Dendritic Spines

Dendritic spines are important sites of excitatory synaptic transmission and changes in the strength of these synapses are likely to underlie important higher brain functions such as learning and memory. Spines form biochemical compartments for isolating reactions that occur at one synapse from those at other synapses thereby providing a possible way to ensure the specificity of connections between neurons in the brain.

Keywords: brain; neuron; dendrite; synapse; ultrastructure; plasticity

Figure 1. Pyramidal cell in hippocampal area CA1 of the rat brain showing the typical appearance of the principal excitatory neurons that occur throughout the brain. (a) The low-magnification view of these cells shows an apical dendrite projecting towards the upper left quadrant of the figure. Many lateral dendrites emerge from the large apical dendrite. Several basillar dendrites project from the base of the cell soma. Dendritic spines are the tiny projections that stud the surface of these dendrites in both the apical and basillar dendritic arbour. (b) A higher magnification view of the dendrites reveals the tiny spines a thin axon (arrow) passing through the dentritic arbour.
Figure 2. (a) Electron micrograph of a section through dendritic spines in stratum radiatum of hippocampal area CA1 that has been colour coded to identify dendrites (yellow), axons (green) and astroglia (purple). In this fortuitous section, three spines were sectioned parallel to their longitudinal axis revealing spines of the stubby (S), mushroom (m) and thin (t) morphologies. The postsynaptic density (PSD) occurs on the spine head (see t) immediately adjacent the synaptic cleft (c) and to a presynaptic axonal bouton that is filled with round vesicles (v). This t spine contains a small tube of smooth endoplasmic reticulum (ser) in its neck. In the m spine a spine apparatus (sa) is visible. A perforated postsynaptic density (perf) is evident on the head of another mushroom spine. Near to this spine is a large astrocytic process (ast) identified by the glycogen granules and clear cytoplasm. (b) A three-dimensional reconstruction of a dendrite showing a variety of spine and synapse shapes and the presence of polyribosomes (black spheres) at the base of the spines. Scale cube=1 m3.
Figure 3. (a) Spines increase the packing density of synapses. Schematic illustrates a cross-section through two dendrites, one without and one with dendrites spines. Convolution and interdigitation of dendrite, axon and spine membranes support more synapses. (b) The presence of spines also allows for an increase in synaptic density without increasing the overall volume of the brain. (c) Spines exist to amplify electrical potential at the synapse and promote associativity among neighbouring synapses. Spine shape and resistance of the spine neck may influence potential (V) generated by synaptic activation. (d) Spines exist as molecular compartments. Smooth endoplasmic reticulum (tubules), calcium and a myriad of other signalling mechanisms (stippling) are recruited in response to synaptic activation (asterisk).
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 Further Reading
    book Bliss T, Collingridge G and Morris R (eds) (2004) Long-term Potentiation: Enhancing Neuroscience for 30 Years. New York: Oxford.
    Ethell IM and Pasquale EB (2005) Molecular mechanisms of dendritic spine development and remodeling. Progress in Neurobiology 75: 161–205.
    Harris KM and Kater SB (1994) Dendritic spines: cellular specializations imparting both stability and flexibility to synaptic function. Annual Review of Neuroscience 17: 341–371.
    Harris KM and Stevens JK (1988) Dendritic spines of rat cerebellar Purkinje cells: serial electron microscopy with reference to their biophysical characteristics. Journal of Neuroscience 8: 4455–4469.
    Harris KM and Stevens JK (1989) Dendritic spines of CA1 pyramidal cells in the rat hippocampus: serial electron microscopy with reference to their biophysical characteristics. Journal of Neuroscience 9: 2982–2997.
    Nimchinsky EA, Sabatini BL and Svoboda K (2002) Structure and function of dendritic spines. Annual of Review of Physiology 64: 313–353.
    book Peters A, Palay SL and Webster HDeF (1991) The Fine Structure of the Nervous System: Neurons and Their Supporting Cells, 3rd edn New York: Oxford.
    book Sorra KE and Harris KM (2000) "Overview on the structure, composition, function, development, and plasticity of hippocampal dendritic spines". Eichenbaum HB (ed.) with Harris KM and Sorra KE (special issue eds) Dendritic Spines of the Hippocampus. Hippocampus 10: 501–511.
    book Stuart G, Spruston N and Hausser M (eds) (2001) Dendrites. New York: Oxford.
    Yuste R and Bonhoeffer T (2001) Morphological changes in dendritic spines associated with long-term synaptic plasticity. Annual Review of Neuroscience 24: 1071–1089.
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Bourne, Jennifer N, and Harris, Kristen M(Apr 2007) Dendritic Spines. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0000093.pub2]