Chapter 3

Cells as
Units of
Life

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3.1. Cell Concept
A. History

1. English scientist Robert Hooke described “little boxes of cells” in 1663

using a compound microscope.
2. Dutch microscopist Anton van Leeuwenhoek made extensive
observations and reported them in letters to Royal Society of London.
3. Advanced high-quality lenses in the early 19th century made it
possible to examine cells.

B. Cell Theory

1. The cell theory asserts that all living organisms are composed of cells.
2. In 1838, Matthias Schleiden announced plant tissue was made of cells.
3. In 1839, Theodore Schwann concluded animals were made of cells.
4. In 1840, J. Purkinje described cell contents as protoplasm; modern
understanding of cell organelles makes “cytoplasm” the preferred term.
5. 1858, another German, Rudolf Virchow, recognized that all cells
came from pre-existing cells.
Figure 03.01
C. How Cells Are Studied

1. Light microscopes use light rays; they are limited in magnification

and resolution.
2. The transmission electron microscope (TEM) uses electrons
passing through the specimen.
a. The wavelength of the electron is 0.00001 that of light, allowing
greater magnification.
b. Specimens must be prepared in thin section; the electrons pass
through to a photographic plate.
3. Scanning electron microscope (SEM) scans electrons across a
metal‑coated specimen; it has a lower magnification than TEM.
4. X-ray crystallography and nuclear magnetic resonance (NMR)
spectroscopy may reveal the shape of molecules.
5. Cytology, the study of cells, has its own methods.
a. Cells are disrupted in a blender and separated by a centrifuge;
organelles are then recovered.
b. Use of radioisotopes allow for tracing of metabolic pathways.
c. Proteins are extracted and purified; antibodies prepared against
the protein can be combined with fluorescent substances to detect
the location of the protein.
Figure 03.02
 3.03
Separation of cell organelles by
centrifugation
3.2. Cell Organization: Complex Organelles
and Energy Transformations
A. Prokaryotic and Eukaryotic Cells

1. Prokaryotes lack a membrane-bound nucleus found in eukaryotes.

2. However, both have DNA, use the same genetic code,
synthesize proteins and use ATP.

B. Major Components of Eukaryotic Cells and Their Functions

1. The cell membrane is the outermost membrane and regulates

the entrance and exit of molecules.
2. A double membrane that separates the nucleus from cytoplasm
encloses the nucleus.
3. Plant cells usually contain plastids for photosynthesis and have a
cellulose-based cell wall.
Figure 03.01
A bacterial cell
Figure 03.05
C. Cell Membrane

1. The current model of membrane structure is the fluid‑mosaic

model.
2. The cell membrane is phospholipid bilayer in which protein
molecules are partially or wholly embedded.
3. The phospholipid molecules have their water-soluble hydrophilic
ends toward the outside and their fat-soluble hydrophobic
portions toward the inside of the membrane.
4. The layer is liquid, providing flexibility; embedded cholesterols
decrease this fluidity.
5. Glycoproteins are proteins with carbohydrates attached.
6. Some proteins catalyze transport of substances such as ions
across the membrane.
7. Others are receptors for specific molecules.
Figure 03.06
D. Nucleus (Figure 3.7)

1. Nuclear envelopes contain less cholesterol than cell membranes;

pores allow relatively large molecules to readily move through.
2. Chromatin is a threadlike material that coils into chromosomes
just before cell division occurs; it contains DNA, protein
histones and nonhistone proteins.
3. Nucleoli are dark‑staining spherical bodies in nucleus and
synthesize ribosomal RNA.
4. After transcription from DNA, ribosomal RNA joins proteins to form
ribosomes.
5. The outer membrane of nucleus is continuous with endoplasmic
reticulum
E. Endoplasmic reticulum

1. The endoplasmic reticulum (ER) is a system of membrane channels

continuous with outer membrane of the nuclear envelope.
2. The space between membranes of nuclear envelope communicates
with channels (cisternae) in ER.
3. The rough ER is studded with ribosomes on cytoplasm side;
products enter cisternae for transport to the Golgi apparatus.
4. Ribosomes on the rough ER synthesize peptides or proteins that
enter the ER cisternae or membrane.
5. Some ribosome products are destined for incorporation into the
cell membrane or for export from the cell.
6. The smooth ER functions to synthesize lipids and phospholipids.
 Endoplasmic reticulum Electron micrograph
showing endoplasmic
Rough ER - studded with ribosomes
reticulum
Smooth ER - few ribosomes
 Figure 03.09

Golgi Complex
Electron micrograph of
a Golgi Complex
Figure 03.10
F. Lysosomes
1. Lysosomes are membrane‑bound vesicles produced by the
Golgi
complex.
2. Lysosomes contain digestive enzymes.
3. These enzymes help digest foreign material or engulfed bacteria:
lysosome vesicles pour enzymes into a food vacuole or
phagosome.
4. They destroy injured or diseased cells; a healthy cell must
maintain the membrane.

G.Contractile vacuoles contain fluid and regulate ions and water.

H. Mitochondria are present in nearly most eukaryotic cells.

1. Mitochondria are bound by a double membrane; inner
membrane
folds (cristae) project into the inner space (matrix).
2. Enzymes on the cristae break down carbohydrate‑derived
products; ATP production occurs here.
3. Mitochondria are self-replicating; their own DNA specifies some
proteins; nuclear DNA codes other proteins.
 Figure 03.11

Electron micrograph of a
mitochondrion
Structure of a typical mitochondrion
Cytoskeleton

The cytoskeleton is a network of filaments and tubules that maintain

support and form.
In many cells, they provide locomotion and translocation of organelles.
The cytoskeleton is composed of microfilaments, microtubules
and intermediate filaments.
Microfilaments are thin, linear structures first recognized in muscle cells.
Actin filaments are long, thin protein fibers that act with several
dozen other proteins.
One of these is myosin; the interaction causes contraction in muscle
and other cells.
Microtubules are composed of the protein tubulin; they move
chromosomes during cell division.
Microtubules radiate out from a microtubule organizing center:
a centrosome.
Centrosomes are not membrane bound. (Figure 3.14)
10. Centrioles are short cylinders with 9 triplets of microtubules;
centrosomes contain two centrioles lying at right angles
to each other.
11. Intermediate filaments are larger than microfilaments and smaller
than microtubules in size.
12. Study of the type of intermediate filaments in cancerous cells can
help in the identification of the original cell type and in the
determination of treatment options.
 Figure 03.12

Electron micrograph
of a cytoskeleton
Figure 03.13

A microtubule
composed of
tubulin molecules
Figure 03.14
Figure 03.14a
J. Surfaces of Cells and Their Specializations

1. Free surfaces of epithelial cells of tubes and cavities sometimes bear

cilia or flagella.
a. Single celled organisms may use cilia or flagella to propel forward.
b. Flagella provide locomotion for male reproductive cells (sperm).
c. Locomotory cilia and flagella both have a cylinder of nine pairs of
microtubules encircling two single microtubules (9 + 2 pattern of
microtubules).
d. At the base of each cilium or flagellum is a basal body (kinetosome).

2. Ameboid movement uses pseudopodia.

a. Ameboid movement is seen in embryonic cells, white blood cells
and
protozoa.
b. Cytoplasmic streaming utilizes actin microfilaments to extend
pseudopodia.
c. Some specialized pseudopodia have cores of microtubules that
assemble and dissemble.
. Cell Junctions (Figure 3.15)

a. In tight junctions, cell membrane proteins fuse to each other;

they hold cells together so tightly that tissues (e.g., epithelial lining
of stomach) form barriers.
b. Desmosomes are “spot welds”; firmly attached to the cytoskeleton
within each cell, are joined by intermediate filaments, and hold cells
together with linker proteins to increase the strength of tissues
in the heart, stomach and bladder. Hemidesmosomes are found at
the base of cells and anchor them to underlying tissue layers.
c. Gap junctions allow cells to communicate; tiny canals between cells
allow the cytoplasm to be continuous in epithelial, nervous and
muscle tissues.
d. Some cell surfaces are laced together with membranes infolding
like a zipper (e.g., epithelia of kidney tubules).

. Microvilli are small finger-like projections of cell membranes

such as those that line the intestine; also called brush borders,
they increase absorptive area. (Figure 3.16)
Figure 03.15
Figure 03.16
Fluid Mosaic Model
 Plasma membrane is composed of both
lipids and globular proteins.

Membrane proteins are not very soluble in
water.
Components of the Cell Membrane

1. Lipid bilayer
2. Transmembrane proteins - proteins that float on or in the lipid
bilayer; provide passageway that allow substances and
information to cross the membrane
3. Network of supporting proteins - intracellular proteins that
reinforce the membrane’s shape
4.Exterior proteins and glycolipids - membrane proteins from
the ER to the golgi complex are transported, and then to
the plasma membrane. ER adds sugar molecules to
membrane proteins creating a “sugar coating”
or “glycocalyx”.
3.3. Membrane Function
A. Plasma Membrane; Dynamic and Selective
1. Also called the plasmalemma, it maintains cellular integrity.
2. It separates the interior environment from the exterior and regulates
molecule traffic flow.
3. It provides many unique functional properties of specialized cells.
4. Internal membranes divide a cell into compartments; they are sites
for most enzymatic reactions.

B. Cell Membrane Function

1. The membrane is the gatekeeper to substances that enter and
exit a cell.
2. Because the interior and exterior are different, the membrane is a
critical controller.
3. Three principal methods are used for crossing a cell membrane:
a. Diffusion along a concentration gradient,
b. Substances bind to a site in a mediated transport system, and
c. Endocytosis encloses a particle in a vesicle that is engulfed.
4. Diffusion and Osmosis (Figure 3.17)

a. Diffusion is movement of particles from higher to lower concentration,

or along a concentration gradient.
b. If a membrane is permeable to a solute, diffusion will continue until
concentrations are equal.
c. Most membranes are selectively permeable, only allowing some
molecules to pass.
d. Most membranes allow free passage of water, gases, urea, and
lipid-soluble solutes.
e. Water-soluble molecules (e.g., sugar), electrolytes and some
macromolecules move across by carrier-mediated processes.
Movement Across Membrane

1. Simple Diffusion - movement of molecules randomly from areas

of greater concentration to areas of lesser concentration until
a state of equilibrium is achieved
e.g. diffusion of a sugar molecule away from a sugar cube
placed in water

2. Facilitated Diffusion - movement of molecules from area of

high concentration to low concentration and requires a
carrier protein
e.g. Polar molecules (not soluble in lipids) may diffuse
through protein channels (pores) in the lipid bilayer.
The protein channels offer a continuous pathway for
specific molecules so that they never come in contact
with the hydrophobic layer of the membrane’s polar
surface.
 i. Marine fish have one-third the solute concentration as seawater; they
are hyposmotic to seawater.
j. A marine fish swimming up a river delta would pass through a region
where external and internal solutes were equal or isosmotic.
k. In freshwater, its blood solutes would be hyperosmotic to the
freshwater.

5. Diffusion through Channels (Figure 3.18)

a. Water and dissolved ions cannot pass through the phospholipid

component of the plasma membrane.
b. Water and ions pass through the membrane by diffusion through
pores
created by transmembrane proteins.
c. Some channels are gated and require a signal to open or close them.
d. Gated ion channels may open or close in response to a signaling
molecule (chemically-gated ion channels) or to the change of an
ionic charge across the plasma membrane (voltage-gated ion
channels).
e. Water channels are called aquaporins.
 
f. Osmosis is movement of water molecules down a concentration
gradient across a membrane.
g. A salt solution in a cell will cause diffusion of water inward until the
increase in weight (hydrostatic or osmotic pressure) of the solution
causes it to be in equilibrium.
h. Osmotic potential is a term used to avoid confusion of term
“osmotic pressure” in the absence of membrane and pure water
reference.
Figure 03.17
i. Marine fish have one-third the solute concentration as seawater; they
are hyposmotic to seawater.
j. A marine fish swimming up a river delta would pass through a region
where external and internal solutes were equal or isosmotic.
k. In freshwater, its blood solutes would be hyperosmotic to the
freshwater.
5. Diffusion through Channels

a. Water and dissolved ions cannot pass through the phospholipid

component of the plasma membrane.
b. Water and ions pass through the membrane by diffusion through
pores created by transmembrane proteins.
c. Some channels are gated and require a signal to open or close
them.
d. Gated ion channels may open or close in response to a signaling
molecule (chemically-gated ion channels) or to the change of an
ionic charge across the plasma membrane (voltage-gated ion
channels).
e. Water channels are called aquaporins.
 
Figure 03.18
6. Carrier-Mediated Transport (Figure 3.19)

a. Special proteins (transporters or permeases) move nutrients and

wastes across the membrane.
b. Permeases form a small passageway for very specific solute molecules.
c. When all transporters become saturated with solutes, the rate of influx
does not increase with more solute; this measures the amount of
transporter molecules.
d. With simple diffusion, the greater the difference in solute concentrations,
the higher the flux.
e. Two mediated transport mechanisms are recognized.
1) Facilitated diffusion permeases assist a molecule (e.g., sugar)
to diffuse that otherwise cannot. (Figure 3.20)
2) Active transport uses energy to transport molecules against the
concentration gradient.
f. Most animal cells require internal potassium levels 20–50 times
outside levels; outside sodium levels may be ten times inside levels.
g. In many cells, sodium and potassium pumping are linked using
the same transporter molecule.
Figure 03.19b
7. Endocytosis (Figure 3.21)

a. All processes (phagocytosis, pinocytosis and receptor-mediated

endocytosis) require energy.
b. Phagocytosis is common among protozoa and lower metazoa.
1) An area of cell membrane coated internally with actin-and-myosin
forms a pocket to engulf material.
2) The membrane-enclosed vesicle detaches from the cell surface for
internal digestion.
c. Pinocytosis
1) Small areas of surface membrane invaginate into tiny vesicles
called caveolae.
2) Specific binding receptors for the molecule or ion are on this cell
surface.
3) It is involved in taking in some vitamins, hormones and growth
factors.
 d. Receptor-mediated endocytosis
1) Plasma membrane proteins bind specific particles called
ligands.
2) This occurs in clathrin-coated pits coated with receptors.
3) Brought within the cell, the pit is uncoated and the ligand
disassociated to be recycled.
4) Some proteins and peptide hormones are brought into cells by
this method.

8.Exocytosis

a. This is the reverse of the invagination and formation of a vesicle.

b. It removes indigestible residues, secretes hormones and
transport substances
Figure 03.21
3.4. Mitosis and Cell Division

A. Cell Types
1. All cells in nearly all multicellular organisms originated from division
of a single cell, the zygote.
2. A zygote is formed from union of egg and sperm, the gametes.
3. Formation of body (somatic) cells by nuclear division is mitosis.
4. Mitosis delivers chromosomes and their DNA to the cell lineage.
5. Although the cell lineage differentiates, the genes not expressed
are still present.
6. Mitosis ensures the equality of genetic material.
7. In animals that reproduce asexually, mitosis transfers
genetic information to progeny.
8. In animals that reproduce sexually, parents produce sex cells
(known as gametes or germ cells) with half the number of chromosomes.
a. This prevents the union of gametes from doubling the number
of parental chromosomes.
b. This requires reduction division or meiosis.
B. Structure of Chromosomes

1. Chromatin
a. Eukaryotic DNA is in the form of chromatin when a cell is not dividing.
b. Chromatin is a complex of DNA with histone and nonhistone proteins.

2. Chromosomes
a. Chromosomes stain deeply with biological dyes.
b. They are of set but varied lengths.
c. A species will have a specific number of chromosomes in all cells
except gametes.
d. Chromosomes are shortened chromatin; they are constricted
at the centromere, which is the location of the kinetochore.
e. DNA packaging allows the cell to fit long strands of DNA into the
small nuclear space; however, when the DNA is packaged
it is not acessible.
C. Phases in Mitosis

1. Cell division involves division of nuclear chromosomes (mitosis) and

cytoplasm (cytokinesis).
a. When a nucleus divides without cytokinesis, a multinucleate cell
results.
b. When several cells fuse, it can also form a multinucleate syncytium.
2. Mitosis is a four-step process with each step merging into the next.
3. When the cell is not actively dividing, it is in interphase during which
DNA replicates and genes are transcribed.
4. Prophase (Figures 3.21–3.23)
a. In early prophase, centrosomes replicate and the two centrosomes
migrate to opposite sides of the nucleus.
b. Microtubules form a football-shaped spindle between the
centrosomes.
c. Other microtubules radiate outward to form asters.
d. Nuclear chromatin condenses into chromosomes; the sister
chromatids were actually formed during interphase.
e. Spindle fibers reach the centrosome and bind to the kinetochore.
5. Metaphase (Figure 3.24)
a. Kinetochore fibers pull condensed sister chromatids to the central
metaphasic plate.
b. Centromeres line up precisely on the equatorial plate; arms of
chromatids dangle.
6. Anaphase
a. A single centromere that holds the chromatids together now splits.
b. Chromosomes move toward their respective poles, pulled by
kinetochore fibers.
c. At the end of anaphase A, microtubules shorten by disassembly of
tubulin units at the kinetochore end.
d. In anaphase B, chromosomes approach respective centrosomes as
the spindle lengthens.
e. Tubulin in microtubules intermingles and pushes the ends of the
spindle apart.
7. Telophase
a. Phase begins as daughter chromosomes reach each pole.
b. Spindle fibers disappear.
c. Chromosomes lose identity and diffuse into chromatin network in
nucleus.
d. Nuclear membranes re-form in daughter cells.
D. Cytokinesis: Cytoplasmic Division

1. During the final stage of nuclear division, a cleavage

furrow appears on cell surface.
2. Microfilaments of actin just beneath the surface draw
the furrow inward.
3. Infolding edges of cytoskeleton meet and fuse,
completing cell division.
E. Cell Cycle (Figure 3.25)

1. Cells undergo cycles of growth and replication.

2. A cell cycle is a mitosis-to-mitosis cycle, the interval between one
cell generation and the next.
3. Interphase
a. Nuclear division occupies 5–10% of the cell cycle; the rest is
interphase.
b. Early concepts of interphase as an inactive stage of rest are
incorrect.
c. DNA replication occurs during interphase.
d. S (for synthesis) stage lasts about 6 of the 18–24 hours of a cell
cycle in a human.
e. Both strands of DNA replicate a complimentary strand.
f. The G1 period precedes the S stage; transfer RNA, ribosomes,
messenger RNA and enzymes are synthesized.
g. The G2 period follows the S stage; spindle and aster proteins
form in preparation for chromosome separation.
4. Embryonic Cells

a. Embryonic cells divide rapidly with no cell growth between

divisions, just subdivision.
b. DNA synthesis may be hundreds of times faster in embryonic cells
than in adult cells.

5. As organisms develop, the cell cycle of most cells lengthens.

a. A nonproliferative phase or G0 ends the cycle.
b. Neurons do not divide further after birth and are in a permanent
G0.
Figure 03.25
6. Cell Cycle Control (Figure 3.26)

a. Regulation of the cell cycle is mediated by cyclin-dependent

kinases (cdk’s).
b. Cyclins are activating subunits of cdk’s.
c. Kinase enzymes add phosphate groups to other proteins to activate
or inactivate them.
d. The passage from one cell cycle to the next is likely regulated by
phosphorylation and dephosphorylation of specific cdk’s and their
interaction with phase specific cyclins.
Figure 03.26
7. Flux of Cells

a. Cell division is very rapid during early development of an organism.

b. The human infant has 2 trillion cells that originated from one
fertilized egg; this represents 42 cell divisions.
c. Five more cell divisions produce an adult with 60 trillion cells.
d. Various cells divide in days, months or years: muscle and nerve
cells stop dividing in childhood or before.
e. A human sheds about 1–2% of the total number of cells daily from
skin, digestive tract, sperm and short-lived red blood cells.

8. Apoptosis
a. Apoptosis is programmed cell death.
b. Apoptosis is necessary for normal development and suicide of
unhealthy cells.
c. Cells shrink, fragment, and then the remains are taken up by
surrounding cells.