Chapter 4 - Cell Structure
Chapter 4 - Cell Structure
Chapter 4 - Cell Structure
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)
Chapter Outline
4.1: Studying Cells
4.2: Prokaryotic Cells
4.3: Eukaryotic Cells
4.4: The Endomembrane System and Proteins
4.5: The Cytoskeleton
4.6: Connections between Cells and Cellular Activities
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.
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 cells protect
the body's surface and cover the organs and body cavities within. Bone cells help to support 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 plays a vital role during the body's
growth, development, and day-to-day maintenance. In spite of their enormous variety, however, cells from all
organisms—even ones as diverse as bacteria, onion, and human—share certain fundamental characteristics.
A cell is the smallest unit of a living thing. Whether comprised of one cell (like bacteria) or many cells (like a
human), we call it an organism. Thus, cells are the basic building blocks of all organisms.
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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.
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.
(a) (b)
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)
For another perspective on cell size, try the HowBig interactive at this site (http://openstaxcollege.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.
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.
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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)
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”).
A prokaryote is a simple, mostly single-celled (unicellular) organism that lacks a nucleus, or any other
membrane-bound 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.
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
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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.
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. Another way
is to develop organelles that perform specific tasks. These adaptations lead to developing more sophisticated
cells, which we call eukaryotic cells.
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?
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.
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(a)
(b)
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?
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 = 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
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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.
(a) (b)
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.
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,
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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) the cell's “powerhouses” or “energy factories”
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 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.
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 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.
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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, while
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.
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 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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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?
its cytoplasmic 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.
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.
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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 salt intake and a supervised
exercise program.
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.
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.
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).
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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.
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.
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.
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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.
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. (This multimedia resource will open in a
browser.) (http://cnx.org/content/m66454/1.3/#eip-id3083425)
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.
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).
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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.
Table 4.1
Table 4.1
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.
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).
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
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desmosomes. The cadherins connect two 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.
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.
To conduct a virtual microscopy lab and review the parts of a cell, work through the steps of this interactive
assignment (http://openstaxcollege.org/l/microscopy_lab) .