• The plasma membrane is the boundary that

separates the living cell from its nonliving

surroundings
• The plasma membrane exhibits selective
permeability, allowing some substances to cross
it more easily than others
Cellular membranes are fluid mosaics of lipids and
proteins
• Phospholipids are the most abundant lipid in the
plasma membrane
• Phospholipids are amphipathic molecules,
containing hydrophobic and hydrophilic regions
• The fluid mosaic model states that a membrane
is a fluid structure with a “mosaic” of various
proteins embedded in it
 WATER

Hydrophilic
head

Hydrophobic
tail

WATER
 Hydrophilic region
of protein

Phospholipid
bilayer

Hydrophobic region of protein

The Fluidity of Membranes

• Phospholipids in the plasma membrane can move

within the bilayer
• Most of the lipids, and some proteins, drift
laterally
• Rarely does a molecule flip-flop transversely
across the membrane
 Lateral movement Flip-flop
(~107 times per second) (~ once per month)
Movement of phospholipids
• As temperatures cool, membranes switch from a
fluid state to a solid state
• The temperature at which a membrane solidifies
depends on the types of lipids
• Membranes rich in unsaturated fatty acids are
more fluid than those rich in saturated fatty acids
• Membranes must be fluid to work properly; they
are usually about as fluid as salad oil
 Fluid Viscous

Unsaturated hydrocarbon Saturated hydro-

tails with kinks carbon tails
Membrane fluidity
• The steroid cholesterol has different effects on
membrane fluidity at different temperatures
• At warm temperatures (such as 37°C), cholesterol
restrains movement of phospholipids
• At cool temperatures, it maintains fluidity by
preventing tight packing
 Cholesterol
Cholesterol within the animal cell membrane
• Some proteins in the plasma membrane can drift
within the bilayer
• Proteins are much larger than lipids and move
more slowly
• Proteins are anchored in lipid rafts

• Can move in one direction

 Membrane proteins

Mouse cell Mixed

proteins
Human cell after
Hybrid cell 1 hour
Membrane Proteins and Their Functions

• A membrane is a collage of different proteins

embedded in the fluid matrix of the lipid bilayer
• Proteins determine most of the membrane’s
specific functions
• Peripheral proteins are not embedded

• Integral proteins penetrate the hydrophobic core

and often span the membrane
Fibers of
extracellular
matrix (ECM)

Glycoprotein

Carbohydrate

Glycolipid
EXTRACELLULAR
SIDE OF
MEMBRANE

Cholesterol

Microfilaments Peripheral
of cytoskeleton proteins Integral
protein
CYTOPLASMIC SIDE
OF MEMBRANE
• Integral proteins that span the membrane are
called transmembrane proteins.
• Are built of three segments.
– The cytosolic and exoplasmic domains have
hydrophilic exterior surfaces that interact with the
aqueous solutions on the cytosolic and
exoplasmic faces of the membrane. These
domains resemble other water-soluble proteins
in their amino acid composition and structure.
• Membrane-spanning domain (3-nm-thick )
contains many hydrophobic amino acids whose
side chains protrude outward and interact with
the hydrocarbon core of the phospholipid
bilayer.
 EXTRACELLULAR
SIDE
N-terminus

C-terminus
CYTOPLASMIC
SIDE
 Helix
• In addition, most transmembrane proteins are
glycosylated with a complex branched sugar
group attached to one or several amino acid
side chains.

• Invariably these sugar chains are localized to

the exoplasmic domains.
• Six major functions of membrane proteins:
– Transport
– Enzymatic activity
– Signal transduction
– Cell-cell recognition
– Intercellular joining
– Attachment to the cytoskeleton and
extracellular matrix (ECM)
 Signal
Enzymes

Receptor
ATP

Transport Enzymatic activity Signal transduction

 Glyco-
protein

Cell-cell recognition Intercellular joining Attachment to the

cytoskeleton and extra-
cellular matrix (ECM)
The Role of Membrane Carbohydrates in Cell-Cell
Recognition
• Cells recognize each other by binding to surface
molecules, often carbohydrates, on the plasma
membrane
• Membrane carbohydrates may be covalently
bonded to lipids (forming glycolipids) or more
commonly to proteins (forming glycoproteins)
• Carbohydrates on the external side of the plasma
membrane vary among species, individuals, and
even cell types in an individual
Synthesis and Sidedness of Membranes

• Membranes have distinct inside and outside faces

• The asymmetrical distribution of proteins, lipids

and associated carbohydrates in the plasma
membrane is determined when the membrane is
built by the ER and Golgi apparatus
 ER

Transmembrane
glycoproteins

Secretory
protein

Glycolipid

Golgi
apparatus

Vesicle

Plasma membrane:
Cytoplasmic face
Extracellular face
Transmembrane
Secreted glycoprotein
protein

Plasma membrane:
Membrane structure results in selective
permeability
• A cell must exchange materials with its
surroundings, a process controlled by the plasma
membrane
• Plasma membranes are selectively permeable,
regulating the cell’s molecular traffic
The Permeability of the Lipid Bilayer

• Hydrophobic (nonpolar) molecules, such as

hydrocarbons, can dissolve in the lipid bilayer
and pass through the membrane rapidly
• Polar molecules, such as sugars, do not cross
the membrane easily
Transport Proteins

• Transport proteins allow passage of hydrophilic

substances across the membrane
• Some transport proteins, called channel proteins,
have a hydrophilic channel that certain molecules
or ions can use as a tunnel
• Channel proteins called aquaporins facilitate the
passage of water
• Other transport proteins, called carrier proteins,
bind to molecules and change shape to shuttle
them across the membrane
• A transport protein is specific for the substance it
moves
Passive transport is diffusion of a substance across a
membrane with no energy investment

• Diffusion is the tendency for molecules to

spread out evenly into the available space
• Although each molecule moves randomly,
diffusion of a population of molecules may exhibit
a net movement in one direction
• At dynamic equilibrium, as many molecules
cross one way as cross in the other direction
Molecules of dye Membrane (cross section)

WATER

Net diffusion Net diffusion Equilibrium

Diffusion of one solute

• Substances diffuse down their concentration
gradient, the difference in concentration of a
substance from one area to another
• No work must be done to move substances down
the concentration gradient
• The diffusion of a substance across a biological
membrane is passive transport because it
requires no energy from the cell to make it happen
 Net diffusion Net diffusion Equilibrium

Net diffusion Net diffusion Equilibrium

Diffusion of two solutes

Effects of Osmosis on Water Balance

• Osmosis is the diffusion of water across a

selectively permeable membrane
• The direction of osmosis is determined only by a
difference in total solute concentration
• Water diffuses across a membrane from the region
of lower solute concentration to the region of
higher solute concentration
Lower Higher Same concentration
concentration concentration of sugar
of solute (sugar) of sugar

H2O

Selectively
permeable mem-
brane: sugar mole-
cules cannot pass
through pores, but
water molecules can

Osmosis
Water Balance of Cells Without Walls

• Tonicity is the ability of a solution to cause a cell

to gain or lose water
• Isotonic solution: solute concentration is the
same as that inside the cell; no net water
movement across the plasma membrane
• Hypertonic solution: solute concentration is
greater than that inside the cell; cell loses water
• Hypotonic solution: solute concentration is less
than that inside the cell; cell gains water
• Animals and other organisms without rigid cell walls
have osmotic problems in either a hypertonic or
hypotonic environment
• To maintain their internal environment, such
organisms must have adaptations for
osmoregulation, the control of water balance
• The protist Paramecium, which is hypertonic to its
pond water environment, has a contractile vacuole
that acts as a pump
 50 µm
Filling vacuole

50 µm
Contracting vacuole
Water Balance of Cells with Walls

• Cell walls help maintain water balance

• A plant cell in a hypotonic solution swells until the wall
opposes uptake; the cell is now turgid (firm)
• If a plant cell and its surroundings are isotonic, there is
no net movement of water into the cell; the cell
becomes flaccid (limp), and the plant may wilt
• In a hypertonic environment, plant cells lose water;
eventually, the membrane pulls away from the wall, a
usually lethal effect called plasmolysis
 Hypotonic solution Isotonic solution Hypertonic solution
Animal
cell
H2O H2O H2O H2O

Lysed Normal Shriveled

Plant H2O H2O H2O H2O

cell

Turgid (normal) Flaccid Plasmolyzed

Facilitated Diffusion: Passive Transport Aided by
Proteins

• In facilitated diffusion, transport proteins speed

movement of molecules across the plasma
membrane
• Channel proteins provide corridors that allow a
specific molecule or ion to cross the membrane
• Carrier proteins undergo a subtle change in
shape that translocates the solute-binding site
across the membrane
EXTRACELLULAR
FLUID

Channel protein Solute

CYTOPLASM
Carrier protein Solute
Active transport uses energy to move solutes
against their gradients
• Facilitated diffusion is still passive because
the solute moves down its concentration
gradient
• Some transport proteins, however, can move
solutes against their concentration gradients
The Need for Energy in Active Transport

• Active transport moves substances against

their concentration gradient
• Active transport requires energy, usually in the
form of ATP
• Active transport is performed by specific proteins
embedded in the membranes
• The sodium-potassium pump is one type of
active transport system
 EXTRACELLULAR [Na+] high Na+
FLUID Na +
[K+] low

Na+ Na+ Na+

Na+ Na+

Na+

[Na+] low ATP

Na+ P P
CYTOPLASM [K+] high
ADP
Cytoplasmic Na+ bonds to Na+ binding stimulates Phosphorylation causes
the sodium-potassium pump phosphorylation by ATP. the protein to change its
conformation, expelling Na+
to the outside.

K+

K+

K+
K+
K+

P
P K+

Extracellular K+ binds Loss of the phosphate K+ is released and Na+

to the protein, triggering restores the protein’s sites are receptive again;
release of the phosphate original conformation. the cycle repeats.
group.
 Passive transport Active transport

ATP
Diffusion Facilitated diffusion
Maintenance of Membrane Potential by Ion Pumps

• Membrane potential is the voltage difference

across a membrane
• Two combined forces, collectively called the
electrochemical gradient, drive the diffusion of
ions across a membrane:
– A chemical force (the ion’s concentration
gradient)
– An electrical force (the effect of the
membrane potential on the ion’s movement)
• An electrogenic pump is a transport protein that
generates the voltage across a membrane
• The main electrogenic pump of plants, fungi, and
bacteria is a proton pump
 – EXTRACELLULAR
+
FLUID

ATP –
+ H+

H+
Proton pump
H+

– + H+

H+
– +
CYTOPLASM
H+
– +
Cotransport: Coupled Transport by a Membrane
Protein

• Cotransport occurs when active transport of a

solute indirectly drives transport of another solute
• Plants commonly use the gradient of hydrogen
ions generated by proton pumps to drive active
transport of nutrients into the cell
 –
+
ATP H+
H+
–
+

Proton pump H+
H+

–
+

H+
–
+
H+ Diffusion
of H+
Sucrose-H+
cotransporter
H+

–
+

–
+ Sucrose
Bulk transport across the plasma membrane
occurs by exocytosis and endocytosis
• Small molecules and water enter or leave the cell
through the lipid bilayer or by transport proteins
• Large molecules, such as polysaccharides and
proteins, cross the membrane via vesicles
Exocytosis

• In exocytosis, transport vesicles migrate to the

membrane, fuse with it, and release their contents
• Many secretory cells use exocytosis to export their
products
Endocytosis

• In endocytosis, the cell takes in macromolecules

by forming vesicles from the plasma membrane
• Endocytosis is a reversal of exocytosis, involving
different proteins
• Three types of endocytosis:
– Phagocytosis (“cellular eating”): Cell engulfs
particle in a vacuole
– Pinocytosis (“cellular drinking”): Cell creates
vesicle around fluid
– Receptor-mediated endocytosis: Binding of
ligands to receptors triggers vesicle formation
 RECEPTOR-MEDIATED ENDOCYTOSIS
Coat protein
Receptor Coated
vesicle

Coated
pit
Ligand

A coated pit
Coat and a coated
protein vesicle formed
during
receptor-
mediated
endocytosis
(TEMs).
Plasma
membrane
0.25 µm