Blood-gas properties of plateau zokor (Myospalax baileyi)

Plateau zokor (Myospalax baileyi) is one of the blind subterranean mole rats that spend their life solely underground in sealed burrows. It is one of the special species of the Qinghai-Tibet plateau. In their burrows, oxygen is low and carbon dioxide is high and their contents fluctuate with the change of seasons, soil types, rain and depth of burrows. However, plateau zokors show successful adaptation to that extreme environment. In this study, their adapting mechanisms to the hypoxic hypercapnic environment were analyzed through the comparison of their blood-gas properties with that of pikas (Ochotona curzniae) and Sprague-Dawley rats. The results indicated that plateau zokors had higher red blood corpuscle counts (8.11+/-0.59 (10(12)/L)) and hemoglobin concentrations (147+/-9.85 g/L), but hematocrit (45.9+/-3.29%) and mean corpuscular volume (56.67+/-2.57 fL) were lower than the other rodents. Their arterial blood and venous blood pH were 7.46+/-0.07 and 7.27+/-0.07. Oxygen pressure in arterial blood of plateau zokors was about 1.5 times higher than that of pikas and rats, and it was 0.36 and 0.26 times in their venous blood. Partial pressure for carbon dioxide in arterial and venous blood of plateau zokors was 1.5-fold and 2.0-fold higher, respectively, than in rats and pikas. Oxygen saturation of plateau zokors was 5.7 and 9.3 times lower in venous blood than that of pikas and rats, respectively. As result, the difference of oxygen saturation in arterial blood to venous blood was 2- and 4.5-fold higher in plateau zokors as that of pikas and rats, respectively. In conclusion, plateau zokors had a high tolerance to pH changes in tissues, together with strong capabilities to obtain oxygen from their hypoxic-hypercapnic environment.     

Diet selection in overwinter caches of plateau zokor (Eospalax baileyi)

Subterranean herbivores affect the plant community by plant consumption and burrowing activities. However, diet selection of subterranean herbivores has not been studied in detail in complex natural fields, mainly for lack of an accurate method to determine diet species, frequency, and biomass. Plateau zokors' (Eospalax baileyi) caching habit for the long inclement winter makes it possible to solve this problem. We studied the diet composition and biomass in caches and vicinity of plateau zokors' burrow systems. We found that plateau zokors are dietary generalist but show a significant selection among the available food items, plant parts, functional groups, organ types, and habitats. These results suggested that plateau zokors strictly selected their diet and were able to adjust their foraging strategy according to the different conditions of food abundance and quality. Plateau zokors' selective foraging can directly reduce the proportion of toxicity forbs; plateau zokors can be recognized as an important element to influence the alpine meadow plant community and cattle husbandry.     

Mitochondrial genome of Plateau zokor Myospalax baileyi

Plateau zokors Myospalax baileyi are subterranean rodents and endemic to the cold and hypoxic (2800-4300 m above sea level) climate of the farm, prairie, alpine prairie, and meadow habitats across the Tibetan plateau. The complete mitochondrial (mt) genome of M. baileyi has been determined. Our results showed that the total length of the mitogenome was 16,351 bp, and had a gene content of 13 protein coding, 22 tRNAs and 2 rRNAs. Except for the seven tRNA and Nd6 genes, all other mt genes are encoded on the heavy strand. The overall base composition of the heavy strand is 33.65% A, 29.65% T, 24.42% C, and 12.28% G, with an AT content of 63.3%.     

青藏高原5种害鼠D型肉毒素抗性相关基因VAMP1的序列分析

突触小泡相关膜蛋白1基因(VAMP1)的变异是导致鼠类对D型肉毒梭毒素灭鼠剂产生抗性的主要原因。本研究利用转录组测序的方法,分析青藏高原地区5种主要害鼠:高原鼠兔(Ochotona curzoniae)、高原鼢鼠(Eospalax baileyi)、长尾仓鼠(Cricetulus longicaudatus)、青海松田鼠(Neodon fuscus)和喜马拉雅旱獭(Marmota himalayana)的VAMP1序列信息。同时,分别采集来自5个地理种群的58只高原鼠兔和59只高原鼢鼠,对VAMP1基因第二外显子区序列进行分析。结果显示,从转录组组装文件中成功获得5种动物的VAMP1基因全序列,长度均为357 bp,共检测到46个核苷酸变异位点和4个氨基酸变异位点,但未发现与D型肉毒素抗性相关的氨基酸位点。对高原鼠兔群体和高原鼢鼠群体的VAMP1基因第二外显子序列的分析显示,高原鼠兔所有个体的序列高度保守,而在高原鼢鼠中则存在一个同义突变位点,但两物种在D型肉毒素抗性相关位点上都未监测出位点变异。该研究结果提示,D型肉毒杀鼠剂在青藏高原地区害鼠防治方面应该可以长期发挥重要作用。     

高原鼢鼠对高寒草甸植被特征及生产力的影响

本研究结果表明,高原鼢鼠栖息10年的斑块,植物群落的物种数减少,植物物种多样性指数下降,地上、地下总生物量显降低,单子叶和可利用双子叶植物生物量极显减少,但不可利用双子叶植物生物量显增加。高原鼢鼠去除5年后,斑块内植物群落的单子叶植物物种数增加,而双子叶植物下降,植物群落物种多样性指数下降,地上、地下总生物量显增加,单子叶和可利用双子叶植物生物量增加极显,不可利用双子叶植物生物量显降低。高原鼢鼠栖息10年的斑块,净初级生产量较未栖息地区减少68.98%。高原鼢鼠去除5年后,净初级生产量增加,但仅达到未栖息地区的58.69%。     

高原鼢鼠prestin基因的组织特异性表达与适应地下生活的关系

高原鼢鼠是青藏高原特有的地下鼠,地下鼠具有准确的空间定位能力。prestin蛋白是在耳蜗特异性表达且与回声定位有关的蛋白分子。为了探讨prestin基因与高原鼢鼠地下空间定位之间的关系,我们克隆了高原鼢鼠prestin基因的编码区序列,并与其他物种的prestin基因序列进行序列比对;运用PAML软件对高原鼢鼠的prestin基因进行进化分析;根据已克隆的序列,应用实时荧光定量的方法测定高原鼢鼠耳蜗、尾部、足垫和鼻垫组织中prestin基因mRNA的表达水平。研究结果表明,与人、大鼠、小鼠、裸鼢鼠、家兔和牛6种哺乳类动物相比,高原鼢鼠prestin基因编码的氨基酸序列显示存在9个氨基酸残基突变;Test2模型未检测到统计上显著的正选择位点;高原鼢鼠耳蜗prestin基因mRNA的表达水平显著高于高原鼠兔(P<0.05),高原鼢鼠耳蜗和尾部prestin基因mRNA的表达水平显著高于足垫和鼻垫（P<0.01）。以上结果说明,prestin基因不仅在高原鼢鼠耳蜗中表达,而且在尾部、足垫和鼻垫组织中也有表达。高原鼢鼠在地下洞道生活过程中,可能利用尾巴、前后足和鼻子辅助感知低频声波,从而准确地进行空间定位。     

Function‐related Drivers of Skull Morphometric Variation and Sexual Size Dimorphism in a Subterranean Rodent,Plateau Zokor (Eospalax baileyi)

Sexual dimorphism is prevalent in most living organisms. The difference in size between sexes of a given species is generally known as sexual size dimorphism (SSD). The magnitude of the SSD is determined by Rensch's rule where size dimorphism increases with increasing body size when the male is the larger sex and decreases with increasing average body size when the female is the larger sex. The unique underground environment that zokors (Eospalax baileyi) live under in the severe habitat of the Qinghai‐Tibetan Plateau (QTP) could create SSD selection pressures that may or may not be supported by Rensch's rule, making this scientific question worthy of investigation. In this study, we investigated the individual variation between sexes in body size and SSD of plateau zokors using measurements of 19 morphological traits. We also investigated the evolutionary mechanisms underlying SSD in plateau zokors. Moreover, we applied Rensch's rule to all extant zokor species. Our results showed male‐biased SSD in plateau zokors: The body‐ and head‐related measurements were greater in males than in females. Linear regression analysis between body length, body weight, and carcass weight showed significant relationships with some traits such as skull length, lower incisor length, and tympanic bulla width, which might support our prediction that males have faster growth rates than females. Further, the SSD pattern corroborated the assumption of Rensch's rule in plateau zokors but not in the other zokor species. Our findings suggest that the natural underground habitat and behavioral differences between sexes can generate selection pressures on male traits and contribute to the evolution of SSD in plateau zokors.     

甘南草原高原鼢鼠年龄划分及其组成分析

建立高原鼢鼠(Eospalax baileyi)种群年龄鉴定划分标准,可为对其的监测及防控提供科学依据。研究于2015年5月、10月分别在甘南草原碌曲县采用夹捕法捕捉高原鼢鼠469只(雄鼠209只,雌鼠260只),在对样本进行逐个解剖和数据记录之后,参考其体重、体长、胴体重及毛色等,对高原鼢鼠的年龄划分标准进行研究。依据胴体重将高原鼢鼠分为6组:亚成年组(雄144 g,雌106 g),成年Ⅰ组(雄144~210 g,雌106~148 g)、成年Ⅱ组(雄211~276 g,雌149~190 g)、成年Ⅲ组(雄277~342 g,雌191~232 g)、成年Ⅳ组(雄343~408 g,雌233~274 g)和老年组(雄409 g,雌275 g)。年龄结构表显示,不同的季节和不同地理区域高原鼢鼠的种群年龄结构类型均为增长型。     

高原鼢鼠、高原鼠兔和大鼠肺表面活性物质组成和含量的比较

本研究通过分析肺表面活性物质(pulmonary surfactant,PS)组成和含量探讨高原鼢鼠(Myospalax baileyi)和高原鼠兔(Ochotona curzoniae)的低氧适应机制.高原鼢鼠和高原鼠兔各36只,捕捉于海拔3 600 m左右的青海省海南州贵德县拉脊山地区,36只Sprague-Daw...     

高原鼢鼠肺组织细胞凋亡相关 基因的进化分析及其在不同 海拔条件下的表达模式

高原鼢鼠（Myospalax baileyi）是青藏高原特有的地下鼠,其地下洞道严重缺氧。一般来说,低氧会促进细胞凋亡。为了探讨高原鼢鼠适应低氧环境的分子机制,本文应用生物信息学方法对p53下游凋亡促进基因Pidd、Fas、Bax、Puma、Apaf-1、Scotin、Perp、Igfbp3和凋亡抑制基因Bcl-2的序列和编码的氨基酸序列进行了进化分析,并以SD大鼠（Rattus norvegicus）为对照,研究了这些基因在不同海拔环境条件下（3 300 m和2 260 m）的表达模式。结果表明：（1）高原鼢鼠细胞凋亡基因的序列与以色列鼹鼠（Nannospalax galili）同源性最高;预测的PIDD、PUMA、Apaf-1、IGFBP3和BCL-2编码蛋白结构域与以色列鼹鼠的存在明显的趋同进化位点;SIFT评估发现,高原鼢鼠和以色列鼹鼠与其他物种相比,p53、PIDD、PUMA、Apaf-1和IGFBP3氨基酸序列分别在78、853、157、320和285号位点的变异对其功能有显著影响;（2）在高海拔条件下（3 300 m）,高原鼢鼠肺组织中凋亡促进基因Pidd、Bax、Puma和Apaf-1表达水平显著下降,凋亡抑制基因Bcl-2表达水平显著升高,而在SD大鼠中凋亡促进基因和凋亡抑制基因的表达水平均没有变化;高原鼢鼠中Bcl-2/Bax比值随海拔的升高显著上升,而在SD大鼠中没有变化。以上结果提示,高原鼢鼠p53结构变异可能导致其下游基因表达模式与SD大鼠不同,其中凋亡促进基因Pidd、Bax、Puma和Apaf-1表达水平下降,凋亡抑制基因Bcl-2表达水平上升,从而抑制了细胞在低氧条件下的凋亡;在长期低氧的作用下,高原鼢鼠p53下游基因产物PIDD、PUMA、Apaf-1和IGFBP3产生了影响其功能的变异位点,这可能改变了它们与发挥功能的复合物的结合力,从而抑制了细胞凋亡。因此,通过长期的低氧适应,高原鼢鼠肺组织中与细胞凋亡相关的基因产物结构发生变异,导致其基因表达水平发生变化,从而抑制细胞凋亡,这是高原鼢鼠适应地下低氧洞道生境的分子机制之一。     

The activity of a subterranean small mammal alters Afroalpine vegetation patterns and is positively affected by livestock grazing

Subterranean rodents can act as ecosystem engineers by shaping the landscape due to soil perturbation and herbivory. At the same time, their burrow density is affected by environmental conditions, vegetation and anthropogenic factors. Disentangling this complex interplay between subterranean rodents and their environment remains challenging. In our study, we analysed the interplay of abiotic conditions, vegetation patterns and human land-use and the burrow density of the giant root-rat (GRR; Tachyoryctes macrocephalus), a subterranean rodent endemic to the Afroalpine ecosystem of the Bale Mountains in south-east Ethiopia. Specifically, we examined the effects of GRR on plant species richness and vegetation cover and vice-versa, and how these reciprocal effects might be modulated by temperature, habitat wetness and grazing. Our results showed that increasing GRR burrow density led to decreased vegetation cover, and that effects of GRR on vegetation cover were slightly stronger than vice-versa. Considering the reciprocal causation models, we found that increasing plant species richness led to increased GRR burrow density, while GRR burrow density decreased as vegetation cover increased. Increases in habitat wetness and livestock grazing intensity also directly led to increased GRR burrow density. Our results stress the importance of subterranean ecosystem engineers on vegetation and highlight the vulnerability of these complex interactions to human activity.     

基于线粒体控制区的序列变异分析青海东部甘肃鼢鼠遗传多样性

甘肃鼢鼠(Myospalax cansus)是一种终年营地下独居生活的小型掘土类动物。本文通过测定mt DNA的控制区部分序列(530 bp)变异,分析青海东部地区8个甘肃鼢鼠地理种群遗传多样性与遗传结构。158个样本共发现26个变异位点,定义了39种单倍型,整体的平均单倍型多样性高(h=0.953 2)、核苷酸多样性低(π=0.006 36)。歧点分布和中性检验均说明青海东部甘肃鼢鼠种群在历史上存在着快速扩张的事件。基于邻接法构建的网络关系图中,单倍型呈星状分布,没有按地理位置形成对应类群。基因流(Nm)数据显示多数地理种群间基因交流贫乏,AMOVA结果显示种群内与种群间遗传变异分别为48.82%和51.18%,遗传分化明显。IBD分析表明,甘肃鼢鼠的遗传分化与地理距离呈正相关,说明距离隔离对甘肃鼢鼠种群分化具有重要作用。甘肃鼢鼠的这种遗传多样性与种群遗传结构特点,可能是地下生活方式靠挖掘迁移带来的较小扩散能力的结果。     

Climate Change-Induced Range Expansion of a Subterranean Rodent: Implications for Rangeland Management in Qinghai-Tibetan Plateau

Disturbances, both human-induced and natural, may re-shape ecosystems by influencing their composition, structure, and functional processes. Plateau zokor (Eospalax baileyi) is a typical subterranean rodent endemic to Qinghai-Tibetan Plateau (QTP), which are considered ecosystem engineers influencing the alpine ecosystem function. It is also regarded as a pest aggravating the degradation of overgrazed grassland and subject to regular control in QTP since 1950s. Climate change has been predicted in this region but little research exists exploring its impact on such subterranean rodent populations. Using plateau zokor as a model, through maximum entropy niche-based modeling (Maxent) and sustainable habitat models, we investigate zokor habitat dynamics driven by the future climate scenarios. Our models project that zokor suitable habitat will increase by 6.25% in 2050 in QTP. The predication indicated more threats in terms of grassland degradation as zokor suitable habitat will increase in 2050. Distribution of zokors will shift much more in their southern range with lower elevation compare to northern range with higher elevation. The estimated distance of shift ranges from 1 km to 94 km from current distribution. Grassland management should take into account such predictions in order to design mitigation measures to prevent further grassland degradation in QTP under climate change scenarios.     

Phylogeny of Helicobacter felis sp. nov., Helicobacter mustelae, and related bacteria.

Strain CS1T (T = type strain) is a gram-negative, microaerophilic, urease-positive, spiral-shaped bacterium that was isolated from the gastric mucosa of a cat. Additional strains which possessed biochemical and ultrastructural characteristics similar to those of strain CS1T were isolated from the gastric mucosa of cats and dogs. The guanine-plus-cytosine content of the DNA of strain CS1T was 42.5 mol%. The 16S rRNA sequences of strain CS1T, strain DS3 (a spiral-shaped isolate from a dog), and Helicobacter mustelae were determined by direct RNA sequencing, using a modified Sanger method. These sequences were compared with the 16S rRNA sequences of Helicobacter pylori, "Flexispira rappini," Wolinella succinogenes, and 11 species of campylobacters. A dendrogram was constructed based upon sequence similarities. Strains CS1T and DS3 were very closely related (level of similarity, 99.3%). Two major phylogenetic groups were formed; one group consisted of strains CS1T and DS3, H. mustelae, H. pylori, "F. rappini," and W. succinogenes, and the other group contained the true campylobacters. The average level of similarity between members of these two groups was 84.9%. Within the first group, strains CS1T and DS3, H. pylori, and H. mustelae formed a cluster of organisms with an interspecies similarity level of 94.5%. The phylogenetic positions of W. succinogenes and "F. rappini" were just outside this cluster. On the basis of the results of this study, we believe that strains CS1T (= ATCC 49179T) and DS3 represent a new species of the genus Helicobacter, for which we propose the name Helicobacter felis.     

营地下生活鼠的种类及其分布

地下鼠是一类生活在特殊生境中的啮齿动物,是研究动物适应进化的理想材料。全世界现存地下鼠主要隶属于5科,3亚科,25属,约160种。在现有研究的基础上,结合资料对地下鼠的八个主要类群及其主要的分布区作以综述,以期为开展地下鼠生理生态学和行为学研究提供基础资料。     

高原鼢鼠和高原鼠兔红细胞低氧适应特征

为探讨高原鼢鼠对低氧高二氧化碳洞道生境及高原鼠兔对高海拔低氧生境的适应机制,用Sysmex SF-3000血细胞分析仪及聚丙烯酰胺凝胶电泳对两种高原动物的血常规及血红蛋白类型进行分析,后者采用聚丙烯酰胺凝胶电泳法。结果表明,高原鼢鼠和高原鼠兔的红细胞数(RBC)、红细胞压积(HCT)及平均红细胞容积(MCV)组间无显著差异(P>0.05),但高原鼢鼠和高原鼠兔的红细胞数显著高于SD大鼠,红细胞压积及平均红细胞容积均显著低于SD大鼠(P<0.05);高原鼢鼠的血红蛋白浓度(HBC)与SD大鼠无显著差异(P>0.05),但显著高于高原鼠兔的HBC(P<0.05)。高原鼢鼠血红蛋白主要有2种类型,高原鼠兔血红蛋白主要有3种类型,而SD大鼠血红蛋白主要有5种类型。从血红蛋白电泳迁移来看,2种高原动物血红蛋白类型有明显的趋同特征并与SD大鼠具有明显的差异。上述结果提示,长期适应高海拔低氧环境的高原动物的红细胞和血红蛋白表现出趋同进化,同时因生境和习性的差异又表现出各自的特异性。     

The impact of digging on craniodental morphology and integration

The relationship between the form and function of the skull has been the subject of a great deal of research, much of which has concentrated on the impact of feeding on skull shape. However, there are a number of other behaviours that can influence craniodental morphology. Previous work has shown that subterranean rodents that use their incisors to dig (chisel‐tooth digging) have a constrained cranial shape, which is probably driven by a necessity to create high bite forces at wide gapes. Chisel‐tooth‐digging rodents also have an upper incisor root that is displaced further back into the cranium compared with other rodents. This study quantified cranial shape and upper incisors of a phylogenetically diverse sample of rodents to determine if chisel‐tooth‐digging rodents differ in craniodental morphology. The study showed that the crania of chisel‐tooth‐digging rodents shared a similar place in morphospace, but a strong phylogenetic signal within the sample meant that this grouping was nonsignificant. It was also found that the curvature of the upper incisor in chisel‐tooth diggers was significantly larger than in other rodents. Interestingly, most subterranean rodents in the sample (both chisel‐tooth and scratch diggers) had upper incisors that were better able to resist bending than those of terrestrial rodents, presumably due to their similar diets of tough plant materials. Finally, the incisor variables and cranial shape were not found to covary consistently in this sample, highlighting the complex relationship between a species’ evolutionary history and functional morphology.     

利用转录组数据分析高原鼢鼠和裸鼹鼠基因的趋同进化

地下鼠由于长期适应于地下洞道生活,在诸多生物学特征上都产生了一定的趋同进化现象,但目前尚未在分子水平分析其趋同进化特征。本研究利用高通量测序数据,从氨基酸位点变化和基因表达量水平,分析两种亲缘关系较远的地下鼠--高原鼢鼠和裸鼹鼠的趋同进化。氨基酸位点变化分析显示,两种地下鼠的54 个基因都存在至少一个理论趋同进化位点,其中13 个基因可以找到相对明确的表型功能注释。基因表达量分析显示,两种地下鼠在转录组水平的总体表达模式上存在趋同效应;有103 个基因在两种地下鼠脑中的表达量是作为对照的地上鼠(小鼠、豚鼠)表达量的3 倍以上,其中有20 个基因可以找到相对明确的表型功能注释。本研究的结果表明,高原鼢鼠和裸鼹鼠为了适应地下生活方式,在氨基酸序列和基因表达量水平上都产生了一定程度的趋同进化。其中,低氧适应相关基因(氧运输、氧代谢、DNA 修复等)是地下鼠进化适应的最重要组成部分,挖掘器官(尤其是肌肉)功能相关基因的重要性次之,能量代谢及平衡相关基因也是地下鼠适应性进化的关键要素。     

祁连山东段高原鼢鼠(Eospalax baileyi)土丘空间分布格局及其与环境因子的空间关联性

高原鼢鼠推土造丘行为对高寒草地生态系统的生产和生态功能有重要影响,研究高原鼢鼠土丘空间分布格局及其与环境因子的关系,可以揭示高原鼢鼠栖息地利用和选择规律,为合理控制鼠害及保护草地生物多样性提供科学依据。于2014年8月在祁连山东段选取面积为140m×100m的高原鼢鼠栖息地,消除景观尺度取样带来的气候、地形和土壤的异质性,利用地统计学方法,分析高原鼢鼠土丘的空间分布格局、并揭示其与环境因子中土壤容重、土壤水分、植物地上、地下生物量、根系营养物质含量(可溶性糖、粗蛋白、粗脂肪)以及各功能群丰富度(禾本科、莎草科、杂类草)的空间关系。半方差函数及普通克里格插值表明,高原鼢鼠土丘存在中等程度的空间变异且呈现聚集分布,各环境因子均存在不同程度的空间异质性。交方差函数分析表明,高原鼢鼠分布虽与各环境因子在多种尺度下表现出复杂的空间关联性(正的或负的),但mantel检验发现土壤容重、莎草科丰富度与高原鼢鼠土丘分布呈现显著的负空间关联性,杂类草丰富度和根系粗脂肪含量与高原鼢鼠土丘分布存在显著正空间关联性。综上所述,高原鼢鼠主要栖息利用在土壤疏松、莎草科丰富度较低、杂类草较多和根系粗脂肪含量较高的地方。