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The invention of the microscope opened up a previously unknown world. Before the invention of the microscope,
very little was known about what made up living things and non-living things, or where living things came from.
During the discovery of cells, spontaneous generation the belief that living organisms grow directly from
decaying organic substances was the accepted explanation for the appearance of small organisms. For example,
people accepted that mice spontaneously appeared in stored grain, and maggots formed in meat with no apparent
external influence. Once cells were discovered, the search for answers to such questions as "What are cells made
of?" and "What is the function of cells?" became the focus of study.
Cell Function
Cells share the same needs: the need to get energy from their environment, the need to respond to their environment,
and the need to reproduce. Cells must also be able to separate their relatively stable interior from the ever-changing
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FIGURE 3.14
The structure and contents of a typical
animal cell. Every animal cell has a cell
membrane, cytoplasm, ribosomes, and a
nucleus, but not all cells have every structure shown here. For example, some cells
such as red blood cells do not have any
mitochondria, yet others such as muscle
cells may have thousands of mitochondria.
external environment. They do this by coordinating many processes that are carried out within organelles, or other
cellular structures. Structures that are common to many different cells indicate the common history shared by cellbased life. Examples of these common structures include the components of both the cell (or plasma) membrane
and the cytoskeleton, and other structures shown in Figure 3.14.
Is there a relationship between the cell structure and its function? Of course there is. The structure-function
relationship describes a pattern evident throughout biological systems. This relationship is evident in proteins
(protein structure determines its function), nucleic acids (nucleic acid structure results in a genetic code), anatomy
(longer necked giraffes are more functional than short neck giraffes), as well as cells. Using the human body as an
example, specialized cells perform many diverse functions, from digestion and excretion to message transmission
and oxygen distribution. The structure of each type of human cell depends on what function it will perform. This
structure-function relationship can be extended to all other organisms, from the largest whale to the smallest bacteria.
The variability between cell function is related to the proteins expressed in a particular type of cell. For example,
though they do have many proteins in common, a neuron is going to use select different proteins than muscle cell.
A direct relationship exists between the proteins expressed, the size and shape of every cell and the tasks it needs to
accomplish. Examples can easily be seen in red blood cells, neurons muscle cells and sperm cells.
Red blood cells are flat, round, and very small. Their small size allows easy maneuverability through the
capillaries, the narrowest blood vessels, where oxygen is transferred into body cells.
Neurons have a long, thin cellular extension, allowing for very quick and accurate communication and
responses. The long length allows a neuron to send electrical messages extremely quickly.
Skeletal muscle cells have an arrangement of linear protein fibers. The elongated shape allows for muscle
contraction.
Sperm cells are the only human cell with flagella. This is because of their need to "swim" long distances to
reach an egg for fertilization.
Vocabulary
flagella (singular, flagellum): A tail-like appendage that protrudes from the cell body of certain prokaryotic
and eukaryotic cells; used for locomotion.
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neuron: An electrically excitable cell that processes and transmits information by electrical and chemical
signaling; a nerve cell.
organelle: A structure within the cytoplasm of a cell; may be enclosed within a membrane; performs a specific
function.
spontaneous generation: An obsolete principle regarding the origin of life from inanimate matter.
structure-function relationship: Principle that states the function of a biological item (molecule, protein,
cell) is determined by its structure.
Summary
A cells function is usually directly related to its structure; this is known as the structure-function relationship.
The structure-function relationship is evident throughout biology.
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All cells have a plasma membrane. This membrane surrounds the cell. So what is its role?
Can molecules enter and leave the cell? Yes. Can anything or everything enter or leave? No. So, what determines
what can go in or out? Is it the nucleus? The DNA? Or the plasma membrane?
Plasma Membrane
The plasma membrane (also called the cell membrane) is a lipid bilayer that is common to all living cells. Its
function is to keep the cell as a distinct entity in a water-based environment. A phospholipid bilayer is a double
layer of closely-packed phospholipid molecules. It is this orientation of the phospholipids into the bilayer that
biochemically gives the membrane its specific functional characteristics.
Organelle membranes are also composed of phospholipids. For example, mitochondria are bounded by a double
membrane. Each membrane has a phospholipid bilayer with embedded proteins. This division from the rest of the
cell makes the mitochondria only partially dependent on the cell. More on the structure of the phospholipid bilayer
will be presented in the The Plasma Membrane: The Phospholipid Bilayer (Advanced) concept.
Along the phospholipid bilayer, numerous proteins are embedded within the membrane. This structure is called the
Fluid Mosaic Model which will be discussed in the The Plasma Membrane: The Fluid Mosaic Model (Advanced)
concept. These proteins have a variety of important roles; hormone binding sites, electron carriers, pumps for active
transport, channels for passive transport, enzymes, cell signaling and cell adhesion.
The plasma membranes allows only certain molecules, such as ions and small organic molecules, into and out of
the cell. The ability to allow only certain molecules in or out of the cell is referred to as selective permeability or
semipermeability. This characteristic helps the cell to regulate its interactions between the internal machinery and
the external surroundings, helping to maintain homeostasis.
The plasma membrane also acts as the attachment point for both the intracellular cytoskeleton and, if present, the
cell wall.
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Vocabulary
cell membrane: Thin coat of lipids (phospholipids) that surrounds and encloses a cell; physical boundary
between the intracellular space and the extracellular environment; also called the plasma membrane.
Fluid Mosaic Model: Model of the plasma membrane; proposes that the membrane behaves like a fluid with
an embedded mosaic of proteins.
phospholipid bilayer: A bilayer (2 layers) of phospholipids that surrounds and encloses a cell; physical
boundary between the intracellular space and the extracellular environment.
plasma membrane: Thin coat of lipids (phospholipids) that surrounds and encloses a cell; physical boundary
between the intracellular space and the extracellular environment; also called the cell membrane.
selective permeability: The ability to allow only certain molecules in or out of the cell; characteristic of the
cell membrane; also called semipermeability.
semipermeability: The ability to allow only certain molecules in or out of the cell; characteristic of the cell
membrane; also called selective permeability.
Summary
The plasma membrane forms a barrier between the cytoplasm and the environment outside the cell.
A main characteristic of the plasma membrane is selective permeability.
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Why a bilayer?
Whats on the inside of the cell and on the outside? Mostly water. As you can see here, the water-based interior of
the cell has lots of components. These need to be kept inside the cell. And it is the nature of the phospholipid bilayer
to keep the inside of the cell separate from the outside.
Phospholipids
The cell membrane (or plasma membrane) is composed mainly of phospholipids with embedded proteins. The
membrane is a lipid bilayer, with the phospholipids oriented in a distinct manner to provide qualities necessary to
maintain a cell in a water-based environment.
A phospholipid is made up of a polar, phosphorus-containing head, and two long fatty acid (hydrocarbon), non-polar
"tails." That is, the head of the molecule is hydrophilic (water-loving), and the tail is hydrophobic (water-fearing).
Cytosol and extracellular fluid - the insides and outsides of the cell - are made up of mostly water. In this watery
environment, the water loving heads point out towards the water, and the water fearing tails point inwards, and push
the water out. The resulting double layer is called a phospholipid bilayer. A phospholipid bilayer is made up of
two layers of phospholipids, in which hydrophobic fatty acids are in the middle of the plasma membrane, and the
hydrophilic heads are on the outside. An example of a simple phospholipid bilayer is illustrated in Figure 3.15.
The cell membrane also decides what may enter or leave a cell. The membrane is said to be semipermeable
or selectively permeable, allowing only certain ions and organic molecules to cross the membrane. The plasma
membrane contain many proteins, as well as other lipids called sterols. The proteins have various functions, such as
channels (channel proteins) that allow certain molecules into the cell, and receptors (receptor proteins) that bind to
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The phospholipid
bilayer consists of two layers of phospholipids, with a hydrophobic, or waterhating, interior and a hydrophilic, or waterloving, exterior. The hydrophilic (polar)
head group and hydrophobic tails (fatty
acid chains) are depicted in the single
phospholipid molecule. The polar head
group and fatty acid chains are attached
by a 3-carbon glycerol unit.
The hy-
The
signal molecules. In Figure 3.15, the smaller (green) molecules shown between the phospholipids are cholesterol
molecules. Cholesterol helps keep the plasma membrane firm and stable over a wide range of temperatures. At least
ten different types of lipids are commonly found in plasma membranes. Each type of cell or organelle will have a
different percentage of each lipid, protein and carbohydrate.
Vocabulary
cholesterol: A steroid alcohol that is present in animal cells and body fluids, regulates membrane fluidity, and
functions as a precursor molecule in various metabolic pathways.
hydrophilic: Characteristic of the phospholipid head group; water-loving.
hydrophobic: Characteristic of the phospholipid tails; water-hating.
phospholipid: A major component of the cell membrane; consists of two hydrophobic tails and a hydrophilic
phosphate head group.
phospholipid bilayer: A bilayer (2 layers) of phospholipids that surrounds and encloses a cell; physical
boundary between the intracellular space and the extracellular environment.
plasma membrane: Thin coat of lipids (phospholipids) that surrounds and encloses a cell; physical boundary
between the intracellular space and the extracellular environment; also called the cell membrane.
Summary
A phospholipid is a lipid molecule with a polar head group ( a phosphate group) and two non-polar hydrocarbon tails.
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The plasma membrane is a selectively permeable lipid bilayer that contains mostly lipids and proteins. These
lipids and proteins are involved in many cellular processes.
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1. Why can hydrophobic (water-hating) molecules easily cross the plasma membrane, while hydrophilic (waterloving) molecules cannot?
2. Describe the composition of the plasma membrane.
3. Describe the orientation of the phospholipids in the cell membrane.
4. What is the role of cholesterol in the plasma membrane?
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Membrane Proteins
The second main component of plasma membranes are the variety of proteins. A membrane protein is a protein
molecule that is attached to, or associated with the membrane of a cell or an organelle. Membrane proteins can be
put into two groups based on how the protein is associated with the membrane: (1) integral membrane proteins and
(2) peripheral membrane proteins.
Integral membrane proteins, also called intrinsic proteins, are permanently embedded within the plasma membrane. Structurally, the integral proteins contain residues with hydrophobic side chains that penetrate the fatty acyl
regions of the phospholipid bilayer, thus anchoring the protein to the membrane. The only way to remove the integral
proteins from the membrane are with synthetic detergents, nonpolar solvents and denaturing agents that disrupt the
hydrophobic interactions of the bilayer.
Integral membrane proteins can be classified according to their relationship with the bilayer:
Transmembrane proteins span the entire plasma membrane. Their function is mainly to regulate the transport
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of specific molecules across the membrane. There are two basic types of transmembrane proteins, alphahelical and beta-barrels, which are discussed in Organic Compounds: Proteins (Advanced).
Integral monotopic proteins are permanently attached to the membrane from only one side.
Examples of integral membrane proteins and their functions are:
1. glycoprotein (cell-to-cell interactions)
2. Na+/K+ ATPase (responsible for establishing and maintaining the electrochemical gradients of Sodium and
Potassium ions across the plasma membrane)
3. glucose permease (the reversible transporter protein of glucose)
4. ion channels gates (the flow of ions across the cell membrane)
5. gap junction proteins (a direct connection between the cytoplasm of two cells, which allows various molecules
and ions to pass freely between cells)
6. Bacterial rhodopsins (a protein in archaeans that uses the energy of light to pump protons across the membrane)
Peripheral membrane proteins, also called extrinsic proteins, are only temporarily associated with the membrane.
Most peripheral membrane proteins are hydrophilic so usually they are either attached to integral membrane proteins,
or they can directly bound to a polar head group of the bilayer. This way they can be easily removed, which allows
them to be involved in cell signaling. Peripheral membrane proteins are often associated with ion channels and
transmembrane receptors.
Examples of peripheral membrane proteins and their functions are:
1. spectrin (links the plasma membrane to the actin cytoskeleton for determination of cell shape, arrangement of
transmembrane proteins, and organization of organelles)
2. Kinase C (enzyme that helps mediate signal transduction cascades by hydrolyzing lipids)
3. phospholipases (hydrolyze various bonds in the the polar head group of phospholipids which are vital to the
degredation of damaged or aged cell membranes)
4. hormone receptors (binds a hormone outside the cell membrane and activates a protein kinase inside the cell)
Glycoproteins and glycolipids, in particular, have a carbohydrate chain that acts as a label to identify the cell
type. Specifically, A, B, O blood groups result from having different carbohydrate chains on the cell surface of
red blood cells and other types of cells. Everyone has glycolipids and glycoproteins with the particular type of
carbohydrate chain that signals type O. However, people with type A also have an additional carbohydrate called
N-Acetylgalactosamine and those with type B have an added galactose. Those with type AB have some glycolipids
and glycoproteins with N-Acetylgalactosamine added and others with galactose added.
Shown in Figure 3.16 are two different types of membrane proteins and associated molecules.
Vocabulary
integral membrane proteins: Proteins that are permanently embedded within the plasma membrane of a cell
or organelle.
membrane protein: A protein molecule that is attached to, or associated with, the membrane of a cell or
organelle.
peripheral membrane proteins: Proteins that are only temporarily associated with the cell membrane; can
be easily removed.
plasma membrane: Thin coat of lipids (phospholipids) that surrounds and encloses a cell; physical boundary
between the intracellular space and the extracellular environment; also called the cell membrane.
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FIGURE 3.16
Some of the membrane proteins make up a major transport system that moves molecules and ions through the
polar phospholipid bilayer.
Summary
The plasma membrane has many proteins that assist other substances in crossing the membrane.
Membrane proteins may be permanently attached/embedded (integral membrane proteins) to the membrane,
or just temporarily associated with the membrane (peripheral membrane proteins).
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1. What are the main differences between the types of proteins associated with the plasma membrane?
2. Name three membrane protein functions.
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In 1972, S. J. Singer and G. L. Nicolson proposed the now widely accepted Fluid Mosaic Model of the structure
of cell membranes (Science, 175: 720-731). Remember, it is the cell membrane that keeps the cells internal
environment separate from its surroundings, but it is also this membrane that constantly and consistently allows
the cell to interact and exchange materials with its environment.
The Fluid Mosaic Model proposes that integral membrane proteins are embedded in the phospholipid bilayer, as
seen in the opening image. The bilayer results from the chemical nature of the phospholipids in a polar environment.
The phospholipids create a double layer - or bilayer - when placed in a polar environment like water. Some of these
proteins associated with the membrane extend all the way through the bilayer, and some only partially across it. In
this model, the integral membrane proteins have their polar groups protruding from the membrane into the aqueous
environment, while the non polar regions of the protein are buried within the hydrophobic interior of the membrane.
This model also proposed that the membrane behaves like a fluid. Scanning electron microscope images demonstrated that the embedded molecules can move sideways throughout the membrane, meaning the membrane is not
solid, but more like a fluid. The membrane proteins and lipids of the membrane can move laterally around the
membrane, much like buoys in water, or sideways throughout the membrane. Such movement causes a constant
change in the "mosaic pattern" of the plasma membrane. The mosaic pattern results from the many different components of the bilayer. These components include the phospholipids, integral and peripheral proteins, glycoproteins
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and glycolipids, which aid in their location and identification of food, water, waste, and other membrane traffic.
Each cell has a particular glycoprotein structure extruding from the cell membrane, based on its need to attract or
repel membrane traffic. The cell is constantly interacting with its environment, bring certain molecules such as ions,
hormones and food into the cell, and exporting materials, such as wastes, out of the cell.
A further description of the fluid mosaic model can be viewed in Fluid Mosaic Model of the Cell Membrane at http
://www.youtube.com/watch?v=LKN5sq5dtW4 (1:27).
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Vocabulary
Fluid Mosaic Model: Model of the plasma membrane; proposes that the membrane behaves like a fluid with
an embedded mosaic of proteins.
integral membrane proteins: Proteins that are permanently embedded within the plasma membrane of a cell
or organelle.
phospholipid bilayer: A bilayer (2 layers) of phospholipids that surrounds and encloses a cell; physical
boundary between the intracellular space and the extracellular environment.
Summary
The Fluid Mosaic Model depicts the biological nature of the plasma membrane, with a fluid phospholipid
bilayer and a mosaic of proteins.
Review
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Cytoplasm is one component of cells that is common to all cells. Cytoplasm is the gel-like material between the cell
membrane and the nucleus. The cytoplasm plays an important role in a cell, serving as a "jelly" in which organelles
are suspended and held together by the cell membrane. Though prokaryotic cells do not have organelles (though
they do have ribosomes), they still have cytoplasm. It is within the cytoplasm that most cellular activities occur,
including the many metabolic pathways that occur within organelles, such as photosynthesis and aerobic respiration.
The cytosol, which is the watery substance that does not contain organelles, is made up of 80% to 90% water. The
cytosol plays a mechanical role by exerting pressure against the cells plasma membrane. This helps keep the shape
of the cell. Cytosol also acts as the site of biochemical reactions such as anaerobic glycolysis and protein synthesis.
In prokaryotes all chemical reactions take place in the cytosol.
Cytoskeleton
The cytoskeleton is a cellular "scaffolding" or "skeleton" that crisscrosses the cytoplasm. All eukaryotic cells
have a cytoskeleton, and recent research has shown that prokaryotic cells also have a cytoskeleton. The eukaryotic
cytoskeleton is made up of a network of long, thin protein fibers and has many functions. It helps to maintain cell
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shape, holds organelles in place, and for some cells, it enables cell movement. The cytoskeleton plays important roles
in both the intracellular movement of substances and in cell division. Certain proteins act like a path that vesicles and
organelles move along within the cell. The threadlike proteins that make up the cytoskeleton continually rebuild to
adapt to the cells constantly changing needs. Three main kinds of cytoskeleton fibers are microtubules, intermediate
filaments, and microfilaments.
Microtubules, shown as (a) in Figure 3.17, are hollow cylinders and are the thickest of the cytoskeleton
structures. They are most commonly made of filaments which are polymers of alpha and beta tubulin, and
radiate outwards from an area near the nucleus called the centrosome. Tubulin is the protein that forms
microtubules. Two forms of tubulin, alpha and beta, form dimers (pairs) which come together to form the
hollow cylinders. The cylinders are twisted around each other to form the microtubules. Microtubules help
the cell keep its shape. They hold organelles in place and allow them to move around the cell, and they form
the mitotic spindle during cell division. Microtubules also make up parts of cilia and flagella, the organelles
that help a cell move.
Microfilaments, shown as (b) in Figure 3.17, are made of two thin actin chains that are twisted around
one another. Microfilaments are mostly concentrated just beneath the cell membrane, where they support the
cell and help the cell keep its shape. Microfilaments form cytoplasmatic extentions, such as pseudopodia
and microvilli, which allow certain cells to move. The actin of the microfilaments interacts with the protein
myosin to cause contraction in muscle cells. Microfilaments are found in almost every cell, and are numerous
in muscle cells and in cells that move by changing shape, such as phagocytes (white blood cells that search
the body for bacteria and other invaders).
Intermediate filaments differ in make-up from one cell type to another. Intermediate filaments organize the
inside structure of the cell by holding organelles and providing strength. They are also structural components
of the nuclear envelope. Intermediate filaments made of the protein keratin are found in skin, hair, and nails
cells.
FIGURE 3.17
(a) The eukaryotic cytoskeleton. Microfilaments are shown in red, microtubules
in green, and the nuclei are in blue. By
linking regions of the cell together, the
cytoskeleton helps support the shape of
the cell. (b) Microscopy of microfilaments
(actin filaments), shown in green, inside
cells. The nucleus is shown in blue.
Shape
Microtubules
About 25 nm
Tubulin, with two subunits, alpha and beta tubulin
Hollow cylinders made of
two protein chains twisted
around each other
Intermediate Filaments
8 to 11 nm
One of different types of
proteins such as lamin, vimentin, and keratin
Protein fiber coils twisted
into each other
Microfilaments
Around 7 nm
Actin
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Microtubules
Organelle and vesicle
movement; form mitotic
spindles
during
cell
reproduction;
cell
motility (in cilia and
flagella)
Intermediate Filaments
Organize cell shape; positions organelles in cytoplasm structural support
of the nuclear envelope
and sarcomeres; involved
in cell-to-cell and cell-tomatrix junctions
Microfilaments
Keep cellular shape; allows movement of certain
cells by forming cytoplasmatic extensions or contraction of actin fibers; involved in some cell-tocell or cell-to-matrix junctions
Representation
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Vocabulary
actin: A thin, threadlike protein filament found in muscle; the protein component of microfilaments.
cytoplasm: The gel-like material inside the plasma membrane of a cell; holds the cells organelles (excluding
the nucleus).
cytoskeleton: The structure of filaments and tubules in the cytoplasm; provides a cell with an internal
framework.
cytosol: A watery cytoplasmic fluid that contains cytoskeletal fragments, dissolved particles and organelles.
intermediate filaments: Intermediate component of the cytoskeleton; made of protein fiber coils twisted into
each other.
microfilaments: Smallest component of the cytoskeleton; made of two actin chains twisted around one
another.
microtubules: Largest component of the cytoskeleton; hollow protein cylinders made of alpha and beta
tubulin; also found in flagella.
microvilli: Cellular membrane protrusions that increase the surface area of cells.
tubulin: Protein component of microtubules; alpha-tubulin and beta-tubulin combine to form components of
microtubules.
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Summary
The cytoplasm consists of everything between the plasma membrane of the cell and the nucleus (of an
eukaryotic cell).
The cytoskeleton is a cellular "skeleton" that crisscrosses the cytoplasm. Three main cytoskeleton fibers are
microtubules, intermediate filaments, and microfilaments.
Microtubules are the thickest of the cytoskeleton structures and are most commonly made of filaments which
are polymers of alpha and beta tubulin.
Microfilament are the thinnest of the cytoskeleton structures and are made of two thin actin chains that are
twisted around one another.
Review
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Flagella ( flagellum, singular) are long, thin structures that protrude from the cell membrane. Both eukaryotic and
prokaryotic cells can have flagella. Flagella help single-celled organisms move or swim towards food. The flagella
of eukaryotic cells are normally used for movement too, such as in the movement of sperm cells, which have only
a single flagellum. The flagella of either group are very different from each other. Prokaryotic flagella, shown in
Figure 3.18, are spiral-shaped and stiff. They spin around in a fixed base much like a screw does, which moves the
cell in a tumbling fashion. Eukaryotic flagella are made of microtubules that bend and flex like a whip.
Cilia ( cilium, singular) are made up of microtubule containing extensions of the cell membrane. Although both
cilia and flagella are used for movement, cilia are much shorter than flagella. Cilia cover the surface of some singlecelled organisms, such as paramecium. Their cilia beat together to move the little animal-like protists through the
water. In multicellular animals, including humans, cilia are usually found in large numbers on a single surface of
cells. Multicellular animals cilia usually move materials inside the body. For example, the mucociliary escalator of
the respiratory system is made up of mucus-secreting ciliated cells that line the trachea and bronchi. These ciliated
cells, shown in Figure 3.19, move mucus away from the lungs. This mucus catches spores, bacteria, and debris and
moves to the esophagus, where it is swallowed.
A video showing flagella and cilia can be viewed at http://www.youtube.com/watch?v=QGAm6hMysTA (3:12).
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FIGURE 3.18
Bacterial flagella spin about in place,
which causes the bacterial cell to "tumble."
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FIGURE 3.19
Left:
Vocabulary
cilia (singular, cilium): Short, hairlike projection, similar to flagella, that allows some cells to move.
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flagella (singular, flagellum): A "tail-like" appendage that protrudes from the cell body of certain prokaryotic
and eukaryotic cells; used for locomotion.
microtubules: Largest component of the cytoskeleton; hollow protein cylinders made of alpha and beta
tubulin; also found in flagella.
Summary
Cilia and flagella are extensions of the cell membrane that contain microtubules, and are usually used for
movement.
Cilia cover the surface of some single-celled animals, such as paramecium, but cover only one side of cells in
some multicellular organisms.
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Review
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