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Cell Membrane Structure and Organelles
Cell Membrane Structure and Organelles
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Part II
Principles of Individual Cell Function
Chapter 5
Cell Membrane Structure and Organelles
Cell structures consist of biological membranes essentially mobile lipid bilayers to
which many membrane proteins at t ach. The cell membrane separates the interior
of the cell from the outer environment, acts as a barrier against exterior forces, and
regulates the ow of materials and information across the membrane.
Many bacteria, including E. coli, have one biological membrane the cell
membrane. Nucleated cells of organisms such as yeasts, animals and plants
have organelles surrounded by biological membranes intert wined with particular
proteins, forming a unique intercellular environment. Many intracellular functions
operate smoothly through the collaboration of organelles. In addition, biological
membranes are not static; they are continuously created and move/fuse together,
thus creating ow bet ween membranes. Cells can grow, divide and perform a
range of movements thanks to the uid structure of their membranes.
I . Cel l Membrane St ruct ure
Prokaryotic and Eukaryotic Cells
Membranes found in cells are called biological membranes. Their basic structure
consists of a lipid bilayer to which many proteins are bound. The cell membrane
surrounding a cell is also a biological membrane. It is believed that primitive cells
were created as a result of genetic material and its reproduction structures
becoming surrounded by membranes. As shown in Figure 5-1, bacteria (an
example of prokaryotic cells) have a simple structure with just a plasma
membrane. Conversely, animal and plant cells (examples of eukaryotic cells) are
surrounded by a plasma membrane containing as shown in Table 5-1
organelles surrounded by double lipid bilayers consisting of inner and outer
membranes (such as the nucleus, mitochondria and chloroplasts) and organelles
surrounded by a single lipid bilayer (such as the plasma membrane, endoplasmic
reticula, Golgi apparatuses and lysosomes).
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Fi gure 5- 1 Prokaryot i c and eukaryot i c cel l s
Tabl e 5- 1
Mai n f unct i ons of eukaryot i c cel l compart ment s
di vi ded by cel l membranes
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Organelles of Eukaryotic Cells
Plasma membrane
The biological structure that separates the interior of a cell from its outer environment
is called the plasma membrane. Bacteria and animal cells are separated from
the outside by a lipid bilayer plasma membrane, whereas plant cells have a
strong cell wall outside the plasma membrane (Fig. 5-1). The plasma membrane
is characteristic in that many of the membrane proteins located on its outer surface
are modied by sugar chains. The plasma membrane has uidit y, and parts of it
continually diffuse into the cell. In this membrane (as discussed later in the chapter),
many channels and transporters carry materials, and receptors pass information
from the outer environment to the interior of the cell.
Nucleus and Nuclear Envelope
Eukaryotic cells normally have one nucleus, which contains a genome a
complete set of hereditary information for an organism. In the nucleus, DNA
replication and RNA transcription occur. Linear DNA and binding proteins (e.g.,
histones) form a complex (i.e., chromatin) in the nucleus. There are t wo t ypes of
nuclear chromatin: euchromatin a lightly packed form under active transcription
and heterochromatin, a tightly packed form in which transcription is limited.
During cell division, DNA in the nucleus becomes increasingly condensed until it
forms rod-like structures called chromosomes that are then distributed to the t wo
daughter cells. The nuclear envelope has many holes known as nuclear pores,
which control the movement of materials across the envelope. As an example,
mRNA generated by transcription passes through the nuclear pores out into the
cytoplasm, where it is translated into proteins (see Chapter 3).
Endoplasmic Reticula and Golgi Apparatuses
Endoplasmic reticulam and Golgi apparatuses are involved in the synthesis and
transport of secretory proteins and the constituents of membranes. Endoplasmic
reticula synthesize and process proteins by sorting, adding sugar chains and
providing other modications. Such endoplasmic reticula have a ribosome-rich
surface, and are called rough endoplasmic reticula. Those without at t ached
ribosomes are known as smooth endoplasmic reticula.
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Endoplasmic reticula are connected to the outer membrane of the nuclear
envelope and form a mesh-like structure. Consisting of a stack of at t ened
membrane structures, Golgi apparatuses are located near endoplasmic reticula,
adding sugar chains to membrane proteins and secretory proteins as well as
sorting proteins. Transportation of lipids and membrane proteins bet ween
organelles is performed by small bag-like structures made of biological membranes
called transport vesicles (see Fig. 5-10).
Endosomes and Lysosomes
Endosomes and lysosomes play a role in the incorporation and digestion of
extracellular materials. Part of the membrane invaginates and pinches off to form
an endosome inside the cell. This process is called endocytosis. Endocytosed
proteins and lipids are transported to organelles aft er being sorted within
endosomes or digested within lysosomes. Plant vacuoles have functions similar to
those of lysosomes, and regulate cell turgor. Lysosomes contain enzymes that
digest nucleic acids, proteins and lipids, and the interior of lysosomes is kept
acidic (pH 5) by proton pumps. Vesicular transport also plays an important role
in the movement of lipids and membrane proteins bet ween these organelles.
Mitochondria, Chloroplasts and Peroxisomes
The inner mitochondrial membrane is compartmentalized into many cristae. The
space it encloses is called a matrix (see Fig. 8-2 in Chapter 8), and contains
DNA unique to mitochondria. Mitochondria are found in almost all eukaryotic
cells, and perform oxidative phosphorylation via the electron transport chain to
synthesize ATP.
Chloroplasts are at t ened, disk-shaped organelles found in plants and algae, and
play a role in photosynthesis. The material inside the inner membrane is called the
stroma, which contains stacks of at t ened, bag-like structures called thylakoids that
perform photosynthesis. The stroma contains DNA unique to chloroplasts.
Peroxisomes contain many oxidases, and perform lipid oxidization and the
metabolism of various materials. As an example, peroxisomes in plants synthesize
carbohydrates from stored lipids.
Vesicular transport does not occur bet ween mitochondria and chloroplasts or
bet ween peroxisomes and other organelles.
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I I . Li pi ds and Membrane Prot ei ns i n Bi ol ogi cal Membranes
Characteristics of a Lipid Bilayer
Lipid is a collective term for materials that do not dissolve readily in water but do
so easily in organic solvents. The lipids that constitute biological membranes are
made of hydrocarbon chains with hydrophilic heads and hydrophobic tails. The
most abundant constituents of biological membranes are phospholipids (Fig.
5-2A), and other constituents include sterols and glycolipids.
When placed in water, the hydrophilic heads of phospholipids face the water,
while the hydrophobic tails line up against one another, thus forming a bylayer
or a ball-shaped structure (i.e., a micelle) (Fig. 5-2B).
Lipid molecules in the bilayer continually move in a horizontal direction, but do
not move bet ween the inner and outer sides of the membrane except under
special conditions. As shown in Figure 5-2C, lipid composition differs bet ween
the inner and outer sides of the membrane, and in animal cells glycolipids are
known to be found more on the outside.
Fi gure 5- 2 Li pi ds of bi ol ogi cal membranes
(A) The phospholipid molecule that constitutes membranes
(B) Model of lipid bilayer and ball-shaped micelles
(C) Membrane lipids readily flow in a horizontal direction, but not bet ween the inside and outside of the membrane.
( A)
( B) ( C)
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Membrane Proteins
In cells, the plasma membrane which has a lipid bilayer as its basic framework
forms a ball-shaped structure and separates the cytoplasm from the aqueous
system outside the cell. Electron microscopy shows that the plasma membrane
consists of a lipid bilayer to which numerous membrane proteins are at t ached.
Membrane proteins take various forms; as shown in Figure 5-3, single- or
multipass transmembrane proteins, tunnel-shaped proteins similar to ion channels,
and proteins that at t ach to one side of the lipid membrane.
For transmembrane proteins, t wo major polypeptide structures are known. One of
these is the -helix structure (Fig. 5-3A). Its highly hydrophobic surface, which has
many amino acids at t ached that have hydrophobic side chains (e.g., leucine and
Fi gure 5- 3 Types of at t achment f or cel l - membrane prot ei ns
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isoleucine), faces the lipids. Multiple -helices can form a path that penetrates the
plasma membrane. In this case, as shown in Figure 5-3B, many hydrophobic
amino acids are located on the outside facing the lipids, whereas many hydrophilic
amino acids are found on the inside facing the aqueous solution.
Membrane proteins can also form a hole that penetrates the plasma membrane
using a -sheet structure (Fig. 5-3C). In this case, hydrophobic amino acids are
concentrated on the outside facing the lipid bilayer, and hydrophilic amino acids
are located on the inside of the tunnel. This sheet structure is called a -barrel
because of its barrel-like shape.
These membrane proteins are involved in the transport of ions and chemical
compounds through the membrane. They also stabilize the membrane by
undercoating it, act as receptors that pass extracellular information to the cell (see
Chapter 9), and bind to the cytoskeleton by connecting to specic lipids in the
plasma membrane (see Chapter 6).
Some proteins, as shown in Figure 5-3E, do not have a structure that penetrates the
membrane; rather, they bind to it via fat t y acids (see the Column on p.96). Many
proteins that form complexes with membrane proteins also accumulate on the
membrane (Fig. 5-3F).
Proteins located on the outside of the plasma membrane of eukaryotic cells oft en
have oligosaccharide chains at t ached. Since extracellular uid, such as blood,
contains many proteases, the at t achment of sugar chains makes it difcult for
proteases to digest the proteins, thus stabilizing the proteins outside the cell.
Sugar chains oft en found in the proteins located on the cell surface are also used
as cell markings, and play an important role when cells recognize and at t ach
themselves to each other.
I I I . Funct i ons of Bi ol ogi cal Membranes
Barrier Functions and Selective Transport of Materials
A cell is a compartment separated from its outer environment by a plasma
membrane. The interior of a eukaryotic cell is also compartmentalized into many
organelles, and different reactions occur in each compartment.
Since lipids are the basic constituents of biological membranes (as shown in Fig.
5-4A), small, electrically non-charged solutes such as ethanol, oxygen and
carbon dioxide can pass through a lipid bilayer by simple diffusion following the
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concentration gradient. Water-soluble ions (Na
+
, K
+
and Cl
), sugars (e.g.,
glucose) and amino acids, however, cannot pass through the membrane. Proteins
high molecules are also unable to penetrate. By rigorously regulating the
transport of these materials, the plasma membrane keeps the intracellular
environment relatively stable even when the outside environment changes.
As shown in Figure 5-4B, the plasma membrane has membrane proteins such as
transporters that transport specic molecules, and channels that let specic
molecules pass. Most solutes can pass through the membrane only when
transported by membrane proteins. In this case, passive transport (i.e., transport
following the concentration gradient) occurs without relying on energy, whereas
Fi gure 5- 4
The pl asma membrane a l i pi d
bi l ayer t hat serves as a barri er
agai nst sol ut es and regul at es t he
passi ng of mat eri al by t ransport ers
and channel s
The plasma membrane lets small, electrically
non-charged solutes with a low molecular weight
pass, but forms a barrier against ions and large
solutes that have a high molecular weight (A) and
regulates transport by transporters and channels.
Passive transport occurs when the concentration
gradient is followed, but active transport, which
requires energy, takes place using ATP-derived or
other energies when transport occurs against the
concentration gradient (B).
( B)
( A)
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active transport (i.e., transport against the concentration gradient) requires
energy. Active transport is performed by transporters.
The mechanism of active transport, which uses energy, is shown in Figure 5-5.
Animal cells have a higher K
+
concentration and a lower Na
+
concentration than
blood. To maintain these conditions, an Na
+
/K
+
pump a t ype of transporter
transports Na
+
ions to the outside and K
+
ions to the inside of the cell against
the concentration gradient using the energy generated when ATP is hydrolyzed
into ADP. A pump protein containing Na
+
at t ached to (1) is phosphorylated
using the energy generated through the hydrolysis of ATP and changes its structure
(2), releases Na
+
to the outside of the cell (3), and catches Ka
+
instead (4). The
pump protein is then dephosphorylated (5) and changes its structure, releasing
K
+
into the cell (6). These reactions occur continually, using 30% of the energy
generated within the cell in some animal cells.
In this case, the energy of ATP is used t wice for the structural changes of pump
proteins caused by phosphorylation and dephosphorylation, which allows the
transport of Na
+
and K
+
.
Fi gure 5- 5
Cont i nual t ransport of i ons by t he
Na
+
/K
+
pump usi ng ATP
Each time the Na
+
/K
+
pump hydrolyzes one
molecule of ATP, three Na
+
ions leave the cell and
t wo K
+
ions enter it.
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The nature of the plasma membrane varies greatly depending on its
lipid composition. Main lipids of the plasma membrane include
phospholipids, sterols and glycolipids. A t ypical phospholipid is
phosphatidylcholine (Column Fig. 5-1A). The head, consisting of
choline and phosphate, is connected by glycerol with hydrocarbon
tails that look like t wo legs. The uidit y of the membrane changes
signicantly depending on how many double bonds these
hydrocarbon tails have. A double bond formed bet ween carbon
and carbon bends the hydrocarbon chain from that point.
The most abundant constituent in the membrane of animal cells is
cholesterol (Column Fig. 5-1B). This short, hard molecule is mainly
located on the inside of the plasma membrane, and lls the gaps
created by the double-bond bending of phospholipids.
The plasma membrane has sites called rafts where cholesterol and
glycolipids are concentrated. In raft s, membrane lipids become like
liquid crystal and hence have low uidit y. It is known that membrane
proteins involved in signaling tend to be concentrated in raft s.
Lipid-modied proteins move from the cytoplasm to the inside of
raft s, whose outside has many glycolipids and at t racts glycolipid-
modied proteins.
Membrane regions rich in cholesterol are associated with
neurological and immunological functions, Alzheimer's disease and
viral infection, and are therefore quite well known. Lipid-rich regions
of the plasma membrane, such as raft s, are known as the micro-
domains of the membrane.
( B)
( A)
Col umn
Chol esterol i n the Pl asma membrane
( C)
Col umn Fi gure 5- 1
Chol est erol i n
bi ol ogi cal membranes
(C)Cholesterol located inside the curve of
a phospholipid tail created by a
hydrocarbon double bond makes the
membrane rigid.
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Membrane Potential
In a resting (non-excited) cell, electrical potential difference exists whereby the
interior of the cell separated by the plasma membrane has negative potential
(i.e., resting membrane potential). This is due to the difference in ion concentration
bet ween the inside and the outside of the cell (Table 5-2) and the selective
permeabilit y of the plasma membrane for ions (Fig. 5-6). With the plasma
membrane in a resting state, certain K
+
channels are kept open; the membrane
can therefore be thought of as a semipermeable membrane that allows K
+
ions to
pass. When a difference in the concentration of K
+
exists bet ween either side of
such a semipermeable membrane, the resting membrane potential is calculated
as approximately 90 mV by the Nernst equation (see the Column on p.98). This
potential is similar to the value for an actual cell, but differs slightly because small
amounts of Na
+
and Cl