东北地区牡丹生态适应性及抗寒性研究
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摘要
牡丹(Paeonia Sect. Moutan DC)是原产中国的传统名花,被誉为“百花之王”,具有“繁荣昌盛、幸福美满”等文化寓意。牡丹经过一千多年的发展,在我国的自然栽培应用区不断扩大,在东北地区也表现出良好的推广应用前景。然而,在东北地区寒冷的特殊气候条件下,牡丹引种、栽培以及育种等方面的研究还相当缺乏,制约这一地区牡丹发展的一个技术瓶颈还无法突破。为此,本研究基于东北地区的自然生态条件,对东北地区现有的牡丹品种资源进行系统的调查,研究了牡丹的生理生态特性,提出牡丹抗寒评价指标,并对不同的品种群的生态适应性与抗寒性进行了对比分析,为牡丹寒地引种、栽培及其发展提供重要参考依据。主要研究内容包括：
1.从2007年9月至2011年3月间,采用实地调查与史料考证相结合,对东北地区牡丹引种及生态适应性进行了调查与研究,发现：1)东北地区牡丹引种地分布在N47°线以南,共计约200个品种和类型、2.7万余株；2)随着纬度的升高,东北地区牡丹生长发育物候期逐步延迟,枝叶衰老、休眠期逐步提前。纬度越高,生理活动时间越短；开花数量逐步减少,花型退化,以单瓣少瓣为主；枝条木质化逐渐变短,芽数减少；叶片形态指标变化不大；3)品种群间存在明显差异,紫斑牡丹有相对良好的适应性,在现有引种最北地(大庆)仍可露地自然越冬并开花；传统品种牡丹在长春以北地区需人工防护才能越冬后开花,多数品种冻害严重。
2.以长春牡丹园为试验点,对13个品种牡丹进行叶片形态指标和光合特性测定,结果表明：牡丹品种间及品种群间叶片比叶质量差异显著,传统品种牡丹叶大且厚,比叶质量高；紫斑牡丹叶片薄,比叶质量低,但叶绿素含量高。净光合速率(Pn)品种间有差异,但品种群间差异不显著；光合日变化显示少数品种有光合“午休”现象,多数品种为单峰曲线；气孔导度(Gs)、胞间二氧化碳浓度(Ci)、蒸腾速率(Tr)日变化结果与光合日变化吻合；品种群间水分利用效率(WUE)差异显著,紫斑牡丹有更高的水分利用效率,表明抗旱适应性好。
3.在牡丹越冬期间以长春牡丹园12个品种为试材,开展了牡丹自然越冬、低温胁迫下枝条电导率测定及半致死温度(LT50)计算、脯氨酸及多糖含量、枝条水分蒸腾及生长恢复观察试验。结果表明：自然越冬条件下,传统品种牡丹地上枝条全部枯死,仅有少数品种从地面下抽发新枝开花；而紫斑牡丹多数品种枝条存活率在60%以上,且能正常开花3朵以上。低温胁迫下相对电导率拟合Logistic方程,拟合度R2在0.90以上；LT50随外界温度变化而变化,紫斑牡丹品种在越冬期间LT50一直低于传统品种且差异显著；采用方程参数估算组织损害区间,表明牡丹抗寒性还有很大潜力；冷冻后恢复生长结果与方程推算LT50结果一致；首次取得牡丹枝条冬季含水量动态变化情况,明确了牡丹枝条冬季水分散失速率以及变化规律,初步解释“抽条”现象是由低温造成枝条组织受损,引起水分散失,进而造成枝条枯死,不是越冬期间枝条自身蒸腾造成的“抽条”；越冬期间枝条脯氨酸含量、可溶性多糖含量,生长期叶片膜透性测定,可以作为牡丹抗寒性的参考指标。牡丹抗寒能力的体现是个复杂的系统,受到自身遗传特性及外界环境因子的综合作用,以越冬期间最冷月份的枝条半致死温度为测定指标,判断牡丹抗寒性较为可靠。
总之,本研究表明东北地区牡丹引种生态适应性主导因子是极端低温；随纬度的升高牡丹生长发育积温降低,枝条木质化降低,开花质量和数量下降,需人工保护措施辅助栽培；光合特性能反映牡丹对光能利用率及光合产物积累对生长发育的影响；越冬期间采用枝条电导率测定,Logistic方程拟合估算LT50的方法评价抗寒性具有可靠性和可行性；牡丹生态适应性及抗寒性在品种(群)间存在差异,紫斑牡丹应是东北地区主要推广应用的品种和类型。
Tree peony (Paeonia Sect. Moutan DC) is Chinese traditional flower, and it originates from China, as " The king of flowers ", and connotes the meanings of "prosperity and happiness" and other cultural meanings. After more than 1,000 years of development, the application of natural cultivated area of Chinese tree peony is constantly expanding and also shows good prospect in the Northeast China. However, because of cold weather conditions in Northeast China, it still lacks of the research on introduction, cultivation and breeding of tree peony, so that the development of the regional tree peony can not break through a technical bottleneck. For this purpose, this research is based on the natural ecological conditions in Northeast China, carrying out a systematic investigation to the existing tree peony cultivars resources in Northeast region, studying the physiological and ecological characteristics of tree peony, putting forward the cold resistance evaluation index about tree peony, and conducting a comparative analysis on different cultivar groups about the ecological adaptability and cold resistance, providing an important reference for the introduction, cultivation and development of the tree peony. The main contents include:
1. From September 2007 to March 2011, thus study took the combination of field survey and historical textural, and conducted a survey and research on the introduction and the ecological adaptability of the tree peony in the Northeast China region. Then, this study found that:1) The introduction of Northeast China tree peony is distributed below latitude 47 degrees north, a total of about 200 cultivars and types,27,000 plants; 2) Increasing with the latitude, growth and development of Northeast China tree peony phenology period progressively delay (the phenology of growth and development of tree peony would progressively delaied), and foliage senescence, dormancy gradually advanced. The higher the latitude is, the shorter the time of physiological activity is, the number of flowers gradually reduces, floral type degradation/degenerates, mainly with simple flower and few petals of flower. Lignified branches are gradually shorten, the number of buds reduces. Little has changed on leaf morphological index; 3) Significant differences between the cultivar groups, and the flare mudan have relatively good adaptability. The existed introducing can naturally expose overwintering and flower in northernmost region (Daqing); traditional cultivars of tree peony need artificial protection to overwinter to the north of Changchun, most cultivars often suffer from seriously frozen injury.
2. Test point is located in Changchun Mudan Garden. This study chooses 13 cultivars to determine the leaf morphological index and photosynthetic features. The results show that:the leaf mass, among the tree peony cultivars, varieties of inter-group difference is significant, the traditional varieties of peony leaves are large and thick, higher specific leaf weight; flare mudan leaves are thin, lower specific leaf weight, but high chlorophyll content. Net photosynthetic rate (Pn) varieties are different, but the variety of inter-group difference is not significant; diurnal variation shows a small number of varieties of photosynthesis "midday depression", the majority of species as a single peak curve; stomatal conductance (Gs), intercellular carbon dioxide concentration (Ci), and transpiration rate (Tr) on the result of changes are consistent with changes in photosynthesis; water use efficiency between species groups (WUE) are significantly different. Flare mudan has higher utilization efficiency of water. It show good adaptability of drought resistance.
3. In the winter period, test points is located in Changchun Mudan Garden. The author selects 12 cultivars of tree peony for the test materials, carries out tree peony natural winter observation, measurement under low temperature stress branches and semi-lethal temperature conductivity (LT50) calculation. The polysaccharide content of proline is measured to determine the branch transpiration and growth recovery observed test. The results show that:In the natural winter conditions, the branches of traditional cultivars of tree peony on the ground are all dead, and only a few of cultivars grow from the ground under the branches and flowers, and flare peony survival rate of most cultivars is 60% or more branches, and can flower 3 or above normal. Relative conductivity under low temperature stresses on fitting Logistic equation, goodness of fit R2 above 0.90. LT50 changes with the outside temperature. During the winter LT50 of the cultivars of flare mudan has been lower than traditional cultivars, and the difference is significant. Tissue damage estimates range equation parameters, and shows that there is great potential for cold resistance of tree peony. Freezing peony recovers the growth and it consistents with the estimated LT50. For the first time i have got the dynamic changes of water content of tree peony branches in winter, knowing the changes of water of tree peony branches loss rate in winter. Preliminary interpretation of " Shoot drying" phenomenon is caused by the low-temperature branch tissue damage, leading to water loss, which is resulted into dead branches, not the branches themselves during winter transpiration caused by the " air-dried ". During the winter determined proline content and soluble polysaccharide content in shoots, leaf growth stage determines the membrane permeability, and can be used as reference for the evaluation tree peony cold resistance. It is a complex system that cold resistance capacity of the tree peony is expressed by their genetic characteristics and the combined effects of environmental factors. During the coldest month of winter, branches semi-lethal temperature for the determination of indicators determine the cold resistance of peony more reliable. Selected branches of the lethal temperature measured as an index to evaluate the cold resistance of tree peony is more reliable.
In summary, this study shows that extreme low temperature is the dominant factor on ecological adaptation of tree peony introduction to Northeast China. As latitude increases, effective temperature in need decreases, branches lignification become lower, flowering quality and quantity decline. Protection measures should be assisted by artificial cultivation. The photosynthetic characteristics of light energy utilization efficiency can reflect the accumulation of photosynthetic products and the impact on the tree peony growth and development; during the overwinter period, the author uses branches to measure the conductivity and fits the Logistic equation, then estimates LT50, and this method is reliable and feasible to evaluate cold resistance. There are the differences of the ecological adaptability and cold resistance in tree peony cultivars (group); flare mudan should be the main cultivars popularized and applied in Northeast China.
1.从2007年9月至2011年3月间,采用实地调查与史料考证相结合,对东北地区牡丹引种及生态适应性进行了调查与研究,发现：1)东北地区牡丹引种地分布在N47°线以南,共计约200个品种和类型、2.7万余株；2)随着纬度的升高,东北地区牡丹生长发育物候期逐步延迟,枝叶衰老、休眠期逐步提前。纬度越高,生理活动时间越短；开花数量逐步减少,花型退化,以单瓣少瓣为主；枝条木质化逐渐变短,芽数减少；叶片形态指标变化不大；3)品种群间存在明显差异,紫斑牡丹有相对良好的适应性,在现有引种最北地(大庆)仍可露地自然越冬并开花；传统品种牡丹在长春以北地区需人工防护才能越冬后开花,多数品种冻害严重。
2.以长春牡丹园为试验点,对13个品种牡丹进行叶片形态指标和光合特性测定,结果表明：牡丹品种间及品种群间叶片比叶质量差异显著,传统品种牡丹叶大且厚,比叶质量高；紫斑牡丹叶片薄,比叶质量低,但叶绿素含量高。净光合速率(Pn)品种间有差异,但品种群间差异不显著；光合日变化显示少数品种有光合“午休”现象,多数品种为单峰曲线；气孔导度(Gs)、胞间二氧化碳浓度(Ci)、蒸腾速率(Tr)日变化结果与光合日变化吻合；品种群间水分利用效率(WUE)差异显著,紫斑牡丹有更高的水分利用效率,表明抗旱适应性好。
3.在牡丹越冬期间以长春牡丹园12个品种为试材,开展了牡丹自然越冬、低温胁迫下枝条电导率测定及半致死温度(LT50)计算、脯氨酸及多糖含量、枝条水分蒸腾及生长恢复观察试验。结果表明：自然越冬条件下,传统品种牡丹地上枝条全部枯死,仅有少数品种从地面下抽发新枝开花；而紫斑牡丹多数品种枝条存活率在60%以上,且能正常开花3朵以上。低温胁迫下相对电导率拟合Logistic方程,拟合度R2在0.90以上；LT50随外界温度变化而变化,紫斑牡丹品种在越冬期间LT50一直低于传统品种且差异显著；采用方程参数估算组织损害区间,表明牡丹抗寒性还有很大潜力；冷冻后恢复生长结果与方程推算LT50结果一致；首次取得牡丹枝条冬季含水量动态变化情况,明确了牡丹枝条冬季水分散失速率以及变化规律,初步解释“抽条”现象是由低温造成枝条组织受损,引起水分散失,进而造成枝条枯死,不是越冬期间枝条自身蒸腾造成的“抽条”；越冬期间枝条脯氨酸含量、可溶性多糖含量,生长期叶片膜透性测定,可以作为牡丹抗寒性的参考指标。牡丹抗寒能力的体现是个复杂的系统,受到自身遗传特性及外界环境因子的综合作用,以越冬期间最冷月份的枝条半致死温度为测定指标,判断牡丹抗寒性较为可靠。
总之,本研究表明东北地区牡丹引种生态适应性主导因子是极端低温；随纬度的升高牡丹生长发育积温降低,枝条木质化降低,开花质量和数量下降,需人工保护措施辅助栽培；光合特性能反映牡丹对光能利用率及光合产物积累对生长发育的影响；越冬期间采用枝条电导率测定,Logistic方程拟合估算LT50的方法评价抗寒性具有可靠性和可行性；牡丹生态适应性及抗寒性在品种(群)间存在差异,紫斑牡丹应是东北地区主要推广应用的品种和类型。
Tree peony (Paeonia Sect. Moutan DC) is Chinese traditional flower, and it originates from China, as " The king of flowers ", and connotes the meanings of "prosperity and happiness" and other cultural meanings. After more than 1,000 years of development, the application of natural cultivated area of Chinese tree peony is constantly expanding and also shows good prospect in the Northeast China. However, because of cold weather conditions in Northeast China, it still lacks of the research on introduction, cultivation and breeding of tree peony, so that the development of the regional tree peony can not break through a technical bottleneck. For this purpose, this research is based on the natural ecological conditions in Northeast China, carrying out a systematic investigation to the existing tree peony cultivars resources in Northeast region, studying the physiological and ecological characteristics of tree peony, putting forward the cold resistance evaluation index about tree peony, and conducting a comparative analysis on different cultivar groups about the ecological adaptability and cold resistance, providing an important reference for the introduction, cultivation and development of the tree peony. The main contents include:
1. From September 2007 to March 2011, thus study took the combination of field survey and historical textural, and conducted a survey and research on the introduction and the ecological adaptability of the tree peony in the Northeast China region. Then, this study found that:1) The introduction of Northeast China tree peony is distributed below latitude 47 degrees north, a total of about 200 cultivars and types,27,000 plants; 2) Increasing with the latitude, growth and development of Northeast China tree peony phenology period progressively delay (the phenology of growth and development of tree peony would progressively delaied), and foliage senescence, dormancy gradually advanced. The higher the latitude is, the shorter the time of physiological activity is, the number of flowers gradually reduces, floral type degradation/degenerates, mainly with simple flower and few petals of flower. Lignified branches are gradually shorten, the number of buds reduces. Little has changed on leaf morphological index; 3) Significant differences between the cultivar groups, and the flare mudan have relatively good adaptability. The existed introducing can naturally expose overwintering and flower in northernmost region (Daqing); traditional cultivars of tree peony need artificial protection to overwinter to the north of Changchun, most cultivars often suffer from seriously frozen injury.
2. Test point is located in Changchun Mudan Garden. This study chooses 13 cultivars to determine the leaf morphological index and photosynthetic features. The results show that:the leaf mass, among the tree peony cultivars, varieties of inter-group difference is significant, the traditional varieties of peony leaves are large and thick, higher specific leaf weight; flare mudan leaves are thin, lower specific leaf weight, but high chlorophyll content. Net photosynthetic rate (Pn) varieties are different, but the variety of inter-group difference is not significant; diurnal variation shows a small number of varieties of photosynthesis "midday depression", the majority of species as a single peak curve; stomatal conductance (Gs), intercellular carbon dioxide concentration (Ci), and transpiration rate (Tr) on the result of changes are consistent with changes in photosynthesis; water use efficiency between species groups (WUE) are significantly different. Flare mudan has higher utilization efficiency of water. It show good adaptability of drought resistance.
3. In the winter period, test points is located in Changchun Mudan Garden. The author selects 12 cultivars of tree peony for the test materials, carries out tree peony natural winter observation, measurement under low temperature stress branches and semi-lethal temperature conductivity (LT50) calculation. The polysaccharide content of proline is measured to determine the branch transpiration and growth recovery observed test. The results show that:In the natural winter conditions, the branches of traditional cultivars of tree peony on the ground are all dead, and only a few of cultivars grow from the ground under the branches and flowers, and flare peony survival rate of most cultivars is 60% or more branches, and can flower 3 or above normal. Relative conductivity under low temperature stresses on fitting Logistic equation, goodness of fit R2 above 0.90. LT50 changes with the outside temperature. During the winter LT50 of the cultivars of flare mudan has been lower than traditional cultivars, and the difference is significant. Tissue damage estimates range equation parameters, and shows that there is great potential for cold resistance of tree peony. Freezing peony recovers the growth and it consistents with the estimated LT50. For the first time i have got the dynamic changes of water content of tree peony branches in winter, knowing the changes of water of tree peony branches loss rate in winter. Preliminary interpretation of " Shoot drying" phenomenon is caused by the low-temperature branch tissue damage, leading to water loss, which is resulted into dead branches, not the branches themselves during winter transpiration caused by the " air-dried ". During the winter determined proline content and soluble polysaccharide content in shoots, leaf growth stage determines the membrane permeability, and can be used as reference for the evaluation tree peony cold resistance. It is a complex system that cold resistance capacity of the tree peony is expressed by their genetic characteristics and the combined effects of environmental factors. During the coldest month of winter, branches semi-lethal temperature for the determination of indicators determine the cold resistance of peony more reliable. Selected branches of the lethal temperature measured as an index to evaluate the cold resistance of tree peony is more reliable.
In summary, this study shows that extreme low temperature is the dominant factor on ecological adaptation of tree peony introduction to Northeast China. As latitude increases, effective temperature in need decreases, branches lignification become lower, flowering quality and quantity decline. Protection measures should be assisted by artificial cultivation. The photosynthetic characteristics of light energy utilization efficiency can reflect the accumulation of photosynthetic products and the impact on the tree peony growth and development; during the overwinter period, the author uses branches to measure the conductivity and fits the Logistic equation, then estimates LT50, and this method is reliable and feasible to evaluate cold resistance. There are the differences of the ecological adaptability and cold resistance in tree peony cultivars (group); flare mudan should be the main cultivars popularized and applied in Northeast China.
引文
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