Ultrastructure of synapses in the first-evolved nervous systems

The phylum Cnidaria represents the first group of animals to evolve a recognizable nervous system. A comparison of the ultrastructural features of synaptic loci in animals representing all four classes of the cnidaria has provided an overview of the first-evolved synapses that can be compared morphologically to synapses in higher forms. Synapses in these watery jellylike animals with unmyelinated axons are sparse and difficult to fix well. However, we now have sufficient evidence to define an early synapse as one with paired electron dense plasma membranes separated by a 13-25 nm gap containing intracleft filaments and with vesicles on one or both sides of the synaptic cleft. The vesicles are of three types: dense-cored, clear, and opaque. Neuromuscular synapses resemble neuronal synapses and lack the postsynaptic specializations of higher animals. However, some coelenterates, such as the jellyfish Chrysaora, have a postsynaptic cisterna in the muscle. Neuromuscular and neuronematocyte synapses can have either clear or dense-cored vesicles. Opaque vesicles at two-way interneuronal synapses and at neuromuscular synapses in the oral sphincter muscle of sea anemones can be labelled with antisera to the neuropeptides Antho-RFamide (Antho-Arg-Phe-NH2) and Antho-RWamides (Antho-Arg-Trp-NH2) I and II, respectively. That suggests that neuropeptides evolved as neurotransmitters early in the animal kingdom. The basic differences between first evolved synapses and synapses of higher animals are the lack of postjunctional folds at neuromuscular synapses and the presence of fewer and somewhat larger synaptic vesicles, generally containing granular cores, in the more primitive animals.