Dendrite

A dendrite (from Greek δένδρον déndron, "tree") or dendron is a branched cytoplasmic process that extends from a nerve cell that propagates the electrochemical stimulation received from other neural cells to the cell body, or soma, of the neuron from which the dendrites project. Electrical stimulation is transmitted onto dendrites by upstream neurons (usually via their axons) via synapses which are located at various points throughout the dendritic tree.

Dendrites play a critical role in integrating these synaptic inputs and in determining the extent to which action potentials are produced by the neuron.

Dendrites are one of two types of cytoplasmic processes that extrude from the cell body of a neuron, the other type being an axon. Axons can be distinguished from dendrites by several features including shape, length, and function. Dendrites often taper off in shape and are shorter, while axons tend to maintain a constant radius and can be very long. Typically, axons transmit electrochemical signals and dendrites receive the electrochemical signals, although some types of neurons in certain species lack specialized axons and transmit signals via their dendrites. Dendrites provide an enlarged surface area to receive signals from axon terminals of other neurons. The dendrite of a large pyramidal cell receives signals from about 30,000 presynaptic neurons. Excitatory synapses terminate on dendritic spines, tiny protrusions from the dendrite with a high density of neurotransmitter receptors. Most inhibitory synapses directly contact the dendritic shaft.

Synaptic activity causes local changes in the electrical potential across the plasma membrane of the dendrite. This change in membrane potential will passively spread along the dendrite, but becomes weaker with distance without an action potential. To generate an action potential, many excitatory synapses have to be active at the same time, leading to strong depolarization of the dendrite and the cell body (soma). The action potential, which typically starts at the axon hillock, propagates down the length of the axon to the axon terminals where it triggers the release of neurotransmitters, but also backwards into the dendrite (retrograde propagation), providing an important signal for spike-timing-dependent plasticity (STDP).

Most synapses are axodendritic, involving an axon signaling to a dendrite. There are also dendrodendritic synapses, signaling from one dendrite to another. An autapse is a synapse in which the axon of one neuron transmits signals to its own dendrite.

The general structure of the dendrite is used to classify neurons into multipolar, bipolar and unipolar types. Multipolar neurons are composed of one axon and many dendritic trees. Pyramidal cells are multipolar cortical neurons with pyramid-shaped cell bodies and large dendrites that extend towards the surface of the cortex (apical dendrite). Bipolar neurons have two main dendrites at opposing ends of the cell body. Many inhibitory neurons have this morphology. Unipolar neurons, typical for insects, have a stalk that extends from the cell body that separates into two branches with one containing the dendrites and the other with the terminal buttons. In vertebrates, sensory neurons detecting touch or temperature are unipolar. Dendritic branching can be extensive and in some cases is sufficient to receive as many as 100,000 inputs to a single neuron.

The term dendrites was first used in 1889 by Wilhelm His to describe the number of smaller "protoplasmic processes" that were attached to a nerve cell. German anatomist Otto Friedrich Karl Deiters is generally credited with the discovery of the axon by distinguishing it from the dendrites.

Some of the first intracellular recordings in a nervous system were made in the late 1930s by Kenneth S. Cole and Howard J. Curtis. Swiss Rüdolf Albert von Kölliker and German Robert Remak were the first to identify and characterize the axonal initial segment. Alan Hodgkin and Andrew Huxley also employed the squid giant axon (1939) and by 1952 they had obtained a full quantitative description of the ionic basis of the action potential, leading to the formulation of the Hodgkin–Huxley model. Hodgkin and Huxley were awarded jointly the Nobel Prize for this work in 1963. The formulas detailing axonal conductance were extended to vertebrates in the Frankenhaeuser–Huxley equations. Louis-Antoine Ranvier was the first to describe the gaps or nodes found on axons and for this contribution these axonal features are now commonly referred to as the Nodes of Ranvier. Santiago Ramón y Cajal, a Spanish anatomist, proposed that axons were the output components of neurons. He also proposed that neurons were discrete cells that communicated with each other via specialized junctions, or spaces, between cells, now known as a synapse. Ramón y Cajal improved a silver staining process known as Golgi's method, which had been developed by his rival, Camillo Golgi.