华北驼绒藜种群生殖生态学研究
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摘要
华北驼绒藜属于藜科驼绒藜属旱生雌雄同株异花半灌木,为中国特有植物种,主要分布在内蒙古中东部和华北地区,其营养丰富,抗逆性强,是生态建设和改良草地的优良植物。近年来其种群严重退化和消失,部分种群退化率高达70%以上。然而,目前国内外有关该物种的生殖特性及其种群退化机制的研究还很少。基于此,本研究对分布于内蒙古野生华北驼绒藜种群进行生殖生态学方面的研究,包括种群动态特征、传粉生态学特征、胚胎发育及生理学特征、种子特性、生殖分配、遗传多样性等内容,主要结果如下:
1.种群数量动态研究表明,多数华北驼绒藜种群幼龄级个体数较少,中老龄级个体数较大,纺锤型的大小级结构表明种群属于衰退型。种群生命表和存活曲线的分析表明,不同华北驼绒藜种群尽管生境条件差异,但存活曲线基本接近DeeveyⅢ型,不同种群偏离典型存活曲线的程度与幼苗缺乏程度有关。对种群数量的时间序列预测显示,随着时间的推移,不同华北驼绒藜种群均会呈现老龄级株数先增加后减少的趋势,缺少幼龄株数补充是共同特征。
2.传粉生态学特征研究表明,华北驼绒藜具有典型的适应风媒传粉的花部特征和性状。华北驼绒藜种群花期较长表现为“渐进式集中开花”的开花模式;雄花单株花期18～21d,雌花20～26d;雄花单花花期1～2d,雌花6～8d,不同种群间差异不显著。华北驼绒藜的花粉活力高,可以持续6d;雌花柱头可授性6～7d,可授期滞后于散粉期。华北驼绒藜主要以风媒传粉为主,传粉效率对结实率影响较大。
3.胚胎发育及生理学特性研究表明,华北驼绒藜胚胎发育方式为藜型,人工栽培华北驼绒藜成熟胚分化时间(21d～23d)短于野生华北驼绒藜(26d～30d),球形胚以后的发育阶段出现败育现象。华北驼绒藜胚胎发育过程中过氧化物酶和酸性磷酸酯酶活性从总体上呈持续增强的趋势,不同生境华北驼绒藜种群间该酶活性无明显差异。硝酸还原酶活性呈先上升后下降的趋势,在胚胎发育前期人工栽培种群种子硝酸还原酶活性极显著高于野生种群。华北驼绒藜胚胎发育过程中人工栽培种子IAA、GA3和ZR含量明显高于野生种群种子;种子的(IAA+GA3+ZR)/ABA比值从总体上呈胚胎发育前期高,而后期较低的趋势,且胚胎发育前期人工栽培种子(IAA+GA3+ZR)/ABA比值要显著高于野生种群种子。胚胎发育的各个时期,野生种群种子营养成分(全氮、全磷、可溶性糖)含量均低于人工栽培种群。
4.种子特性研究表明,华北驼绒藜种子特性在种群内和种群间均存在广泛变异,种群内的方差分量均大于种群间,表明华北驼绒藜种子性状受到较强的遗传控制。种子形态特征间有极显著相关性(P<0.01);萌发率、萌发指数和活力指数间有极显著相关性(r>0.836,P<0.01)且与结实率有显著相关性(r>0.896,P<0.01)。种子性状与地理气候因素相关性表现为,种子背毛长与海拔,降雨量,种子宽与温度、降雨量有显著相关(r>0.801,P<0.01)。
5.生殖分配特征研究表明,从蕾期、花期到结实期,华北驼绒藜不同种群生物量、全氮、全磷、全钾生殖分配逐渐增加。在蕾期和花期生物量、全氮、全磷、全钾的分配均是以茎和叶占较大比例,结实期以生殖器官占优势比例。华北驼绒藜以生物量、全氮、全磷、全钾的生殖分配与植株大小的变化趋势基本一致,可划分为幼龄生殖期(径级1～2cm)、过渡生殖期(2～3cm)、稳定生殖期(3～7cm)和生殖衰退期(>7cm)4个时期。对种群生殖分配有重要影响的14个环境因子的主成分分析表明,海拔、坡向、群落植被盖度、华北驼绒藜盖度、土壤厚度、年蒸发量、年降雨量和人为干扰度是影响种群生殖分配的重要因素。运用生殖分配的有关学说对华北驼绒藜生殖分配格局进行了解释,生境稳定性学说和生活史理论假说可以全部和部分解释华北驼绒藜的生殖分配格局。
6.遗传多样性研究表明,华北驼绒藜的遗传多样性水平很高。在物种水平上,多态位点百分率(PPL)为98.05%,Nei′s指数(h)为0.298,Shannon信息指数(I)为0.456;在种群水平上,PPL、h和I分别平均为80.62%、0.268和0.403。基于Nei′s遗传多样性指数和AMOVA分析均表明华北驼绒藜种群的遗传变异主要存在于种群内。结合聚类分析和地理变异规律把种群划分为两个大的种群组:荒漠化草原种群组和典型草原种群组。
Ceratoides arborescens belongs to Chenopodiaceae, Ceratoides genera, monoecism subshrub. It is endemic to China and mainly distributed in Mid-Eastern Inner Mongolia and North China. It has rich nutrition and strong resistance and is an important plant that is used for constructing ecology and improving grassland. Up to date, wild C. arborescens population degraded seriously, some population degraded up to 70%.However, the study on the reproduction peculiarity and degraded mechanism of this species is still unknown.Therefore,the paper focuses on reproductive ecology of C.arborescens population in population dynamics, pollination ecology, embryonic development and physiology, seed characteristics,reproductive allocation and genetic diversity,the main results were as follows:
1. The analysis on the age structure of C.arborescens populations showed that there were fewer young individuals, but middle-aged and old individuals were relatively rich. The analysis on the life tables and survival curves showed that the survival curve of C.arborescens populations belonged to DeeveyⅢ, and the death peak of different populations was in the period ofⅧandⅨ.The number difference among populations reflected the population habitat. Time sequence prediction indicated the numbers of old individuals would be increased at the beginning, and decreased finally in some years later. It was difficult to maintain the population stability.
2. The study of pollination ecology showed that C.arborescens adapted to anemophily. The duration of flowering in a population lasted 45～52 days; male flower male flower 18～21 and female flower lasted 20～26d at individual plant level; flower male flower 1～2d and female flower 6～8d at individual flower level, and there were no significant differences among populations. The pollen viability and stigmatic receptivity could last 6d and 6 to 7d respectively. The effective pollination way was wind and the efficiency of pollination had significant effect on seed setting rate.
3. The embryo development speed of the cultivate C.arborescens is faster than the feral C.arborescens and the embryo development modes are both chenopodiad; There are abortion phenomenons in feral C.arborescens embryo development. The peroxidase and acpase activity in C. arborescens had a tendency of ascent with the embryo development. The nitrate reductase activity during early period of embryo development increased significantly and then decreased, and that in cultivate C.arborescens seeds was lower significantly than feral C.arborescens seeds. The contents of IAA,GA3 and ZR in cultivate C.arborescens seeds were higher than two types of feral C.arborescens seeds; The ratio of (IAA+GA3+ZR)/ABA in two types of C. arborescens seeds was high during the early period of embryo development, and then had a tendency of descent with the development of embryo, and the ratio in cultivate C.arborescens seeds was higher significantly than two types of feral C.arborescens seeds. During the embryo development, the nutrient(N,P,Soluble sugar) contents of the cultivate C.arborescens is higher than the feral C.arborescens.
4. The seed characteristics showed that there were significant differences among population and among individual plants within populations.The variance factors analysis indicated that the variation within a population was the main part of the phenotypic variation. The morphological traits of seed had significant positive correlation with each other (P<0.01). The pappus length and latitude and rainfall, as well as, between seed width and temperature and rainfall had significant correlations(r>0.801, P<0.01). The three germination-related indices had significant positive correlations with each other (r > 0.836, P<0.01) and with percentage of seed fill(r>0.896, P<0.01), however, possessing weak correlations with morphological traits.
5. The characteristics of reproductive allocation showed that reproductive allocation of biomass, total nitrogen, total phosphorus,total potassium in C.arborescens increased gradually from the flower bud phase to the flower and fruit phase.Allocation of stem and leaf took advantage ratio in the flower bud and the flower phase, and reproductive organs took advantage ratio in fruit phase. The reproductive age of C. arborescens can be divided into four stages in terms of the relationship between the reproductive allocation of biomass and the age of the reproductive individuals. These are the junior, non-stable, stable and declining phases. The most important components to affect pattern and development of C.arborescens population were altitude, slope face, vegetation cover, C. arborescens cover, thickness of soil, annual mean precipitation, annual evaporation and intensity of human disturbance. Three hypotheses were evaluated with regard to the observed patterns of reproductive allocation.One hypothesis, that reproductive allocation patterns were a function of differences in resource availability, did not provide an explanation for our results. A second hypothesis, that reproductive allocation was negatively correlated with successional maturity of the habitat could explain the observed patterns of reproductive allocation. The third, life history hypothesis which assumed the existence of a trade off between current reproduction and vegetative growth and/or survival rate, could explain patterns of reproductive allocation in part.
6. The research of genetic diversity showed that at species level, percentage of polymorphic loci PPB is 98.05%, Nei′s gene diversity is 0.298, and Shannon′s information index is 0.456.At population level, PPB is 80.62%, Nei′s gene diversity is 0.268 and Shannon′s information index is 0.403. C.arborescens population had an extraordinarily high level of genetic diversity. The analysis of molecular variance (AMOVA) showed that 86.79% of genetic variance was found within populations, and 13.21% of it was resided within populations. The coefficient of gene differentiation (Gst) was 0.1035.The level of genetic differentiation between populations is lower than that among populations. According to cluster analysis and the law of geographic variation, the populations were classified into two large groups: desert grassland group and typical grassland group.
1.种群数量动态研究表明,多数华北驼绒藜种群幼龄级个体数较少,中老龄级个体数较大,纺锤型的大小级结构表明种群属于衰退型。种群生命表和存活曲线的分析表明,不同华北驼绒藜种群尽管生境条件差异,但存活曲线基本接近DeeveyⅢ型,不同种群偏离典型存活曲线的程度与幼苗缺乏程度有关。对种群数量的时间序列预测显示,随着时间的推移,不同华北驼绒藜种群均会呈现老龄级株数先增加后减少的趋势,缺少幼龄株数补充是共同特征。
2.传粉生态学特征研究表明,华北驼绒藜具有典型的适应风媒传粉的花部特征和性状。华北驼绒藜种群花期较长表现为“渐进式集中开花”的开花模式;雄花单株花期18～21d,雌花20～26d;雄花单花花期1～2d,雌花6～8d,不同种群间差异不显著。华北驼绒藜的花粉活力高,可以持续6d;雌花柱头可授性6～7d,可授期滞后于散粉期。华北驼绒藜主要以风媒传粉为主,传粉效率对结实率影响较大。
3.胚胎发育及生理学特性研究表明,华北驼绒藜胚胎发育方式为藜型,人工栽培华北驼绒藜成熟胚分化时间(21d～23d)短于野生华北驼绒藜(26d～30d),球形胚以后的发育阶段出现败育现象。华北驼绒藜胚胎发育过程中过氧化物酶和酸性磷酸酯酶活性从总体上呈持续增强的趋势,不同生境华北驼绒藜种群间该酶活性无明显差异。硝酸还原酶活性呈先上升后下降的趋势,在胚胎发育前期人工栽培种群种子硝酸还原酶活性极显著高于野生种群。华北驼绒藜胚胎发育过程中人工栽培种子IAA、GA3和ZR含量明显高于野生种群种子;种子的(IAA+GA3+ZR)/ABA比值从总体上呈胚胎发育前期高,而后期较低的趋势,且胚胎发育前期人工栽培种子(IAA+GA3+ZR)/ABA比值要显著高于野生种群种子。胚胎发育的各个时期,野生种群种子营养成分(全氮、全磷、可溶性糖)含量均低于人工栽培种群。
4.种子特性研究表明,华北驼绒藜种子特性在种群内和种群间均存在广泛变异,种群内的方差分量均大于种群间,表明华北驼绒藜种子性状受到较强的遗传控制。种子形态特征间有极显著相关性(P<0.01);萌发率、萌发指数和活力指数间有极显著相关性(r>0.836,P<0.01)且与结实率有显著相关性(r>0.896,P<0.01)。种子性状与地理气候因素相关性表现为,种子背毛长与海拔,降雨量,种子宽与温度、降雨量有显著相关(r>0.801,P<0.01)。
5.生殖分配特征研究表明,从蕾期、花期到结实期,华北驼绒藜不同种群生物量、全氮、全磷、全钾生殖分配逐渐增加。在蕾期和花期生物量、全氮、全磷、全钾的分配均是以茎和叶占较大比例,结实期以生殖器官占优势比例。华北驼绒藜以生物量、全氮、全磷、全钾的生殖分配与植株大小的变化趋势基本一致,可划分为幼龄生殖期(径级1～2cm)、过渡生殖期(2～3cm)、稳定生殖期(3～7cm)和生殖衰退期(>7cm)4个时期。对种群生殖分配有重要影响的14个环境因子的主成分分析表明,海拔、坡向、群落植被盖度、华北驼绒藜盖度、土壤厚度、年蒸发量、年降雨量和人为干扰度是影响种群生殖分配的重要因素。运用生殖分配的有关学说对华北驼绒藜生殖分配格局进行了解释,生境稳定性学说和生活史理论假说可以全部和部分解释华北驼绒藜的生殖分配格局。
6.遗传多样性研究表明,华北驼绒藜的遗传多样性水平很高。在物种水平上,多态位点百分率(PPL)为98.05%,Nei′s指数(h)为0.298,Shannon信息指数(I)为0.456;在种群水平上,PPL、h和I分别平均为80.62%、0.268和0.403。基于Nei′s遗传多样性指数和AMOVA分析均表明华北驼绒藜种群的遗传变异主要存在于种群内。结合聚类分析和地理变异规律把种群划分为两个大的种群组:荒漠化草原种群组和典型草原种群组。
Ceratoides arborescens belongs to Chenopodiaceae, Ceratoides genera, monoecism subshrub. It is endemic to China and mainly distributed in Mid-Eastern Inner Mongolia and North China. It has rich nutrition and strong resistance and is an important plant that is used for constructing ecology and improving grassland. Up to date, wild C. arborescens population degraded seriously, some population degraded up to 70%.However, the study on the reproduction peculiarity and degraded mechanism of this species is still unknown.Therefore,the paper focuses on reproductive ecology of C.arborescens population in population dynamics, pollination ecology, embryonic development and physiology, seed characteristics,reproductive allocation and genetic diversity,the main results were as follows:
1. The analysis on the age structure of C.arborescens populations showed that there were fewer young individuals, but middle-aged and old individuals were relatively rich. The analysis on the life tables and survival curves showed that the survival curve of C.arborescens populations belonged to DeeveyⅢ, and the death peak of different populations was in the period ofⅧandⅨ.The number difference among populations reflected the population habitat. Time sequence prediction indicated the numbers of old individuals would be increased at the beginning, and decreased finally in some years later. It was difficult to maintain the population stability.
2. The study of pollination ecology showed that C.arborescens adapted to anemophily. The duration of flowering in a population lasted 45～52 days; male flower male flower 18～21 and female flower lasted 20～26d at individual plant level; flower male flower 1～2d and female flower 6～8d at individual flower level, and there were no significant differences among populations. The pollen viability and stigmatic receptivity could last 6d and 6 to 7d respectively. The effective pollination way was wind and the efficiency of pollination had significant effect on seed setting rate.
3. The embryo development speed of the cultivate C.arborescens is faster than the feral C.arborescens and the embryo development modes are both chenopodiad; There are abortion phenomenons in feral C.arborescens embryo development. The peroxidase and acpase activity in C. arborescens had a tendency of ascent with the embryo development. The nitrate reductase activity during early period of embryo development increased significantly and then decreased, and that in cultivate C.arborescens seeds was lower significantly than feral C.arborescens seeds. The contents of IAA,GA3 and ZR in cultivate C.arborescens seeds were higher than two types of feral C.arborescens seeds; The ratio of (IAA+GA3+ZR)/ABA in two types of C. arborescens seeds was high during the early period of embryo development, and then had a tendency of descent with the development of embryo, and the ratio in cultivate C.arborescens seeds was higher significantly than two types of feral C.arborescens seeds. During the embryo development, the nutrient(N,P,Soluble sugar) contents of the cultivate C.arborescens is higher than the feral C.arborescens.
4. The seed characteristics showed that there were significant differences among population and among individual plants within populations.The variance factors analysis indicated that the variation within a population was the main part of the phenotypic variation. The morphological traits of seed had significant positive correlation with each other (P<0.01). The pappus length and latitude and rainfall, as well as, between seed width and temperature and rainfall had significant correlations(r>0.801, P<0.01). The three germination-related indices had significant positive correlations with each other (r > 0.836, P<0.01) and with percentage of seed fill(r>0.896, P<0.01), however, possessing weak correlations with morphological traits.
5. The characteristics of reproductive allocation showed that reproductive allocation of biomass, total nitrogen, total phosphorus,total potassium in C.arborescens increased gradually from the flower bud phase to the flower and fruit phase.Allocation of stem and leaf took advantage ratio in the flower bud and the flower phase, and reproductive organs took advantage ratio in fruit phase. The reproductive age of C. arborescens can be divided into four stages in terms of the relationship between the reproductive allocation of biomass and the age of the reproductive individuals. These are the junior, non-stable, stable and declining phases. The most important components to affect pattern and development of C.arborescens population were altitude, slope face, vegetation cover, C. arborescens cover, thickness of soil, annual mean precipitation, annual evaporation and intensity of human disturbance. Three hypotheses were evaluated with regard to the observed patterns of reproductive allocation.One hypothesis, that reproductive allocation patterns were a function of differences in resource availability, did not provide an explanation for our results. A second hypothesis, that reproductive allocation was negatively correlated with successional maturity of the habitat could explain the observed patterns of reproductive allocation. The third, life history hypothesis which assumed the existence of a trade off between current reproduction and vegetative growth and/or survival rate, could explain patterns of reproductive allocation in part.
6. The research of genetic diversity showed that at species level, percentage of polymorphic loci PPB is 98.05%, Nei′s gene diversity is 0.298, and Shannon′s information index is 0.456.At population level, PPB is 80.62%, Nei′s gene diversity is 0.268 and Shannon′s information index is 0.403. C.arborescens population had an extraordinarily high level of genetic diversity. The analysis of molecular variance (AMOVA) showed that 86.79% of genetic variance was found within populations, and 13.21% of it was resided within populations. The coefficient of gene differentiation (Gst) was 0.1035.The level of genetic differentiation between populations is lower than that among populations. According to cluster analysis and the law of geographic variation, the populations were classified into two large groups: desert grassland group and typical grassland group.
引文
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