Brand S, Rakic P
Anat Embryol (Berl). 1984;169(1):21-34. doi: 10.1007/BF00300583.
Cytodifferentiation and synaptogenesis in the neostriatum (caudate nucleus and putamen) were analyzed by the Golgi impregnation method and electron microscopy in 14 fetuses and 8 postnatal rhesus monkeys. During the second fetal month the neostriatum consists primarily of simple, mostly bipolar, immature cells and a small number of undefined profiles ending with growth cones. The first morphologically defined synapses appear in the putamen at embryonic day 60 ( E60 ) and in the head of the caudate nucleus at E65 . Synaptic density in both structures is less than one per 1000/micron2 of neuropil at this stage; synapses are characterized by asymmetric junctions between axonal profiles and immature dendritic shafts, accumulation of an intermembrane web and aggregation of round clear vesicles in presynaptic profiles. During the third fetal month neuronal cell bodies and glial cells enlarge, and axonal and dendritic processes in Golgi preparations become more complex. Although the basic morphology of synapses remains unchanged, their density increases to 9/1000 micron2 in the putamen and 3.7/1000 micron2 in the caudate. During the fourth fetal month the four principal cell classes of the neostriatum emerge. Spines on the shafts of dendrites are followed closely by the appearance of axospinous synapses. Synaptic density in the putamen is still significantly higher (10.1/1000 micron2) than in the caudate (5.4/1000 micron2), but by the end of the fifth fetal month ( E150 ) it is the same (80/1000 micron2) in both structures. A dramatic increase in synaptic density to 125/1000 micron2 occurs before term ( E165 ) with the emergence of the first asymmetric synapses as well as symmetric synapses with flat or pleomorphic vesicles that terminate predominately on dendritic shafts. Synaptic density continues to increase after birth, reaching a plateau of approximately 190/1000 micron2 at the end of the first postnatal month. Throughout postnatal development the proportions of symmetric and asymmetric synapses on the smooth dendritic shafts undergo systematic fluctuations which may reflect the ingrowth of various afferents as well as local cytological differentiation including the formation of cellular compartments.
运用高尔基染色法和电子显微镜技术,对14例胎儿及8只出生后的恒河猴新纹状体(尾状核和壳核)的细胞分化和突触形成进行了分析。在胎儿第二个月时,新纹状体主要由简单的、大多为双极的未成熟细胞以及少量以生长锥结尾的未明确形态的结构组成。最早在形态上确定的突触于胚胎第60天(E60)出现在壳核,于E65出现在尾状核头部。在此阶段,这两个结构中的突触密度均低于每1000平方微米神经毡1个;突触的特征是轴突结构与未成熟树突干之间的不对称连接、膜间网的积聚以及突触前结构中圆形清亮小泡的聚集。在胎儿第三个月时,神经元细胞体和胶质细胞增大,高尔基染色标本中的轴突和树突过程变得更加复杂。虽然突触的基本形态保持不变,但其密度在壳核中增加到每1000平方微米9个,在尾状核中增加到每1000平方微米3.7个。在胎儿第四个月时,新纹状体的四种主要细胞类型出现。树突干上的棘突出现后紧接着出现轴棘突触。壳核中的突触密度仍显著高于尾状核(分别为每1000平方微米10.1个和5.4个),但到胎儿第五个月末(E150),两者相同(均为每1000平方微米80个)。在足月前(E165),随着第一批不对称突触以及主要终止于树突干的具有扁平或多形小泡的对称突触的出现,突触密度急剧增加至每1000平方微米125个。出生后突触密度持续增加,在出生后第一个月末达到约每1000平方微米190个的平台期。在整个出生后发育过程中,光滑树突干上对称和不对称突触的比例经历系统性波动,这可能反映了各种传入纤维的长入以及包括细胞区室形成在内的局部细胞学分化。