Donoghue J P, Ebner F F
J Comp Neurol. 1981 May 20;198(3):389-420. doi: 10.1002/cne.901980303.
The projections of the ventrobasal complex (VB), the ventrolateral complex (VL), and the central intralaminar nucleus (CIN) to the somatic sensory-motor (SSM) cortex of the Virginia opossum were studied with light and electron microscopic autoradiographic methods. VB, VL, and CIN have overlapping projections to SSM cortex and each one also projects to an additional cortical area. Unit responses to somatic sensory stimulation and the areal and laminar distribution of axons in cortex is different for VB, VL, and CIN, but the axons from each form similar round asymmetrical synapses, predominantly with dendritic spines. As in other mammals, VB units in the opossum have discrete, contralateral cutaneous receptive fields. VB projects somatotopically to SSM cortex and also projects to the second somatic sensory representation. Within the cortex, VB axons terminate densely in layer IV and the adjacent part of layer III. A few axons also terminate in the outermost part of layer I and the upper part of layer VI. Most VB axons terminate upon dendritic spines (86.6%), but they also contact dendritic shafts (10%) and neuronal cell bodies (3%). Neurons in VL have no reliable response to somatic stimulation under our recording conditions. VL projects to the SSM cortex and to the posterior parietal area. Throughout this entire projection field VL fibers terminate in layers I, III, and IV most densely, and sparsely in the other cortical layers. The density of termination in the mid-cortical laminae is quite sparse compared to VB, but the projection to layer I is considerably greater. Nearly all (93%) of VL axons contact dendritic spines, the remainder (7%) end on dendritic shafts. CIN is a thalamic target of ascending medial lemniscal, cerebellar, spinal, and reticular formation axons. Neurons in CIN respond to stimulation restricted to a particular body part, but typically responses may be evoked from larger areas and at longer latencies than neurons in VB that are related to the same body part. CIN neurons require a firm tap or electrical stimulation within their receptive field to elicit a response in the anesthetized preparation. CIN axons terminate throughout the entire parietal cortex, but unlike VB and VL, CIN fibers end almost exclusively in the outer part of layer I. Approximately 21% of CIN fibers contact dendritic shafts in layer I, which is twice the percentage of shafts contacted by VL or VB axons. All of the other CIN synapses are formed with dendritic spines. These experiments demonstrate three different pathways to SSM cortex. The results suggest that each projection has a unique role in controlling the patterns of activity of neurons within the SSM cortex.
采用光镜和电镜放射自显影方法,研究了弗吉尼亚负鼠腹侧基底复合体(VB)、腹外侧复合体(VL)和中央板内核(CIN)向躯体感觉运动(SSM)皮层的投射。VB、VL和CIN对SSM皮层有重叠投射,且各自还投射到一个额外的皮层区域。VB、VL和CIN对躯体感觉刺激的单位反应以及皮层中轴突的区域和层状分布有所不同,但来自每种结构的轴突形成相似的圆形不对称突触,主要与树突棘相连。与其他哺乳动物一样,负鼠的VB单位具有离散的对侧皮肤感受野。VB向SSM皮层进行躯体定位投射,也投射到第二躯体感觉代表区。在皮层内,VB轴突密集地终止于IV层和III层的相邻部分。少数轴突也终止于I层的最外层和VI层的上部。大多数VB轴突终止于树突棘(86.6%),但它们也与树突干(10%)和神经元胞体(3%)接触。在我们的记录条件下,VL中的神经元对躯体刺激没有可靠反应。VL投射到SSM皮层和顶叶后区。在整个这个投射区域,VL纤维最密集地终止于I、III和IV层,在其他皮层层中则较稀疏。与VB相比,皮层中层的终止密度相当稀疏,但对I层的投射则大得多。几乎所有(93%)的VL轴突与树突棘接触,其余(7%)终止于树突干。CIN是内侧丘系、小脑、脊髓和网状结构上行轴突的丘脑靶点。CIN中的神经元对局限于特定身体部位的刺激有反应,但通常与同一身体部位相关的CIN神经元相比,其反应可能从更大区域诱发,且潜伏期更长。在麻醉的标本中,CIN神经元需要在其感受野内进行有力的轻敲或电刺激才能引发反应。CIN轴突终止于整个顶叶皮层,但与VB和VL不同,CIN纤维几乎完全终止于I层的外部。大约21%的CIN纤维在I层与树突干接触,这是VL或VB轴突接触树突干百分比的两倍。所有其他CIN突触都与树突棘形成。这些实验证明了通往SSM皮层的三种不同途径。结果表明,每种投射在控制SSM皮层内神经元的活动模式中都有独特作用。