33 resultados para Juniperus przewalskii
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Pharmacologically active xanthone compounds isolated from Swertia przewalskii pissjauk were well separated by capillary electrophoresis (CE) within 5 min, using a running buffer of 25 mM disodium tetraborate at pH 9.0. Quantitative determination was shown to be possible because regression equations revealed a linear relationship between the peak area of each constituent and its concentration, with correlation coefficients of 0.9972-0.9994. The relative standard deviations were between 0.44%-0.73% for migration times and 2.52%-4.28% for peak areas. The dissociation constant of 1,7-O-beta-D-glucopyranosyl-8-hydroxy-3,7-dimethoxyxanthone, 1,8-dihydroxy-3, 7-dimethoxy-xanthone and 1,7-dihydroxy-3,8-dimethoxyxanthone were also measured by the CE method, giving a value of 9.04, 8.94, and 8.59, respectively.
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根据中国麻黄属植物分类及麻黄属起源和演化研究中存在的争论,本论文主要进行了分类学、形态学、微形态学、解剖学、形态发生学、孢粉学、古植物学、植物生态学和植物地理学等方面的研究。 1、分类学 从性状变异式样的分析入手,结合标本馆工作和野外调查,确认了中国麻黄属植物共有13个种;合并了丽江麻黄(Ephedra likiangensis)、雌雄麻黄(E. fedtschenkoae);澄清了前人在细子麻黄(E. regeliana)、灰麻黄(E. glauca)、草麻黄(E. sinica)和双穗麻黄(E. distachya)等种类鉴定中混乱的问题,并描述了一个新种——日土麻黄(Ephedra rituensis Y. Yang, D. Z. Fu et G. Zhu)。 2、形态学 提供了中国麻黄属种类的标本(包括模式标本)照片资料及中国麻黄属植物的雌球花和雄球花的照片资料,为麻黄属分类和系统学研究提供了证据。 3、微形态学 首次报道了中国麻黄属植物的种子表面微形态特征,聚合囊表面纹饰、小枝表面特征。 种子表面特征可分为4种类型。类型I:种子表面平滑,包括:中麻黄(Ephedra intermedia)、双穗麻黄、细子麻黄等;类型II:种子表面具疣状突起,仅发现木贼麻黄(E. equisetina)一种;类型III:种子表面具横向片层状突起,仅有斑子麻黄(E. rhytidosperma)一种;类型IV:种子表面具纵条纹,包括喜马拉雅特有的3个大种子的种类,即矮麻黄(E. minuta)、山岭麻黄(E. gerardiana)和藏麻黄(E. saxatilis)。 对小枝表面扫描电镜下的微形态构造的研究并没有发现前人报道的毛状体特征,而只有一些疣状角质突起和有变异的气孔,同时注意到,麻黄属植物的小枝表面特征受到生境、发育阶段的影响。 麻黄属聚合囊顶孔开口处具脑波状纹饰,而聚合囊表面纹饰具二级纹饰。麻黄属聚合囊表面纹饰非常独特,为该属在地层中雄性生殖构造的发现提供了可参考的性状。 4、孢粉学 补充了麻黄属中非常特化的种——斑子麻黄的孢粉资料,同时也对膜果麻黄(Ephedra przewalskii)、中麻黄、细子麻黄和木贼麻黄的孢粉重新进行了研究。结果表明斑子麻黄的花粉形态也非常特殊,其纵肋数目5条左右,纵肋波状扭曲,肋间沟槽内透明线发达且分枝。膜果麻黄的花粉与前人描述的一致,但是,中麻黄、细子麻黄和木贼麻黄的花粉形态可能受到发育的影响而与前人报道中描述的有些差异。讨论了麻黄属4种花粉类型的演化趋势。 5、形态发生学 对麻黄属雌球花三种代表类型的个体发育研究表明,三胚珠球花发育早期球花主轴的顶芽发育,双胚珠球花的顶芽没有发育,而在单胚珠球花中,雌性生殖单位直接占据顶芽的位置发育。这三种类型也代表了麻黄属球花演化的三个阶段,三胚珠球花代表麻黄属雌球花的原始式样,而单胚珠雌球花则为麻黄属雌球花最为特化的式样。总之,麻黄属雌球花有简化的趋势,伴随着这种趋势,雌球花苞片数目减少、种子数目减少、种子体积增大、表面复杂度增加。 6、解剖学 对斑子麻黄种子表面微构造的形态发生及解剖研究表明,斑子麻黄雌性生殖单位发育早期,其表面为平滑型,因此,突起为次生的。这些突起是由雌性生殖单位外盖被表皮细胞的外壁向外突出形成,这种突出生长常常在相邻细胞之间同时发生,因此,出现片层状构造。石蜡切片显示,这些突起在发育早期有完整的细胞核、细胞质等结构,但在种子发育晚期和成熟后,这些外壁突起的细胞成为空腔隙,本文对这种构造的生态学意义进行了分析。 7、古植物学 本文描述了早白垩麻黄科的2个大化石种。 古斑子麻黄(Ephedra archaeorhytidosperma Y. Yang, B. Y. Geng et D. L. Dilcher et Z. D. Chen)发现于辽西义县组尖山沟层,时代为早白垩的芭雷姆期。该种的雌球花具2对苞片、1或2枚种子、珠孔管短直或顶部稍弯、种子表面具横列片层状纹饰等特征而与现存的斑子麻黄特征最为相近。二者之间如此多的相似之处只有一种解释,那就是它们曾经有过共同祖先。因此,该种应归属于麻黄属的肉苞组。古斑子麻黄雌球花特征的稳定性同时也表明,在麻黄属中也存在所谓的“形态演化停滞现象(morphological stasis)”,而这种现象也是首次在买麻藤类植物中报道。 星学异麻黄(Alloephedra xingxueii J. R. Tao et Y. Yang)产于吉林大拉子组,时代属早白垩的阿普特期——阿尔布期。该种雌球花具2对苞片、2枚种子,种子长椭圆形且具短的珠孔管与现存麻黄科植物最为相近,但是,其分枝式样与麻黄属不同,因此,本文在麻黄科中建立了一新属新种。此外,麻黄科植物常被看作是干旱区植被的代表,但是,对星学异麻黄当时所处的环境研究发现,大拉子组当时具有较明显的亚热带气候条件下的植被性质,因此,星学异麻黄即使需要干旱的条件,可能也只是局部小生境的条件。 8、生态学和植物地理学 本文首次对麻黄属植物生存环境中的水热条件进行了调查。从所选择的8个分布点来看,除了拉萨外,在麻黄属生殖生长期,其余7个分布点的湿度均是一年中最底的时期,而温度在这个时期逐渐升高。 对各地区内麻黄属种类的分布情况的分析发现,麻黄属的种类主要集中在三个地区,一是我国西北、前苏联的中亚地区及邻近地区,二是美国西南部、墨西哥西北部的地区,三是南美的智利、阿根廷等地。
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麻黄属(Ephedra)起源较早,该属植物的形态性状极度退化或简化,可用于经典分类学的有价值的性状非常有限。分布于青藏高原的藏麻黄(E. saxatilis)和丽江麻黄(E. likiangensis)二者间形态相似,性状变异连续,很难分辨,但被《中国植物志》中、英文版作两个种处理。 本文对藏麻黄和丽江麻黄的七个居群、151个个体的叶、雌球花和节间长等形态学性状进行了分析,发现各性状的变异情况在群体间无明显差异。同时我们还对154个个体的叶绿体DNA trnT-trnF和 trnS-trnfM区进行了序列分析,两个片段的联合矩阵长1382bp,共有29个变异位点,其中有9个碱基变异和2个indel,可划分为H1、H2和H3三种单倍型。这3种单倍型在丽江麻黄中均有分布,但藏麻黄仅含H1和H2。 综合来自形态学和分子方面的证据,我们发现藏麻黄和丽江麻黄的关系非常近缘,因此建议予以合并。同时本文还以膜果麻黄(E. przewalskii)为外类群,从谱系生物地理学角度探讨了三种叶绿体单倍型的进化关系,发现H2最原始,分布最广;H1与其它两种单倍型间的序列差异较大,可能是较进化的类型。此外,无性克隆的繁殖方式可能是导致Ephedra单倍型非常简单的重要原因之一。
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树木年轮的生长与气候因子、立地环境条件以及树木自身生长节律等密切相关。一方面,干旱半干旱区的树木年轮宽窄能够比较真实地记录过去的气候变化,据此,可以利用树木年轮资料重建干旱半干旱区过去几百年甚至上千年的气候变化。另一方面,立地环境条件也可能改变年轮生长与气候因子之间的响应关系。 有“世界屋脊”之称的青藏高原地区的气候变化与亚洲季风及全球气候系统密切相关。青藏高原也是世界上地形最复杂的高原,具有极其复杂的地形、地貌。因此,在青藏高原尤其是其东北部的干旱半干旱区开展树木年轮气候学及生态学研究具有极其重要的意义。本研究的目标就是利用树木年轮生态学方法研究树龄、立地条件等对青藏高原东北部祁连圆柏树木年轮宽度生长的可能影响。 为实现以上目标,本研究拟订了以下主要研究内容:探究祁连圆柏树种的年轮宽度生长对气候的响应关系中的年龄问题;比较每个样方内的个体径向生长变化以及三个样方内祁连圆柏个体径向生长规律;探讨导致个体径向生长差异的可能原因。 本文的研究材料为青藏高原常见的长寿树种祁连圆柏,研究方法是国际上通用的树木年轮学研究方法和手段。主要结果归纳如下: 1)依据国际树木年轮库的标准,建立了青藏高原东北部包括德令哈、乌兰、香日德、油葫芦、青羊沟、芒扎、东沟七个采样点的树木年轮年表。 2)运用响应函数、相关函数及单因素方差分析等方法分析了年龄因素对树木年轮生长与气候关系的可能影响。结果为:年龄在200-500年之间的祁连圆柏径向生长对气候变化的响应关系比较好。幼树对气候变化的响应相对差。而当树龄大于500年时,尽管由于树木内部生理变化,对气候变化的响应相对减弱,但是单因素方差分析结果显示这种减弱未能达到显著水平。因此,总体而言,我们认为长寿的祁连圆柏树种是重建古气候的理想材料。 3)通过设立样方,分析了样方内个体径向生长之间的相关性和差异性,以及生长释放与抑制事件发生的规律。同时还探讨了树间距离、立地微环境、限制性气候因子等对树木年轮生长的可能影响。主要研究结果:青藏高原东北部祁连圆柏年轮宽度序列中生长抑制事件发生的频率相对生长释放事件低很多,且存在个体间的差异;不同样方内个体径向生长之间的相关性有差别,乌兰样方内单棵树年轮宽度指数序列之间的相关性比香日德样方以及德令哈样方内单棵树年轮宽度指数序列之间的相关性差;乌兰样方内树木年轮生长受立地微环境的影响比较大。 本论文中年龄相关的树木年轮生长与气候因子响应关系的研究进一步证实了祁连圆柏在重建古气候方面的应用价值;通过设立样方进行祁连圆柏种群内个体差异及立地条件对比方面的研究结果对重建青藏高原地区的气候变化、了解采样地的干扰历史以及树木年轮采样工作等也同样具有重要的参考意义。
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Complete sets of chromosome-specific painting probes, derived from flow-sorted chromosomes of human (HSA), Equus caballus (ECA) and Equus burchelli (EBU) were used to delineate conserved chromosomal segments between human and Equits burchelli, and among four equid species, E. przewalskii (EPR), E. caballus, E. burchelli and E. zebra hartmannae (EZH) by cross-species chromosome painting. Genome-wide comparative maps between these species have been established. Twenty-two human autosomal probes revealed 48 conserved segments in E. burchelli. The adjacent segment combinations HSA3/21, 7/16p, 16q/19q, 14/15, 12/22 and 4/8, presumed ancestral syntenies for all eutherian mammals, were also found conserved in E. burchelli. The comparative maps of equids allow for the unequivocal characterization of chromosomal rearrangements that differentiate the karyotypes of these equid species. The karyotypes of E. przewalskii and E. caballus differ by one Robertsonian translocation (ECA5 = EPR23 + EPR24); numerous Robertsonian translocations and tandem fusions and several inversions account for the karyotypic differences between the horses and zebras. Our results shed new light on the karyotypic evolution of Equidae. Copyright (C) 2003 S. Karger AG, Basel.
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程海共有土著鱼类15种,它位于青藏高原横断山区的东缘,云南高原的西北,鱼类区系却独具一格,表现出江河平原鱼类区系的特点,显示程海鱼类区系来自长江的事实。程海自与金沙江隔离之后,湖内存在空白生态灶(niche)是产生物种分化的重要因素,尤以鲤鱼Cyprinus carpio L.的形态,表现出一系列的适应性变异,这些变异发生在近300年内,与处于海拔和纬度较高的甘子河裸鲤Gymnocypris przewalskii ganzihoensis Zhu et Wu形成的时间相比,相对而言是比较快的,然而却慢于
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149 complete mitochondrial DNA (mtDNA) cytochrome b (Cyt b) genes (1140 bp) of Gymnocypris przewalskii, Gymnocypris eckloni and Gymnocyptis scolistomus from the Lake Qinghai, Yellow River and Qaidam Basin were sequenced and analyzed. Consistent dendrogram indicated that the samples collected from the same species do not constitute a separate monophyletic group and all the samples were grouped into three highly divergent lineages (A, B and C). Among them, Lineage A contained all samples of G. przewalskii from the Lake Qinghai and partial samples of the G. eckloni from the Yellow River. Lineage B contained the remaining samples of G. eckloni from the Yellow River. Lineage C was composed of a monophyletic group by G. eckloni from the Qaidam Basin. Analysis of molecular variance (AMOVA) indicated that most of genetic variations were detected within these three mtDNA lineages (93.12%), suggesting that there are three different lineages of Gymnocypris in this region. Our Cyt b sequence data showed that G. przewalskii was not a polytypic species, and G. scolistomus was neither an independent species nor a subspecies of G. eckloni. The divergent mtDNA lineages of G. eckloni from the Yellow River suggested that gene flow between the different populations was restricted to a certain extent by several gorges on the upper reach of the Yellow River. Lineage B of G. eckloni might be the genetic effect from the ancestor which was incorporated with the endemic schizothoracine fishes when the headward erosion of the Yellow River reached to its current headwaters of late. The G. eckloni from Basin Qaidam was a monophyletic group (lineage C) and F-st values within G. eckloni from the Yellow River were higher than 0.98, suggesting that the gene flow has been interrupted for a long time and the G. eckloni from Basin Qaidam might have been evolved into different species by ecology segregation. The correlation between the rakers number of Gymnocypris and population genetic variation was not significant. All Gymnocypris populations exhibited a low nucleotide diversity (pi = 0.00096-0.00485). Therefore the Gymnocyptis populations from Basin Qaidam could have experienced severe bottleneck effect in history. Our result suggested Gym-nocypris populations of Basin Qaidam should give a high priority in conservation programs.
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岷江上游地区高山/亚高山植被分布的坡向性分异显著,阴阳坡高山林线不仅物种组成差异明显,并且分布海拔呈现出阴坡高阳坡低的格局.阳坡林线树种主要是圆柏属乔木,林线类型多为渐变型,海拔高度大约在3 400m~3 800m;阴坡林线树种主要是冷杉,林线类型多为骤变型,海拔高度约在3 800m~4 400m.本研究采用土壤种子库物理筛选、室内萌发实验及野外群落调查等方法,对岷江上游地区阴坡岷江冷杉和阳坡祁连圆柏两类林线树种不同海拔梯度上土壤种子库以及幼苗库特征进行了调查,从土壤种子库和幼苗更新特征的角度对林线乔木树种种群更新特征进行了分析,进而对该地区高山林线在阴阳坡分布差异的原因进行了探讨,结果显示: 1.土壤种子库 阴坡:阴坡高山林线附近岷江冷杉土壤种子的平均密度大约为50.96粒/m2,其中树线以上10m处土壤种子密度为1.00粒/m2,树线处大约19.33粒/m2,林线交错带内土壤种子密度最高为136.83粒/m2,郁闭林内种子密度小于林线交错带,只有30.50粒/m2,种子平均空壳率为52%,霉变率达34%,完好种子只有6%.土壤种子库垂直分布特征为地被物层含种子比重最大,大约在67.50%左右;其次为0~2cm层,约18.84%左右;2~5cm层所占种子比例最小,约13.66%左右.霉变种子数量与土壤深度呈负相关. 阳坡:阳坡祁连圆柏土壤种子的平均密度为60.16粒/m2.树线以上10m处密度为1.92粒/m2,树线位置大约108.16粒/m2,林线交错带内平均为75.80粒/m2,郁闭林内种子密度小于林线交错带,只有20.00粒/m2.种子平均空壳率为36%,完好种子占49%,霉变率较低,大约为10%.阴阳坡林线树种土壤种子库垂直分布特征为:地被物层含种子最多,其次为0~4cm层,4~10cm层所占种子比例最小,霉变种子数量与土壤深度也呈负相关. 2. 幼苗库调查 阳坡:在树线以上区域没有发现幼苗,林线交错带内幼苗密度平均达3 250株/hm2,郁闭林内仅2 750株/ hm2.整个样地内1~2a幼苗很少甚至没有出现,3~10a的幼苗相对较多.空间分布上,祁连圆柏幼苗在林线交错带内接近随机分布,郁闭林内则介于随机分布和均匀分布之间. 阴坡:在树线以上幼苗密度为1 250株/ hm2,全部为1~2a幼苗,林线交错带内幼苗密度平均达7 000株/ hm2,郁闭林内达6 250株/ hm2.林线附近岷江冷杉幼苗丰富度以及幼苗的出现频率明显高于祁连圆柏,年龄结构也较祁连圆柏完整.岷江冷杉幼苗空间分布除了树线处幼苗的分布为随机分布,其他海拔则为集群分布. 3.从不同土壤深度的种子总量和幼苗数量的相关性检验发现,当年生幼苗数量跟表层种子总量相关性极显著, 但是两年生幼苗的数量与底层种子数量相关性显著.土壤种子在土壤中的垂直分布格局从一定程度上可以反映种子库的年际特征.岷江冷杉土壤种子库较丰富,种子散布后的存活力随着时间的变化逐渐下降,属于季节性瞬时种子库;祁连圆柏土壤种子散布格局为集群型分布,成熟种子大部分散布在母株冠幅内,属于永久性土壤种子库. 4.在阴坡林线交错带及以上区域还存在较为丰富的乔木土壤种子,并且在树线以上区域还发现了少量的岷江冷杉幼苗.从样地乔木的年龄结构发现,在林线交错带内上部到树线位置主要以幼龄林为主,且年龄结构完整,基本符合入侵性林线特征;阳坡林线交错带内幼苗出现频率很低,树线以上区域虽然存在种子库,但是没有幼苗出现,在林线交错带内乔木径级差距很大,年龄结构异常不完整,这种特征的林线将会面临两个可能结果:一种是维持现有状态,保持平衡;另外一种就是退化,但阳坡林线的实际动态趋势还有待长期定点研究. Treelines on the upper region of Minjiang River differ between the north aspect and the south aspect in their appearances, altitudes and tree species. On the north aspect, trees of Abies form a sharp and abrupt treeline ranging from 3800m to 4400m, while on the south the treeline is generally lower(3 400~3 800m), more open and gradual and mostly composed of Sabina. In this study, we examined the altitudinal gradients of soil seed banks and seedling recruitments at the treeline ecotones of a N-aspect and a S-aspect by using soil sieving, germination experiment and field investigations, analyzed the characteristics of population regeneration of tree species at the transitional zone and presented a analysis of the causes to the aspect-related difference in treeline patterns in the study area. Major results of our study include: 1. Soil seed bank N-aspect: Of the 50 plots investigated, the average density of soil seeds is 50.96/m2, in which well-formed seeds account for 6%, empty seeds 52%, parasitized seeds34%, and seeds damaged by animals 8%. The size of soil seed bank varies along altitude, being 1.00 seeds /m2 at the 10m above the treeline and ca.19.33 seeds/m2 at the upper limit of treeline. The highest density (136.83 seeds/m2) occurs at the treeline ecotone. By contrast, the density of soil seed for the closed forest is only 30.50 seeds/m2. In terms of vertical strata, 67.50% of the total seeds are at the surface layer, 18.84% at the middle layer (0~2cm) and 13.66% at deeper layer (2~5cm). The number of parasitized seeds is negatively correlated to soil depth. S-aspect: Of the 50 plots investigated, the average density of soil seeds is 60.16 seeds/m2, and the well-formed seeds account for 49%, empty seeds 36%, parasitized seeds10%, and seeds damaged by animals 1%. The size of soil seed bank varies along altitude, with 1.92 seeds/m2 recorded at the10m above the treeline,108.16 seeds/m2 at the upper limit of treeline, and 75.80 seeds/m2 at the treeline ecotone, while that for the closed forest is 20.00 seeds/m2. The number of seeds decreases with the depth of soil. As is on the N-aspect, the size of soil bank, from large to small, follows the order of the surface layer, the middle layer (0~4cm) and the bottom layer (4~10cm). The number of parasitized seeds is also negatively correlated to the depth of the soil. 2. Seedling bank N-aspect: A mean maximum seedling abundance of 31 000 seedlings/hm2 was recorded near alpine treeline at growing season. The density of seedlings is 1 250 seedlings/ha (all being 1 or 2 years old) at the alpine meadow 10m away above treeline, 7 000 seedlings/ha at treeline ecotone and 6 250 seedlings/ha for closed forest.The spatial distribution of Abies faxoniana seedlings is random at the upper limit of the treeline but clumped at other altitudes. S-aspect: No seedlings were found at the alpine meadow 10m away from the treeline. The density of seedlings was 3 250 seedlings/ha at treeline ecotone and 2 750 seedlings/ha for the closed forest.Hardly any 1 year current and 2 year-old seedlings appeared at the plots. The spatial distribution of Sabina przewalskii seedlings is random at treeline ecotone and between “random” and “even” forest closed forest. 3.Correlation tests of seedling population and seed bank at different soil layers indicated that the emergents were strongly correlated to seed bank at surface layer while the number of two-year seedlings was significantly correlated to the seed bank at the bottom of soil layer, indicating that germination mainly occurs at the soil surface while the middle or bottom layer was the reserve for non-germination or dead seeds. It can thus be postulated that Abies faxoniana soil seed bank is of seasonal transient type. By contrast, the soil seed bank of Sabina przewalskii is of persistent type and the soil seeds and seedlings of this species occurred more frequently near the islands of adult trees. 4.A good many soil seeds of both tree species were found near the treeline ecotone and above at N- and S-aspects. A few young seedlings were found above the Abies treeline. Investigation of five altitudinal transects respectively on N- and S-aspects indicated that Abies faxoniana has a more complete age structure than the stands of Sabina przewalskii. The age of firs decreased from closed forest to the upper limit of treeline, which suggests that the Abies treeline is advancing to higher altitude. While on the south aspect, only few Sabina przewalskii soil seeds and nearly no seedlings were found above the treeline ecotone. The stands exhibit extremely great difference in diameter classes with significantly incomplete age structure. This would lead to two possible results for the treelines: maintaining an equilibrium state at the current position or degenerating. But more studies should be carried out at longer time scales or larger spatial scales to understand whether the Sabina treeline is degenerating.
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土壤是人类赖以生存的自然环境和农业生产的重要资源,世界面临的粮食、资源和环境问题与土壤密切相关,目前危害土壤的主要因素是干旱和重金属污染。杨树具有适应性强、生长快和丰产等特性,本论文以青杨组杨树为模式植物,采用植物生态、生理及生物化学等方法,研究杨树对土壤干旱和锰胁迫的生态生理反应以及种群间差异,研究成果可为我国干旱半干旱地区营造人工林、防止沙漠化提供理论依据,也为恢复与重建重金属污染地区退化生态系统提供科学指导。主要研究结果如下: 1. 青海杨不同种群对干旱胁迫的响应差异 干旱胁迫显著降低了两个青海杨种群(干旱种群和湿润种群)生物量积累,包括株高、基径、干物质积累等,通过植物结构的调整,有更多的生物量向根部分配。干旱胁迫还显著降低了叶绿素和类胡萝卜素含量,增加了游离脯氨酸和总氨基酸含量。另一方面,干旱胁迫诱导了活性氧的积累,作为第二信使,激活了抗氧化系统,包括抗坏血酸(ASA)含量和酶系统如超氧化物歧化酶(SOD),愈创木酚过氧化物酶(GPX),抗坏血酸过氧化物酶(APX)和谷胱甘肽还原酶(GR)。这样,杨树既有避旱机制又有耐旱机制,使其在干旱胁迫下有相当程度的可塑性。与湿润种群相比,干旱种群杨树有更多的生物量分配到根部,积累了更多的游离脯氨酸和总氨基酸来进行渗透调节,并且有更有效的抗氧化系统,包括更高含量的ASA 和更高活性的APX 和GR,这些使得干旱种群杨树比湿润种群杨树对干旱有更好的耐性。 2. 喷施硝普钠(SNP)对青海杨阿坝种群干旱胁迫耐性的影响 干旱胁迫显著的降低了青海杨阿坝种群的生长和生物量积累以及叶片相对含水量,还诱导了脯氨酸的合成以进行渗透调节。干旱胁迫下过氧化氢(H2O2)显著累积从而造成对膜脂和蛋白的伤害,使得丙二醛和蛋白羰基含量升高。干旱胁迫下喷施SNP可以减轻干旱胁迫造成的伤害,包括增加叶片的相对含水量,增加脯氨酸和总氨基酸的积累,并激活抗氧化酶系统如SOD,GPX和APX,从而减少丙二醛(MDA)和蛋白羰基(C=O)的积累,但是在水分良好情况下SNP的效果不显著。 3. 青杨不同种群对锰胁迫的生长与形态响应差异 在同一锰浓度下,干旱种群的耐性指数都要高于湿润种群,这表明青杨对干旱和高锰胁迫具有交叉耐性。两个种群的株高,生物量和叶绿素含量都随锰浓度的升高而逐渐下降。就累积浓度而言,0 和0.1 mM 锰胁迫下,干旱种群积累的锰浓度要高于湿润种群,而在高浓度锰胁迫下(0.5 和1 mM),湿润种群要高于干旱种群。在0,0.1 和0.5 mM下,锰大多积累在根中,叶片次之,茎中最少。而在1 mM,锰更多的积累在叶片中。就累积总量而言,在各个锰浓度胁迫下,根,茎和叶相比,两个种群青杨都是叶片累积的锰总量要高于根和茎。两个种群间比较,对照中没有显著区别,0.1 mM 锰胁迫下,湿润种群根中累积的锰要高于干旱种群,而在地上部中,干旱种群要高于湿润种群。而0.5 和1 mM 锰胁迫下,根、叶、茎+叶、根+茎+叶中,锰累积总量都是湿润种群高于干旱种群。锰胁迫下,青杨叶片数和叶面积包括总叶面积和平均叶面积都显著降低。叶片横切面的光学显微观察结果表明未经锰胁迫的栅栏组织的细胞饱满,海绵组织发达、清晰;胁迫后杨树叶片栅栏组织细胞出现不同程度的皱缩,海绵组织几乎不可见,此外还发现输导组织在胁迫下密度变小和分生组织严重割裂等现象。 4. 青杨不同种群对锰胁迫的生理与生化响应差异 青杨两个种群脱落酸(ABA)含量在锰胁迫下都显著增加,干旱种群的增幅更大。三种多胺含量在锰胁迫下显示了不同的响应趋势:腐胺在两个种群的各个锰处理下都增加,亚精胺只在干旱种群中显著增加,而精胺除了干旱种群在1 mM 下有所增加外,在锰胁迫下变化很小。谷胱甘肽含量随锰浓度升高而增加,在0.5 mM 锰时达到最高值,1mM 时有所下降。植物络合素(PCs)含量与非蛋白巯基(NP-SH)趋势相似,随锰浓度的升高而增加,且干旱种群中含量要高于湿润种群。锰处理还引起氧化胁迫,表现为过氧化氢和丙二醛含量增加。SOD 活性在湿润种群中,在0 到0.5 mM 锰胁迫下活性升高,但在1 mM 锰胁迫时,其活性有所下降。而在干旱种群中,SOD 活性变化较小,并始终维持在一个较高的水平。APX 活性在两个种群中都随锰浓度的升高而增加,干旱种群活性要高于湿润种群。锰胁迫还显著增加了酚类物质的含量,同时GPX 和多酚氧化酶(PPO)活性也随锰浓度的升高而增加。干旱种群的酚类含量和GPX 与PPO 活性都要高于湿润种群。锰胁迫还改变了氨基酸的含量和构成,根据锰胁迫下浓度变化的不同,可以将游离氨基酸分为三组:第一组包括,谷氨酸,丙氨酸和天门冬氨酸,这一组氨基酸含量在锰胁迫下有所下降。第二组包括缬氨酸,亮氨酸和苏氨酸含量在锰胁迫下基本不变化或变化很小。剩下的氨基酸为第三组,这组氨基酸含量在锰胁迫下显著增加,而根据增加的幅度又可以将它们分为两个亚组,丝氨酸,酪氨酸,苯丙氨酸,组氨酸和脯氨酸,在1 mM 下的含量是对照的4 倍以上。异亮氨酸,赖氨酸,精氨酸和甘氨酸含量在1 mM 下是对照含量的2 倍以下。同时,同一锰浓度下,干旱种群比湿润种群积累的氨基酸含量要高。 Soil is the indispensable environment for human survival and important resource foragriculture development. Food and environmental problems facing the world are all closelyrelated to soil and nowadays it is threatened by many factors, among which drought stress andheavy metal pollution are the most serious ones. Poplars (Populus spp.) are importantcomponents of ecosystem and suitable as a source of fuel, fiber and lumber due to their fastgrowth. In this study, different populations of Section Tacamahaca spach were used as modelplants to investigate the adaptability to drought stress and manganese toxicity and differencesbetween populations from dry and wet climate regions. Our results can provide theoreticalevidence for the afforestation and prevention of desertification in the arid and semi-arid areas,and also can supply scientific direction for the reconstruction and rehalibitation of ecosystemscontaminated by heavy metals. The results are as follows: 1. Differences in ecophysiological responses to drought stress in two contrastingpopulations of Populus przewalskii Drought stress not only significantly affected dry mass accumulation and allocation, butalso significantly decreased chlorophyll pigment contents and accumulated free proline andtotal amino acids. On the other hand, drought also significantly increased the levels ofabscisic acid and reactive oxygen species, as secondary messengers, to induce the entire set ofantioxidative systems including the increase of reduced ascorbic acid content and the activities of superoxide dismutase, guaiacol peroxidase, ascorbate peroxidase and glutathioneredutase. Thus the combination of drought avoidance and tolerance mechanisms conferredpoplar a high degree of plasticity in response to drought stress. Compared with the wetclimate population, the dry climate population showed lower dry matter accumulation andallocated more biomass to root systems, and accumulated more free proline and total aminoacids for osmotic adjustment. The dry climate population also showed more efficientantioxidant systems with higher content of ascorbic acid and higher activities of ascorbateperoxidase and glutathione redutase than the wet climate population. All these made the dryclimate population superior in adaptation to drought stress than the wet climate population. 2. Effect of exogenous applied SNP on drought tolerance in Populus przewalskii Drought stress significantly increased hydrogen peroxide content and caused oxidativestress to lipids and proteins assessed by the increase in malondialdehyde and total carbonylcontents, respectively. The cuttings of P. przewalskii accumulated proline and other aminoacids for osmotic adjustment to lower water potential, and activated the antioxidant enzymes such as superoxide dismutase, guaiacol peroxidase and ascorbate peroxidase to maintain thebalance of generation and quenching of reactive oxygen species. Moreover, exogenous SNPapplication significantly heightened the growth performance of P. przewalskii cuttings underdrought treatment by promotion of proline accumulation and activation of antioxidant enzymeactivities, while under well-watered treatment the effect of SNP application was very little. 3. Morphological responses to manganese toxicity in the two contrasting populations ofPopulus cathayana High concentration of manganese caused significant decrease in shoot height andbiomass accumulation. The tolerance index of the dry climate population was significantlyhigher than that of the wet climate population, suggesting the superior Mn tolerance in theformer and the existence of cross-tolerance of drought stress and high Mn toxicity. Injuries tothe leaf anatomical features were also found as the reduced thickness in palisade and spongyparenchyma, the decreased density in the conducting tissue and the collapse and split in themeristematic tissue in the central vein. As for the Mn concentrations in the plant tissues, under0, 0.1 and 0.5 mM, most of the Mn accumulated in the roots, then leaves, and stem the least, while under 1 mM, most of the Mn accumulated in the leaves. As far as the total amounts ofMn extraction are concerned, the leaf extracted more Mn than the root and stem in the twopopulations under various Mn concentrations. There is no difference between the twopopulations under control. Under 0.1 mM, the wet climate population extracted higher Mn inthe root than the dry climate population, while in the shoot, the dry climate populationextracted much more Mn. Under 0.5 and 1 mM, the wet climate population translocated moreMn both in the root and the shoot than the dry climate population. 4. Physiological and biochemical responses to manganese toxicity in the two contrastingpopulations of Populus cathayana Mn treatment resulted in oxidative stress indicated by the oxidation to lipids, proteinsand DNA. A regulated network of defence strategies was employed for the chelation,detoxification and tolerance of Mn including the enhanced synthesis of ABA and polyamines,the accumulation of free amino acids, especially His and Pro, and the activation of theenzymes superoxide dismutase and guaiacol peroxidase. Contents of non-protein thiol,reduced glutathione, phytochelatins and phenolics compounds and activities of superoxide dismutase, guaiacol peroxidase and polyphenol oxidase also increased significantly forantioxidant or chelation functions. The wet climate population not only accumulated lessabscisic acid, free amino acids, phytochelatins and phenolics compounds, but also exhibitedlower activities of superoxide dismutase, guaiacol peroxidase and polyphenol oxidase thusresulting in more serious oxidative damage and more curtained growth.
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青杨组(Section Tacamahaca Spach)杨树是我国重要的乡土经济树种,目前对其分子遗传变异和系统进化的研究还很少,尤其是在青杨组杨树遗传资源极为丰富的川西地区,杨树的分子进化及亲缘关系的研究极为缺乏,非常不利于该树种遗传资源的开发和利用。本研究从川西地区收集了青杨(Populus cathayana)、青海杨(P. prezewalskii)、滇杨(P. yunnanensis)、康定杨(P. kangdingensis)、西南杨(P. schneideri)、小叶杨(P.simonii)和三脉青杨(P. trinervis)这7 个青杨组树种的10 个群体,利用多种分子标记手段对其种间的亲缘关系进行比较,并结合形态和地史资料进行了全面的研究和评价,得到了如下的主要研究结果: 1. SSR 和ISSR 位点变异丰富。通过10 对引物对50 个杨树个体的DNA 样品进行了SSR 分析,所有位点展现了丰富的群体间和种间的多态性,多态位点率达到了100%,每位点的等位基因数变化范围为5 ~ 17,平均为11.9 个;通过11 条ISSR 随机引物对供试的混合DNA 样品进行分析,共检测到130 个标记,其中多态性标记为119 个,多态百分率为91.5%。研究认为,SSR 单个标记能展现高水平信息,而ISSR 单个引物能探测更多数量多态性。通过两个标记的遗传距离、聚类图和PCA 分析,表明:同一种内不同群体间的同源性最高;康定杨和西南杨有较近的亲缘关系;小叶杨和三脉青杨聚合在一起,显示了其相互较近的亲缘关系;滇杨与其它杨树种可能存在着较远的亲缘关系。 2. 采用4 对选择性引物对7 个青杨组杨树种10 个群体进行AFLP 分析,总共扩增出284 个位点,其中200 个位点显示出了多态性,多态位点百分比为70.4%,平均多态带为50 条。TE-AFLP 的分析总共扩增出192 个位点,其中139 个位点显示出了多态性,多态位点百分比为72.4%,平均多态带为34.7 条。比较的结果表明AFLP、TE-AFLP 的遗传信息含量比较接近,略小于ISSR,大约仅为SSR 的1/3;但这两个基于AFLP 的标记系统的信息探察能力也远大于ISSR 和SSR 标记系统。这两个分子标记的聚类结果,显示小叶杨、三脉青杨和滇杨三个种聚为一组,其中小叶杨与三脉青杨的亲缘关系更近;其它几个杨树种聚为一类,西南杨与青杨表现出较近的亲缘关系。 3. 所有7 对cpSSR 引物中,仅有4 个叶绿体位点在种间具有多态性,而在种内群体中并不具有多态性,共检测出13 个条带,组合成了4 种不同的单倍型;对于cpDNA的5 对引物,共检测出了73 条酶切片段,其中52 条是多态带,组合成了9 种不同的单倍型;而5 对mtDNA 通用引物未能检测出多态性的条带,表现出线粒体的保守性。叶绿体的聚类分析认为,小叶杨、三脉青杨和滇杨有较近的母性起源,且依次聚合;其余四种杨树聚为一类,并且康定杨与西南杨表现出最近的亲缘关系,并依次与青杨和青海杨聚合。 4. 根据本文的分子数据,结合形态和生境分布资料分析认为:青杨组杨树种内群体间的遗传变异程度是小于种间的遗传差异,显示了与传统分类一致的结果;三脉青杨和小叶杨有很近的亲缘关系,可能拥有相同的祖先类群;滇杨与小叶杨和三脉青杨之间具有一定的亲缘关系,特别是在其母性祖先的起源上有着一定的同源性;西南杨与青杨和康定杨均保持着较近的亲缘关系,且有可能是这两个种原始祖先杂交后所形成的。 Although western Sichuan is regarded as a natural distribution and variation center forthe Section Tacahamaca of the Populus species in China, little is currently known about themajority of poplar species occurring in this region. In the present study, molecular data wereutilized to determine the genetic relationships among Populus species in Section Tacamahacain western Sichuan including P. cathayana, P. prezewalskii, P. yunnanensis, P. kangdingensis,P. schneideri, P. simonii and P.trinervis. The results are as fellows: 1. The genetic variation at SSR and ISSR loci was abundant. All the 10 SSR loci werepolymorphic, and the number of alleles per locus varied from 5 to 17 with a mean valueequaling 11.9. Based on the 11 ISSR primers, 130 clear and reproducible DNA fragmentswere generated, of which 119 (91.5%) were polymorphic. Our results reveal that single SSRlocus can present more genetic information, while more polymorphic bands can be detectedby single ISSR primer. Moreover, the genetic distance, cluster and PCA analysisdemonstrated that: a close relationship among accessions of the same species and suggestedmonophyly in P. przewalskii and P. cathayana; P. schneideri is genetically highly similar to P.kangdingensis; P. trinervis and P. simonii have a close genetic affinity; P. yunnanensis isdistinct from the other species. 2. Genetic relationships of poplar species in Section Tacamahaca from western Sichuanwere evaluated by means of AFLP and TE-AFLP. For four AFLP primer combinations, atotal of 284 bands were obtained of which 200 (70.4%) were polymorphic with the average of50 polymorphic bands. For four TE-AFLP primer combinations, a total of 192 band wereobtained of which 139 (72.4%) were polymorphic with the average of 34.7 polymorphicbands. Our results indicate that the genetic information of AFLP is similar to that ofTE-AFLP, and little less than that of ISSR, but only about 1/3 of that of SSR. However, theability of information detection of the two AFLP-based markers is much higher than that ofISSR and SSR. In addition, the cluster analysis of AFLP, TE-AFLP and combined data revealthat: P. yunnanensis, P. trinervis and P. simonii clustered together, and P. trinervis and P.simonii showed more closed affinity; the other four poplar species clustered together, P.cathayana and P. schneideri showed more closed origin especially. 3. The cpSSR analysis for seven Populus species belonging to the Section Tacamahaca.Four out of the seven analyzed chloroplast loci were polymorphic, whereas none of the lociwere polymorphic across the accessions within a species. 13 bands and 4 different kinds ofhaplotypes were reduced. Based on 5 pairs of cpDNA primers, 73 fragments (52 polymorphic)and 9 kinds of haplotypes were produced. However, none of the polymorphic was detected bythe 5 mtDNA primer pairs, revealing conservation of mtDNA region. The cluster analysis ofcpDNA revealed that: similar maternal phylogeny among P. yunnanensis, P. trinervis and P.simonii; the other four species clustered together, P. schneideri and P. kangdingensis showedmore closed maternal lineage especially. 4. Our molecular data, morphological characters and nature habitat revealed that: sameto the traditional taxonomy assignment, genetic variation within a same Populus species islower than that among Populus species in Section Tacamahaca; P. yunnanensis may share itschloroplast ancestor with P. trinervis and P. simonii; moreover, sister genetic relationship of P.trinervis and P. simonii indicated their similar origin; P. schneideri clustered with P.kangdingensis and P. cathayana, respectively, and may have derived from an ancienthybridization event involving the ancestors of the two species.
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沙蜥属Phrynocephalus Kaup,1825隶属于爬行纲(Reptilia)有鳞目(Squamata)蜥蜴亚目(Lacertilia)鬣蜥科(Agamidae),是欧亚大陆荒漠和稀疏草原常见蜥蜴。沙蜥属的分类及系统演化关系、地理分布格局与新生代第三纪以来古地中海的变迁、青藏高原的抬升及亚洲内陆干旱荒漠化的过程有密切的关系,长期以来有关沙蜥属的研究一直受到中外学者们的关注。由于沙蜥属地理分布广、形态变异大、体色和斑纹变化复杂,虽然前人使用过许多形态性状来描述和分类沙蜥属物种,但是仍然存在许多问题。性状的分类学意义不明确是造成这些问题的主要原因之一,因此本研究针对沙蜥属常用的鉴别性状进行分类意义的分析,希望能对沙蜥属物种鉴定及分类学其它研究有所裨益。 中国沙蜥属物种主要分布于西北的干旱荒漠区域及青藏高原的大部分地区,大约为18种。 本文研究了中国境内12种沙蜥:青海沙蜥(Phrynocephalus vlangalii)、西藏沙蜥(P. theobaldi)、南疆沙蜥(P. forsythii)、变色沙蜥(P. versicolor)、旱地沙蜥(P. helioscopus)、荒漠沙蜥(P. przewalskii)、乌拉尔沙蜥(P. guttatus)、草原沙蜥(P. frontalis)、叶城沙蜥(P. axillaris)、白稍沙蜥(P. koslowi)、无斑沙蜥(P. immaculatus)和白条沙蜥(P. albolineatus),对它们的65项外部形态性状进行了观察和测量,其中数量性状29项、质量性状36项。评价了这些性状的序级性、间断性和代表性,结论如下: 1. 对于数量性状,得出了适合各级分类的数值区间; 2. 给出了在不同序级上适合分类的质量性状。 并利用各性状评价的结果,给出12种沙蜥的检索表,以及对中国沙蜥物种某些尚存在争议的问题进行了探讨。 详细记录了青海沙蜥红原亚种的骨骼系统,首次发现并命名了肘骨(elbow bone)和垫骨(stepping bone),为沙蜥属系统学研究补充了骨骼方面的证据;解剖了乌拉尔沙蜥、旱地沙蜥、荒漠沙蜥的雌体和雄体的骨骼系统,并在14项骨骼形态性状上对这3种沙蜥进行了比较。 Phrynocephalus (Squamata,Agamidae) is a familiar genus of lizards inhabited desert and sparse steppes in Eurasia. The taxonomics, phylogenetics and distribution pattern of Phrynocephalus are relative intensely to these events: the vicissitudes of the archaic Mediterranean sea since the Cainozoic, the uplift of Qingzang Plateau and the expending arid areas in the inland of Asia. Owing to the wide distribution, the large variability of the morphology and the different colors in Phrynocephalus, it is difficult to identify them. Tough many morphological characters are used to describe and discriminate them,a lot of questions still exist. One of the most important reasons is the confusion in the morphological characters. In this study, we demonstrate the validity and the invalidity of the familiar characters. There are about 18 species of the genus Phrynocephalus in China, which exist in arid desert in Northwest China and Qingzang Plateau. Twelve Chinese species was analyzed in this paper. They are P. vlangalii,P. theobaldi,P. forsythia,P. versicolor,P. helioscopus,P. przewalskii,P. guttatus,P. frontalis,P. axillaris,P. koslowi,P. immaculatus,P. albolineatus. We measure 29 quantitative characters and observe 36 qualitative characters in each individual. Through analyzing these characters, we made some conclusions as follows: 1. to every quantitative character, we get a clear numeric area to discriminate the different operational taxonomic units. 2. we chose the valid qualitative characters in these operational taxonomic units. This paper is the first to describe the “elbow bone”, which is a bone in pectoral appendage equivalent to patella,and “stepping bone”, which is a bone under carpal. A detailed description of the skeletal system of female Phrynocephalus vlangalii hongyuanensis was conducted. We also anatomise the skeletal systems of three species: P. guttatus,P. przewalskii,P. helioscopus, and compare or contrast 14 skeletal characters in them. What’s more,this paper offers some suggestions to the questions of Chinese Phrynocephalus species and keys to 12 species of Phrynocephalus basing on our conclusions on the evaluation of the morphological characters.
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雌雄异熟和异型花在一些被子植物中的共同存在被认为是一种有效避免两性花植物自交并促进异交的机制。本文报道青藏高原特有植物祁连獐牙菜Swertia przewalskii pissjauk.的花同时具有雌雄异熟和异型花这两种特征。在此基础上,我们详细观察了该植物访花昆虫的访花行为,主要探讨这种促进异花传粉的机制是否能完全避免株内异花传粉的自交。繁育系统试验证明祁连獐牙菜单花套袋不结实,说明雌雄异熟和异型花的花特征完全避免了单花的自花传粉,自然结实必须依赖昆虫传粉才能完成。然而,该植物却是高度自交亲和的,因为人工株内异花自交具有较高的种子结实率。每个植株在盛花期开放1-6朵,平均具有3朵开放的雄性期或者雌性期的花,二者的比例为2:1,在植株上的分布没有规律。我们观察到多种昆虫访问该植物的花,其中,克什米尔熊蜂Bumbuskeshimirensis Friese和蜜蜂如Apis mellifera L.是该植物最常见与最有效的传粉者。进一步的观察发现,这两类传粉昆虫对雌性阶段和雄性阶段的花的访问没有选择,但在一个植株上访花顺序的总体趋势是由下向上的。它们在同一植株上访花数超过1朵的访问比例约占37.1%。因此可以初步推断,祁连獐牙菜中异型花和雌雄异熟现象的共同存在并没有完全避免同株异花传粉导致的自交。结果支持两性花中的这种雌雄异熟和异型花现象可能在避免两性冲突上具有更为重要的功能。
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目的 研究祁连獐牙菜Swertia przewalskii全草的化学成分。方法 采用硅胶柱层析进行分离纯化,通过波谱方法及化学关联进行结构鉴定;结果 分离纯化出15个化合物,分别鉴定为:1,7-二羟基-3,8-二甲氧基San酮(Ⅰ),1,8-二羟基-3,7-二甲氧基San酮(Ⅱ),1,7,8-三羟基-3-甲氧基San酮(Ⅲ),1,3,7,8-四羟基San酮(Ⅳ),1-O-β-D-吡喃葡萄糖基-7-羟基-3,8-二甲氧基San酮(Ⅴ),1-O-β-D-吡喃葡萄糖基-8-羟基-3,7-二甲氧基San酮(Ⅵ),7-O-β-D-吡喃葡萄糖基-1,8-二羟基-3-甲氧基San酮(Ⅶ),1-O-[β-D-吡喃木糖(1→6)-β-D-吡喃葡萄糖]-7-羟基-3,8-二甲氧基San酮(Ⅷ),1-O-[β-D-吡喃葡萄糖]-1,7-二羟基-3-甲氧基S sgmk(X),木犀草素(Ⅺ),齐墩果酸(Ⅻ),乌苏酸(XⅣ)和龙胆苦苷(XV)。结论 化合物Ⅳ-Ⅷ,Ⅺ,Ⅻ,XⅣ和XV均为首次从该种植物中分得。
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分析和报道了马先蒿属(Pedicularis Linn.)分布于青藏高原东北部6个特有种的核型,并根据核型及其有关参数,分析和比较了该6种马先蒿核型的不对称性和相对进化程度。6个种的体细胞染色体数目都是2n=16。核型分别为:绵德马先蒿P.pilostachya Maxim.,核型公式K(2n)=16=4m+12sm,染色体相对长度组成2L+6M2+6M1+2S,核型不对称系数As·K=65.29%,属于2A型;青海马先蒿P.przewalskii Maxim.,K(2n)=16=8m(SAT)+4sm+2s+2t,2L+8M2+2M1+4S,As·K=65.02%,2B型;华马先蒿P.oederi Vahl.var.sinensis(Maxim.)Hurus.,K(2n)=16=12m+4sm,2L+4M2+8M1+2 S,As·K=59.89%,2B型;粗野马先蒿P.rudis Maxim.,K(2n)=16=4m+10sm+2st,4L+4M2+4M1+2 S,As·K=68.10%,2B型;甘肃马先蒿P.kansuensis Maxim. Subsp. kansuensis,K(2n)=16=6m+6sm+2st+2t,2L+6M2+6M1+2 S,As·K=68.92%,2A型;藓生马先蒿P.muscicola Maxim. K(2n)=16=8m(SAT)+8sm,2L=8M2+4M1+2S,As·K=62.64%,2B型。根据这6个种的核型和已有资料,认为该属的染色体基数x=8,极少数种有多倍体。通过对以上6种核型及进化程度的比较,该属核型变异较大,以中部着丝粒染色体为组成基础(较原始的种类,如华马先蒿和绵穗马先蒿),端部或近端部着丝粒染色体存在与否与该属内种的进化程度有关。核型不对称性所表示的进化程度似乎与花冠的演化有联系。