山东省常用恢复树种径向生长与气候关系的研究
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摘要
应用植被生态学理论恢复森林生态系统，提倡和强调用乡土树种重建乡土森林植被(Native forest with native trees)，容易在较短的时间内恢复当地森林生态系统。本地乡土种作为生态恢复和重建的目标，和探索关键技术的研究对象，加强对其生长特性的了解意义重大。研究表明树木的生长尤其是径向生长和立地环境密切相关并受气候变化的影响，通过研究树轮宽度的生长变异，不但可以了解树木历年的生长情况及其所在地历年气候的变化情况，还能够帮助人们预测未来气候变化以及研究气候变化对树木生长和森林结构与功能的影响，为探讨生态恢复与重建措施提供令人信服的证据。本研究选择外来树种刺槐(Robinia pseudoacacia L．)和两种乡土树种赤松(Pinus densiflora Sieb．et Zucc．)、麻栎(Quercus acutissima Carr．)作为研究对象，旨在为适宜树种选择提供重要的理论依据。刺槐作为一个较常用的混交树种和植被重建的先锋树种对我国温带大部分地区植被恢复和生态重建具有重大的意义，并且刺槐林已成为山东省分布最广和面积最大的落叶阔叶林。赤松林作为山东省地带性的针叶林，也是面积较大的针叶林。已有的研究表明赤松作为一个亚顶级乡土针叶种，不但成为主要的森林建群树种，也能够为其他地带性落叶阔叶树种入侵、生长提供基础，创造适生条件，这对于在荒山秃岭科学地营造赤松林以及重建山东省地带性落叶阔叶林群落有重大意义。而麻栎作为山东省的典型的、地带性的乡土落叶阔叶树种，其生长、发育、繁衍和分布与当地的生态条件，特别是气候和土壤条件相适应。这三种树种作为山东乃至中国华北生态恢复与重建的先锋种、顶级优势种代表，研究其树木径向生长与环境气候因子之间的关系，能从生理生态学方面为山东乃至中国北方的生态恢复引种造林、森林抚育、演替提供依据，具有非常重要的理论价值和实践意义。
本研究考虑到地理位置的代表意义和三种树种的发展现状，选取莱芜房干(LF)、枣庄抱犊崮(ZB)、东营仙河(DX)、青岛崂山(QL)、德州临邑(DL)、菏泽曹县(HC)六个较典型地区作为此次研究样地，基于国际树木年轮数据库(ITRDB)的标准进行野外取样，对样本进行交叉定年和宽度测量。参数结果显示样本平均敏感度较高，刺槐分布范围在0.25～0.5之间，赤松分布范围在0.20～0.30之间，麻栎分布范围在0.25～0.30之间，均符合平均敏感度取值要求(＞0.1)，表明三种树种受气候因子的限制作用明显，年轮宽度足以反映样点地区环境变化。研究选用指数函数曲线来模拟三个树种径向生长趋势，经过检验是可靠的，都达到极显著水平，据此求得树轮指数序列(RWI)。选择当年生长季(CG：4、5、6、7、8、9、10月)，生长季外(OG：11、12、1、2、3月)，上一个生长季(PG：4、5、6、7、8、9、10月)的月平均气温(T_m)和月降水量(P_m)作为气象因子与树轮指数序列进行相关分析。
刺槐在6个样点的相关分析显示，刺槐径向生长在莱芜房干地区主要受上一年11月降水、当年2、3月的降水和温度以及当年7月的温度限制；在青岛崂山地区主要受当年5月的温度，以及当年2、10月降水的控制；在东营仙河地区主要受当年3月的温度、上一年11月和当年9月的降水影响；在德州临邑地区主要受上一年5月和当年3月温度、当年10月降水限制；在菏泽曹县地区主要受上一年12月的温度以及当年7月的温度和降水控制；在枣庄抱犊崮地区则主要受上一年12月和当年7月的温度显著影响。由此可以看出，刺槐的生长在生长季前期和生长旺季都主要受到温度的限制，而夏季降水高峰过去之后，刺槐生长对降水的需求则明显表现出来，并且由于温度和降水的滞后效应，上一年秋冬季的温度和降水对刺槐生长也具有显著影响。具体到各个地区由于气候差异，温度和降水的限制作用也有所差异。
赤松在莱芜房干和青岛崂山的相关分析显示，莱芜房干当年2月降水和10月温度对赤松径向生长有正相关作用，10月的降水对赤松径向生长有负相关作用；在青岛崂山，赤松径向生长与当年2-5月、9月温度有负相关。这表明生长季前期和末期的气候因子对赤松生长影响较大，但两个地区的影响作用也不同，这可能是由赤松自身生理特性以及地区气候差异而导致的结果。赤松为喜光树种，在莱芜房干生长季末温度升高伴随降水增加能够增强光合作用，仍能促进赤松生长，但降水过度增加造成光照减少，影响光合作用反而制约赤松生长。在青岛崂山，赤松生长与春季低温相适应，温度升高使树木呼吸作用加强，而光合作用受阻，从而抑制赤松生长。
麻栎在莱芜房干和枣庄抱犊崮的相关分析显示，麻栎径向生长在莱芜房干主要受上一年12月的温度，上一年5、11月和当年5、6月降水的限制；在枣庄抱犊崮主要受上一年6、12月和当年1、2、9、10月的温度，以及上一年11月降水的显著影响。这是因为枣庄抱犊崮的降水丰富从而降低对树木生长的限制作用，温度的影响表现的更加明显；莱芜房干的降水远低于枣庄抱犊崮，在温度与枣庄抱犊崮较接近的情况下，降水成为麻栎生长的主要限制因子。
对刺槐、赤松、麻栎分别进行单年分析，对每年的温度和降水因子求距平，检查与其相应的树轮指数变化，研究显示相关分析的结果能够充分解释产生宽轮和窄轮的原因，进一步验证了相关分析的可靠性和稳定性。
应用多重逐步线性回归模型描述树木径向生长与环境因子之间关系，结果表明各线性回归方程的方差解释比值较高，F检验显著，这说明模型整体拟合效果较好，同时又采用岭回归分析建立多重回归方程，尽管岭回归分析比多重逐步线性回归分析的精确度减小，但更符合实际，并且方程整体高度显著。
综上所述，本研究结果得到了刺槐、赤松、麻栎在不同地区生长的不同气候限制因子，对三个树种径向生长与气候因子间的关系进行了定量描述，为植被生态恢复与重建过程中人工造林和封山育林的经营管理和保护提供了科学指导依据，对生态恢复与重建的定量化和模型化研究提供了补充，具有重要的理论和实践意义。
Applying Vegetation ecology theory to reconstruct forest ecosystem is a new focus of restoration ecology, which emphasizes reconstruction of the native forest with the native trees to restore local forest ecosystem. Therefore, the native trees become the aim of ecological restoration as well as the study object of developing key techniques. Thorough study on the growth traits should be done. The growth especially radial growth of trees has great relations to site conditions and climate changes. Base on the growth variance of tree ring width, we can get some information of the annual growth of the trees and the annual climatic changes in their locations. The effects of the future climatic changes on tree growth and forest structure and function can be predicted, which can provide a credible evidence for ecological restoration measures. In this article three important north species are selected, Robinia psendoacacia L. (black locust) , Pinus densiflora Sieb. et Zucc. (red pine) and Quercus acutissima Carr.(sawtooth oak) to study the relationships between climatic changes and tree radial growth. Black locust, as a perfect hybrid species and a pioneer species in restoring vegetation, is very valuable in restoring vegetation and ecological construction in temperate zone of China In the drought and semi-drought areas, such as Loess Plateau, applying black locust has made better progress. And the largest and broadest broadleaf tree in Shandong province is black locust. Red pine, as the native conifer, has the largest afforestation areas in Shandong province. Some previous studies show that red pine can provide foundation and create conditions for invading and growing of other zonal deciduous broadleaf trees, which is significant for constructing restoring red pines scientifically and restoring deciduous broadleaf trees. Sawtooth oak, as the native deciduous broadleaf tree, its growth, reproduction and distribution accommodate to local environment such as climate and soil conditions. The relationship between tree radial growth and climatic factors of black locust, red pine and sawtooth oak are investigated in this study, which can provide the foundation for ecological restoration from physiological ecology of Shandong province and north China. There is great theoretic value and practical value of artificial afforestation, forest tending and succession in this study.
Considering geographical meaning and developing conditions of three species, Laiwu Fanggan (LF ), Zaozhuang Baodugu (ZB ), Dongying Xianhe (DX), Qingdao Laoshan (QL), Dezhou Linyi (DL) and Heze Caoxian (HC) are chosen as sampling plots. We carry out the fieldwork basing on ITRDB. The result indicates that MS is high enough. The MS of black locusts, red pines and sawtooth Oak are respectively from 0.25 to 0.5, 0.2 to 0.3 and 0.25 to 0.3, which shows that the growth of the three species are limited by climatic factors. We choose exponential function to simulate growth trend of three species and get the RWI. Monthly mean temperature and monthly precipitation of current growth season (Apr, May, Jun., Jul., Aug., Sep., Oct.) , out of growth season (Nov., Dec, Jan., Feb., Mar.) , and previous growth season (Apr., May, Jun., Jul., Aug., Sep., Oct.) are selected as climatic factors to process correlation analysis between climatic factors and RWI.
Correlation analyses of black locusts in six locations indicate that the radial growth is mostly limited by precipitation in previous winter, precipitation and temperature in current spring and temperature of Jul. in LF. In QL it is mostly controlled by temperature and precipitation in current spring and precipitation of Oct.. In DX it is limited by temperature in current spring and precipitation of previous autumn and Sep.. In DL it is effected mostly by temperature of previous May and current Mar. and precipitation of Oct.. It is controlled by temperature of previous winter, temperature and precipitation of Jul. in HC. Also it is affected mostly by temperature of previous winter and current summer. Therefore, the growth of black locust is mostly limited by temperature in its prophase and midseason of growth season. After peak precipitation in summer, the positive needs of the precipitation for growth show obviously. And for the Lagging effect of temperature and precipitation, the temperature and precipitation in previous autumn and winter influence greatly on the growth of black locust.
Correlation analyses of red pines in LF and QL indicate that precipitation of February, temperature and precipitation of Oct. have prominent effect on the radial growth in LF, and in QL the growth is mostly limited by temperature of Feb., Mar., Apr., May and Sep.. As a species fond of light, red pine can be accelerated because rising temperature at the end of growing season with precipitation increasing could strengthen photosynthesis in LF. But excessive precipitation can affect photosynthesis leading to the reduction of the growth. In QL low temperature in spring can be satisfied with the growth. On the contrary rising temperature may aggravate evaporation so as to restrain the growth.
Correlation analyses of sawtooth oak in LF and ZB indicate that the radial growth is controlled mostly by the precipitation of previous spring, temperature and precipitation of previous autumn and winter, precipitation of May and Jun. in LF. In ZB the growth is effected obviously by temperature of previous Jun., temperature and precipitation of previous autumn and winter, temperature of Jan., Feb., Sep. and Oct.. It shows that the rich precipitation can reduce the limitation on the growth in ZB, which make the temperature effect more obviously. Because the precipitation in LF is far lower than that in ZB, it becomes the primary limiting factor on the growth in LF with the case of temperature being nearly in both two locations.
The Stepwise Multiple Regression is applied to describe the relation between the radial growth and environmental factors. The results show that the variance explanation ratio is much higher in each equation and F test is prominent. Moreover, the Ridge Regression Analysis also is used to build the equation to get better results.
Based on the results, the climatic limiting factors are shown and the relationships between the radial growth of three species and the climatic factors are quantitatively described in this study, which can provide scientific foundation for vegetation ecological restoration and provide supplement for the quantitative and modeling studies in ecological restoration, and it is valuable in theory and practice.
本研究考虑到地理位置的代表意义和三种树种的发展现状，选取莱芜房干(LF)、枣庄抱犊崮(ZB)、东营仙河(DX)、青岛崂山(QL)、德州临邑(DL)、菏泽曹县(HC)六个较典型地区作为此次研究样地，基于国际树木年轮数据库(ITRDB)的标准进行野外取样，对样本进行交叉定年和宽度测量。参数结果显示样本平均敏感度较高，刺槐分布范围在0.25～0.5之间，赤松分布范围在0.20～0.30之间，麻栎分布范围在0.25～0.30之间，均符合平均敏感度取值要求(＞0.1)，表明三种树种受气候因子的限制作用明显，年轮宽度足以反映样点地区环境变化。研究选用指数函数曲线来模拟三个树种径向生长趋势，经过检验是可靠的，都达到极显著水平，据此求得树轮指数序列(RWI)。选择当年生长季(CG：4、5、6、7、8、9、10月)，生长季外(OG：11、12、1、2、3月)，上一个生长季(PG：4、5、6、7、8、9、10月)的月平均气温(T_m)和月降水量(P_m)作为气象因子与树轮指数序列进行相关分析。
刺槐在6个样点的相关分析显示，刺槐径向生长在莱芜房干地区主要受上一年11月降水、当年2、3月的降水和温度以及当年7月的温度限制；在青岛崂山地区主要受当年5月的温度，以及当年2、10月降水的控制；在东营仙河地区主要受当年3月的温度、上一年11月和当年9月的降水影响；在德州临邑地区主要受上一年5月和当年3月温度、当年10月降水限制；在菏泽曹县地区主要受上一年12月的温度以及当年7月的温度和降水控制；在枣庄抱犊崮地区则主要受上一年12月和当年7月的温度显著影响。由此可以看出，刺槐的生长在生长季前期和生长旺季都主要受到温度的限制，而夏季降水高峰过去之后，刺槐生长对降水的需求则明显表现出来，并且由于温度和降水的滞后效应，上一年秋冬季的温度和降水对刺槐生长也具有显著影响。具体到各个地区由于气候差异，温度和降水的限制作用也有所差异。
赤松在莱芜房干和青岛崂山的相关分析显示，莱芜房干当年2月降水和10月温度对赤松径向生长有正相关作用，10月的降水对赤松径向生长有负相关作用；在青岛崂山，赤松径向生长与当年2-5月、9月温度有负相关。这表明生长季前期和末期的气候因子对赤松生长影响较大，但两个地区的影响作用也不同，这可能是由赤松自身生理特性以及地区气候差异而导致的结果。赤松为喜光树种，在莱芜房干生长季末温度升高伴随降水增加能够增强光合作用，仍能促进赤松生长，但降水过度增加造成光照减少，影响光合作用反而制约赤松生长。在青岛崂山，赤松生长与春季低温相适应，温度升高使树木呼吸作用加强，而光合作用受阻，从而抑制赤松生长。
麻栎在莱芜房干和枣庄抱犊崮的相关分析显示，麻栎径向生长在莱芜房干主要受上一年12月的温度，上一年5、11月和当年5、6月降水的限制；在枣庄抱犊崮主要受上一年6、12月和当年1、2、9、10月的温度，以及上一年11月降水的显著影响。这是因为枣庄抱犊崮的降水丰富从而降低对树木生长的限制作用，温度的影响表现的更加明显；莱芜房干的降水远低于枣庄抱犊崮，在温度与枣庄抱犊崮较接近的情况下，降水成为麻栎生长的主要限制因子。
对刺槐、赤松、麻栎分别进行单年分析，对每年的温度和降水因子求距平，检查与其相应的树轮指数变化，研究显示相关分析的结果能够充分解释产生宽轮和窄轮的原因，进一步验证了相关分析的可靠性和稳定性。
应用多重逐步线性回归模型描述树木径向生长与环境因子之间关系，结果表明各线性回归方程的方差解释比值较高，F检验显著，这说明模型整体拟合效果较好，同时又采用岭回归分析建立多重回归方程，尽管岭回归分析比多重逐步线性回归分析的精确度减小，但更符合实际，并且方程整体高度显著。
综上所述，本研究结果得到了刺槐、赤松、麻栎在不同地区生长的不同气候限制因子，对三个树种径向生长与气候因子间的关系进行了定量描述，为植被生态恢复与重建过程中人工造林和封山育林的经营管理和保护提供了科学指导依据，对生态恢复与重建的定量化和模型化研究提供了补充，具有重要的理论和实践意义。
Applying Vegetation ecology theory to reconstruct forest ecosystem is a new focus of restoration ecology, which emphasizes reconstruction of the native forest with the native trees to restore local forest ecosystem. Therefore, the native trees become the aim of ecological restoration as well as the study object of developing key techniques. Thorough study on the growth traits should be done. The growth especially radial growth of trees has great relations to site conditions and climate changes. Base on the growth variance of tree ring width, we can get some information of the annual growth of the trees and the annual climatic changes in their locations. The effects of the future climatic changes on tree growth and forest structure and function can be predicted, which can provide a credible evidence for ecological restoration measures. In this article three important north species are selected, Robinia psendoacacia L. (black locust) , Pinus densiflora Sieb. et Zucc. (red pine) and Quercus acutissima Carr.(sawtooth oak) to study the relationships between climatic changes and tree radial growth. Black locust, as a perfect hybrid species and a pioneer species in restoring vegetation, is very valuable in restoring vegetation and ecological construction in temperate zone of China In the drought and semi-drought areas, such as Loess Plateau, applying black locust has made better progress. And the largest and broadest broadleaf tree in Shandong province is black locust. Red pine, as the native conifer, has the largest afforestation areas in Shandong province. Some previous studies show that red pine can provide foundation and create conditions for invading and growing of other zonal deciduous broadleaf trees, which is significant for constructing restoring red pines scientifically and restoring deciduous broadleaf trees. Sawtooth oak, as the native deciduous broadleaf tree, its growth, reproduction and distribution accommodate to local environment such as climate and soil conditions. The relationship between tree radial growth and climatic factors of black locust, red pine and sawtooth oak are investigated in this study, which can provide the foundation for ecological restoration from physiological ecology of Shandong province and north China. There is great theoretic value and practical value of artificial afforestation, forest tending and succession in this study.
Considering geographical meaning and developing conditions of three species, Laiwu Fanggan (LF ), Zaozhuang Baodugu (ZB ), Dongying Xianhe (DX), Qingdao Laoshan (QL), Dezhou Linyi (DL) and Heze Caoxian (HC) are chosen as sampling plots. We carry out the fieldwork basing on ITRDB. The result indicates that MS is high enough. The MS of black locusts, red pines and sawtooth Oak are respectively from 0.25 to 0.5, 0.2 to 0.3 and 0.25 to 0.3, which shows that the growth of the three species are limited by climatic factors. We choose exponential function to simulate growth trend of three species and get the RWI. Monthly mean temperature and monthly precipitation of current growth season (Apr, May, Jun., Jul., Aug., Sep., Oct.) , out of growth season (Nov., Dec, Jan., Feb., Mar.) , and previous growth season (Apr., May, Jun., Jul., Aug., Sep., Oct.) are selected as climatic factors to process correlation analysis between climatic factors and RWI.
Correlation analyses of black locusts in six locations indicate that the radial growth is mostly limited by precipitation in previous winter, precipitation and temperature in current spring and temperature of Jul. in LF. In QL it is mostly controlled by temperature and precipitation in current spring and precipitation of Oct.. In DX it is limited by temperature in current spring and precipitation of previous autumn and Sep.. In DL it is effected mostly by temperature of previous May and current Mar. and precipitation of Oct.. It is controlled by temperature of previous winter, temperature and precipitation of Jul. in HC. Also it is affected mostly by temperature of previous winter and current summer. Therefore, the growth of black locust is mostly limited by temperature in its prophase and midseason of growth season. After peak precipitation in summer, the positive needs of the precipitation for growth show obviously. And for the Lagging effect of temperature and precipitation, the temperature and precipitation in previous autumn and winter influence greatly on the growth of black locust.
Correlation analyses of red pines in LF and QL indicate that precipitation of February, temperature and precipitation of Oct. have prominent effect on the radial growth in LF, and in QL the growth is mostly limited by temperature of Feb., Mar., Apr., May and Sep.. As a species fond of light, red pine can be accelerated because rising temperature at the end of growing season with precipitation increasing could strengthen photosynthesis in LF. But excessive precipitation can affect photosynthesis leading to the reduction of the growth. In QL low temperature in spring can be satisfied with the growth. On the contrary rising temperature may aggravate evaporation so as to restrain the growth.
Correlation analyses of sawtooth oak in LF and ZB indicate that the radial growth is controlled mostly by the precipitation of previous spring, temperature and precipitation of previous autumn and winter, precipitation of May and Jun. in LF. In ZB the growth is effected obviously by temperature of previous Jun., temperature and precipitation of previous autumn and winter, temperature of Jan., Feb., Sep. and Oct.. It shows that the rich precipitation can reduce the limitation on the growth in ZB, which make the temperature effect more obviously. Because the precipitation in LF is far lower than that in ZB, it becomes the primary limiting factor on the growth in LF with the case of temperature being nearly in both two locations.
The Stepwise Multiple Regression is applied to describe the relation between the radial growth and environmental factors. The results show that the variance explanation ratio is much higher in each equation and F test is prominent. Moreover, the Ridge Regression Analysis also is used to build the equation to get better results.
Based on the results, the climatic limiting factors are shown and the relationships between the radial growth of three species and the climatic factors are quantitatively described in this study, which can provide scientific foundation for vegetation ecological restoration and provide supplement for the quantitative and modeling studies in ecological restoration, and it is valuable in theory and practice.
引文
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