. Our previous studies have shown that when slices of the rat superior colliculus (SC) are exposed to a solution containing 10 M bicuculline and a low concentration of Mg 2ϩ (0.1 mM), most neurons in the intermediate gray layer (stratum griseum intermediale; SGI), wide-field vertical (WFV) cells in the optic layer (stratum opticum; SO), and a minor population of neurons in the superficial gray layer (stratum griseum superficiale; SGS) exhibit spontaneous depolarization and burst firing, which are synchronous among adjacent neurons. These spontaneous and synchronous depolarizations were thought to share common mechanisms with presaccadic burst activity in SGI neurons. In the present study, we explored the site responsible for generation of synchronous depolarization of SGI neurons by performing dual whole cell recordings under different slice conditions. A pair of SGI neurons recorded in a small rectangular piece of the SGI punched out from the SC slice showed synchronous depolarization but far less frequently than those recorded in a small rectangular piece including SGS and SO. This suggests that the superficial layers are needed for triggering synchronous depolarization in the SGI. Furthermore, we recorded spontaneous depolarizations in pairs of neurons belonging to the different layers. Analysis of their synchronicity revealed that WFV cells in the SO exhibit synchronous depolarizations with both SGS and SGI neurons, and the onset of spontaneous depolarization in WFV cells precedes those of neurons in other layers. Further, when SGS and SGI neurons exhibit synchronous depolarizations, SGI neurons usually precede the SGS neurons. These observations give further evidence to the existence of interlaminar interaction between superficial and deeper layers of the SC. In addition, it is suggested that WFV cells can trigger burst activity in other layers of the SC and that there is an excitatory signal transmission from the deeper layers to the superficial layers.
I N T R O D U C T I O NOrienting behaviors such as saccades are preceded by burst firing of neurons in the deeper layers (intermediate and deep layers) of the mammalian superior colliculus (SC). The burst activity determines both the metrics and timing of saccades (Munoz and Wurtz 1995;Schiller and Koerner 1971;Schiller and Stryker 1972; Sparks and May 1980;Sparks et al. 1976;Wurtz and Goldberg 1972). Previous studies have suggested that the burst activity is attributed to local excitatory interactions by recurrent collaterals of deep layer neurons (Bozis and Moschovakis 1998; Moschovakis et al. 1988a,b).In our previous studies, using whole cell patch-clamp technique in rat brain stem slice preparations, we demonstrated the presence of local excitatory interactions among a large number of SGI neurons that communicate through excitatory recurrent connections (Isa et al. 1998; Isa 2003, 2004). Recruitment of these neurons to excitatory interactions was dependent on both activation of N-methyl D-aspartate (NMDA) receptors and release from GABAergic inhibition...