CHAPTER 4

Cell Structure

Figure 4.1 (a) Nasal sinus cells (viewed with a light microscope), (b) onion cells (viewed with a
light microscope), and (c) Vibrio tasmaniensis bacterial cells (seen through a scanning electron
microscope) are from very different organisms, yet all share certain basic cell structure
characteristics. (credit a: modification of work by Ed Uthman, MD; credit b: modification of work by
Umberto Salvagnin; credit c: modification of work by Anthony D'Onofrio, William H. Fowle, Eric J.
Stewart, and Kim Lewis of the Lewis Lab at Northeastern University; scale-bar data from Matt
Russell)

INTRODUCTION Close your eyes and picture a brick wall. What is the wall's basic building block? It is a
single brick. Like a brick wall, cells are the building blocks that make up your body.
Chapter Outline
Your body has many kinds of cells, each specialized for a specific purpose. Just as we use a variety of
materials to build a home, the human body is constructed from many cell types. For example, epithelial 4.1 Studying Cells
cells protect the body's surface and cover the organs and body cavities within. Bone cells help to support 4.2 Prokaryotic Cells
and protect the body. Immune system cells fight invading bacteria. Additionally, blood and blood cells
carry nutrients and oxygen throughout the body while removing carbon dioxide. Each of these cell types 4.3 Eukaryotic Cells
plays a vital role during the body's growth, development, and day-to-day maintenance. In spite of their 4.4 The Endomembrane
enormous variety, however, cells from all organisms—even ones as diverse as bacteria, onion, and System and Proteins
human—share certain fundamental characteristics.
4.5 The Cytoskeleton

4.6 Connections
between Cells and
Cellular Activities

4.1 Studying Cells

By the end of this section, you will be able to do the following:
• Describe the role of cells in organisms
• Compare and contrast light microscopy and electron microscopy
• Summarize cell theory

A cell is the smallest unit of a living thing. Whether comprised of one cell (like bacteria) or many cells (like a
96 Chapter 4 • Cell Structure

human), we call it an organism. Thus, cells are the basic building blocks of all organisms.

Several cells of one kind that interconnect with each other and perform a shared function form tissues.
These tissues combine to form an organ (your stomach, heart, or brain), and several organs comprise an
organ system (such as the digestive system, circulatory system, or nervous system). Several systems that
function together form an organism (like a human being). Here, we will examine the structure and function
of cells.

There are many types of cells, which scientists group into one of two broad categories: prokaryotic and
eukaryotic. For example, we classify both animal and plant cells as eukaryotic cells; whereas, we classify
bacterial cells as prokaryotic. Before discussing the criteria for determining whether a cell is prokaryotic or
eukaryotic, we will first examine how biologists study cells.

Microscopy
Cells vary in size. With few exceptions, we cannot see individual cells with the naked eye, so scientists use
microscopes (micro- = “small”; -scope = “to look at”) to study them. A microscope is an instrument that
magnifies an object. We photograph most cells with a microscope, so we can call these images micrographs.

The optics of a microscope’s lenses change the image orientation that the user sees. A specimen that is right-
side up and facing right on the microscope slide will appear upside-down and facing left when one views
through a microscope, and vice versa. Similarly, if one moves the slide left while looking through the
microscope, it will appear to move right, and if one moves it down, it will seem to move up. This occurs
because microscopes use two sets of lenses to magnify the image. Because of the manner by which light
travels through the lenses, this two lens system produces an inverted image (binocular, or dissecting
microscopes, work in a similar manner, but include an additional magnification system that makes the final
image appear to be upright).

Light Microscopes
To give you a sense of cell size, a typical human red blood cell is about eight millionths of a meter or eight
micrometers (abbreviated as eight μm) in diameter. A pin head is about two thousandths of a meter (two
mm) in diameter. That means about 250 red blood cells could fit on a pinhead.

Most student microscopes are light microscopes (Figure 4.2a). Visible light passes and bends through the
lens system to enable the user to see the specimen. Light microscopes are advantageous for viewing living
organisms, but since individual cells are generally transparent, their components are not distinguishable
unless they are colored with special stains. Staining, however, usually kills the cells.

Light microscopes that undergraduates commonly use in the laboratory magnify up to approximately 400
times. Two parameters that are important in microscopy are magnification and resolving power.
Magnification is the process of enlarging an object in appearance. Resolving power is the microscope's
ability to distinguish two adjacent structures as separate: the higher the resolution, the better the image's
clarity and detail. When one uses oil immersion lenses to study small objects, magnification usually
increases to 1,000 times. In order to gain a better understanding of cellular structure and function,
scientists typically use electron microscopes.

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 4.1 • Studying Cells 97

Figure 4.2 (a) Most light microscopes in a college biology lab can magnify cells up to approximately 400 times and have a resolution of
about 200 nanometers. (b) Electron microscopes provide a much higher magnification, 100,000x, and a have a resolution of 50 picometers.
(credit a: modification of work by "GcG"/Wikimedia Commons; credit b: modification of work by Evan Bench)

Electron Microscopes
In contrast to light microscopes, electron microscopes (Figure 4.2b) use a beam of electrons instead of a beam of light. Not only
does this allow for higher magnification and, thus, more detail (Figure 4.3), it also provides higher resolving power. The method
to prepare the specimen for viewing with an electron microscope kills the specimen. Electrons have short wavelengths (shorter
than photons) that move best in a vacuum, so we cannot view living cells with an electron microscope.

In a scanning electron microscope, a beam of electrons moves back and forth across a cell’s surface, creating details of cell
surface characteristics. In a transmission electron microscope, the electron beam penetrates the cell and provides details of a
cell’s internal structures. As you might imagine, electron microscopes are significantly more bulky and expensive than light
microscopes.

Figure 4.3 (a) These Salmonella bacteria appear as tiny purple dots when viewed with a light microscope. (b) This scanning electron
microscope micrograph shows Salmonella bacteria (in red) invading human cells (yellow). Even though subfigure (b) shows a different
Salmonella specimen than subfigure (a), you can still observe the comparative increase in magnification and detail. (credit a: modification
of work by CDC/Armed Forces Institute of Pathology, Charles N. Farmer, Rocky Mountain Laboratories; credit b: modification of work by
NIAID, NIH; scale-bar data from Matt Russell)

LINK TO LEARNING
For another perspective on cell size, try the HowBig interactive at this site (http://openstax.org/l/cell_sizes) .

Cell Theory
The microscopes we use today are far more complex than those that Dutch shopkeeper Antony van Leeuwenhoek, used in the
1600s. Skilled in crafting lenses, van Leeuwenhoek observed the movements of single-celled organisms, which he collectively
termed “animalcules.”

In the 1665 publication Micrographia, experimental scientist Robert Hooke coined the term “cell” for the box-like structures he
observed when viewing cork tissue through a lens. In the 1670s, van Leeuwenhoek discovered bacteria and protozoa. Later
advances in lenses, microscope construction, and staining techniques enabled other scientists to see some components inside
cells.
98 Chapter 4 • Cell Structure

By the late 1830s, botanist Matthias Schleiden and zoologist Theodor Schwann were studying tissues and proposed the unified
cell theory, which states that one or more cells comprise all living things, the cell is the basic unit of life, and new cells arise from
existing cells. Rudolf Virchow later made important contributions to this theory.

CAREER CONNECTION

Cytotechnologist
Have you ever heard of a medical test called a Pap smear (Figure 4.4)? In this test, a doctor takes a small sample of cells from the
patient's uterine cervix and sends it to a medical lab where a cytotechnologist stains the cells and examines them for any
changes that could indicate cervical cancer or a microbial infection.

Cytotechnologists (cyto- = “cell”) are professionals who study cells via microscopic examinations and other laboratory tests. They
are trained to determine which cellular changes are within normal limits and which are abnormal. Their focus is not limited to
cervical cells. They study cellular specimens that come from all organs. When they notice abnormalities, they consult a
pathologist, a medical doctor who interprets and diagnoses changes that disease in body tissue and fluids cause.

Cytotechnologists play a vital role in saving people’s lives. When doctors discover abnormalities early, a patient’s treatment can
begin sooner, which usually increases the chances of a successful outcome.

Figure 4.4 These uterine cervix cells, viewed through a light microscope, are from a Pap smear. Normal cells are on the left. The cells on the
right are infected with human papillomavirus (HPV). Notice that the infected cells are larger. Also, two of these cells each have two nuclei
instead of one, the normal number. (credit: modification of work by Ed Uthman, MD; scale-bar data from Matt Russell)

4.2 Prokaryotic Cells

By the end of this section, you will be able to do the following:
• Name examples of prokaryotic and eukaryotic organisms
• Compare and contrast prokaryotic and eukaryotic cells
• Describe the relative sizes of different cells
• Explain why cells must be small

Cells fall into one of two broad categories: prokaryotic and eukaryotic. We classify only the predominantly single-celled
organisms Bacteria and Archaea as prokaryotes (pro- = “before”; -kary- = “nucleus”). Animal cells, plants, fungi, and protists are
all eukaryotes (eu- = “true”).

Components of Prokaryotic Cells

All cells share four common components: 1) a plasma membrane, an outer covering that separates the cell’s interior from its
surrounding environment; 2) cytoplasm, consisting of a jelly-like cytosol within the cell in which there are other cellular
components; 3) DNA, the cell's genetic material; and 4) ribosomes, which synthesize proteins. However, prokaryotes differ from
eukaryotic cells in several ways.

A prokaryote is a simple, mostly single-celled (unicellular) organism that lacks a nucleus, or any other membrane-bound

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 4.2 • Prokaryotic Cells 99

organelle. We will shortly come to see that this is significantly different in eukaryotes. Prokaryotic DNA is in the cell's central
part: the nucleoid (Figure 4.5).

Figure 4.5 This figure shows the generalized structure of a prokaryotic cell. All prokaryotes have chromosomal DNA localized in a nucleoid,
ribosomes, a cell membrane, and a cell wall. The other structures shown are present in some, but not all, bacteria.

Most prokaryotes have a peptidoglycan cell wall and many have a polysaccharide capsule (Figure 4.5). The cell wall acts as an
extra layer of protection, helps the cell maintain its shape, and prevents dehydration. The capsule enables the cell to attach to
surfaces in its environment. Some prokaryotes have flagella, pili, or fimbriae. Flagella are used for locomotion. Pili exchange
genetic material during conjugation, the process by which one bacterium transfers genetic material to another through direct
contact. Bacteria use fimbriae to attach to a host cell.

CAREER CONNECTION

Microbiologist
The most effective action anyone can take to prevent the spread of contagious illnesses is to wash his or her hands. Why?
Because microbes (organisms so tiny that they can only be seen with microscopes) are ubiquitous. They live on doorknobs,
money, your hands, and many other surfaces. If someone sneezes into his hand and touches a doorknob, and afterwards you
touch that same doorknob, the microbes from the sneezer’s mucus are now on your hands. If you touch your hands to your
mouth, nose, or eyes, those microbes can enter your body and could make you sick.

However, not all microbes (also called microorganisms) cause disease; most are actually beneficial. You have microbes in your
gut that make vitamin K. Other microorganisms are used to ferment beer and wine.

Microbiologists are scientists who study microbes. Microbiologists can pursue a number of careers. Not only do they work in the
food industry, they are also employed in the veterinary and medical fields. They can work in the pharmaceutical sector, serving
key roles in research and development by identifying new antibiotic sources that can treat bacterial infections.

Environmental microbiologists may look for new ways to use specially selected or genetically engineered microbes to remove
pollutants from soil or groundwater, as well as hazardous elements from contaminated sites. We call using these microbes
bioremediation technologies. Microbiologists can also work in the bioinformatics field, providing specialized knowledge and
insight for designing, developing, and specificity of computer models of, for example, bacterial epidemics.

Cell Size
At 0.1 to 5.0 μm in diameter, prokaryotic cells are significantly smaller than eukaryotic cells, which have diameters ranging from
10 to 100 μm (Figure 4.6). The prokaryotes' small size allows ions and organic molecules that enter them to quickly diffuse to
other parts of the cell. Similarly, any wastes produced within a prokaryotic cell can quickly diffuse. This is not the case in
eukaryotic cells, which have developed different structural adaptations to enhance intracellular transport.
100 Chapter 4 • Cell Structure

Figure 4.6 This figure shows relative sizes of microbes on a logarithmic scale (recall that each unit of increase in a logarithmic scale
represents a 10-fold increase in the quantity measured).

Small size, in general, is necessary for all cells, whether prokaryotic or eukaryotic. Let’s examine why that is so. First, we’ll
consider the area and volume of a typical cell. Not all cells are spherical in shape, but most tend to approximate a sphere. You
may remember from your high school geometry course that the formula for the surface area of a sphere is 4πr2, while the
formula for its volume is 4πr3/3. Thus, as the radius of a cell increases, its surface area increases as the square of its radius, but
its volume increases as the cube of its radius (much more rapidly). Therefore, as a cell increases in size, its surface area-to-
volume ratio decreases. This same principle would apply if the cell had a cube shape (Figure 4.7). If the cell grows too large, the
plasma membrane will not have sufficient surface area to support the rate of diffusion required for the increased volume. In
other words, as a cell grows, it becomes less efficient. One way to become more efficient is to divide. Other ways are to increase
surface area by foldings of the cell membrane, become flat or thin and elongated, or develop organelles that perform specific
tasks. These adaptations lead to developing more sophisticated cells, which we call eukaryotic cells.

VISUAL CONNECTION

Figure 4.7 Notice that as a cell increases in size, its surface area-to-volume ratio decreases. When there is insufficient surface area to
support a cell’s increasing volume, a cell will either divide or die. The cell on the left has a volume of 1 mm3 and a surface area of 6 mm2,
with a surface area-to-volume ratio of 6 to 1; whereas, the cell on the right has a volume of 8 mm3 and a surface area of 24 mm2, with a
surface area-to-volume ratio of 3 to 1.

Prokaryotic cells are much smaller than eukaryotic cells. What advantages might small cell size confer on a cell? What
advantages might large cell size have?

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 4.3 • Eukaryotic Cells 101

4.3 Eukaryotic Cells

By the end of this section, you will be able to do the following:
• Describe the structure of eukaryotic cells
• Compare animal cells with plant cells
• State the role of the plasma membrane
• Summarize the functions of the major cell organelles

Have you ever heard the phrase “form follows function?” It’s a philosophy that many industries follow. In architecture, this
means that buildings should be constructed to support the activities that will be carried out inside them. For example, a
skyscraper should include several elevator banks. A hospital should have its emergency room easily accessible.

Our natural world also utilizes the principle of form following function, especially in cell biology, and this will become clear as
we explore eukaryotic cells (Figure 4.8). Unlike prokaryotic cells, eukaryotic cells have: 1) a membrane-bound nucleus; 2)
numerous membrane-bound organelles such as the endoplasmic reticulum, Golgi apparatus, chloroplasts, mitochondria, and
others; and 3) several, rod-shaped chromosomes. Because a membrane surrounds eukaryotic cell’s nucleus, it has a “true
nucleus.” The word “organelle” means “little organ,” and, as we already mentioned, organelles have specialized cellular functions,
just as your body's organs have specialized functions.

At this point, it should be clear to you that eukaryotic cells have a more complex structure than prokaryotic cells. Organelles
allow different functions to be compartmentalized in different areas of the cell. Before turning to organelles, let’s first examine
two important components of the cell: the plasma membrane and the cytoplasm.

VISUAL CONNECTION
102 Chapter 4 • Cell Structure

Figure 4.8 These figures show the major organelles and other cell components of (a) a typical animal cell and (b) a typical eukaryotic plant
cell. The plant cell has a cell wall, chloroplasts, plastids, and a central vacuole—structures not in animal cells. Most cells do not have
lysosomes or centrosomes.

If the nucleolus were not able to carry out its function, what other cellular organelles would be affected?

The Plasma Membrane

Like prokaryotes, eukaryotic cells have a plasma membrane (Figure 4.9), a phospholipid bilayer with embedded proteins that
separates the internal contents of the cell from its surrounding environment. A phospholipid is a lipid molecule with two fatty
acid chains and a phosphate-containing group. The plasma membrane controls the passage of organic molecules, ions, water,
and oxygen into and out of the cell. Wastes (such as carbon dioxide and ammonia) also leave the cell by passing through the
plasma membrane.

Figure 4.9 The eukaryotic plasma membrane is a phospholipid bilayer with proteins and cholesterol embedded in it.

The plasma membranes of cells that specialize in absorption fold into fingerlike projections that we call microvilli (singular =

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 4.3 • Eukaryotic Cells 103

microvillus); (Figure 4.10). Such cells typically line the small intestine, the organ that absorbs nutrients from digested food. This
is an excellent example of form following function. People with celiac disease have an immune response to gluten, which is a
protein in wheat, barley, and rye. The immune response damages microvilli, and thus, afflicted individuals cannot absorb
nutrients. This leads to malnutrition, cramping, and diarrhea. Patients suffering from celiac disease must follow a gluten-free
diet.

Figure 4.10 Microvilli, as they appear on cells lining the small intestine, increase the surface area available for absorption. These microvilli
are only on the area of the plasma membrane that faces the cavity from which substances will be absorbed. (credit "micrograph":
modification of work by Louisa Howard)

The Cytoplasm
The cytoplasm is the cell's entire region between the plasma membrane and the nuclear envelope (a structure we will discuss
shortly). It is comprised of organelles suspended in the gel-like cytosol, the cytoskeleton, and various chemicals (Figure 4.8).
Even though the cytoplasm consists of 70 to 80 percent water, it has a semi-solid consistency, which comes from the proteins
within it. However, proteins are not the only organic molecules in the cytoplasm. Glucose and other simple sugars,
polysaccharides, amino acids, nucleic acids, fatty acids, and derivatives of glycerol are also there. Ions of sodium, potassium,
calcium, and many other elements also dissolve in the cytoplasm. Many metabolic reactions, including protein synthesis, take
place in the cytoplasm.

The Nucleus
Typically, the nucleus is the most prominent organelle in a cell (Figure 4.8). The nucleus (plural = nuclei) houses the cell’s DNA
and directs the synthesis of ribosomes and proteins. Let’s look at it in more detail (Figure 4.11).

Figure 4.11 The nucleus stores chromatin (DNA plus proteins) in a gel-like substance called the nucleoplasm. The nucleolus is a condensed
chromatin region where ribosome synthesis occurs. We call the nucleus' boundary the nuclear envelope. It consists of two phospholipid
bilayers: an outer and an inner membrane. The nuclear membrane is continuous with the endoplasmic reticulum. Nuclear pores allow
substances to enter and exit the nucleus.
104 Chapter 4 • Cell Structure

The Nuclear Envelope

The nuclear envelope is a double-membrane structure that constitutes the nucleus' outermost portion (Figure 4.11). Both the
nuclear envelope's inner and outer membranes are phospholipid bilayers.

The nuclear envelope is punctuated with pores that control the passage of ions, molecules, and RNA between the nucleoplasm
and cytoplasm. The nucleoplasm is the semi-solid fluid inside the nucleus, where we find the chromatin and the nucleolus.

Chromatin and Chromosomes

To understand chromatin, it is helpful to first explore chromosomes, structures within the nucleus that are made up of DNA,
the hereditary material. You may remember that in prokaryotes, DNA is organized into a single circular chromosome. In
eukaryotes, chromosomes are linear structures. Every eukaryotic species has a specific number of chromosomes in the nucleus
of each cell. For example, in humans, the chromosome number is 46, while in fruit flies, it is eight. Chromosomes are only
visible and distinguishable from one another when the cell is getting ready to divide. When the cell is in the growth and
maintenance phases of its life cycle, proteins attach to chromosomes, and they resemble an unwound, jumbled bunch of
threads. We call these unwound protein-chromosome complexes chromatin (Figure 4.12). Chromatin describes the material
that makes up the chromosomes both when condensed and decondensed.

Figure 4.12 (a) This image shows various levels of chromatin's organization (DNA and protein). (b) This image shows paired chromosomes.
(credit b: modification of work by NIH; scale-bar data from Matt Russell)

The Nucleolus
We already know that the nucleus directs the synthesis of ribosomes, but how does it do this? Some chromosomes have sections
of DNA that encode ribosomal RNA. A darkly staining area within the nucleus called the nucleolus (plural = nucleoli) aggregates
the ribosomal RNA with associated proteins to assemble the ribosomal subunits that are then transported out through the pores
in the nuclear envelope to the cytoplasm.

Ribosomes
Ribosomes are the cellular structures responsible for protein synthesis. When we view them through an electron microscope,
ribosomes appear either as clusters (polyribosomes) or single, tiny dots that float freely in the cytoplasm. They may be attached
to the plasma membrane's cytoplasmic side or the endoplasmic reticulum's cytoplasmic side and the nuclear envelope's outer
membrane (Figure 4.8). Electron microscopy shows us that ribosomes, which are large protein and RNA complexes, consist of
two subunits, large and small (Figure 4.13). Ribosomes receive their “orders” for protein synthesis from the nucleus where the
DNA transcribes into messenger RNA (mRNA). The mRNA travels to the ribosomes, which translate the code provided by the
sequence of the nitrogenous bases in the mRNA into a specific order of amino acids in a protein. Amino acids are the building
blocks of proteins.

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 4.3 • Eukaryotic Cells 105

Figure 4.13 A large subunit (top) and a small subunit (bottom) comprise ribosomes. During protein synthesis, ribosomes assemble amino
acids into proteins.

Because protein synthesis is an essential function of all cells (including enzymes, hormones, antibodies, pigments, structural
components, and surface receptors), there are ribosomes in practically every cell. Ribosomes are particularly abundant in cells
that synthesize large amounts of protein. For example, the pancreas is responsible for creating several digestive enzymes and
the cells that produce these enzymes contain many ribosomes. Thus, we see another example of form following function.

Mitochondria
Scientists often call mitochondria (singular = mitochondrion) “powerhouses” or “energy factories” of both plant and animal cells
because they are responsible for making adenosine triphosphate (ATP), the cell’s main energy-carrying molecule. ATP
represents the cell's short-term stored energy. Cellular respiration is the process of making ATP using the chemical energy in
glucose and other nutrients. In mitochondria, this process uses oxygen and produces carbon dioxide as a waste product. In fact,
the carbon dioxide that you exhale with every breath comes from the cellular reactions that produce carbon dioxide as a
byproduct.

In keeping with our theme of form following function, it is important to point out that muscle cells have a very high
concentration of mitochondria that produce ATP. Your muscle cells need considerable energy to keep your body moving. When
your cells don’t get enough oxygen, they do not make much ATP. Instead, producing lactic acid accompanies the small amount of
ATP they make in the absence of oxygen.

Mitochondria are oval-shaped, double membrane organelles (Figure 4.14) that have their own ribosomes and DNA. Each
membrane is a phospholipid bilayer embedded with proteins. The inner layer has folds called cristae. We call the area
surrounded by the folds the mitochondrial matrix. The cristae and the matrix have different roles in cellular respiration.

Figure 4.14 This electron micrograph shows a mitochondrion through an electron microscope. This organelle has an outer membrane and
an inner membrane. The inner membrane contains folds, called cristae, which increase its surface area. We call the space between the two
membranes the intermembrane space, and the space inside the inner membrane the mitochondrial matrix. ATP synthesis takes place on
106 Chapter 4 • Cell Structure

the inner membrane. (credit: modification of work by Matthew Britton; scale-bar data from Matt Russell)

Peroxisomes
Peroxisomes are small, round organelles enclosed by single membranes. They carry out oxidation reactions that break down
fatty acids and amino acids. They also detoxify many poisons that may enter the body. (Many of these oxidation reactions release
hydrogen peroxide, H2O2, which would be damaging to cells; however, when these reactions are confined to peroxisomes,
enzymes safely break down the H2O2 into oxygen and water.) For example, peroxisomes in liver cells detoxify alcohol.
Glyoxysomes, which are specialized peroxisomes in plants, are responsible for converting stored fats into sugars. Plant cells
contain many different types of peroxisomes that play a role in metabolism, pathogene defense, and stress response, to mention
a few.

Vesicles and Vacuoles

Vesicles and vacuoles are membrane-bound sacs that function in storage and transport. Other than the fact that vacuoles are
somewhat larger than vesicles, there is a very subtle distinction between them. Vesicle membranes can fuse with either the
plasma membrane or other membrane systems within the cell. Additionally, some agents such as enzymes within plant vacuoles
break down macromolecules. The vacuole's membrane does not fuse with the membranes of other cellular components.

Animal Cells versus Plant Cells

At this point, you know that each eukaryotic cell has a plasma membrane, cytoplasm, a nucleus, ribosomes, mitochondria,
peroxisomes, and in some, vacuoles, but there are some striking differences between animal and plant cells. While both animal
and plant cells have microtubule organizing centers (MTOCs), animal cells also have centrioles associated with the MTOC: a
complex we call the centrosome. Animal cells each have a centrosome and lysosomes; whereas, most plant cells do not. Plant
cells have a cell wall, chloroplasts and other specialized plastids, and a large central vacuole; whereas, animal cells do not.

The Centrosome
The centrosome is a microtubule-organizing center found near the nuclei of animal cells. It contains a pair of centrioles, two
structures that lie perpendicular to each other (Figure 4.15). Each centriole is a cylinder of nine triplets of microtubules.

Figure 4.15 The centrosome consists of two centrioles that lie at right angles to each other. Each centriole is a cylinder comprised of nine
triplets of microtubules. Nontubulin proteins (indicated by the green lines) hold the microtubule triplets together.

The centrosome (the organelle where all microtubules originate) replicates itself before a cell divides, and the centrioles appear
to have some role in pulling the duplicated chromosomes to opposite ends of the dividing cell. However, the centriole's exact
function in cell division isn’t clear, because cells that have had the centrosome removed can still divide, and plant cells, which
lack centrosomes, are capable of cell division.

Lysosomes
Animal cells have another set of organelles that most plant cells do not: lysosomes. The lysosomes are the cell’s “garbage
disposal.” In plant cells, the digestive processes take place in vacuoles. Enzymes within the lysosomes aid in breaking down
proteins, polysaccharides, lipids, nucleic acids, and even worn-out organelles. These enzymes are active at a much lower pH
than the cytoplasm's. Therefore, the pH within lysosomes is more acidic than the cytoplasm's pH. Many reactions that take place
in the cytoplasm could not occur at a low pH, so again, the advantage of compartmentalizing the eukaryotic cell into organelles

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 4.3 • Eukaryotic Cells 107

is apparent.

The Cell Wall

If you examine Figure 4.8, the plant cell diagram, you will see a structure external to the plasma membrane. This is the cell wall,
a rigid covering that protects the cell, provides structural support, and gives shape to the cell. Fungal and some protistan cells
also have cell walls. While the prokaryotic cell walls' chief component is peptidoglycan, the major organic molecule in the plant
(and some protists') cell wall is cellulose (Figure 4.16), a polysaccharide comprised of glucose units. Have you ever noticed that
when you bite into a raw vegetable, like celery, it crunches? That’s because you are tearing the celery cells' rigid cell walls with
your teeth.

Figure 4.16 Cellulose is a long chain of β-glucose molecules connected by a 1-4 linkage. The dashed lines at each end of the figure indicate
a series of many more glucose units. The size of the page makes it impossible to portray an entire cellulose molecule.

Chloroplasts
Like the mitochondria, chloroplasts have their own DNA and ribosomes, but chloroplasts have an entirely different function.
Chloroplasts are plant cell organelles that carry out photosynthesis. Photosynthesis is the series of reactions that use carbon
dioxide, water, and light energy to make glucose and oxygen. This is a major difference between plants and animals. Plants
(autotrophs) are able to make their own food, like sugars used in cellular respiration to provide ATP energy generated in the
plant mitochondria. Animals (heterotrophs) must ingest their food.

Like mitochondria, chloroplasts have outer and inner membranes, but within the space enclosed by a chloroplast’s inner
membrane is a set of interconnected and stacked fluid-filled membrane sacs we call thylakoids (Figure 4.17). Each thylakoid
stack is a granum (plural = grana). We call the fluid enclosed by the inner membrane that surrounds the grana the stroma.

Figure 4.17 The chloroplast has an outer membrane, an inner membrane, and membrane structures - thylakoids that are stacked into
grana. We call the space inside the thylakoid membranes the thylakoid space. The light harvesting reactions take place in the thylakoid
membranes, and sugar synthesis takes place in the fluid inside the inner membrane, which we call the stroma. Chloroplasts also have their
own genome, which is contained on a single circular chromosome.

The chloroplasts contain a green pigment, chlorophyll, which captures the light energy that drives the reactions of
photosynthesis. Like plant cells, photosynthetic protists also have chloroplasts. Some bacteria perform photosynthesis, but their
chlorophyll is not relegated to an organelle.
108 Chapter 4 • Cell Structure

EVOLUTION CONNECTION

Endosymbiosis
We have mentioned that both mitochondria and chloroplasts contain DNA and ribosomes. Have you wondered why? Strong
evidence points to endosymbiosis as the explanation.

Symbiosis is a relationship in which organisms from two separate species depend on each other for their survival.
Endosymbiosis (endo- = “within”) is a mutually beneficial relationship in which one organism lives inside the other.
Endosymbiotic relationships abound in nature. We have already mentioned that microbes that produce vitamin K live inside the
human gut. This relationship is beneficial for us because we are unable to synthesize vitamin K. It is also beneficial for the
microbes because they are protected from other organisms and from drying out, and they receive abundant food from the
environment of the large intestine.

Scientists have long noticed that bacteria, mitochondria, and chloroplasts are similar in size. We also know that bacteria have
DNA and ribosomes, just like mitochondria and chloroplasts. Scientists believe that host cells and bacteria formed an
endosymbiotic relationship when the host cells ingested both aerobic and autotrophic bacteria (cyanobacteria) but did not
destroy them. Through many millions of years of evolution, these ingested bacteria became more specialized in their functions,
with the aerobic bacteria becoming mitochondria and the autotrophic bacteria becoming chloroplasts.

The Central Vacuole

Previously, we mentioned vacuoles as essential components of plant cells. If you look at Figure 4.8b, you will see that plant cells
each have a large central vacuole that occupies most of the cell's area. The central vacuole plays a key role in regulating the cell’s
concentration of water in changing environmental conditions. Have you ever noticed that if you forget to water a plant for a few
days, it wilts? That’s because as the water concentration in the soil becomes lower than the water concentration in the plant,
water moves out of the central vacuoles and cytoplasm. As the central vacuole shrinks, it leaves the cell wall unsupported. This
loss of support to the plant's cell walls results in the wilted appearance.

The central vacuole also supports the cell's expansion. When the central vacuole holds more water, the cell becomes larger
without having to invest considerable energy in synthesizing new cytoplasm.

4.4 The Endomembrane System and Proteins

By the end of this section, you will be able to do the following:
• List the components of the endomembrane system
• Recognize the relationship between the endomembrane system and its functions

The endomembrane system (endo = “within”) is a group of membranes and organelles (Figure 4.18) in eukaryotic cells that works
together to modify, package, and transport lipids and proteins. It includes the nuclear envelope, lysosomes, and vesicles, which
we have already mentioned, and the endoplasmic reticulum and Golgi apparatus, which we will cover shortly. Although not
technically within the cell, the plasma membrane is included in the endomembrane system because, as you will see, it interacts
with the other endomembranous organelles. The endomembrane system does not include either mitochondria or chloroplast
membranes.

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 4.4 • The Endomembrane System and Proteins 109

VISUAL CONNECTION

Figure 4.18 Membrane and secretory proteins are synthesized in the rough endoplasmic reticulum (RER). The RER also sometimes
modifies proteins. In this illustration, an attachment of a (purple) carbohydrate modifies a (green) integral membrane protein in the ER.
Vesicles with the integral protein bud from the ER and fuse with the Golgi apparatus' cis face. As the protein passes along the Golgi’s
cisternae, the addition of more carbohydrates further modifies it. After its synthesis is complete, it exits as an integral membrane protein of
the vesicle that buds from the Golgi’s trans face. When the vesicle fuses with the cell membrane, the protein becomes an integral portion of
that cell membrane. (credit: modification of work by Magnus Manske)

If a peripheral membrane protein were synthesized in the lumen (inside) of the ER, would it end up on the inside or outside of
the plasma membrane?

The Endoplasmic Reticulum

The endoplasmic reticulum (ER) (Figure 4.18) is a series of interconnected membranous sacs and tubules that collectively
modifies proteins and synthesizes lipids. However, these two functions take place in separate areas of the ER: the rough ER and
the smooth ER, respectively.

We call the ER tubules' hollow portion the lumen or cisternal space. The ER's membrane, which is a phospholipid bilayer
embedded with proteins, is continuous with the nuclear envelope.

Rough ER
Scientists have named the rough endoplasmic reticulum (RER) as such because the ribosomes attached to its cytoplasmic
110 Chapter 4 • Cell Structure

surface give it a studded appearance when viewing it through an electron microscope (Figure 4.19).

Figure 4.19 This transmission electron micrograph shows the rough endoplasmic reticulum and other organelles in a pancreatic cell.
(credit: modification of work by Louisa Howard)

Ribosomes transfer their newly synthesized proteins into the RER's lumen where they undergo structural modifications, such as
folding or acquiring side chains. These modified proteins incorporate into cellular membranes—the ER or the ER's or other
organelles' membranes. The proteins can also secrete from the cell (such as protein hormones, enzymes). The RER also makes
phospholipids for cellular membranes.

If the phospholipids or modified proteins are not destined to stay in the RER, they will reach their destinations via transport
vesicles that bud from the RER’s membrane (Figure 4.18).

Since the RER is engaged in modifying proteins (such as enzymes, for example) that secrete from the cell, you would be correct
in assuming that the RER is abundant in cells that secrete proteins. This is the case with liver cells, for example.

Smooth ER
The smooth endoplasmic reticulum (SER) is continuous with the RER but has few or no ribosomes on its cytoplasmic surface
(Figure 4.18). SER functions include synthesis of carbohydrates, lipids, and steroid hormones; detoxification of medications and
poisons; and storing calcium ions.

In muscle cells, a specialized SER, the sarcoplasmic reticulum, is responsible for storing calcium ions that are needed to trigger
the muscle cells' coordinated contractions.

LINK TO LEARNING
You can watch an excellent animation of the endomembrane system here (http://openstax.org/l/insane_in_the_endomembrane)
. At the end of the animation, there is a short self-assessment.

CAREER CONNECTION

Cardiologist
Heart disease is the leading cause of death in the United States. This is primarily due to our sedentary lifestyle and our high
trans-fat diets.

Heart failure is just one of many disabling heart conditions. Heart failure does not mean that the heart has stopped working.
Rather, it means that the heart can’t pump with sufficient force to transport oxygenated blood to all the vital organs. Left
untreated, heart failure can lead to kidney failure and other organ failure.

Cardiac muscle tissue comprises the heart's wall. Heart failure occurs when cardiac muscle cells' endoplasmic reticula do not
function properly. As a result, an insufficient number of calcium ions are available to trigger a sufficient contractile force.

Cardiologists (cardi- = “heart”; -ologist = “one who studies”) are doctors who specialize in treating heart diseases, including
heart failure. Cardiologists can diagnose heart failure via a physical examination, results from an electrocardiogram (ECG, a test
that measures the heart's electrical activity), a chest X-ray to see whether the heart is enlarged, and other tests. If the
cardiologist diagnoses heart failure, he or she will typically prescribe appropriate medications and recommend a reduced table

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 4.4 • The Endomembrane System and Proteins 111

salt intake and a supervised exercise program.

The Golgi Apparatus

We have already mentioned that vesicles can bud from the ER and transport their contents elsewhere, but where do the vesicles
go? Before reaching their final destination, the lipids or proteins within the transport vesicles still need sorting, packaging, and
tagging so that they end up in the right place. Sorting, tagging, packaging, and distributing lipids and proteins takes place in
the Golgi apparatus (also called the Golgi body), a series of flattened membranes (Figure 4.20).

Figure 4.20 The Golgi apparatus in this white blood cell is visible as a stack of semicircular, flattened rings in the lower portion of the image.
You can see several vesicles near the Golgi apparatus. (credit: modification of work by Louisa Howard)

We call the Golgi apparatus' the cis face. The opposite side is the trans face. The transport vesicles that formed from the ER travel
to the cis face, fuse with it, and empty their contents into the Golgi apparatus' lumen. As the proteins and lipids travel through
the Golgi, they undergo further modifications that allow them to be sorted. The most frequent modification is adding short
sugar molecule chains. These newly modified proteins and lipids then tag with phosphate groups or other small molecules in
order to travel to their proper destinations.

Finally, the modified and tagged proteins are packaged into secretory vesicles that bud from the Golgi's trans face. While some
of these vesicles deposit their contents into other cell parts where they will be used, other secretory vesicles fuse with the plasma
membrane and release their contents outside the cell.

In another example of form following function, cells that engage in a great deal of secretory activity (such as salivary gland cells
that secrete digestive enzymes or immune system cells that secrete antibodies) have an abundance of Golgi.

In plant cells, the Golgi apparatus has the additional role of synthesizing polysaccharides, some of which are incorporated into
the cell wall and some of which other cell parts use.

CAREER CONNECTION

Geneticist
Many diseases arise from genetic mutations that prevent synthesizing critical proteins. One such disease is Lowe disease (or
oculocerebrorenal syndrome, because it affects the eyes, brain, and kidneys). In Lowe disease, there is a deficiency in an enzyme
localized to the Golgi apparatus. Children with Lowe disease are born with cataracts, typically develop kidney disease after the
first year of life, and may have impaired mental abilities.

A mutation on the X chromosome causes Lowe disease. The X chromosome is one of the two human sex chromosomes, as these
chromosomes determine a person's sex. Females possess two X chromosomes while males possess one X and one Y
chromosome. In females, the genes on only one of the two X chromosomes are expressed. Females who carry the Lowe disease
gene on one of their X chromosomes are carriers and do not show symptoms of the disease. However, males only have one X
chromosome and the genes on this chromosome are always expressed. Therefore, males will always have Lowe disease if their X
chromosome carries the Lowe disease gene. Geneticists have identified the mutated gene's location, as well as many other
mutation locations that cause genetic diseases. Through prenatal testing, a woman can find out if the fetus she is carrying may
be afflicted with one of several genetic diseases.
112 Chapter 4 • Cell Structure

Geneticists analyze prenatal genetic test results and may counsel pregnant women on available options. They may also conduct
genetic research that leads to new drugs or foods, or perform DNA analyses for forensic investigations.

Lysosomes
In addition to their role as the digestive component and organelle-recycling facility of animal cells, lysosomes are part of the
endomembrane system. Lysosomes also use their hydrolytic enzymes to destroy pathogens (disease-causing organisms) that
might enter the cell. A good example of this occurs in macrophages, a group of white blood cells which are part of your body’s
immune system. In a process that scientists call phagocytosis or endocytosis, a section of the macrophage's plasma membrane
invaginates (folds in) and engulfs a pathogen. The invaginated section, with the pathogen inside, then pinches itself off from the
plasma membrane and becomes a vesicle. The vesicle fuses with a lysosome. The lysosome’s hydrolytic enzymes then destroy the
pathogen (Figure 4.21).

Figure 4.21 A macrophage has engulfed (phagocytized) a potentially pathogenic bacterium and then fuses with lysosomes within the cell to
destroy the pathogen. Other organelles are present in the cell but for simplicity we do not show them.

4.5 The Cytoskeleton

By the end of this section, you will be able to do the following:
• Describe the cytoskeleton
• Compare the roles of microfilaments, intermediate filaments, and microtubules
• Compare and contrast cilia and flagella
• Summarize the differences among the components of prokaryotic cells, animal cells, and plant cells

If you were to remove all the organelles from a cell, would the plasma membrane and the cytoplasm be the only components left?
No. Within the cytoplasm, there would still be ions and organic molecules, plus a network of protein fibers that help maintain
the cell's shape, secure some organelles in specific positions, allow cytoplasm and vesicles to move within the cell, and enable
cells within multicellular organisms to move. Collectively, scientists call this network of protein fibers the cytoskeleton. There
are three types of fibers within the cytoskeleton: microfilaments, intermediate filaments, and microtubules (Figure 4.22). Here,
we will examine each.

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 4.5 • The Cytoskeleton 113

Figure 4.22 Microfilaments thicken the cortex around the cell's inner edge. Like rubber bands, they resist tension. There are microtubules in
the cell's interior where they maintain their shape by resisting compressive forces. There are intermediate filaments throughout the cell
that hold organelles in place.

Microfilaments
Of the three types of protein fibers in the cytoskeleton, microfilaments are the narrowest. They function in cellular movement,
have a diameter of about 7 nm, and are comprised of two globular protein intertwined strands, which we call actin (Figure 4.23).
For this reason, we also call microfilaments actin filaments.
114 Chapter 4 • Cell Structure

Figure 4.23 Two intertwined actin strands comprise microfilaments.

ATP powers actin to assemble its filamentous form, which serves as a track for the movement of a motor protein we call myosin.
This enables actin to engage in cellular events requiring motion, such as cell division in eukaryotic cells and cytoplasmic
streaming, which is the cell cytoplasm's circular movement in plant cells. Actin and myosin are plentiful in muscle cells. When
your actin and myosin filaments slide past each other, your muscles contract.

Microfilaments also provide some rigidity and shape to the cell. They can depolymerize (disassemble) and reform quickly, thus
enabling a cell to change its shape and move. White blood cells (your body’s infection-fighting cells) make good use of this
ability. They can move to an infection site and phagocytize the pathogen.

LINK TO LEARNING
To see an example of a white blood cell in action, watch a short time-lapse video of the cell capturing two bacteria. It engulfs one
and then moves on to the other.

Click to view content (https://www.openstax.org/l/chasing_bcteria)

Intermediate Filaments
Several strands of fibrous proteins that are wound together comprise intermediate filaments (Figure 4.24). Cytoskeleton
elements get their name from the fact that their diameter, 8 to 10 nm, is between those of microfilaments and microtubules.

Figure 4.24 Intermediate filaments consist of several intertwined strands of fibrous proteins.

Intermediate filaments have no role in cell movement. Their function is purely structural. They bear tension, thus maintaining
the cell's shape, and anchor the nucleus and other organelles in place. Figure 4.22 shows how intermediate filaments create a
supportive scaffolding inside the cell.

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 4.5 • The Cytoskeleton 115

The intermediate filaments are the most diverse group of cytoskeletal elements. Several fibrous protein types are in the
intermediate filaments. You are probably most familiar with keratin, the fibrous protein that strengthens your hair, nails, and
the skin's epidermis.

Microtubules
As their name implies, microtubules are small hollow tubes. Polymerized dimers of α-tubulin and β-tubulin, two globular
proteins, comprise the microtubule's walls (Figure 4.25). With a diameter of about 25 nm, microtubules are cytoskeletons'
widest components. They help the cell resist compression, provide a track along which vesicles move through the cell, and pull
replicated chromosomes to opposite ends of a dividing cell. Like microfilaments, microtubules can disassemble and reform
quickly.

Figure 4.25 Microtubules are hollow. Their walls consist of 13 polymerized dimers of α-tubulin and β-tubulin (right image). The left image
shows the tube's molecular structure.

Microtubules are also the structural elements of flagella, cilia, and centrioles (the latter are the centrosome's two perpendicular
bodies). In animal cells, the centrosome is the microtubule-organizing center. In eukaryotic cells, flagella and cilia are quite
different structurally from their counterparts in prokaryotes, as we discuss below.

Flagella and Cilia

The flagella (singular = flagellum) are long, hair-like structures that extend from the plasma membrane and enable an entire cell
to move (for example, sperm, Euglena, and some prokaryotes). When present, the cell has just one flagellum or a few flagella.
However, when cilia (singular = cilium) are present, many of them extend along the plasma membrane's entire surface. They are
short, hair-like structures that move entire cells (such as paramecia) or substances along the cell's outer surface (for example, the
cilia of cells lining the Fallopian tubes that move the ovum toward the uterus, or cilia lining the cells of the respiratory tract that
trap particulate matter and move it toward your nostrils.)

Despite their differences in length and number, flagella and cilia share a common structural arrangement of microtubules called
a “9 + 2 array.” This is an appropriate name because a single flagellum or cilium is made of a ring of nine microtubule doublets,
surrounding a single microtubule doublet in the center (Figure 4.26).
116 Chapter 4 • Cell Structure

Figure 4.26 This transmission electron micrograph of two flagella shows the microtubules' 9 + 2 array: nine microtubule doublets surround
a single microtubule doublet. (credit: modification of work by Dartmouth Electron Microscope Facility, Dartmouth College; scale-bar data
from Matt Russell)

You have now completed a broad survey of prokaryotic and eukaryotic cell components. For a summary of cellular components
in prokaryotic and eukaryotic cells, see Table 4.1.

Components of Prokaryotic and Eukaryotic Cells

Present
Present
Cell Present in in
Function in Plant
Component Prokaryotes? Animal
Cells?
Cells?

Separates cell from external environment; controls passage

Plasma
of organic molecules, ions, water, oxygen, and wastes into Yes Yes Yes
membrane
and out of cell

Provides turgor pressure to plant cells as fluid inside the

Cytoplasm central vacuole; site of many metabolic reactions; medium in Yes Yes Yes
which organelles are found

Darkened area within the nucleus where ribosomal subunits

Nucleolus No Yes Yes
are synthesized.

Cell organelle that houses DNA and directs synthesis of

Nucleus No Yes Yes
ribosomes and proteins

Ribosomes Protein synthesis Yes Yes Yes

Mitochondria ATP production/cellular respiration No Yes Yes

Oxidize and thus break down fatty acids and amino acids,
Peroxisomes No Yes Yes
and detoxify poisons

Vesicles and
Storage and transport; digestive function in plant cells No Yes Yes
vacuoles

Table 4.1

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 4.6 • Connections between Cells and Cellular Activities 117

Present
Present
Cell Present in in
Function in Plant
Component Prokaryotes? Animal
Cells?
Cells?

Unspecified role in cell division in animal cells; microtubule

Centrosome No Yes No
source in animal cells

Digestion of macromolecules; recycling of worn-out

Lysosomes No Yes Some
organelles

Yes,
Protection, structural support, and maintenance of cell Yes, primarily
Cell wall No primarily
shape peptidoglycan
cellulose

Chloroplasts Photosynthesis No No Yes

Endoplasmic
Modifies proteins and synthesizes lipids No Yes Yes
reticulum

Golgi Modifies, sorts, tags, packages, and distributes lipids and

No Yes Yes
apparatus proteins

Maintains cell’s shape, secures organelles in specific

Cytoskeleton positions, allows cytoplasm and vesicles to move within cell, Yes Yes Yes
and enables unicellular organisms to move independently

No, except
for some
Flagella Cellular locomotion Some Some
plant
sperm cells

Cellular locomotion, movement of particles along plasma

Cilia Some Some No
membrane's extracellular surface, and filtration

Table 4.1

4.6 Connections between Cells and Cellular Activities

By the end of this section, you will be able to do the following:
• Describe the extracellular matrix
• List examples of the ways that plant cells and animal cells communicate with adjacent cells
• Summarize the roles of tight junctions, desmosomes, gap junctions, and plasmodesmata

You already know that tissue is a group of similar cells working together. As you might expect, if cells are to work together, they
must communicate with each other, just as you need to communicate with others if you work on a group project. Let’s take a
look at how cells communicate with each other.

Extracellular Matrix of Animal Cells

While cells in most multicellular organisms release materials into the extracellular space, animal cells will be discussed as an
example. The primary components of these materials are proteins, and the most abundant protein is collagen. Collagen fibers
are interwoven with proteoglycans, which are carbohydrate-containing protein molecules. Collectively, we call these materials
the extracellular matrix (Figure 4.27). Not only does the extracellular matrix hold the cells together to form a tissue, but it also
118 Chapter 4 • Cell Structure

allows the cells within the tissue to communicate with each other. How can this happen?

Figure 4.27 The extracellular matrix consists of a network of proteins and carbohydrates.

Cells have protein receptors on their plasma membranes' extracellular surfaces. When a molecule within the matrix binds to the
receptor, it changes the receptor's molecular structure. The receptor, in turn, changes the microfilaments' conformation
positioned just inside the plasma membrane. These conformational changes induce chemical signals inside the cell that reach
the nucleus and turn “on” or “off” the transcription of specific DNA sections, which affects the associated protein production,
thus changing the activities within the cell.

Blood clotting provides an example of the extracellular matrix's role in cell communication. When the cells lining a blood vessel
are damaged, they display a protein receptor, which we call tissue factor. When tissue factor binds with another factor in the
extracellular matrix, it causes platelets to adhere to the damaged blood vessel's wall, stimulates the adjacent smooth muscle cells
in the blood vessel to contract (thus constricting the blood vessel), and initiates a series of steps that stimulate the platelets to
produce clotting factors.

Intercellular Junctions
Cells can also communicate with each other via direct contact, or intercellular junctions. There are differences in the ways that
plant and animal and fungal cells communicate. Plasmodesmata are junctions between plant cells; whereas, animal cell contacts
include tight junctions, gap junctions, and desmosomes.

Plasmodesmata
In general, long stretches of the plasma membranes of neighboring plant cells cannot touch one another because the cell wall
that surrounds each cell separates them (Figure 4.8). How then, can a plant transfer water and other soil nutrients from its
roots, through its stems, and to its leaves? Such transport uses the vascular tissues (xylem and phloem) primarily. There also
exist structural modifications, which we call plasmodesmata (singular = plasmodesma). Numerous channels that pass between
adjacent plant cells' cell walls connect their cytoplasm, and enable transport of materials from cell to cell, and thus throughout
the plant (Figure 4.28).

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 4.6 • Connections between Cells and Cellular Activities 119

Figure 4.28 A plasmodesma is a channel between two adjacent plant cells' cell walls. Plasmodesmata allow materials to pass from one
plant cell's cytoplasm to an adjacent cell's cytoplasm.

Tight Junctions
A tight junction is a watertight seal between two adjacent animal cells (Figure 4.29). Proteins (predominantly two proteins called
claudins and occludins) tightly hold the cells against each other.

Figure 4.29 Tight junctions form watertight connections between adjacent animal cells. Proteins create tight junction adherence. (credit:
modification of work by Mariana Ruiz Villareal)

This tight adherence prevents materials from leaking between the cells; tight junctions are typically found in epithelial tissues
that line internal organs and cavities, and comprise most of the skin. For example, the tight junctions of the epithelial cells
lining your urinary bladder prevent urine from leaking out into the extracellular space.

Desmosomes
Also only in animal cells are desmosomes, which act like spot welds between adjacent epithelial cells (Figure 4.30). Cadherins,
short proteins in the plasma membrane connect to intermediate filaments to create desmosomes. The cadherins connect two
120 Chapter 4 • Cell Structure

adjacent cells and maintain the cells in a sheet-like formation in organs and tissues that stretch, like the skin, heart, and
muscles.

Figure 4.30 A desmosome forms a very strong spot weld between cells. Linking cadherins and intermediate filaments create it. (credit:
modification of work by Mariana Ruiz Villareal)

Gap Junctions
Gap junctions in animal cells are like plasmodesmata in plant cells in that they are channels between adjacent cells that allow for
transporting ions, nutrients, and other substances that enable cells to communicate (Figure 4.31). Structurally, however, gap
junctions and plasmodesmata differ.

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 4.6 • Connections between Cells and Cellular Activities 121

Figure 4.31 A gap junction is a protein-lined pore that allows water and small molecules to pass between adjacent animal cells. (credit:
modification of work by Mariana Ruiz Villareal)

Gap junctions develop when a set of six proteins (connexins) in the plasma membrane arrange themselves in an elongated
donut-like configuration - a connexon. When the connexon's pores (“doughnut holes”) in adjacent animal cells align, a channel
between the two cells forms. Gap junctions are particularly important in cardiac muscle. The electrical signal for the muscle to
contract passes efficiently through gap junctions, allowing the heart muscle cells to contract in tandem.

LINK TO LEARNING
To conduct a virtual microscopy lab and review the parts of a cell, work through the steps of this interactive assignment
(http://openstax.org/l/microscopy_lab) .
122 Chapter 4 • Key Terms

KEY TERMS
cell theory see unified cell theory flagellum (plural = flagella) long, hair-like structure that
cell wall rigid cell covering comprised of various molecules extends from the plasma membrane and moves the cell
that protects the cell, provides structural support, and gap junction channel between two adjacent animal cells
gives shape to the cell that allows ions, nutrients, and low molecular weight
central vacuole large plant cell organelle that regulates the substances to pass between cells, enabling the cells to
cell’s storage compartment, holds water, and plays a communicate
significant role in cell growth as the site of Golgi apparatus eukaryotic organelle comprised of a series
macromolecule degradation of stacked membranes that sorts, tags, and packages
centrosome region in animal cells made of two centrioles lipids and proteins for distribution
that serves as an organizing center for microtubules intermediate filament cytoskeletal component, comprised
chlorophyll green pigment that captures the light energy of several fibrous protein intertwined strands, that bears
that drives the light reactions of photosynthesis tension, supports cell-cell junctions, and anchors cells to
chloroplast plant cell organelle that carries out extracellular structures
photosynthesis light microscope an instrument that magnifies an object
chromatin protein-DNA complex that serves as the using a beam of visible light that passes and bends
chromosomes' building material through a lens system to visualize a specimen
chromosome structure within the nucleus that comprises lysosome organelle in an animal cell that functions as the
chromatin that contains DNA, the hereditary material cell’s digestive component; it breaks down proteins,
cilium (plural = cilia) short, hair-like structure that extends polysaccharides, lipids, nucleic acids, and even worn-out
from the plasma membrane in large numbers and organelles
functions to move an entire cell or move substances along microfilament the cytoskeleton system's narrowest
the cell's outer surface element; it provides rigidity and shape to the cell and
cytoplasm entire region between the plasma membrane enables cellular movements
and the nuclear envelope, consisting of organelles microscope an instrument that magnifies an object
suspended in the gel-like cytosol, the cytoskeleton, and microtubule the cytoskeleton system's widest element; it
various chemicals helps the cell resist compression, provides a track along
cytoskeleton protein fiber network that collectively which vesicles move through the cell, pulls replicated
maintains the cell's shape, secures some organelles in chromosomes to opposite ends of a dividing cell, and is
specific positions, allows cytoplasm and vesicles to move the structural element of centrioles, flagella, and cilia
within the cell, and enables unicellular organisms to mitochondria (singular = mitochondrion) cellular
move independently organelles responsible for carrying out cellular
cytosol the cytoplasm's gel-like material in which cell respiration, resulting in producing ATP, the cell’s main
structures are suspended energy-carrying molecule
desmosome linkages between adjacent epithelial cells that nuclear envelope double-membrane structure that
form when cadherins in the plasma membrane attach to constitutes the nucleus' outermost portion
intermediate filaments nucleoid central part of a prokaryotic cell's central part
electron microscope an instrument that magnifies an where the chromosome is located
object using an electron beam that passes and bends nucleolus darkly staining body within the nucleus that is
through a lens system to visualize a specimen responsible for assembling ribosome subunits
endomembrane system group of organelles and nucleoplasm semi-solid fluid inside the nucleus that
membranes in eukaryotic cells that work together contains the chromatin and nucleolus
modifying, packaging, and transporting lipids and nucleus cell organelle that houses the cell’s DNA and
proteins directs ribosome and protein synthesis
endoplasmic reticulum (ER) series of interconnected organelle compartment or sac within a cell
membranous structures within eukaryotic cells that peroxisome small, round organelle that contains hydrogen
collectively modify proteins and synthesize lipids peroxide, oxidizes fatty acids and amino acids, and
eukaryotic cell cell that has a membrane-bound nucleus detoxifies many poisons
and several other membrane-bound compartments or plasma membrane phospholipid bilayer with embedded
sacs (integral) or attached (peripheral) proteins, and separates
extracellular matrix material secreted from animal or the cell's internal content from its surrounding
fungal cells that provides mechanical protection and environment
anchoring for the cells in the tissue plasmodesma (plural = plasmodesmata) channel that

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 Chapter 4 • Chapter Summary 123

passes between adjacent plant cells' cell walls, connects lipids, and steroid hormones; detoxifies certain chemicals
their cytoplasm, and allows transporting of materials (like pesticides, preservatives, medications, and
from cell to cell environmental pollutants), and stores calcium ions
prokaryote unicellular organism that lacks a nucleus or any tight junction protein adherence that creates a firm seal
other membrane-bound organelle between two adjacent animal cells
ribosome cellular structure that carries out protein unified cell theory a biological concept that states that one
synthesis or more cells comprise all organisms; the cell is the basic
rough endoplasmic reticulum (RER) region of the unit of life; and new cells arise from existing cells
endoplasmic reticulum that is studded with ribosomes vacuole membrane-bound sac, somewhat larger than a
and engages in protein modification and phospholipid vesicle, which functions in cellular storage and transport
synthesis vesicle small, membrane-bound sac that functions in
smooth endoplasmic reticulum (SER) region of the cellular storage and transport; its membrane is capable of
endoplasmic reticulum that has few or no ribosomes on fusing with the plasma membrane and the membranes of
its cytoplasmic surface and synthesizes carbohydrates, the endoplasmic reticulum and Golgi apparatus

CHAPTER SUMMARY
4.1 Studying Cells hydrolyze fatty acids, amino acids, and some toxins. Vesicles
and vacuoles are storage and transport compartments. In
A cell is the smallest unit of life. Most cells are so tiny that we
plant cells, vacuoles also help break down macromolecules.
cannot see them with the naked eye. Therefore, scientists use
microscopes to study cells. Electron microscopes provide Animal cells also have a centrosome and lysosomes. The
higher magnification, higher resolution, and more detail centrosome has two bodies perpendicular to each other, the
than light microscopes. The unified cell theory states that centrioles, and has an unknown purpose in cell division.
one or more cells comprise all organisms, the cell is the basic Lysosomes are the digestive organelles of animal cells.
unit of life, and new cells arise from existing cells.
Plant cells and plant-like cells each have a cell wall,
4.2 Prokaryotic Cells chloroplasts, and a central vacuole. The plant cell wall, whose
primary component is cellulose, protects the cell, provides
Prokaryotes are single-celled organisms of the domains structural support, and gives the cell shape. Photosynthesis
Bacteria and Archaea. All prokaryotes have plasma takes place in chloroplasts. The central vacuole can expand
membranes, cytoplasm, ribosomes, and DNA that is not without having to produce more cytoplasm.
membrane-bound. Most have peptidoglycan cell walls and
many have polysaccharide capsules. Prokaryotic cells range 4.4 The Endomembrane System and
in diameter from 0.1 to 5.0 μm. Proteins
As a cell increases in size, its surface area-to-volume ratio The endomembrane system includes the nuclear envelope,
decreases. If the cell grows too large, the plasma membrane lysosomes, vesicles, the ER, and Golgi apparatus, as well as
will not have sufficient surface area to support the rate of the plasma membrane. These cellular components work
diffusion required for the increased volume. together to modify, package, tag, and transport proteins and
lipids that form the membranes.
4.3 Eukaryotic Cells
The RER modifies proteins and synthesizes phospholipids in
Like a prokaryotic cell, a eukaryotic cell has a plasma
cell membranes. The SER synthesizes carbohydrates, lipids,
membrane, cytoplasm, and ribosomes, but a eukaryotic cell
and steroid hormones; engages in the detoxification of
is typically larger than a prokaryotic cell, has a true nucleus
medications and poisons; and stores calcium ions. Sorting,
(meaning a membrane surrounds its DNA), and has other
tagging, packaging, and distributing lipids and proteins take
membrane-bound organelles that allow for
place in the Golgi apparatus. Budding RER and Golgi
compartmentalizing functions. The plasma membrane is a
membranes create lysosomes. Lysosomes digest
phospholipid bilayer embedded with proteins. The nucleus’s
macromolecules, recycle worn-out organelles, and destroy
nucleolus is the site of ribosome assembly. We find
pathogens.
ribosomes either in the cytoplasm or attached to the
cytoplasmic side of the plasma membrane or endoplasmic 4.5 The Cytoskeleton
reticulum. They perform protein synthesis. Mitochondria
The cytoskeleton has three different protein element types.
participate in cellular respiration. They are responsible for
From narrowest to widest, they are the microfilaments (actin
the majority of ATP produced in the cell. Peroxisomes
filaments), intermediate filaments, and microtubules.
124 Chapter 4 • Visual Connection Questions

Biologists often associate microfilaments with myosin. They and are connected to each other via tight junctions,
provide rigidity and shape to the cell and facilitate cellular desmosomes, and gap junctions. Plant cells are connected
movements. Intermediate filaments bear tension and anchor and communicate with each other via plasmodesmata.
the nucleus and other organelles in place. Microtubules help
When protein receptors on the plasma membrane's surface
the cell resist compression, serve as tracks for motor
of an animal cell bind to a substance in the extracellular
proteins that move vesicles through the cell, and pull
matrix, a chain of reactions begins that changes activities
replicated chromosomes to opposite ends of a dividing cell.
taking place within the cell. Plasmodesmata are channels
They are also the structural element of centrioles, flagella,
between adjacent plant cells, while gap junctions are
and cilia.
channels between adjacent animal cells. However, their
4.6 Connections between Cells and structures are quite different. A tight junction is a watertight
Cellular Activities seal between two adjacent cells, while a desmosome acts like
a spot weld.
Animal cells communicate via their extracellular matrices

VISUAL CONNECTION QUESTIONS

1. Figure 4.7 Prokaryotic cells are much smaller than 2. Figure 4.8 If the nucleolus were not able to carry out its
eukaryotic cells. What advantages might small cell size function, what other cellular organelles would be
confer on a cell? What advantages might large cell size affected?
have?
3. Figure 4.18 If a peripheral membrane protein were
synthesized in the lumen (inside) of the ER, would it end
up on the inside or outside of the plasma membrane?

REVIEW QUESTIONS
4. When viewing a specimen through a light microscope, 8. Which of the following organisms is a prokaryote?
scientists use ________ to distinguish the individual a. amoeba
components of cells. b. influenza A virus
a. a beam of electrons c. charophyte algae
b. radioactive isotopes d. E. coli
c. special stains
d. high temperatures 9. Which of the following is surrounded by two
phospholipid bilayers?
5. The ________ is the basic unit of life.
a. the ribosomes
a. organism
b. the vesicles
b. cell
c. the cytoplasm
c. tissue
d. the nucleoplasm
d. organ
10. Peroxisomes got their name because hydrogen peroxide
6. Prokaryotes depend on ________ to obtain some is:
materials and to get rid of wastes. a. used in their detoxification reactions
a. ribosomes b. produced during their oxidation reactions
b. flagella c. incorporated into their membranes
c. cell division d. a cofactor for the organelles’ enzymes
d. diffusion
11. In plant cells, the function of the lysosomes is carried out
7. Bacteria that lack fimbriae are less likely to ________. by __________.
a. adhere to cell surfaces a. vacuoles
b. swim through bodily fluids b. peroxisomes
c. synthesize proteins c. ribosomes
d. retain the ability to divide d. nuclei

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 Chapter 4 • Review Questions 125

12. Which of the following is both in eukaryotic and 18. Congenital disorders of glycosylation are a growing class
prokaryotic cells? of rare diseases. Which organelle would be most
a. nucleus commonly involved in the glycoprotein disorder portion
b. mitochondrion of the group?
c. vacuole a. RER
d. ribosomes b. ribosomes
c. endosomes
13. Tay-Sachs disease is a genetic disorder that results in the
d. Golgi apparatus
destruction of neurons due to a buildup of sphingolipids
in the cells. Which organelle is malfunctioning in Tay-
19. Which of the following have the ability to disassemble
Sachs?
and reform quickly?
a. lysosome
a. microfilaments and intermediate filaments
b. endoplasmic reticulum
b. microfilaments and microtubules
c. peroxisome
c. intermediate filaments and microtubules
d. mitochondria
d. only intermediate filaments

14. Which of the following is not a component of the 20. Which of the following do not play a role in intracellular
endomembrane system? movement?
a. mitochondrion a. microfilaments and intermediate filaments
b. Golgi apparatus b. microfilaments and microtubules
c. endoplasmic reticulum c. intermediate filaments and microtubules
d. lysosome d. only intermediate filaments

15. The process by which a cell engulfs a foreign particle is 21. In humans, _____ are used to move a cell within its
known as: environment while _____ are used to move the
a. endosymbiosis environment relative to the cell.
b. phagocytosis a. cilia, pseudopodia
c. hydrolysis b. flagella; cilia
d. membrane synthesis c. microtubules; flagella
d. microfilaments; microtubules
16. Which of the following is most likely to have the greatest
concentration of smooth endoplasmic reticulum?
22. Which of the following are only in plant cells?
a. a cell that secretes enzymes
a. gap junctions
b. a cell that destroys pathogens
b. desmosomes
c. a cell that makes steroid hormones
c. plasmodesmata
d. a cell that engages in photosynthesis
d. tight junctions
17. Which of the following sequences correctly lists in order
23. The key components of desmosomes are cadherins and
the steps involved in the incorporation of a
__________.
proteinaceous molecule within a cell?
a. actin
a. protein synthesis of the protein on the ribosome;
b. microfilaments
modification in the Golgi apparatus; packaging in
c. intermediate filaments
the endoplasmic reticulum; tagging in the vesicle
d. microtubules
b. synthesis of the protein on the lysosome; tagging in
the Golgi; packaging in the vesicle; distribution in 24. Diseased animal cells may produce molecules that
the endoplasmic reticulum activate death cascades to kill the cells in a controlled
c. synthesis of the protein on the ribosome; manner. Why would neighboring healthy cells also die?
modification in the endoplasmic reticulum; tagging a. The death molecule is passed through
in the Golgi; distribution via the vesicle desmosomes.
d. synthesis of the protein on the lysosome; packaging b. The death molecule is passed through
in the vesicle; distribution via the Golgi; tagging in plasmodesmata.
the endoplasmic reticulum c. The death molecule disrupts the extracellular
matrix.
d. The death molecule passes through gap junctions.
126 Chapter 4 • Critical Thinking Questions

CRITICAL THINKING QUESTIONS

25. In your everyday life, you have probably noticed that 34. Why are plasma membranes arranged as a bilayer rather
certain instruments are ideal for certain situations. For than a monolayer?
example, you would use a spoon rather than a fork to eat
35. In the context of cell biology, what do we mean by form
soup because a spoon is shaped for scooping, while soup
follows function? What are at least two examples of this
would slip between the tines of a fork. The use of ideal
concept?
instruments also applies in science. In what situation(s)
36. In your opinion, is the nuclear membrane part of the
would the use of a light microscope be ideal, and why?
endomembrane system? Why or why not? Defend your
26. In what situation(s) would the use of a scanning electron
answer.
microscope be ideal, and why?
27. In what situation(s) would a transmission electron 37. What are the similarities and differences between the
microscope be ideal, and why? structures of centrioles and flagella?
28. What are the advantages and disadvantages of each of 38. How do cilia and flagella differ?
these types of microscopes? 39. Describe how microfilaments and microtubules are
29. Explain how the formation of an adult human follows involved in the phagocytosis and destruction of a
the cell theory. pathogen by a macrophage.
40. Compare and contrast the boundaries that plant,
30. Antibiotics are medicines that are used to fight bacterial
animal, and bacteria cells use to separate themselves
infections. These medicines kill prokaryotic cells
from their surrounding environment.
without harming human cells. What part or parts of the
bacterial cell do you think antibiotics target? Why? 41. How does the structure of a plasmodesma differ from
31. Explain why not all microbes are harmful. that of a gap junction?
42. Explain how the extracellular matrix functions.
32. You already know that ribosomes are abundant in red
43. Pathogenic E. coli have recently been shown to degrade
blood cells. In what other cells of the body would you
tight junction proteins during infection. How would
find them in great abundance? Why?
this provide an advantage to the bacteria?
33. What are the structural and functional similarities and
differences between mitochondria and chloroplasts?

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