2.

Organization of cells

Difference between Eukaryotes & Prokaryote Cells

https://www.youtube.com/watch?v=-ivGgYgAB0A

Eukaryotic cells are enclosed within a thin, selectively permeable plasma membrane. The most
prominent organelle is a spherical or ovoid nucleus, enclosed within two membranes to form a double
layered nuclear envelope Cellular material located between the plasma membrane and the nuclear
envelope is collectively called cytoplasm. Within the cytoplasm are many organelles, such as
mitochondria, Golgi complexes, centrioles, and endoplasmic reticulum. In addition, plant cells
typically contain plastids, some of which are photosynthetic organelles, and plant cells bear a cell wall
containing cellulose outside the plasma membrane.

The fluid mosaic model was first proposed by S.J. Singer and Garth L. Nicolson in 1972 to
explain the structure of the plasma membrane. The model has evolved somewhat over time, but it still
best accounts for the structure and functions of the plasma membrane as we now understand them. The
fluid mosaic model describes the structure of the plasma membrane as a mosaic of components —
including phospholipids, cholesterol, proteins, and carbohydrates—that gives the membrane a fluid
character.
 The main fabric of the membrane is composed of amphiphilic or dual-loving, phospholipid
molecules. The hydrophilic or water-loving areas of these molecules are in contact with the aqueous fluid
both inside and outside the cell. Hydrophobic, or water-hating molecules, tend to be non- polar. A
phospholipid molecule consists of a three-carbon glycerol backbone with two fatty acid molecules
attached to carbons 1 and 2, and a phosphate-containing group attached to the third carbon. This
arrangement gives the overall molecule an area described as its head (the phosphate-containing group),
which has a polar character or negative charge, and an area called the tail (the fatty acids), which has no
charge. They interact with other non-polar molecules in chemical reactions, but generally do not interact
with polar molecules. When placed in water, hydrophobic molecules tend to form a ball or cluster. The
hydrophilic regions of the phospholipids tend to form hydrogen bonds with water and other polar
molecules on both the exterior and interior of the cell. Thus, the membrane surfaces that face the interior
and exterior of the cell are hydrophilic. In contrast, the middle of the cell membrane is hydrophobic and
will not interact with water. Therefore, phospholipids form an excellent lipid bilayer cell membrane that
separates fluid within the cell from the fluid outside of the cell.
 A B

A) The structure of a phospholipid molecule: This phospholipid molecule is composed of a

hydrophilic head and two hydrophobic tails. The hydrophilic head group consists of a phosphate-
containing group attached to a glycerol molecule. The hydrophobic tails, each containing either a
saturated or an unsaturated fatty acid, are long hydrocarbon chains. B) Phospholipid aggregation: In an
aqueous solution, phospholipids tend to arrange themselves with their polar heads facing outward and
their hydrophobic tails facing inward.

Proteins make up the second major component of plasma membranes. Integral proteins (some
specialized types are called integrins) are, as their name suggests, integrated completely into the
membrane structure, and their hydrophobic membrane-spanning regions interact with the hydrophobic
region of the the phospholipid bilayer. Single-pass integral membrane proteins usually have a
hydrophobic transmembrane segment that consists of 20–25 amino acids. Some span only part of the
membrane—associating with a single layer—while others stretch from one side of the membrane to the
other and are exposed on either side. Some complex proteins are composed of up to 12 segments of a
single protein, which are extensively folded and embedded in the membrane. This type of protein has a
hydrophilic region or regions, and one or several mildly hydrophobic regions. This arrangement of regions
of the protein tends to orient the protein alongside the phospholipids, with the hydrophobic region of the
protein adjacent to the tails of the phospholipids and the hydrophilic region or regions of the protein
protruding from the membrane and in contact with the cytosol or extracellular fluid.

Carbohydrates are the third major component of plasma membranes. They are always found on
the exterior surface of cells and are bound either to proteins (forming glycoproteins) or to lipids (forming
glycolipids). These carbohydrate chains may consist of 2–60 monosaccharide units and can be either
straight or branched. Along with peripheral proteins, carbohydrates form specialized sites on the cell
surface that allow cells to recognize each other. This recognition function is very important to cells, as it
allows the immune system to differentiate between body cells (called “self”) and foreign cells or tissues
(called “non-self”). Similar types of glycoproteins and glycolipids are found on the surfaces of viruses and
may change frequently, preventing immune cells from recognizing and attacking them. These
carbohydrates on the exterior surface of the cell—the carbohydrate components of both glycoproteins and
glycolipids—are collectively referred to as the glycocalyx (meaning “sugar coating”). The glycocalyx is
highly hydrophilic and attracts large amounts of water to the surface of the cell. This aids in the interaction
of the cell with its watery environment and in the cell’s ability to obtain substances dissolved in the water.
Key points

 The main fabric of the membrane is composed of amphiphilic or dual-loving, phospholipid

molecules.
 Integral proteins, the second major component of plasma membranes, are integrated completely
into the membrane structure with their hydrophobic membrane-spanning regions interacting with
the hydrophobic region of the phospholipid bilayer.
 Carbohydrates, the third major component of plasma membranes, are always found on the
exterior surface of cells where they are bound either to proteins (forming glycoproteins ) or to
lipids (forming glycolipids).

Key terms
 amphiphilic: Having one surface consisting of hydrophilic amino acids and the opposite surface
consisting of hydrophobic (or lipophilic) ones.
 hydrophilic: Having an affinity for water; able to absorb, or be wetted by water, “water-loving.”
 hydrophobic: Lacking an affinity for water; unable to absorb, or be wetted by water, “water-
fearing.”

For cell structure and process, please watch this summary video
https://www.youtube.com/watch?v=URUJD5NEXC8
Title: Biology: Cell Structure l Nucleus Medical Media

Deep dive into fluid-mosaic model

The fluid-mosaic model is the currently accepted concept describing

plasma membrane structure. By electron microscopy, a plasma
membrane appears as two dark lines, each approximately 3 nm thick,
at each side of a light zone (Figure 3.5). The entire membrane is 8 to
10 nm thick. This image is the result of a phospholipid bilayer, two
layers of phospholipid molecules, all oriented with their water-soluble
(hydrophilic) head regions toward the outside and their fat soluble
(hydrophobic) tail regions toward the inside of the membrane (Figure
3.6). An important characteristic of the phospholipid bilayer is that it is
fluidlike, giving the membrane flexibility and allowing the phospholipid
molecules to move sideways freely within their own monolayer.
Molecules of cholesterol are interspersed in the lipid portion of the
bilayer (Figure 3.6). They make the membrane even less permeable to
water-soluble ions and molecules and decrease membrane flexibility.
One way that cells acclimate to changes in temperature is to add or
remove cholesterol to make the membrane less or more flexible,
respectively, in an effort to maintain function. Glycoproteins (proteins with carbohydrates attached) are
essential components of plasma membranes (Figure 3.6).

Some of polypeptides destined for the plasma membrane,

for lysosomes, or for export from the cell. The outer
membrane of the nuclear envelope is continuous with a
cytoplasmic endomembrane system composed of the ER
(Figure 3.7 and Figure 3.8), the Golgi complex or
apparatus, lysosomes, the plasma membrane, and the
vesicles that pass between them. The space between the
membranes of the nuclear envelope communicates with
the space between the ER membranes (cisterna, pl.,
cisternae). The ER membranes may be covered on their
outer surfaces with ribosomes and are thus designated
rough ER. This is one of the sites of polypeptide
synthesis, mentioned on page 39. Polypeptides
synthesized on rough ER enter the ER cisternae or
membrane and are destined for incorporation into the
plasma membrane (Figure 3.9), for export from the cell, or
for use by the lysosomes. When ER lacks a ribosomal
covering, it is called smooth ER. Smooth ER functions in
synthesis of lipids and phospholipids and as a
detoxification site within cells. The Golgi complex or apparatus (Figure 3.9 and Figure 3.10) is composed
of a stack of membranous cisternae that function in modification and packaging of polypeptide and protein
products produced by rough ER. Small vesicles of ER membrane containing polypeptide or protein
detach and then fuse with sacs on the cis or “forming face” of a Golgi complex. During the modification
process, polypeptides or proteins may be cleaved to form smaller polypeptides or proteins, or they may
have complex carbohydrates added to them as they move through the Golgi cisternae until they reach the
trans or “maturing face” of the complex (Figure 3.9 and Figure 3.10). Finally, vesicles detach from the
trans face of the complex and their contents may be expelled to the outside of the cell, as secretory
products such as from a glandular cell. Some vesicles may carry transmembrane polypeptides or proteins
for incorporation into the plasma membrane, such as receptors or carrier proteins. Others may contain
enzymes that remain in the same cell that produced them. Such vesicles are called lysosomes (literally
“loosening body,” a body capable of causing lysis, or disintegration) (Figure 3.10). Enzymes that they
contain are involved in the breakdown of foreign material, including bacteria engulfed by a cell.
Lysosomes also destroy injured or diseased cells and worn-out cellular components. Their enzymes are
so powerful that they kill the cell that formed them if enough of the lysosome membranes rupture. In
normal cells the enzymes remain safely enclosed within the protective vesicle membranes. Lysosomal
vesicles may pour their enzymes into a larger membrane-bound body containing an ingested food
particle, a food vacuole or phagosome, as is the case in unicellular eukaryotes.

Mitochondria (sing., mitochondrion) (Figure 3.11) are conspicuous organelles present in nearly all
eukaryotic cells. They are diverse in size, number, and shape; some are rodlike, and others are nearly
spherical. They may be scattered uniformly throughout the cytoplasm or localized near regions of high
metabolic activity. A mitochondrion is composed of a double membrane. The outer membrane is smooth,
whereas the inner membrane is folded into numerous platelike or fingerlike projections called cristae
(sing., crista; Figure 3.11), which increase the internal surface area where chemical reactions occur.
These characteristic features make mitochondria easy to identify among organelles. Mitochondria are
often called “powerhouses of cells,” because enzymes located on the cristae catalyze the energy-yielding
steps of aerobic metabolism. Most of a cell’s ATP, the most important energy- transfer molecule of all
cells, is produced in this organelle. Mitochondria are self-replicating. They have a tiny, circular genome,
much like genomes of prokaryotes except much smaller, which contains DNA specifying some, but not all,
proteins of a mitochondrion.
Eukaryotic cells characteristically have a
system of tubules and filaments that
form a cytoskeleton (Figure 3.12 and
Figure 3.13). These provide support and
maintain the form of cells, as well as a
means of cellular locomotion and
movement of macromolecules and
organelles within a cell. The cytoskeleton
is composed of microfilaments,
microtubules, and intermediate
filaments. Microfilaments are thin,
linear

structures, observed distinctly in some protozoan groups and macrophages of the immune system, where
they facilitate cellular locomotion, and in some cell types, such as muscle cells, where they cause cellular
contraction. They are made of a protein called actin. Several dozen other proteins (called actin-binding
proteins or ABPs) bind actin and determine its configuration and behavior in particular cell types. One
ABP is myosin, whose interaction with actin causes contraction in muscle and other cells. Actin is also
instrumental in cytokinesis, the process of cytoplasmic division that takes place at the end of mitosis and
meiosis, as well as in endocytosis and exocytosis. Microtubules, larger than microfilaments, are hollow,
tubular structures composed of a protein called tubulin (Figure 3.13). Each tubulin molecule is actually a
doublet, or dimer, composed of two globular proteins. The molecules are attached head-to-tail to form a
strand, and 13 strands aggregate to form a microtubule. Because the tubulin subunits in a microtubule are
always attached head- to tail, the ends of the microtubule differ chemically and functionally. One end
(called the plus end) both adds and deletes tubulin subunits more rapidly than the other end (the minus
end). Microtubules play a vital role in moving chromosomes toward daughter cells during cell division, and
they are important in intracellular architecture, organization, and transport. They provide a means for
movement of molecules and organelles through the cytoplasm, as well as movement of messenger RNA
from the nucleus to particular positions within the cytoplasm. Microtubules and associated motor proteins
are also important in movement of vesicles between the ER, the Golgi complex, and the plasma
membrane or lysosomes. In addition, microtubules form essential parts of the structures of cilia and
flagella. Microtubules radiate from a microtubule organizing center, the centrosome, near the nucleus.
Centrosomes are not membrane bound. Within centrosomes are found a pair of centrioles (Figure 3.4
and Figure 3.14), which are themselves composed of microtubules. Each centriole of a pair lies at right
angles to the other and is a short cylinder of nine triplets of microtubules. They replicate before cell
division. Although cells of higher plants do not have centrioles, a microtubule organizing center is present.
Intermediate filaments are larger than microfilaments but smaller than microtubules. There are six
biochemically distinct subtypes of intermediate filaments, and their composition and arrangement depend
on the cell type in which they occur. These filaments resist cell stretching and help to hold adjacent cells
together. They are particularly prominent in skin cells associated with desmosomes.
 Surfaces of Cells and Their Specializations

The free surface of epithelial cells sometimes bears either cilia or flagella (sing., cilium, flagellum).
These are motile extensions of the cell surface that may be used to sweep materials past the cell. This
technique is used during feeding in some unicellular eukaryotes and in sponges. In many single-celled
eukaryotes and some small multicellular organisms, they propel the entire organism through a liquid
medium Flagella provide the means of locomotion for male reproductive cells (sperm) of most animals
and many plants. In addition to their function in cell movement and fluid movement around cells, cilia are
proposed to play an integral role in cell signaling, both during development and in the adult organism,
from simple eukaryotes to mammals. Cilia and flagella have different beating patterns, but their internal
structure is the same. With few exceptions, the internal structures of locomotory cilia and flagella are
composed of a long cylinder of nine pairs of microtubules enclosing a central pair. At the base of each
cilium or flagellum is a basal body kinetosome), which is identical in structure to a single centriole. Cilia
and flagella movement mechanisms Many cells move neither by cilia nor by flagella but by ameboid
movement using pseudopodia. Some groups of unicellular eukaryotes, migrating cells in embryos of
multicellular animals, and some cells of adult multicellular animals, such as lymphocytes and
macrophages, show ameboid movement. Cytoplasmic streaming through the assembly and disassembly
of actin microfilaments extends a cytoplasmic process (pseudopodium) outward from the surface of the
cell. Continued streaming in the direction of a pseudopodium brings cytoplasmic organelles into the
process, followed by the rest of the cell, and accomplishes movement of the entire cell. Some specialized
pseudopodia have cores of microtubules, and movement is affected by assembly and disassembly of the
tubulin subunits. Cells covering the surface of a structure (epithelial cells), or cells packed together in a
tissue may have specialized junctional complexes between them. Nearest the free or exposed surface,
the membranes of two cells next to each other appear to fuse, forming a tight junction (Figure 3.15A).
They are formed from rows of transmembrane proteins that bind tightly between adjacent cells. There is
usually a space of about 20 nm between the plasma membranes of intermediate filaments extends into
the cytoplasm connecting desmosomes within a cell, and transmembrane linker proteins extend through
the plasma membrane into the intercellular space to bind the desmosomal discs of adjacent cells
together. Desmosomes are not seals but seem to increase the strength of the tissue. Many are found
between the cells of the skin in vertebrates. Hemidesmosomes (Figure 3.15A) occur at the base of cells
and anchor them to underlying connective tissue layers. Gap junctions (Figure 3.15A), rather than
serving as points of attachment, provide a means of intercellular communication. They form tiny canals
between cells, so that their cytoplasm becomes continuous, and small molecules and ions can pass from
one cell to the other. Gap junctions may occur between cells of epithelial, nervous, and muscle tissues
(see Chapter 9, p. 190). Another specialization of cell surfaces occurs where plasma membranes of
adjacent cells infold and interdigitate very much like a zipper. These infoldings are especially common in
epithelial cells of kidney tubules and serve to increase the surface area of the cells for absorption or
secretion. The distal or apical boundaries of some epithelial cells, as seen by electron microscopy, show
regularly arranged microvilli (sing., microvillus). They are small, fingerlike projections consisting of
tubelike evaginations of the plasma membrane with a core of cytoplasm containing bundles of actin
microfilaments (Figure 3.15A and B). They are seen clearly in the lining of the intestine where they greatly
increase the absorptive and digestive surface. Such specializations are often termed brush borders due to
their appearance when seen by light microscopy.

Cilia and Flagella

https://www.youtube.com/watch?v=sVHUO89-sXg
 Membrane Function

The incredibly thin, yet sturdy, plasma membrane that encloses every cell is vitally important in
maintaining cellular integrity. Plasma membranes (also called the plasmalemma) form dynamic structures
having remarkable activity and selectivity. They are a permeability barrier that separates the interior from
the external environment. They regulate the flow of molecules into and out of the cell, and provide many
of the unique functional properties of specialized cells, such as allowing communication with the
surrounding extracellular fluid and other cells. Membranes within a cell surround a variety of organelles,
and thus divide the cell into numerous compartments. If all membranes present in one gram of liver tissue
were spread out flat, they would cover 30 square meters! Internal membranes share many structural
features of plasma membranes and are the site for many of a cell’s enzymatic reactions and internal
communication systems.
A plasma membrane acts as a selective gatekeeper for entrance and exit of many substances
involved in cell metabolism. Some substances can pass through with ease, others enter slowly and with
difficulty, and still others cannot enter at all. Because conditions outside a cell are different from and more
variable than conditions within a cell, it is necessary that passage of substances across the membrane be
rigorously controlled. We recognize three principal ways that a substance may enter across a cell
membrane: (1) by diffusion along a concentration gradient; (2) by a mediated transport system, in
which the substance uses a specific transmembrane protein that assists it across the membrane; and (3)
by endocytosis, in which the substance is enclosed within a vesicle that forms from the membrane
surface and detaches inside the cell.

Diffusion
Diffusion is a movement of particles or molecules from an area of higher concentration to an area of lower
concentration of the particles or molecules, thus tending to equalize the concentration throughout the area
of diffusion. If a living cell surrounded by a membrane is
immersed in a solution having a higher concentration of solute molecules than the fluid inside the cell, a
concentration gradient instantly exists between the two fluids across the membrane. If the membrane is
permeable to the solute, there is a net movement of solute toward the inside of the cell, the side having
the lower concentration. The solute diffuses “downhill” across the membrane until its concentrations on
each side are equal. Most cell membranes are selectively permeable, often, permeable to water but
variably permeable or impermeable to solutes. In free diffusion it is this selectiveness that regulates
molecular traffic. As a rule, gases (such as oxygen and carbon dioxide), urea, and lipid- soluble solutes
(such as fats, fatlike substances, and alcohol) are the only solutes that can diffuse through biological
membranes with any degree of freedom. Water and many water-soluble molecules readily pass-through
membranes, but movements cannot be explained by simple diffusion. Sugars, water, many electrolytes,
and macromolecules are moved across membranes by mediated transport systems.

Osmosis

If we place a membrane between two unequal

concentrations of solutes to which the membrane is
impermeable, water flows through the membrane
from the more dilute to the more concentrated
solution. The water molecules move across the
membrane down a concentration gradient from an
area where the water molecules are more
concentrated to an area on the other side of the
membrane where they are less concentrated. This is
osmosis—diffusion of water molecules across a
membrane. Water flows across the plasma
membrane via osmosis because the cytoplasm and
surrounding external environment are often at
differing concentrations. The process of osmosis can
be readily demonstrated using red blood cells. If red
blood cells are placed in a container of pure water,
after a short time the red blood cells will swell and
burst (lyse). This occurs because of a pressure
buildup inside the cell due to a net movement of
water into the cells—the cell membrane is unable to
withstand this high pressure and ruptures. Inside the
cells the cytoplasm contains large macromolecules,
as well as salts and water molecules, while the
container contains only water molecules. Thus, the
concentration of water is less inside the cells
because some of the available space is
occupied by the nondiffusible
macromolecules and salt ions. A
concentration gradient therefore exists for
water molecules in the system. Water diffuses from the region of greater concentration of
water (pure water in the container) to the region of lesser water concentration inside the
cells (Figure 3.17A). In the experiment just described, the region of highest water
concentration (in the container) is termed hypotonic to the cytoplasm due to low levels of
(or no) salt ions or macromolecules in the container, while the cytoplasm is hyperosmotic
to the water in the container due to the presence of high levels of macromolecules and salt
ions. The pressure that resists the flow of water into the cytoplasm is termed the osmotic
pressure. If red blood cells were placed in a solution of water and salts that was similar in
concentration to the cytoplasm, then the solution would be isotonic or isoosmotic to the
cytoplasm and there would be no net movement of water (Figure 3.17B). If red blood cells
were placed in a solution of water and salt that was at a higher concentration (hypertonic)
than that of the red blood cell cytoplasm, then water molecules would flow out of the
cytoplasm and the cell would shrink (Figure 3.17C). The cytoplasm would be hypoosmotic
to the solution in the container due to the presence of fewer macromolecules and salt ions
than in the container. The concept of osmosis is very important in understanding how
animals control their internal fluid and solute environment. For example, unicellular
eukaryotes possess a contractile vacuole that functions in osmoregulation. In particular,
freshwater forms accumulate water via osmosis because their cytoplasm is hyperosmotic
(a higher solute concentration) compared with the surrounding environment. The
contractile vacuole rapidly fills with this excess water and expels it through the plasma
membrane by exocytosis. Osmoregulation is critical for marine bony fishes that maintain a
solute concentration in their blood about one-third of that in seawater; they are
hypoosmotic to seawater. If a fish swims into a river mouth and then up a freshwater
stream, as salmon do, it passes through a region where its blood solutes are equal in
concentration to those in its environment (isoosmotic), then enters freshwater, where its
blood solutes are hyperosmotic to those in its environment. It must have physiological
mechanisms to avoid net loss of water in the sea and gain of water in the river.
 Diffusion and Osmosis
https://www.youtube.com/watch?v=6DsaD3LBjUc
 Carrier-Mediated Transport

The plasma membrane is an effective barrier

to free diffusion of most molecules of
biological significance, yet it is essential that
such materials enter and leave a cell.
Nutrients such as sugars and materials for
growth such as amino acids must enter a cell,
and wastes of metabolism must leave. Such
molecules utilize a mediated transport
system composed of transmembrane
proteins called transporters, or carriers.
Transporters enable solute molecules to
cross the phospholipid bilayer (Figure
3.18A). They are usually quite specific,
recognizing and transporting only a limited
group of chemical substances or perhaps
even a single substance. At high
concentrations of solute, mediated transport
systems show a saturation effect. This means
simply that the rate of influx reaches a
plateau beyond which increasing the solute
concentration has no further effect on influx
rate (Figure 3.18B). This is evidence that the
number of transporters available in a
membrane is limited When all transporters
become occupied by solutes, the rate of transport is at a maximum and it cannot be
increased. Simple diffusion shows no such limitation; the greater the difference in solute
concentrations on the two sides of the membrane, the faster the influx. Two distinctly
different kinds of carrier-mediated transport mechanisms are recognized: (1) facilitated
diffusion or facilitated transport, in which a transporter assists a molecule to diffuse
through the membrane that it cannot otherwise penetrate, and (2) active transport, in
which energy from ATP is supplied to the transporter system to transport molecules in the
direction opposite to a concentration gradient (Figure 3.19). Facilitated diffusion or
transport therefore differs from active transport in that it supports movement only in a
downhill direction (in the direction of a concentration gradient) and requires no metabolic
energy from ATP to drive the transport system.
In many animals facilitated diffusion aids transport of glucose (blood sugar) into
body cells that oxidize it as a principal energy source for the synthesis of ATP. The
concentration of glucose is greater in blood than in the cells that consume it, favoring
inward diffusion, but glucose is a water-soluble molecule that does not, by itself, penetrate
cell membranes rapidly enough to support the metabolism of many cells; the carrier-
mediated transport system increases the inward flow of glucose. In active transport,
molecules are moved against the forces of passive diffusion. Active transport always
involves an expenditure of energy (from ATP) because materials are transported against a
concentration gradient. Among the most important active-transport systems in all animals
are those that maintain sodium and

potassium ion gradients between cells and the surrounding extracellular fluid or external
environment. Most animal cells require a high internal concentration of potassium ions for
protein synthesis at the ribosome and for certain enzymatic functions. The potassium ion
concentration may be 20 to 50 times greater inside a cell than outside. Sodium ions, on the
other hand, may be 10 times more concentrated outside a cell than inside. This sodium
gradient forms the basis for electrical signal generation within the nervous system of
animals. Both of these ionic gradients are maintained by active transport of potassium ions
into and sodium ions out of the cell. In many cells outward transport of sodium is linked to
inward transport of potassium; the same transporter protein does both. As much as 10% to
40% of all energy produced by cells is consumed by the sodium-potassium pump (Figure
3.19).

Endocytosis

Endocytosis, the ingestion of material by cells, is a collective term that describes three
similar processes: phagocytosis, pinocytosis, and receptor-mediated endocytosis (Figure
3.20). They are pathways for specifically internalizing solid particles, small molecules and
ions, and macromolecules, respectively. All require energy and thus are forms of active
transport.

Phagocytosis, which literally means “cell eating,” is a common method of feeding

among unicellular forms: sponges, cnidaria, and flatworms. It is also the way in which
white blood cells (leukocytes) and macrophages engulf cellular debris and invading
microbes or other pathogens in the blood. During phagocytosis, an area of the plasma
membrane, coated externally with specific receptors and internally with actin and actin-
binding proteins, forms a pocket that engulfs the solid material. The membrane-enclosed
vesicle, a food vacuole or phagosome, then detaches from the cell surface and moves into
the cytoplasm where it fuses with lysosomes, its contents are digested by lysosomal
enzymes, and useful products are absorbed from the lysosome across its membrane by
diffusion or carrier-mediated transport.

Pinocytosis is similar to phagocytosis except that small areas of the surface

membrane are invaginated into cells to form tiny vesicles. The invaginated pits and vesicles
are called caveolae (ka-vēə-lē). Specific binding receptors for the molecule or ion to be
internalized are concentrated on the cell surface of caveolae. Pinocytosis apparently
functions for intake of at least some vitamins, hormones, and growth factors. Lysosomes
fuse with these caveolae, causing digestion of their contents prior to absorption into the
cytoplasm. Similar mechanisms may be important in translocating substances from one
side of a cell to the other (see Exocytosis, next section), as occurs during some capillary
exchange. This variation of pinocytosis is often called transcytosis. In this case, contents of
the caveolae would remain largely unchanged as they are translocated across the cell.
Receptor-mediated endocytosis is a specific mechanism for bringing large molecules within
the cell. Proteins of the plasma membrane specifically bind particular molecules (termed
ligands), which may be present in the extracellular fluid in very low concentrations. The
invaginations of the cell surface that bear the receptors are coated on the cytoplasmic side
of the membrane with a protein. One example of a coating protein is called clathrin; hence,
they are called clathrin-coated pits. As a clathrincoated pit with its receptor- bound ligand
invaginates and is brought within the cell, it is uncoated, the receptor and the ligand are
dissociated, and the receptor and surrounding membrane are recycled back to the surface
membrane. Lysosomes fuse with the remaining vesicle, now called an endosome, and its
contents are digested and absorbed into the cytoplasm. Some important proteins, peptide
hormones, and cholesterol are brought into cells in this manner. In phagocytosis,
pinocytosis, and receptor-mediated endocytosis, some amount of extracellular fluid is
necessarily trapped in the vesicle and nonspecifically brought within the cell. We call this
bulk-phase endocytosis.

Exocytosis

Just as materials can be brought into a cell by membrane invagination and formation of a
vesicle, the embrane of a vesicle within a cell can fuse with the plasma membrane and
extrude its contents to the surrounding medium. This is the process of exocytosis. This
process occurs in various cells to remove undigestible residues of substances brought in by
endocytosis, to secrete substances such as hormones (see Figure 3.10), to recycle
membrane receptors and membranes, as mentioned in receptor-mediated endocytosis
(Figure 3.20), and to transport a substance completely across a cellular barrier
(transcytosis). Actin and actin-binding proteins are essential cytoskeletal components in
the processes of endocytosis and exocytosis.