植物生态学报 ›› 2013, Vol. 37 ›› Issue (9): 793-802.DOI: 10.3724/SP.J.1258.2013.00083
• 研究论文 • 下一篇
收稿日期:
2013-03-06
接受日期:
2013-06-28
出版日期:
2013-03-06
发布日期:
2013-09-02
通讯作者:
史作民
作者简介:
*E-mail: shizm@caf.ac.cn基金资助:
LI Dong-Sheng1, SHI Zuo-Min1,*(), FENG Qiu-Hong2, LIU Feng3
Received:
2013-03-06
Accepted:
2013-06-28
Online:
2013-03-06
Published:
2013-09-02
Contact:
SHI Zuo-Min
摘要:
植物叶片的形态性状能够有效地反映生存环境的变化, 并且影响植物的基本行为和功能。该研究通过获取植物标本提供的叶片形态信息, 结合相关分析和标准化主轴分析, 探讨了南北样带暖温带区栎属(Quercus)树种叶片形态性状对气候条件的响应及适应策略。结果表明: 在南北样带暖温带区, 随着气候条件的变化, 栎属树种的叶片形态性状发生显著的变化。随着年平均气温的降低和年日照时数的增加, 栎属树种叶面积增加, 以利于吸收更多的光照辐射, 并增加叶片的边界层阻力, 减少叶片热量的散失; 而叶片分裂程度的增加不仅可以降低热量的散失, 也可以提高叶片液流的波动以增强叶片的生理活动; 叶脉密度随温度的升高、光照强度和降水量的增加而增加, 以响应叶片蒸腾作用的增强, 提高水分的运输能力和叶片的支撑能力。此外, 为适应南北样带暖温带区气候条件的变化, 栎属树种的叶片形成了一系列的形态性状组合, 随着叶面积的增加, 叶柄长度和叶片分裂程度逐渐增加, 而叶脉密度降低; 随着叶片倾向于向长条状发展, 叶柄长度和叶脉密度也随之增加。
李东胜, 史作民, 冯秋红, 刘峰. 中国东部南北样带暖温带区栎属树种叶片形态性状对气候条件的响应. 植物生态学报, 2013, 37(9): 793-802. DOI: 10.3724/SP.J.1258.2013.00083
LI Dong-Sheng, SHI Zuo-Min, FENG Qiu-Hong, LIU Feng. Response of leaf morphometric traits of Quercus species to climate in the temperate zone of the North-South Transect of Eastern China. Chinese Journal of Plant Ecology, 2013, 37(9): 793-802. DOI: 10.3724/SP.J.1258.2013.00083
地点 Site | 纬度 Latitude (N) | 经度 Longitude (E) | 树种 Species | 拉丁名 Latin name |
---|---|---|---|---|
吉林敦化市牡丹岭 Mudanling, Dunhua, Jilin | 43°21′ | 128°11′ | 蒙古栎 | Quercus mongolica |
辽宁营口市熊岳植物园 Xiongyue Botanical Garden, Yingkou, Liaoning | 40°40′ | 122°14′ | 蒙古栎 | Q. mongolica |
麻栎 | Q. acutissima | |||
河北兴隆县雾灵山 Wuling Mountain, Xinglong, Hebei | 40°26′ | 117°28′ | 蒙古栎 | Q. mongolica |
河北遵化市清东陵 Eastern Tombs of the Qing Dynasty, Zunhua, Hebei | 40°12′ | 117°58′ | 槲树 | Q. dentata |
麻栎 | Q. acutissima | |||
北京西山 Xishan Mountain, Beijing | 39°54′ | 116°10′ | 槲栎 | Q. aliena |
河北蔚县小五台山 Xiaowutai Mountain, Yu Xian, Hebei | 39°50′ | 114°26′ | 蒙古栎 | Q. mongolica |
辽东栎 | Q. liaotungensis | |||
河北秦皇岛北戴河 Beidaihe, Qinhuangdao, Hebei | 39°49′ | 119°29′ | 麻栎 | Q. acutissima |
山西五台县五台山 Wutai Mountain, Wutai, Shanxi | 38°43′ | 113°12′ | 粗齿蒙古栎 | Q. mongolica var. grosseserrata |
山西交城县关帝山 Guandi Mountain, Jiaocheng, Shanxi | 37°33′ | 112°08′ | 辽东栎 | Q. liaotungensis |
山西左权县太行山 Taihang Mountain, Zuoquan, Shanxi | 37°04′ | 113°21′ | 蒙古栎 | Q. mongolica |
栓皮栎 | Q. variabilis | |||
山西沁源县太岳山 Taiyue Mountain, Qinyuan, Shanxi | 36°30′ | 112°18′ | 蒙古栎 | Q. mongolica |
山西临汾市 Linfen, Shanxi | 36°04′ | 111°30′ | 辽东栎 | Q. liaotungensis |
山西陵川县 Lingchuan, Shanxi | 35°47′ | 113°16′ | 锐齿槲栎 | Q. alienavar.acuteserrata |
栓皮栎 | Q. variabilis | |||
槲树 | Q. dentata | |||
辽东栎 | Q. liaotungensis | |||
陕西韩城县黄龙山 Huanglong Mountain, Hancheng, Shaanxi | 35°28′ | 110°27′ | 辽东栎 | Q. liaotungensis |
山西垣曲县中条山 Zhongtiao Mountain, Yuanqu, Shanxi | 35°18′ | 111°38′ | 麻栎 | Q. acutissima |
河南焦作市云台山 Yuntai Mountain, Jiaozuo, Henan | 35°14′ | 113°25′ | 白栎 | Q. fabri |
山西夏县 Xia Xian, Shanxi | 35°07′ | 111°13′ | 槲栎 | Q. aliena |
河南济源 Jiyuan, Henan | 35°15′ | 112°07′ | 橿子栎 | Q. baronii |
山西永济市雪花山 Xuehua Mountain, Yongji, Shanxi | 34°55′ | 110°25′ | 辽东栎 | Q. liaotungensis |
陕西华阴市华山 Huashan Mountain, Huayin, Shaanxi | 34°35′ | 110°05′ | 橿子栎 | Q. baronii |
陕西眉县太白山 Taibai Mountain, Mei Xian, Shaanxi | 34°17′ | 107°45′ | 辽东栎 | Q. liaotungensis |
陕西西安市南五台山 Nanwutai Mountain, Xi’an, Shaanxi | 34°10′ | 108°56′ | 槲栎 | Q. aliena |
刺叶高山栎 | Q. spinosa | |||
橿子栎 | Q. baronii | |||
锐齿槲栎 | Q. alienavar.acuteserrata | |||
河南栾川县 Luanchuan, Henan | 33°45′ | 111°38′ | 橿子栎 | Q. baronii |
匙叶栎 | Q. dolicholepis |
表1 标本采集地点和树种
Table 1 Sites and species of specimen sampling
地点 Site | 纬度 Latitude (N) | 经度 Longitude (E) | 树种 Species | 拉丁名 Latin name |
---|---|---|---|---|
吉林敦化市牡丹岭 Mudanling, Dunhua, Jilin | 43°21′ | 128°11′ | 蒙古栎 | Quercus mongolica |
辽宁营口市熊岳植物园 Xiongyue Botanical Garden, Yingkou, Liaoning | 40°40′ | 122°14′ | 蒙古栎 | Q. mongolica |
麻栎 | Q. acutissima | |||
河北兴隆县雾灵山 Wuling Mountain, Xinglong, Hebei | 40°26′ | 117°28′ | 蒙古栎 | Q. mongolica |
河北遵化市清东陵 Eastern Tombs of the Qing Dynasty, Zunhua, Hebei | 40°12′ | 117°58′ | 槲树 | Q. dentata |
麻栎 | Q. acutissima | |||
北京西山 Xishan Mountain, Beijing | 39°54′ | 116°10′ | 槲栎 | Q. aliena |
河北蔚县小五台山 Xiaowutai Mountain, Yu Xian, Hebei | 39°50′ | 114°26′ | 蒙古栎 | Q. mongolica |
辽东栎 | Q. liaotungensis | |||
河北秦皇岛北戴河 Beidaihe, Qinhuangdao, Hebei | 39°49′ | 119°29′ | 麻栎 | Q. acutissima |
山西五台县五台山 Wutai Mountain, Wutai, Shanxi | 38°43′ | 113°12′ | 粗齿蒙古栎 | Q. mongolica var. grosseserrata |
山西交城县关帝山 Guandi Mountain, Jiaocheng, Shanxi | 37°33′ | 112°08′ | 辽东栎 | Q. liaotungensis |
山西左权县太行山 Taihang Mountain, Zuoquan, Shanxi | 37°04′ | 113°21′ | 蒙古栎 | Q. mongolica |
栓皮栎 | Q. variabilis | |||
山西沁源县太岳山 Taiyue Mountain, Qinyuan, Shanxi | 36°30′ | 112°18′ | 蒙古栎 | Q. mongolica |
山西临汾市 Linfen, Shanxi | 36°04′ | 111°30′ | 辽东栎 | Q. liaotungensis |
山西陵川县 Lingchuan, Shanxi | 35°47′ | 113°16′ | 锐齿槲栎 | Q. alienavar.acuteserrata |
栓皮栎 | Q. variabilis | |||
槲树 | Q. dentata | |||
辽东栎 | Q. liaotungensis | |||
陕西韩城县黄龙山 Huanglong Mountain, Hancheng, Shaanxi | 35°28′ | 110°27′ | 辽东栎 | Q. liaotungensis |
山西垣曲县中条山 Zhongtiao Mountain, Yuanqu, Shanxi | 35°18′ | 111°38′ | 麻栎 | Q. acutissima |
河南焦作市云台山 Yuntai Mountain, Jiaozuo, Henan | 35°14′ | 113°25′ | 白栎 | Q. fabri |
山西夏县 Xia Xian, Shanxi | 35°07′ | 111°13′ | 槲栎 | Q. aliena |
河南济源 Jiyuan, Henan | 35°15′ | 112°07′ | 橿子栎 | Q. baronii |
山西永济市雪花山 Xuehua Mountain, Yongji, Shanxi | 34°55′ | 110°25′ | 辽东栎 | Q. liaotungensis |
陕西华阴市华山 Huashan Mountain, Huayin, Shaanxi | 34°35′ | 110°05′ | 橿子栎 | Q. baronii |
陕西眉县太白山 Taibai Mountain, Mei Xian, Shaanxi | 34°17′ | 107°45′ | 辽东栎 | Q. liaotungensis |
陕西西安市南五台山 Nanwutai Mountain, Xi’an, Shaanxi | 34°10′ | 108°56′ | 槲栎 | Q. aliena |
刺叶高山栎 | Q. spinosa | |||
橿子栎 | Q. baronii | |||
锐齿槲栎 | Q. alienavar.acuteserrata | |||
河南栾川县 Luanchuan, Henan | 33°45′ | 111°38′ | 橿子栎 | Q. baronii |
匙叶栎 | Q. dolicholepis |
变量 Variable | 定义 Definition |
---|---|
叶长 Leaf length (cm) | 叶片基部到叶尖的最长轴长度 Longest line from leaf base to leaf apex |
叶宽 Leaf width (cm) | 垂直于主脉的最大宽度 Longest line perpendicular to the major axis |
叶面积 Leaf area (cm2) | |
叶片周长 Leaf perimeter (cm) | |
叶柄长 Leaf petiole length (cm) | |
叶脉密度 Leaf vein density | 叶片侧脉数量/叶面积 Leaf lateral veins numbers / leaf area |
松散度 Compactness | 叶片周长2/叶片面积 Leaf perimeter2 / leaf area |
长宽比 Length-width ratio | 叶片长度/叶片宽度 Leaf length / leaf width |
形态指数 Shape factor | 4π×叶面积/叶片周长2 4π × leaf area / leaf perimeter2 |
表2 叶片形态变量定义
Table 2 Definition of leaf physiognomic variables
变量 Variable | 定义 Definition |
---|---|
叶长 Leaf length (cm) | 叶片基部到叶尖的最长轴长度 Longest line from leaf base to leaf apex |
叶宽 Leaf width (cm) | 垂直于主脉的最大宽度 Longest line perpendicular to the major axis |
叶面积 Leaf area (cm2) | |
叶片周长 Leaf perimeter (cm) | |
叶柄长 Leaf petiole length (cm) | |
叶脉密度 Leaf vein density | 叶片侧脉数量/叶面积 Leaf lateral veins numbers / leaf area |
松散度 Compactness | 叶片周长2/叶片面积 Leaf perimeter2 / leaf area |
长宽比 Length-width ratio | 叶片长度/叶片宽度 Leaf length / leaf width |
形态指数 Shape factor | 4π×叶面积/叶片周长2 4π × leaf area / leaf perimeter2 |
气候因子 Climate factor | 最小值 Minimum | 最大值 Maximum | 平均值 Mean |
---|---|---|---|
年平均气温 Annual mean air temperature (℃) | 3.30 | 14.80 | 9.05 |
平均年降水量 Mean annual precipitation (mm) | 408.00 | 985.00 | 696.50 |
平均年日照时数 Mean annual sunlight (h) | 1 646.10 | 3 032.00 | 2 474.05 |
表3 气候因子的变化范围
Table 3 Variation ranges of climate factor
气候因子 Climate factor | 最小值 Minimum | 最大值 Maximum | 平均值 Mean |
---|---|---|---|
年平均气温 Annual mean air temperature (℃) | 3.30 | 14.80 | 9.05 |
平均年降水量 Mean annual precipitation (mm) | 408.00 | 985.00 | 696.50 |
平均年日照时数 Mean annual sunlight (h) | 1 646.10 | 3 032.00 | 2 474.05 |
叶片形态性状 Leaf morphological trait | 年平均气温 Annual mean air temperature (℃) | 平均年降水量 Mean annual precipitation (mm) | 平均年日照时数 Mean annual sunlight (h) | |||||
---|---|---|---|---|---|---|---|---|
相关系数 Correlation coefficient | 显著性Significance | 相关系数 Correlation coefficient | 显著性Significance | 相关系数 Correlation coefficient | 显著性 Significance | |||
Log叶面积 Log leaf area (cm2) | -0.590 | 0.000 | -0.303 | 0.051 | 0.527 | 0.000 | ||
Log叶柄长 Log leaf petiole length (cm) | 0.012 | 0.938 | -0.028 | 0.862 | 0.138 | 0.384 | ||
Log叶脉密度 Log leaf vein density | 0.527 | 0.000 | 0.360 | 0.019 | -0.495 | 0.001 | ||
Log松散度 Log compactness | -0.532 | 0.000 | -0.263 | 0.092 | 0.478 | 0.001 | ||
Log长宽比 Log length-width ratio | 0.078 | 0.621 | 0.122 | 0.442 | 0.129 | 0.415 | ||
Log形态指数 Log shape factor | 0.537 | 0.000 | 0.261 | 0.096 | -0.483 | 0.001 |
表4 栎属树种叶片形态性状与气候因子的关系
Table 4 Relationships between leaf morphological traits of Quercus trees and climate factors
叶片形态性状 Leaf morphological trait | 年平均气温 Annual mean air temperature (℃) | 平均年降水量 Mean annual precipitation (mm) | 平均年日照时数 Mean annual sunlight (h) | |||||
---|---|---|---|---|---|---|---|---|
相关系数 Correlation coefficient | 显著性Significance | 相关系数 Correlation coefficient | 显著性Significance | 相关系数 Correlation coefficient | 显著性 Significance | |||
Log叶面积 Log leaf area (cm2) | -0.590 | 0.000 | -0.303 | 0.051 | 0.527 | 0.000 | ||
Log叶柄长 Log leaf petiole length (cm) | 0.012 | 0.938 | -0.028 | 0.862 | 0.138 | 0.384 | ||
Log叶脉密度 Log leaf vein density | 0.527 | 0.000 | 0.360 | 0.019 | -0.495 | 0.001 | ||
Log松散度 Log compactness | -0.532 | 0.000 | -0.263 | 0.092 | 0.478 | 0.001 | ||
Log长宽比 Log length-width ratio | 0.078 | 0.621 | 0.122 | 0.442 | 0.129 | 0.415 | ||
Log形态指数 Log shape factor | 0.537 | 0.000 | 0.261 | 0.096 | -0.483 | 0.001 |
图1 栎属树种叶片形态性状与年平均气温(MAT)的关系。SMA, 标准化主轴法的斜率。
Fig. 1 Relationships between leaf morphological traits of Quercus species and annual mean air temperature (MAT). SMA, slope for standardized major axis.
图2 栎属树种叶脉密度与平均年降水量(MAP)的关系。SMA, 标准化主轴法的斜率。
Fig. 2 Relationships between leaf vein density of Quercus species and mean annual precipitation (MAP). SMA, slope for standardized major axis.
图3 栎属树种叶片形态性状与平均年日照时数(MAS)的关系。SMA, 标准化主轴法的斜率。
Fig. 3 Relationships between leaf morphological traits of Quercus species and mean annual sunlight (MAS). SMA, slope for standardized major axis.
叶片形态性状 Leaf morphological trait | 系数 Coefficient | Log叶面积 Log leaf area (cm2) | Log叶柄长 Log leaf petiole length (cm) | Log叶脉密度 Log leaf vein density | Log松散度 Log compactness | Log长宽比 Log length- width ratio | Log形态指数 Log shape factor |
---|---|---|---|---|---|---|---|
Log叶面积 Log leaf area (cm2) | 相关系数 Correlation coefficient | 1.000 | |||||
显著性 Significance | 0.000 | ||||||
Log叶柄长 Log leaf petiole length (cm) | 相关系数 Correlation coefficient | 0.318 | 1.000 | ||||
显著性 Significance | 0.040 | 0.000 | |||||
Log叶脉密度 Log leaf vein density | 相关系数 Correlation coefficient | -0.931 | -0.134 | 1.000 | |||
显著性 Significance | 0.000 | 0.397 | 0.000 | ||||
Log松散度 Log compactness | 相关系数 Correlation coefficient | 0.537 | -0.047 | -0.512 | 1.000 | ||
显著性 Significance | 0.000 | 0.765 | 0.001 | 0.000 | |||
Log长宽比 Log length-width ratio | 相关系数 Correlation coefficient | -0.138 | 0.351 | 0.398 | 0.111 | 1.000 | |
显著性 Significance | 0.384 | 0.023 | 0.009 | 0.485 | 0.000 | ||
Log形态指数 Log shape factor | 相关系数 Correlation coefficient | -0.546 | 0.058 | 0.524 | -0.998 | -0.102 | 1.000 |
显著性 Significance | 0.000 | 0.717 | 0.000 | 0.000 | 0.519 | 0.000 |
表5 栎属树种叶片形态性状之间的关系
Table 5 Relationships among leaf morphological traits of Quercus species
叶片形态性状 Leaf morphological trait | 系数 Coefficient | Log叶面积 Log leaf area (cm2) | Log叶柄长 Log leaf petiole length (cm) | Log叶脉密度 Log leaf vein density | Log松散度 Log compactness | Log长宽比 Log length- width ratio | Log形态指数 Log shape factor |
---|---|---|---|---|---|---|---|
Log叶面积 Log leaf area (cm2) | 相关系数 Correlation coefficient | 1.000 | |||||
显著性 Significance | 0.000 | ||||||
Log叶柄长 Log leaf petiole length (cm) | 相关系数 Correlation coefficient | 0.318 | 1.000 | ||||
显著性 Significance | 0.040 | 0.000 | |||||
Log叶脉密度 Log leaf vein density | 相关系数 Correlation coefficient | -0.931 | -0.134 | 1.000 | |||
显著性 Significance | 0.000 | 0.397 | 0.000 | ||||
Log松散度 Log compactness | 相关系数 Correlation coefficient | 0.537 | -0.047 | -0.512 | 1.000 | ||
显著性 Significance | 0.000 | 0.765 | 0.001 | 0.000 | |||
Log长宽比 Log length-width ratio | 相关系数 Correlation coefficient | -0.138 | 0.351 | 0.398 | 0.111 | 1.000 | |
显著性 Significance | 0.384 | 0.023 | 0.009 | 0.485 | 0.000 | ||
Log形态指数 Log shape factor | 相关系数 Correlation coefficient | -0.546 | 0.058 | 0.524 | -0.998 | -0.102 | 1.000 |
显著性 Significance | 0.000 | 0.717 | 0.000 | 0.000 | 0.519 | 0.000 |
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