摘要
沙冬青(Ammopiptanthus mongolicus(Maxim.)Cheng f.)是西北荒漠地区珍稀濒危的常绿阔叶灌木,为国家二级保护植物。本论文系统研究了沙冬青的群落生态学、种群生态学、及沙冬青种子的发芽生物学,为将来其它的后续研究奠定了良好的背景基础;因其为豆科植物,可与根瘤菌建立共生关系,根瘤菌对其生长发育和种群恢复有重要作用;因此调查沙冬青群落中根瘤的分布,收集并分离沙冬青及其伴生豆科植物的根瘤菌菌株,研究其生理抗性。该研究对沙冬青主要研究成果如下:
(1)综合沙冬青分布区内气候、土壤和水分等因子,选取内蒙古磴口县作为研究样地,并于样地内根据不同生境选取3个典型样点,研究了3个样点沙冬青群落的物种组成、群落结构、群落多样性等方面。在240个灌木样方和480个草本样方中共计录到灌木7科13属14种,多年生草本3科4属4种,一年生草本3科5属5种,最大的科是菊科(5种)和蒺藜科(5种),其次是豆科(3种),蓼科(2种)和禾本科(2种)居第三位。沙冬青群落植物组成成分因生境异质性而表现出很大的差异性。根据群落中植物成分和其在群落中的作用,对三个样地的沙冬青群落类型划分如下:Plot1群落属于暖温型草原化荒漠亚带;Plot2和Plot3群落属于典型荒漠亚带。沙冬青群落结构因生境异质性也表现出一定的差异性。Plot1群落3个亚层发育较好;Plot2和Plot3草本层没有发育,而且两个亚层成分单调。沙冬青的盖度、最大高度、最大幅度、平均高度和平均幅度与群落丰富度指数成负相关;而沙冬青的密度与丰富度成正相关,相关系数为0.508。用所选用的物种多样性指数(H′,D)、丰富度指数(S,R)和均匀度指数(J_(sw),J_(si),E)测度磴口县沙冬青植物群落的多样性,都反映出基本一致的趋势,其多样性顺序为:Plot1(沙冬青—刺旋花—沙葱)>Plot2(沙冬青—油蒿)>Plot3(沙冬青—旱蒿);均匀度指数则近于相反。
(2)采用相邻格子法收集数据,对沙冬青的高度和幅度进行分级和方差分析,研究了磴口县3个沙冬青种群的大小结构和年龄结构;并用矩阵法对沙冬青种群大小结构和群落物种多样性的关系进行了分析。结果表明,不同生境条件下的沙冬青种群的大小结构有差异,在高度和幅度上分别有极显著差异(P<0.01)和显著差异(P<0.05),但都呈现中间大两头小的态势,年龄结构呈现衰退趋势;群落的物种多样性水平与沙冬青种群的平均高度和幅度成负相关,物种间的竞争主要影响了沙冬青种群的整体高度和幅度,种群内个体间的竞争主要影响了影响了沙冬青种群内个体的高度和幅度大小的差异。
(3)对沙冬青的株高和冠幅进行分级,应用扩散系数C、负二次指数K、平均拥挤度m~*、丛生指数,、Cassie指数1/K、聚块性指数m~*/m和扩散型指数7种聚集度指标确定不同生境条件下沙冬青种群的空间分布格局类型和动态,考察沙冬青种群在不同尺度上的空间分布格局。结果表明:不同生境条件下的沙冬青种群结构有差异,但都呈现衰退趋势。不同生境的沙冬青种群空间分布格局类型和聚集强度不同,聚集强度在不同尺度上表现出基本一致的变化趋势。样地2在25和100 m~2的范围内集群分布,样地3在150 m~2范围内集群分布,样地1在7个取样面积下,均成随机分布。不同发育阶段种群的分布格局为:幼龄和老龄植株均成随机分布,中龄植株呈聚集分布。环境是沙冬青种群空间分布格局形成和发展的决定因子之一。
(4)为确定沙冬青最佳发芽温度,在15℃、28℃和37℃下进行发芽试验。结果发现,15℃下种子发芽延迟,37℃的高温使种子发芽率降低,且部分种子失活。28℃下种子发芽迅速整齐,胚根生长旺盛。然后测定了发芽过程中可溶性糖、糖组分、以及激素含量的变化。在28℃下,可溶性糖、糖组分、激素含量变化明显;激素在发芽4小时后出现峰值;20小时后多糖含量急剧下降,而四糖、二糖葡萄糖含量急剧上升,此时间与大部分胚根突破种皮的时间相近。28℃促进激素的活化和糖的代谢,有利于种子的发芽。
(5)综合土壤、水分条件,选定宁夏中卫沙坡头、内蒙古阿拉善左旗、内蒙古磴口县、内蒙古乌拉特后旗4个地区作为调查样区,研究了沙冬青植物群落和沙冬青根瘤的特征。沙冬青植物群落组成较为丰富;因水分、土壤类型及地形差异,不同样区内沙冬青群落成分、结构有一定变化;水分是沙冬青植物群落的决定性生态因子。沙冬青根瘤的最佳采集时间是在结果期之前,根瘤的外部形态呈现多样性,不同样区根瘤的着生部位有差异;水分是根瘤菌侵染沙冬青根系并形成根瘤的主要限制因子。沙冬青群落其他豆科植物的根瘤与沙冬青根瘤具有相似的外部形态。
(6)分离得到根瘤菌17株,对其耐盐性、耐酸碱性、生长温度范围和抗生素抗性进行了研究。结果表明,64.7%的菌株可以在含3%NaCl的YMA培养基上生长;94.1%的菌株可以在pH5~11的范围内生长;全部菌株在60℃处理10min后仍能生长;不同菌株对不同抗生素表现出不同的抗性,Zw_4和Wh_4~1对各抗生素表现出较强的抗性。分离自磴口县的根瘤菌普遍表现出较强的抗酸碱和抗高温的能力,这是对其环境的适应。
以Medicago truncatula为材料,鉴定出共生受体下游潜在的信号蛋白RopGEF,见附录。
Ammopiptanthus mongolieus(Maxim.) Cheng f.is an endangered species of evergreen broad-leaf plant in northwest desert zone of China.In this work,community ecology,population ecology,and germination biology of A.mongolicus were studied,providing good background and basic knowledge for the research in this plant in the future.As a legume specieses,A.mongolicus can construct a symbiotic relationship with rhizobium,which is helpful to its development and population recovering.With above novel information,the distribution of nodules in A.mongolicus was analyzed.Subsequently,we studied the resistance of rhizobium isolated from A.mongolicus. The main experimental results are as following:
(1) According to soil,water and other ecological factors,Dengkou was defined as research regions,where 3 research plots were selected according different habitats.Plot1 was on the bank of Yellow River,Plot2 was the old route of Yellow River,and Plot3 was on the east of Wulanbuhe Desert.Floristic composition,community structure and species diversity of A.mongolicus communities were studied in the research plots.There were 7 families,13 genera and 14 species shrubs,3 families,4 genera and 4 species perennial herbs,3 families,5 genera and 5 species annual herbs.The biggest families were Compositae(5 species) and Zygophyllaceae(5 species).The second families was Leguminosae(3 species),and the third families were Gramineae(2 species) and Polygonaceae(2 species).The floristic composition of A:mongolicus communities was different under different environmental conditions.According to the floristic composition and their function in the communities,the types of the 3 A.mongolicus communities were defined.The community of Plot1(A.mongolicus—Convolvulus tragacanthoides—Allium mongolicum) was belonged to warm-temperate steppe desert.The communities of Plot2(A. mongolicus—Artemisia ordosica) and of Plot3(A.mongolicus—Artemisia xerophytica) were belonged to typical desert.The community structure of the 3 A.mongolicus communities was also of dissimilitude under different environmental conditions.3 layers of Plot1 community developed well,but Plot2 and Plot3 communities hadn't herb layer,and floristic composition of their only 2 layers was very exiguous.The coverage(C),maximal height(Hmax),maximal width(Wmax), mean height(MH),and mean width(MW) of A.mongolicus were negatively correlated to richness index(S),while the destiny(D) positively correlated to S,and the coefficient of correlation was 0.508.Plant community diversity of A.mongolicus was estimated by species diversity index(H′, D),richness index(S,R) and evenness index(J_(sw),J_(st),E),which represented accordant trend.The order of diversity was Plot1>Plot2>Plot3,while the result of evenness was opposite.
(2) The population structure of A.mongolicus in Dengkou,Inner Mongolia was studied. According to the date which was obtained by contiguous grid quadrate method,the height and width of A.mongolicus were classified and analyzed by ANOVA method.The relation of population structure of A.mongolicus and species diversity of A.mongolicus community was analyzed with correlation matrix.The result indicated that the population structure of A. mongolicus was different under different environmental conditions.The mean individual height of different population were significantly different(P<0.01),while the mean width were markedly different(P<0.05).The age structure of the 3 A.mongolicus populations presented senescent type. The mean height(MH) and mean width(MW) of A.mongolicus population were negatively correlated to richness index(S) of A.mongolicus community and their coefficient of correlation were -0.994 and -1 in turn.The competition among species influenced the unitary population height and width,while the competition among individuals of A.mongolicus population influenced the height and width of individuals in the same population.
(3) According to the data,the height and width of A.mongolicus were classified.The results indicated that the population structures of A.mongolicus were different in diferent environmental conditions.The age structures of three A.mongolicus populations showed the decline trend.The spatial distribution patterns and pattern dynamics of A.mongolicus populations were studied by applying seven aggregate indices(C,K,m~*,I,1/K,m~*/m and I_δ)in different environmental conditions.And the spatial distribution pattern with diferent quadrat scale was examined.The results indicated that the spatial distribution pattern and aggregation intensity were different in diferent environmental conditions,while the tendency of pattern aggregation was generally paralle1.The figure of pattern scale and pattern intensity showed that plot 2 clumped in 25 and 100 m~2,and plot 3 clumped in 150 m~2,while plot 1 performed the pattern of random distribution in all quadrat scale.With the population age increased,the distribution pattern had a trend from random to clustering and finally to random.The young and old individuals performed the pattern of random distribution,while the individuals of middle age stage clumped.The environmental factors principally influenced the formation and development of the spatial distribution pattern of A.mongolicuus populations.
(4) In order to ascertain optimal temperature for germination,seeds were germinated under 15℃、28℃、37℃.It was found that seeds growed slowly under 15℃,while the temperature of 37℃had a detrimental effect on seedling growth,some seeds were less likely to survive in the germination test.Seeds could bourgeond quickly with strong radicel at 28℃.Subsequently the change of saccharide and hormone were tested duing seed germination.Saccharide and hormone changed intensively under 28℃.The level of disaccharide、tetrasaccharide and glucose in seeds increased intensively at 20h,while the peak value of IAA and cytokinin appeared at 4h.28℃is the best temperature for seed germination of A.mongolicus.
(5) According to soil,water and other ecological factors,we defined Shapotou,Alashan, Dengkou and Wulatehouqi as our research regions where we studied characters of A.mongolicus plant communities and nodules.The plant community was rich.The components and structure were different in different research regions because of the diversity of ecological factors.Water is the decisive ecological factor which influenced the components and structure of A.mongolicus. plant communities.Nodules morphology of A.mongolicus was various.The best time to collect nodules should be before fruit stage of its host.Nodules were in different root parts in different research regions.Water was the primary ecological factor which influenced the infection of rhizobia and the generation of nodules.Nodules isolated from other legumes had similar morphology with nodules of A.mongolicus.
(6) Seventeen rhizobia strains were isolated from A.mongolicus.It was found that nodules were various in their attachment mode,size,shape and color,which were related to the differences of their eco-environment.And water may be the principal influencing factor.Several biochemical characteristics were detected,including resistance to salt,acid-alkali,temperature variation and intrinsic antibiotics.The results indicated that 64.7%strains could tolerate NaCl stress at 3% concentration,94.1%strains could grow during pH 5-11,and all strains could grow after disposed at 60℃C for 10 min.Differences in resistance to different intrinsic antibiotics existed among strains, ZW_4 and Wh_4~1 had high resistance to different intrinsic antibiotics.Rhizobia strains from Dengkou had higher resistance to acid-alkali and temperature,which was the adaptation of rhizobia to its environment
Identify the potential signaling proteins in downstram of symbiotic receptors from legume model system Medicago truncatula(Appendix).
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