华北地区3个种源文冠果苗期水分利用特性及抗旱性研究
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摘要
当今世界能源枯竭、环境污染、粮食安全等问题已日益严重，作为一种可再生绿色能源---生物柴油备受关注，而阻碍其产业化的瓶颈则是原料供应及成本。因此，挖掘适宜非耕地种植的木本油料植物，发展生态能源林，丰富生物柴油原料的供应链，显得尤为重要。文冠果（Xanthocera sorbifolia Bunge）以其分布范围广、适应性强、种子含油率高、燃油品质优、开发利用价值高等特点，成为近年我国北方地区大力推广的生物柴油树种，具有巨大的开发和利用潜力。本文是在对我国文冠果资源的系统调查、分布区划分以及生物学特性等研究的基础上，开展文冠果集中分布的华北地区3个不同种源的室内盆栽控水试验，研究不同梯度水分胁迫对苗木生长、生理特性、水分利用效率、耗水规律以及其叶片形态解剖结构等方面的影响，进而探究不同种源文冠果对水分胁迫的响应机制；同时，开展文冠果抗旱节水综合评价指标体系的研建，并对不同种源文冠果抗旱性进行综合评价与筛选。主要研究结果如下：
（1）在水分胁迫条件下，3个不同种源文冠果的苗高和地径生长量均表现出山西最高、河北居中、河南最差的趋势；而且在重度水分胁迫处理的后期，河北和河南种源文冠果出现叶片发黄及脱落现象，而山西种源苗木未出现；
随着处理时间的增加，充足水分处理下的3个种源文冠果叶片的保水能力均逐渐降低、叶绿素含量并未出现明显差异，但山西种源保持相对较高的叶片保水能力；在重度水分胁迫条件下，3个种源文冠果叶片保水能力均有所增强、叶绿素含量明显降低，其中山西种源叶片保水力增大最多，且在复水后仍保持较高的叶片保水能力，并具有相对较低的蒸腾速率。山西种源不论其生长状况、抗旱能力均好于河北和河南两种源。
（2）不同种源文冠果的碳稳定同位素分馏、水分利用效率和耗水量对水分胁迫响应程度存在一定差异。与河南及河北两种源相比，山西种源文冠果在各种梯度水分胁迫下均能保持相对较高的碳同位素比率（δ13C）值、水分利用效率（WUE）值以及较低的耗水量，表明山西种源文冠果在水分胁迫下对水分亏缺响应较为敏感，能通过短期内迅速减少耗水量来适应水分胁迫环境，这也是山西种源文冠果具有较高抗旱能力的体现。
（3）不同种源文冠果的叶片形态结构对水分胁迫响应程度存在一定差异，与河南及河北两种源相比，山西种源文冠果在各种胁迫时期具有较高的比叶重与较厚的各组织结构，且能形成较多的晶体以及小而多的导管等，结合耗水量最少的实验结果，表明山西种源文冠果更趋向于旱生结构。
（4）与河南及河北两种源相比，山西种源文冠果在各种梯度胁迫时期均能保持相对较高的水分利用效率和叶片保水能力以及较低的蒸腾速率、叶绿素含量和耗水量，表明山西种源文冠果具有较高的抗旱能力，是形态及解剖构造和水分生理生态特性对干旱环境适应性响应的结果。
（5）在不同梯度水分胁迫对文冠果苗木生长及生理特性和叶片形态解剖结构的分析研究的基础上，应用主成分分析和多维空间坐标综合评定值累加方法，建立文冠果抗旱节水综合评价指标体系，并对3种源文冠果苗木的抗旱性进行综合比较评价。结果发现，山西种源文冠果的抗旱性最强、河北种源次之、河南种源最差，这与线性回归模型验证结果以及水分胁迫下的叶片形态解剖结构和细胞水分利用特性的研究结果相一致。
以上研究结果表明，不同种源文冠果苗木生长、叶片的形态解剖结构以及生理特性对水分胁迫的响应程度存在一定差异，并表现出相应的适应性改变，进而导致其抗旱性生理功能的差异；对不同种源文冠果抗旱性进行综合评价，筛选出山西种源文冠果为抗旱性最强种源。这些研究结果可为我国干旱地区文冠果栽培以及优良种源筛选提供理论依据，具有重大的现实意义。
Problems of fossil energy depletion, environmental pollution and food security, have gonefrom bad to worse all over the world in recent years. Considerable attention has been paid tothe bio-diesel as renewable green energy. However, the provision and high cost of bio-dieselraw materials are the major barriers for its commercialization. Therefore, implementation ofthe bio-diesel industry must get a plentiful supply of raw materials. Use of woody bio-dieselfeedstock can reduce bio-diesel production cost and increase the supply. The investigation ofwoody oil plants suitable for non-arable land cultivation, development of bio-energy forest,and improvement of bio-diesel feedstock supply chain, is particularly important. Xanthocerasorbifolia Bunge has many advantages, including its wide distribution area, wide ecologicalamplitude, high seed oil, excellent fuel property, development and utilization of high value,and others. It was bio-diesel species promoted in northern China, and has enormous potentialfor development and utilization in recent years. In this paper, indoor potted water control testof3different X. sorbifolia Bunge provenances was worked out on the basis of systematicinvestigation, categorization of the resources distribution and biological characteristics research.We studied different gradients of drought stress on seedling growth, physiologicalcharacteristics, water use efficiency, water consumption law and its morphological anatomicalstructure of leaves, etc., and then explored X. sorbifolia of different provenances responsemechanism on water stress; Simultaneously, the X.sorbifolia resisting drought and saving watercomprehensive evaluation index system has been build, and the drought resistance of X.sorbifolia of different provenances was been comprehensive evaluated and screened. Theresults are as follows:
（1）The height and diameter growth of X. sorbifolia of three different provenancesshowed the trends which the growth of Shanxi provenance is the highest and the Henan is theworst under water stress conditions; the X. sorbifolia leaves of Hebei and Henan provenances appear yellow or fall off, and Shanxi provenance seedlings did not appear in the severe waterstress treatment.
With increasing the treatment time, water retention capacity of3provenances X.sorbifolia leaves was gradually reduced, and the chlorophyll content did not show significantdifferences. However, the X. sorbifolia leaves of Shanxi provenance maintained at a relativelyhigh water retention capacity; Water retention capacity of3provenances X. sorbifolia leaveshas been strengthened, and the chlorophyll content decreased under severe water stressconditions. Among them, water retention capacity of Shanxi provenance X. sorbifolia leaveshas the largest increases, remained higher after re-water, and has a relatively low transpirationrate. Regardless of their growing conditions or drought resistance, the X. sorbifolia Shanxiprovenances of was better than Henan and Hebei.
（2）The X. sorbifolia of different provenances response degree of stable carbon isotopefractionation, water use efficiency and water consumption on water stress exists a certaindifferences. Compared with the henan and hebei provenance, relatively high carbon isotoperatio value, water use efficiency and low water consumption of shanxi provenance X. sorbifoliacould maintain in all sorts of gradient water stress. This indicates the response of Shanxiprovenance is more sensitive to water deficit under drought stress, and to adapt to droughtstress environment by rapidly reducing water consumption in the short term, which was alsothe high drought tolerance expression of Shanxi provenance X. sorbifolia.
（3）There are some differences to water stress responsiveness among morphology ofleaves of different provenances X. sorbifolia. Compared with the Henan and Hebeiprovenances, X. sorbifolia of Shanxi provenances has higher specific leaf weight (SLW)andthicker organizations structures in various stress period, and can form more crystal sand vesselsof small and many and etc. Combined with experimental results of least water usage, X.sorbifolia of Shanxi provenances tend to xeromorphism.
（4）Compared with the Henan and Hebei provenances, X. sorbifolia of Shanxiprovenances can maintain a relatively higher water use efficiency (WUE), water-holding capacity and lower transpiration rate of leaf, chlorophyll content and water consumption invarious kinds of stress gradients period. These indicate X. sorbifolia of Shanxi provenances hashigher drought resistance capacity. That was the result of morphological and anatomicalstructure and water physiological ecology in response to drought environmental adaptability.
（5）On the basis of different gradients of water stress on seedling growth andphysiological property of X. sorbifolia and analysis of leaf morphological and anatomicalstructure, the drought resistance system of comprehensive evaluation of X. sorbifolia has beencreated by principal component analysis and comprehensive assessment of the value ofmulti-dimensional spatial coordinates accumulation method. And3X.sorbifolia provenanceshave been evaluated. It was found that X. sorbifolia of Shanxi provenances has strongestdrought resistance, followed Hebei’s, and Henan’s worst, consisting with the results of linearregression model validation and leaves morphological and anatomical structure under droughtstress and property of cell water-using.
The above findings suggest that there are some differences about seedling growth, leafmorphological and anatomical structures and physiological property of X. sorbifolia ofdifferent provenances in the response to drought stress, and exhibit appropriate adaptivechanges, leading to differences in their physiological functions of drought resistance.Comprehensively evaluating drought resistance of X.sorbifolia of different provenances, X.sorbifolia of Shanxi provenance has been screened as the provenances of strongest droughtresistance. These findings may provide a theoretical basis for X. sorbifolia cultivation in aridregions of China and excellent provenance screening. It is of great practical significance.
（1）在水分胁迫条件下，3个不同种源文冠果的苗高和地径生长量均表现出山西最高、河北居中、河南最差的趋势；而且在重度水分胁迫处理的后期，河北和河南种源文冠果出现叶片发黄及脱落现象，而山西种源苗木未出现；
随着处理时间的增加，充足水分处理下的3个种源文冠果叶片的保水能力均逐渐降低、叶绿素含量并未出现明显差异，但山西种源保持相对较高的叶片保水能力；在重度水分胁迫条件下，3个种源文冠果叶片保水能力均有所增强、叶绿素含量明显降低，其中山西种源叶片保水力增大最多，且在复水后仍保持较高的叶片保水能力，并具有相对较低的蒸腾速率。山西种源不论其生长状况、抗旱能力均好于河北和河南两种源。
（2）不同种源文冠果的碳稳定同位素分馏、水分利用效率和耗水量对水分胁迫响应程度存在一定差异。与河南及河北两种源相比，山西种源文冠果在各种梯度水分胁迫下均能保持相对较高的碳同位素比率（δ13C）值、水分利用效率（WUE）值以及较低的耗水量，表明山西种源文冠果在水分胁迫下对水分亏缺响应较为敏感，能通过短期内迅速减少耗水量来适应水分胁迫环境，这也是山西种源文冠果具有较高抗旱能力的体现。
（3）不同种源文冠果的叶片形态结构对水分胁迫响应程度存在一定差异，与河南及河北两种源相比，山西种源文冠果在各种胁迫时期具有较高的比叶重与较厚的各组织结构，且能形成较多的晶体以及小而多的导管等，结合耗水量最少的实验结果，表明山西种源文冠果更趋向于旱生结构。
（4）与河南及河北两种源相比，山西种源文冠果在各种梯度胁迫时期均能保持相对较高的水分利用效率和叶片保水能力以及较低的蒸腾速率、叶绿素含量和耗水量，表明山西种源文冠果具有较高的抗旱能力，是形态及解剖构造和水分生理生态特性对干旱环境适应性响应的结果。
（5）在不同梯度水分胁迫对文冠果苗木生长及生理特性和叶片形态解剖结构的分析研究的基础上，应用主成分分析和多维空间坐标综合评定值累加方法，建立文冠果抗旱节水综合评价指标体系，并对3种源文冠果苗木的抗旱性进行综合比较评价。结果发现，山西种源文冠果的抗旱性最强、河北种源次之、河南种源最差，这与线性回归模型验证结果以及水分胁迫下的叶片形态解剖结构和细胞水分利用特性的研究结果相一致。
以上研究结果表明，不同种源文冠果苗木生长、叶片的形态解剖结构以及生理特性对水分胁迫的响应程度存在一定差异，并表现出相应的适应性改变，进而导致其抗旱性生理功能的差异；对不同种源文冠果抗旱性进行综合评价，筛选出山西种源文冠果为抗旱性最强种源。这些研究结果可为我国干旱地区文冠果栽培以及优良种源筛选提供理论依据，具有重大的现实意义。
Problems of fossil energy depletion, environmental pollution and food security, have gonefrom bad to worse all over the world in recent years. Considerable attention has been paid tothe bio-diesel as renewable green energy. However, the provision and high cost of bio-dieselraw materials are the major barriers for its commercialization. Therefore, implementation ofthe bio-diesel industry must get a plentiful supply of raw materials. Use of woody bio-dieselfeedstock can reduce bio-diesel production cost and increase the supply. The investigation ofwoody oil plants suitable for non-arable land cultivation, development of bio-energy forest,and improvement of bio-diesel feedstock supply chain, is particularly important. Xanthocerasorbifolia Bunge has many advantages, including its wide distribution area, wide ecologicalamplitude, high seed oil, excellent fuel property, development and utilization of high value,and others. It was bio-diesel species promoted in northern China, and has enormous potentialfor development and utilization in recent years. In this paper, indoor potted water control testof3different X. sorbifolia Bunge provenances was worked out on the basis of systematicinvestigation, categorization of the resources distribution and biological characteristics research.We studied different gradients of drought stress on seedling growth, physiologicalcharacteristics, water use efficiency, water consumption law and its morphological anatomicalstructure of leaves, etc., and then explored X. sorbifolia of different provenances responsemechanism on water stress; Simultaneously, the X.sorbifolia resisting drought and saving watercomprehensive evaluation index system has been build, and the drought resistance of X.sorbifolia of different provenances was been comprehensive evaluated and screened. Theresults are as follows:
（1）The height and diameter growth of X. sorbifolia of three different provenancesshowed the trends which the growth of Shanxi provenance is the highest and the Henan is theworst under water stress conditions; the X. sorbifolia leaves of Hebei and Henan provenances appear yellow or fall off, and Shanxi provenance seedlings did not appear in the severe waterstress treatment.
With increasing the treatment time, water retention capacity of3provenances X.sorbifolia leaves was gradually reduced, and the chlorophyll content did not show significantdifferences. However, the X. sorbifolia leaves of Shanxi provenance maintained at a relativelyhigh water retention capacity; Water retention capacity of3provenances X. sorbifolia leaveshas been strengthened, and the chlorophyll content decreased under severe water stressconditions. Among them, water retention capacity of Shanxi provenance X. sorbifolia leaveshas the largest increases, remained higher after re-water, and has a relatively low transpirationrate. Regardless of their growing conditions or drought resistance, the X. sorbifolia Shanxiprovenances of was better than Henan and Hebei.
（2）The X. sorbifolia of different provenances response degree of stable carbon isotopefractionation, water use efficiency and water consumption on water stress exists a certaindifferences. Compared with the henan and hebei provenance, relatively high carbon isotoperatio value, water use efficiency and low water consumption of shanxi provenance X. sorbifoliacould maintain in all sorts of gradient water stress. This indicates the response of Shanxiprovenance is more sensitive to water deficit under drought stress, and to adapt to droughtstress environment by rapidly reducing water consumption in the short term, which was alsothe high drought tolerance expression of Shanxi provenance X. sorbifolia.
（3）There are some differences to water stress responsiveness among morphology ofleaves of different provenances X. sorbifolia. Compared with the Henan and Hebeiprovenances, X. sorbifolia of Shanxi provenances has higher specific leaf weight (SLW)andthicker organizations structures in various stress period, and can form more crystal sand vesselsof small and many and etc. Combined with experimental results of least water usage, X.sorbifolia of Shanxi provenances tend to xeromorphism.
（4）Compared with the Henan and Hebei provenances, X. sorbifolia of Shanxiprovenances can maintain a relatively higher water use efficiency (WUE), water-holding capacity and lower transpiration rate of leaf, chlorophyll content and water consumption invarious kinds of stress gradients period. These indicate X. sorbifolia of Shanxi provenances hashigher drought resistance capacity. That was the result of morphological and anatomicalstructure and water physiological ecology in response to drought environmental adaptability.
（5）On the basis of different gradients of water stress on seedling growth andphysiological property of X. sorbifolia and analysis of leaf morphological and anatomicalstructure, the drought resistance system of comprehensive evaluation of X. sorbifolia has beencreated by principal component analysis and comprehensive assessment of the value ofmulti-dimensional spatial coordinates accumulation method. And3X.sorbifolia provenanceshave been evaluated. It was found that X. sorbifolia of Shanxi provenances has strongestdrought resistance, followed Hebei’s, and Henan’s worst, consisting with the results of linearregression model validation and leaves morphological and anatomical structure under droughtstress and property of cell water-using.
The above findings suggest that there are some differences about seedling growth, leafmorphological and anatomical structures and physiological property of X. sorbifolia ofdifferent provenances in the response to drought stress, and exhibit appropriate adaptivechanges, leading to differences in their physiological functions of drought resistance.Comprehensively evaluating drought resistance of X.sorbifolia of different provenances, X.sorbifolia of Shanxi provenance has been screened as the provenances of strongest droughtresistance. These findings may provide a theoretical basis for X. sorbifolia cultivation in aridregions of China and excellent provenance screening. It is of great practical significance.
引文
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