We studied the anatomical pathway underlying the nictitating reflex in the monitor lizard Varanus exanthematicus by the anterograde degeneration technique combined with retrograde transport of horseradish peroxidase (HRP) and electron microscopy. After application of HRP to the abducens nerve, retrogradely labeled neurons were observed in the ipsilateral principal and accessory abducens motor nuclei. The transection, in the same experiments, of the root of the trigeminal nerve resulted in massive degeneration of myelinated fibers in the descending trigeminal tract. In the ipsilateral accessory abducens nucleus, we observed electron-dense degenerating axon terminals that formed asymmetric synaptic contacts with the primary and secondary dendrites of large neurons retrogradely labeled with HRP. A few of the degenerating terminals could be traced in serial sections to myelinated axons. No terminal degeneration was found in the contralateral accessory abducens nucleus or in the ipsilateral and contralateral principal abducens nuclei. The present results are complementary with the findings of previous light microscopic experimental tracing studies (Barbas-Henry, H.A., and A.H.M. Lohman, J. Comp. Neurol. 1986, 254:314-329; see also J. Comp. Neurol. 1988, 267:370-386), and strongly suggest the existence in Varanus of a monosynaptic, unilateral reflex pathway in which trigeminal fibers, presumably originating from the cornea, synapse with motoneurons of the bursalis and retractor bulbi muscles, which are located in the accessory abducens nucleus. This monosynaptic pathway may mediate a rapid unilateral eyeball retraction and nictitating membrane extension.
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