雌激素受体β（ERβ）对雌性棕色田鼠社会行为及其脑区OTR和AVPV1aR表达的影响
|
摘要
动物的社会行为与神经系统、内分泌系统等密切相关。神经肽、性激素及很多内分泌因子,与动物的生存环境共同影响着动物的社会行为。大量实验证据表明,雌激素(E)、中枢加压素(AVP)和催产素(OT)是调控社会识别的关键物质。它们可以影响动物的社会识别、繁殖行为、性行为、攻击行为、配偶选择行为、亲密行为和个体社会互作行为等等,并且,ERβ可以调控OTmRNA和AVPmRNA的转录。ERβ在社会行为中是否可以调节OTRmRNA和AVPV1aRmRNA还未见报道。雌激素可以调控下丘脑中的OT和AVP基因的表达,这种调控作用在雌激素受体β基因敲除的小鼠中完全不存在,即在此调控中雌激素受体β(ERβ)是否被激活是个关键因素。而AVP在神经系统中发挥作用主要是依赖于V1a受体的存在,OT的作用是通过与其受体结合,经下丘脑—垂体—肾上腺(HPA)轴或自主神经等系统来改变动物的生理和行为。但是关于ERβ在OTR和AVPV1aR调控棕色田鼠(Microtus mandarinus)社会行为中的作用还不清楚。因此,本实验将重点研究ERβ如何影响田鼠的部分社会行为和OTR和AVPVlaR的表达,以探讨ERβ与OTR和AVPV1aR共同作用下对田鼠社会行为的调控。
本实验利用反义核酸技术和行为学检测平台相结合进行研究,通过给雌性棕色田鼠侧脑室中枢连续注射微量的雌激素受体βmRNA反义核酸,然后对实验鼠进行相关的行为检测;同时利用反义核酸技术和原位杂交组织化学技术相结合,探讨雌激素及其受体和催产素、加压素及其受体在雌性棕色田鼠脑内如何共同调节复杂的社会行为。
1.选择成体雌性棕色田鼠侧脑室中枢连续注射微量的ERβ受体mRNA反义核酸,分别在第一日、第三日及第五日同一时间麻醉注射。六小时后,利用攻击行为学检测平台进行实验,历时10分钟。实验结果显示:注射反义核酸组攻击行为的频次和持续时间明显高于注射随机序列组,而潜伏期没有显著性的差异。推断ERβmRNA的反义核酸在mRNA水平上阻止其翻译形成ERβ,进而阻碍了E及受体调控雌性棕色田鼠的攻击行为。
2.选择成体雌性棕色田鼠侧脑室中枢连续注射微量的ERβ受体mRNA反义核酸,分别在第一日、第三日及第五日同一时间麻醉注射。六小时后,利用配偶选择行为学检测平台进行实验,历时60分钟。实验结果显示:雌鼠在对陌生异性鼠箱、配偶鼠箱和中立箱中访问时间上有显著差异。说明给成年雌性棕色田鼠注射ERβmRNA的反义核酸后,使BST、CE、LSD、PVN和ME核团AVPV1aRmRNA和OTRmRNA表达增强,导致与其形成的神经投射功能发挥,从而表现出与之相应的行为活性增强,以致于增强了雌性棕色田鼠和配偶鼠之间的配偶联系。
3.选择成体雌性棕色田鼠侧脑室中枢连续注射微量的ERβ受体mRNA反义核酸,分别在第一日、第三日及第五日同一时间麻醉注射。六小时后,利用亲密行为学检测平台进行实验,历时60分钟。实验结果显示:雌鼠对配偶鼠的善待行为的增加,防御行为减弱。对陌生异性鼠的探究行为减弱,防御行为的增强。这些变化说明ERβ在田鼠偏好测试中,对配偶鼠和陌生异性鼠的社会识别可能存在着根本不同的神经通路,推测ERβ反义核酸可加强配偶关系。
4.选择成体雌性棕色田鼠侧脑室中枢连续注射微量的ERβmRNA反义核酸,分别在第一日、第三日及第五日同一时间麻醉注射。六小时后,利用原位杂交组织化学技术进行OTmRNA检测。实验结果显示:与对照组相比在棕色田鼠脑区核团中终纹床核(BST)和杏仁中央核(CE)的OTR的表达有显著性增强,在杏仁内侧核(ME)、下丘脑视前区(AH)和下丘脑室旁核(PVN)的OTR的表达有显著性减弱。说明在这些核团ERβ的反义核酸与OTR相互作用影响了雌性田鼠的社会行为,提示我们在理解E在介导雌性棕色田鼠的社会识别时,可能是通过催产素及其受体来发挥作用的。
5.选择成体雌性棕色田鼠侧脑室中枢连续注射微量的ERβ受体mRNA反义核酸,分别在第一日、第三日及第五日同一时间麻醉注射。六小时后,利用原位杂交组织化学技术进行AVPV1RmRNA检测。实验结果显示:与对照组相比在棕色田鼠脑区核团中终纹床核(BST)、下丘脑室旁核(PVN)和隔外侧核背部(LSD)核团中AVPV1aR的表达有显著性增加,在杏仁中央核(CE)的AVPV1aR表达显著性减弱,由此推测在这些核团ERβ的反义核酸与AVPR相互作用影响了雌性田鼠的社会行为。
由以上结论可得出:ERβ、OTR和AVPV1aR与雌性棕色田鼠的社会行为密切相关。ERβ可在一定程度上降低其攻击行为。ERβ对雌性棕色田鼠的配偶关系的调节及社会识别活动的调控是与OTR和AVPV1aR相关联的,ERβ反义核酸可提高雌性棕色田鼠的配偶识别能力,加强配偶之间的联系,即任何一种行为都受到复杂的神经内分泌机制的调控。
Animal social behavior,was controlled and regulated by centre nervous system and neuroendocrine system.Neuropeptide,sex hormone,a number of endocrine factors and environments where animal lives cooperate to affect social behavior of animals.Many experimental evidences indicate that E,OT and AVP acting in the brain appear to be critical for social discrimination.They could affect many social behaviors,such as social recognition,aggression, sexual behavior,partner preference,affinity behavior and individual behavior.Whether ERβcan regulate the OTRmRNA and AVPV1aRmRNA has not been reported.We preit that estrogen may show much regulation on the levels of OT and AVP messenger RNA.So,the present study will aim at the two problems:1,how E could affect some social behaviors in voles;2,Is there effects to OTR and AVPV1a expression by E receptor?
we used monogamous female mandarin voles(Microtus mandarinus) as research objects. Combining antisense nucleotides technic with behavioral observation,we examined the correlative behavior after i.c.v,continuous injection of antisense nucleotides of estrogen receptor-β.Meanwhile, Combined antisense nucleotides technic with in situ hybridization,we investigated how ERβ, vasopressin,oxytocin and its receptors controlled complex social behaviors in female voles.
1.Selected female adult mandarin voles were given i.c.v,continuous injections of antisense nucleotides of estrogen receptor-βrespectively in first day,third day and fifth day.After six hours, we examined aggression behavior during 10 minutes.The results were following:Compared with group of injection of scramble nucleotides,anisense group showed significant increase in frequency and total duration in aggression behavior.We suggested that antisense nucleotides of ERβinhibit the translation of ERβin mRNA levels.Furthermore,it blocked E to modulate aggression behavior in male voles.
2.Selected female adult mandarin voles were given i.c.v,continuous injections of antisense nucleotides of estrogen receptor-βin first day,third day and fifth day.After six hours,we examined partner preference behavior during 60 minutes.The results were following:There were significant differences in visiting duration in mate's compartment,strange heterosexual vole's compartment and neutral compartment,and significant increase in the number of AVPV1RmRNA and OTRmRNA in BST、CE、LSD、PVN and ME,It is suggested that continuous injection of antisense nucleotides of ERβcan enhance social recognition,and can enhance the pair bond formation of male and female voles.
3.Selected female adult mandarin voles were given i.c.v,continuous injections of antisense nucleotides of estrogen receptor-βrespectively in first day,third day and fifth day.After six hours, we examined affinity behavior during 60 minutes.The results were following:There were significant enhance in amicable behavior and decrease in defensive behavior with its mate.At the same time,there were significant enhance in defensive behavior and decrease in investigating behavior with the strange heterosexual voles.It suggested that there were different nervous circuit, which female voles recognized socially its mate and stranger voles in the preference behavior examination.
4.Selected female adult mandarin voles were given i.c.v,continuous injections of antisense nucleotides of estrogen receptor-βin first day,third day and fifth day.After six hours,OTR mRNA was examnined using in situ hybridization method in female voles.The results were following: There were significant differences in the BST,CE,ME,PVN and AH.Compared with injection of scramble nucleotides and injection of isotonic saline treatment,there were significant reduction in the number of OTR mRNA in AH,PVN and ME,and significant increase in the number of OTR mRNA in BST and CE Compared with injection of saline.These data suggested that the roles between antisense nucleotides of ERβand OTR affected social behavior in the female voles.It explained that ERβmodulated social behavior in female voles possibly through oxytocin and its receptor.
5.Selected female adult mandarin voles were given i.c.v,continuous injections of antisense nucleotides of estrogen receptor-βin first day,third day and fifth day.After six hours, AVPV1RmRNA was examnined using in situ hybridization method in female voles.The results were following:There were significant differences in the BST,CE,PVN and LSD.Compared with injection of scramble nucleotides and injection of isotonic saline treatment,there were significant increase in the number of AVPV1RmRNA in BST、PVN and LSD,and significant reduction in the number of AVPV1R mRNA in CE.These data suggested that the roles between antisense nucleotides of ERβand AVPV1R affected social behavior in the female voles.
Results from those conclusion that ERβ、OTR and AVPV1aR have nearly relation with female adult mandarin voles.ERβcan reduce the aggression behavior at a certain extent.The antisense nucleotides of estrogen receptorβcan enhance the preference of mandarin voles for mate or strange heterosexual vole,and strengthen the relation with mate in female adult mandarin voles.So,all behaviors were regulated by the nerve internal secretion mechanism.
本实验利用反义核酸技术和行为学检测平台相结合进行研究,通过给雌性棕色田鼠侧脑室中枢连续注射微量的雌激素受体βmRNA反义核酸,然后对实验鼠进行相关的行为检测;同时利用反义核酸技术和原位杂交组织化学技术相结合,探讨雌激素及其受体和催产素、加压素及其受体在雌性棕色田鼠脑内如何共同调节复杂的社会行为。
1.选择成体雌性棕色田鼠侧脑室中枢连续注射微量的ERβ受体mRNA反义核酸,分别在第一日、第三日及第五日同一时间麻醉注射。六小时后,利用攻击行为学检测平台进行实验,历时10分钟。实验结果显示:注射反义核酸组攻击行为的频次和持续时间明显高于注射随机序列组,而潜伏期没有显著性的差异。推断ERβmRNA的反义核酸在mRNA水平上阻止其翻译形成ERβ,进而阻碍了E及受体调控雌性棕色田鼠的攻击行为。
2.选择成体雌性棕色田鼠侧脑室中枢连续注射微量的ERβ受体mRNA反义核酸,分别在第一日、第三日及第五日同一时间麻醉注射。六小时后,利用配偶选择行为学检测平台进行实验,历时60分钟。实验结果显示:雌鼠在对陌生异性鼠箱、配偶鼠箱和中立箱中访问时间上有显著差异。说明给成年雌性棕色田鼠注射ERβmRNA的反义核酸后,使BST、CE、LSD、PVN和ME核团AVPV1aRmRNA和OTRmRNA表达增强,导致与其形成的神经投射功能发挥,从而表现出与之相应的行为活性增强,以致于增强了雌性棕色田鼠和配偶鼠之间的配偶联系。
3.选择成体雌性棕色田鼠侧脑室中枢连续注射微量的ERβ受体mRNA反义核酸,分别在第一日、第三日及第五日同一时间麻醉注射。六小时后,利用亲密行为学检测平台进行实验,历时60分钟。实验结果显示:雌鼠对配偶鼠的善待行为的增加,防御行为减弱。对陌生异性鼠的探究行为减弱,防御行为的增强。这些变化说明ERβ在田鼠偏好测试中,对配偶鼠和陌生异性鼠的社会识别可能存在着根本不同的神经通路,推测ERβ反义核酸可加强配偶关系。
4.选择成体雌性棕色田鼠侧脑室中枢连续注射微量的ERβmRNA反义核酸,分别在第一日、第三日及第五日同一时间麻醉注射。六小时后,利用原位杂交组织化学技术进行OTmRNA检测。实验结果显示:与对照组相比在棕色田鼠脑区核团中终纹床核(BST)和杏仁中央核(CE)的OTR的表达有显著性增强,在杏仁内侧核(ME)、下丘脑视前区(AH)和下丘脑室旁核(PVN)的OTR的表达有显著性减弱。说明在这些核团ERβ的反义核酸与OTR相互作用影响了雌性田鼠的社会行为,提示我们在理解E在介导雌性棕色田鼠的社会识别时,可能是通过催产素及其受体来发挥作用的。
5.选择成体雌性棕色田鼠侧脑室中枢连续注射微量的ERβ受体mRNA反义核酸,分别在第一日、第三日及第五日同一时间麻醉注射。六小时后,利用原位杂交组织化学技术进行AVPV1RmRNA检测。实验结果显示:与对照组相比在棕色田鼠脑区核团中终纹床核(BST)、下丘脑室旁核(PVN)和隔外侧核背部(LSD)核团中AVPV1aR的表达有显著性增加,在杏仁中央核(CE)的AVPV1aR表达显著性减弱,由此推测在这些核团ERβ的反义核酸与AVPR相互作用影响了雌性田鼠的社会行为。
由以上结论可得出:ERβ、OTR和AVPV1aR与雌性棕色田鼠的社会行为密切相关。ERβ可在一定程度上降低其攻击行为。ERβ对雌性棕色田鼠的配偶关系的调节及社会识别活动的调控是与OTR和AVPV1aR相关联的,ERβ反义核酸可提高雌性棕色田鼠的配偶识别能力,加强配偶之间的联系,即任何一种行为都受到复杂的神经内分泌机制的调控。
Animal social behavior,was controlled and regulated by centre nervous system and neuroendocrine system.Neuropeptide,sex hormone,a number of endocrine factors and environments where animal lives cooperate to affect social behavior of animals.Many experimental evidences indicate that E,OT and AVP acting in the brain appear to be critical for social discrimination.They could affect many social behaviors,such as social recognition,aggression, sexual behavior,partner preference,affinity behavior and individual behavior.Whether ERβcan regulate the OTRmRNA and AVPV1aRmRNA has not been reported.We preit that estrogen may show much regulation on the levels of OT and AVP messenger RNA.So,the present study will aim at the two problems:1,how E could affect some social behaviors in voles;2,Is there effects to OTR and AVPV1a expression by E receptor?
we used monogamous female mandarin voles(Microtus mandarinus) as research objects. Combining antisense nucleotides technic with behavioral observation,we examined the correlative behavior after i.c.v,continuous injection of antisense nucleotides of estrogen receptor-β.Meanwhile, Combined antisense nucleotides technic with in situ hybridization,we investigated how ERβ, vasopressin,oxytocin and its receptors controlled complex social behaviors in female voles.
1.Selected female adult mandarin voles were given i.c.v,continuous injections of antisense nucleotides of estrogen receptor-βrespectively in first day,third day and fifth day.After six hours, we examined aggression behavior during 10 minutes.The results were following:Compared with group of injection of scramble nucleotides,anisense group showed significant increase in frequency and total duration in aggression behavior.We suggested that antisense nucleotides of ERβinhibit the translation of ERβin mRNA levels.Furthermore,it blocked E to modulate aggression behavior in male voles.
2.Selected female adult mandarin voles were given i.c.v,continuous injections of antisense nucleotides of estrogen receptor-βin first day,third day and fifth day.After six hours,we examined partner preference behavior during 60 minutes.The results were following:There were significant differences in visiting duration in mate's compartment,strange heterosexual vole's compartment and neutral compartment,and significant increase in the number of AVPV1RmRNA and OTRmRNA in BST、CE、LSD、PVN and ME,It is suggested that continuous injection of antisense nucleotides of ERβcan enhance social recognition,and can enhance the pair bond formation of male and female voles.
3.Selected female adult mandarin voles were given i.c.v,continuous injections of antisense nucleotides of estrogen receptor-βrespectively in first day,third day and fifth day.After six hours, we examined affinity behavior during 60 minutes.The results were following:There were significant enhance in amicable behavior and decrease in defensive behavior with its mate.At the same time,there were significant enhance in defensive behavior and decrease in investigating behavior with the strange heterosexual voles.It suggested that there were different nervous circuit, which female voles recognized socially its mate and stranger voles in the preference behavior examination.
4.Selected female adult mandarin voles were given i.c.v,continuous injections of antisense nucleotides of estrogen receptor-βin first day,third day and fifth day.After six hours,OTR mRNA was examnined using in situ hybridization method in female voles.The results were following: There were significant differences in the BST,CE,ME,PVN and AH.Compared with injection of scramble nucleotides and injection of isotonic saline treatment,there were significant reduction in the number of OTR mRNA in AH,PVN and ME,and significant increase in the number of OTR mRNA in BST and CE Compared with injection of saline.These data suggested that the roles between antisense nucleotides of ERβand OTR affected social behavior in the female voles.It explained that ERβmodulated social behavior in female voles possibly through oxytocin and its receptor.
5.Selected female adult mandarin voles were given i.c.v,continuous injections of antisense nucleotides of estrogen receptor-βin first day,third day and fifth day.After six hours, AVPV1RmRNA was examnined using in situ hybridization method in female voles.The results were following:There were significant differences in the BST,CE,PVN and LSD.Compared with injection of scramble nucleotides and injection of isotonic saline treatment,there were significant increase in the number of AVPV1RmRNA in BST、PVN and LSD,and significant reduction in the number of AVPV1R mRNA in CE.These data suggested that the roles between antisense nucleotides of ERβand AVPV1R affected social behavior in the female voles.
Results from those conclusion that ERβ、OTR and AVPV1aR have nearly relation with female adult mandarin voles.ERβcan reduce the aggression behavior at a certain extent.The antisense nucleotides of estrogen receptorβcan enhance the preference of mandarin voles for mate or strange heterosexual vole,and strengthen the relation with mate in female adult mandarin voles.So,all behaviors were regulated by the nerve internal secretion mechanism.
引文
[1]Nomura M,McKenna E,Korach K.S,Pfaff D.W,Ogawa S,Estrogen receptor-βregulates transcript levels for oxytocin and arginine vasopressin in the hypothalamic paraventricular nucleus of male mice[J].Molecular Brain Research.109(2002)84-94.
[2]袁斌,丁丽华,褚春雨,叶棋浓.雌激素受体β的研究进展,生物技术通讯,2005,16(6),661-664
[3]Caldwell JD,Prange AJ,Jr.and Pedersen CA,Oxytocin facilitates the sexual receptivity of estrogen-treated female rats[J].Neuropeptides.2004,7(2):175-89
[4]唐业忠,王祖望.田鼠婚配制度的神经内分泌基础[J].兽类学报,2000,5,20[2]135-140.
[5]Kavaliers M,Choleris E,Agmo A,and Donald W.Pfaff[J].Hormones and Behavior.46(2004) 272-283.
[6]Herynk MH,Fuqua SA1 Est rogen receptor mutations in human disease[J].Endocr Rev.2004,25(6):8692-8981
[7]Muramatsu M,Inoue S1 Est rogen receptors:how do they control reproductive and nonreproductive functions[J].Biochem Biophys Res Commun.2000,270(1):12101
[8]Ogawa S,Inoue S,Watanabe T,et all The complete primary st ructure of human est rogen receptor beta(hER beta) and its heterodimerization with ER alpha in vivo and in vit ro[J].1 Biochem Biophys Res Commun.1998,243(1):1222-1261
[9]Krege JH,Hkdgin JB,Couse JF et al.Generation and reproductive phenotypes of mice lacking estrogen receptor[3[J].Proc Natl Acad USA.1998;95:15677-15682
[10]Shughrue PJ,Lane MV,Merchenthaler I.Comparative distribution of es-trogen receptor-alpha and beta mRNA in the rat central nervous system[J].Comp Neurol.1997,388(4):507-525.
[11]Li X,Schwartz PE,Rissman EF.Distribution of estrogen receptor-beta-like immunoreactivity in rat forebrain[J].Neuroendocrinology.1997,66(2):63-67.
[12]Hrabovszky E,Kallo I,Hajszan T,et al.Expression of estrogen receptor-beta messenger ribonucleic acid in oxytocin and vasopressin neurons of the rat supraoptic and paraventricular nuclei[J].Endocrinology.1998,139(5):2600-2604.
[13]张吉强,蔡文琴雌激素β受体免疫阳性神经元在成年雌性大鼠脑内的分布[J].解剖学报,2002,33(2):118-120.
[14]Ogawa S.Lubahn DB,Korach KS,Pfaff DW.Behavioral effects of estrogen receptor gene disruption in male mice [J]. Proc Natl Acad Sci U S A. 1997, 94:1476-1481.
[15]Ogawa S, Chester AE, Hewitt SC, Walker VR, Gustafsson JA, Smithies O, Korach KS, Pfaff DW. Abolition of male sexual behaviors in mice lacking estrogen receptors α and β(αβERKO) [J]. Natl Acad Sci U S A. 2000, 97:14737-14741.
[16]McEwen BS,Akama K,Alves S.Tracking the estrogen receptor in neurons:implications for estrogen -induced synapse formation[J].proc Natl Acad Sci USA.2001,98:7093-7100
[17]Wang CN,Chi CW,Lin YL et al.The neuroprotective effects of Phytoestrogens on Amyloid pprotein-induced toxicity are mediated by abrogating the activation of caspase cascade in rat cortical neurons [J]. Biol Chen.2001,276:5287-5295
[18]Xu H,Gouras GK,Greenfield JP et al.estrogen reduces neuronal generation of Alzheimer beta-amyloid peptide [J] .Nat Med. 1998,4:447—451
[19]Pike C,Estrogen modulates neuronal Bcl-XL expression and β-Amyloid-induced apoptosis:relevance to Alzheimer's disease[J]. Neurochem.1999;72:1552—1563.
[20]Kelly MJ,Levin ER.Rapid action of plasma membrane estrogen receptors [J]. Trends Endocrind Metab.2001;12:152-156
[21]Hlinak Z. Social recognition in ovariectomized and estradiol-treated female rats[J]. Horm. Behav.1993, 27:159-166.
[22]Imwalle DB, Scordalakes EM, Rissman EF. Estrogen receptor alpha influences socially motivated behaviors[J]. Horm. Behav.2002, 42:484—491.
[23]Clark AS, Kelton MC, Guarraci FA, Clyons EQ. Hormonal status and test condition, but not sexual experience, modulate partner preference in female rats[J]. Horm. Behav. 2004,45(5):314-323.
[24]Tang AC, Nakazawa M, Romeo RD, Reeb BC, Sisti H, McEwen BS. Effects of long-term estrogen replacement on social investigation and social memory in ovariectomized C57BL/6 mice[J]. Hormones and Behavior.2005, 47:350—357.
[25]Ferguson JN, Young LJ, Hearn EF, Matzuk MM, Insel TR, Winslow JT. Social amnesia in mice lacking the oxytocin gene[J]. Nat. Genet. 2000, 25:284—288.
[26]Ferguson JN, Aldag JM, Insel TR, Young LJ. Oxytocin in the medial amygdala is essential for social recognition in the mouse[J]. Neurosci. 2001, 21:8278— 8285.
[27]Simon N., Lu S., McKenna S., Chen X., Clifford A., Sexual dimorphisms in regulatiory systems for 24 sggression. In: Haug M, Whalen R, Aron C. Olsen K. editors.The development of sex differences and similarities in behavior[J].Boston:Kluwer Academic Publishers.1993,389-408.
[28]Lindzey J.,Wetsel W.C.,Couse J.F.,Stoker T.,Cooper R.,Korach K.S.,Effects castration and chronic steroid treatments on hypothalamic gonadotropin-releasing hommone content and pituitary gonadotropins in male wild-type and estrogen receptor-alpha knockout mice[J].Endocrinology.1998,139,4092-4101.
[29]Temple J.L.,Fugger H.N.,Li X.,Gustafsson J.A.,Rissman E.F.,Estrogen receptor β hypothalamic pituitary gonadal axis sensitivity to negative feedback[J].Soc.Neurosci.Abstr.,2000,26,540.20.
[30]Pendaries C.,Darblade B.,Rochaix P.,Krust A.,Chambon P.,Korache K.S.,Bayard F.,Arnal J.F.,The AF- 1 activation-function of ERα may be dispensable to mediate the effect of estradiol on endothelial NO production in mice[J].Proc.Natl.Acad.Sci.USA.,2002,99,2205-2210.
[31]Ogawa S.,Chester A.E.,Hewitt S.C.,Walker V.R.,Gustafsson J.A.,Smithies O.,Korach K.S.,Pfaff D.W.,Abolition of male sexual behaviors in mice lacking estrogen receptors α and β[J].Proc.Natl.Acad.Sci.U.S.A.,2000,97,14737-14741.
[32]Lu H.,Ozawa H.,Nishi M.,Ito T.,Kawata M.,Serotonergic neurones in the dorsal raphe nucleus that project into the medial preoptic area contain oestrogen receptor beta[J].Neuroendocrinol.2001,13(10),839-45.
[33]谢启文.神经肽[M].复旦大学出版社,2004年第一版,106-108.
[34]Ishunina TA,Dolzhikov AA,Grinevich VV.Activity of vasopressin neurons in the human supraoptic nucleus:estrogen inhibitory effect[J].Usp Fiziol Nauk,2001,32:48-59.
[35]何风琴,邰发道,张育辉,石靖.加压素和催产素在中枢神经系统中的作用[J].生物学通报,2006,41(8):10-13.
[36]Carter CS.Developmental consequences of oxytocin[J].Physiology and Behavior,2003,79:383-397.
[37]Cassoni P,Sapino A,Fortunati N,Munaron L,Chini B,Bussolati G.Oxytocin inhibits the proliferation of MDA-MB231 human breastcancer cells via cyclic adenosine monophosphate and protein kinase A[J].IntJ Cancer,1997,72:340-344.
[38]Paquin LDanalache BA,Jankowski M,McCann SM,Gutkowski J.Oxytocin induces differentiation of P19 embryonic stem cells to cardiomyocytes[M].Proc Natl Acad Sci USA.2002.99:9550-9555.
[39]Benelli A,Bertolini A,Poggioli R,Menozzi B,Basaglia R,Arletti R.Polymodal dose-response curve for oxytocin in the social recognition test[J].Neuropeptides.1995,28:251-5.
[40]Engelmann M,Ebner K,Wotjak CT,Landgraf R.Endogenous oxytocin is involved in short-term olfactory memory in female rats[J].Behav Brain Res,1998,90:89-94.
[41]Isadora F.Bielsky,Larry J.Young.Oxytocin,vasopressin,and social recognition in mammals[J].Peptides,2004,25:1565-1574.
[42]Van Wimersma Greidanus TB.Disturbed behavior and memory of the Brattelbororats[J].Ann NYAcad Sci,1982,394:655-662.
[43]Engelmann M,Landgraf R.Microdialysis administration of vasopressin into the septum improves social recognition in Brattlebororats[J].Physiol Behav 1994,55:145-149.
[44]Carter CS.Developmental consequences of oxytocin[J].Physiology and Behavior,2003,79:383-397.
[45]Winslow JT,Hastings N,Carter CS,Harbaugh CR,Insel YR.A role for central vasopressin in pair bonding in monogamous prairie voles[J].Nature,1993,365:545-548.
[46]Carter CS.Oxytocin and sexual behavior[J].Neurosci Biobehav Rev,1992,16:131-44.
[47]Keverne EB,Kendrick KM.Oxytocin facilitation of maternal behavior in sheep[J].Ann N Y Acad Sci,1992,652:83-101.
[48]Ferris C.Role of vasopressin in aggressive and dominate/subordinate behaviors[J].Ann N Y Acad Sci,1992,652:212-76.
[49]Irvin RW,Szot P,Dorsa DM,Potegal M,Ferris CF.Vasopressin in the septal area of the golden hamster controls scent marking and grooming[J].Physiol Behav,1990,48:693-9.
[50]Carter CS,De Vries AC,Getz LL.Physiological substrates of mammalian monogamy:the prairie vole model[J].Neurosci Biobehav Rev,1995,19:303-14.
[51]梁伟,袁崇刚.反义核酸技术在神经生物学上的应用[J].安徽农业大学学报,2004,31(3):278-281.
[52]Joseph Christopher Probstl.Antisense Oligodeoxynucleotide and Ribozyme design [J].METHODS 2000,22:271-281.
[53]Falker B,Herlitse S,Amthor B.Short antisense oligonucleotide-mediated inhibiton is strongly dependent oligo on length and concentration but almost independent of location of the targeted sequence[J].Biol Chem.,1994,269:16187-16194.
[54]Probst JC.Antisense oligodeoxynucleotide and ribozyme design[J].Methods.2000,22(3):271
[55]Chiasson B J,Armstrong J N,Hooper M L,Robertson H A.The appli cation of antisense oligonucleotide technology to the brain:some pitfalls[J].Cell Mol.Neurobiol.1994,14:507-521.
[56]Peyman A,Ryte A,Helsberg M,Mag M.Ehanced cellular uptake of G-rich oligonucleotides[J].Nucleosides Nucletides.1995,14:3-5.
[57]Stein C A,Mori K,Loke S L,Necker L M.Phosphorothioate and normal oligodeoxyribonucleotides with 5'-linked acridine:characterization and preliminary kinetics of cellular uptake[J].Gene,1988,72:333-341.
[58]Cossum PA,Sasmor H,Dellinger D,et al.Disposition of the 14C labeled phosphorothioate oligonucleotide ISIS 2105 after intravenous administration to rats[J].Phamacol Exp Ther,1993,267:1181-1190
[59]McCarthy MM,Auger AP,Mong JA,et al.Antisense oligodeoxynucleotides as a tool in developmental neuroendocrinology[J].Methods,.2000,22:239-248
[60]Wahlestedt C,Pich E M,Koob G F.Modulation of anxiety and neuropeptide Y-Y1receptors by antisense oligodeoxyribonucleotides[J].Sicence,1993,259:528-531.
[61]Standier K M,Chien C C,Wahlestedt C,Brown G P.Selective loss of-opioid analgesia and binding by antisense oligonucleotides to a-opioid receptor[J].Neuron,1994,12:805-810
[62]Gyurko R,Wielbo D,Phillips M I.Antisense inhibition of AT1 receptor m RNA and angiotensinogen m RNA in the brain of spontaneously hypertensive rats hypertension of neurogenic origion[J].Regul.Petides,1993,49:167-174.
[63]Neuman L,Kremarik P,Pittman Q J.Acute,sequence-specific effects of oxytocin and vasopressin antisense oligonucleotides on neuronal responses[J].Neuroscience,1995,69:997-1003.
[64]McCarthy M M,Books P J,Brown H E.Antisense oligonucleotides in behavior neuroscience[J].Methods Neurosci.1993,2:67-74.
[65]Ogawa S,Parhar L S.Pfaff D W.Effects of intrahypothalamic administration of antisense DNA for progesterone receptor m RNA on reproductive behavior and progesterone receptor immunoreactivity[J].Neurosci,1994,14:1766-1774.
[66]Abraham WC,Logan B,Thompson VL,et al.Sequence-independent effects of phosphorothiolated oligonucleotides on synaptic transmission and excitability in the hippocampus in vivo[J].Neurophamacology,1997,36:345-352.
[67]邰发道,王廷正.豫西黄土塬农作区棕色田鼠[J].陕西师范大学学报(自然科学版),1998a,26(4):82-86.
[68]邰发道,王廷正.棕色田鼠的种群年龄结构和密度关系分析[J].陕西师范大学学报(自然科学版),1997,25:95-97.
[69]邰发道,孙儒泳,王廷正,周宏伟.五种鼠的脑和头骨形态及其生态关系[J].动物学研究,2001a,22:472-477.
[70]邰发道,王廷正.棕色田鼠种群空间格局的研究[J].陕西师范大学学报(自然科学版),1998b,26(1):65-70.
[71]张育辉,刘加坤.七种啮齿动物视觉器官形态结构得比较研究[J].兽类学报,1994,14(3):189-194.
[72]李晓晨,王廷正.棕色田鼠与甘肃鼢鼠咀嚼肌结构和功能的比较[J].兽类学报,1999,19(4):308-314.
[73]邰发道,孙儒泳,王廷正,周宏伟.五种鼠的脑和头骨形态及其生态关系[J].动物学研究,2001 c,22(6):472-477.
[74]邰发道,王廷正,张育辉,郝琳,孙濡泳.棕色田鼠与沼泽田鼠犁鼻器和副嗅球的组织结构[J].动物学报,2003,49:248-255.
[75]何建平,郭进,安书成,史秀超,邰发道.棕色田鼠心电图的测定与分析[J].陕西师范大学学报(自然科学版),2001,29(3):91-94.
[76]徐金会,安书成,邰发道.棕色田鼠消化道形态变化与能量需求的关系[J].动物学报,2003,49(1):32-39.
[77]刘莹,李晓晨.棕色田鼠肾脏的显微结构[J].陕西师范大学学报(自然科学版),2003,31:223-225.
[78]李晓晨,王廷正.棕色田鼠与甘肃鼢鼠咀嚼肌结构和功能的比较[J].兽类学报,1999,19(4):308-314.
[79]黄惠敏,王廷正.棕色田鼠食性食量的研究[J].陕西师范大学学报(自然科学版),1999,27(3):88-92.
[80]邰发道,王廷正,闵一建.棕色田鼠的发声极其频谱分析[J].动物学研究,1999,20(4):278-283.
[81]邰发道,王廷正,张育辉,郝琳,孙濡泳.棕色田鼠与沼泽田鼠犁鼻器和副嗅球的组织结构[J].动物学报,2003,49:248-255.
[82]Tai FD,Wang TZ,Zhang YH,Sun RY.Sexual dimorphises of vomeralnasal organ in mandarin vole and reed vole[J].Acta Theriologica.2004,49(1):33-42.
[83]Tai FD,Broders HG,Lian Y,Wang TZ,Sun RY.Cellular activation patterns of the main olfactory bulb and accessory olfactory bulb following exposure to different bedding of mandarin vole[J].Can.J.Zool.2006,84:1138-1145.
[84]Tai ED.,Wang T.Z.,Zhao Y.J.,Mate choice and related characteristics of Mandarin voles(Microtus Mandarinus ),Acta Zoologica Sinica.2001b,47,260-267.
[85]金玫蕾,莫韫,刘留,郭宁,谢青莲,林桢,王星,李葆明,赵国屏,景乃禾,于雷.筛选神经 系统基因功能的行为学检测平台的探索[J].生理学报,2001,53(4):247-251.
[86]曹长姝,隋鸿锦。反义核酸技术及应用[J].大连医科大学学报,2003,4:300-302.
[87]金玫蕾,郭宁,李晨,丁澄,谢青莲,邹洪,王凌燕,于军,景乃禾.用不同实验小鼠品系对10个基因进行功能复筛的研究[J].上海实验动物科学,2002,3:133-137.
[88]Arnaud Nicot,Donald W Pfaff.Antisense oligodeoxynucleotides as specific tools for studying neuroendocrine and behavioral functions:Some prospects and problems[J].Journal of Neuroscience Methods.1997,71:45-53.
[89]金玫蕾,莫韫,刘留,郭宁,谢青莲,林桢,王星,李葆明,赵国屏,景乃禾,于雷.筛选神经系统基因功能的行为学检测平台的探索[J].生理学报,2001,53(4):247-251.
[90]金玫蕾,刘留,郭宁,莫韫,谢青莲,李葆明,赵国屏,景乃禾.BALBPc和C57BLP6小鼠在基因功能检测中行为学的比较研究[J].中国实验动物学报,2001,9:3-9.
[100]金玫蕾,莫韫,刘留,郭宁,谢青莲,林桢,王星,李葆明,赵国屏,景乃禾,于雷.筛选神经系统基因功能的行为学检测平台的探索[J].生理学报,2001,53(4):247-251.
[101]Zuoxin Wang,Brandon J Aragona.Neurochemical regulation of pairbonding in male prairie voles[J].Physiology & Behavior,2004,83:319-328.
[102]R Lucille Roberts,Amy Zullo,Eric A Gustafson,C Sue Carter.Perinatala steroid treatments alter alloparental and affiliative behavior in Prairie Voles[J].Hormones and Behavior.1996,30:576-582.
[103]孔雀.邰发道.攻击行为神经机制的研究进展[J].现代生物医学进展.2006.6(8):55-58.
[104]Lindzey J.,Wetsel W.C..Couse J.F.,Stoker T.,Cooper R.,Korach K.S..Effects castration and chronic steroid treatments on hypothalamic gonadotropin -releasing hommone content and pituitary gonadotropins in male wild-type and estrogen receptor-alpha knockout mice[J]. Endocrinology. 1998, 139,4092-4101.
[105]Temple J.L., Fugger H.N., Li X., Gustafsson J. A., Rissman E.F., Estrogen receptor β hypothalamic pituitary gonadal axis sensitivity to negative feedback, Soc[J]. Neurosci., Abstr.2000,26,540.20.
[106]Tai FD,Wng TZ,Zhou HW,Mating system of mandarin vole (Microtus mandarinus) [J]. Acta Zoologica Sinica .2001,47: 260-267
[107]He F. Q., Tai F. D., Zhang Y. H., An SH., CH. 2004. The relationship between social behavior and the expression of estrogen receptor β and androgen receptor in olfactory-related brain regions of the male mandarin vole Microtus mandarinus and reed vole M.fostis[J]. Acta Zoologica Sinica. 50(2): 165-175.
[108] Zuoxin Wang, Brandon J Aragona. Neurochemical regulation of pairbonding in male prairie voles [J]. Physiology & Behavior, 2004, 83: 319-328.
[109]Getz L L, Carter S C, Gavish L. The mating system of the prairie vole, Microtus ochrogaster: field and laboratory evidence for pair bonding [J].Behav Ecol Sociobiol, 1981,8: 189-194.
[110]Kramer K, Cushing BS, Carter CS, Wu J, Ottinger MA. Chemical and biological validation of an enzyme immunoassay for plasma oxytocin[J]. Horm. Behav, 2002, 41:476.
[111] Insel TR, Hulihan T. A gender-specific mechanism for pair bonding: oxytocin and partner preference formation in monogamous voles[J]. Behav Neurosci, 1995, 109:782-9.
[112]Lim MM, Young LJ. Vasopressin-dependent neural circuits underlying pair bond formation in the monogamous prairie vole[J]. 2004, 125(1):35-45.
[113]Nomura M, McKenna E, Korach K. S ,Pfaff D.W, Ogawa S, Estrogen receptor-β regulates transcript levels for oxytocin and arginine vasopressin in the hypothalamic paraventricular nucleus of male mice[J]. Molecular Brain Research 109(2002)84—94
[114]Michael J. Weiser, Chad D. Foradori, Robert J. Handa Estrogen receptor beta in the brain[J]. From form to function.brain research reviews. 2007.100509.1 - 12
[115]Scordalakes E.M, Imwalle D.B, Rissman E.F., Oestrogen's masculine side: mediation of mating in male mice[J]. Reproduction, 2002,124(3),331 -8.
[116]Warembourg M., Leroy D., Comparative distribution of estrogen receptor a and β immunoreactivities in the forebrain and the midbrain of the female guinea pig[J].Brain Research, 2004,1002,55—66.
[117]Zhang J.X., Zhang Z.B., Wang Z.W., Scent, social status, and reproductive condition in rat-like hamsters(Cricetulus triton) [J].Physiol Behav. 2001. 74(4-5),415-20.
[118]Fugger H.N.,Foster T.C.,Gustafsson J.A.,Rissman E.F.,Novel effects of estradiol and estrogen receptor α and β on cognitive function,Brain Research,2000,883,258-264.
[119]梁伟,袁崇刚.反义核酸技术在神经生物学上的应用[J].安徽农业大学学报,2004,31(3):278-281
[120]Shibata S,Yamamoto TY,Ueki S.Differential effects of medial,central and basolateral amygdaloid lesions on four models of experimentally-induced aggression in rats[J].Physiol Behav,1982,28:289-94.
[121]Gammie SC,Nelson RJ.cFOS and pCREB activation and maternal aggression in mice[J].Brain Res,2001,898:23-41.
[122]Wang Z,Hulihan TJ,Insel TR.Sexual and social experience is associated with different patterns of behavior and neural activation in male prairie voles[J].Brain Res,1997,767:31-32.
[123]Bunnell BN,Sodetz F J,Shalloway DI.Amygdaloid lesions and social behavior in the golden hamster[J].Physiol Behav,1970,5:51-61.
[124]Meisel,Sachs B.Cellular and molecular mechanisms of female reproductive behaviors.The Physiology of Reproduction[M].2th-end.USA.New York.1994,2:5-6.
[125]Yahr P,Finn PD,Hoffman NW,Sayag N.Sexually dimorphic cell groups in the medial preoptic area that are essential for male sex behavior and the neural pathways needed for their effects[J].Psychoneuroendocrinology.1994,19:463-470.
[126]Ferguson JN,Aldag JM,Insel TR,Young LJ.Oxytocin in the medial amygdala is essential for social recognition in the mouse[J].Neurosci,2001,21:8278-8285.
[127]Newman SW.The medial extended amygdala in male reproductive behavior- A node in the mammalian social behavior network[J].Ann.N.Y.Acad.Sci,1999,877:242-257.
[128]Rasia-Filho AA,Londero RG,Achaval M.Functional activities of the amygdala:an overview[J].J.Psych.Neurosci,2000,25:14-23.
[129]Cushing BS.Kramer KM.Mechanisms underlying epigenetic effects of early social experience:The role of neruopeptides and steroids[J].Neuroscience and Biobehavioral Reviews,2005,29:1089-1105.
[130]Jackson D,Burns R,Trksak G,Simeone B,DeLeon KR,Connor DF,Harrison RJ,Melloni RH Jr.Anterior hypothalamic vasopressin modulates the aggression-stimulating effects of adolescent cocaine exposure in Syrian hamsters[J]. Neuroscience.2005, 133:635—646.
[131]Liu Y, Curtis J T, Wang Z X. Vasopressin in the lateral septum regulates pair bond formation in male prairie voles (Microtus ochrogaster) [J]. Behav Neurosci, 2001, 115:910-919.
[132]Cho M M, DeVries A C, Williams J R, Carter C S. The effects of oxytocin and vasopressin on partner preferences in male and female prairie voles (Microtus ochrogaster) [J]. Behav Neurosci. 1999, 113: 1071-1079.
[133]Landgraf R, Frank E, Aldag J, Neumann I, Sharer C, Ren X, Terwilliger E, Niwa M, Wigger A, Young L. Viral vector-mediated gene transfer of the vole V1a vasopressin receptor in the rat septum: improved social discrimination and active social behaviour [J]. Eur J Neurosci. 2003, 18: 403-411.
[2]袁斌,丁丽华,褚春雨,叶棋浓.雌激素受体β的研究进展,生物技术通讯,2005,16(6),661-664
[3]Caldwell JD,Prange AJ,Jr.and Pedersen CA,Oxytocin facilitates the sexual receptivity of estrogen-treated female rats[J].Neuropeptides.2004,7(2):175-89
[4]唐业忠,王祖望.田鼠婚配制度的神经内分泌基础[J].兽类学报,2000,5,20[2]135-140.
[5]Kavaliers M,Choleris E,Agmo A,and Donald W.Pfaff[J].Hormones and Behavior.46(2004) 272-283.
[6]Herynk MH,Fuqua SA1 Est rogen receptor mutations in human disease[J].Endocr Rev.2004,25(6):8692-8981
[7]Muramatsu M,Inoue S1 Est rogen receptors:how do they control reproductive and nonreproductive functions[J].Biochem Biophys Res Commun.2000,270(1):12101
[8]Ogawa S,Inoue S,Watanabe T,et all The complete primary st ructure of human est rogen receptor beta(hER beta) and its heterodimerization with ER alpha in vivo and in vit ro[J].1 Biochem Biophys Res Commun.1998,243(1):1222-1261
[9]Krege JH,Hkdgin JB,Couse JF et al.Generation and reproductive phenotypes of mice lacking estrogen receptor[3[J].Proc Natl Acad USA.1998;95:15677-15682
[10]Shughrue PJ,Lane MV,Merchenthaler I.Comparative distribution of es-trogen receptor-alpha and beta mRNA in the rat central nervous system[J].Comp Neurol.1997,388(4):507-525.
[11]Li X,Schwartz PE,Rissman EF.Distribution of estrogen receptor-beta-like immunoreactivity in rat forebrain[J].Neuroendocrinology.1997,66(2):63-67.
[12]Hrabovszky E,Kallo I,Hajszan T,et al.Expression of estrogen receptor-beta messenger ribonucleic acid in oxytocin and vasopressin neurons of the rat supraoptic and paraventricular nuclei[J].Endocrinology.1998,139(5):2600-2604.
[13]张吉强,蔡文琴雌激素β受体免疫阳性神经元在成年雌性大鼠脑内的分布[J].解剖学报,2002,33(2):118-120.
[14]Ogawa S.Lubahn DB,Korach KS,Pfaff DW.Behavioral effects of estrogen receptor gene disruption in male mice [J]. Proc Natl Acad Sci U S A. 1997, 94:1476-1481.
[15]Ogawa S, Chester AE, Hewitt SC, Walker VR, Gustafsson JA, Smithies O, Korach KS, Pfaff DW. Abolition of male sexual behaviors in mice lacking estrogen receptors α and β(αβERKO) [J]. Natl Acad Sci U S A. 2000, 97:14737-14741.
[16]McEwen BS,Akama K,Alves S.Tracking the estrogen receptor in neurons:implications for estrogen -induced synapse formation[J].proc Natl Acad Sci USA.2001,98:7093-7100
[17]Wang CN,Chi CW,Lin YL et al.The neuroprotective effects of Phytoestrogens on Amyloid pprotein-induced toxicity are mediated by abrogating the activation of caspase cascade in rat cortical neurons [J]. Biol Chen.2001,276:5287-5295
[18]Xu H,Gouras GK,Greenfield JP et al.estrogen reduces neuronal generation of Alzheimer beta-amyloid peptide [J] .Nat Med. 1998,4:447—451
[19]Pike C,Estrogen modulates neuronal Bcl-XL expression and β-Amyloid-induced apoptosis:relevance to Alzheimer's disease[J]. Neurochem.1999;72:1552—1563.
[20]Kelly MJ,Levin ER.Rapid action of plasma membrane estrogen receptors [J]. Trends Endocrind Metab.2001;12:152-156
[21]Hlinak Z. Social recognition in ovariectomized and estradiol-treated female rats[J]. Horm. Behav.1993, 27:159-166.
[22]Imwalle DB, Scordalakes EM, Rissman EF. Estrogen receptor alpha influences socially motivated behaviors[J]. Horm. Behav.2002, 42:484—491.
[23]Clark AS, Kelton MC, Guarraci FA, Clyons EQ. Hormonal status and test condition, but not sexual experience, modulate partner preference in female rats[J]. Horm. Behav. 2004,45(5):314-323.
[24]Tang AC, Nakazawa M, Romeo RD, Reeb BC, Sisti H, McEwen BS. Effects of long-term estrogen replacement on social investigation and social memory in ovariectomized C57BL/6 mice[J]. Hormones and Behavior.2005, 47:350—357.
[25]Ferguson JN, Young LJ, Hearn EF, Matzuk MM, Insel TR, Winslow JT. Social amnesia in mice lacking the oxytocin gene[J]. Nat. Genet. 2000, 25:284—288.
[26]Ferguson JN, Aldag JM, Insel TR, Young LJ. Oxytocin in the medial amygdala is essential for social recognition in the mouse[J]. Neurosci. 2001, 21:8278— 8285.
[27]Simon N., Lu S., McKenna S., Chen X., Clifford A., Sexual dimorphisms in regulatiory systems for 24 sggression. In: Haug M, Whalen R, Aron C. Olsen K. editors.The development of sex differences and similarities in behavior[J].Boston:Kluwer Academic Publishers.1993,389-408.
[28]Lindzey J.,Wetsel W.C.,Couse J.F.,Stoker T.,Cooper R.,Korach K.S.,Effects castration and chronic steroid treatments on hypothalamic gonadotropin-releasing hommone content and pituitary gonadotropins in male wild-type and estrogen receptor-alpha knockout mice[J].Endocrinology.1998,139,4092-4101.
[29]Temple J.L.,Fugger H.N.,Li X.,Gustafsson J.A.,Rissman E.F.,Estrogen receptor β hypothalamic pituitary gonadal axis sensitivity to negative feedback[J].Soc.Neurosci.Abstr.,2000,26,540.20.
[30]Pendaries C.,Darblade B.,Rochaix P.,Krust A.,Chambon P.,Korache K.S.,Bayard F.,Arnal J.F.,The AF- 1 activation-function of ERα may be dispensable to mediate the effect of estradiol on endothelial NO production in mice[J].Proc.Natl.Acad.Sci.USA.,2002,99,2205-2210.
[31]Ogawa S.,Chester A.E.,Hewitt S.C.,Walker V.R.,Gustafsson J.A.,Smithies O.,Korach K.S.,Pfaff D.W.,Abolition of male sexual behaviors in mice lacking estrogen receptors α and β[J].Proc.Natl.Acad.Sci.U.S.A.,2000,97,14737-14741.
[32]Lu H.,Ozawa H.,Nishi M.,Ito T.,Kawata M.,Serotonergic neurones in the dorsal raphe nucleus that project into the medial preoptic area contain oestrogen receptor beta[J].Neuroendocrinol.2001,13(10),839-45.
[33]谢启文.神经肽[M].复旦大学出版社,2004年第一版,106-108.
[34]Ishunina TA,Dolzhikov AA,Grinevich VV.Activity of vasopressin neurons in the human supraoptic nucleus:estrogen inhibitory effect[J].Usp Fiziol Nauk,2001,32:48-59.
[35]何风琴,邰发道,张育辉,石靖.加压素和催产素在中枢神经系统中的作用[J].生物学通报,2006,41(8):10-13.
[36]Carter CS.Developmental consequences of oxytocin[J].Physiology and Behavior,2003,79:383-397.
[37]Cassoni P,Sapino A,Fortunati N,Munaron L,Chini B,Bussolati G.Oxytocin inhibits the proliferation of MDA-MB231 human breastcancer cells via cyclic adenosine monophosphate and protein kinase A[J].IntJ Cancer,1997,72:340-344.
[38]Paquin LDanalache BA,Jankowski M,McCann SM,Gutkowski J.Oxytocin induces differentiation of P19 embryonic stem cells to cardiomyocytes[M].Proc Natl Acad Sci USA.2002.99:9550-9555.
[39]Benelli A,Bertolini A,Poggioli R,Menozzi B,Basaglia R,Arletti R.Polymodal dose-response curve for oxytocin in the social recognition test[J].Neuropeptides.1995,28:251-5.
[40]Engelmann M,Ebner K,Wotjak CT,Landgraf R.Endogenous oxytocin is involved in short-term olfactory memory in female rats[J].Behav Brain Res,1998,90:89-94.
[41]Isadora F.Bielsky,Larry J.Young.Oxytocin,vasopressin,and social recognition in mammals[J].Peptides,2004,25:1565-1574.
[42]Van Wimersma Greidanus TB.Disturbed behavior and memory of the Brattelbororats[J].Ann NYAcad Sci,1982,394:655-662.
[43]Engelmann M,Landgraf R.Microdialysis administration of vasopressin into the septum improves social recognition in Brattlebororats[J].Physiol Behav 1994,55:145-149.
[44]Carter CS.Developmental consequences of oxytocin[J].Physiology and Behavior,2003,79:383-397.
[45]Winslow JT,Hastings N,Carter CS,Harbaugh CR,Insel YR.A role for central vasopressin in pair bonding in monogamous prairie voles[J].Nature,1993,365:545-548.
[46]Carter CS.Oxytocin and sexual behavior[J].Neurosci Biobehav Rev,1992,16:131-44.
[47]Keverne EB,Kendrick KM.Oxytocin facilitation of maternal behavior in sheep[J].Ann N Y Acad Sci,1992,652:83-101.
[48]Ferris C.Role of vasopressin in aggressive and dominate/subordinate behaviors[J].Ann N Y Acad Sci,1992,652:212-76.
[49]Irvin RW,Szot P,Dorsa DM,Potegal M,Ferris CF.Vasopressin in the septal area of the golden hamster controls scent marking and grooming[J].Physiol Behav,1990,48:693-9.
[50]Carter CS,De Vries AC,Getz LL.Physiological substrates of mammalian monogamy:the prairie vole model[J].Neurosci Biobehav Rev,1995,19:303-14.
[51]梁伟,袁崇刚.反义核酸技术在神经生物学上的应用[J].安徽农业大学学报,2004,31(3):278-281.
[52]Joseph Christopher Probstl.Antisense Oligodeoxynucleotide and Ribozyme design [J].METHODS 2000,22:271-281.
[53]Falker B,Herlitse S,Amthor B.Short antisense oligonucleotide-mediated inhibiton is strongly dependent oligo on length and concentration but almost independent of location of the targeted sequence[J].Biol Chem.,1994,269:16187-16194.
[54]Probst JC.Antisense oligodeoxynucleotide and ribozyme design[J].Methods.2000,22(3):271
[55]Chiasson B J,Armstrong J N,Hooper M L,Robertson H A.The appli cation of antisense oligonucleotide technology to the brain:some pitfalls[J].Cell Mol.Neurobiol.1994,14:507-521.
[56]Peyman A,Ryte A,Helsberg M,Mag M.Ehanced cellular uptake of G-rich oligonucleotides[J].Nucleosides Nucletides.1995,14:3-5.
[57]Stein C A,Mori K,Loke S L,Necker L M.Phosphorothioate and normal oligodeoxyribonucleotides with 5'-linked acridine:characterization and preliminary kinetics of cellular uptake[J].Gene,1988,72:333-341.
[58]Cossum PA,Sasmor H,Dellinger D,et al.Disposition of the 14C labeled phosphorothioate oligonucleotide ISIS 2105 after intravenous administration to rats[J].Phamacol Exp Ther,1993,267:1181-1190
[59]McCarthy MM,Auger AP,Mong JA,et al.Antisense oligodeoxynucleotides as a tool in developmental neuroendocrinology[J].Methods,.2000,22:239-248
[60]Wahlestedt C,Pich E M,Koob G F.Modulation of anxiety and neuropeptide Y-Y1receptors by antisense oligodeoxyribonucleotides[J].Sicence,1993,259:528-531.
[61]Standier K M,Chien C C,Wahlestedt C,Brown G P.Selective loss of-opioid analgesia and binding by antisense oligonucleotides to a-opioid receptor[J].Neuron,1994,12:805-810
[62]Gyurko R,Wielbo D,Phillips M I.Antisense inhibition of AT1 receptor m RNA and angiotensinogen m RNA in the brain of spontaneously hypertensive rats hypertension of neurogenic origion[J].Regul.Petides,1993,49:167-174.
[63]Neuman L,Kremarik P,Pittman Q J.Acute,sequence-specific effects of oxytocin and vasopressin antisense oligonucleotides on neuronal responses[J].Neuroscience,1995,69:997-1003.
[64]McCarthy M M,Books P J,Brown H E.Antisense oligonucleotides in behavior neuroscience[J].Methods Neurosci.1993,2:67-74.
[65]Ogawa S,Parhar L S.Pfaff D W.Effects of intrahypothalamic administration of antisense DNA for progesterone receptor m RNA on reproductive behavior and progesterone receptor immunoreactivity[J].Neurosci,1994,14:1766-1774.
[66]Abraham WC,Logan B,Thompson VL,et al.Sequence-independent effects of phosphorothiolated oligonucleotides on synaptic transmission and excitability in the hippocampus in vivo[J].Neurophamacology,1997,36:345-352.
[67]邰发道,王廷正.豫西黄土塬农作区棕色田鼠[J].陕西师范大学学报(自然科学版),1998a,26(4):82-86.
[68]邰发道,王廷正.棕色田鼠的种群年龄结构和密度关系分析[J].陕西师范大学学报(自然科学版),1997,25:95-97.
[69]邰发道,孙儒泳,王廷正,周宏伟.五种鼠的脑和头骨形态及其生态关系[J].动物学研究,2001a,22:472-477.
[70]邰发道,王廷正.棕色田鼠种群空间格局的研究[J].陕西师范大学学报(自然科学版),1998b,26(1):65-70.
[71]张育辉,刘加坤.七种啮齿动物视觉器官形态结构得比较研究[J].兽类学报,1994,14(3):189-194.
[72]李晓晨,王廷正.棕色田鼠与甘肃鼢鼠咀嚼肌结构和功能的比较[J].兽类学报,1999,19(4):308-314.
[73]邰发道,孙儒泳,王廷正,周宏伟.五种鼠的脑和头骨形态及其生态关系[J].动物学研究,2001 c,22(6):472-477.
[74]邰发道,王廷正,张育辉,郝琳,孙濡泳.棕色田鼠与沼泽田鼠犁鼻器和副嗅球的组织结构[J].动物学报,2003,49:248-255.
[75]何建平,郭进,安书成,史秀超,邰发道.棕色田鼠心电图的测定与分析[J].陕西师范大学学报(自然科学版),2001,29(3):91-94.
[76]徐金会,安书成,邰发道.棕色田鼠消化道形态变化与能量需求的关系[J].动物学报,2003,49(1):32-39.
[77]刘莹,李晓晨.棕色田鼠肾脏的显微结构[J].陕西师范大学学报(自然科学版),2003,31:223-225.
[78]李晓晨,王廷正.棕色田鼠与甘肃鼢鼠咀嚼肌结构和功能的比较[J].兽类学报,1999,19(4):308-314.
[79]黄惠敏,王廷正.棕色田鼠食性食量的研究[J].陕西师范大学学报(自然科学版),1999,27(3):88-92.
[80]邰发道,王廷正,闵一建.棕色田鼠的发声极其频谱分析[J].动物学研究,1999,20(4):278-283.
[81]邰发道,王廷正,张育辉,郝琳,孙濡泳.棕色田鼠与沼泽田鼠犁鼻器和副嗅球的组织结构[J].动物学报,2003,49:248-255.
[82]Tai FD,Wang TZ,Zhang YH,Sun RY.Sexual dimorphises of vomeralnasal organ in mandarin vole and reed vole[J].Acta Theriologica.2004,49(1):33-42.
[83]Tai FD,Broders HG,Lian Y,Wang TZ,Sun RY.Cellular activation patterns of the main olfactory bulb and accessory olfactory bulb following exposure to different bedding of mandarin vole[J].Can.J.Zool.2006,84:1138-1145.
[84]Tai ED.,Wang T.Z.,Zhao Y.J.,Mate choice and related characteristics of Mandarin voles(Microtus Mandarinus ),Acta Zoologica Sinica.2001b,47,260-267.
[85]金玫蕾,莫韫,刘留,郭宁,谢青莲,林桢,王星,李葆明,赵国屏,景乃禾,于雷.筛选神经 系统基因功能的行为学检测平台的探索[J].生理学报,2001,53(4):247-251.
[86]曹长姝,隋鸿锦。反义核酸技术及应用[J].大连医科大学学报,2003,4:300-302.
[87]金玫蕾,郭宁,李晨,丁澄,谢青莲,邹洪,王凌燕,于军,景乃禾.用不同实验小鼠品系对10个基因进行功能复筛的研究[J].上海实验动物科学,2002,3:133-137.
[88]Arnaud Nicot,Donald W Pfaff.Antisense oligodeoxynucleotides as specific tools for studying neuroendocrine and behavioral functions:Some prospects and problems[J].Journal of Neuroscience Methods.1997,71:45-53.
[89]金玫蕾,莫韫,刘留,郭宁,谢青莲,林桢,王星,李葆明,赵国屏,景乃禾,于雷.筛选神经系统基因功能的行为学检测平台的探索[J].生理学报,2001,53(4):247-251.
[90]金玫蕾,刘留,郭宁,莫韫,谢青莲,李葆明,赵国屏,景乃禾.BALBPc和C57BLP6小鼠在基因功能检测中行为学的比较研究[J].中国实验动物学报,2001,9:3-9.
[100]金玫蕾,莫韫,刘留,郭宁,谢青莲,林桢,王星,李葆明,赵国屏,景乃禾,于雷.筛选神经系统基因功能的行为学检测平台的探索[J].生理学报,2001,53(4):247-251.
[101]Zuoxin Wang,Brandon J Aragona.Neurochemical regulation of pairbonding in male prairie voles[J].Physiology & Behavior,2004,83:319-328.
[102]R Lucille Roberts,Amy Zullo,Eric A Gustafson,C Sue Carter.Perinatala steroid treatments alter alloparental and affiliative behavior in Prairie Voles[J].Hormones and Behavior.1996,30:576-582.
[103]孔雀.邰发道.攻击行为神经机制的研究进展[J].现代生物医学进展.2006.6(8):55-58.
[104]Lindzey J.,Wetsel W.C..Couse J.F.,Stoker T.,Cooper R.,Korach K.S..Effects castration and chronic steroid treatments on hypothalamic gonadotropin -releasing hommone content and pituitary gonadotropins in male wild-type and estrogen receptor-alpha knockout mice[J]. Endocrinology. 1998, 139,4092-4101.
[105]Temple J.L., Fugger H.N., Li X., Gustafsson J. A., Rissman E.F., Estrogen receptor β hypothalamic pituitary gonadal axis sensitivity to negative feedback, Soc[J]. Neurosci., Abstr.2000,26,540.20.
[106]Tai FD,Wng TZ,Zhou HW,Mating system of mandarin vole (Microtus mandarinus) [J]. Acta Zoologica Sinica .2001,47: 260-267
[107]He F. Q., Tai F. D., Zhang Y. H., An SH., CH. 2004. The relationship between social behavior and the expression of estrogen receptor β and androgen receptor in olfactory-related brain regions of the male mandarin vole Microtus mandarinus and reed vole M.fostis[J]. Acta Zoologica Sinica. 50(2): 165-175.
[108] Zuoxin Wang, Brandon J Aragona. Neurochemical regulation of pairbonding in male prairie voles [J]. Physiology & Behavior, 2004, 83: 319-328.
[109]Getz L L, Carter S C, Gavish L. The mating system of the prairie vole, Microtus ochrogaster: field and laboratory evidence for pair bonding [J].Behav Ecol Sociobiol, 1981,8: 189-194.
[110]Kramer K, Cushing BS, Carter CS, Wu J, Ottinger MA. Chemical and biological validation of an enzyme immunoassay for plasma oxytocin[J]. Horm. Behav, 2002, 41:476.
[111] Insel TR, Hulihan T. A gender-specific mechanism for pair bonding: oxytocin and partner preference formation in monogamous voles[J]. Behav Neurosci, 1995, 109:782-9.
[112]Lim MM, Young LJ. Vasopressin-dependent neural circuits underlying pair bond formation in the monogamous prairie vole[J]. 2004, 125(1):35-45.
[113]Nomura M, McKenna E, Korach K. S ,Pfaff D.W, Ogawa S, Estrogen receptor-β regulates transcript levels for oxytocin and arginine vasopressin in the hypothalamic paraventricular nucleus of male mice[J]. Molecular Brain Research 109(2002)84—94
[114]Michael J. Weiser, Chad D. Foradori, Robert J. Handa Estrogen receptor beta in the brain[J]. From form to function.brain research reviews. 2007.100509.1 - 12
[115]Scordalakes E.M, Imwalle D.B, Rissman E.F., Oestrogen's masculine side: mediation of mating in male mice[J]. Reproduction, 2002,124(3),331 -8.
[116]Warembourg M., Leroy D., Comparative distribution of estrogen receptor a and β immunoreactivities in the forebrain and the midbrain of the female guinea pig[J].Brain Research, 2004,1002,55—66.
[117]Zhang J.X., Zhang Z.B., Wang Z.W., Scent, social status, and reproductive condition in rat-like hamsters(Cricetulus triton) [J].Physiol Behav. 2001. 74(4-5),415-20.
[118]Fugger H.N.,Foster T.C.,Gustafsson J.A.,Rissman E.F.,Novel effects of estradiol and estrogen receptor α and β on cognitive function,Brain Research,2000,883,258-264.
[119]梁伟,袁崇刚.反义核酸技术在神经生物学上的应用[J].安徽农业大学学报,2004,31(3):278-281
[120]Shibata S,Yamamoto TY,Ueki S.Differential effects of medial,central and basolateral amygdaloid lesions on four models of experimentally-induced aggression in rats[J].Physiol Behav,1982,28:289-94.
[121]Gammie SC,Nelson RJ.cFOS and pCREB activation and maternal aggression in mice[J].Brain Res,2001,898:23-41.
[122]Wang Z,Hulihan TJ,Insel TR.Sexual and social experience is associated with different patterns of behavior and neural activation in male prairie voles[J].Brain Res,1997,767:31-32.
[123]Bunnell BN,Sodetz F J,Shalloway DI.Amygdaloid lesions and social behavior in the golden hamster[J].Physiol Behav,1970,5:51-61.
[124]Meisel,Sachs B.Cellular and molecular mechanisms of female reproductive behaviors.The Physiology of Reproduction[M].2th-end.USA.New York.1994,2:5-6.
[125]Yahr P,Finn PD,Hoffman NW,Sayag N.Sexually dimorphic cell groups in the medial preoptic area that are essential for male sex behavior and the neural pathways needed for their effects[J].Psychoneuroendocrinology.1994,19:463-470.
[126]Ferguson JN,Aldag JM,Insel TR,Young LJ.Oxytocin in the medial amygdala is essential for social recognition in the mouse[J].Neurosci,2001,21:8278-8285.
[127]Newman SW.The medial extended amygdala in male reproductive behavior- A node in the mammalian social behavior network[J].Ann.N.Y.Acad.Sci,1999,877:242-257.
[128]Rasia-Filho AA,Londero RG,Achaval M.Functional activities of the amygdala:an overview[J].J.Psych.Neurosci,2000,25:14-23.
[129]Cushing BS.Kramer KM.Mechanisms underlying epigenetic effects of early social experience:The role of neruopeptides and steroids[J].Neuroscience and Biobehavioral Reviews,2005,29:1089-1105.
[130]Jackson D,Burns R,Trksak G,Simeone B,DeLeon KR,Connor DF,Harrison RJ,Melloni RH Jr.Anterior hypothalamic vasopressin modulates the aggression-stimulating effects of adolescent cocaine exposure in Syrian hamsters[J]. Neuroscience.2005, 133:635—646.
[131]Liu Y, Curtis J T, Wang Z X. Vasopressin in the lateral septum regulates pair bond formation in male prairie voles (Microtus ochrogaster) [J]. Behav Neurosci, 2001, 115:910-919.
[132]Cho M M, DeVries A C, Williams J R, Carter C S. The effects of oxytocin and vasopressin on partner preferences in male and female prairie voles (Microtus ochrogaster) [J]. Behav Neurosci. 1999, 113: 1071-1079.
[133]Landgraf R, Frank E, Aldag J, Neumann I, Sharer C, Ren X, Terwilliger E, Niwa M, Wigger A, Young L. Viral vector-mediated gene transfer of the vole V1a vasopressin receptor in the rat septum: improved social discrimination and active social behaviour [J]. Eur J Neurosci. 2003, 18: 403-411.