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Introduction: Beyond the Synapse Cambridge University Press 978-1-107-41156-2 - Beyond the Synapse: Cell–Cell Signaling in Synaptic Plasticity Edited by R. Douglas Fields Excerpt More information Introduction: Beyond the synapse R. Douglas Fields Powerful new imaging techniques can now reveal Much like the work of Rose and his colleagues on synapses changing structure and track neurotransmitter learning in the chick, the involvement of cell adhesion receptors shuttling into and out of the synaptic mem- molecules and extracellular matrix molecules in brane. Even as synaptic plasticity is studied at a finer long-term potentiation (LTP), is the focus of the chapter scale than could be imagined previously, it is important by Richard LeBaron and colleagues. Here, the investi- to remember that learning and memory are behaviors, gators show that the specialized cell surface molecules not molecules, cells, or synapses. Synaptic plasticity is and the associated intracellular signaling enzymes of focal central, but in widening the scope of consideration, the adhesions in non-neuronal cells provide an important contributors to this book reveal that there is much to foundation for plasticity in synaptic connections between learning and memory which lies beyond the synapse. neurons in the hippocampus. All cells in the body communicate with one Lisa Boulanger’s research concerns molecules another, and they all change physiologically from past mediating cell–cell signaling in the immune system, experience – an elemental form of learning. Many the major histocompatibility complex (MHC) class 1 mechanisms of intercellular communication, intercel- proteins, in the context of structural and functional lular messenger systems, cell adhesion molecules, plasticity in the nervous system. This work is an growth factors, interactions with the immune system example of how broadening thinking beyond the cells, and gene expression, for example, are relevant to synaptic cleft may lead to fundamental insights into how neural circuits change their structure and function how the brain changes structure and function through from experience. Moreover, system- and circuit-level experience. Molecular mechanisms of cell–cell com- properties of nervous system operation are funda- munication used by other cells in the body are likely to mental to understanding the behaviors of learning and be adopted by neurons in several aspects of synaptic memory. communication and plasticity. Also, work such as Steven Rose begins with the Hebb hypothesis, the this, at the intersection between the immune and fulcrum which has leveraged the great mass of research nervous systems, may further illuminate how immune on learning and memory in modern times, but, as he responses and exposure to disease may affect devel- makes clear in his chapter, the Hebb hypothesis is a opment and plasticity of neural circuits. fulcrum, not an endpoint. His chapter also reminds us Information processing in the hippocampus that memory is a process, not a single event like the underlying episodic memory is examined in the chapter sudden jump in voltage of a synaptic potential in a slice by Howard Eichenbaum. “I woke, put on my pants, of brain triggered by the flick of a switch. Memory has ran down the stairs, and ate eggs while reading the paper multiple phases, spanning from milliseconds to weeks, in the kitchen.” Without this sequential coherence, and it results from a sequence of very different cellular fragmentary impressions of events – no matter how processes. Many engage machinery well outside the indelibly recorded in the altered strengths of individual synapse to reach into the nucleus of the cell. Tran- synaptic connections – could not yield a useful memory scription factors, cell adhesion molecules, and growth any more than the frames of a filmstrip sliced into frag- factors all come into play over the course of hours or ments could yield a meaningful sequence. Although the days in learning the simplest task: a chick avoiding entire record may be preserved, without the vital episodic selecting a distasteful bead after a single experience connections, the record cannot provide a comprehensible with it. memory. What are the cognitive processes and circuitry 1 © in this web service Cambridge University Press www.cambridge.org Cambridge University Press 978-1-107-41156-2 - Beyond the Synapse: Cell–Cell Signaling in Synaptic Plasticity Edited by R. Douglas Fields Excerpt More information 2 R. DOUGLAS FIELDS that preserves the sequence of records in an episodic pioneering studies of glia in regulating information- memory? Eichenbaum argues that three cognitive processing in the retina show that calcium signaling, processes supported by the hippocampus: associative glutamate, and purinergic signaling between neurons representation; sequential organization; and relational and glia contribute to light-evoked responses in the networking, must be integrated to provide a coherent retina. The retina, being the most accessible part of the episodic memory. CNS, is a window into how circuitry in the brain William Greenough and colleagues approach the operates, and through that window we see glia as a vital subject of memory from the opposite direction of most: part of the mechanism. beginning from the behavior and tracing to the roots of Behavioral states of arousal are fundamental to cellular changes induced by experience. His research learning, and Korz and Frey combine behavioral shows that there are indeed changes in neurons and experiments with cellular electrophysiology to untan- neuronal structure with experience, but the cellular gle the hormonal effects of stress and novelty on changes in the brain sculpted by experience are hardly learning. Cain, Debiec, and LeDoux examine con- limited to neurons. Glia of many different types (and solidation and reconsolidation of Pavlovian fear con- blood vessels) are altered by functional experience, ditioning, and isolate in detail the mechanisms of along with changes in synapse number, synapse emotional memory storage at a molecular level, with morphology, and neurogenesis. One particularly important implications for treating fear disorders in intriguing finding is that the myelin insulation on axons humans. The relationship between memory consoli- changes in animals brought up in impoverished or dation and reconsolidation emerges from studies of enriched environments, a subject that I consider briefly fear conditioning, with a wealth of information on the in a separate chapter. Operating well beyond the syn- circuitry, receptors, ion channels, intracellular signal- apse, changes in myelin may be an underappreciated ing cascades mobilized to consolidate short-term form of nervous system plasticity, affecting infor- memories into long-term memories, and reconsolidate mation-processing in the brain by regulating the speed the memory once it is retrieved. Neurotrophin sig- of conduction in neural circuits, and thus the degree of naling, nitric oxide signaling, and neuromodulators temporal summation at synapses. and hormones as well as regulation of gene tran- Several other chapters are devoted to consider- scription are considered in this comprehensive chapter ation of glia, brain cells which have been ignored in of fear conditioning memory. the majority of studies of learning and memory. The phases of memory consolidation extend Glia remain absent from all computational neuro- through cycles of sleep and wakefulness, and in their science and that will remain so for some time to come. chapter, Walker and Stickgold review the largely mys- At present our knowledge of glia and their interactions terious role of sleep in supporting memory consoli- with neurons is simply too limited to incorporate into dation and reconsolidation. Neuroimaging studies a quantitative theory of brain function in plasticity, yet show that a task learned in the daytime re-emerges in the these cells have a powerful influence on synaptic hippocampus during slow wave sleep. Cellular studies and neuronal function. Dmitri Leonoudakis and col- show that sleep can contribute as much to changes in leagues consider the involvement of cytokines released synaptic connectivity as visual experience does to visual from astrocytes in regulating a-amino-3-hydroxy-5- cortical neurons. Gene array studies show that sleep is methyl-4-isoxazolepropionic acid (AMPA) receptor hardly an idle period in brain function, transcription of trafficking in central nervous system (CNS) neurons. approximately 100 genes is increased during sleep. This Sarina Elmariah and colleagues examine the involve- includes many of the same immediate early genes ment of neurotrophin signaling among neurons and associated with memory formation. glia during synaptogenesis. Nedergaard and colleagues Sex hormones are powerful agents driving behav- have pioneered studies on the involvement of astrocytes ior, and their effects on the brain are dramatic. Changes in hippocampal synaptic transmission, and the chapter in synapse number and morphology are encountered by Liu and colleagues considers how inhibitory synaptic during the phases of hormonal cycles, and male and transmission is regulated through calcium-dependent female hormones have different effects on cognition release of glutamate from astrocytes. Eric Newman’s and neuronal protection and plasticity. Foy, Baudry, © in this web service Cambridge University Press www.cambridge.org Cambridge University Press 978-1-107-41156-2 - Beyond the Synapse: Cell–Cell Signaling in Synaptic Plasticity Edited by R. Douglas Fields
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