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cAMP CAN BE VIEWED AS SUBSERVING TWO MAJOR FUNCTIONS IN THE NERVOUS SYSTEM AND MOST OF THE EFFECTS OF cAMP ON CELL FUNCTION ARE MEDIATED VIA PROTEIN PHOSPHORYLATION First, cAMP mediates some short-term aspects of synaptic transmission: some rapid actions of certain neurotransmitters on ion channels that do not involve ligand-gated channels are mediated through cAMP. Second, cAMP, along with other intracellular messengers, plays a central role in mediating other aspects of synaptic transmission: virtually all other effects of neurotransmitters on target neuron functioning, both short-term and long-term, are achieved through intracellular messengers. This includes regulation of the general metabolic state of the target neurons, as well as modulatory effects on neurotransmitter synthesis, storage, release and receptor sensitivity; cytoskeletal organization and structure; and neuronal growth and differentiation. This also includes those long-term actions of neurotransmitters that are mediated through alterations in neuronal gene expression. membrane depolarization and more slowly to increases in cellular levels of cAMP by activation of Ca2+/ calmodulin-sensitive forms of adenylyl cyclase. cAMP then mediates several other effects of glutamate on the neurons. By virtue of numerous interactions between cAMP and other intracellular messenger pathways, these pathways play the central role in coordinating a myriad neuronal processes and adjusting neuronal function to environmental cues [43]. It is important to emphasize that such a role for cAMP and other intracellular messengers is not limited to actions of neurotransmitters mediated via G proteincoupled receptors. Thus, although activation of ligandgated ion channels leads to initial changes in membrane potential independent of intracellular messengers, it also leads to numerous additional, albeit slower, effects that are mediated via intracellular messengers. For example, activation of certain glutamate receptors, which are ligand-gated ion channels, leads rapidly to 64 THE CLAPPER 2015 - 2016 By far the most important mechanism by which cAMP exerts its myriad physiological effects is through the specific activation of cAMP-dependent protein kinase. This was demonstrated first by Krebs and coworkers for cAMP regulation of glycogenolysis, and shortly thereafter it was shown to be a widespread mechanism by Paul Greengard and his colleagues. Indeed, cAMP-dependent protein kinase is now known to phosphorylate virtually every major class of neural protein; this accounts for the ability of cAMP to influence so many diverse aspects of neuronal function. The ability of cAMP to activate protein kinases and the role of protein phosphorylation in the regulation of neuronal function are covered. Specificity is achieved at two levels: at the level of tissuespecific receptors for the neurotransmitter or hormone and at the level of tissue-specific substrate proteins for the protein kinase. Only tissues which possess specific receptors will respond to a certain neurotransmitter or hormone. Moreover, since all cells contain very similar catalytic subunits of cAMP-dependent protein kinase, the nature of the proteins that are phosphorylated in