海拔高度对青海沙蜥成体神经发生影响的研究
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摘要
在二十世纪九十年代,人们自成年哺乳动物脑中分离出一种能不断进行自我更新分化的细胞,即神经干细胞,它能终生不断地产生新的神经元和神经胶质细胞。而到口前为止,有关爬行类动物成体神经发生的研究较少。青海沙蜥作为我国西北地区特有的蜥蜴品种,在我国青藏高原海拔2000米到4500米均有分布,生活于不同海拔高度对个体发育具有很大影响。
目的：以青藏高原特有物种青海沙蜥为动物模型,探索和研究我国青藏高原不同海拔生存环境对于动物成体神经干细胞增殖和新生细胞存活的影响。
方法：分别在我国青藏高原海拔2500m、3500m、4500m的地区各捕获30只健康成年雄性青海沙蜥,随机分为三组,每组10只。通过进行BrdU活体注射标记新生细胞,分别让其存活24小时、7天、28天,结合免疫组织化学DAB显色法等技术,对新生细胞的增殖和存活进行定量研究,分析和比较青藏高原不同海拔环境中青海沙蜥成体神经干细胞的增殖速率和存活情况。
结果：高海拔组青海沙蜥端脑AON区域中的阳性细胞数目比中海拔组和低海拔组分别减少了48.9%和46.9%(P<0.05),表明高原环境抑制了青海沙蜥神经干细胞的增殖。新生细胞存活7天至28天过程中,高海拔组神经干细胞数量出现了显著上升至显著下降的变化,表明高原环境影响青海沙蜥新生细胞存活。
结论：在我国的高海拔环境中生活的青海沙蜥,其成体神经干细胞的增殖明显受抑制,同时伴随着新生细胞的存活受到明显的影响。
In the nineteen nineties, people have found that neural stem cells still exist in human and animals, and these neural stem cells can constantly generate new neurons and astrozytes in life span. But so far, the study of reptiles' adult neurogenesis is less. As a typical lizard species in Qinghai-Tibetan Plateau in northwest China, Phrynocephalus vlangalii locate at an altitude from2000m to4500m, Living at different altitude has great influence on individual development.
Objective:The aim of present study is to investigate the influence of high-altitude living environment on the proliferation of adult neural stem cells and survival of newborn cells in Phrynocephalus vlangalii.
Methods:At altitude of2500m,3500m,4500m in Qinghai-Tibetan Plateau in northwest China, we captured30healthy adult male Phrynocephalus vlangalii in each area, and divided them into three groups randomly, each group has ten experimental animals. Phrynocephalus vlangalii were intraperitoneally injected with BrdU in order to label dividing cells, respectively let them survive for24h,7d and28d. DAB-immunocytochemistry was performed. Microscope imaging was used to observe the location newborn cells and to count the number of BrdU-positive cells.
Results:In the region of AON of Phrynocephalus vlangalii, the BrdU-positive cells in high-altitude group significantly reduced by48.9%and46.9%compared with that of low-altitude group and intermediate-altitude group(P<0.05), indicating that the high-altitude environment inhibits the proliferation of adult neural stem cells in Phrynocephalus vlangalii. In the process of survival of the newborn cells from7days to28days, the number of neural stem cells in high-altitude group showed significantly increasing and declining, indicating that the high-altitude environment influence the survival of newborn cells in phrynocephalus vlangalii.
Conclusion:In the high-altitude environment, the proliferation of adult neural stem cells in Phrynocephalus vlangalii is significantly decreased, and the survival of newborn cells is significantly affected.
目的：以青藏高原特有物种青海沙蜥为动物模型,探索和研究我国青藏高原不同海拔生存环境对于动物成体神经干细胞增殖和新生细胞存活的影响。
方法：分别在我国青藏高原海拔2500m、3500m、4500m的地区各捕获30只健康成年雄性青海沙蜥,随机分为三组,每组10只。通过进行BrdU活体注射标记新生细胞,分别让其存活24小时、7天、28天,结合免疫组织化学DAB显色法等技术,对新生细胞的增殖和存活进行定量研究,分析和比较青藏高原不同海拔环境中青海沙蜥成体神经干细胞的增殖速率和存活情况。
结果：高海拔组青海沙蜥端脑AON区域中的阳性细胞数目比中海拔组和低海拔组分别减少了48.9%和46.9%(P<0.05),表明高原环境抑制了青海沙蜥神经干细胞的增殖。新生细胞存活7天至28天过程中,高海拔组神经干细胞数量出现了显著上升至显著下降的变化,表明高原环境影响青海沙蜥新生细胞存活。
结论：在我国的高海拔环境中生活的青海沙蜥,其成体神经干细胞的增殖明显受抑制,同时伴随着新生细胞的存活受到明显的影响。
In the nineteen nineties, people have found that neural stem cells still exist in human and animals, and these neural stem cells can constantly generate new neurons and astrozytes in life span. But so far, the study of reptiles' adult neurogenesis is less. As a typical lizard species in Qinghai-Tibetan Plateau in northwest China, Phrynocephalus vlangalii locate at an altitude from2000m to4500m, Living at different altitude has great influence on individual development.
Objective:The aim of present study is to investigate the influence of high-altitude living environment on the proliferation of adult neural stem cells and survival of newborn cells in Phrynocephalus vlangalii.
Methods:At altitude of2500m,3500m,4500m in Qinghai-Tibetan Plateau in northwest China, we captured30healthy adult male Phrynocephalus vlangalii in each area, and divided them into three groups randomly, each group has ten experimental animals. Phrynocephalus vlangalii were intraperitoneally injected with BrdU in order to label dividing cells, respectively let them survive for24h,7d and28d. DAB-immunocytochemistry was performed. Microscope imaging was used to observe the location newborn cells and to count the number of BrdU-positive cells.
Results:In the region of AON of Phrynocephalus vlangalii, the BrdU-positive cells in high-altitude group significantly reduced by48.9%and46.9%compared with that of low-altitude group and intermediate-altitude group(P<0.05), indicating that the high-altitude environment inhibits the proliferation of adult neural stem cells in Phrynocephalus vlangalii. In the process of survival of the newborn cells from7days to28days, the number of neural stem cells in high-altitude group showed significantly increasing and declining, indicating that the high-altitude environment influence the survival of newborn cells in phrynocephalus vlangalii.
Conclusion:In the high-altitude environment, the proliferation of adult neural stem cells in Phrynocephalus vlangalii is significantly decreased, and the survival of newborn cells is significantly affected.
引文
[1]Eriksson P S, Perfilieva E, Thomas Bjork-Eriksson, et al. Neurogenesis in the adult human hippocampus[J]. Nature Medicine,1998,4(11):1313-1317.
[2]Egger B, SGold K, Brand A H. Notch regulates the switch from symmetric to asymmetric neural stem cell division in the Drosophila optic lobe [J]. Development,2010, 137(18):2981-2987.
[3]Ferron S R, Pozo N, Laguna A, et al. Regulated Segregation of Kinase Dyrk1 A during Asymmetric Neural Stem Cell Division Is Critical for EGFR-Mediated Biased Signaling[J]. Cell Stem Cell,2010,7(3):367-379.
[4]Nishimura T. Molecular mechanisms for asymmetric cell division in Drosophila neural stem cells[J]. Tanpakushitsu Kakusan Koso,2009,54(11):1363-1369.
[5]Walczak P, Kedziorek D, Gilad A, et al. Applicability and limitations of MR tracking of neural stem cells with asymmetric cell division and rapid turnover:the case of the shiverer dysmyelinated mouse brain[J], Magnetic Resonance in Medicine,2007,58(2):261-269.
[6]Pagano S F, Impagnatiello F, Girelli M, et al. Isolation and Characterization of Neural Stem Cells from the Adult Human Olfactory Bulb[J]. Stem Cells,2002,18(4):295-300.
[7]Duan X, Kang E, Liu C Y, et al. Development of neural stem cell in the adult brain[J]. Current Opinion in Neurobiology,2008,18(1):108-115.
[8]Ming G, Song H. Adult neurogenesis in the mammalian central nervous system[J]. Annual Review of Neuroscience,2005(28):223-250.
[9]Zhao C, Deng W, Gage F H. Mechanisms and functional implications of adult neurogenesis[J]. Cell,2008,132(4):645-660.
[10]Cameron H A, Mckay R D. Adult neurogenesis produces a large poo! of new granule cells in the dentate gyrus[J]. The Journal of Comparative Neurology,2001,435(4):406-417.
[11]Gage F H. Mammalian neural stem ceIls[J]. Science,2000,287(5457):1433-1438.
[12]Lledo P, Alonso M, Grubb M S. Adult neurogenesis and functional plasticity in neuronal circuits[J]. Nature Reviews Neuroscience,2006,7(3):179-193.
[13]Carleton A, Petreanu L T, Lansford R, et al. Becoming a new neuron in the adult olfactory bulb[J]. Nature Neuroscience,2003,6(5):507-518.
[14]AlvarezBuylla A, Jose Manuel Garcia-Verdugo. Neurogenesis in adult subventricular zone[J]. The Journal of Neuroscience,2002,22(3):629-634.
[15]Goldman S A, Nottebohm F. Neuronal production, migration, and differentiation in a vocal control nucleus of the adult female canary brain[J]. Proceeding of the National Academy of Sciences of the United States of America,1983,80(8):2390-2394.
[16]AlvarezBuylla A, Ling C, Yu W S. Contribution of neurons born during embryonic and adult life to the brain of adult canaries:Regional specificity and delayed birth of neurons in the song-control nuclei[J]. Journal of Comparative neurology,1994,347(2):233-248.
[17]Bernocchi, G, Scherini, E., Nano, R. Developmental patterns in the rat cerebellum after cis-dichlorodiammineplatinum treatment[J]. Neuroscience.1990,39(1):179-188.
[18]Polenov A, Chetverukhin V. Ultrastructural radioautographic analysis of neurogenesis in the hypothalamus of the adult frog, Rana temporaria, with special reference to physiological regeneration of the preoptic nucleus.I.Ventricular zone cell proliferation[J]. Cell and tissue research,1993,291 (2):341-350.
[19]EM D, Fingerlin A, Hwang D, et al. Seasonal Cell Proliferation in the Chemosensory Epithelium and Brain of Red-Backed Salamanders, Plethodon cinereus[J]. Brain,Behavior and Evolution,2000,56(1):1-13.
[20]Raucci F, Fiore M M D, Pinelli C, et al. Proliferative activity in the frog brain:a PCNA-immunohistochemistry analysis[J]. Journal of Chemical Neuroanatomy,2006, 32(2-4):127-142.
[21]Scoville W, Milner B. Loss of recent memory after bilateral hippocampal lesions[J]. Neurol Neurosurg Psychiatry,1957,20(1):11-21.
[22]Corkin S, Sullivan E, Carr F. Prognostic factors for life expectancy after penetrating head injury[J]. Archives of neurology,1984,41(9):975-977.
[23]Schmolck H, Kensinger E A, Corkin S, et al. Semantic knowledge in patient HM and other patients with bilateral medial and lateral temporal lobe lesions[J]. Hippocampus,2002, 12(4):520-533.
[24]Corkin S. What's new with the amnesic patient HM?[J]. Nature Reviews Neuroscience, 2002,3(2):153-160.
[25]Rakic P. Neurogenesis in adult primate neocortex:an evaluation of the evidence[J]. Nature Reviews Neuroscience,2002,3(1):65-71.
[26]Hayes N, Nowakowski R. Dynamics of cell proliferation in the adult dentate gyrus of two inbred strains of mice[J]. Developmental Brain Research,2002,134(1-2):77-85.
[27]Selden D J R, Dolbeare F, Clair J H, et al. Statistical confirmation that immunofluorescent detection of DNA repair in human fibroblasts by measurement of bromodeoxyuridine incorporation is stoichiometric and sensitive[J]. Cytometry,1993,14(2):154-167.
[28]Eng L F. Glial fibrillary acidic protein (GFAP):the major protein of glial intermediate filaments in differentiated astrocytes[J]. Journal of Neuroimmunology,1985(8):203-214.
[29]Mullen R, Buck C, Smith, et al. NeuN, a neuronal specific nuclear protein in vertebrates [J]. Development,1992(161):201-211.
[30]Dyer M A, Livesey F J, Cepko C L, et al. Proxl function controls progenitor cell proliferation and horizontal cell genesis in the mammalian retina[J]. Nature Genetics, 2003(34):53-58.
[31]Gdalyahu A, Levy I G. DCX, a new mediator of the JNK pathway[J]. The EMBO Journal, 2004(23):823-832.
[32]Gerdes J, Lemke H, Baisch H, et al. Cell cycle analysis of a cell proliferation-associated human nuclear antigen defined by the monoclonal antibody Ki-67[J]. The Journal of Immunology,1984,133(4):1710-1715.
[33]Hall D P A, Levison D A, Woods A L, et al. Proliferating cell nuclear antigen (PCNA) immunolocalization in paraffin sections:An index of cell proliferation with evidence of deregulated expression in some, neoplasms[J]. The Journal of Pathology,1990,162(4):285-294.
[34]Lewis P F, Emerman M. Passage through mitosis is required for oncoretroviruses but not for the human immunodeficiency virus.[J]. Journal of Virology,1994,68(1):510-516.
[35]Prendergast F G, Mann K. G. Chemical and physical properties of aequorin and the green fluorescent protein isolated from Aequorea forskalea[J]. Biochemistry,1978,17(17):3448-3453.
[36]Tsien R Y. The green fluorescent protein[J]. Annu. Rev. Biochem,1998(67):509-544.
[37]姚檀栋,刘晓东,王宁练.青藏高原地区的气候变化幅度问题[J].科学通报,2000,45(1)：98-106.
[38]林振耀,赵昕奕.青藏高原气温降水变化的空间特征[J].中国科学,1996,26(4)：354-358.
[39]张静.高原病研究的新进展(综述)[J].中国城乡企业卫生,2009,12(06)：17-19.
[40]1. Bojana B. Beleslin-Cokic, Cokic V P, Yu X, etal. Erythropoietin and hypoxia stimulate erythropoietin receptor and nitric oxide production by endothelial cells[J]. Blood,2004, 104(7):2073-2080.
[41]Li, Y.X., Ding, S.J.,et al. Desferoxamine preconditioning protects against cerebral ischemia in rats by inducing expressions of hypoxia inducible factor 1α and erythropoietin[J]. Neurosci Bull,2008,24(2):89-95.
[42]杨燕,马慧萍,张汝学,等.急性高原病发病机制的研究进展[J].医学综述,2010,16(17)：2561-2563.
[43]于永才.高原病的发病及防治[J].社区医学杂志,2010,8(24)：10-13.
[44]周其全,刘福玉,郑必海,等.急性重症高原病患者脑功能损伤与多器官功能障碍[J].世界急危重病医学杂志,2007,4(3)：1836-1840.
[45]吕国蔚.医学神经生物学[M].第2版.北京：高等教育出版社,2000：13-399.
[46]Michael Ward(国家体委体育科学研究所运动医学研究所等译)Mountain Medicine[M]第1版.北京：人民卫生出版社,1978：203-312.
[47]刘重庆.高原医学[M].1983,(2)：19.
[48]Reite M, Jackson D, Cahoon R L, et al. Sleep physiology at high altitude[J]. Electroencephalography and Clinical Neurophysiology,1975,38(5):463-471.
[49]MA L P, Oxfd B, BA M W, etal. Some effects of high altitude on man[J]. The Lancet,1956, 268(6953):1115-1121.
[50]蒲小燕,沈国平.高原低氧SD大鼠海马神经细胞的结构改变[J].医学信息,2010,23(1)：234.
[51]尹宏,张先钧,张晓岩.急性高原低氧对大鼠海马神经细胞形态结构的影响[J].青海医学院学报,2007,28(1)：29-31.
[52]张先钧,张晓岩,蒲小燕,等.急性和亚急性高原低氧大鼠海马神经细胞的电镜研究[J].青海医学院学报,2005,26(4)：236-239.
[53]赵霞,张先钧,张晓岩,等.业急性高原低氧大鼠海马神经细胞的电镜研究[J].高原医学杂志,2006,16(2)：59-60.
[54]吴穹,郭新庆.高原低氧应激诱导大鼠海马神经元凋亡[J].菏泽医学专科学校学报,2009,21(3)：1-2.
[55]李文林,沈娟志.高原环境对大鼠海马神经元凋亡的影响[J].包头医学院学报,2010,26(4)：9-10.
[56]高原,李源,蔡晓红,等.慢性问歇低氧对幼鼠海马神经细胞凋亡及慢性问歇低氧对幼鼠海马神经细胞凋亡及慢性问歇低氧对幼鼠海马神经细胞凋亡及caspase-12表达的影响[J].温州医学院学报,2009,39(5)：422-425.
[57]Lu X, Chen X, Weng J, et al. Hippocampal spine-associated Rap-specific GTPase-activating protein induces enhancement of learning and memory in postnatally hypoxia-exposed mice[J]. Neuroscience,2009,162(2):404-414.
[58]Zhang J, Chen X, Du J, et al. Neonatal exposure to intermittent hypoxia enhances mice performance in water maze and 8-arm radial maze tasks[J]. Journal of Neurobiology,2005, 65(1):72-84.
[59]余小燕,马福海.低氧诱导因子-1在高原低氧训练中的研究现状与进展[J].辽宁体育科技,2010,32(2)：19-21.
[60]Poon E, Harris A L, Ashcroft M. Targeting the hypoxia-inducible factor (HIF) pathway in cancer[J]. Expert Reviews in Molecular Medicine,2009(11):1-23.
[61]王雪莲,李占全,冀林华.HIF-1 α,VEGF在慢性高原低氧环境下SD大鼠骨髓,心脏中的表达及意义[J].山东医药,2009,49(9)：34-35.
[62]刘丽愉,刘力芳,曹亚.低氧诱导因子信号通路的新进展[J].国际病理科学与临床杂志,2011,31(3)：218-222.
[63]Lathia J D, Mattson M P, Cheng A. Notch:from neural development to neurological disorders[J]. Journal of Neurochemistry,2008,107(6):1471-1481.
[64]Nefedova Y, Cheng P, Alsina M, et al. Involvement of Notch-1 signaling in bone marrow stroma-mediated de novo drug resistance of myeloma and other malignant lymphoid cell lines[J]. Blood,2009,29(10):1644-1654.
[65]Arumugam T V, Chan S L, Jo D, et al. Gamma secretase-mediated Notch signaling worsens brain damage and functional outcome in ischemic stroke[J]. Nature Medicine,2006, 12(6):621-623.
[66]Gustafsson M V, Zheng X, Pereira T, et al. Hypoxia requires notch signaling to maintain the undifferentiated cell state[J]. Developmental Cell,2005,9(5):617-628.
[67]沈晓玲,映红.高原低氧对白主神经系统的影响[J].高原医学杂志,2008,18(3)：54-57.
[68]马晓庆,李琳娜.高原缺氧对神经系统的影响与护理[J].中华护理杂志,1991,36(10)：439-441.
[69]田开新,覃军,黄岚,等.急进高原自主神经功能变化与急性高原反应的关系[J].解放军预防医学杂志,2008,26(1)：9-13.
[70]隆敏,覃军.急进高原自主神经系统功能变化研究进展[J].解放军预防医学杂志,2008,26(2)：147-149.
[71]张勤文.不同海拔地区牦牛的肺组织形态学比较[J].中国兽医科技,2004,34(2)：73-74.
[72]陈秋生,冯霞,姜生成.牦牛肺脏高原适应性的结构研究[J].中国农业科学,2006,39(10)：2107-2113.
[73]江家椿,嘎玛仁增,何玛丽.不同海拔高度西藏高原牦牛若干血液生理常值的比较[J].畜牧兽医学报,1991,22(1)：20-26.
[74]司本辉,张庆成,李积荣,等.久居高原地区缺氧环境对红细胞膜蛋白结构和血液流变学的影响[J].高原医学杂志,2001,11(3)：26-27.
[75]Jin Y, Liu N, Li J. Elevational variation in body size of Phrynocephalus vlangalii in the North Qinghai-Xizang (Tibetan) Plateau[J]. Belg. J. Zool.,2007,137(2):197-202.
[76]金园庭,田仁荣,刘迺发.四种沙蜥的形态地理变异Bergmann和Allen规律的检验[J].动物学报,2006,52(5)：838-845.
[77]赵肯堂.中国的沙蜥属研究[J].动物学杂志,1997,32(1)：15-19.
[78]李俊,郭宪光,王跃招.沙蜥属Phrynocephalus生物地理学研究进展[J].四川动物,2010,29(1)：144-150.
[79]王跃招,曾晓茂,方白立,等.沙蜥属一有效种贵德沙蜥及红原沙蜥的分类研究(蜥蜴亚目：鬣蜥科)[J].动物分类学报,2002,27(2)：372-383.
[80]赵肯堂.巾国胎生爬行纲动物[J].生物学通报,1988(1)：1-3.
[81]吴鹏飞,王跃招,王硕果,等.青海沙蜥(蜥蜴亚目：鬣蜥科)种群的年龄结构与性比[J].四川大学学报(自然科学版),2002,39(6)：1134-1139.
[82]江耀明,黄庆云,赵尔宓.青海沙蜥一新亚种及其生态初步观察[J].动物学报,1980,26(2)：178-183.
[83]吴鹏飞,王跃招,郭海燕,等.青海沙蜥的生长及两性生长差异[J].四川大学学报(自然科学版),2005,42(6)：1252-1257.
[84]赵肯堂.沙蜥的泄殖系统及其繁殖习性[J].动物学杂志,1964(4)：158-186.
[85]鲍敏,曾阳,马建滨,等.青海沙蜥和密点麻蜥的分布状况和食性[J].青海师范大学学报(自然科学版),1998,4(4)：42-45.
[86]吾玛尔阿布力孜.东疆沙蜥和密点麻蜥的食性研究[J].干旱区研究,1992,16(4)：10-13.
[87]邵虹.青海沙蜥成体神经发生的特征描述[D].兰州大学,2011.
[88]H.Shao, L.Fan, X.J.Xu, et al. Characterization of adult neurogenesis in lizard Phrynocephalus vlangalii (Agamidae:Reptilia)[J]. Italian Journal of Zoology. 2012,79(4):547-558.
[89]伍冠一,廖锦锋,唐宗湘.爬行动物皮层的细胞构筑与纤维联系[J].四川动物,2006,25(2)：416-419.
[90]农万廷,石哪.石龙子脑核团定位研究[J].南宁师范高等专科学校学报,2002,19(2)：45-47.
[91]Marchioro M, Nunes J D A M, Ramalho A R, etal. Postnatal neurogenesis in the medial cortex of the tropical lizard Tropidurus hispidus[J]. Neuroscience,2005,134(2):407-413.
[92]DelgadoGonzalez F, GonzalezGranero S, TrujilloTrujillo C, etal. Study of adult neurogenesis in the gallotia gal lot i lizard during different seasons[J]. Brain Research, 2011(1390):50-58.
[93]颜坤,王荫亭,王书荣.蜥蜴中脑的Golgi-Cox研究[J].科学通报,1982(6)：371-373.
[94]王书荣,颜坤,王荫亭,等.蜥蜴中脑神经通路和起源细胞的形态[J].解剖学报,1989,20(2)：159-164.
[95]伍冠一.蛤蚧皮质加厚区的细胞构筑与纤维联系研究[D].广西师范大学,2006.
[96]伍冠一,廖锦锋,唐宗湘.爬行动物脑皮质加厚区的结构与功能[J].动物医学进展,2006,27(3)：105-107.
[97]F.J. Delgado-Gonzalez, AlonsoFuentes A, DelgadoFumero A, etal. Seasonal differences in ventricular proliferation of adult Gallotia galloti lizards[J]. Brain Research,2008(1191):39-46.
[98]王典群.四种荒漠蜥蜴脑和脑神经的比较解剖[J].兰州大学学报(自然科学版).1992,38(8)：145-149.
[99]张法奎.似哺乳类爬行动物和哺乳类动物脑颅侧壁构造类型的演替[J].古脊椎动物学报,1989,27(3)：209-218.
[100]LOPEZGARCIA C, MOLOWNY A, NACHER J, etal. The lizard cerebral cortex as a model to study neuronal regeneration[J]. Anais da Academia Brasileira de Ciencias,2002, 74(1):85-104.
[1]Reynolds BA, Weiss S.Generation of neurons and astrocytes from isolated cells of the adult mammalian central nervous system[J].Science,1992,255(5052):1707-1710.
[2]Guo-li Ming,Hongjun Song.Adult neurogenesis in the mammalian central nervous system[J].Annual Review of Neuroscience,2005,28:223-250.
[3]Elizabeth Gould.How widespread is adult neurogenesis in mammalsfJ].Nature Reviews Neuroscience,2007,8(6):481-488.
[4]Xin Duan,Eunchai Kang,Cindy Y Liu,et al.Development of neural stem cell in the adult brain[J].Current Opinion in Neurobiology,2008,18(1):108-115.
[5]Fred H.Gage.Mammalian neural stem cells[J].Science,2000,287(5457):1433-1438.
[6]Chunmei Zhao, Wei Deng,Fred H.Gage.Mechanisms and functional implications of adult neurogenesis[J].Cell.2008,132(4):645-660.
[7]Heather A.Cameron,Ronald D.GMckay.Adult neurogenesis produces a large pool of new granule cells in the dentate gyrus[J].Journal of Comparitive Neurology,2001,435(4):406-417.
[8]Pierre-Marie Lledo,Mariana Alonso & Matthew S,Grubb.Adult neurogenesis and functional plasticity in neuronal circuits[J].Nature Reviews Neuroscience,2006,7(3):179-193.
[9]Alan Carleton,Leopoldo T.Petreanu,et al.Becoming a new neuron in the adult olfactory bulb[J].Natural Neuroscience,2003,6(5):507-518.
[10]Arturo Alvarez-Buylla,Jose Manuel Garcia-Verdugo.Neurogenesis in Adult Subventricular Zone[J].The Journal of Neuroscience,2002,22(3),629-634.
[11]R.Christie,Nicolas Toni,Theo D.et al.Gage.Functional neurogenesis in the adult hippocampus[J].Nature,2002,415(6875):1030-1034.
[12]Zaal Kokaia,Olle Lindvall.Neurogenesis after ischaemic brain insults[J].Current Opinion in Neurobiology,2003,13(1),127-132.
[13]左明雪,向斌.成年白腰文鸟端脑新生神经元的光镜和电镜研究[J].动物学报,1999,45(4)：378-384.
[14]史兆春,吴婷,乔莉,董海蓉,李敏.有氧训练对阿尔茨海默病模型大鼠海马齿状回神经发生的影响[J].中国临床神经科学.2011,19(3)：255-259.
[15]Xinjie Bao,Ming Feng,Junji Wei,et al.Transplantation of Flk-1+human bone marrow-derived mesenchymal stem cells promotes angiogenesis and neurogenesis after cerebral ischemia in rats[J].European Journal of Neuroscience,2011,34(l):87-98.
[16]Sun,Xuelian Li,Qingjie Zhao.Limited hippocampal neurogenesis in SAMP8 mouse model of Alzheimer's disease[J].Brain Research,2011,1389:183-193.
[17]M.Mar Perez-Canellas,JoseM.Garcia-Verdugo.Adult neurogenesis in the telencephalon of a lizard:a [3H]thymidine autoradiographic and bromodeoxyuridine immunocytochemical study[J].Developmental Brain Research,1999,93:49-61.
[18]Font E.,Desfilis E.,Perez-Canellas M.M.,et al.Neurogenesis and Neuronal Regeneration in the Adult Reptilian Brain[J].Brain,Behavior and Evolution,2001,58:276-295.
[19]Garcia.Postnatal neurogenesis in the medial cortex of the tropical lizard Tropidurus hispidus[J].Neuroscience,2005,134:407-413.
[20]Yuanting Jin,Naifa Liu,Jinlu Li.Elevational variation in body size of Phrynocephalus Vlangalii in North Qinghai-Xizang(Tibetan)Plateau[J].Belgian Journal of Zoology,2007,137(2):197-202.
[21]Gratzner HGMonoclonal antibody to 5-bromo-and 5-iododeoxyuridine:A new reagent for detection of DNA replication[J].Science,1982,218(4571):474-475.
[22]Pasko Rakic.Neurogenesis in adult primate neocortex:an evaluation of the evidence[J].Nature Reviews Neuroscience,2002,3(1):65-71.
[23]HG Kuhn,H Dickinson-Anson,FH Gage.Neurogenesis in the dentate gyrus of the adult rat:age-related decrease of neuronal progenitor proliferation[J].The Journal of Neuroscience,1996,16(6):2027-2033.
[24]姚檀栋,刘晓东,工宁练.青藏高原地区的气候变化幅度问题[J].科学通报.1996,45(1)：98-106.
[25]Kuan Zhang,Lingling Zhu,Ming Fan.Oxygen,a Key Factor Regulating Cell Behavior during Neurogenesis and Cerebral Diseases[J].Frontiers in molecular neuroscience,2011,4:5.
[26]CARLOS LOPEZ-GARCIA,ASUNCION MOLOWNY,JUAN NACHER,et al.The lizard cerebral cortex as a model to study neuronal regeneration[J].Anais da Academia Brasileira de Cicncias,2002,74(1):85-104.
[27]H.Shao,L.Fan,X.J.Xu,et al.Characterization of adult neurogenesis in lizard Phrynocephalus vlangalii (Agamidae:Reptilia)[J].Italian Journal of Zoology.2012,79(4):547-558.
[28]Lorenz Studer,Marie Csete,Sang-Hun Lee,et al.Enhanced proliferation,survival,and dopaminergic differentiation of CNS precursors in lowered oxygen[J].The Journal of Neuroscience,2000,20(19):7377-7383.
[29]Sean J.Morrison,Marie Csete,Andrew K.Groves,et al.Culture in reduced levels of oxygen promotes clonogenic sympathoadrenal differentiation by isolated neural crest stem cells[J].The Journal of Neuroscience,2000,20(19):7370-7376.
[30]shi Ding,Wen-hong Fan,Ming Fan.Neurogenesis in the adult rat brain after intermittent hypoxia[J].Brain Research,2005,1055(1-2):1-6.
[31]Maria V. Gustafsson, Xiaowei Zheng, Teresa Pereira,et al.Hypoxia requires notch signaling to maintain the undifferentiated cell state[J].Developmental Cell,2005,9(5):617-628.
[32]Xiaofeng Zheng,Sarah Linke,Jose M.Dias,et al.Interaction with factor inhibiting HIF-1 defines an additional mode of cross-coupling between the Notch and hypoxia signaling pathways[J].Proceedings of the National Academy of Sciences of the United States of America,2008,105(9):3368-3373.
[33]Till Acker,Helmut Acker.Cellular oxygen sensing need in CNS function:physiological and pathological implications[J].The Journal of Experimental Biology,2004,207:3171-3188.
[34]F.J.Delgado-Gonzaleza,A.Alonso-Fuentesa,A.Delgado-Fumeroa,et al.Seasonal differences in ventricular proliferation of adult Gallotia galloti lizards[J].Brain Research,2008,1191:39-46.
[35]鲍敏,曾阳,马建滨,等.青海沙蜥和密点麻蜥的分布状况、生活规律及食性[J].青海帅范大学学报(自然科学版),1998,4：42-45.
[2]Egger B, SGold K, Brand A H. Notch regulates the switch from symmetric to asymmetric neural stem cell division in the Drosophila optic lobe [J]. Development,2010, 137(18):2981-2987.
[3]Ferron S R, Pozo N, Laguna A, et al. Regulated Segregation of Kinase Dyrk1 A during Asymmetric Neural Stem Cell Division Is Critical for EGFR-Mediated Biased Signaling[J]. Cell Stem Cell,2010,7(3):367-379.
[4]Nishimura T. Molecular mechanisms for asymmetric cell division in Drosophila neural stem cells[J]. Tanpakushitsu Kakusan Koso,2009,54(11):1363-1369.
[5]Walczak P, Kedziorek D, Gilad A, et al. Applicability and limitations of MR tracking of neural stem cells with asymmetric cell division and rapid turnover:the case of the shiverer dysmyelinated mouse brain[J], Magnetic Resonance in Medicine,2007,58(2):261-269.
[6]Pagano S F, Impagnatiello F, Girelli M, et al. Isolation and Characterization of Neural Stem Cells from the Adult Human Olfactory Bulb[J]. Stem Cells,2002,18(4):295-300.
[7]Duan X, Kang E, Liu C Y, et al. Development of neural stem cell in the adult brain[J]. Current Opinion in Neurobiology,2008,18(1):108-115.
[8]Ming G, Song H. Adult neurogenesis in the mammalian central nervous system[J]. Annual Review of Neuroscience,2005(28):223-250.
[9]Zhao C, Deng W, Gage F H. Mechanisms and functional implications of adult neurogenesis[J]. Cell,2008,132(4):645-660.
[10]Cameron H A, Mckay R D. Adult neurogenesis produces a large poo! of new granule cells in the dentate gyrus[J]. The Journal of Comparative Neurology,2001,435(4):406-417.
[11]Gage F H. Mammalian neural stem ceIls[J]. Science,2000,287(5457):1433-1438.
[12]Lledo P, Alonso M, Grubb M S. Adult neurogenesis and functional plasticity in neuronal circuits[J]. Nature Reviews Neuroscience,2006,7(3):179-193.
[13]Carleton A, Petreanu L T, Lansford R, et al. Becoming a new neuron in the adult olfactory bulb[J]. Nature Neuroscience,2003,6(5):507-518.
[14]AlvarezBuylla A, Jose Manuel Garcia-Verdugo. Neurogenesis in adult subventricular zone[J]. The Journal of Neuroscience,2002,22(3):629-634.
[15]Goldman S A, Nottebohm F. Neuronal production, migration, and differentiation in a vocal control nucleus of the adult female canary brain[J]. Proceeding of the National Academy of Sciences of the United States of America,1983,80(8):2390-2394.
[16]AlvarezBuylla A, Ling C, Yu W S. Contribution of neurons born during embryonic and adult life to the brain of adult canaries:Regional specificity and delayed birth of neurons in the song-control nuclei[J]. Journal of Comparative neurology,1994,347(2):233-248.
[17]Bernocchi, G, Scherini, E., Nano, R. Developmental patterns in the rat cerebellum after cis-dichlorodiammineplatinum treatment[J]. Neuroscience.1990,39(1):179-188.
[18]Polenov A, Chetverukhin V. Ultrastructural radioautographic analysis of neurogenesis in the hypothalamus of the adult frog, Rana temporaria, with special reference to physiological regeneration of the preoptic nucleus.I.Ventricular zone cell proliferation[J]. Cell and tissue research,1993,291 (2):341-350.
[19]EM D, Fingerlin A, Hwang D, et al. Seasonal Cell Proliferation in the Chemosensory Epithelium and Brain of Red-Backed Salamanders, Plethodon cinereus[J]. Brain,Behavior and Evolution,2000,56(1):1-13.
[20]Raucci F, Fiore M M D, Pinelli C, et al. Proliferative activity in the frog brain:a PCNA-immunohistochemistry analysis[J]. Journal of Chemical Neuroanatomy,2006, 32(2-4):127-142.
[21]Scoville W, Milner B. Loss of recent memory after bilateral hippocampal lesions[J]. Neurol Neurosurg Psychiatry,1957,20(1):11-21.
[22]Corkin S, Sullivan E, Carr F. Prognostic factors for life expectancy after penetrating head injury[J]. Archives of neurology,1984,41(9):975-977.
[23]Schmolck H, Kensinger E A, Corkin S, et al. Semantic knowledge in patient HM and other patients with bilateral medial and lateral temporal lobe lesions[J]. Hippocampus,2002, 12(4):520-533.
[24]Corkin S. What's new with the amnesic patient HM?[J]. Nature Reviews Neuroscience, 2002,3(2):153-160.
[25]Rakic P. Neurogenesis in adult primate neocortex:an evaluation of the evidence[J]. Nature Reviews Neuroscience,2002,3(1):65-71.
[26]Hayes N, Nowakowski R. Dynamics of cell proliferation in the adult dentate gyrus of two inbred strains of mice[J]. Developmental Brain Research,2002,134(1-2):77-85.
[27]Selden D J R, Dolbeare F, Clair J H, et al. Statistical confirmation that immunofluorescent detection of DNA repair in human fibroblasts by measurement of bromodeoxyuridine incorporation is stoichiometric and sensitive[J]. Cytometry,1993,14(2):154-167.
[28]Eng L F. Glial fibrillary acidic protein (GFAP):the major protein of glial intermediate filaments in differentiated astrocytes[J]. Journal of Neuroimmunology,1985(8):203-214.
[29]Mullen R, Buck C, Smith, et al. NeuN, a neuronal specific nuclear protein in vertebrates [J]. Development,1992(161):201-211.
[30]Dyer M A, Livesey F J, Cepko C L, et al. Proxl function controls progenitor cell proliferation and horizontal cell genesis in the mammalian retina[J]. Nature Genetics, 2003(34):53-58.
[31]Gdalyahu A, Levy I G. DCX, a new mediator of the JNK pathway[J]. The EMBO Journal, 2004(23):823-832.
[32]Gerdes J, Lemke H, Baisch H, et al. Cell cycle analysis of a cell proliferation-associated human nuclear antigen defined by the monoclonal antibody Ki-67[J]. The Journal of Immunology,1984,133(4):1710-1715.
[33]Hall D P A, Levison D A, Woods A L, et al. Proliferating cell nuclear antigen (PCNA) immunolocalization in paraffin sections:An index of cell proliferation with evidence of deregulated expression in some, neoplasms[J]. The Journal of Pathology,1990,162(4):285-294.
[34]Lewis P F, Emerman M. Passage through mitosis is required for oncoretroviruses but not for the human immunodeficiency virus.[J]. Journal of Virology,1994,68(1):510-516.
[35]Prendergast F G, Mann K. G. Chemical and physical properties of aequorin and the green fluorescent protein isolated from Aequorea forskalea[J]. Biochemistry,1978,17(17):3448-3453.
[36]Tsien R Y. The green fluorescent protein[J]. Annu. Rev. Biochem,1998(67):509-544.
[37]姚檀栋,刘晓东,王宁练.青藏高原地区的气候变化幅度问题[J].科学通报,2000,45(1)：98-106.
[38]林振耀,赵昕奕.青藏高原气温降水变化的空间特征[J].中国科学,1996,26(4)：354-358.
[39]张静.高原病研究的新进展(综述)[J].中国城乡企业卫生,2009,12(06)：17-19.
[40]1. Bojana B. Beleslin-Cokic, Cokic V P, Yu X, etal. Erythropoietin and hypoxia stimulate erythropoietin receptor and nitric oxide production by endothelial cells[J]. Blood,2004, 104(7):2073-2080.
[41]Li, Y.X., Ding, S.J.,et al. Desferoxamine preconditioning protects against cerebral ischemia in rats by inducing expressions of hypoxia inducible factor 1α and erythropoietin[J]. Neurosci Bull,2008,24(2):89-95.
[42]杨燕,马慧萍,张汝学,等.急性高原病发病机制的研究进展[J].医学综述,2010,16(17)：2561-2563.
[43]于永才.高原病的发病及防治[J].社区医学杂志,2010,8(24)：10-13.
[44]周其全,刘福玉,郑必海,等.急性重症高原病患者脑功能损伤与多器官功能障碍[J].世界急危重病医学杂志,2007,4(3)：1836-1840.
[45]吕国蔚.医学神经生物学[M].第2版.北京：高等教育出版社,2000：13-399.
[46]Michael Ward(国家体委体育科学研究所运动医学研究所等译)Mountain Medicine[M]第1版.北京：人民卫生出版社,1978：203-312.
[47]刘重庆.高原医学[M].1983,(2)：19.
[48]Reite M, Jackson D, Cahoon R L, et al. Sleep physiology at high altitude[J]. Electroencephalography and Clinical Neurophysiology,1975,38(5):463-471.
[49]MA L P, Oxfd B, BA M W, etal. Some effects of high altitude on man[J]. The Lancet,1956, 268(6953):1115-1121.
[50]蒲小燕,沈国平.高原低氧SD大鼠海马神经细胞的结构改变[J].医学信息,2010,23(1)：234.
[51]尹宏,张先钧,张晓岩.急性高原低氧对大鼠海马神经细胞形态结构的影响[J].青海医学院学报,2007,28(1)：29-31.
[52]张先钧,张晓岩,蒲小燕,等.急性和亚急性高原低氧大鼠海马神经细胞的电镜研究[J].青海医学院学报,2005,26(4)：236-239.
[53]赵霞,张先钧,张晓岩,等.业急性高原低氧大鼠海马神经细胞的电镜研究[J].高原医学杂志,2006,16(2)：59-60.
[54]吴穹,郭新庆.高原低氧应激诱导大鼠海马神经元凋亡[J].菏泽医学专科学校学报,2009,21(3)：1-2.
[55]李文林,沈娟志.高原环境对大鼠海马神经元凋亡的影响[J].包头医学院学报,2010,26(4)：9-10.
[56]高原,李源,蔡晓红,等.慢性问歇低氧对幼鼠海马神经细胞凋亡及慢性问歇低氧对幼鼠海马神经细胞凋亡及慢性问歇低氧对幼鼠海马神经细胞凋亡及caspase-12表达的影响[J].温州医学院学报,2009,39(5)：422-425.
[57]Lu X, Chen X, Weng J, et al. Hippocampal spine-associated Rap-specific GTPase-activating protein induces enhancement of learning and memory in postnatally hypoxia-exposed mice[J]. Neuroscience,2009,162(2):404-414.
[58]Zhang J, Chen X, Du J, et al. Neonatal exposure to intermittent hypoxia enhances mice performance in water maze and 8-arm radial maze tasks[J]. Journal of Neurobiology,2005, 65(1):72-84.
[59]余小燕,马福海.低氧诱导因子-1在高原低氧训练中的研究现状与进展[J].辽宁体育科技,2010,32(2)：19-21.
[60]Poon E, Harris A L, Ashcroft M. Targeting the hypoxia-inducible factor (HIF) pathway in cancer[J]. Expert Reviews in Molecular Medicine,2009(11):1-23.
[61]王雪莲,李占全,冀林华.HIF-1 α,VEGF在慢性高原低氧环境下SD大鼠骨髓,心脏中的表达及意义[J].山东医药,2009,49(9)：34-35.
[62]刘丽愉,刘力芳,曹亚.低氧诱导因子信号通路的新进展[J].国际病理科学与临床杂志,2011,31(3)：218-222.
[63]Lathia J D, Mattson M P, Cheng A. Notch:from neural development to neurological disorders[J]. Journal of Neurochemistry,2008,107(6):1471-1481.
[64]Nefedova Y, Cheng P, Alsina M, et al. Involvement of Notch-1 signaling in bone marrow stroma-mediated de novo drug resistance of myeloma and other malignant lymphoid cell lines[J]. Blood,2009,29(10):1644-1654.
[65]Arumugam T V, Chan S L, Jo D, et al. Gamma secretase-mediated Notch signaling worsens brain damage and functional outcome in ischemic stroke[J]. Nature Medicine,2006, 12(6):621-623.
[66]Gustafsson M V, Zheng X, Pereira T, et al. Hypoxia requires notch signaling to maintain the undifferentiated cell state[J]. Developmental Cell,2005,9(5):617-628.
[67]沈晓玲,映红.高原低氧对白主神经系统的影响[J].高原医学杂志,2008,18(3)：54-57.
[68]马晓庆,李琳娜.高原缺氧对神经系统的影响与护理[J].中华护理杂志,1991,36(10)：439-441.
[69]田开新,覃军,黄岚,等.急进高原自主神经功能变化与急性高原反应的关系[J].解放军预防医学杂志,2008,26(1)：9-13.
[70]隆敏,覃军.急进高原自主神经系统功能变化研究进展[J].解放军预防医学杂志,2008,26(2)：147-149.
[71]张勤文.不同海拔地区牦牛的肺组织形态学比较[J].中国兽医科技,2004,34(2)：73-74.
[72]陈秋生,冯霞,姜生成.牦牛肺脏高原适应性的结构研究[J].中国农业科学,2006,39(10)：2107-2113.
[73]江家椿,嘎玛仁增,何玛丽.不同海拔高度西藏高原牦牛若干血液生理常值的比较[J].畜牧兽医学报,1991,22(1)：20-26.
[74]司本辉,张庆成,李积荣,等.久居高原地区缺氧环境对红细胞膜蛋白结构和血液流变学的影响[J].高原医学杂志,2001,11(3)：26-27.
[75]Jin Y, Liu N, Li J. Elevational variation in body size of Phrynocephalus vlangalii in the North Qinghai-Xizang (Tibetan) Plateau[J]. Belg. J. Zool.,2007,137(2):197-202.
[76]金园庭,田仁荣,刘迺发.四种沙蜥的形态地理变异Bergmann和Allen规律的检验[J].动物学报,2006,52(5)：838-845.
[77]赵肯堂.中国的沙蜥属研究[J].动物学杂志,1997,32(1)：15-19.
[78]李俊,郭宪光,王跃招.沙蜥属Phrynocephalus生物地理学研究进展[J].四川动物,2010,29(1)：144-150.
[79]王跃招,曾晓茂,方白立,等.沙蜥属一有效种贵德沙蜥及红原沙蜥的分类研究(蜥蜴亚目：鬣蜥科)[J].动物分类学报,2002,27(2)：372-383.
[80]赵肯堂.巾国胎生爬行纲动物[J].生物学通报,1988(1)：1-3.
[81]吴鹏飞,王跃招,王硕果,等.青海沙蜥(蜥蜴亚目：鬣蜥科)种群的年龄结构与性比[J].四川大学学报(自然科学版),2002,39(6)：1134-1139.
[82]江耀明,黄庆云,赵尔宓.青海沙蜥一新亚种及其生态初步观察[J].动物学报,1980,26(2)：178-183.
[83]吴鹏飞,王跃招,郭海燕,等.青海沙蜥的生长及两性生长差异[J].四川大学学报(自然科学版),2005,42(6)：1252-1257.
[84]赵肯堂.沙蜥的泄殖系统及其繁殖习性[J].动物学杂志,1964(4)：158-186.
[85]鲍敏,曾阳,马建滨,等.青海沙蜥和密点麻蜥的分布状况和食性[J].青海师范大学学报(自然科学版),1998,4(4)：42-45.
[86]吾玛尔阿布力孜.东疆沙蜥和密点麻蜥的食性研究[J].干旱区研究,1992,16(4)：10-13.
[87]邵虹.青海沙蜥成体神经发生的特征描述[D].兰州大学,2011.
[88]H.Shao, L.Fan, X.J.Xu, et al. Characterization of adult neurogenesis in lizard Phrynocephalus vlangalii (Agamidae:Reptilia)[J]. Italian Journal of Zoology. 2012,79(4):547-558.
[89]伍冠一,廖锦锋,唐宗湘.爬行动物皮层的细胞构筑与纤维联系[J].四川动物,2006,25(2)：416-419.
[90]农万廷,石哪.石龙子脑核团定位研究[J].南宁师范高等专科学校学报,2002,19(2)：45-47.
[91]Marchioro M, Nunes J D A M, Ramalho A R, etal. Postnatal neurogenesis in the medial cortex of the tropical lizard Tropidurus hispidus[J]. Neuroscience,2005,134(2):407-413.
[92]DelgadoGonzalez F, GonzalezGranero S, TrujilloTrujillo C, etal. Study of adult neurogenesis in the gallotia gal lot i lizard during different seasons[J]. Brain Research, 2011(1390):50-58.
[93]颜坤,王荫亭,王书荣.蜥蜴中脑的Golgi-Cox研究[J].科学通报,1982(6)：371-373.
[94]王书荣,颜坤,王荫亭,等.蜥蜴中脑神经通路和起源细胞的形态[J].解剖学报,1989,20(2)：159-164.
[95]伍冠一.蛤蚧皮质加厚区的细胞构筑与纤维联系研究[D].广西师范大学,2006.
[96]伍冠一,廖锦锋,唐宗湘.爬行动物脑皮质加厚区的结构与功能[J].动物医学进展,2006,27(3)：105-107.
[97]F.J. Delgado-Gonzalez, AlonsoFuentes A, DelgadoFumero A, etal. Seasonal differences in ventricular proliferation of adult Gallotia galloti lizards[J]. Brain Research,2008(1191):39-46.
[98]王典群.四种荒漠蜥蜴脑和脑神经的比较解剖[J].兰州大学学报(自然科学版).1992,38(8)：145-149.
[99]张法奎.似哺乳类爬行动物和哺乳类动物脑颅侧壁构造类型的演替[J].古脊椎动物学报,1989,27(3)：209-218.
[100]LOPEZGARCIA C, MOLOWNY A, NACHER J, etal. The lizard cerebral cortex as a model to study neuronal regeneration[J]. Anais da Academia Brasileira de Ciencias,2002, 74(1):85-104.
[1]Reynolds BA, Weiss S.Generation of neurons and astrocytes from isolated cells of the adult mammalian central nervous system[J].Science,1992,255(5052):1707-1710.
[2]Guo-li Ming,Hongjun Song.Adult neurogenesis in the mammalian central nervous system[J].Annual Review of Neuroscience,2005,28:223-250.
[3]Elizabeth Gould.How widespread is adult neurogenesis in mammalsfJ].Nature Reviews Neuroscience,2007,8(6):481-488.
[4]Xin Duan,Eunchai Kang,Cindy Y Liu,et al.Development of neural stem cell in the adult brain[J].Current Opinion in Neurobiology,2008,18(1):108-115.
[5]Fred H.Gage.Mammalian neural stem cells[J].Science,2000,287(5457):1433-1438.
[6]Chunmei Zhao, Wei Deng,Fred H.Gage.Mechanisms and functional implications of adult neurogenesis[J].Cell.2008,132(4):645-660.
[7]Heather A.Cameron,Ronald D.GMckay.Adult neurogenesis produces a large pool of new granule cells in the dentate gyrus[J].Journal of Comparitive Neurology,2001,435(4):406-417.
[8]Pierre-Marie Lledo,Mariana Alonso & Matthew S,Grubb.Adult neurogenesis and functional plasticity in neuronal circuits[J].Nature Reviews Neuroscience,2006,7(3):179-193.
[9]Alan Carleton,Leopoldo T.Petreanu,et al.Becoming a new neuron in the adult olfactory bulb[J].Natural Neuroscience,2003,6(5):507-518.
[10]Arturo Alvarez-Buylla,Jose Manuel Garcia-Verdugo.Neurogenesis in Adult Subventricular Zone[J].The Journal of Neuroscience,2002,22(3),629-634.
[11]R.Christie,Nicolas Toni,Theo D.et al.Gage.Functional neurogenesis in the adult hippocampus[J].Nature,2002,415(6875):1030-1034.
[12]Zaal Kokaia,Olle Lindvall.Neurogenesis after ischaemic brain insults[J].Current Opinion in Neurobiology,2003,13(1),127-132.
[13]左明雪,向斌.成年白腰文鸟端脑新生神经元的光镜和电镜研究[J].动物学报,1999,45(4)：378-384.
[14]史兆春,吴婷,乔莉,董海蓉,李敏.有氧训练对阿尔茨海默病模型大鼠海马齿状回神经发生的影响[J].中国临床神经科学.2011,19(3)：255-259.
[15]Xinjie Bao,Ming Feng,Junji Wei,et al.Transplantation of Flk-1+human bone marrow-derived mesenchymal stem cells promotes angiogenesis and neurogenesis after cerebral ischemia in rats[J].European Journal of Neuroscience,2011,34(l):87-98.
[16]Sun,Xuelian Li,Qingjie Zhao.Limited hippocampal neurogenesis in SAMP8 mouse model of Alzheimer's disease[J].Brain Research,2011,1389:183-193.
[17]M.Mar Perez-Canellas,JoseM.Garcia-Verdugo.Adult neurogenesis in the telencephalon of a lizard:a [3H]thymidine autoradiographic and bromodeoxyuridine immunocytochemical study[J].Developmental Brain Research,1999,93:49-61.
[18]Font E.,Desfilis E.,Perez-Canellas M.M.,et al.Neurogenesis and Neuronal Regeneration in the Adult Reptilian Brain[J].Brain,Behavior and Evolution,2001,58:276-295.
[19]Garcia.Postnatal neurogenesis in the medial cortex of the tropical lizard Tropidurus hispidus[J].Neuroscience,2005,134:407-413.
[20]Yuanting Jin,Naifa Liu,Jinlu Li.Elevational variation in body size of Phrynocephalus Vlangalii in North Qinghai-Xizang(Tibetan)Plateau[J].Belgian Journal of Zoology,2007,137(2):197-202.
[21]Gratzner HGMonoclonal antibody to 5-bromo-and 5-iododeoxyuridine:A new reagent for detection of DNA replication[J].Science,1982,218(4571):474-475.
[22]Pasko Rakic.Neurogenesis in adult primate neocortex:an evaluation of the evidence[J].Nature Reviews Neuroscience,2002,3(1):65-71.
[23]HG Kuhn,H Dickinson-Anson,FH Gage.Neurogenesis in the dentate gyrus of the adult rat:age-related decrease of neuronal progenitor proliferation[J].The Journal of Neuroscience,1996,16(6):2027-2033.
[24]姚檀栋,刘晓东,工宁练.青藏高原地区的气候变化幅度问题[J].科学通报.1996,45(1)：98-106.
[25]Kuan Zhang,Lingling Zhu,Ming Fan.Oxygen,a Key Factor Regulating Cell Behavior during Neurogenesis and Cerebral Diseases[J].Frontiers in molecular neuroscience,2011,4:5.
[26]CARLOS LOPEZ-GARCIA,ASUNCION MOLOWNY,JUAN NACHER,et al.The lizard cerebral cortex as a model to study neuronal regeneration[J].Anais da Academia Brasileira de Cicncias,2002,74(1):85-104.
[27]H.Shao,L.Fan,X.J.Xu,et al.Characterization of adult neurogenesis in lizard Phrynocephalus vlangalii (Agamidae:Reptilia)[J].Italian Journal of Zoology.2012,79(4):547-558.
[28]Lorenz Studer,Marie Csete,Sang-Hun Lee,et al.Enhanced proliferation,survival,and dopaminergic differentiation of CNS precursors in lowered oxygen[J].The Journal of Neuroscience,2000,20(19):7377-7383.
[29]Sean J.Morrison,Marie Csete,Andrew K.Groves,et al.Culture in reduced levels of oxygen promotes clonogenic sympathoadrenal differentiation by isolated neural crest stem cells[J].The Journal of Neuroscience,2000,20(19):7370-7376.
[30]shi Ding,Wen-hong Fan,Ming Fan.Neurogenesis in the adult rat brain after intermittent hypoxia[J].Brain Research,2005,1055(1-2):1-6.
[31]Maria V. Gustafsson, Xiaowei Zheng, Teresa Pereira,et al.Hypoxia requires notch signaling to maintain the undifferentiated cell state[J].Developmental Cell,2005,9(5):617-628.
[32]Xiaofeng Zheng,Sarah Linke,Jose M.Dias,et al.Interaction with factor inhibiting HIF-1 defines an additional mode of cross-coupling between the Notch and hypoxia signaling pathways[J].Proceedings of the National Academy of Sciences of the United States of America,2008,105(9):3368-3373.
[33]Till Acker,Helmut Acker.Cellular oxygen sensing need in CNS function:physiological and pathological implications[J].The Journal of Experimental Biology,2004,207:3171-3188.
[34]F.J.Delgado-Gonzaleza,A.Alonso-Fuentesa,A.Delgado-Fumeroa,et al.Seasonal differences in ventricular proliferation of adult Gallotia galloti lizards[J].Brain Research,2008,1191:39-46.
[35]鲍敏,曾阳,马建滨,等.青海沙蜥和密点麻蜥的分布状况、生活规律及食性[J].青海帅范大学学报(自然科学版),1998,4：42-45.