During the first 10 days after birth in the rat there are a succession of major developmental stages in the retinotectal pathway. During most of this time, the only recordable event in the superior colliculus is spontaneous activity. We studied and characterized this spontaneous activity, hypothesizing that it could play an important role in pathway development. The spontaneous discharges are detectable on postnatal day 5 (P5). After P5, the number of spontaneously active cells per penetration increases up to P10, after which they decrease to adult-like levels by P14-P15. Between P5 and P10, the spontaneous discharges exhibit several patterns of activity, from constant firing to intermittent bursts with periods of quiescence, without any bearing to age. We isolated the retina and superior colliculus by injecting xylocaine onto the optic nerve and found no change in collicular activity. While this suggests that the spontaneous activity in the colliculus is independent of the retina at the ages studied, the opposite experiment, i.e., electrically stimulating the optic nerve, resulted in increased firing by collicular neurons, perhaps via nonclassical synaptic transmission. Finally, we compared interval histograms for spontaneously active cells between P5 and P15. The histograms suggest that at certain ages, spontaneous firing is more regular; moreover, these ages precede major functional advances, e.g., onset of numerous spontaneously firing cells at P6, the first response to optic nerve stimulation at P10, and the first light-evoked response at P12-P13. Our results support the hypothesis that spontaneous activity in the neonatal superior colliculus has a role in development of the retinotectal pathway, but the data also indicate that classical synaptic transmission is not involved.
The present study examined, in the superior colliculus (SC) of anaesthetised cats, the functional connectivity between superficial-layer neurones (SLNs) and tectoreticular neurones (TRNs: collicular output cells). TRNs were antidromically identified by electrical stimulation of the predorsal bundle. The auto- and cross-correlation histograms of visual responses of both types of neurones were recorded and analysed. A delayed, sharp peak in cross-correlograms allowed us to verify whether SLN and TRN cells were coupled; in addition, oscillatory activities were compared to verify if rhythmic responses of SLN sites were transmitted to TRN sites. We found that oscillatory activity was rarely observed in spontaneous activity of superficial (1/74) and TRN sites (1/48). Moving light bars induced oscillation in 31% (23/74) of the superficial-layer and in 23% (11/48) of the TRN sites. The strength of the rhythmic responses was determined by specific ranges of stimulus velocity in 83% (19/23) and 64% (7/11) of oscillating SLN and TRN sites, respectively. Frequencies of oscillations ranged between 5 and 125 Hz and were confined, for 53% of the cells, to the 5-20 Hz band. Thus, the band-width of frequencies of the stimulus-related oscillations in the superior colliculus was broader than the gamma range. Analysis of cross-correlation histograms revealed a significant predominant peak with a mean delay of 2.7+/-0.9 ms in 46% (17/37) of SLN-TRN pairs. Most correlated SLN-TRN pairs (88%: 15/17) had superimposed receptive fields, suggesting they were functionally interconnected. However, individual oscillatory frequencies of correlated and oscillatory SLN and TRN cells were never the same (0/8). Together, these results suggest that the neurones in collicular superficial layer contact TRNs and, consequently, support the idea that the superficial layers contribute to collicular outputs producing eye- and head-orienting movements.
The present investigation is aimed at characterizing the development of ON and OFF visually evoked responses in the rat superior colliculus from postnatal day 13 (P13) to postnatal day 25. Depth profiles of field potentials reveal that ON and OFF long latency biphasic field potentials are already present when collicular cells are first responsive to light (P13). There is an inversion in the polarity of these responses as the electrode penetrates the collicular layers, suggesting a synaptic organization similar to the one found in adult animals. At P15, OFF field potentials begin to exhibit oscillatory activity. Local cobalt injections within the superior colliculus abolishes these OFF oscillations, suggesting a postsynaptic origin. Fast Fourier transform (FFT) analysis of the OFF field potentials demonstrates that oscillatory activity increases in frequency during development. This increase is thought to reflect the myelination and stabilization of synaptic connections that occur during this period. To our knowledge, this is the first report of OFF oscillatory responses in the superior colliculus.
Simple and complex cells of visual areas of cats may be reliably classified according to the modulatory index (MI) of their responses. This investigation is aimed at analysing the MI in area 18 when a small region (about 200-400 microm in diameter) of area 17 was inactivated with a microinjection of GABA, in anesthetized cats. Cells were stimulated with sine-wave gratings whose orientation, spatial, and temporal frequencies were optimal for the studied unit. The AC and DC response components, and the MI were computed along with fast Fourier transforms of evoked discharges recorded as peristimulus time histograms. Results showed that these response components were relatively unaffected in simple cells, whereas complex cells exhibited large changes when area 17 was silenced. In particular, a large proportion of complex cells showed a MI greater than 1, thereby adopting a response pattern resembling simple cells. It is suggested that this subpopulation of complex cells receives a direct input from geniculate X cells.
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