125 resultados para Kalamis, active 5th century B.C.


Relevância:

100.00% 100.00%

Publicador:

Resumo:

长链脂肪酸、醇及其酯与多种功能材料密切相关,如表面活性剂、生物膜、昆虫激素、储能材料等。链结构的变化对材料性能的影响,引起化学家的广泛兴趣。弄清链结构变化与光谱之间的关系,对探讨材料结构与性能的关系、异构体的鉴别有着重要的理论意义和实际用途。长链单不饱和脂肪醇乙酸酯作为鳞翅目昆虫的性外激素,有着重要的实用价值。有关该类化和物的红外光谱和拉曼光谱以及简正坐标分析在国内外尚未见报导。我们详细探讨了该类系列化和物共11个的红外光谱和拉曼光谱振动频率及其强度与结构的变化关系,并以顺-3-已烯-1-醇乙酸酯为模形进行了简正坐标分析。第一部分研究了系列化合物的红外和拉曼光谱对11个化合物在室温(液态)和低温(固态)的红外光谱及室温时拉曼光谱进行了观测和研究。一、谱带归属参考有观文献及我们所作的简正作标分析,将本系列化合物振动光谱谱带进行归属。C=O为极性基团,其伸缩振动在红外光谱中呈现强吸收带。双键C=C伸缩和和甲基、亚甲基中C-H伸缩、亚甲基变形在拉曼光谱中为强散射带。因此红外与拉曼光谱相配和有助于区分这类化和物。二、系列化合物的光谱差异在室温下,各异构体的红外和拉曼光谱都相似,难以区分不同的异构体。但低温固态的红外光谱则有明显差别,可用于异构体的结构判别。低温红外光谱中1500~1300cm~(-1)甲基、亚甲基的变角振动区,1050cm~(-1)附近的C-O伸缩振动区1000cm~(-1)以下亚甲基平面摇摆振动及C=O面外变形振动有差别,这些差别不仅表现为谱带分裂数目不同,而且相对强度也不同。三、谱带位移双键插入分子链中引起某些红外特征频率移动。在室温下,亚甲基平面摇摆振动720cm~(-1)区随亚甲基链-(-CH_2-)-_m、-(-CH_2-)-_nm,n的增加向低频方向移动。而在低温固态时,除亚甲基平面摇摆振动频移外,C-O-C反对称伸缩振动1250cm~(-1)区随m,n的增大移向高频,与饱和脂肪酸酯中该频率位移方向相反。=C-C反对称伸缩970cm~(-1)区也随n的增加移向高频,与m的增加无关。四、红外光谱强度双键的存在也影响了亚甲基伸缩振动的红外吸收强度。双键的嵌入破坏了亚甲基链的连续性,亚甲基对称伸缩的相对强度I_(2857)/I_(2957)与亚甲基链长总数无直接关系,而与连续亚甲基链长m,n有线性关系。m,n之一增大时,相对强度增强,当链长一定,双键位置在链中移动时,双键位于碳链中间时,相对强度最低。因此亚甲基对称伸缩红外吸收相对强度由连续亚甲基链-(-CH_2-)-_m -(-CH_2-)-_n中m、n的大小和双键所在位置决定。五、拉曼光谱强度系列化合物碳链长度及双键位置对拉曼光谱散射强度也有规律性影响。双键的存在使亚甲基的振动模式的散射强度不是单个亚甲基的简单加和,而与m,n有关。m,n之一增大时,亚甲基对称伸缩相对强度I_(2851)/I_(2871)增强。m,n的增大对相对强度的影响程度不同,m增大时所引起的相对强度增强速度更快,是n的1.7倍左右。因此,利用这些工作曲线,接合某些物性资料,可以估计该类未知化合物的双键位置和链长。第二部分对该类化合物进行了简正坐标分析。我们以顺-3-已烯醇-1-乙酸酯为模型,对长链单不饱和脂肪醇乙酸酯进行了简正坐标分析,选取局部对称的U-B力场,应用NRCC程序,得到一套使分子中各特征基团的计算频率与观测值同时符合得很好的力常数和相互作用力常数。顺-3-已烯-1-醇乙酸酯的结构式:CH_3CH_2HC=CHCH_2CH_2-O-C = O 模型化合物分子含有三种基团:C = C、 -C = O、CH_3-CH_2-。无整体对称性。将它们进行了局部对称化,胺局部对称的U-B力场选取了相应的初始力常数。C-H伸缩振动,计算计算频率与观测值符合的非常好,势能分布很集中。各振动模式间偶合作用很小。甲基的两种简并伸缩模式,Takehike等认为不能区分,只给出一个力常数。实际上这是两种不同的振动模式。我们给出了各自的力常数。振动频率分别为2952cm~(-1)、2930cm~(-1)。甲基和亚甲基各振动模式,计算结果与实验值吻合的很好。应用局部对称坐标组合后,各振动模式之间相互作用较小,势能分部较集中。甲基变形振动与伸缩振动一样,Takahiko对两种简并的弯曲和摇摆振动只分别给出了一个力常数,指认其振动频率为1460cm~(-1)、1150cm~(-1)。我们的结果区分了这四种不同的振动模式,相应得到四个力常数和各自的振动频率14680cm~(-1)、1460cm~(-1)及1088cm~(-1)、1026cm~(-1)。亚甲基的剪式变形,面内摇摆及面外摇摆分别为1439cm~(-1)、1302cm~(-1)、726cm~(-1)。其扭曲变形和弯曲变形不是强峰,不作为鉴别亚甲基的特征峰。饱和烷烃中C-C伸缩模式,指认很不统一,Snyder等认为在1060cm~(-1)左右;Takehiko等认为在830cm~(-1)附近。我们的结果为901cm~(-1),红外和拉曼光谱峰都很弱,不宜作为特征基团频率使用 。双键部分的振动,计算频率与观测值符合的很好。我们给出了双键部分与对称与反对称振动的力常数。=C-H面内对称与反对称变形振动频率分别为1263cm~(-1)、1397cm~(-1)。面外对称与反对称变形分别是784cm~(-1)、887cm~(-1)。它们的势能分布较低。=C-C对称与反对称伸缩分别为859cm~(-1)、975cm~(-1)。势能分布较文献的结果高得多。有关C=O部分观测频率数目较少,计算值与实验值符合得很好。但其中一个C-O伸缩模式相差较大,且势能分布很低。

Relevância:

100.00% 100.00%

Publicador:

Resumo:

动物的适应进化是生物学研究的最基本的问题之一。虽然,人们早从形态学上研究了动物的适应现象,但是对适应的遗传学机制知之甚少。动物感觉系统对周围环境的适应是动物适应进化中的一个关键问题。因此,我们选取了动物感觉系统中苦味受体基因家族(T2R)和犁鼻器受体1基因家族(VIR)为研究对象,对哺乳动物适应性进化的分子机制进行探讨。通过生物信息学和分子生物学手段相结合,我们在哺乳动物6个目共16个物种中获得了157个的苦味味觉受体基因,并对这些TZR基因的系统发育关系进行分析,结果显示哺乳动物的TZR基因家族经历了"生一和一灭"的进化模式,即频繁的基因重复和假基因化。另外,结果还显示这些基因可以被分为3个主要类群,分别命名为A,B和C。有趣的是,B和C类群的基因在所研究的物种间普遍是一对一的直系同源的,而A类群基因则显示了种属或世系特异性。有可能B和C类群的基因是识别哺乳动物共同的苦味物质所必需的,而A类群基因则是用于识别具有种属特异性的苦味物质。这个分析还揭示了在系统发育关系上近相关的基因在它们的染色体位置上也是靠近的,这证明了串联重复是新的TZR基因产生的主要方式。此外,通过核昔酸的异义替换数和同义替换数的比较显示不同物种新近产生的基因所编码的受体蛋白在膜外区的异义替换数显著地大于同义替换数,提示着这些通过基因重复产生的新基因受到了正选择的作用。在自然界中,许多的天然有毒物质一般都是苦的,因此,我们推测哺乳动物TZR基因的分"化选择是为了使其在探索新的生活环境和寻找新食物时能够辨别出更多不同的有毒物质,更好地适应新环境。此外,我们还研究了苦味受体基因和甜味/鲜味受体基因的进化途径。结果显示苦味受体基因和甜味/鲜味受体基因在进化上具有远相关,并具有不同的进化途径,提示着这可能是导致了这些受体基因具有不同功能,传导不同味觉的原因。犁鼻受体噬因家族(VIR)是哺乳动物的信息素受体。应用生物信息学手,我们从大鼠和小鼠的基因组中分别得到了152和115个VIR基因。大鼠VIR基因家族包含11个亚家族,其中10个是与小鼠共享的,而M家族是大鼠特有的;另夕卜大鼠缺少了H和I亚家族,而这两个亚家族存在于小鼠的基因组中。系统发育关系分析发现,"生一和一灭"进化模式也在V1R基因的进化过程中占了主导地位。所有检测到的亚家族都出现于啮齿目和灵长目分歧之后,这说明V1R基因的多样性反映了这一基因家族在啮齿目内基因重复、丢失,基因漂变及自然选择等作用的动态过程。我们的分析还表明大部分不同亚家族下的基因簇爆发的时间接近于大、小鼠分歧的时间。此外,用最大似然法分析的结果表明在这些基因簇中异义替换和同义替换的比值远大于1,揭示了正选择在这些基因的分化过程的作用。一般认为V1R在动物识别信息素过程中起重要的作用,因此我们推测V1R基因的适应性进化是为了使不同的哺乳动物能够识别不同的、复杂的信息素。

Relevância:

100.00% 100.00%

Publicador:

Resumo:

精子从男性生殖道释放后必须经过获能和顶体反应,才能与卵子结合。获能和顶体反应后,精子细胞内和精子表面的许多大分子都会发生改变。利用这些变化,可探索新的避孕方法和男性不育诊断及治疗的新途径。本文将正常人精子于体外在BWW-BSA培养基中获能,用钙离子载体A23187诱导人精子进行顶体反应,以三染和金霉素荧光染色两种方法检测这些精子的顶体反应率为50%左右,然后用这些经顶体反应处理的新鲜人精子(AR sperm)作为抗复,腹腔免疫Balb/C鼠,利用最近发展起来的半固体培养及普通的液体培养方法制备单抗。分别用未经任何处理的人精子(NT sperm)及上述顶体反应的人精子包被酶标板,用ELISA法检测所得杂交瘤细胞分泌的抗体,得到了近百株阳性杂交瘤,其中已克隆并得到腹水的有23株,并对这些单抗进行了以下鉴定:腹水滴度测定;抗体分类;与人白细胞系U937,Raji和Jurkat的交叉反应;在NT和AR人精子、树(左鼠右句)和小鼠精子上的荧光定位;单抗对人精子的凝集(SA)和制动(SI)试验;免疫印迹测定单抗相应抗原的分子量。主要结果如下:1.单抗与NT和AR精子反应的ELISA结果表明,有12个单抗主要与AR精子抗原发生反应(A组抗体),6个单抗主要与NT精子抗原反应(B组抗体),而另外5个则与这两种精子抗原都呈阳性反应(C组抗体)。2.在得到的23个单抗中,绝大多数(21个)为IgM,只有两个分别是IgC_1和IgG_(2a)。3.23个单抗与白细胞系的交叉反应强度不同。A组单抗的交叉反应有的较强,有的较弱,有的居中;B组单抗的交叉反应为弱阳性或阴性;C组单抗则呈现要么很强、要么很弱的交叉反应。4.所得单抗的荧光定位主要在赤道板和中段,未发现定位于顶体及顶体后的单抗,而国内外其他实验室已获得的单抗,主要定位在顶体。某些单抗在两种不同精子(NT sperm, AR sperm)上有不同的荧光定位。这些结果表明,AR精子的免疫原性是十分特殊的,它明显不同于NT精子。5.有9个单抗显示较强的精子凝集作用,另有9个单抗的凝集作用稍弱,未发现有精子制动效应的单抗。6.免疫印迹结果表明,有9个单抗的靶抗原是蛋白质类物质,其分子量为16-146kDa,其余14个单抗的免疫印迹呈阴性反应。有关这些单抗及其抗原的鉴定仍在进行中。其中10个单抗已送世界卫生组织(WHO),参与WHO的抗人精子抗原的单克隆抗体的研究计划。

Relevância:

100.00% 100.00%

Publicador:

Resumo:

The microstructures in iron- and sulphur-doped InP crystals were studied using both electron microscopy and electron diffraction. A modulated structure has been found in S-doped InP crystal, where the commensurate modulations corresponded to periodicities of 0.68 nm and 0.7 nm in real space and were related to the reflections of the cubic lattice in [111] and [113BAR] directions; they were indexed as q111* = 1/2(a* + b* + c*) and q113BAR* = 1/4(-a* - b* + 3c*), respectively. Single atomic layers of iron precipitate were observed, with preferred orientations along which precipitates are formed. Simulated calculations by means of the dynamical theory of electron diffraction using models for the precipitate structure were in good agreement with our experimental results. The relation between the modulated structure and the precipitates is also discussed.

Relevância:

100.00% 100.00%

Publicador:

Resumo:

A multi-finger structure power SiGe HBT device (with an emitter area of about 166μm^2) is fabricated with very simple 2μm double-mesa technology. The DC current gain β is 144.25. The B-C junction breakdown voltage reaches 9V with a collector doping concentration of 1 × 10^17cm^-3 and a collector thickness of 400nm. Though our data are influenced by large additional RF probe pads, the device exhibits a maximum oscillation frequency fmax of 10.1GHz and a cut-off frequency fτ of 1.8GHz at a DC bias point of IC=10mA and VCE = 2.5V.

Relevância:

100.00% 100.00%

Publicador:

Resumo:

制备了3种结构的器件:A:ITO/SiO2/Alq3/Al,B:ITO/Alq3/SiO2/Al,C:ITO/SiO2/Alq3/SiO2/Al。对于器件A和B,在正向偏压(ITO接正极)下才能观察到发光;而对于器件C,在正向和反向偏压下都可以观察到发光。随着电压升高,器件B和C产生的蓝色发光相对绿光逐渐增强。这主要是由于SiO2中的加速电子碰撞激发Alq3发光层产生热电子,并与空穴形成电子空穴对,复合产生蓝光;而对于器件A,在反向偏压下被热电子碰撞激发出的空穴与正向偏压下从Al电极进入的电子复合形成激子,产生绿色发光。这些结构的器件发光不但可来源于电子与积累的空穴复合,而且也来自固态阴极射线发光。

Relevância:

100.00% 100.00%

Publicador:

Resumo:

OLAND两阶段生物脱氮系统是一项正在开发且极具应用前景的处理高氨氮、低COD废水的新技术。在实验室水平,对OLAND系统限氧亚硝化阶段MBR反应器和厌氧氨氧化阶段SBR反应器的启动和运行进行了系统研究。MBR反应器控制参数在DO0.1-0.3mg/L,pH7.8±0.1,温度30±0.5℃,SRT无穷大的条件下,可实现在较高的容积负荷By=IO00mgN/L,水力停留时间HRT:ld下稳定的亚硝酸型硝化,使NH4+-N和NO2-N的出水比例达到理想的比值(1:1.20±0.20),保证厌氧氨氧化阶段SBR反应器的理想进水;SBR反应器在完全厌氧、pH7.8-8.2,温度30±0.5℃,SRT无穷大的条件下,无需外加任何有机碳源,在较高总氮负荷550mgN/L下,可实现NH4+-N和NO2--N同时稳定的去除,二者的消耗比例为1:(1.21±0.05),总氮去除率高达92%。采用巢式PCR、DGGE、 FISH等分子技术对MBR反应器硝化菌群随溶解氧的动态变化规律和SBR反应器厌氧氨氧化菌群结构、组成进行了研究,并探讨了硝化菌群与氮素组成变化之间的内在联系。结果表明:在限氧亚硝化阶段硝化菌群中氨氧化菌受溶解氧浓度的影响较大,其种群结构从反应器启动初期到稳定后期发生了非常明显的变化。亚硝酸氧化菌NOB的种群组成受溶氧影响并不明显,从启动初期到稳定后期其种群结构无明显变化。硝化菌群中氨氧化菌 AOB与亚硝酸氧化菌NOB的数量比例关系随溶解氧的降低而不断升高,从最初的3.6:1升高到稳定后期的5.5:1。MBR反应器优势硝化菌群主要由A、B、C三类氨氧化菌和硝化杆菌D、硝化螺菌F组成,其中优势菌C为维持MBR反应器稳定出水比例的主要功能菌。硝化菌群组成和结构的变化,带来了不同N素之间组成和比例的规律性变化。厌氧氨氧化阶段基本由厌氧氨氧化菌AnAOB和ANAMMOX两类菌组成,二者空间结构紧密,在反应器中的活菌数量比例分别为55%和42%。厌氧氨氧化菌AnAOB种群多样性相对比较丰富,主要由条带I和H所代表的两种优势菌组成;ANAMMOX菌种群多样性变化较小,其种群主要由优势菌K和J组成。对MBR和SBR反应器中的优势菌进行了克隆、测序和系统发育学分析,结果表明:限氧亚硝化阶段MBR反应器启动初期的优势菌A属于Nitrosomonadaceae科,是否是一个新属,还有待于进一步鉴定。运行中期优势菌B与Nitroso)nonaseurooaea亲缘关系最近,同源性高达99.1%,暂命名为Nilrosomonas sp.BI。稳定后期优势菌C与Nitrosomonas eutroPha亲缘关系最近,同源性为96.3%,暂命名为Nifrosomonas sp.cl。硝化杆菌属优势菌D与Nitrobacter alkalicus、Nitrobacter hambllrgensts和Nitlobacrwinograsky 亲缘关系较近,同源性分别为95.5-97%、96.5~97%和95.8~96.8%。SBR反应器中AnAOB优势菌I与MBR反应器优势菌B亲缘关系最近,同源性高达98.7%,与Nitlosomonas euroPaea同源性为98.3%。根据序列比较和生理特性分析,优势菌I与优势菌B应为Nitrosomonas属两个不同的种,暂将优势菌I命名为Nitrosomonas sp.II。优势菌H与MBR反应器优势菌C亲缘关系最近,同源性达97.9%,与Nitrosomonas eutropha同源性为96.3%。结合其生理特性分析,二者应为Nitrosomonas属两个不同的种,暂将优势菌H命名为Nitrosomollas sp.Hl。ANAMMox优势菌K与未培养的 Planctomycete和已鉴定的另一种ANAMMoX菌尤uenenia sf况ttgartiensis亲缘关系较近,同源性分别为99.8%和96.6%。优势菌J与Gen bank收录的所有菌的相似性均低于76%,说明该菌是OLAND系统厌氧氨氧化阶段比较独特的菌,是迄今为止在厌氧反应过程中未发现的一个新菌。

Relevância:

100.00% 100.00%

Publicador:

Resumo:

三面压缩式高超声速进气道在侧压式进气道的基础上加入顶面压缩,理论上具有的更短的压缩距离,从而有利于超声速飞行器的整体设计。该进气道不同于传统的顶压式进气道和侧压式进气道,由于其压缩激波在竖直和水平方向都存在,因此出现三维空间的激波相交和反射,造成流场复杂度大大增加,而其也直接影响了进气道设计构型与波系的相对位置关系及进气道性能。\newline 三面压缩进气道由于顶板与侧板两个方向压缩,在顶板与侧板相交线附近区域,激波并不是简单的二维平面激波结构。顶压激波和侧压激波相交后,在波型上产生了明显的变化。在角区的激波相干后的波系结构:顶板激波与侧壁激波相交后,都变得不再连续,即在相互"穿越"的过程中发生了"断裂",出现了过渡的桥波2。然而对该相干结构的认识目前也仅限于此,其流动的特征,对进气道性能的影响以及如何规划三面压缩进气道设计构型都还需要深入的探索。忽略激波边界层干扰,专注于激波相干现象本身,对于这种三维激波相干结构开展了无黏数值分析研究,探索了其关键影响因素,理论分析了其相干特征。\newline 本文分析认为角区波系可分成$A$,$B$,$C$,$D$,$E$5个部分,$A$为未经激波压缩的区域;$B$为只经过顶压激波3压缩的区域:$C$为只经过侧压激波1压缩的区域;$D$为经过顶压激波3和侧压激波1共同压缩的区域;$E$为过桥波2压缩的区域。对于顶压激波来说,$B$区域的气体与$D$区域的气体参数并不相同,因为$D$区域的气体还经过了侧压激波的压缩,因此在相同的气流转角下,$D$区域的顶板激波角大于$B$区域的顶板激波角,所以顶压激波在穿越侧压激波后发生了"断裂",7的位置高于3的位置;同理,侧压激波穿越顶压激波后,4的位置也会向对称面移动,侧压激波也发生"断裂"。基于以上物理模型,应用二维激波关系,探索性给出的三维激波相干位置的无黏近似计算方法。\newline 数值研究进一步发现相干结构产生的桥波区域为低总压区,对进气道总压恢复系数不利。

Relevância:

100.00% 100.00%

Publicador:

Resumo:

毛壳霉属(Chaetomium)和曲霉属(Aspergillus)真菌产生多种具有生物活性的化合物。为系统阐明两属微生物的次生代谢物,对三种毛壳霉、两种曲霉真菌分别进行固态发酵,以色谱和波谱技术研究发酵物中的成分,分离鉴定了51个化合物,其中23个为新化合物,测试了部分化合物对肿瘤细胞的活性。 1、从螺卷毛壳霉(C. cochloides)固态发酵物中分离鉴定了11个化合物,3个新化合物为螺卷毛壳霉素A~C(1~3)。化合物1、3及dethio-tetra (methylthio) chetomin(4)对Bre-04、Lu-04和N-04细胞株生长抑制的GI50值为0.05~7.0 μg/mL。 2、从印度毛壳霉(C. indicum)固态发酵物中鉴定的三个异喹啉生物碱印度毛壳霉素A~C(12~14)代表两类骨架新颖的异喹啉生物碱。 3、从巴西毛壳霉(C. brasiliense)固态发酵物中鉴定了11个化合物,其中Mollicellins I~J(15~16)、2-Hydroxymethyl-6-methylmethyleugenin(19)为新化合物。化合物16和Mollicellin H(18)对Bre-04、Lu-04、N-04细胞株生长抑制的GI50值在2.5~8.6 μg/mL。 4、从土曲霉(A. terreus)固态发酵物中鉴定了18个化合物。5个为新化合物为Terretonin A~D(24~27)和Asterrelenin(28),24~27为二倍半萜化合物,28为吲哚生物碱。 5、从杂色曲霉(A. versicolor)固态发酵物中鉴定了16个化合物。9个新的化合物Brevianamides K~N (40~43)、Averins A~C (44~46)和Glyanphenines A~B (47~48)代表三种类型的生物碱。 6、综述了1997-2007年间新的二倍半萜的研究进展。 The fungi of the genera Chaetomium and Aspergillus produce various secondary metabolites with biological activities. In order to systematically study the secondary metabolites, the solid-state fermented rice culture of three species of Chaetomium and two of Aspergillus were chemically studied. By the means of chromatograhy and spectroscopy, 55 compounds were isolated and identified, among of them 23 were new ones. The biological activities of some compounds were investigated. 1. From the fungus C. cochliodes, three new epipolythiodioxopiperazines, chaetocochins A-C (1-3) were isolated, together with 8 known ones (4-11). Compounds 1, 3 and 4 showed growth inhibitory effects against cancer cell lines Bre-04, Lu-04 and N-04 with GI50 values from 0.05 to 7.0 μg/mL. 2. Three novel isoquinolines Chaetoindicins A-C (12-14) were isolated and identified from the fungus C. indicum. Chaetoindicin A, Chaetoindicins B-C represented two classes of novel carbon skeletons. 3. Three new compounds, Mollicellins I-J (15-16), and 2-hydroxymethyl-6-methylmethyleugenin (19), were isolated from C. brasiliense. Compound 16 and Mollicellin H (18) showed growth inhibitory effects against cancer cell lines Bre-04, Lu-04 and N-04 with GI50 values from 2.5 to 8.6 μg/mL. 4. Eighteen compounds were isolated from the fungus A. terreus. Terretonin A-D(24 - 27)and Asterrelenin(28) are new compounds belonging to sesterterpoids and indole-ralated alkaloid, respectively. 5. From the fungus A. versicolor, sixteen secondary metabolites, including nine new ones, Brevianamides K-N (40-43), Averins A-C (44-46), and Glyanphenines A-B (47-48), were isolated and identified. Brevianamides K-N (40-43), Averins A-C (44-46), and Glyanphenines A-B (47-48) represented three classes of alkaloids. 6. New sesterterpenes and their bioactivities reported from 1997 to 2007 were summarized.

Relevância:

100.00% 100.00%

Publicador:

Resumo:

青藏高原东缘的亚高山针叶林是长江上游重要的生态屏障,经过近六十年的采伐后,取而代之的是大量人工种植的云杉纯林。目前,这些人工林已经表现出树种单一,结构层次简单等生态问题,其物种多样性及生态效益与同地带天然林相比差距较明显。如何丰富该地区物种多样性,完善人工林生态系统的生态功能是一个十分重要的课题。林下植物是人工林群落的重要组成部分,对维持群落的生物多样性及完善生态系统功能具有明显的作用。因此,研究该地区人工针叶林的林下植被对不同生境的适应性对于理解人工林生态系统物种多样性的形成和维持机制都具有重要的意义。 本文以青藏高原东部亚高山针叶林的主要森林类型----云杉人工林为研究对象,选择林下11种具有不同喜光特性的常见植物,分别设置人工林林冠下及成熟林窗为研究样地,通过对各种植物叶片形态与物质分配特征、叶片解剖学特征、叶片光合生理特性、植物自然分布特征等方面的比较分析,研究林下植物对不同光生境的适应策略及其适应能力,揭示不同物种对人工林生境的适应共性,为西南亚高山地区植被恢复及人工林的经营管理提供科学依据。具体研究结果如下: 在叶片形态和物质分配特征方面:在林窗光生境中,11种林下植物叶片比叶重(LMA)显著高于林下光生境的同种植物。同时,林窗下生长的植物叶片叶片厚度及栅栏细胞长度显著增加,这是影响叶片比叶重变化的直接原因。而多数植物叶重比在两种生境中无明显变化。说明在长期适应自然生境之后,植物可能更多地采取调节叶片组织细胞水平(即叶片功能细胞形态)及叶片器官水平(即单个叶片形态)特征的策略来适应各类生境,而非整株水平上的叶片总比重的增减。 在叶片解剖结构特征方面:多数阔叶物种栅栏组织厚度(PT)、栅栏组织厚度/海绵组织厚度(PT/ST)、栅栏细胞层数及近半数种的气孔密度(SD)在林窗生境中更大或更多,而叶片表皮细胞厚度(UET、LET)气孔长径(SL)及海绵组织厚度(ST)受两种生境影响不大。喜光特性相似的物种在生境适应策略上具有一定的趋同性。 在光合生理特征方面:在林窗生境中多数种植物的最大光合速率(Amax)、暗呼吸速率(Rd)及喜光植物光补偿点(LCP)显著或极显著高于林内生境同种植物。且在同一生境条件下,多数深度耐荫植物比喜光及轻度喜光植物有稍低的Rd和LCP。各植物在林内低光生境中具有更大的内禀光能转化效率,并在中午12:00~14:00之间光强最大的时刻发生了的最深程度的光抑制。多数种能通过调节自身某种光合素含量或色素之间的比例来适应不同的光生境,即通过增加叶绿素含量或降低Chla/b值来适应林内弱光生境,通过提高类胡萝卜素含量或单位叶绿素的类胡萝卜素含量降低强光带来的伤害。绝大多数物种并不采取调节叶片C、N含量的策略来适应不同的光生境。总之,植物部分光合参数(Amax、Rd、LCP)受生境的影响与其自身喜光特性有关,但另一些参数(Fv/Fm日变化、色素含量及比例、叶氮相对含量)受生境影响与其自身喜光特性无明显关联。 在表型可塑性方面:在叶片各表型参数中,器官水平及细胞水平的形态特征参数平均可塑性大于整株水平形态和物质分配特征参数可塑性;叶片光合组织的可塑性大于非光合组织可塑性;反映植物光合能力的参数可塑性大于叶片色素含量参数可塑性。植物叶片形态和物质分配、解剖学特征参数平均可塑性大小与其自身喜光特性基本吻合,即喜光种及轻度耐荫种各参数可塑性最高,深度耐荫种可塑性最小,而这种规律并未在光合生理参数的可塑性大小上体现出来。但是叶片形态和物质分配参数、光合生理参数的平均可塑性水平却大于叶片解剖学参数。 在植物自然分布特征方面:喜光物种云杉幼苗及歪头菜在林内生境中分布密度明显降低,深度耐荫种疏花槭却恰恰相反,更多数物种(7种植物)在两种生境中密度变化趋势不明显。从分布格局来看,7种植物在两种生境中均为聚集分布,但聚集强度为林窗>林内;少数物种桦叶荚迷、直穗小檗、冰川茶藨、黄背勾儿茶在林窗中为聚集型,在林内生境中的分布型发生改变而成为随机型,说明光生境的差异能影响到植物种群的分布特征。但这种影响程度与植物自身的喜光特性无关,同时与各物种叶片表型平均可塑性的大小也无明显关联。 The subalpine coniferous forest area in eastern Qinghai-Tibet Plateau is important ecology-barrier of upriver Yangtze. In past sixty years, those forests had been cut down and replaced with a lot of spruce plantations. At now, there are many ecology problems presenting to us such as singleness species, simple configuration, lower species diversity and ecological benefit than natural forests at the same belt. How to restore the species diversity and enhance the eco-function of the plantations is a very important issue. The understory plants are important part of plantation community, which improved the bio-diversity and eco-function distinctly of forests. So, it is very significance to study the adaptation of understory plants to different environment in plantation, and this study would helping us to understand how plantations to develop and remain their biodiversity. This study was conducted in a 60a spruce plantation in Miyaluo located in western Sichuan, China, and spruce plantation is major types of subalpine coniferous forest in eastern Qinghai-Tibet Plateau. In this paper, the leaf morphological and biomass-distributed characteristics, the anatomical characteristics, the photosynthetic characteristics and the distribution patterns characteristics of eleven different light-requirement understory species grown in two different environments (forest gaps and underneath close canopy) were studied and compared. The purpose of this study was to analyze the adaptation of this forest understory plants, to show up the commonness of these different light-requirement understory species in light acclimation, and to provide some scientific reference to manage and restore the vegetation of subalpine plantation of southwest China. The results were as follows: The leaf morphological and biomass-distributed characteristics: These eleven species in forest gaps had significantly higher dry weight per leaf area (LMA) than those under close canopy. The palisade parenchyma cells of the broad-leaved species in gaps were significantly longer than those grown under the canopy, which been a directed factor for the change of leaf mass per unit area (LMA) in different environment. But the leaf weight ratio (LWR) of most plants species were not evidently changed by the contrasted environments in our study. It was shown the morphological characteristics changing been adopted as a strategy of light acclimation for plants wasn’t on whole plant level (leaf weight ratio) but cellular level (the function cells morphological characteristics) and organic level (the leaf morphological and biomass-distributed characteristics) mostly. The leaf anatomical characteristics: Most broad-leaved plants in gaps increased palisade parenchyma thickness (PT), the palisade parenchyma cell layers and the ratio of palisade to spongy parenchyma (PT/ST). So did as almost about half species in this study in stomatal density (SD). No significant differences in thickness of leaf epidermal cells (UET, LET), stomatal length (SL) and spongy parenchyma (ST) between two environments of most species were observed. The results suggested that species with light-requirement approximately had convergent evolution on adaptation to light condition. The leaf photosynthetic characteristics: The dark respiration rate (Rd) of most plants species, the light compensation point (LCP) of light-demanding plants species in gaps were significantly increased than under close canopy in this study. In a same habitat, most deep-shade-tolerant plants had lower Rd and LCP than those light-demanding plants and slight-shade-tolerant plants. Each species has bigger inherent electron transport rate under close canopy than in gaps, and the greatest photoinhibition happened during 12 to 14 in the daytime. Most species could adapt different light environment by the way of changing their photosynthetic pigments content or the ratio of pigments content. For example, some plants under close canopy increased chlorophyll (Chl) or reduced the values of the ratio Chla/b to adapted the low light condition, some plants in gaps increased carotenoid (Car) or reduced the weight ratio CarChl to avoid been hurt in high light. For most plants, changing the content of C and N in leaf wasn’t a strategy of light acclimation. In conclusion, the variation of some leaf photosynthetic parameters in different light environment such as Fv/Fm, pigments, C and N in leaf related with the light-requirmnet of species, but the others such as Amax, Rd, LCP did not. The leaf plasticity indexes: Among those leaf plasticity indexes, the leaf morphological and biomass-distributed parameters on cellular and organic level were greater than on whole plant level for same species, and the photosynthetic parenchyma parameters were greater than non-photosynthetic parenchyma parameters in same leaf, and photosynthetic capability parameters were greater than photosynthetic pigments content parameters for same species. The average plasticity indexes of leaf morphological and biomass-distributed and anatomical parameters were accordant with plants’ light-requirement approximately: those light-demanding plants and slight-shade-tolerant plants had bigger plasticity indexes than deep-shade-tolerant plants. But this regular wasn’t observed in physiological plasticity indexes for most plants, though the average leaf plasticity indexes of leaf morphological and biomass-distributed, photosynthetic characteristics parameters was greater than the anatomical characteristics parameters. The distribution patterns characteristics: Oppositely to the deep-shade-tolerant specie Acer laxiflorum Pax., the density of light-demanding species Picea asperata Mast. and Vicia unijuga A. Br. in gaps was bigger than under close canopy. Each of the other species has the approximately density in two different environment. The spatial patterns of seven species were aggregated distribution in two environments, but the trend of aggregation of population under close canopy was decrease from in gaps. A few species such as Viburnum betulifoium Batal., Berberis dasystachya Maxim., Ribes glaciale Wall. and Berchemia flavescens Brongn. were aggregated distribution in gaps while random distribution under close canopy. It was shown that the difference between two light environments could affect the distribution pattern of plant population, and the effect didn’t relate with the light-requirement or plasticity indexes of species.

Relevância:

100.00% 100.00%

Publicador:

Resumo:

近十年,植物群体遗传学的研究飞速发展,然而与海拔相关的植物群体遗传结构和遗传变异研究却相对较少。到目前为止,还不清楚遗传变异与海拔之间是否有一个通用的格局。在山区,各种生态因子,如温度、降水、降雪、紫外线辐射强度以及土壤成分都随海拔梯度急剧变化,造成了即使在一个小的空间区域,植被类型变化显著,这种高山环境的异质性和复杂性为我们研究植物群体遗传结构和分化提供了方便。沙棘(Hippophea)属于胡颓子科(Elaeagnaceae)为多年生落叶灌木或乔木,雌雄异株,天然种群分布极为广泛。中国沙棘(H. rhamnoides subsp. sinensis)是沙棘属植物中分布较广的一个亚种,种内形态变异非常丰富,加之其具有独特的繁育系统和广泛的生态地理分布,是研究沙棘属植物遗传变异和系统分化的理想材料。本文从1,800 m 到3,400 m 分5 个海拔梯度进行取样,用RAPD 和cpSSR 分子标记研究了卧龙自然保护区中国沙棘天然群体的遗传结构和遗传变异。5 个取样群体依次标记为A、B、C、D 和E,它们分别代表分布在海拔1,800,2,200,2,600,3,000 和3,400 m 的5 个天然群体。RAPD实验用11 条寡核苷酸引物,扩增得到151 个重复性好的位点,其中143 个多态位点,多态率达94.7%。在5 个沙棘群体中,总遗传多样性值(HT)为0.289,B群体内的遗传多样性值为0.315,这完全符合沙棘这种多年生、远交的木本植物具有高遗传变异的特性。5 个群体内遗传多样性随海拔升高呈低-高-低变异趋势,在2,200 m海拔处的B群体遗传多样性达最大值0.315,3,400 m海拔处的E群体则表现最小仅0.098。5 个群体间的遗传分化值GST=0.406,也即是说有40.6%的遗传变异存在于群体间,1,800 m海拔处的A群体与其它群体的明显分离是造成群体间遗传分化大的原因。UPGMA聚类图和PCoA散点图进一步确证了5 个群体间的关系和所有个体间的关系。最后,经过Mantel检测,遗传距离与海拔表现了明显的相关性(r = 0.646, P = 0.011)。cpSSR 实验中,经过对24 对cpSSR 通用引物筛选,11 对引物能扩增出特异性条带,只有2 对引物(ccmp2 和ARCP4)呈现多态性。4 个等位基因共组合出4 种单倍型,单倍型Ⅰ出现在A 群体的所有个体和B 群体的8 个个体中,C、D、E 三个群体均不含有,而单倍型Ⅱ出现在C、D、E 三个群体的所有个体及B 群体的18 个个体中,A 群体不含有。另外两种单倍型Ⅲ和Ⅳ为稀有类型,仅B 群体中的4 个个体拥有。这种单倍型分布模式和TFPGA 群体聚类图揭示了,C、D、E 群体可能来源于同一祖先种,而A 群体却是由另一祖先种发展起来的,B 群体则兼具了这两种起源种的信息,这可能是因为在历史上的某一时期,在中国沙棘群体高山分化的过程中,B 群体处某个或者某些个体发生了基因突变,具备了适应高海拔环境的能力,产生了高海拔沙棘群体的祖先种。 In recent ten years, studies about population genetics of plants developed rapidly,whereas their genetic structure and genetic variation along altitudinal gradients have beenstudied relatively little. So far, it is uncleared whether there is a common pattern betweengenetic variation and altitudinal gradients. In the mountain environments, importantecological factors, e.g., temperature, rainfall, snowfall, ultraviolet radiation and soil substratesetc., change rapidly with altitudes, which cause the vegetation distribution varying typically,even on a small spatial scale. The mountain environments, which are heterogeneous andcomplex, facilitate and offer a good opportunity to characterize population genetic structureand population differentiation.The species of the genus Hippophae L. (Elaeagnaceae) are perennial deciduous shrubs ortrees, which are dioecious, wind-pollinated pioneer plants. The natural genus has a widedistribution extending from Northern Europe through Central Europe and Central Asia toChina. According to the latest taxonomy, the genus Hippophae is divided into six species and12 subspecies. The subspecies H. rhamnoides ssp. sinensis shows significant morphologicalvariations, large geographic range and dominantly outcrossing mating system. Thesecharacteristics of the subspecies are favourable to elucidate genetic variation and systemevolution. To estimate genetic variation and genetic structure of H. rhamnoides ssp. sinensisat different altitudes, we surveyed five natural populations in the Wolong Natural Reserve at altitudes ranging from 1,800 to 3,400 m above sea level (a.s.l.) using random amplifiedpolymorphic DNA markers (RAPDs) and cpSSR molecular methods. The five populations A,B, C, D, and E correspond to the altitudes 1,800, 2,200, 2,600, 3,000 and 3,400 m,respectively.Based on 11 decamer primers, a total of 151 reproducible DNA loci were yielded, ofwhich 143 were polymorphic and the percentage of polymorphic loci equaled 94.7%. Amongthe five populations investigated, the total gene diversity (HT) and gene diversity within population B equaled 0.289 and 0.315, respectively, which are modest for a subspecies of H.rhamnoides, which is an outcrossing, long-lived, woody plant. The amount of geneticvariation within populations varied from 0.098 within population E (3,400 m a.s.l.) to 0.315within population B (2,200 m a.s.l.). The coefficient of gene differentiation (GST) amongpopulations equaled 0.406 and revealed that 40.6% of the genetic variance existed amongpopulations and 59.4% within populations. The population A (1,800 m a.s.l.) differed greatlyfrom the other four populations, which contributes to high genetic differentiation. A UPGMAcluster analysis and principal coordinate analyses based on Nei's genetic distances furthercorroborated the relationships among the five populations and all the sampling individuals,respectively. Mantel tests detected a significant correlation between genetic distances andaltitudinal gradients (r = 0.646, P = 0.011).Eleven of the original 24 cpSSR primer pairs tested produced good PCR products, onlytwo (ccmp2 and ARCP4) of which were polymorphic. Four total length variants (alleles) werecombined resulting in 4 haplotypes. The haplotype was present in all individuals of Ⅰpopulation A and 8 individuals of populations B, the other three populations (C, D and Epopulations) did not share. The haplotype was present in all individuals of populations C, D Ⅱand E and 18 individuals of populations B, population A did not share. The other twohaplotypes and were rare haplotypes, which were only shared in 4 individuals of Ⅲ Ⅳpopulation B. The distribution of haplotypes and TFPGA population clustering map showedthat the populations C, D and E might be origined from one ancestor seed and population Amight be from another, whereas population B owned information of the two ancestor seeds. Itwas because that gene mutation within some individual or seed in the location of population Bwas likely to happen in the history of H. rhamnoides, which was the original ancestor of thehigh-altitude populations.

Relevância:

100.00% 100.00%

Publicador:

Resumo:

群落结构的确定是群落研究的基础。本文立足于个体、种群、群落三级水平,从空间和时间两种概念出发,对贡嘎山麦吊杉(Picea brachytyla)群落的结构进行了研究。经初步统计,在麦吊杉林分布的主要地段共有维管束植物93科、263属、约560种,其中种子植物71科、220属、479种。通过种子植物区系地理成份的分析表明,麦吊杉群落的区系组成具有区系成分复杂、特有现象明显和成分古老的特点。在生活型组成上,麦吊杉群落的高位芽植物所占比例最大,其中小高位芽植物最多,而在叶级谱中小叶类型占有较明显的优势。通过种—面积曲线和重要值—面积曲线两种方法求得的麦吊杉群落的最小面积在800-900平方米之间,实际应用时可选用1000平方米的保守面积。在物种的综合特征分析中,群落表现出优势种明显的特征,而存在度II占有最大的比例。物种多样性的研究表明,贡嘎山麦吊杉群落的Shannon-Wiener指数在2.33-3.26之间,Simpson指数在0.43-0.70之间,种间相遇(PIE)在0.42-0.79之间,而以Shannon-Wiener指数为基础的均匀度在34.45%-62.90%之间。麦吊杉群落的垂直结构较为复杂,分层明显。一般可划分出乔木层、灌木层、草本层和活地被层四个基本层次,乔木层通常还可划分为2-3个亚层,由高海拔至低海拔成层现象有逐渐复杂的趋势。构成麦吊杉群落的层片类型较多,其中以常绿针叶大高位芽植物为其建群层片。麦吊杉林中的层外植物较丰富,种子植物、蕨类植物、以及苔藓植物和地衣植物皆可构成其空中层片,其中苔藓植物生长繁茂,反映出群落结构的完整性。麦吊杉群落结构的水平分化较明显,经研究,其密度值在0.32-0.41之间,平均距离在1.56-1.41之间。应用Raunkiaer的频度指数方法对整个麦吊杉群落的种群频度进行调查后表明,其频度分布情况与Raunkiaer的频度定律基本上是符合的,即A>B> C

Relevância:

100.00% 100.00%

Publicador:

Resumo:

从空间、时间、食物3个方面研究了若尔盖湿地3种两栖类的种间竞争,运用生态位理论探讨了3种两栖类利用环境资源的能力以及它们之间的共存模式,研究了3种两栖类年周期食性,并考察了畜牧业对3种两栖类食性及食物竞争格局的影响;此外,还通过实验室研究对2种两栖类幼体的种间竞争策略进行了考察。主要结果如下: 1、两栖类的空间资源利用状况:在3种两栖类成体生态位宽度的比较中,岷山蟾蜍(Bufo minshanicus)成体在牧场性质(0.41)、牛粪数量(0.42)、植被盖度(0.20)、地表温度(0.50)、地表湿度(0.51) 5个维度上的生态位宽度最窄;而倭蛙(Nanorana pleskei)成体在牛粪数量(0.81)、植被高度(0.63)、植被盖度(0.47)、小水体距离(0.68) 4个维度上的生态位宽度最宽。在3种两栖类亚成体生态位宽度的比较中,岷山蟾蜍亚成体在牧场性质(0.66)、牛粪数量(0.58)、植被高度(0.64)、小水体距离(0.51)、地表湿度(0.79) 5个维度的上生态位宽度最宽;倭蛙亚成体在牧场性质(0.39)、牛粪数量(0.30)、地表温度(0.18)、地表湿度(0.33) 4个维度上的生态位宽度最窄。高原林蛙(Rana kukunoris)在地表温度(成体:0.62;亚成体:0.56)、地表湿度(成体:0.84;亚成体:0.60)两个维度上具有较大的生态位宽度值,而在小水体距离维度上(成体:0.27;亚成体:0.14)的生态位宽度值则很小。比较3种无尾两栖类在不同生长阶段(成体、亚成体)的生态位宽度,发现高原林蛙和倭蛙的亚成体对栖息环境的要求更高。3种两栖类空间资源利用的相似程度很高,高原林蛙与倭蛙之间的生态重叠度(0.87)较之它与岷山蟾蜍(0.81)的生态位重叠度更大。 2、两栖类的日活动节律:高原林蛙成体、亚成体、岷山蟾蜍亚成体活动的最低气温为0℃、2℃、8℃;岷山蟾蜍和高原林蛙亚成体出现的数量与气温成极显著的正相关(r=0.797, p<0.001;r=0.794, p<0.001),高原林蛙成体出现的数量与气温有一定相关性(r=0.456, p<0.05);晴天时两栖类的活动性明显高于阴天(p<0.001);多云转晴天气,高原林蛙和岷山蟾蜍亚成体出现两次日活动高峰,分别为中午12:30左右和下午15:30~16:30之间;多云天气,高原林蛙和岷山蟾蜍亚成体出现两次日活动高峰,分别为9:30~10:30之间和15:30~16:30之间。 3、两栖类的食物资源利用状况:春、秋两季,高原林蛙最主要的食物是蜉金龟科(Aphodiidae)昆虫,相对重要性指数(IRI)最高(春季:35.28%,秋季:28.57%),其次为昆虫的幼虫,以及双翅目的毛蚊科(Bibionidae)、蝇科(Muscidae)、丽蝇科(Calliphoridae)昆虫,秋季,蝗虫是高原林蛙食物组成中的重要部分;岷山蟾蜍最主要的食物是蚂蚁(IRI,春季:85.54%,秋季:49.70%),其次为蜉金龟科、象甲科(Curculionidae)、步甲科(Carabidae)、粪金龟科(Geotrupidae) 等鞘翅目昆虫;倭蛙春季的最主要食物也是蜉金龟科昆虫(IRI,春季:13.41%),其次为蚂蚁、毛蚊科昆虫、昆虫的幼虫以及狼蛛科(Lycosidae)。3种两栖类中,倭蛙的食性生态位宽度相对较宽(0.43),而岷山蟾蜍(0.09)和高原林蛙(0.22)的生态位宽度较窄,与春季相比,两栖类在秋季的食谱更宽。以利用食物种类为标准,春季高原林蛙与倭蛙的生态位重叠度(0.40)比它与岷山蟾蜍的生态位重叠度(0.33)更大。 4、畜牧业对两栖类食性及食物竞争格局的影响:以藏牦牛粪为食物或寄居场所的昆虫,如蜉金龟科、粪金龟科、毛蚊科、蝇科、丽蝇科昆虫和某些昆虫幼虫,是3种两栖类食物谱中最主要的组成部分,蜉金龟科昆虫在高原林蛙食谱中的比例更高,高原林蛙可能从畜牧业发展中获得更多的好处,使之在食物竞争方面处于优势地位。与无放牧样地相比,在有放牧样地的中,两栖类食谱中的蜉金龟科昆虫数量更多(有放牧:31.94%;无放牧:21.32%)、出现频率更高(有放牧:76.38%;无放牧:44%)。然而在不同样地上(有放牧/无放牧),两栖类的食物组成无显著性差异(P=0.188),两栖类的数量(P=0.075)、肥满度(P=0.537)均没有显著差别。 5、两栖类幼体的竞争策略:实验室条件下,通过活动性水平,变态时的体重、增长率和完成变态所需时间考察自然条件下常同水塘分布的中华蟾蜍(Bufo gargarizans)和高原林蛙蝌蚪的竞争策略。结果表明:中华蟾蜍蝌蚪在不同食物资源条件下,所选择的生存策略可能不同,即食物资源充足时,增加活动性获取更多食物,食物资源有限时,降低活动性且提前完成变态;与中华蟾蜍蝌蚪相比,在食物资源有限时高原林蛙蝌蚪获取食物能力可能更强。 This paper presented the study of competition of three amphibians (Rana kukunoris, Nanorana pleskei, Bufo minshanicus) based on spatial, temporal and dietary scales in Zoige wetland. We measured coexistence patterns of three amphibians and analyzed their ability of exploiting resource. Effects of grazing on the diet composition and diet competition of amphibians were analyzed by their diet composition during spring and autumn. Furthermore, we examined the competitive ability of larval common frogs (Rana kukunoris)and common toads(Bufo gargarizans) in a laboratory experiment, and analyzed their competitive strategies respectively. The results were as follows: 1 .The status of using spatial resource Niche breadths of B. minshanicus adults on 5 dimensional axes including character of pasture(0.41), number of yaks dung(0.42), vegetation coverage(0.20), temperature (0.50)and humidity(0.51) of ground surface were narrower than adults of R. kukunoris and N. pleskei. Niche breadths of B. minshanicus subadults were broader than R.kukunoris subadults and N.pleskei subadults on 5 dimensional axes including character of pasture (0.66), number of yaks dung (0.58), vegetation height (0.64), distance to small waterbodies (0.51), humidity of ground surface (0.79). Niche breadths of N. pleskei subadults were the narrowest in three anurans subadults on 4 dimensional axes including character of pasture (0.39), number of yaks dung (0.30), temperature (0.18) and humidity (0.33) of ground surface, niche breadths of N. pleskei adults were the broadest in three anurans adults on 4 dimensional axes including number of yaks dung (0.81), vegetation height (0.63) and coverage(0.47), distance to small waterbodies(0.68).Comparatively, niche breadths of R. kukunoris were broader on the two microclimate factors including temperature(adults:0.62;subadults:0.56) and humidity (adults:0.84;subadults:0.60)of ground surface, but was narrow on distance to small waterbodies(adults:0.27;subadults:0.14). Strategies for using habitat resource of adults and subadults of the three species anuran were different. Generally, subadults of R. kukunoris and N. pleskei needs better habitat condition. It was quite similar that three anurans exploited spatial resource, Niche overlap between R. kukunoris and N. pleskei (0.87) was greater than that between R. kukunoris and B.minshanicus(0.81). 2.Daily activity rhythm R. kukunoris audlts were active when air temperatures were as low as 0℃, R. kukunoris subadults were active at 2℃, B.minshanicus subaudlts were active at 8℃. Positive correlation was found between activities of amphibians and air temperature, Subadults of R.kukunoris, (r=0.797, p<0.001), Subadults,of,B.minshanicus, (r=0.794, p<0.001), andbadults,of,R.kukunoris(r=0.456, p<0.05).Amphibians were more active during sunny days than cloudy days. In cloudy turning into sunny, R. kukunoris and B.minshanicus subadults had two active peak: at noon about 12:30 and 15:30~16:30 pm; in cloudy, R. kukunoris and B.minshanicus subadult had two active peak too : 9:30~10:30am,15:30~16:30pm. 3.Diet analysis Aphodiidae was the most commonly consumed food item by R. kukunoris based on index of relative importance (IRI) during spring (35.28%) and autumn (28.57%) in Zogie wetland. Besides Aphodiidae, larval insect, dipterans such as Bibionidae, Muscidae, Calliphoridae also were important food item for R. kukunoris, in autumn, locust was one of important food item for R. kukunoris. The most important food item for B.minshanicus during spring (IRI:85.54%) and autumn (IRI:49.70%) was ants, following, was coleopterans, such as Aphodiidae, dung beetle. Aphodiidae (IRI:13.41%) were the most important consumed food item by N. pleskei during spring too, following, was ants and Bibionidae. Dietary breadth of N. pleskei (0.43) were greater than R. kukunoris (0.22) and B. minshanicus (0.09). As a whole, Dietary breadth of amphibians during aurumn were greater than spring. Based on prey item, dietary overlap between R. kukunoris and N. pleskei (0.40) was greater than that between R. kukunoris and B.minshanicus (0.33) during spring. 4.Effects of grazing on the diet composition and diet competition of amphibians Amphibians are an important part of the pasture ecosystems as prey and predator. In Zogie wetland, major diet of amphibians was closely associated with dung of yaks, for example, Aphodiidae, Bibionidae, Muscidae, dung beetle. Dung of yaks was major diet and habitat of these insects. Proportion of Aphodiidae was higher in diet composition of R. kukunoris than N. pleskei and B.minshanicus, with development of pasturage, R. kukunoris may have a diet competitive advantage over N. pleskei and B.minshanicus. Number of Aphodiidae in diet composition of amphibians was higher in samples with grazing (31.94%) than in those without grazing (21.32%). Occurrence Frequency of Aphodiidae in diet composition of amphibians was higher in samples with grazing (76.38%) than in those without grazing (44%). However, There was not significantly different on diet composition (P=0.188), and number (P=0.075) and the relative fatness (P=0.537) of amphibians between grazing samples and without grazing. 5.Competitive strategies of amphibian larvae I examined the competitive ability of larval toads (Bufo gargarizans) and frogs (Rana kukunoris) which co-occur in the nature pond by activity level, the growth rate and mass at metamorphosis and larval period in a laboratory experiment. The results suggest: In laborary, B.gargarizans adapted himself to different food level by changing activity. At high food level, B. gargarizans increased activity to gain more diet. At low food level, B. gargarizans decreased activity and achieved early metamorphosis. When food resource was limit, R. kukunoris could gain more food than B. gargarizans.

Relevância:

100.00% 100.00%

Publicador:

Resumo:

角蟾科(Megophryidae)是以角蟾属(Megophrys Kuhl and Van Hasselt, 1822)为模式属而建立的,隶于无尾目(Anura),变凹型亚目(Anomocoela)。角蟾科包括2 亚科11 属142 种,分布于东洋界,从巴基斯坦、中国西部向东直到菲律宾和苏达群岛;中国有9 属75 种分布于华中和华南地区。角蟾科被认为是原始的两栖动物之一,其分类学、系统学、生态学、动物地理学的研究均深受中外科学家的瞩目。近年来,通过形态学、古生物学、细胞学、生态学、支序系统学的研究,角蟾科的分类与系统学研究取得了较大进展。与成体形态和分子系统学研究结果相比较,蝌蚪的研究存在更多的问题和挑战,尚需深入研究:(1)角蟾科蝌蚪的形态多样性分析;(2)角蟾科的系统发育关系与蝌蚪的演化,以及口漏斗的起源;(3)角蟾科蝌蚪表型分化与栖息环境和觅食行为的适应演化。针对上述问题,本文对角蟾科9 属30 种蝌蚪的形态特征,包括外部宏观形态和口器外部结构特征、口器内部显微结构、唇齿和角质颌的亚显微结构作了深入细致、多层次的比较研究;通过12s rRNA 和cytochrome b 基因构建最大简约树,采用贝叶斯系统发育进行分析,蝌蚪型的演化采用祖先性状的重建方法分析;得到如下结论:1)初步将角蟾科蝌蚪分为4 种类型;并且建立了2 种新的角蟾科蝌蚪类型。A 型:拟髭蟾型蝌蚪,该型蝌蚪包括拟髭蟾属、髭蟾属、齿蟾属和齿突蟾属的物种;B 型:新类型,掌突蟾型蝌蚪,该型蝌蚪在本文中包括掌突蟾属、小臂蟾属的物种;C 型:新类型,短腿蟾型蝌蚪,一种特化类型,该型蝌蚪在本文中仅包括短腿蟾属的物种;D 型:角蟾型蝌蚪,该型蝌蚪在本文中包括无耳蟾属、小口拟角蟾属和异角蟾属的物种。2)对角蟾科的分类进行了修订:(1)支持角蟾科两个亚科的分类系统;(2)角蟾亚科包括拟角蟾属、异角蟾属、无耳蟾属和短腿蟾属;该亚科形态差异小,系统学关系比较复杂,暂不作族级分类的再划分;(3)拟髭蟾亚科分为2 个族:拟髭蟾族,该族物种具有类型A 的蝌蚪,包括4 个属:拟髭蟾属、髭蟾属、齿蟾属、齿突蟾属;掌突蟾族,该族物种具有类型B 的蝌蚪,包括2 个属:掌突蟾属和小臂蟾属。3)结合分子系统进化关系探讨了4 种蝌蚪类型的演化。(1)角蟾科蝌蚪的最近共同祖先来自于一类具有拟髭蟾型蝌蚪性状的蝌蚪;(2)掌突蟾型蝌蚪和角蟾亚科的蝌蚪是由具有拟髭蟾型蝌蚪性状的祖先蝌蚪分别演化而来;(3)短腿蟾型蝌蚪是角蟾型蝌蚪的一种特化类型;(4)外群蝌蚪具有与拟髭蟾型蝌蚪相似的性状,进一步印证了类拟髭蟾型蝌蚪是角蟾科蝌蚪的最近共同祖先的假说;(5)具有口漏斗的蝌蚪类型是由不具口漏斗的蝌蚪类型演化而来,在角蟾科中口漏斗是一种衍生性状。4)分析了角蟾科四种蝌蚪类型与栖息环境的适应演化。(1)角蟾科蝌蚪的口部和体形的变化反映了该科蝌蚪由缓流向类似静水生境的回水凼的渐变式适应,角蟾科蝌蚪的形态显示了多方面的适应变化;(2)随着蝌蚪类型由A 向D的演化,当水速较大时,拟髭蟾型的蝌蚪营流水攀吸型生活方式;当水速递减时,掌突蟾型蝌蚪营流水附着型生活方式;当水速进一步递减时,具有较小口漏斗的短腿蟾型蝌蚪和具有大漏斗的角蟾型蝌蚪营流水浮泳型生活。角蟾科蝌蚪对于水流递减的适应演化说明蝌蚪的生态学适应是具有进化意义的;(3)蝌蚪口器内部结构的分化揭示了蝌蚪和食性的适应关系,蝌蚪以口部的唇齿与角质颌刮取或吞吸水中的物质,然后,通过口乳突有选择地过滤进入口腔中食物。拟髭蟾亚科蝌蚪的唇齿多而窄,唇齿间距宽,颌鞘粗而稀,反映了其植食性为主的特点;它们的舌前乳突一般为指状,在口腔入口处所占面积小,其机械过滤的作用很多被唇齿和角质颌分担了;而角蟾亚科的蝌蚪,其角质颌弱,其舌前乳突一般为匙状,几乎填满了口腔入口处,因此舌前乳突起了主要的机械过滤作用。The family Megophryidae is the largest and most diverse families inArchaeobatrachia, and most of its species occur in India, Pakistan, and eastward intoChina, Southeast Asia, Borneo and the Philippines to the Sunda Islands. Currently thefamily includes 142 species have been grouped into two subfamilies, Megophryinaeand Leptobrachiinae. The mountains of central and southern China are rich in speciesof Megophryidae, 75 species belong to 9 genera and two subfamilies.The family was supposed to be ideal materials of studies in many fields of biology,such as taxonomy, evolution, systematics, ecology, and biogeography. Recently, therehave a great development in taxonomy and systematics of megophryids throughstudied by morphology, paleontology, cytology, ecology, and cladistics. However,larvae of megophryids were generally unknown, although the tadpoles might be veryimportant for above studies.In this paper, we examined the evolutionary scenario of the tadpoles’ morphologyin the context of a phylogenetic framework. Our objectives are (1) to evaluate thedivergence of larval body shape and oral discs in the family Megophryidae, (2) toexplore the evolutionary trends of the larvae in megophryidae, and test if thefunnel-shaped oral disc is apomorphic, and (3) to explore the relationship of the larvalstructure, diet and microhabitat.We examined larval morphology of 30 megophryid species, the larval body shape,oral discs, the buccopharyngeal cavity, and jaw sheaths and denticles of the Chinesemegophryid frogs were re-examined. We constructed a phylogeny of the species on thebasis of published mitochondrial cytochrome b and 16S rRNA gene segments usingpartitioned Bayesian analyses. Furthermore, hypothetical changes of larval morphologywere inferred using parsimony principle on the phylogeny. The results showed that:1) Four tadpole types in Megophryidae. The larval morphological charactersseries in Chinese megophryids fall into four general categories according to the bodyshape and oral discs: (A) Leptobrachiini type, species from genera Leptobrachium,Oreolalax, Scutiger and, Vibrissaphora share this type of tadpoles. (B) Leptolalax type,species of genus Leptolalax have this type of tadpoles. (C) Brachytarsophrys type,species of the genus Brachytarsophrys have this type of tadpoles. (D) Megophryinitype, species of the genera Atympanophrys, Ophryophryne, and Xenophrys share this type of tadpoles. Of which B and C are two novel types.2)Taxonomic implications. The present study leads us to reconsider the generalclassification of tribes attributed to members of Megophryidae. More specifically,concerning the phylogenetic relationships and the two novel tadpole types describedherein, we propose a provisional taxonomy for the family but suggest that further taxasampling of other megophryids be performed to confirm this taxonomic change. TheMegophryidae is composed of two subfamilies (Leptobrachiinae and Megophryinae).The Leptobrachiinae was recogonized the two tribes: (1) tribe Leptobrachiini sensuDubois, corresponding to the tadpole of type A, including four genera, i.e.,Leptobrachium, Oreolalax, Scutiger and, Vibrissaphora; (2) tribe Leptolalaxini,corresponding to the tadpole of novel type B, including two genera, i.e., Leptolalaxand Leptobrachella. However, the relationships among the genera of Megophryinaewere largely unresolved, they recognized no monophyletic groups above the generalevel. A more thorough sampling will likely foster a better taxonomic solution.3) The larval evolutionary scenario in Megophryidae.Type A is characteristicof normal-mouthed with multiple tooth rows, representing the tadpole type of theMRCA of Chinese megophryids. Type B is characteristic of normal-mouthed withreduced tooth rows, prolonging labium, and integumetary glands. Type C ischaracteristic of no labial teeth and smaller umbeliform oral disc. Type D ischaracteristic of no labial teeth, enlarged umbeliform oral disc, representing the tadpoleof the MRCA of subfamily Megophryinae. A previous hypothesis, referring tofunnel-shaped oral discs as an apomorphy, is supported.4) The larval adaptation to habitats in Megophryidae. Tadpoles generallyadhere to substrates using their mouths, and the microhabitat that the tadpoles occupyreflects the degree of adhesion and oral complexity. The morphological changes inmegophryid tadpoles virtually allow a progressive adaptation to a changing habitatfrom faster water to slower water. Within the tadpoles of Type A to type D, the TOTbecomes smaller and smaller, and the oral disc orientates from anteroventral toumbelliform upturned, and eye position orientates from dorsal to lateral, and the trunkis more and more depressed and tail becomes relatively longer and slender. Within therunning water, the normal-mouthed with multiple tooth rows of Leptobrachiini tadpoles are correlated with lotic-suctorial, benthic feeders with anteroventral oraldisc and the largest body. With the water’s velocity decreasing, the lotic-adherentfeeders of Leptolalax tadpoles have tube-shaped labium with reduced tooth rows andintegumetary glands. And then, the smaller umbeliform in Brachytarsophrys tadpolesand the enlarged umbeliform oral disc in the Megophryini tadpoles are inhabitmicrohabitats of non-flowing backwaters of rivers, indicative of adaptive traits oflotic-neustonic surface feeders. The scheme of megophryid tadpoles andmicrohabitats provided the first clear evidence which congruent with the hypothesis ofAltig and Johnston (1989). The ecological divergence plays a general role in thedivergence and evolution of megophrid larvae. There is a definite correlation amongthe buccopharyngeal cavity, diet and feeding mechanisms, the tadpole graze orswallow the food particles, then through papillae which like a sieve and sort out foodparticles to the oesophagus. The tadpole of Leptobrachiinae possess multiple toothrows, wide intertooth distance as well as thick and sparse jaw sheath, these tadpolesinhabit bottom of the streams and graze on epiphyton or major detritus of organicmatter on the substrates, their prelingual papillae like single finger, the mechanicalpurpose of papillae served share in by tooth and jaw. The tadpoles of Megophryinaeoccur near the water surface of small streams and are the filter feeder, their dietincludes plankton and organic debris floating on the water surface, those tadpolepossess weak jaw, their prelingual papillae like spoon, the mechanical purpose ofpapillae served mostly for sieve.