26.1 Neurons and Glial Cells

Learning Objectives

Learning Objectives

In this section, you will explore the following questions:

  • What are the functions of the structural components of a neuron?
  • What are the four main types of neurons?
  • What are the functions of different types of glial cells?

Connection for AP® Courses

Connection for AP® Courses

Much information about the various organ systems of animals is not within the scope of AP®. The nervous system, however, was selected for in-depth study because an animal’s ability to detect, transmit, and respond to information is critical for survival. The nervous system interacts with all other organ systems to coordinate responses. The information in this chapter allows us to apply concepts we explored previously, including structure and function relationships, homeostasis, the movement of substances across cell membranes, cell signaling and communication, and the use of ATP.

Nervous systems in animals range from relatively simple nerve nets in jellyfish to a complex brain, spinal cord, and peripheral nervous system in humans. (For the purpose of AP®, you do not need to have detailed information about myriad types of nervous systems in other animals. Instead, just focus on the complex nervous system of humans.) The basic structure of the nervous system that reflects function is the neuron, of which there are three types: sensory, motor, and interneuron. A typical neuron consists of dendrites, a cell body, and an axon to detect, generate, transmit, and integrate signal information. Many neurons are surrounded by a type of glial cell called Schwann cells that form a myelin sheath, which acts as an electrical insulator, like the plastic wrap that surrounds the copper wires in a household appliance cord. The Schwann cells are separated by gaps of unmyelinated fibers called nodes of Ranvier over which the nerve impulse travels as the signal passes along the neuron, increasing the speed of transmission. As we will learn in the Nervous System Disorders section, some diseases of the nervous system result from the loss of myelin. Glial cells, often thought of as the supporting cast of the nervous system, outnumber neurons and play a role in the development of neurons, buffer harmful ions and chemicals, and provide myelin sheaths around neurons. Most brain tumors are caused by mutations in glial cells.

Information presented and the examples highlighted in the section support concepts outlined in Big Idea 3 of the AP® Biology Curriculum Framework. The AP® Learning Objectives listed in the Curriculum Framework provide a transparent foundation for the AP® Biology course, an inquiry-based laboratory experience, instructional activities, and AP® exam questions. A learning objective merges required content with one or more of the seven science practices.

Big Idea 3 Living systems store, retrieve, transmit, and respond to information essential to life processes.
Enduring Understanding 3.E Transmission of information results in changes within and between biological systems.
Essential Knowledge 3.E.2 Animals have nervous systems that detect external and internal signals, transmit and integrate information, and produce responses.
Science Practice 1.2 The student can describe representations and models of natural or man-made phenomena and systems in the domain.
Learning Objective 3.44 The student is able to describe how nervous systems detect external and internal signals.
Essential Knowledge 3.E.2 Animals have nervous systems that detect external and internal signals, transmit and integrate information, and produce responses.
Science Practice 1.1 The student can create representations and models of natural or man-made phenomena and systems in the domain.
Learning Objective 3.48 The student is able to create a visual representation to describe how nervous systems detect external and internal signals.

Nervous systems throughout the animal kingdom vary in structure and complexity, as illustrated by the variety of animals shown in Figure 26.2. Some organisms, like sea sponges, lack a true nervous system. Others, like jellyfish, lack a true brain and instead have a system of separate but connected nerve cells (neurons) called a nerve net. Echinoderms such as sea stars have nerve cells that are bundled into fibers called nerves. Flatworms of the phylum Platyhelminthes have both a central nervous system (CNS), made up of a small brain and two nerve cords, and a peripheral nervous system (PNS) containing a system of nerves that extend throughout the body. The insect nervous system is more complex but also fairly decentralized. It contains a brain, ventral nerve cord, and ganglia, which are clusters of connected neurons. These ganglia can control movements and behaviors without input from the brain. Octopi may have the most complicated of invertebrate nervous systems—they have neurons that are organized in specialized lobes and eyes that are structurally similar to vertebrate species.

Illustration A shows the nerve net of a hydra, which resembles a fish net surrounding the body. Illustration B shows the nervous system of a sea star. A nerve ring is present in the center of the body. Radiating out from this ring into the five arms are radial nerves. Illustration C shows the nervous system of a planarian, or flatworm. The flatworm has centralized ganglia, or brains, around each eye in the anterior end, and two nerve cords that run along the sides of the body. Transverse nerves connect th
Figure 26.2 Nervous systems vary in structure and complexity. In (a) cnidarians, nerve cells form a decentralized nerve net. In (b) echinoderms, nerve cells are bundled into fibers called nerves. In animals exhibiting bilateral symmetry such as (c) planarians, neurons cluster into an anterior brain that processes information. In addition to a brain, (d) arthropods have clusters of nerve cell bodies, called peripheral ganglia, located along the ventral nerve cord. Mollusks such as squid and (e) octopi, which must hunt to survive, have complex brains containing millions of neurons. In (f) vertebrates, the brain and spinal cord comprise the central nervous system, while neurons extending into the rest of the body comprise the peripheral nervous system. (credit e: modification of work by Michael Vecchione, Clyde F.E. Roper, and Michael J. Sweeney, NOAA; credit f: modification of work by NIH)

Compared to invertebrates, vertebrate nervous systems are more complex, centralized, and specialized. While there is great diversity among different vertebrate nervous systems, they all share a basic structure: a CNS that contains a brain and spinal cord and a PNS made up of peripheral sensory and motor nerves. One interesting difference between the nervous systems of invertebrates and vertebrates is that the nerve cords of many invertebrates are located ventrally whereas the vertebrate spinal cords are located dorsally. There is debate among evolutionary biologists as to whether these different nervous system plans evolved separately or whether the invertebrate body plan arrangement somehow flipped during the evolution of vertebrates.

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Watch this video of biologist Marc Kirschner discussing the flipping phenomenon of vertebrate evolution.

The nervous system is made up of neurons, specialized cells that can receive and transmit chemical or electrical signals, and glia, cells that provide support functions for the neurons by playing an information processing role that is complementary to neurons. A neuron can be compared to an electrical wire because it transmits a signal from one place to another. Glia can be compared to the workers at the electric company who make sure wires go to the right places, maintain the wires, and take down wires that are broken. Although glia have been compared to workers, recent evidence suggests that they also usurp some of the signaling functions of neurons.

There is great diversity in the types of neurons and glia that are present in different parts of the nervous system. There are four major types of neurons and they share several important cellular components.

Neurons

Neurons

The nervous system of the common laboratory fly, Drosophila melanogaster, contains around 100,000 neurons, the same number as a lobster. This number compares to 75 million in the mouse and 300 million in the octopus. A human brain contains around 86 billion neurons. Despite these very different numbers, the nervous systems of these animals control many of the same behaviors, from basic reflexes to more complicated behaviors like finding food and courting mates. The ability of neurons to communicate with each other as well as with other types of cells underlies all of these behaviors.

Most neurons share the same cellular components. But neurons are also highly specialized—different types of neurons have different sizes and shapes that relate to their functional roles.

Parts of a Neuron

Like other cells, each neuron has a cell body, or soma, that contains a nucleus, smooth and rough endoplasmic reticulum, Golgi apparatus, mitochondria, and other cellular components. Neurons also contain unique structures, illustrated in Figure 26.3 for receiving and sending the electrical signals that make neuronal communication possible. Dendrites are tree-like structures that extend away from the cell body to receive messages from other neurons at specialized junctions called synapses. Although some neurons do not have any dendrites, some types of neurons have multiple dendrites. Dendrites can have small protrusions called dendritic spines, which further increase surface area for possible synaptic connections.

Once a signal is received by the dendrite, it then travels passively to the cell body. The cell body contains a specialized structure, the axon hillock that integrates signals from multiple synapses and serves as a junction between the cell body and an axon. An axon is a tube-like structure that propagates the integrated signal to specialized endings called axon terminals. These terminals in turn synapse on other neurons, muscle, or target organs. Chemicals released at axon terminals allow signals to be communicated to these other cells. Neurons usually have one or two axons, but some neurons, like amacrine cells in the retina, do not contain any axons. Some axons are covered with myelin, which acts as an insulator to minimize dissipation of the electrical signal as it travels down the axon, greatly increasing the speed on conduction. This insulation is important as the axon from a human motor neuron can be as long as a meter, from the base of the spine to the toes. The myelin sheath is not actually part of the neuron. Myelin is produced by glial cells. Along the axon there are periodic gaps in the myelin sheath. These gaps are called nodes of Ranvier and are sites where the signal is recharged as it travels along the axon.

It is important to note that a single neuron does not act alone—neuronal communication depends on the connections that neurons make with one another, as well as with other cells, like muscle cells. Dendrites from a single neuron may receive synaptic contact from many other neurons. For example, dendrites from a Purkinje cell in the cerebellum are thought to receive contact from as many as 200,000 other neurons.

Visual Connection

Illustration shows a neuron. The main part of the cell body, called the soma, contains the nucleus. Branch-like dendrites project from three sides of the soma. A long, thin axon projects from the fourth side. The axon branches at the end. The tip of the axon is in close proximity to dendrites of an adjacent nerve cell. The narrow space between the axon and dendrites is called the synapse. Cells called oligodendrocytes are located next to the axon. Projections from the oligodendrocytes wrap around the axon
Figure 26.3 Neurons contain organelles common to many other cells, such as a nucleus and mitochondria. They also have more specialized structures, including dendrites and axons.

Types of Neurons

There are different types of neurons, and the functional role of a given neuron is intimately dependent on its structure. There is an amazing diversity of neuron shapes and sizes found in different parts of the nervous system and across species, as illustrated by the neurons shown in Figure 26.4.

Part A shows a pyramidal cell with two long, branched projections on either end of the soma. Dendrites branch from either side. Part B shows a Purkinje cell with highly branched dendrites opposite the axon. Part C shows cells with long, thin axons. The dendrites are less branched than in pyramidal or Purkinje cells.
Figure 26.4 There is great diversity in the size and shape of neurons throughout the nervous system. Examples include (a) a pyramidal cell from the cerebral cortex, (b) a Purkinje cell from the cerebellar cortex, and (c) olfactory cells from the olfactory epithelium and olfactory bulb.

While there are many defined neuron cell subtypes, neurons are broadly divided into four basic types: unipolar, bipolar, multipolar, and pseudounipolar. Figure 26.5 illustrates these four basic neuron types. Unipolar neurons have only one structure that extends away from the soma. These neurons are not found in vertebrates but are found in insects where they stimulate muscles or glands. A bipolar neuron has one axon and one dendrite extending from the soma. An example of a bipolar neuron is a retinal bipolar cell, which receives signals from photoreceptor cells that are sensitive to light and transmits these signals to ganglion cells that carry the signal to the brain. Multipolar neurons are the most common type of neuron. Each multipolar neuron contains one axon and multiple dendrites. Multipolar neurons can be found in the CNC, comprised of the brain and spinal cord. An example of a multipolar neuron is a Purkinje cell in the cerebellum, which has many branching dendrites but only one axon. Pseudounipolar cells share characteristics with both unipolar and bipolar cells. A pseudounipolar cell has a single process that extends from the soma, like a unipolar cell, but this process later branches into two distinct structures, like a bipolar cell. Most sensory neurons are pseudounipolar and have an axon that branches into two extensions: one connected to dendrites that receive sensory information and another that transmits this information to the spinal cord.

The unipolar cell has a single, long axon extending from the cell body. The bipolar neuron has two axons projecting from opposite sides of the cell body. The multipolar neuron has one long axon and several short, highly branched axons extending in all directions. The pseudounipolar neuron has one axon that forms two branches a short distance from the cell body, each of which extends in a different direction.
Figure 26.5 Neurons are broadly divided into four main types based on the number and placement of axons: unipolar, bipolar, multipolar, and pseudounipolar.

Everyday Connection

Neurogenesis

At one time, scientists believed that people were born with all the neurons they would ever have. Research performed during the last few decades indicates that neurogenesis, the birth of new neurons, continues into adulthood. Neurogenesis was first discovered in songbirds that produce new neurons while learning songs. For mammals, new neurons also play an important role in learning: Each day, about 1,000 new neurons develop in the hippocampus, which is the part of the brain involved in learning and memory. While most of the new neurons will die, researchers found that an increase in the number of surviving new neurons in the hippocampus correlated with how well rats learned a new task. Interestingly, exercise also promotes neurogenesis in the hippocampus. Stress has the opposite effect. While neurogenesis is quite limited compared to regeneration in other tissues, research in this area may lead to new treatments for disorders such as Alzheimer’s, stroke, and epilepsy.

How do scientists identify new neurons? A researcher can inject a compound called bromodeoxyuridine (BrdU) into the brain of an animal. While all cells will be exposed to BrdU, BrdU will only be incorporated into the DNA of newly generated cells that are in S phase. A technique called immunohistochemistry can be used to attach a fluorescent label to the incorporated BrdU, and a researcher can use fluorescent microscopy to visualize the presence of BrdU, and thus new neurons, in brain tissue. Figure 26.6 is a micrograph which shows fluorescently labeled neurons in the hippocampus of a rat.

In the micrograph, several cells are fluorescently labeled green only. Three cells are labeled red only, and four cells are labeled green and red. The cells labeled green and red are astrocytes, and the cells labeled red are neurons. The neurons are oval and about ten microns long. Astrocytes are slightly larger and irregularly shaped.
Figure 26.6 This micrograph shows fluorescently labeled new neurons in a rat hippocampus. Cells that are actively dividing have BrdU incorporated into their DNA and are labeled in red. Cells that express glial fibrillary acidic protein (GFAP) are labeled in green. Astrocytes, but not neurons, express GFAP. Thus, cells that are labeled both red and green are actively dividing astrocytes, whereas cells labeled red only are actively dividing neurons. (credit: modification of work by Dr. Maryam Faiz, et. al., University of Barcelona; scale-bar data from Matt Russell)

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This site contains more information about neurogenesis, including an interactive laboratory simulation and a video that explains how BrdU labels new cells.

Science Practice Connection for AP® Courses

Think About It

How does the unique structure of the neuron allow it to detect and ultimately transmit incoming signals?

Glia

Glia

While glia are often thought of as the supporting cast of the nervous system, the number of glial cells in the brain actually outnumbers the number of neurons by a factor of ten. Neurons would be unable to function without the vital roles that are fulfilled by these glial cells. Glia guide developing neurons to their destinations, buffer ions and chemicals that would otherwise harm neurons, and provide myelin sheaths around axons. Scientists have recently discovered that they also play a role in responding to nerve activity and modulating communication between nerve cells.

Types of Glia

There are several different types of glia with different functions, two of which are shown in Figure 26.7. Astrocytes, shown in Figure 26.8a make contact with both capillaries and neurons in the CNS. They provide nutrients and other substances to neurons, regulate the concentrations of ions and chemicals in the extracellular fluid, and provide structural support for synapses. Astrocytes also form the blood-brain barrier—a structure that blocks entrance of toxic substances into the brain. Astrocytes, in particular, have been shown through calcium imaging experiments to become active in response to nerve activity, transmit calcium waves between astrocytes, and modulate the activity of surrounding synapses. Satellite glia provide nutrients and structural support for neurons in the PNS. Microglia scavenge and degrade dead cells and protect the brain from invading microorganisms. Oligodendrocytes, shown in Figure 26.8b form myelin sheaths around axons in the CNS. One axon can be myelinated by several oligodendrocytes, and one oligodendrocyte can provide myelin for multiple neurons. This is distinctive from the PNS where a single Schwann cell provides myelin for only one axon as the entire Schwann cell surrounds the axon. Radial glia serve as scaffolds for developing neurons as they migrate to their end destinations. Ependymal cells line fluid-filled ventricles of the brain and the central canal of the spinal cord. They are involved in the production of cerebrospinal fluid, which serves as a cushion for the brain, moves the fluid between the spinal cord and the brain, and is a component for the choroid plexus.

Illustration A shows various types of glial cells surrounding a multipolar nerve of the central nervous system. Oligodendrocytes have an oval body and protrusions that wrap around the axon. Astrocytes are round and slightly larger than neurons, with many extensions projecting outward to neurons and other cells. Microglia are small and rectangular, with many fine projections. Ependymal cells have small, round bodies lined up in a row. Long extensions connect with an astrocyte. Illustration B shows a pseudo
Figure 26.7 Glial cells support neurons and maintain their environment. Glial cells of the (a) CNS include oligodendrocytes, astrocytes, ependymal cells, and microglial cells. Oligodendrocytes form the myelin sheath around axons. Astrocytes provide nutrients to neurons, maintain their extracellular environment, and provide structural support. Microglia scavenge pathogens and dead cells. Ependymal cells produce cerebrospinal fluid that cushions the neurons. Glial cells of the (b) PNS include Schwann cells, which form the myelin sheath, and satellite cells, which provide nutrients and structural support to neurons.
Astrocytes, fluorescently labeled green, are irregularly shaped with long extensions that provide support to nerve cells. Oligodendrocytes, also labeled green, are round with long, branched extensions that form the myelin sheath of nerve cells.
Figure 26.8 (a) Astrocytes and (b) oligodendrocytes are glial cells of the CNS. (credit a: modification of work by Uniformed Services University; credit b: modification of work by Jurjen Broeke; scale-bar data from Matt Russell)

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