmental factors. Theresponse of leaf traits to environmental changes were also examined by commongarden experiment with seedlings of15populations from different provenances grownin a common garden. The main results were as follows:
1) Leaf morphogy traits of oriental oak significantly differed among different insitu populations (P <0.001), and the differences among common garden populationsreduced after two years growing in common garden except for leaf petiole length.Particularly, leaf length（P=0.160） and leaf area （P=0.105）had no significantdifference among common garden populations. Leaf length to width was negativelycorrelated with latitude for in situ populations (r~2=0.134, P=0.015). Leaf length,width and petiole length showed negative correlations with longitude for in situpopulations (r~2=0.132, P=0.016for leaf length; r~2=0.109, P=0.028for leaf width;r~2=0.289, P <0.001for leaf petiole length). While leaf morphology traits wereindependent of latitude and longitude of origins (P=0.081–0.949). Leaf morphologytraits were invariant with MAP (P=0.070–0.511). Leaf length to width rationegatively correlated with MAT (r~2=0.135, P=0.014). Leaf length and leaf length towidth ratio were significantly negatively correlated with MDSH (r~2=0.207, P=0.002for leaf length and r~2=0.278, P <0.001for leaf length to width ratio). Additionally,leaf length and leaf length to width were negatively correlated with soil Kconcentrations (r~2=0.131, P=0.033for leaf length and r~2=0.235, P=0.003for leaflength to width). We concluded that the main factor that affected leaf morphology wasMDSH, secondly was MAT by multiple regression analysis. Leaf morphology traits oforiental oak were sensitive to climatic factors. The variances in field observed in thisstudy were mainly due to environmental differences among different sampling sites.The heritability of leaf morphology traits variations caused by environment was low.
2) Minor vein density (≥3rdorder) of oriental oak decreased significantly with theincreasing latitude (r~2=0.63and P <0.001) for the in situ populations, and this patternremained unchanged for the garden-Populations (r~2=0.63and P <0.001). Minor veindensity both positively correlated with mean annual precipitation (MAP) and mean annual temperature (MAT) of the origins for both the garden-Populations (r~2=0.29, P=0.04and r~2=0.35, P=0.02, respectively) and the in situ Populations (r~2=0.30, P <0.001and r~2=0.53, P <0.001, respectively). Leaf vein density significantly positivelycorrelated with mean daily sunshine hours (MDSH) of the origins for the in situpopulations (r~2=0.24, P=0.001), but were invariant for the common gardenpopulations (r~2=0.23, P=0.07). Minor vein density significantly increased with leaflength and leaf length to width for in situ populations (r~2=0.12, P=0.02and r~2=0.36,P <0.001, respectively). Leaf minor vein density were negatively correlated with leaflength (r~2=0.44, P=0.007). Additionally, leaf minor vein density showed a negativelyrelationships with soil K and Ca concentrations (r~2=0.16, P=0.02and r~2=0.21, P=0.007, respectively). MAT and leaf shape were the main factors that had directed effecton leaf minor vein density of oriental oak through Path Coefficient analysis, whileMAP and MDSH had strong in-directed effects. These results implied that leaf veindensity was a genotypic trait for adapting for local climatic factors, and this trait did notsensitively respond to temporal change in growing conditions.
3) Stomatal density and SOI of in situ populations decreased with longitude (r~2=0.098, P=0.039of stomata density; r~2=0.120, P=0.021of SOI), while stomataldensity and SOI were independent of longitude, and have no significant differencesamong15garden populations (P=0.278–0.540). Stomatal density was significantlynegatively correlated with MAP of the origins (r~2=0.157, P=0.008), but wereinvariant with other climatic factors for the in situ populations (P=0.321–0.740).Compared with stomatal density, leaf SOI was more sensitive to climatic factors. SOIshowed a negative relationship with MAT (r~2=0.12, P=0.02) and MAP (r~2=0.39, P <0.001), and a positive correlation with mean monthly solar radiation (MMSR) of theorigins for the in situ populations (r~2=0.16, P=0.011). For the common gardenpopulations, both stomatal density and SOI were invariant with climatic factors oforigins (P=0.101–0.775). Stomatal density was negatively correlated with stomatal length (r~2=0.236, P=0.001) and stomatal width-to-length (r~2=0.266, P <0.001)across in situ populations. Both of stomatal density and SOI were positively correlatedwith leaf dry mass per area (LMA) and leaf petiole length (r~2=0.199–0.299, P=<0.001–0.002) for the in situ populations, while no significant correlation for thegarden populations (P=0.056–0.568). SOI was positively correlated with soil Ca (r~2=0.18, P=0.012) and K concentrations for the in situ populations (r~2=0.15, P=0.019). Stomatal Rpvalue was significantly positively correlated with latitude for the insitu populations (r~2=0.210, P=0.002), while showed no correlations for the commongarden populations (r~2=0.006, P=0.781). Stomatal Rpvalue were negativelycorrelated with MAP of the origins (r~2=0.160, P=0.007), positively correlated withstomatal length and width for the in situ populations (r~2=0.231–0.235, P=0.001).The main factors that affected stomatal density and SOI of oriental oak were stomatalsize, LMA and MAP by multiple regression analysis. The main factors influencingstomatal Rpvalue were stomatal length and MAP. The present results implied thatstomatal density of oriental oak were genetically adaptive to long-term ecologicalenvironments and had high flexibilities in response to temporary climatic changes. Thestomatal Rpvalue was relatively stable, environmental factors influenced the stomatalRpvalue in some extent, but did not change the non-random distribution pattern ofstomata.
4) Leaf trichome density of oriental oak had significant differences among in situpopulations (P <0.001), while had no significant differences among15gardenpopulations (P=0.521). Trichome density was invariant with latitude, longitude andaltitude of the origins for both in situ and common garden populations (P=0.107– 0.650). Leaf trichome density was significantly negatively correlated with MAP (r~2=0.159, P=0.007), positively correlated with MMSR (r~2=0.095, P=0.053) and wasinvariant with other climatic factors for in situ populations (P=0.074–0.613).Trichome density was independent of climatic factors of origins for common gardenpopulations (P=0.404–0.985). Leaf trichome density was significantly positivelycorrelated with leaf petiole length (r~2=0.145, P=0.0110) and LMA (r~2=0.235, P=0.001), and significantly positively correlated with stomatal density (r~2=0.510, P <0.001) for in situ populations. Leaf LMA and MAP were the main factors that affectedleaf trichome density of oriental oak. The present study suggested that the leaf trichomeand stomata may be two leaf traits that have coevolution relationships during long-termenvironmental adaptation. Leaf trichome density of oriental oak was very sensitive toenvironmental changes and had strong phenotypic plasticity.
引文
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