Herr Steve, Klug Karl, Sterling Peter, Schein Stan
Department of Psychology, Franz Hall, University of California, Los Angeles, Los Angeles, California 90095-1563, USA.
J Comp Neurol. 2003 Mar 3;457(2):185-201. doi: 10.1002/cne.10553.
Synaptic terminals of cones (pedicles) are presynaptic to numerous processes that arise from the dendrites of many types of bipolar cell. One kind of process, a central element, reaches deeply into invaginations of the cone pedicle just below an active zone associated with a synaptic ribbon. By reconstruction from serial electron micrographs, we show that L- and M-cone pedicles in macaque fovea are presynaptic to approximately 20 central elements that arise from two types of inner (invaginating) bipolar cell, midget and diffuse. In contrast, S-cone pedicles, with more synaptic ribbons, active zones/ribbon, and central elements/active zone, are presynaptic to approximately 33 central elements. Moreover, all of these arise from one type of bipolar cell, previously described by others, here termed an inner S-cone bipolar cell. Each provides approximately 16 central elements. Thirty-three is twice 16; correspondingly, these bipolar cells are twice as numerous as S cones. (Specifically, each S cone is presynaptic to four inner S-cone bipolar cells; in turn, each bipolar cell provides central elements to two S cones.) These bipolar cells are presynaptic to an equal number of small-field bistratified ganglion cells, giving cell numbers in 2G:2B:1S ratios. Each ganglion cell receives input from two or more inner S-cone bipolar cells and thereby collects signals from three or more S cones. This convergence, along with chromatic aberration of short-wavelength light, suggests that S-cone contributions to this ganglion cell's coextensive blue-ON/yellow-OFF receptive field are larger than opponent L/M-cone contributions via outer diffuse bipolar cells and that opponent L/M-cone signals are conveyed mainly by inner S-cone bipolar cells.
视锥细胞的突触终末(视锥小足)位于许多不同类型双极细胞树突产生的众多突起的突触前。其中一种突起,即中央突起,深入到视锥小足的凹陷处,就在与突触带相关的活性区下方。通过对连续电子显微镜图像的重建,我们发现猕猴中央凹的L-和M-视锥小足突触于大约20个中央突起,这些中央突起来自两种类型的内(凹陷)双极细胞,即侏儒双极细胞和弥散双极细胞。相比之下,具有更多突触带、活性区/突触带和中央突起/活性区的S-视锥小足突触于大约33个中央突起。此外,所有这些中央突起都来自一种双极细胞,此前其他人已对此进行过描述,这里称为内S-视锥双极细胞。每个内S-视锥双极细胞提供大约l6个中央突起。33是16的两倍;相应地,这些双极细胞的数量是S视锥细胞的两倍。(具体而言,每个S视锥细胞突触于四个内S-视锥双极细胞;反过来,每个双极细胞为两个S视锥细胞提供中央突起。)这些双极细胞突触于数量相等的小场双分层神经节细胞,细胞数量比例为2G:2B:1S。每个神经节细胞从两个或更多内S-视锥双极细胞接收输入,从而收集来自三个或更多S视锥细胞的信号。这种汇聚,连同短波长光的色差,表明S视锥细胞对该神经节细胞共同的蓝ON/黄OFF感受野的贡献大于通过外弥散双极细胞的拮抗L/M视锥细胞的贡献,并且拮抗L/M视锥细胞信号主要由内S-视锥双极细胞传递。