黄喉wu高级发声中枢及其壳区的神经投射和P物质在发声听觉中枢的分布

用生物素示踪法和P物质(SP)免疫组化技术研究表明：黄喉wu的高级发声中枢(HVc)接受端脑听区(L)、新纹状体中部界面核、新纹状体巨细胞核(MAN)、丘脑葡萄形核、桥脑蓝斑核的传入，并有神经纤维投射到古纹状体栎核(RA)和嗅叶X区(X)；HVc壳投射到RA壳并接受L的传入。听觉控制与学习通路与发声中枢之间有许多神经联系，提示黄喉wu发声学习依赖于听觉反馈。在HVc、RA和MAN有SP阳性细胞体，在X、中脑背内侧核和延髓舌下神经核气管鸣管部、丘脑卵圆核壳区、中脑背外侧核壳区及中脑丘间核有SP阳性纤维和终末。SP广泛分布于发声-听觉中枢，可能参与了它们的活动。     

黄喉鵐高级发声中枢及其壳区的神经投射和P物质在发声听觉中枢的分布

用生物素示踪法和P物质 (SP)免疫组化技术研究表明 :黄喉的高级发声中枢 (HVc)接受端脑听区 (L)、新纹状体中部界面核、新纹状体巨细胞核 (MAN)、丘脑葡萄形核、桥脑蓝斑核的传入 ,并有神经纤维投射到古纹状体栎核 (RA)和嗅叶X区 (X) ;HVc壳投射到RA壳并接受L的传入。听觉控制与学习通路与发声中枢之间有许多神经联系 ,提示黄喉发声学习依赖于听觉反馈。在HVc、RA和MAN有SP阳性细胞体 ,在X、中脑背内侧核和延髓舌下神经核气管鸣管部、丘脑卵圆核壳区、中脑背外侧核壳区及中脑丘间核有SP阳性纤维和终末。SP广泛分布于发声 -听觉中枢 ,可能参与了它们的活动     

黄喉鹀高级发声中枢及其壳区的神经投射和P物质在发声听觉中枢的分布

用生物素示踪法和P物质(SP)免疫组化技术研究表明:黄喉(巫鸟)的高级发声中枢(HVc)接受端脑听区(L)、新纹状体中部界面核、新纹状体巨细胞核(MAN)、丘脑葡萄形核、桥脑蓝斑核的传入,并有神经纤维投射到古纹状体栎核(RA)和嗅叶X区(X);HVc壳投射到RA壳并接受L的传入.听觉控制与学习通路与发声中枢之间有许多神经联系,提示黄喉(巫鸟)发声学习依赖于听觉反馈.在HVc、RA和MAN有SP阳性细胞体,在X、中脑背内侧核和延髓舌下神经核气管鸣管部、丘脑卵圆核壳区、中脑背外侧核壳区及中脑丘间核有SP阳性纤维和终末.SP广泛分布于发声-听觉中枢,可能参与了它们的活动.     

黄喉Jiu高级发声中枢及其壳区的神经投射和P物质在发声听觉中枢的分布

用生物素示踪法和P物质（SP）免疫组化技术研究表明：黄喉Jiu的高级发声中枢（HVc) 接受端脑听区（L）、新纹状体中部界面核、新纹状体巨细胞核（MAN）、丘脑葡萄形核、桥脑蓝斑核的传入,并有神经纤维投射到古纹状体栎核（RA）和嗅叶X区（X）;HVc壳投射到RA壳并接受L的传入。听觉控制与学习通路与发声中枢之间有许多神经联系,提示黄喉Jiu发声学习依赖于听觉反馈。在HVc、RA和MAN有SP阳性细胞体,在X、中脑背内侧核和延髓舌下神经核气管鸣管部、丘脑卵圆核壳区、中脑背我 核壳区及中脑丘间核有SP阳性纤维和终末。SP广泛分布于发声－听觉中枢,可能参与了它们的活动。     

黄雀发声核团与部分听觉中枢内P物质的分布和性双态比较

用免疫组织化学方法研究P物质在雌雄黄雀发声控制核团和听觉中枢内的分布,结合计算机图像分析仪检测SP免疫阳性细胞和末梢的灰度值,并作雌雄比较。结果如下：1．在发声学习中枢嗅叶X区有大量的SP阳性神经末梢和一些神经细胞。2．在发声控制核团前脑高级发声中枢(HVc)、古纹状体栎核、发声学习中枢新纹状体巨细胞核和丘脑背内侧核外侧部内有许多的SP免疫阳性细胞。3．在发声控制中枢中脑背内侧核和延髓舌下神经核气管鸣管部、听觉中枢丘脑卵圆核的壳区、中脑背外侧核壳区及中脑丘间核等有密集的SP免疫阳性神经末梢和纤维分布;雄性发声中枢内SP的分布比雌性丰富,两者有显著的差异。结果表明：SP的分布在雌雄发声中枢之间存在显著的性双态;SP广泛分布于黄雀发声控制核团和部分听觉中枢内,提示SP可能在发声控制及听觉中枢内具有重要的生理功能。     

黄雀发声核团与部分听觉中枢内P物质的分布和性双态比较

用免疫组织化学方法研究P物质在雌雄黄雀发声控制核团和听觉中枢内的分布,结合计算机图像分析仪检测SP免疫阳性细胞和末梢的灰度值,并作雌雄比较。结果如下:1.在发声学习中枢嗅叶X区有大量的SP阳性神经末梢和一些神经细胞。2.在发声控制核团前脑高级发声中枢(HVc)、古纹状体栎核、发声学习中枢新纹状体巨细胞核和丘脑背内侧核外侧部内有许多的SP免疫阳性细胞。3.在发声控制中枢中脑背内侧核和延髓舌下神经核气管呜管部、听觉中枢丘脑卵圆核的壳区、中脑背外侧核壳区及中脑丘间核等有密集的SP免疫阳性神经末梢和纤维分布;雄性发声中枢内SP的分布比雌性丰富,两者有显著的差异。结果表明:SP的分布在雌雄发声中枢之间存在显著的性双态;SP广泛分布于黄雀发声控制核团和部分听觉中枢内,提示SP可能在发声控制及听觉中枢内具有重要的生理功能。     

鸣禽锡嘴雀(Coccothraustes coccothraustes)发声中枢的定位研究

用常规组织学,HRP 逆行示踪,电生理等方法确定了鸣禽锡嘴雀控制发声的神经核团及这些核团的定位坐标值。锡嘴雀控制发声的神经通路由四级神经核团组成。位于端脑上纹状体腹侧的尾部区域(HVc)是控制鸣禽发声的高位中枢,它发出的神经纤维投射到端脑原纹状体腹内侧的粗核(RA),由 RA 又发出两束纤维,分別投射到中脑丘间核(ICo)和延脑的中间核(IM)。左右侧发声控制神经通路并非严格单侧性,每侧气管鸣管肌群分別受双侧发声中枢的交叉控制。中脑 ICo 在控制发声行为中具有相对独立性。各级发声核团的定位坐标值为,HVc∶p1.3,L/R2.4,H0.8;RA∶1.4,L/R3.2,H6.0;ICo∶p0.3,L/R2.6,H8.5:IM∶P3.1,L/R1.0,H∶7.8。     

血清性激素在白腰文鸟前脑发声控制核团神经元早期性双态分化中的作用

本文究了雌、雄白腰文鸟(Lonchura striata swinhoei)不同发育时期前脑四个控制发声重要核团古纹状体栎核(RA)、新纹状体前部巨细胞核外侧部(LMAN)、X区(Area X)和高级发声中枢(HVC)中神经元数量、体积和体内雌二醇(E2)和睾酮(T)浓度的变化,以揭示性激素对鸣禽发声核团性双态性分化的影响。结果发现：(1)HVC、LMAN和X区在发育早期神经元数量和体积均呈显著性双态性差异,而RA神经元直至30日龄(P30)后才显示出明显性别差异(P＜0．05);(2)除RA外,HVC、LMAN和X区神经元体积的显著性双态性差异均发生在P20左右,P20后雌、雄核团内的神经元体积仅有较小范围的波动;(3)RA和LMAN神经元数量随年龄增长而逐渐减少;雌、雄鸟HVC和雄鸟X区的神经元数量在P20—30间均增长,雄鸟HVC的增长幅度显著大于雌鸟。P30后HVC和X区的神经元数量不再增加,开始小幅度减少;(4)四个发声核团的神经元数量和体积在P5-120期间均出现1—2个急剧变化期,此变化期与体内雌激素水平开始出现显著性差异的临界期及核团间神经联系开始建立的时期相对应;(5)雌、雄鸟血清中E2的水平在核团发育初期(P5)差异显著,雌鸟为雄鸟的7．45倍,P5后则呈相反方向变化趋势,在P15时雄鸟中的E2水平反超过雌鸟,差异显著(P＜0．05)。睾酮仅在发育P50后的雄鸟体内被检测出,雌鸟中始终未能检测出T的存在。结果提示：雌、雄白腰文鸟发育早期体内E2浓度的变化启动了HVC、LMAN和X区早期神经元性双态性的分化和持续发育;睾酮对雌、雄鸣禽发声控制核团中早期神经元的性双态性分化作用较小[动物学报49(3)：353—361,2003]。     

燕雀丘间复合体背内侧核对叫声的调控模式

鸣禽鸟的基本发声中枢——丘间复合体背内侧核(nucleus dorsalis medialis of the intercollicular complex,DM)对叫声的调控模式是诱发单次叫声。应用电生理与声学分析相结合的方法研究鸣禽鸟燕雀(Fringilla montifringilla)DM核团对声音调控的模式。经语图和频谱图分析,结果显示：燕雀左侧DM诱发单次叫声的声长和主能量区的带宽分别为右侧DM诱发叫声的2倍和1．7～3．1倍,提示燕雀DM对叫声声长和声强的调控能力,均呈明显的左侧优势。这与高级发声中枢(high vocal center,HVC)和古纹状体粗核(nucleus robustus archistriatalis,RA)等发声控制核团在控声模式中具有左侧优势的特征相似,从而为鸣禽发声控制通路高、低级中枢具有内源投射关系提供了声学上的证据。     

鸣禽栗 新纹状体L复合区与发声控制系统的神经联系

利用辣根过氧化物酶顺、逆行追踪的方法对鸣禽栗端脑新纹状体Ｌ复合区的神经联系进行了研究。结果表明,新纹状体Ｌ复合区中的Ｌ２区主要接受来自丘脑卵圆核的传入,并与Ｌ１和Ｌ３区有纤维联系;而Ｌ１和Ｌ３区传出纤维投射至高级发声中枢腹侧的架区、古纹状体粗核喙背侧的杯区以及上纹状体腹部尾外侧等处;Ｌ复合区亦接受来自新纹状体前部巨细胞核内侧部的传入投射。     

10种鸣禽控制鸣啭神经核团大小与鸣唱复杂性的相关性

为进一步揭示鸣禽鸣唱行为的神经生物学机制 ,本实验先对 8个科 10种鸣禽的鸣唱行为进行了观察和录音 ,并借助声谱软件分析了每种鸣禽的鸣唱复杂性。鸣唱语句复杂性的评价指标包括 :短语总数、每个短语中所含的平均音节数及音节种类数、所有短语的总音节数及音节种类数、最长短语的音节数及音节种类数。然后 ,测定了前脑三个鸣啭学习控制核团和一个与发声无关的视觉参考核团体积 ,分析了鸣唱语句复杂性和这些核团大小间的相关关系。结果表明 :1)HVC和HVC/Rt与 7种鸣唱语句复杂性指标无关 ;RA和RA/Rt与总音节种类数相关 ;AreaX与总音节数及音节种类数相关 ;2 )HVC/RA和HVC/X比值与多个鸣唱语句复杂性指标相关。结果提示 :鸣禽鸣唱复杂性不同特征可能受不同神经控制     

鸣禽发声学习行为的神经生物学机制

鸣禽发声学习的控制系统主要由一条直接神经通路和一条辅助神经通路组成,由前脑控制发声学习的最高中枢ＨＶＣ、旁嗅叶的Ｘ区和巨细胞核外侧部（ｌＭＡＮ）组成的辅助通路,对鸟类发声学习行为的发育和调制具有重要作用。发声控制系统中神经元类型、数量及再生与更替、神经组构及其重组、神经介质和受体的分布等差异,决定了鸣禽在发声学习行为表现的差异以及性双态性。本文对近年鸟类控制发声学习行煌神经生物学机制的进展作了较为     

白腰文鸟发声行为的神经发育

本文研究了 5～ 15 0日龄雄性白腰文鸟 (Lonchurastriataswinhoei)不同年龄段的声谱变化以及这种变化的神经调制机制。结果如下 :(1)HVC、RA和AreaX三个发声核团的神经联系基本接近成年鸟的水平后 ,幼鸟才开始学习鸣叫 (约 45日龄 ) ;(2 )HVC、RA和AreaX达到成年核团体积时 (约 80日龄 ) ,幼鸟才具有成年雄鸟的鸣叫模式 ;(3)发声控制核团的发育与核团间的神经支配有关 ,而基本不受鸣唱行为的影响 ,HVC、RA和AreaX的最快增长时间段各不相同 ,三个核团随年龄增长而呈现体积增长的显著变化 (one wayANOVA ,P <0 0 5 ) ,但各核团在任意两个时间段的体积差异并不都显著。结果提示 :发声行为产生的时间和发展与发声控制核团的发育、核团间的神经联系有关 ,最终的体积发育程度受内在遗传力的作用 ,同时可能还受神经核团建立正常神经联系时间的影响     

Acetylcholinesterase in central vocal control nuclei of the zebra finch (<Emphasis Type="Italic">Taeniopygia guttata</Emphasis>)

The distribution of acetylcholinesterase (AChE) in the central vocal control nuclei of the zebra finch was studied using enzyme histochemistry. AChE fibres and cells are intensely labelled in the forebrain nucleus area X, strongly labelled in high vocal centre (HVC) perikarya, and moderately to lightly labelled in the somata and neuropil of vocal control nuclei robust nucleus of arcopallium (RA), medial magnocellular nucleus of the anterior nidopallium (MMAN) and lateral magnocellular nucleus of the anterior nidopallium (LMAN). The identified sites of cholinergic and/or cholinoceptive neurons are similar to the cholinergic presence in vocal control regions of other songbirds such as the song sparrow, starling and another genus of the zebra finch (Poephila guttata), and to a certain extent in parallel vocal control regions in vocalizing birds such as the budgerigar. AChE presence in the vocal control system suggests innervation by either afferent projecting cholinergic systems and/or local circuit cholinergic neurons. Co-occurrence with choline acetyltransferase (ChAT) indicates efferent cholinergic projections. The cholinergic presence in parts of the zebra finch vocal control system, such as the area X, that is also intricately wired with parts of the basal ganglia, the descending fibre tracts and brain stem nuclei could underlie this circuitry’s involvement in sensory processing and motor control of song.     

Sexual Differences in Cell Loss during the Post-Hatch Development of Song Control Nuclei in the Bengalese Finch

Birdsongs and the regions of their brain that control song exhibit obvious sexual differences. However, the mechanisms underlying these sexual dimorphisms remain unknown. To address this issue, we first examined apoptotic cells labeled with caspase-3 or TUNEL in Bengalese finch song control nuclei - the robust nucleus of the archopallium (RA), the lateral magnocellular nucleus of the anterior nidopallium (LMAN), the high vocal center (HVC) and Area X from post-hatch day (P) 15 to 120. Next, we investigated the expression dynamics of pro-apoptotic (Bid, Bad and Bax) and anti-apoptotic (Bcl-2 and Bcl-xL) genes in the aforementioned nuclei. Our results revealed that the female RA at P45 exhibited marked cell apoptosis, confirmed by low densities of Bcl-xL and Bcl-2. Both the male and female LMAN exhibited apoptotic peaks at P35 and P45, respectively, and the observed cell loss was more extensive in males. A corresponding sharp decrease in the density of Bcl-2 after P35 was observed in both sexes, and a greater density of Bid was noted at P45 in males. In addition, we observed that RA volume and the total number of BDNF-expressing cells decreased significantly after unilateral lesion of the LMAN or HVC (two areas that innervate the RA) and that greater numbers of RA-projecting cells were immunoreactive for BDNF in the LMAN than in the HVC. We reasoned that a decrease in the amount of BDNF transported via HVC afferent fibers might result in an increase in cell apoptosis in the female RA. Our data indicate that cell apoptosis resulting from different pro- and anti-apoptotic agents is involved in generating the differences between male and female song control nuclei.     

Differential estrogen accumulation among populations of projection neurons in the higher vocal center of male canaries

The higher vocal center (HVC) of adult male canries undergoes a seasonal change in volume that corresponds to seasonal modifications of vocal behavior: HVC is large when birds produce stereotyped song (spring) and is small when birds produce plastic song and add new song syllables into their vocal repertoires (fall). We reported previously that systemic exposure to testosterone (T) produces an increase in the volume of HVC similar to that observed with long-day photoperiods. T-induced growth of HVC occured regardless of wheter the borders of HVC were defined by Nissl-staining, the distribution of androgen-concentrating cells, or the distribution of projection neurons [separate neuronal populations within HVC project to the robust nucleus of the archistriatum (RA) and to Area X of the avian striatum (X)]. In the present study we used steroid autoradiography to determine whether T can influence the distribution of HVC cells that bind estrogen, and we combined estrogen autoradiography with retrograde labeling to determine whether HVC neurons that project to RA versus X differ in their ability to accumulate estrogen. Results showed that T increased the volume of Nissl-defined HVC and although HVC contained a low density of estrogen-concentrating cells, T increased the spatial distribution of these cells to match the Nissl borders of HVC. We also identified a region containing a high density of estrogenconcentrating cells located medial to HVC [we call this region paraHVC (pHVC)], and T also increased the volume of pHVC. pHVC also contained numerous X-projecting neurons, but few if any RA-projecting neurons. Double-labeling analysis revealed the RA-projecting neurons did not accumulate estrogen, a small percentage of X-projecting neurons in HVC accumulated estrogen, and the majority of X-projecting neurons in pHVC showed heavy accumulation of estrogen. The data reported here and in our previous article suggest distinct roles for gonadal steroids within the HVC-pHVC complex: estrogens are concentrated by neurons that project to a striatal region that influences vocal production during song learning (X), whereas androgens are concentrated primarily by neurons that project to a motor region that is involved in vocal production during both song learning and the recitation of already-learned song (RA). © 1995 John Wiley & Sons, Inc.