Chapter 12 · Nervous tissue and neural signaling · Topic 65

Neurons and glial cells

A&P IStructure and functionInteractive lesson

Nervous tissue is built from two kinds of cells: neurons, which carry signals, and glial cells, which make it possible for them to do so. This page covers neuron structure in detail and the types of glial cells: the parts of a neuron from the dendrites to the axon terminals, how neurons are classified by shape and by job, the four kinds of glia in the central nervous system and the two in the peripheral nervous system, how Schwann cells and oligodendrocytes build the myelin sheath, and why a cut nerve in your arm can grow back while damage to your spinal cord usually cannot.

What you already know

From the nervous tissue topic: a neuron receives signals on its dendrites and cell body, and sends its own signal away along a single axon to axon terminals, which release chemicals onto the next cell. Glial cells support and protect neurons, and unlike mature neurons they can divide. From the electrical signaling topics: when a stimulus pushes a neuron's membrane to threshold, it fires an all-or-none action potential that propagates along the axon. From the last topic: the brain and spinal cord form the CNS, and everything else forms the PNS. This page zooms in on the cells themselves.

The cell body in detail

The soma (soma = body) is another name for the cell body. It holds the nucleus and nearly all of the neuron's protein-making machinery. Stain a neuron and its soma is speckled with dark clumps, called Nissl bodies after the scientist who described them. They are stacks of rough ER covered with ribosomes. A neuron needs that much rough ER because its cell body must build proteins for an axon that may hold hundreds of times more cytoplasm than the soma itself.

Those proteins, and organelles such as mitochondria, reach the far end of the axon by axonal transport: motor proteins walk along microtubules, carrying cargo out from the soma and bringing worn-out material and chemical messages back. Fast transport moves vesicles hundreds of millimeters a day; slow transport moves the cytoskeleton and many enzymes only about a millimeter or a few millimeters a day. That slow rate will matter when you get to nerve regrowth.

From dendrite to axon terminal

Figure 1 shows a typical neuron from the CNS. Follow the path a signal takes through it.

Drawing of a single neuron. An irregular cell body holding a nucleus sends out several short, branching dendrites. One long axon leaves the cell body and is wrapped in evenly spaced segments of insulating covering; a glial cell reaches out with two arms to form two of the segments. The axon ends in many branches whose tips meet the dendrites of the next neuron. At the upper left, the axon of another neuron ends on this neuron's dendrites.
Figure 1. A multipolar neuron from the CNS: a cell body with many dendrites, one axon wrapped in segments of myelin with gaps between them, and branched axon terminals meeting the next neuron. One oligodendrocyte forms two of the segments. OpenStax Anatomy and Physiology 2e, Figure 12.8, openstax.org, CC BY 4.0.
DendritesAxon
Number per neuronUsually manyOne (it may branch into collaterals)
Direction of signalToward the somaAway from the soma
Kind of electrical signalGraded potentialsAction potentials
LengthShort, tapering, heavily branchedFrom under a millimeter to over a meter; uniform width
Nissl bodies (rough ER)Present near the somaAbsent, from the hillock on
MyelinNoneOften present

Classifying neurons by shape

Neurons come in many shapes, but anatomists sort them by one simple feature: how many processes (extensions) leave the soma. Figure 2 shows the three main types.

Three line drawings of neurons. In the first, a round cell body hangs below a single short stalk that joins one long process, which ends in branches at both ends. In the second, an oval cell body sits in the middle of a line, with one branched process leaving each end. In the third, a cell body with many short branched processes sends out one long process that ends in branches.
Figure 2. Neurons classified by the number of processes leaving the cell body: one (unipolar), two (bipolar) or many (multipolar). OpenStax Anatomy and Physiology 2e, Figure 12.9, openstax.org, CC BY 4.0.

In a unipolar neuron the signal does not pass through the soma at all. It starts near the sensory receptor ending, runs up the peripheral branch, past the T-junction and on into the CNS, with the soma sitting off to one side like a station beside the track. Both branches carry action potentials and are built like an axon. Some books still call the peripheral branch a dendrite because it carries signals toward the soma's side.

MultipolarBipolarUnipolar (pseudounipolar)
Processes leaving the somaThree or more: one axon, many dendritesTwo: one axon, one dendriteOne, which splits into two branches
How commonThe vast majority of neuronsRareCommon in the PNS
Usual jobMotor neurons and interneuronsSensory, in a few sense organsSensory neurons for touch, pain, temperature, body position and organ stretch
Where the soma sitsMostly CNS gray matter; also ganglia of the autonomic nervous systemIn the sense organ itselfIn a ganglion beside the brain or spinal cord

Classifying neurons by job

You can also sort neurons by which way they carry information, which matches the divisions you met in the last topic.

Put the two classifications together and a pattern appears: a neuron's shape follows its job and its position. Figure 3 traces a signal through all three functional types.

CNS (spinal cord) sensory receptor sensory neuron (unipolar): soma in a ganglion interneuron motor neuron (multipolar) skeletal muscle
Figure 3. Signal flow through the three functional types: a unipolar sensory neuron carries input to the CNS, an interneuron integrates, and a multipolar motor neuron carries the command out. Dashed arrows mean "signals flow to".
Sensory neuronInterneuronMotor neuron
Carries signalsFrom sensory receptors to the CNSBetween neurons within the CNSFrom the CNS to effectors
Also calledAfferent neuronAssociation neuronEfferent neuron
Usual shapeUnipolar (bipolar in some sense organs)MultipolarMultipolar
Where the soma sitsA ganglion outside the CNSCNS gray matterCNS gray matter (somatic); a ganglion outside the CNS for the last neuron of an autonomic chain
Share of all neuronsA small fractionThe vast majorityA small fraction

Glial cells: six kinds

You met glial cells in the nervous tissue topic as the support cells of nervous tissue. There are six kinds: four in the CNS and two in the PNS. Each one has a distinct job, and several of them have a partner on the other side of the CNS–PNS border that does a similar job in a different way.

How many glial cells are there? Direct counts of whole human brains find about 86 billion neurons and a similar number of other cells, most of them glia, so the overall ratio is close to one to one. It varies a great deal by region: glia far outnumber neurons in the white matter, and neurons outnumber glia in some densely packed gray matter.

Glia of the CNS

Figure 4 shows the four kinds of glial cells in the CNS, gathered around two neurons.

A drawing of four kinds of support cells among two large branching neurons. Star-shaped cells with many branching processes sit among the neurons and touch them. Two small cells with fine, bristly processes sit nearby. A row of cube-shaped cells lines a surface at the lower left, sending long thin processes up into the tissue. Pale cells at the lower right send out arms that wrap segments of several different axons in layered sleeves.
Figure 4. The four glial cells of the CNS: a star-shaped astrocyte, small microglial cells, a row of ependymal cells lining a fluid-filled space, and oligodendrocytes whose arms wrap segments of several axons. OpenStax Anatomy and Physiology 2e, Figure 12.11, openstax.org, CC BY 4.0.

Astrocytes

Astrocytes (astr- = star, -cyte = cell) are star-shaped cells with many branching processes. They are the most varied glial cells in what they do:

Oligodendrocytes

Oligodendrocytes (oligo- = few, dendr- = branch, -cyte = cell) have a handful of arms. Each arm flattens into a sheet and wraps itself many times around a segment of an axon, forming the insulating layers of myelin described below. One oligodendrocyte can myelinate segments of dozens of different axons.

Microglia

Microglia (micro- = small) are small cells with fine, constantly moving processes. They are the resident immune cells of the CNS: they patrol the tissue, and when they detect damage or infection they multiply, move to the spot and engulf dead cells, debris and microbes by phagocytosis, acting like the macrophages of connective tissue. They also prune unneeded connections between neurons as the brain develops. Unlike the other glia, which arise from the same embryonic tissue as neurons, microglia come from immune precursor cells that settle in the brain early in development.

Ependymal cells

Ependymal cells (ependyma = upper garment) form a single layer of cube- to column-shaped cells lining the fluid-filled cavities inside the brain and the central canal that runs down the middle of the spinal cord. Many carry cilia, whose beating helps keep the fluid moving. Specialized ependymal cells, together with capillaries, produce that fluid, which cushions the brain and spinal cord; the topic on the brain's protection covers it.

Glia of the PNS

The PNS has only two kinds of glial cells. Figure 5 shows both around a sensory neuron.

A drawing of a sensory neuron's round cell body covered by a layer of flattened cells, like tiles over a ball. A single stalk leaves the cell body and joins an axon that runs across the bottom of the picture. Elongated cells wrapped around the axon sit end to end along it, and the cut end of the axon shows the layers wrapped around it.
Figure 5. The two glial cells of the PNS: satellite cells covering the cell body of a sensory neuron in a ganglion, and Schwann cells lined up along its axon. OpenStax Anatomy and Physiology 2e, Figure 12.12, openstax.org, CC BY 4.0.
CNS gliaPNS glia
KindsFour: astrocytes, oligodendrocytes, microglia, ependymal cellsTwo: Schwann cells, satellite cells
Make myelinOligodendrocytesSchwann cells
Surround neuron cell bodies and regulate their fluidAstrocytesSatellite cells
Clear debris and fight infectionMicroglia, the resident immune cellsSchwann cells, with macrophages that arrive from the blood
Line fluid-filled spacesEpendymal cellsNo equivalent
Response to an axon injuryA glial scar forms; axons rarely regrowSchwann cells guide the regrowing axon

The myelin sheath

In the nervous tissue topic you read that some glial cells wrap axons in many layers of their own plasma membrane. That wrapping is the myelin sheath (myel- = marrow, from its fatty, marrow-like look), and the material itself is myelin. Because it is layer on layer of membrane with almost no cytoplasm between, myelin is mostly lipid, about 70 to 80% of its dry weight, with the rest protein. Lipid is a poor conductor, so myelin works as electrical insulation, and its fat gives white matter its color.

Myelin is not continuous. Each glial wrap covers one segment of axon, up to a millimeter or two long, called an internode. Between neighboring segments is a tiny gap, about a thousandth of a millimeter wide, where the axon's membrane is bare: a node of Ranvier (named after the anatomist Louis-Antoine Ranvier), also called a myelin sheath gap. Voltage-gated sodium channels are packed densely at the nodes and sparse under the myelin. Axons wrapped this way are myelinated; axons with no myelin are unmyelinated.

Myelin makes an action potential travel far faster along an axon, up to about 120 meters per second in the thickest myelinated axons compared with only about 0.5 to 2 meters per second in thin unmyelinated ones. How the insulation and the nodes produce that speed is the subject of the next topic.

How Schwann cells and oligodendrocytes build myelin

Both cells make myelin the same basic way. The glial cell flattens part of itself into a sheet, lays the sheet against the axon, and spirals around it, sometimes a hundred times, like a bandage wound around a finger. As it winds, the cytoplasm is squeezed out of the layers, leaving tightly packed membranes. Figure 6 compares the two cells.

PNS: Schwann cells Schwann cell nucleus, outside the wrap node of Ranvier One Schwann cell = one segment of one axon CNS: an oligodendrocyte one cell body, many axons
Figure 6. Myelin two ways. In the PNS, each Schwann cell wraps one segment of one axon, and its nucleus and cytoplasm stay on the outside of the wrap. In the CNS, each arm of one oligodendrocyte wraps a segment of a different axon.

The differences matter for repair:

Schwann cellOligodendrocyte
LocationPNSCNS
Axons myelinated per cellOne segment of one axonSegments of many axons (often dozens)
Where its nucleus sitsOn the outside of its own wrapIn a cell body away from the axons
Outer living layer (neurilemma)YesNo
Handles unmyelinated axons byCradling several of them in grooves of one cellLeaving them bare
Role after an axon is cutClears debris and forms a tube that guides regrowthLeaves debris that contains growth-blocking proteins

Myelination starts before birth and goes on for years. A newborn's nervous system is only partly myelinated, which is one reason babies' movements are slow and poorly coordinated; some regions of the brain keep adding myelin into your twenties.

Unmyelinated axons

Not every axon is myelinated. Thin axons, such as those carrying dull, aching pain and many autonomic axons, are unmyelinated. In the PNS they are still not bare: a single Schwann cell cradles several of them in grooves in its surface, without wrapping any of them in layers. In the CNS, unmyelinated axons simply run among the other cells. Signals travel along them much more slowly.

Why a cut PNS axon can grow back

A carpenter cuts a nerve at the base of his thumb. The surgeon sews the cut ends together and warns him that feeling will come back slowly, starting near the scar and moving out toward the fingertip over months. Figure 7 shows why.

1. The axon is cut muscle 2. The part beyond the cut breaks down; debris is cleared macrophages and Schwanncells engulf debris cell body swells, makes more protein 3. Schwann cells line the tube; a sprout grows in regeneration tube of Schwann cells 4. The axon reaches its target and is myelinated again
Figure 7. Regeneration of a cut PNS axon. The part beyond the cut dies back, Schwann cells and macrophages clear it, Schwann cells line up into a guiding tube, and the axon regrows along it, about a millimeter a day. The new myelin segments are shorter and thinner than the originals.

Step by step:

  1. The far segment breaks down. The part of the axon beyond the cut has lost its supply of proteins from the soma. Within days it fragments, and its myelin falls apart with it. This is called Wallerian degeneration, after the physician Augustus Waller, who described it.
  2. Debris is cleared. Schwann cells in the far segment break down their own myelin and signal macrophages to come in from the blood. Together they clear the wreckage within a few weeks.
  3. The soma gears up. The neuron's cell body swells and its Nissl bodies disperse as it switches its protein production from signaling to growth.
  4. Schwann cells build a tube. The Schwann cells multiply and line up in a column inside the connective tissue sheath that once surrounded the axon, forming a regeneration tube. They release growth-promoting proteins and coat the tube with adhesive molecules that growing axons cling to.
  5. The axon regrows. Sprouts grow out from the cut end of the stump. One that finds the tube is guided along it at roughly 1 millimeter per day, about the speed of slow axonal transport.
  6. Reconnection. If the axon reaches a target, Schwann cells wrap it in new myelin and it can work again.

Recovery is often incomplete. If the cut ends are far apart, or scar tissue blocks the way, sprouts cannot find the tube and may form a painful tangle. Axons can also enter the wrong tube and reach the wrong target, for example a sensory axon regrowing to a different patch of skin. And if the journey takes too long, the muscle fibers without a nerve supply waste away before the axon arrives. That is why surgeons line up the cut ends precisely and repair nerves quickly.

Worked example: how long will regrowth take?

Problem. A nerve is cut and repaired 20 cm from the tip of the index finger. Axons regrow at about 1 mm per day once they cross the repair site. About how long until feeling can start to return to the fingertip?

  1. Put the distance in the same unit as the rate. 20 cm × 10 mm/cm = 200 mm.
  2. Write the relation. Time = distance ÷ rate.
  3. Substitute. Time = 200 mm ÷ 1 mm/day = 200 days.
  4. Convert to months. 200 days ÷ 30 days per month ≈ 6.7 months.

Answer. About 200 days, roughly six to seven months, and in practice somewhat longer: the sprouts need days to weeks to cross the repair site before steady growth begins. Feeling returns first near the scar and last at the fingertip.

Why CNS axons rarely grow back

Damage to the brain or spinal cord that cuts axons usually leaves permanent loss. It is not that CNS neurons cannot grow axons at all. In classic experiments, CNS axons grew for centimeters into a graft of peripheral nerve placed in the brain or spinal cord. The problem is mainly the surroundings the axon must grow through:

Research on spinal cord repair targets each of these: blocking the myelin proteins, breaking down the scar, and transplanting cells that act like Schwann cells.

PNS axon cutCNS axon cut
Myelinating gliaSchwann cellsOligodendrocytes
Debris clearanceFast (weeks): Schwann cells and macrophages from the bloodSlow (months or longer): microglia
Guidance for regrowthRegeneration tube of Schwann cellsNone
Growth-blocking signalsFewMyelin proteins and the astrocyte scar
Usual outcomeRegrowth at about 1 mm/day, often with partial recoveryLittle or no regrowth; loss is usually permanent

Neurons don't divide; glia do

Regrowing an axon is not the same as replacing a neuron. Mature neurons do not divide, so a neuron whose soma dies is gone. Whether adult human brains make any new neurons at all, for example in a region important for memory, is still unsettled: studies of donated brains disagree, some finding signs of new neurons into old age and others finding almost none after childhood. Either way, adult neuron production is far too small to replace what an injury destroys.

Glial cells, by contrast, keep dividing: astrocytes to form scars, microglia to fight infection, oligodendrocyte precursors to replace lost myelin, Schwann cells to rebuild a nerve. That capacity has a downside. Most tumors that start in brain tissue itself arise from glial cells or their precursors, and are called gliomas; tumors made of neurons are rare.