The two closely related gymnotiform fishes, Apteronotus and Eigenmannia, share many similar communication and electrolocation behaviors that require modulation of the frequency of their electric organ discharges. The premotor linkages between their electrosensory system and their medullary pacemaker nucleus, which controls the repetition rate of their electric organ discharges, appear to function differently, however. In the context of the jamming avoidance response, Eigenmannia can raise or lower its electric organ discharge frequency from its resting level. A normally quiescent input from the diencephalic pre-pacemaker nucleus can be recruited to raise the electric organ discharge frequency above the resting level. Another normally active input, from the sublemniscal pre-pacemaker nucleus, can be inhibited to lower the electric organ discharge frequency below the resting level (Metzner 1993). In contrast, during a jamming avoidance response, Apteronotus cannot lower its electric organ discharge frequency below the resting level. The sublemniscal pre-pacemaker is normally completely inhibited and release of this inhibition allows the electric organ discharge frequency to rise during the jamming avoidance response. Further inhibition of this nucleus cannot lower the electric organ discharge frequency below the resting level. Lesions of the diencephalic pre-pacemaker do not affect performance of the jamming avoidance response. Thus, in Apteronotus, the sublemniscal pre-pacemaker alone controls the changes of the electric organ discharge frequency during the jamming avoidance response.
The diencephalic nucleus electrosensorius (nE) of gymnotiform fish comprises a series of finely tuned neuronal filters for control of the jamming avoidance response (JAR) and probably other electromotor tasks as well. The nE receives electrosensory input from the dorsal torus semicircularis (TSd) and octavolateral input from the ventral torus (TSv). The nE, in turn, projects to various hypothalamic and thalamic nuclei, including the prepacemaker nucleus (PPn), which can modulate the frequency of electric organ discharges (EODs) via its unique input to the medullary pacemaker nucleus. Four subdivisions of the nE can now be recognized: 1) The beat-related area (nEb)--a rostral cluster of tightly packed cells which receives TSd input and projects to the inferior lobe, anterior tuberal nucleus, anterior thalamic nucleus, central posterior thalamic nucleus, and PPn. The nEb contains neurons responsive to beat patterns caused by jamming stimuli. Stimulation of the nEb with L-glutamate, however, fails to induce any EOD-frequency shift. 2) The area causing EOD-frequency rises (nE increases)--a horizontal band of cells at the dorsal aspect of the caudal nE which receives TSd input and projects to the PPn and vicinity and to the cerebellum; nE increases stimulation induces slow EOD-frequency rises characteristic of the JAR. Responses of these cells to jamming stimuli are not yet known. 3) The area causing EOD-frequency falls (nE decreases)--a horizontal band of cells at the ventral aspect of the caudal nE which receives TSd input and projects only to the PPn and vicinity; nE decreases stimulation induces slow EOD-frequency falls characteristic of the JAR. The responses of these cells to jamming stimuli are not yet known. 4) The acousticolateral region (nEar)--a complex medial region of the nE which receives input predominantly from the ventral torus and projects to the inferior lobe, anterior tuberal nucleus, central posterior thalamic nucleus, PPn, and cerebellum; the sensory and motor properties of this region are not known in detail, although auditory and mechanosensory responses have been recorded here. Projections to the PPn and its vicinity suggest direct control of electromotor behaviors by the nE, whereas thalamic and hypothalamic projections may provide a substrate for electrosensory influences on neuroendocrine and motivational control centers. The optic tectum projects strongly to the pretectum and various other diencephalic nuclei in the vicinity of the nE, but it does not innervate the nE itself. Accordingly, ablation of the tectum does not affect the performance of the JAR.
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