Chapter 5

Membranes and Transport

Russell, Biology: The Dynamic Science, 5th edition. © 2021 Cengage. All Rights Reserved. May not be
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Why It Matters. . .
• The plasma membrane separates a cell from its
surroundings.
• The plasma membrane maintains the internal
environment of cells.
• Transport proteins move particular ions and
molecules.

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5.1 Membrane Structure and Function
• Biological membranes consist of lipids and
proteins
• Proportions of lipid and protein molecules in
membranes vary.
• Phospholipids and sterols

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Phospholipids
• Phospholipids have a polar (electrically charged)
and a nonpolar (uncharged) end
• Phospholipids have dual solubility properties
• In an aqueous medium, phospholipid molecules
assemble into a bilayer

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The Phospholipid Bilayer
• In a bilayer, the polar ends of the phospholipid
molecules are located at the surfaces.
• The nonpolar fatty-acid chains assemble in the
membrane interior.
• At low temperatures, the phospholipid bilayer
freezes into a semisolid.
• When a phospholipid bilayer is shaken in water, it
breaks and spontaneously forms small vesicles

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Phospholipid Bilayers

A. Phospholipid molecule B. Fluid bilayer C. Bilayer vesicle (cross section)

(hydrophilic)

Aqueous
Polar end

Choline solution
Aqueous
solution
Phosphate
group
Glycerol Aqueous Aqueous Lipid
solution solution bilayer

D. “Frozen” bilayer
((hydrophobic)
Nonpolar end

Hydrophobic
tail

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Sterols
• Membrane sterols also have dual solubility
characteristics
• Sterols have nonpolar carbon rings with a
nonpolar side chain at one end and a single polar
group (—OH) at the other end
• Sterols aligns (stretches through) in the
phospholipid bilayer
• Cholesterol

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Cholesterol in Bilayers
Cholesterol
OH Hydrophilic
end

Hydrophobic
end

Hydrophobic
tail

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Membrane Proteins
• Transport proteins form channels
• Recognition proteins in the plasma membrane
identify a cell as part of the same individual
• Receptor proteins recognize and bind molecules,
e.g. hormones
• Cell adhesion proteins bind cells together by
recognizing and binding receptors or chemical
groups on other cells

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Membrane Protein Structure

A. Typical membrane protein B. Hydrophilic and hydrophobic surfaces

Outside Channel Membrane Hydrophilic

Hydrophilic Protein surface
cell surface channel
NH2
Alpha
helix

Plasma
membrane
interior

Hydrophobic Hydrophilic
Cytosol COOH protein surface Protein surface

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Membrane Glycolipids and Glycoproteins
• Glycolipids and glycoproteins are found on the
outer surface of cell membranes
• In many animal cells, the carbohydrate groups of
the cell surface glycolipids and glycoproteins form
a surface coat called the glycocalyx

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The Fluid Mosaic Model
• The fluid mosaic model proposes that the
membrane consists of a fluid phospholipid bilayer
in which proteins are embedded and float freely
• “Fluid” refers to the phospholipid molecules,
which constantly move and exchange places
within the same layer
• Unsaturated fatty acid chains
• Cholesterol

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The Fluid Mosaic Model (cont'd.)
• “Mosaic” refers to membrane proteins:
• Particular functional groups
• Integral proteins
• Peripheral proteins

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Membrane Structure and the Fluid Mosaic
Model

Integral
Outside cell
Carbohydrate
proteins
groups

Integral
proteins

Glycolipid

Plasma membrane

Gate Cholesterol Integral protein

(transport protein)

Integral protein Peripheral Glycoprotein Peripheral protein Microfilament Peripheral

(gated channel proteins (linking microtubule of cytoskeleton protein
protein) to membrane)

Cytosol

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Plasma Membrane of an Animal Cell
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Plasma membrane

carbohydrate
extracellular chain Outside cell
Matrix (ECM)

hydrophobic hydrophilic
glycoprotein tails heads
phospholipid
phospholipid glycolipid bilayer

filaments of cytoskeleton Inside cell

peripheral protein integral protein

cholesterol

18
Membrane Functions
1. Membranes define the boundaries of cells
2. Membranes are permeability barriers
3. Some membrane proteins have enzyme activities
4. Membrane-spanning channel proteins selectively regulate
transport of specific ions or water through the membrane
5. Membrane-spanning carrier proteins bind to specific
substances
6. Some membrane proteins are receptors.
7. Membranes have electrical properties and conduct
signals
8. Some membrane proteins facilitate cell adhesion and cell-
to-cell communication (e.g., gap junctions)

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 Plasma Membrane Structure
and Function
• Functions of Membrane Proteins
 Channel Proteins:
• Allow passage of molecules through membrane via a
channel in the protein

 Carrier Proteins:
• Combine with the substance to be transported
• Change shape
• Assist passage of molecules through membrane
• E.g. Potassium and Sodium transport I nerve cell.

 Cell Recognition Proteins:

• Glycoproteins
• Help the body recognize foreign substances or pathogens

26
 Plasma Membrane Structure
and Function
• Functions of Membrane Proteins (continued)
 Receptor Proteins:
• Bind with specific molecules, has a specific shape
• Allow a cell to respond to signals from other cells

 Enzymatic Proteins:
• Carry out metabolic reactions directly

 Junction Proteins:
• Involved in forming various junctions between cells.
• Attach adjacent cells

27
 Membrane Protein Diversity
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Cell Recognition
Protein:
Channel Protein: Carrier Protein: glycoproteins
Allows a particular Selectively interacts are different for each
molecule or ion to with a specific person, so organ
cross the plasma molecule or ion so transplants are difficult
membrane freely. that it can cross the to achieve. Cells with
plasma membrane. foreign
glycoproteins are
attacked by white blood
cells responsible for
immunity.
a. b. c.

Enzymatic Junction Proteins:

Receptor
Protein: Tight junctions join
Protein:
Catalyzes a cells so that a tissue
Is shaped in
specific can fulfill a function,as
such a
reaction. when a tissue pinches
way that a
specific off the neural tube
molecule can during development.
bind to Without this
it. cooperation between
cells, an animal
embryo would have no
nervous system.
d. e. f.
5.2 Functions of Membranes in Transport:
Passive Transport
• Transport is the controlled movement of specific
ions and molecules across a membrane by
membrane proteins
• Passive transport moves ions and molecules
with the concentration gradient
• Active transport moves ions or molecules against
the concentration gradient.

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Diffusion
• Passive transport is a form of diffusion – the net
movement of ions or molecules from a region of
higher concentration to a region of lower
concentration
• Diffusion involves a net movement of molecules or
ions

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Diffusion
• A solution consists of:
• A solvent (liquid), and
• A solute (dissolved solid)
• Diffusion
• Net movement of molecules down a concentration
gradient
• Equilibrium:
• When NET movement stops
• Solute concentration is uniform – no gradient

32
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Selective Permeability
• Biological membranes are selectively permeable
• Hydrophobic (nonpolar) molecules move freely
through the lipid bilayer
• Hydrophilic molecules (ions and polar molecules)
move through slowly
• Charged atoms and molecules are blocked by the
hydrophobic core

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Simple Diffusion
• Simple diffusion
• Diffusion through the lipid part of a biological
membrane
• Depends solely on molecular size and lipid
solubility
• Nonpolar inorganic gases and organic molecules
are transported by simple diffusion
• Water is small enough to slip between the
hydrocarbon tails.

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Facilitated Diffusion
• Facilitated diffusion
• Diffusion of polar and charged molecules through
transport proteins in the hydrophobic lipid bilayer
• Facilitated diffusion is specific

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Characteristics of Transport Mechanisms

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Transport Proteins in Facilitated Diffusion
• Channel proteins
• Integral membrane proteins that form hydrophilic
channels in the membrane through which water
and ions can pass
• Aquaporins
• Ion channels
• Gated channels

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Channel Proteins
A. Channel protein
Channel proteins form hydrophilic channels in the membrane through which water and ions can move.

1 Aquaporin 2 K+ voltage-gated channel

Outside cell Water
moleculeConcentration Direction of K+ channel
Activation
gradient transport Concentration Direction of
gate
(High) gradient transport
(Low)
Lipid
bilayer +
membrane +

(High)
(Low)

Aquaporin

Cytosol

An aquaporin is a water channel. With normal voltage In response to a

Water molecules move through across the voltage change
the channel by being handed off membrane, across the
to a succession of hydrogen- the activation gate of membrane, the
bonding sites on the channel in this the K+ channel is activation gate of the
protein. closed and K + cannot K + channel opens,
move across the and K + moves with its
membrane. concentration
gradient
from the cytoplasm to
outside the cell.

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Transport Proteins in Facilitated Diffusion
(cont'd.)
• Carrier proteins
• Bind a specific single solute and transport it across
the lipid bilayer (uniport transport)
• Undergo conformational changes
• Can become saturated
• Each type of cellular membrane has a
characteristic group of transport proteins

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Carrier Proteins

B. Carrier protein 1 Carrier protein is in

conformation so that binding
Carrier proteins each bind a single solute and site is exposed toward region
transport it across the lipid bilayer. During the of higher concentration.
transport step, the carrier protein undergoes
conformational changes that Solute molecule Concentration
move the solute-binding site progressively from to be transportedgradient
Carrier protein (High)
one side of the membrane to the other,thereby
Binding site
transporting the solute. Shown is the transport Membrane
of glucose. (Low)

Direction of
transport Diffusion
4 Transported
Solute is released
2 Solute
molecule binds
and carrier protein
to carrier protein.
returns to
conformation in step .

3 Solute
molecule binds
to carrier protein.

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5.3 Passive Water Transport and Osmosis
• Osmosis is the diffusion of water across a
selectively permeable membrane in response to
concentration gradients
• Movement of water by osmosis develops forces
• Animal cells expend energy to counteract the
inward or outward movement of water by osmosis

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Osmosis in a Physical System
• Water moves across a semipermeable membrane
• Association of water molecules with solute
molecules reduces the amount of water available
to cross the membrane
• The concentration of free water molecules is lower
on the solute side than on the pure water side

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Osmosis

A. Demonstration of osmosis B. Basis of osmotic water flow

Glucose
solution d
rises in tube Region of lower
free water
Distilled concentration
H2O
Glucose Glucose
Glucose solution molecule
solution
Selectively
in water
permeable
Direction
membrane
of osmotic
water flow Water molecule
Cellophane Region of higher
membrane free water
H2O
concentration

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Osmotic Pressure
• The solution stops rising in the tube when the
pressure created by the weight of the raised
solution exactly balances the tendency of water
molecules to move from the beaker into the tube in
response to the concentration gradient
• This pressure is the osmotic pressure of the
solution

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Tonicity
• Tonicity is a property of a solution with respect to
a particular membrane
• If the solution surrounding a cell contains
nonpenetrating solutes at lower concentrations
than in the cell, the solution is hypotonic to the
cell
• If the solution surrounding a cell contains
nonpenetrating solutes at higher concentrations
than in the cell, the solution is hypertonic to the
cell

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Tonicity (cont'd.)
• If the concentrations of solutes inside and outside
the cell are balanced (equal), the two solutions are
isotonic

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Tonicity and Osmotic Water Movement

2 M sucrose
solution

A. Hypotonic B. Hypertonic C. Isotonic

conditions conditions conditions

Distilled water: 10 M sucrose solution: 2 M sucrose solution:

bag swells bag shrinks bag unchanged

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Tonicity in Animal Cells

Red blood cells:

David M. Phillips/Science
Source
Water diffuses inward; Water diffuses outward; No net movement of
cells swell. cells shrink. water; cells do not
change in size or
shape.
Hypotonic Hypertonic Isotonic

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Tonicity in Plant Cells
• In a hypotonic solution, strong walls prevent plant
cells from bursting – turgor pressure
• In a hypertonic solution, stems and leaves wilt -
plasmolysis

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Tonicity in Plants

Plant cells:

Turgor
pressure

Cell Vacuole Cytoplasm

wall Weakened turgor
Normal turgor pressure Plasmolysis pressure

Hypotonic Hypertonic Isotonic

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5.4 Active Transport
• Transport of substances across a membrane
against a concentration gradient requires active
transport, a process that requires energy input
• Three main functions of active transport
• Uptake of essential nutrients from fluid surrounding
cells
• Removal of secretory or waste materials from cells
or organelles
• Maintenance of intracellular concentrations

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Membrane Potential
• Active transport of ions contributes to an electrical
potential difference across the plasma membrane
- membrane potential
• Neurons and muscle cells use membrane potential
in response to a stimulus.

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Two Kinds of Active Transport
• Primary active transport
• The protein that transports a substance also
hydrolyzes ATP to power the transport directly
• Secondary active transport
• Transport is indirectly driven by ATP hydrolysis

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Primary Active Transport
• Primary active transport pumps move positively
charged ions across membranes
• Gradients of positive ions established by primary
active transport pumps underlie essential functions
of life
• H+ pumps (proton pumps) generate membrane
potential.

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The Sodium/Potassium Pump
• The Na+/K+ pump (Na+/K+-ATPase) in the plasma
membrane of all animal cells moves 3 Na+ out of
the cell and 2 K+ into the cell in the same pumping
cycle
• Positive charges accumulate outside the
membrane, and the inside becomes relatively
negative.
• Forms an electrochemical gradient – a form of
potential energy

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 Outside Cell Na+–K+ pump
Na+ concentration High-affinity
gradient Plasma binding site for
mem- Low-affinity Na+
brane binding site
for K+
K+ concentration
Cytosol gradient
Direction of K+
transport

Low-affinity
binding site
for Na+

High-affinity
Direction of Na+ binding site
transport for K+

Stepped Art
5.5 Exocytosis and Endocytosis
• Eukaryotic cells import and export larger particles
and molecules such as proteins by exocytosis and
endocytosis
• Require energy from ATP

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Exocytosis
• In exocytosis, secretory vesicles bud from the
Golgi complex move through the cytoplasm and
contact the plasma membrane

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Exocytosis (cont’d.)

A. Exocytosis
Vesicle joins plasma membrane, releases contents.

Secretory Cytosol Outside

vesicle cell

Proteins
inside
vesicle
Proteins in
vesicle
Plasma
membrane
membrane

1 Secretory vesicle approaches 2 Vesicle fuses with 3 Proteins inside vesicle

plasma membrane. plasma membrane. are released to the cell
exterior; proteins in vesicle
membrane become part of
plasma membrane.

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Endocytosis
• In endocytosis, proteins and other substances are
trapped in pitlike depressions that bulge inward
from the plasma membrane and pinch off as an
endocytic vesicle
• Endocytosis occurs by one of two pathways:
• 1. bulk endocytosis or
• 2. receptor-mediated endocytosis

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Bulk Endocytosis
• Bulk endocytosis (pinocytosis) takes in a drop
of the aqueous extracellular fluid (ECF)
surrounding the cells
• Pinocytosis is nonspecific – it takes in any solutes
present in ECF

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Bulk Endocytosis
Cytosol Outside
cell
Water
molecule

Solute
molecule
Plasma
membrane

1 Solute molecules and 2 Membrane pockets inward, 3 Pocket pinches off

water molecules are outside enclosing solute molecules and as endocytic vesicle.
the plasma membrane. water molecules.

Receptor-mediated endocytosis: Vesicle imports specific molecules.

Cytosol Outside
cell
Clathrin

Target
molecule
Plasma Receptor
membrane

1 Substances attach to 2 Membrane 3 Pocket pinches off

membrane receptors. pockets inward. as endocytic vesicle.

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Receptor-Mediated Endocytosis
• In receptor-mediated endocytosis, target
molecules are bound to receptor proteins on the
outer cell surface
• The receptors (integral membrane proteins)
recognize and bind only specific molecules in the
ECF
• Receptors with target molecules collect in a
depression in the plasma membrane called a
coated pit

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Receptor-Mediated Endocytosis (cont'd.)
• The pits pinch free of the plasma membrane to
form endocytic vesicles
• Enzymes in the lysosome digest the contents of
the vesicle
• Molecular products cross the vesicle membrane
into the cytoplasm via transport proteins
• Membrane proteins are recycled to the plasma
membrane

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Receptor-Mediated Endocytosis

Micrographs of stages of receptor-mediated endocytosis shown above

Molecules
bound to Plasma
surface membrane
receptors pinching off
Coated
pit
Clathrin Coated pit

jsc.biologists.
coat deepens

Plasma
membrane

org
0.25 µm

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Phagocytosis
• Phagocytes in the bloodstream, and protists such
as Amoeba, take in large particles or whole cells
by phagocytosis
• Surface receptors bind to the materials to be taken
in
• Cytoplasmic lobes extend and surround the
materials
• Enzymes digest the materials.

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Phagocytosis

Endocytic Food vacuole

vesicle

Lebendkulturen.de/Shutterstoc
k.com
Prey
1 Lobes begin to surround prey. 2 Lobes close around prey. 3 Prey is enclosed in endocytic vesicle (called a food vacuole
here) that sinks into cytoplasm.

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How Molecules Cross the
Plasma Membran e
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charged molecules
– + and ions
water outside cell

H2O
noncharged
molecules

+ macromolecule

water insidecell phospholipid

molecule

protein

75
Passage of Molecules Into
and out of the Cell

76