光诱导c-fos基因在母鸡中脑的表达
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摘要
本试验用c-fos法探讨光信息在母鸡中脑的传导通路,为深入了解光照对母鸡生产性能的影响机理打下基础。在养鸡生产中,光照是鸡赖以生存的环境中的重要因素之一;科学利用光照可以进一步提高鸡的生产性能,如:促进鸡的生长发育,调节性成熟,提高产蛋率、受精率、种蛋合格率等。禽类是最依赖视觉系统的脊椎动物。鸟类的视觉系统主要分为两个平行的视觉信息加工通路:离顶盖通路(thetectofugal pahway:视网膜—视顶盖—圆核—外纹体)和离丘脑通路(thethalamofugal pathway:视网膜—丘脑主视核和丘脑背外侧复合体—视厅);而对于禽类来说,离顶盖通路远比离丘脑通路重要的多。在鸡脑离顶盖视觉通路中,光刺激是否引起不同神经元的功能活动;哪些神经元参与某一特定光信息的传递,这些问题还有待于进一步研究。
光线通过鸡的眼睛,刺激视网膜,经视神经叶的视神经途径传到丘脑下部或直接透过颅骨到达松果体及丘脑下部,下丘脑产生和分泌促性腺激素释放激素(Gonadotrophin Releasing Hormone,GnRH),再通过垂体门脉系统传至脑下垂体前叶,引起促卵泡激素(Follicle Stimulating Hormone,FSH)和促黄体激素(LutenizingHormone,LH)的分泌,LH和FSH协同促进卵泡成熟,促进卵巢分泌雌二醇。这些激素共同构成支配生殖机能的内分泌体系,影响生殖器官的发育;促使卵泡生长发育和排卵。但光信息在神经系统内部通过怎样的途径和方式对鸡下丘脑-垂体-性腺轴起调节作用,目前还不清楚。
脑中神经元间的联系错综复杂,利用常规的神经束路追踪标记物方法不能对光信息通路进行完全的描绘,因为其一般不能跨神经元传递。但即早基因c-fos可通过跨突触活化作用诱导表达,在中枢神经系统接受内外刺激后,c-fos的表达产物Fos蛋白可作为单一神经元应答的分辨标记。
c-fos原癌基因是一种即刻早基因(immediately early gene,IEG),耦连细胞外信息与细胞内靶基因的转录,被称为核内“第三信使”(third messenger);普遍存在于神经中枢系统(Central Nervous System,CNS),正常情况下高度保守,表达水平很低,难以检测,在受刺激的情况下可以迅速发生一过性表达。c-fos基因的表达可作为脑对包括光照在内的各种刺激起反应的标志物。疼痛刺激诱导c-fos基因在鸭脊髓的表达作为光诱导c-fos基因在母鸡中脑的表达的免疫组化阳性对照;从而对Fos蛋白的分布、形态有初步的认知。光诱导c-fos基因在母鸡中脑的表达试验:将鸡右眼遮光7天后接受20 1x的光照刺激1.5h,暗适应1.5h后灌流固定,取脑制作石蜡切片,采用免疫组化方法和Western Blot的方法检测c-fos基因在中脑的表达。结果显示:中脑内有大量Fos样免疫反应阳性神经元,主要见于左侧,分布于中脑SGC层、峡核大细胞部(Nucleusisthmi pars magnocellularis,Imc)和小细胞部(Nucleus isthmi pars parvocellularis,Ipc)、峡视核(Nucleus isthmo-opticus,ION)、半月核(Nucleus Semilunaris,SLu)、中脑外侧核背侧部(Nucleus mesencephalicus lateralis pars dorsalis,MLd)、红核(Nucleus raphes,RN)、滑车神经核(Nucleus nervitrochlearis,nⅣ)。中脑两侧峡核的Fos样免疫反应阳性神经元的细胞数密度及Fos样免疫反应阳性神经元所占神经核团细胞数量的百分比差异显著(P＜0.05)。但在未接受光刺激的脑区域内也有少量Fos样免疫反应阳性神经元。上述结果提示:光照信息主要通过视网膜经离顶盖通路传达到母鸡中脑,同时顶盖一峡核回路也起重要作用。非视觉感光系统(non-visual photosensitivesystem,NVPS)及离顶盖通路中光信息的交叉投射造成c-fos基因在未接受光刺激的脑区域中有少量的表达。
The pathway of optical information in hen mesencephalon was investigated using c-fos method, to do some basic work for understanding the mechanism in the effect of light on egg-laying. In poultry production, light is one of the important environmental factors for maintaining the life of the animals. Scientific utilization of light can further enhance chicken's productive performance, for example: promoting chicken's growth and development, regulating sexual maturation, increasing egg production rate, fertility rate, hatching eggs eligible rate and so on. Poultry is one of vertebrate which is the most dependent on the visual system. Their visual system is mainly divided into two parallel processing pathways of visual information: the tectofugal pathway and the thalamofugal pathway. For poultry, the tectofugal pathway is more important than the thalamofugal pathway. In chicken's tectofugal visual pathway, the problems that whether different light stimulation induce functional activities of different neurons or what are the neurons involved in the transmission of a specific light information have not be known yet.
Light stimulates retina and the information is transmitted through optic nerve to hypothalamus or directly permeating skull arrives in conarium. Hypothalamus produces and secretes GnRH, which passing through pituitary portal system to anterior portion of hypophsis, inducing secretion of LH and FSH. LH cooperating with FSH facilitates follicular maturation, promoting ovary to secrete estradiol. These hormones jointly constitute endocrine system dominating procreation enginery and effect genital organs growth. But it is still unclear that in the nervous system, how the light information was adjusted.
In encephalon, connection among neurons is labyrinthine. The conventional method to trace nervous pathway cannot diplay the integrated light pathway, because it generally cannot trace by trans-neuron. But immediate early gene-c-fos can be induced and expressed through trans-synaptic activation. When central nervous system received inside and outside stimulation, the Fos protein of expressive products of c-fos gene can be as a resolving tag of a single neuron response.
c-fos is one of main member of immediate early gene in protooncogene. Coulping extracellular information in intracellular gene transcription, c-fos is described as "third messenger " in cell nucleus. Under normal circumstance, c-fos gene is commonly present in central nervous system, highly conserves and expresses at very low level, while in case of stimulated situation, a transient expression will be rapidly made. The expression of c-fos gene is a sign of response of encephalon receving various stimulations. The expression of c-fos gene induced by ache stimulation in ducks' spinal cord is used as immunoreactive positive picture of the expression of c-fos gene induced by light stimulation in hen mesencephalon, primarily cognizing the distribution and morphology of Fos-like immunoreactive positive neuron in ducks' spinal cord. The expression of c-fos gene was induced by light stimulation in hen mesecephalon: unilateral eyes of hens were kept dark for seven days. Then they accepted light stimulation of 20 lx for 1.5 hours and dark adaptation 1.5 hours. After perfusion with fixative, their brains were removed from the skulls and were made into continuous paraffin sections. Using immunohistochemical method and Western Blot method, expression of c-fos gene in mesencephalon was examined. The result showed that a number of Fos-like immunoreactive positive neurons were observed in mesencephalon, being mainly present in left stratum griseum centrale (SGC) of tectum, isthmi pars principals (Imc, Ipc), nucleus semilunaris (SLu), nucleus isthmo-opticus (ION), nucleus mesencephalicus lateralis (pars dorsalis, MLd), nucleus raphes (RN), and nucleus nervitrochlearis (nIV). And the Fos-like immunoreaction positive neurons were predominantly distributed in Imc and Ipc of the left mesencephalon, and the differentia of the cell density and the percentage chart of Fos-like immunoreactive positive neurons were distinct. But few Fos-like immunoreactive positive neurons were observed in encephalon regions without receiving light stimulation. The results suggested that though the optical information is mainly conveyed to mesencephalon through the tectofugal pathway, the tectoisthmi circus also play an important role in this course. The non-visual photosensitivesystem and light cross-projection in the tectofugal visual pathway cause the few expression of c-fos gene in encephalon regions without receiving light stimulation.
光线通过鸡的眼睛,刺激视网膜,经视神经叶的视神经途径传到丘脑下部或直接透过颅骨到达松果体及丘脑下部,下丘脑产生和分泌促性腺激素释放激素(Gonadotrophin Releasing Hormone,GnRH),再通过垂体门脉系统传至脑下垂体前叶,引起促卵泡激素(Follicle Stimulating Hormone,FSH)和促黄体激素(LutenizingHormone,LH)的分泌,LH和FSH协同促进卵泡成熟,促进卵巢分泌雌二醇。这些激素共同构成支配生殖机能的内分泌体系,影响生殖器官的发育;促使卵泡生长发育和排卵。但光信息在神经系统内部通过怎样的途径和方式对鸡下丘脑-垂体-性腺轴起调节作用,目前还不清楚。
脑中神经元间的联系错综复杂,利用常规的神经束路追踪标记物方法不能对光信息通路进行完全的描绘,因为其一般不能跨神经元传递。但即早基因c-fos可通过跨突触活化作用诱导表达,在中枢神经系统接受内外刺激后,c-fos的表达产物Fos蛋白可作为单一神经元应答的分辨标记。
c-fos原癌基因是一种即刻早基因(immediately early gene,IEG),耦连细胞外信息与细胞内靶基因的转录,被称为核内“第三信使”(third messenger);普遍存在于神经中枢系统(Central Nervous System,CNS),正常情况下高度保守,表达水平很低,难以检测,在受刺激的情况下可以迅速发生一过性表达。c-fos基因的表达可作为脑对包括光照在内的各种刺激起反应的标志物。疼痛刺激诱导c-fos基因在鸭脊髓的表达作为光诱导c-fos基因在母鸡中脑的表达的免疫组化阳性对照;从而对Fos蛋白的分布、形态有初步的认知。光诱导c-fos基因在母鸡中脑的表达试验:将鸡右眼遮光7天后接受20 1x的光照刺激1.5h,暗适应1.5h后灌流固定,取脑制作石蜡切片,采用免疫组化方法和Western Blot的方法检测c-fos基因在中脑的表达。结果显示:中脑内有大量Fos样免疫反应阳性神经元,主要见于左侧,分布于中脑SGC层、峡核大细胞部(Nucleusisthmi pars magnocellularis,Imc)和小细胞部(Nucleus isthmi pars parvocellularis,Ipc)、峡视核(Nucleus isthmo-opticus,ION)、半月核(Nucleus Semilunaris,SLu)、中脑外侧核背侧部(Nucleus mesencephalicus lateralis pars dorsalis,MLd)、红核(Nucleus raphes,RN)、滑车神经核(Nucleus nervitrochlearis,nⅣ)。中脑两侧峡核的Fos样免疫反应阳性神经元的细胞数密度及Fos样免疫反应阳性神经元所占神经核团细胞数量的百分比差异显著(P＜0.05)。但在未接受光刺激的脑区域内也有少量Fos样免疫反应阳性神经元。上述结果提示:光照信息主要通过视网膜经离顶盖通路传达到母鸡中脑,同时顶盖一峡核回路也起重要作用。非视觉感光系统(non-visual photosensitivesystem,NVPS)及离顶盖通路中光信息的交叉投射造成c-fos基因在未接受光刺激的脑区域中有少量的表达。
The pathway of optical information in hen mesencephalon was investigated using c-fos method, to do some basic work for understanding the mechanism in the effect of light on egg-laying. In poultry production, light is one of the important environmental factors for maintaining the life of the animals. Scientific utilization of light can further enhance chicken's productive performance, for example: promoting chicken's growth and development, regulating sexual maturation, increasing egg production rate, fertility rate, hatching eggs eligible rate and so on. Poultry is one of vertebrate which is the most dependent on the visual system. Their visual system is mainly divided into two parallel processing pathways of visual information: the tectofugal pathway and the thalamofugal pathway. For poultry, the tectofugal pathway is more important than the thalamofugal pathway. In chicken's tectofugal visual pathway, the problems that whether different light stimulation induce functional activities of different neurons or what are the neurons involved in the transmission of a specific light information have not be known yet.
Light stimulates retina and the information is transmitted through optic nerve to hypothalamus or directly permeating skull arrives in conarium. Hypothalamus produces and secretes GnRH, which passing through pituitary portal system to anterior portion of hypophsis, inducing secretion of LH and FSH. LH cooperating with FSH facilitates follicular maturation, promoting ovary to secrete estradiol. These hormones jointly constitute endocrine system dominating procreation enginery and effect genital organs growth. But it is still unclear that in the nervous system, how the light information was adjusted.
In encephalon, connection among neurons is labyrinthine. The conventional method to trace nervous pathway cannot diplay the integrated light pathway, because it generally cannot trace by trans-neuron. But immediate early gene-c-fos can be induced and expressed through trans-synaptic activation. When central nervous system received inside and outside stimulation, the Fos protein of expressive products of c-fos gene can be as a resolving tag of a single neuron response.
c-fos is one of main member of immediate early gene in protooncogene. Coulping extracellular information in intracellular gene transcription, c-fos is described as "third messenger " in cell nucleus. Under normal circumstance, c-fos gene is commonly present in central nervous system, highly conserves and expresses at very low level, while in case of stimulated situation, a transient expression will be rapidly made. The expression of c-fos gene is a sign of response of encephalon receving various stimulations. The expression of c-fos gene induced by ache stimulation in ducks' spinal cord is used as immunoreactive positive picture of the expression of c-fos gene induced by light stimulation in hen mesencephalon, primarily cognizing the distribution and morphology of Fos-like immunoreactive positive neuron in ducks' spinal cord. The expression of c-fos gene was induced by light stimulation in hen mesecephalon: unilateral eyes of hens were kept dark for seven days. Then they accepted light stimulation of 20 lx for 1.5 hours and dark adaptation 1.5 hours. After perfusion with fixative, their brains were removed from the skulls and were made into continuous paraffin sections. Using immunohistochemical method and Western Blot method, expression of c-fos gene in mesencephalon was examined. The result showed that a number of Fos-like immunoreactive positive neurons were observed in mesencephalon, being mainly present in left stratum griseum centrale (SGC) of tectum, isthmi pars principals (Imc, Ipc), nucleus semilunaris (SLu), nucleus isthmo-opticus (ION), nucleus mesencephalicus lateralis (pars dorsalis, MLd), nucleus raphes (RN), and nucleus nervitrochlearis (nIV). And the Fos-like immunoreaction positive neurons were predominantly distributed in Imc and Ipc of the left mesencephalon, and the differentia of the cell density and the percentage chart of Fos-like immunoreactive positive neurons were distinct. But few Fos-like immunoreactive positive neurons were observed in encephalon regions without receiving light stimulation. The results suggested that though the optical information is mainly conveyed to mesencephalon through the tectofugal pathway, the tectoisthmi circus also play an important role in this course. The non-visual photosensitivesystem and light cross-projection in the tectofugal visual pathway cause the few expression of c-fos gene in encephalon regions without receiving light stimulation.
引文
[1]张金月,王宜奇,杜艳芬.科学光照提高蛋鸡效益[J].中国家禽,2001,23(3).
[2]戴四发,朱夕婷,黎观红.蛋鸡舍光照的控制与应用[J].饲养管理,当代畜牧,2003,5.
[3]Shimizu T,Karten H J.The avian visual system and the evolution of the neocortex[J].Vision,Brain and Behavior in Birds(Zeigler H P &.Bischof H J ed),MIT Press,Cambridge,MA,1993.
[4]Hu M,Naito J,Chen Y,Ohmori Y,Fukuta K.Afferent and efferent connections of the nucleus rotundus demonstrated by WGA-HRP in the chick.Anat Histol Embryol,2003,32(6):335-340.
[5]Karten HJ,Cox K,Mpodozis J.Two distinct populations of tectal neurons have unique connections within the retinotectorotundal pathway of the pigeon(Columba livia)[J].Comp Neurol,1997,387(3):449-465.
[6]Zeigler,Bischof.Vision,Brain and Behavior in Birds[M].Cambridge,Massachusetts:The MIT Press,Massachusettes Institute of Technology,1993:99-169.
[7]雷秋霞,李善立,杨艳.鸡性成熟研究进展[J].家禽科学,2005,11:45-46.
[8]王光瑛,王寿昆.生殖激素对家禽就巢的调控[J].福建农林大学学报(自然科学版),2005,34(1):82-86.
[9]Muller R.,Tremblay J.M.,Adamson E.D.,Verma I.M.Tissue and cell type specific expression of two human c-onc gene.Nature,1983,304:454-456.
[10]Muller R.,Muller D.,Guilbert L.Differential expression of c-fos in hematopoietic cells:correlation with differentiation of monomyelocytic cells in vitro.EMBO J.1984,3:1887-1890.
[11]Morgan J.I,Curran T.Stimulus--transcription coupling in the nervous system:In-volvement of the inducible protooncogenes fos and jun.Annu.Rev.Neurosci.1991,14:421-51.
[12]Morgan JI,Curran T.Immediate-early genes:ten years on.Trends Nenrosci.1995,18(2):66-67.
[13]Twu JS,Schloemer RH.Transcriptional trans-activating function of hepatitis B virus[J].Virol 1987,61:3448-3453.
[14]Spandau DF,Lee CH.Trans-activation of viral enhancers by the hepatitis B virus X protein [J].Virol 1988,62:427-434.
[15]Zahm P,Hofschneider PH,Koshy R.The HBV X-ORF encodes a transactivator:a potential factor in viral hepatocarcinogenesis[J].Oncogene 1988,3:169-177.
[16]Seto E,Mitchell PJ,Yen TS.Transactivation by the hepatitis B vires X protein depends on AP-2and other transcription factors[J].Nature 1990,344:72-74.
[17]Twu JS,Lai MY,Chen DS,Robinson WS.Activation of protooncogene c-jun by the X protein of hepatitis B virus[J].Virology 1993,192:346-350.
[18]Kekule AS,Lauer U,Weiss L,Luber B,Hofschneider PH.Hepatitis B virus transactivator HBx uses a tumour promoter signalling pathway[J].Nature 1993,361:742-745.
[19]Natoli G,Avantaggiati ML,Chirillo P,Costanzo A,Artini M,Balsano C,Levrero M.Induction of the DNA-binding activity of c-jun/c-fos heterodimers by the hepatitis B virus transactivator pX[J]. Moi Cell Biol 1994,14:989-998.
[20]刘智平.c-fos基因在中枢神经功能网络系统及离体神经元的表达与机理研究[D].中国协和医科大学,1993.
[21]Morgan J.,Cutran T.Stimulus-transcription coupling in neurons:role of cellular immediate-early genes.TINS.1989,12:459-462.
[22]韩济生.神经科学纲要[M].北京:北京医科大学、中国协和医科大学联合出版社,1993:274-278,483.
[23]HEISE C E,MITROFANIS J.Fos immunoreactivity in some locomotor neural centres of GOHDA-lesioned rats[J].Anat Embryol,2006,211(6):659-671.
[24]周清,陈剑,刘珏,等.视觉刺激诱导c-Fos在弱视成年大鼠视皮质神经元表达的研究[J].中国病理生理杂志,2007,23(6):1228-1229,1232.
[25]KAZI J A,GEE C F.Effect of gabapentin on c-Fos expression in the CNS after paw surgery in rats [J].Neurosci,2007,32(3):228-34.
[26]MURPHY M,GREFERATH U,WlLISON Y M.A method for detecting functional activity related expression in gross brain regions,specific brain nuclei and individual neuronal cell bodies and their projections[J].Biol Proced Online.2007,9:1-8.
[27]PUSKAS L,PUSKAS N,BABOVIC S S,et al.C-fos protein expression in the parietal cortex and olfactory tubercle in the hypoxic rat brain[J].Med Preql,2007,60(3-4):128-133.
[28]Bell Dj,BM Freeman.Physiology and Biochemistry of the Domestic Fowl.London;Academic Pr,1971,2:651-661.
[29]都格尔斯仁,刘为民,陈建新,郭和以鸡中脑向小脑的直接投射-HRP法研究①内蒙古农牧学院学报 1995,16(2).
[30]田九畴 主编.畜禽神经解剖学[M],1998:394-397.
[31]Wang S R.The nucleus isthmi and dual modulation of the receptive field of tectal neurons in non-mammals[J].Brain Research Review,2003,41(1):13-25.
[32]肖泉.峡视核-研究中枢神经系统发育及调亡的新模型[J].生物化学与生物物理进展,2000,27(2):119-121.
[33]Wang S R,Wang Y C,Frost B J.Magnocellular and parvocellular divisions of pigeon nucleus isthmi differentially modulate visual responses in the tectum[J].Exp Brain Res,1995,104:376-384.
[34]范海燕,魏熹,于晓艳.金翅中脑峡核的组织学研究[J].阜阳师范学院学报(自然科学版),2007,24(2):39-40.
[35]Wang S R.The nucleus isthmi and dualmodulation of the recetive field of tectal neurons in non-mammals[J].Brain Research Review,2003,41:13-25.
[36]Wang Y,Xiao J,Wang S R.Excitatory and inhibitory receptive fields of tectal cells are differentially modified by magnocellular and parvocellular divisions of the pigeon nucleus isthmi[J].Comp Physio 1A,2000,186:505-511.
[37]Wang Y C,Frost B J.Visual response characteristics of neurons in the nucleus isthmi parvocellularis of pigeons[J].Exp Brain Res,1991,87:624-633.
[38]Yan K,Wang SR.Visual responses of neurons in the avain nucleus isthmi Neurosci Lett.1986,64:340-344.
[39]BURKHARD HELLMAN,MART1NA MANNA.Nucleus Isthmi,Pars Semilunaris as a Key Component of the Tectofugal Visual System in Pigeons[J].The Journal of Comparative Neurology,2001,436:153-16.
[40]PANDA S,SATO T K,CASTRUCCI A M,et al.Melanopsin(Opn4)requirement for normal light-induced circadian phase shifting[J].Science,2002,298(5601):2213-2216.
[41]RUBY N F,BRENNAN T J,XIE X,et al.Role of melanopsin in circadian responses to light[J].Science,2002,298(5601):2211-2213.
[42]熊鲲,黄菊芳,罗学港.非视觉感光系统的研究进展[J].解剖学杂志,2005,28(3):358-360.
[43]额尔敦木图,陈耀星,王子旭,等.单色光对蛋鸡产蛋性能的影响[J].畜牧兽医学报,2007,38(1):40-45.
[44]Simone L.Meddlel and Brian K.FoileR.Photoperiodically Driven Changes in Fos Expression within the Basal Tuberal Hypothalamus and Median Eminence of Japanese Quail[J].Neuroscience,1997,17(22):8909-8918.
[45]战辉,程义勇.重新认识神经胶质细胞[J].生理科学进展,2003,34(3):231-234.
[46]Mikula M,Gotzmann J,Fischer AN,et al.The proto..oncopro-tein c-Fos negatively regulates hepatoeellular tum origene-sis[J].Oneogene,2003,22(43):6725-6738.
[47]Munglani R,Hunt SP.Molecular biology of pain[J].Br JAnaesth,1995,75(2):186-192.
[48]Morgan S,Greenberg ME.The regulation and function of c-fos and other IEGS in the neurous system[J].Neuron,1990,4(2):477-485.
[49]刁文卓,韩影,罗毅男,等,颅脑损伤后c-fos基因表达研究的进展,中国实验临床免疫学杂志,1999,11(6):131-134.
[50]Matsuo H,Schally AVI.Structure of porcine LH- and FSH-releasing hormone I,the proposed amino acid sequence[J].BioeheBiophys Res Commun,1971,43:1334-1339.
[51]Jimenez-Linan M,Rubin BS,King JC.Examination of guinea pig luteinizing hormone-releasing hormone gene reveals a unique decapeptide and existence of two transcripts in the brain.Endocrinology,2001,138:4123-4130.
[52]王强,侯正录,权富.GnRH的研究进展生物学杂志[J].2003,20(5):40-41.
[53]Naor Z,Shacham S.Signal transduction of the GnRH recepter:crosstalk of calcium protein kinase C(PKC)and arachidonic acid[J].Cell MolNeurobiol,1995,15:527-544.
[54]Dubois E A,Zandbergen M A,Peute J,Goos H J.Evolutionary development of three gonadotropin-releasing hormone(GnRH)systems in vertebrates.Brain Res Bull,2002,57:413-418.
[55]孙刚.胎盘内分泌的基础与临床.上海:第二军医大学出版社,2001,110-111.
[56]King J A,Millar R P.Evolution of gonadotropin-releasing hormones.Trends Endocrinol Metab,1992,3:339-346.
[57]Johnson P A,Wang S Y,Brooks C.Characterization of a source and levels of plasma immunoreactive inhibin during the ovulatory cycle of the domestic hen[J].Biol Reprod,1993,48(2):262-267.
[58]李莹辉,汪琳仙,杨传任.催乳素对促性腺激素诱导的鸡卵泡膜细胞类固醇激素合成的抑制作用.中国兽医学报,1996,16(6):598-602.
[59]詹慧琴.鸡产蛋相关基因的SNPs检测及其与蛋用性能关系的研究[D].华中农业大学,2006.
[60]Gorbman A,Sower S A.Evolution of the role of GnRH in animal(Metazoan)biology.Gen Comp Endocrinal,2003,134(3):207-21.
[61]Brooks J,Co leman D L,Cohen S L.Regulation of gonadotrophin releasing hormone receptor expression in the ewe.Anim Reprod Sci,1996,42:89-98.
[62]叶丹,潘建伟,廖鸣娟,张志和,朱睦元.促性腺激素释放激素的结构及其生物学功能[J].生物化学与生物物理进展,2003,30(1):49-53.
[63]郭晓娜,朱永义,朱科学.生物体内γ-氨基丁酸的研究[J].氨基酸和生物资源,2003,25(2):70-72.
[64]Krajnc D,Neff NH,HadjiconstantiouM.Glutamate,glutamine and glutamine synthetase in the neonatal rat brain following hypoxia[J].Brain Res,1996,707(10):134-137.
[65]包华琼,王新庄.γ-氨基丁酸(GABA)的生殖生理作用[J].动物医学进展,2002,23(3):39-40.
[66]杨庆武.二次脑损伤后c-fos基因表达与β-内啡肽的变化及意义[D].福建医科大学,2003.
[67]张世仪,张敏磊,付建军.大鼠尾状核内注射μ或δ型阿片受体激动剂对操作式条件反射的影响[J].基础医学与临床,1993,13(5):28-31.
[68]田野.运动生理学高级教程[M].北京:高等教育出版社,2003,48-223.
[69]Said S I,Mutt V.Polypeptide with broad biological activity isolated from small intestin[J].Science,1970,169:1217.
[70]王华,刘双珍,王平宝.血管活性肠肽及其在眼科学的研究进展[J].国际眼科杂志,2004,4(6):1095-1098.
[71]徐伟,赵亚华,孔洁.血管活性肠肽研究进展[J].药物生物技术,2002,9(6):364-368.
[72]Pitts GR,Youngren OM,Silsby JL,et al.Role of vasoactive intestinal peptide in the control of prolactin-induced turkey incubation behavior.Ⅰ.Acute infusion of vasoactive intestinal peptide[J].Biol Reprod,1994,50(6):1344-1349.
[73]Halawani ME,Silsby JL,Mauro LJ.Vasoactive intestinal peptide is a hypothalamic prolactin-releasing neuropeptide in the turkey(Meleagris gallopavo)[J].Gen Comp Endocrinol,1990,78(1):66-73.
[74]Samson WK,Said SI,Snyder G,et al.Invitro stimulation of prolactin release by vasoactive intestinal peptide[J].Peptides,1980,1(4):325-327.
[75]Sharp P J,Sterling R J,Talbot R T,et al.The role of hypothalamic vasoactive intestinal polypeptide in the maintenance of prolactin secretion in incubating bantam hems:observations using passive immunization,radioimmunoassay and immunohistochemistry[J].Endocrinol,1989,122(1):5-13.
[76]Aronin N,Sagar SM,Sharp FR,Schwartz WJ.Light regulates expression of a Fos-related protein in the rat SCN.Proc Natl Acad Sci USA,1990,87(15):5959-5962.
[77]Aste N,Viglietti-Panzica C,Fasolo A,et al.Mapping of neurochemical markers in quail central nervous system:VIP- and SP-like immunoreactivity[J].J Chem Neuroanat,1995,8(2):87-102.
[78]Kriegsfeld LJ,Silver R,Gore AC,et al.Vasoactive intestinal polypeptide contacts on gonadotropin-releasing hormone neurons increase following puberty in female rats[J].Neuroendocrinol,2002,14(9):685-690.
[79]Piggins HD,Cutler DJ.The roles of vasoactive intestinal polypeptide in the mammalian circadian clock[J].Endocrinol,2003,177(1):7-15.
[80]Aton SJ,Colwell CS,Harmar AJ,et al.Vasoactive intestinal polypeptide mediates circadian rhythmicity and synchrony in mammalian clock neurons[J].Nat Neurosci,2005,8(4):476-483.
[81]Yarnada S,Mikami S,Yanaihara N.Immunohisto-chemical localization of vasoactive intestinal polypeptide(VIP)-containing neurons in the hy-pothalamus of the Japanese quailCoturnix coturnix[J].Cell Tissue Res,1982,226:13-26.
[82]Mikami S,Yamada S.Immunohistochemistry of the hypothalamic neuropeptides and anterior pituitary cells in the Japanese quail[J].J Exp Zool,1984,232:405-417.
[83]王荫亭,许红艳,王书荣.家鸽血管活性肠肽阳性神经元在顶盖和峡核中的分布及形态特征[J].生物物理学报,1993,9(3):447-452.
[84]王荫亭,许红艳,王书荣.血管活性肠肽免疫阳性神经元在家鸽中脑和间脑中的分布[J].生物物理学报,1994,10(1):103-110.
[85]左明雪,李锐.血管活性肠肽在环鸽旁听觉神经通路中的分布[J].解剖学报,1998,29(2):135-138.
[86]蒋书东,周天红,张丽霞.血管活性肠肽在皖南白鹅中脑和脑桥内的分布[J].中国兽医科技,2004,34(1]):59-61.
[87]Gerhold LM,Wise PM.Vasoactive intestinal polypeptide regulates dynamic changes in astrocyte morphometry:Impact on gonadotropin releasing hormone neurons[J].Endocrinology,2006,Feb 9.
[88]Oelschlager HH,Nakamura M,Herzog M,Burda H.Visual system labeled by c-Fos immunohistochemistry after light exposure in the 'blind' subterranean Zambian mole-rat (Cryptomys anselli).Brain Behav.Evo.1,2000,55(4):209-220.
[89]李锦新,马宁芳.光刺激移植视网膜诱导大鼠上丘FOS的表达.中国组织化学与细胞化学杂志,2003,12(1):42-45.
[90]方富贵,章孝荣.P物质研究进展[J].动物医学进展,2005,26(1):6-8.
[91]Battmann T,Parsadaniantz S M,Jeanjean B,et al.In-vivo inhibi-tion of the preovulatoryLHsurge by substance P and in-vitromodula-tion of gonadotrophin-releasing hormone-induced LH release by sub-stance P.oestradiol and progesterone in the female rat[J].Journal of Endocrinology,1991,130:169-175.
[92]Duval P,Lenoir V,Garret C,et al.Reduction of the amplitude of preovalatory LH and FSH surges and of the amplitude of the in vitro GnRH-induced LHrelease by substance P.Reversal of the effect by RP67580[J].Neuropharmacology,1996,35(12):1805-1810.
[93]姜恩魁,李志山,常志杰等.大鼠性周期中下丘脑和垂体P物质含量的变化[J].锦州医学院学报,1987,8(4):315-317.
[94]姜恩魁,李志山,常志杰等.大白鼠垂体P物质含量与卵巢孕酮生成关系的研究[J].锦州医学院学报,1988,9(2):108-109.
[95]Jessop D S,Chowdrey H S,Larsen P J,et al.Substance P:multi-functional peptide in the hypothalamo-pituitary system[J].Journal ofEndocrinology,1992,132:331-337.
[96]唐宗湘,马殷华.鸟类视觉系统的离顶盖通路[J].广西师范大学学报(自然科学版),2004,22(4):79-82.
[97]WANG Y,XIAO J,WANG S R.Excitatory and inhibitory receptive fields of tectal cells are differentially modified by magnocellular and parvocellular divisions of the pigeon nucleus isthmi[J].J Comp Physio[A],2000,186:505-511.
[98]HELLMANN B,MANNS M,GUNTURKUN O.Nucleus isthmi,pars sere ilunaris as a key component of the tectofugal visual system in pigeons[J].Comp Neuro 1,2001,436:153-166.
[99]赵丽杰.鸡中脑视顶盖Ⅰ层细胞构筑及几种神经肽在相关神经核团中的分布研究[D],河北农业大学,2007.
[100]刘佳.雏鸡中脑视顶盖SGC层细胞构筑及几种神经肽在OT-Rt通路的定位研究[D],河北农业大学,2007.
[101]Larsen P J,SaermarkT,Mau S E.Bindingof an iodinated substance P analogue to cultured anterior pituitary prolactin and luteinizing hor-mone-containing cells[J].The Journal of Histochemistry and Cyto-chemistry,1992,40(4):487-493.
[102]Hajihira S et al.Brain Res,1997,751:124.
[103]Presley RW et al.[J].Neurosci,1990,10:23.
[104]Lupi D,Sekaran S,Jones SL,Hankins MW,Foster RG.Light-evoked FOS induction within the suprachiasmatic nuclei(SCN)of melanopsin knockout(Opn4-/-)mice:a developmental study.Chronobiol Int.2006,23(1-2):167-17.
[105]Hattar S,Lucas RJ,Mrosovsky N,et al.Melanopsin and rodcone photoreceptive systems account for all major accessory visual functions in mice[J].Nature,2003,424(6944):76-81.
[106]Panda S,Provencio I,Tu DC,et al.Melanopsin is required for non-image-forming photic responses in blind mice[J].Science,2003,301(5632):525-527.
[107]Benson DM.Strange vision:ganglion cells as circadian photoreceptors[J].Trends Neurosci,2003,26(6):314-320.
[108]Sancar A.Cryptoehrome:the second photoactive pigment in the eye and its role in circadian photoreception[J].Annu Rev Biochem,2000,69:31-67.
[109]Russell N,Van Gelder,Wee R,et al.Reduced pupillary light responses in mice lacking cryptochromes[J].Science,2003,299(5604):222.
[2]戴四发,朱夕婷,黎观红.蛋鸡舍光照的控制与应用[J].饲养管理,当代畜牧,2003,5.
[3]Shimizu T,Karten H J.The avian visual system and the evolution of the neocortex[J].Vision,Brain and Behavior in Birds(Zeigler H P &.Bischof H J ed),MIT Press,Cambridge,MA,1993.
[4]Hu M,Naito J,Chen Y,Ohmori Y,Fukuta K.Afferent and efferent connections of the nucleus rotundus demonstrated by WGA-HRP in the chick.Anat Histol Embryol,2003,32(6):335-340.
[5]Karten HJ,Cox K,Mpodozis J.Two distinct populations of tectal neurons have unique connections within the retinotectorotundal pathway of the pigeon(Columba livia)[J].Comp Neurol,1997,387(3):449-465.
[6]Zeigler,Bischof.Vision,Brain and Behavior in Birds[M].Cambridge,Massachusetts:The MIT Press,Massachusettes Institute of Technology,1993:99-169.
[7]雷秋霞,李善立,杨艳.鸡性成熟研究进展[J].家禽科学,2005,11:45-46.
[8]王光瑛,王寿昆.生殖激素对家禽就巢的调控[J].福建农林大学学报(自然科学版),2005,34(1):82-86.
[9]Muller R.,Tremblay J.M.,Adamson E.D.,Verma I.M.Tissue and cell type specific expression of two human c-onc gene.Nature,1983,304:454-456.
[10]Muller R.,Muller D.,Guilbert L.Differential expression of c-fos in hematopoietic cells:correlation with differentiation of monomyelocytic cells in vitro.EMBO J.1984,3:1887-1890.
[11]Morgan J.I,Curran T.Stimulus--transcription coupling in the nervous system:In-volvement of the inducible protooncogenes fos and jun.Annu.Rev.Neurosci.1991,14:421-51.
[12]Morgan JI,Curran T.Immediate-early genes:ten years on.Trends Nenrosci.1995,18(2):66-67.
[13]Twu JS,Schloemer RH.Transcriptional trans-activating function of hepatitis B virus[J].Virol 1987,61:3448-3453.
[14]Spandau DF,Lee CH.Trans-activation of viral enhancers by the hepatitis B virus X protein [J].Virol 1988,62:427-434.
[15]Zahm P,Hofschneider PH,Koshy R.The HBV X-ORF encodes a transactivator:a potential factor in viral hepatocarcinogenesis[J].Oncogene 1988,3:169-177.
[16]Seto E,Mitchell PJ,Yen TS.Transactivation by the hepatitis B vires X protein depends on AP-2and other transcription factors[J].Nature 1990,344:72-74.
[17]Twu JS,Lai MY,Chen DS,Robinson WS.Activation of protooncogene c-jun by the X protein of hepatitis B virus[J].Virology 1993,192:346-350.
[18]Kekule AS,Lauer U,Weiss L,Luber B,Hofschneider PH.Hepatitis B virus transactivator HBx uses a tumour promoter signalling pathway[J].Nature 1993,361:742-745.
[19]Natoli G,Avantaggiati ML,Chirillo P,Costanzo A,Artini M,Balsano C,Levrero M.Induction of the DNA-binding activity of c-jun/c-fos heterodimers by the hepatitis B virus transactivator pX[J]. Moi Cell Biol 1994,14:989-998.
[20]刘智平.c-fos基因在中枢神经功能网络系统及离体神经元的表达与机理研究[D].中国协和医科大学,1993.
[21]Morgan J.,Cutran T.Stimulus-transcription coupling in neurons:role of cellular immediate-early genes.TINS.1989,12:459-462.
[22]韩济生.神经科学纲要[M].北京:北京医科大学、中国协和医科大学联合出版社,1993:274-278,483.
[23]HEISE C E,MITROFANIS J.Fos immunoreactivity in some locomotor neural centres of GOHDA-lesioned rats[J].Anat Embryol,2006,211(6):659-671.
[24]周清,陈剑,刘珏,等.视觉刺激诱导c-Fos在弱视成年大鼠视皮质神经元表达的研究[J].中国病理生理杂志,2007,23(6):1228-1229,1232.
[25]KAZI J A,GEE C F.Effect of gabapentin on c-Fos expression in the CNS after paw surgery in rats [J].Neurosci,2007,32(3):228-34.
[26]MURPHY M,GREFERATH U,WlLISON Y M.A method for detecting functional activity related expression in gross brain regions,specific brain nuclei and individual neuronal cell bodies and their projections[J].Biol Proced Online.2007,9:1-8.
[27]PUSKAS L,PUSKAS N,BABOVIC S S,et al.C-fos protein expression in the parietal cortex and olfactory tubercle in the hypoxic rat brain[J].Med Preql,2007,60(3-4):128-133.
[28]Bell Dj,BM Freeman.Physiology and Biochemistry of the Domestic Fowl.London;Academic Pr,1971,2:651-661.
[29]都格尔斯仁,刘为民,陈建新,郭和以鸡中脑向小脑的直接投射-HRP法研究①内蒙古农牧学院学报 1995,16(2).
[30]田九畴 主编.畜禽神经解剖学[M],1998:394-397.
[31]Wang S R.The nucleus isthmi and dual modulation of the receptive field of tectal neurons in non-mammals[J].Brain Research Review,2003,41(1):13-25.
[32]肖泉.峡视核-研究中枢神经系统发育及调亡的新模型[J].生物化学与生物物理进展,2000,27(2):119-121.
[33]Wang S R,Wang Y C,Frost B J.Magnocellular and parvocellular divisions of pigeon nucleus isthmi differentially modulate visual responses in the tectum[J].Exp Brain Res,1995,104:376-384.
[34]范海燕,魏熹,于晓艳.金翅中脑峡核的组织学研究[J].阜阳师范学院学报(自然科学版),2007,24(2):39-40.
[35]Wang S R.The nucleus isthmi and dualmodulation of the recetive field of tectal neurons in non-mammals[J].Brain Research Review,2003,41:13-25.
[36]Wang Y,Xiao J,Wang S R.Excitatory and inhibitory receptive fields of tectal cells are differentially modified by magnocellular and parvocellular divisions of the pigeon nucleus isthmi[J].Comp Physio 1A,2000,186:505-511.
[37]Wang Y C,Frost B J.Visual response characteristics of neurons in the nucleus isthmi parvocellularis of pigeons[J].Exp Brain Res,1991,87:624-633.
[38]Yan K,Wang SR.Visual responses of neurons in the avain nucleus isthmi Neurosci Lett.1986,64:340-344.
[39]BURKHARD HELLMAN,MART1NA MANNA.Nucleus Isthmi,Pars Semilunaris as a Key Component of the Tectofugal Visual System in Pigeons[J].The Journal of Comparative Neurology,2001,436:153-16.
[40]PANDA S,SATO T K,CASTRUCCI A M,et al.Melanopsin(Opn4)requirement for normal light-induced circadian phase shifting[J].Science,2002,298(5601):2213-2216.
[41]RUBY N F,BRENNAN T J,XIE X,et al.Role of melanopsin in circadian responses to light[J].Science,2002,298(5601):2211-2213.
[42]熊鲲,黄菊芳,罗学港.非视觉感光系统的研究进展[J].解剖学杂志,2005,28(3):358-360.
[43]额尔敦木图,陈耀星,王子旭,等.单色光对蛋鸡产蛋性能的影响[J].畜牧兽医学报,2007,38(1):40-45.
[44]Simone L.Meddlel and Brian K.FoileR.Photoperiodically Driven Changes in Fos Expression within the Basal Tuberal Hypothalamus and Median Eminence of Japanese Quail[J].Neuroscience,1997,17(22):8909-8918.
[45]战辉,程义勇.重新认识神经胶质细胞[J].生理科学进展,2003,34(3):231-234.
[46]Mikula M,Gotzmann J,Fischer AN,et al.The proto..oncopro-tein c-Fos negatively regulates hepatoeellular tum origene-sis[J].Oneogene,2003,22(43):6725-6738.
[47]Munglani R,Hunt SP.Molecular biology of pain[J].Br JAnaesth,1995,75(2):186-192.
[48]Morgan S,Greenberg ME.The regulation and function of c-fos and other IEGS in the neurous system[J].Neuron,1990,4(2):477-485.
[49]刁文卓,韩影,罗毅男,等,颅脑损伤后c-fos基因表达研究的进展,中国实验临床免疫学杂志,1999,11(6):131-134.
[50]Matsuo H,Schally AVI.Structure of porcine LH- and FSH-releasing hormone I,the proposed amino acid sequence[J].BioeheBiophys Res Commun,1971,43:1334-1339.
[51]Jimenez-Linan M,Rubin BS,King JC.Examination of guinea pig luteinizing hormone-releasing hormone gene reveals a unique decapeptide and existence of two transcripts in the brain.Endocrinology,2001,138:4123-4130.
[52]王强,侯正录,权富.GnRH的研究进展生物学杂志[J].2003,20(5):40-41.
[53]Naor Z,Shacham S.Signal transduction of the GnRH recepter:crosstalk of calcium protein kinase C(PKC)and arachidonic acid[J].Cell MolNeurobiol,1995,15:527-544.
[54]Dubois E A,Zandbergen M A,Peute J,Goos H J.Evolutionary development of three gonadotropin-releasing hormone(GnRH)systems in vertebrates.Brain Res Bull,2002,57:413-418.
[55]孙刚.胎盘内分泌的基础与临床.上海:第二军医大学出版社,2001,110-111.
[56]King J A,Millar R P.Evolution of gonadotropin-releasing hormones.Trends Endocrinol Metab,1992,3:339-346.
[57]Johnson P A,Wang S Y,Brooks C.Characterization of a source and levels of plasma immunoreactive inhibin during the ovulatory cycle of the domestic hen[J].Biol Reprod,1993,48(2):262-267.
[58]李莹辉,汪琳仙,杨传任.催乳素对促性腺激素诱导的鸡卵泡膜细胞类固醇激素合成的抑制作用.中国兽医学报,1996,16(6):598-602.
[59]詹慧琴.鸡产蛋相关基因的SNPs检测及其与蛋用性能关系的研究[D].华中农业大学,2006.
[60]Gorbman A,Sower S A.Evolution of the role of GnRH in animal(Metazoan)biology.Gen Comp Endocrinal,2003,134(3):207-21.
[61]Brooks J,Co leman D L,Cohen S L.Regulation of gonadotrophin releasing hormone receptor expression in the ewe.Anim Reprod Sci,1996,42:89-98.
[62]叶丹,潘建伟,廖鸣娟,张志和,朱睦元.促性腺激素释放激素的结构及其生物学功能[J].生物化学与生物物理进展,2003,30(1):49-53.
[63]郭晓娜,朱永义,朱科学.生物体内γ-氨基丁酸的研究[J].氨基酸和生物资源,2003,25(2):70-72.
[64]Krajnc D,Neff NH,HadjiconstantiouM.Glutamate,glutamine and glutamine synthetase in the neonatal rat brain following hypoxia[J].Brain Res,1996,707(10):134-137.
[65]包华琼,王新庄.γ-氨基丁酸(GABA)的生殖生理作用[J].动物医学进展,2002,23(3):39-40.
[66]杨庆武.二次脑损伤后c-fos基因表达与β-内啡肽的变化及意义[D].福建医科大学,2003.
[67]张世仪,张敏磊,付建军.大鼠尾状核内注射μ或δ型阿片受体激动剂对操作式条件反射的影响[J].基础医学与临床,1993,13(5):28-31.
[68]田野.运动生理学高级教程[M].北京:高等教育出版社,2003,48-223.
[69]Said S I,Mutt V.Polypeptide with broad biological activity isolated from small intestin[J].Science,1970,169:1217.
[70]王华,刘双珍,王平宝.血管活性肠肽及其在眼科学的研究进展[J].国际眼科杂志,2004,4(6):1095-1098.
[71]徐伟,赵亚华,孔洁.血管活性肠肽研究进展[J].药物生物技术,2002,9(6):364-368.
[72]Pitts GR,Youngren OM,Silsby JL,et al.Role of vasoactive intestinal peptide in the control of prolactin-induced turkey incubation behavior.Ⅰ.Acute infusion of vasoactive intestinal peptide[J].Biol Reprod,1994,50(6):1344-1349.
[73]Halawani ME,Silsby JL,Mauro LJ.Vasoactive intestinal peptide is a hypothalamic prolactin-releasing neuropeptide in the turkey(Meleagris gallopavo)[J].Gen Comp Endocrinol,1990,78(1):66-73.
[74]Samson WK,Said SI,Snyder G,et al.Invitro stimulation of prolactin release by vasoactive intestinal peptide[J].Peptides,1980,1(4):325-327.
[75]Sharp P J,Sterling R J,Talbot R T,et al.The role of hypothalamic vasoactive intestinal polypeptide in the maintenance of prolactin secretion in incubating bantam hems:observations using passive immunization,radioimmunoassay and immunohistochemistry[J].Endocrinol,1989,122(1):5-13.
[76]Aronin N,Sagar SM,Sharp FR,Schwartz WJ.Light regulates expression of a Fos-related protein in the rat SCN.Proc Natl Acad Sci USA,1990,87(15):5959-5962.
[77]Aste N,Viglietti-Panzica C,Fasolo A,et al.Mapping of neurochemical markers in quail central nervous system:VIP- and SP-like immunoreactivity[J].J Chem Neuroanat,1995,8(2):87-102.
[78]Kriegsfeld LJ,Silver R,Gore AC,et al.Vasoactive intestinal polypeptide contacts on gonadotropin-releasing hormone neurons increase following puberty in female rats[J].Neuroendocrinol,2002,14(9):685-690.
[79]Piggins HD,Cutler DJ.The roles of vasoactive intestinal polypeptide in the mammalian circadian clock[J].Endocrinol,2003,177(1):7-15.
[80]Aton SJ,Colwell CS,Harmar AJ,et al.Vasoactive intestinal polypeptide mediates circadian rhythmicity and synchrony in mammalian clock neurons[J].Nat Neurosci,2005,8(4):476-483.
[81]Yarnada S,Mikami S,Yanaihara N.Immunohisto-chemical localization of vasoactive intestinal polypeptide(VIP)-containing neurons in the hy-pothalamus of the Japanese quailCoturnix coturnix[J].Cell Tissue Res,1982,226:13-26.
[82]Mikami S,Yamada S.Immunohistochemistry of the hypothalamic neuropeptides and anterior pituitary cells in the Japanese quail[J].J Exp Zool,1984,232:405-417.
[83]王荫亭,许红艳,王书荣.家鸽血管活性肠肽阳性神经元在顶盖和峡核中的分布及形态特征[J].生物物理学报,1993,9(3):447-452.
[84]王荫亭,许红艳,王书荣.血管活性肠肽免疫阳性神经元在家鸽中脑和间脑中的分布[J].生物物理学报,1994,10(1):103-110.
[85]左明雪,李锐.血管活性肠肽在环鸽旁听觉神经通路中的分布[J].解剖学报,1998,29(2):135-138.
[86]蒋书东,周天红,张丽霞.血管活性肠肽在皖南白鹅中脑和脑桥内的分布[J].中国兽医科技,2004,34(1]):59-61.
[87]Gerhold LM,Wise PM.Vasoactive intestinal polypeptide regulates dynamic changes in astrocyte morphometry:Impact on gonadotropin releasing hormone neurons[J].Endocrinology,2006,Feb 9.
[88]Oelschlager HH,Nakamura M,Herzog M,Burda H.Visual system labeled by c-Fos immunohistochemistry after light exposure in the 'blind' subterranean Zambian mole-rat (Cryptomys anselli).Brain Behav.Evo.1,2000,55(4):209-220.
[89]李锦新,马宁芳.光刺激移植视网膜诱导大鼠上丘FOS的表达.中国组织化学与细胞化学杂志,2003,12(1):42-45.
[90]方富贵,章孝荣.P物质研究进展[J].动物医学进展,2005,26(1):6-8.
[91]Battmann T,Parsadaniantz S M,Jeanjean B,et al.In-vivo inhibi-tion of the preovulatoryLHsurge by substance P and in-vitromodula-tion of gonadotrophin-releasing hormone-induced LH release by sub-stance P.oestradiol and progesterone in the female rat[J].Journal of Endocrinology,1991,130:169-175.
[92]Duval P,Lenoir V,Garret C,et al.Reduction of the amplitude of preovalatory LH and FSH surges and of the amplitude of the in vitro GnRH-induced LHrelease by substance P.Reversal of the effect by RP67580[J].Neuropharmacology,1996,35(12):1805-1810.
[93]姜恩魁,李志山,常志杰等.大鼠性周期中下丘脑和垂体P物质含量的变化[J].锦州医学院学报,1987,8(4):315-317.
[94]姜恩魁,李志山,常志杰等.大白鼠垂体P物质含量与卵巢孕酮生成关系的研究[J].锦州医学院学报,1988,9(2):108-109.
[95]Jessop D S,Chowdrey H S,Larsen P J,et al.Substance P:multi-functional peptide in the hypothalamo-pituitary system[J].Journal ofEndocrinology,1992,132:331-337.
[96]唐宗湘,马殷华.鸟类视觉系统的离顶盖通路[J].广西师范大学学报(自然科学版),2004,22(4):79-82.
[97]WANG Y,XIAO J,WANG S R.Excitatory and inhibitory receptive fields of tectal cells are differentially modified by magnocellular and parvocellular divisions of the pigeon nucleus isthmi[J].J Comp Physio[A],2000,186:505-511.
[98]HELLMANN B,MANNS M,GUNTURKUN O.Nucleus isthmi,pars sere ilunaris as a key component of the tectofugal visual system in pigeons[J].Comp Neuro 1,2001,436:153-166.
[99]赵丽杰.鸡中脑视顶盖Ⅰ层细胞构筑及几种神经肽在相关神经核团中的分布研究[D],河北农业大学,2007.
[100]刘佳.雏鸡中脑视顶盖SGC层细胞构筑及几种神经肽在OT-Rt通路的定位研究[D],河北农业大学,2007.
[101]Larsen P J,SaermarkT,Mau S E.Bindingof an iodinated substance P analogue to cultured anterior pituitary prolactin and luteinizing hor-mone-containing cells[J].The Journal of Histochemistry and Cyto-chemistry,1992,40(4):487-493.
[102]Hajihira S et al.Brain Res,1997,751:124.
[103]Presley RW et al.[J].Neurosci,1990,10:23.
[104]Lupi D,Sekaran S,Jones SL,Hankins MW,Foster RG.Light-evoked FOS induction within the suprachiasmatic nuclei(SCN)of melanopsin knockout(Opn4-/-)mice:a developmental study.Chronobiol Int.2006,23(1-2):167-17.
[105]Hattar S,Lucas RJ,Mrosovsky N,et al.Melanopsin and rodcone photoreceptive systems account for all major accessory visual functions in mice[J].Nature,2003,424(6944):76-81.
[106]Panda S,Provencio I,Tu DC,et al.Melanopsin is required for non-image-forming photic responses in blind mice[J].Science,2003,301(5632):525-527.
[107]Benson DM.Strange vision:ganglion cells as circadian photoreceptors[J].Trends Neurosci,2003,26(6):314-320.
[108]Sancar A.Cryptoehrome:the second photoactive pigment in the eye and its role in circadian photoreception[J].Annu Rev Biochem,2000,69:31-67.
[109]Russell N,Van Gelder,Wee R,et al.Reduced pupillary light responses in mice lacking cryptochromes[J].Science,2003,299(5604):222.