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CAMPBELL
BIOLOGYReece • Urry • Cain • Wasserman • Minorsky • Jackson
© 2014 Pearson Education, Inc.
TENTH
EDITION
CAMPBELL
BIOLOGYReece • Urry • Cain • Wasserman • Minorsky • Jackson
© 2014 Pearson Education, Inc.
TENTH
EDITION
7Membrane
Structure and
Function
Lecture Presentation by
Dr Burns
NVC
Outline
I. Biological membranes
II. Plasma membrane
A. Structure and function
B. Proteins
C. Transport
Fig. 4.6
Biological membranes
Membranes all have a similar structure
Plasma membrane surrounds the cell – this
membrane controls what enters and exits
the cell - exhibits selective permeability
There are many membrane bound
organelles within the cell. The membranes
that surround these organelles have the
same basic structure
Functions of the Plasma Membrane
1. Regulates passage of materials
2. Participates in biochemical reactions
3. Receives information about environment – signal transduction
4. Communicates with other cells
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
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© 2011 Pearson Education, Inc.
• In 1935, Hugh Davson and James Danielli proposed
a sandwich model in which the phospholipid bilayer
lies between two layers of globular proteins
Later studies found problems with this model,
particularly the placement of membrane proteins,
which have hydrophilic and hydrophobic regions
In 1972, S. J. Singer and G. Nicolson proposed that
the membrane is a mosaic of proteins dispersed
within the bilayer, with only the hydrophilic regions
exposed to water
Plasma Membrane
Bilayer
Mixture of phospholipids, sterols (cholesterol), and
proteins
Phospholipids arrange to have the polar head on
the outside and the non-polar, lipid portion on the
inside
Proteins arranged on surfaces, some form
channels
Figure 7.2
Hydrophilic
head
Hydrophobic
tail
WATER
WATER
Figure 7.3
Phospholipid
bilayer
Hydrophobic regions
of protein
Hydrophilic
regions 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
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Figure 7.7
Membrane proteins
Mouse cellHuman cell
Hybrid cell
Mixed proteins
after 1 hour
RESULTS
3
Figure 7.6
Lateral movement occurs
107 times per second.
Flip-flopping across the membrane
is rare ( once per month).
Figure 7.5
Glyco-
proteinCarbohydrate
Glycolipid
Microfilaments
of cytoskeleton
EXTRACELLULAR
SIDE OF
MEMBRANE
CYTOPLASMIC SIDE
OF MEMBRANE
Integral
protein
Peripheral
proteins
Cholesterol
Fibers of extra-
cellular matrix (ECM)
Membrane Fluidity
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
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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
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Membrane Fluidity – Cholesterol’s Role
Figure 7.8
Fluid
Unsaturated hydrocarbon
tails
Viscous
Saturated hydrocarbon tails
(a) Unsaturated versus saturated hydrocarbon tails
(b) Cholesterol within the animal
cell membrane
Cholesterol
Plasma Membrane – Lipid Rafts
Cholesterol aids in maintaining the fluidity of
the membrane and making “lipid rafts”
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Figure 7.5
Glyco-
proteinCarbohydrate
Glycolipid
Microfilaments
of cytoskeleton
EXTRACELLULAR
SIDE OF
MEMBRANE
CYTOPLASMIC SIDE
OF MEMBRANE
Integral
protein
Peripheral
proteins
Cholesterol
Fibers of extra-
cellular matrix (ECM)
Five main components in membrane
1. Phospholipid bilayer – controls what passes
through membrane
2. Cholesterol – maintains proper fluidity of
membrane
3. Proteins – transport, support, communication,
recognition, enzymatic, junctions
4. Glycoproteins - form binding sites and function
in cell recognition
5. Glycolipids - form binding sites and function in
cell recognition
Phospholipids
Amphipathic - have both hydrophilic and
hydrophobic regions
Phosphate head (hydrophilic)
Fatty acid tail (hydrophobic)
Glycoproteins and Glycolipids
Short chains of carbohydrates (usually
less than 15 sugars) attached to proteins
or lipids
Glycoproteins
Glycolipids
Cell to Cell Interactions
Indentity Markers:
Glycolipids – Identify tissue types
Glycoproteins – Identify cells as “our cells”.
MHC proteins – major histocompatibility
complex proteins
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Six major functions of membrane proteins
Transport
Enzymatic activity
Signal transduction
Cell-cell recognition
Intercellular joining
Attachment to the cytoskeleton and extracellular
matrix (ECM)
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Membrane Proteins - FunctionsFigure 7.10
Enzymes
Signaling molecule
Receptor
Signal transduction
Glyco-
protein
ATP
(a) Transport (b) Enzymatic activity (c) Signal transduction
(d) Cell-cell recognition (e) Intercellular joining (f) Attachment to
the cytoskeleton
and extracellular
matrix (ECM)
Figure 7.10a
Enzymes
Signaling molecule
Receptor
Signal transductionATP
(a) Transport (b) Enzymatic activity (c) Signal transduction
Figure 7.10b
Glyco-
protein
(d) Cell-cell recognition (e) Intercellular joining (f) Attachment to
the cytoskeleton
and extracellular
matrix (ECM)
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
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Membrane Proteins - Production
Proteins that will be associated with the inner
surface of the membrane are manufactured
by free floating ribosomes
Proteins that will be associated with the outer
surface of the membrane are manufactured
by ribosomes on the rough endoplasmic
reticulum.
6
The proteins destined to be on the outer surface
of the plasma membrane have a sugar chain is
added to these proteins
transmembrane integral
outer surface peripherial proteins
These proteins are glycoproteins.
The sugar chain is added in the lumen of the
RER
Figure 7.12
Transmembraneglycoproteins
ER
ER lumen
Glycolipid
Plasma membrane:
Cytoplasmic face
Extracellular face
Secretoryprotein
Golgiapparatus
Vesicle
Transmembraneglycoprotein Secreted
protein
Membraneglycolipid
Membrane Proteins
Peripheral membrane proteins
Not embedded in the cell membrane, either
found inner or the outer surface of the
membrane
Integral membrane proteins
Embedded in the cell membrane.
Some completely cross the membrane =
transmembrane proteins
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Membrane Proteins - Peripheral
Peripheral membrane proteins
anchored to a phospholipid in one layer of the
membrane
possess nonpolar regions that are inserted in
the lipid bilayer
are free to move throughout one layer of the
bilayer
35 36
Membrane Proteins - Intergral
Integral membrane proteins
May span the lipid bilayer (transmembrane
proteins)
nonpolar regions of the protein are embedded
in the interior of the bilayer
polar regions of the protein protrude from both
sides of the bilayer
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Integral Proteins
Integral proteins penetrate the hydrophobic
core
Integral proteins that span the membrane are
called transmembrane proteins
The hydrophobic regions of an integral protein
consist of one or more stretches of nonpolar
amino acids, often coiled into alpha helices
© 2011 Pearson Education, Inc.
Figure 7.9
N-terminus
helix
C-terminus
EXTRACELLULAR
SIDE
CYTOPLASMIC
SIDE
39 40
Membrane Proteins - integral
Extensive polar regions within a
transmembrane protein can create a pore
through the membrane.
b–pleated sheets in the protein secondary
structure form a cylinder called a b-barrel
b -barrel interior is polar and allows water and
small polar molecules to pass through the
membrane
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Types of membrane proteins
1. Spectrins – give shape to the cell, form a
scaffold.
2. Clatherins – line coated pits to facilitate
receptor mediated endocytosis
3. Adhesion proteins – adhere cells to each other
or to surface
Integrins and cadherins – proteins that anchor
the cell to a substrate or cell to cell, they also
attach to microfilaments inside the cell, also
can play a role in signal transduction
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Types of membrane proteins
4. Cell Surface Receptors – transmit messages
from outside the cell to inside the cell = signal
transduction
5. Cell Surface identity markers – proteins and
glycoproteins characteristic for that cell type
6. Junction proteins – form channels between
cells (Gap junctions)
7. Transport proteins – Carrier and transport
proteins. Transport of items across the cell
membrane
Aquaporins – Transport water, important in
kidneys and in certain bacteria
Ion channels
8. Enzymes – catalyze reactions
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Proteins that anchor the cell to a substrate, they also attach
to microfilaments inside the cell
1. Spectrins
2. Receptors
3. Integrins
4. Enzymes
Junct
ion
Rec
epto
rs
Inte
grins
33% 33%33%
Passage Through the Cell Membrane
The cell membrane is selectively permeable
or semi-permeable.
This means some molecules can pass but
not all molecules.
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
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What can freely pass through the membrane
Hydrophobic compounds (non-polar)
Gases – oxygen, carbon dioxide
Very small uncharged polar compounds
(Water, glycerol) – but they will pass slowly
What can not freely pass through the membrane
Ions
Most hydrophillic compounds
Charged compounds
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
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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
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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 be directional
At dynamic equilibrium, as many molecules cross the
membrane in one direction as in the other
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© 2011 Pearson Education, Inc.
Animation: Membrane Selectivity
Right-click slide / select “Play”
© 2011 Pearson Education, Inc.
Animation: Diffusion
Right-click slide / select “Play”
Figure 7.13a
Molecules of dyeMembrane (cross section)
WATER
(a) Diffusion of one solute
Net diffusion Net diffusion Equilibrium
Figure 7.13b
(b) Diffusion of two solutes
Net diffusion Net diffusion
Net diffusion Net diffusion
Equilibrium
Equilibrium
Movement through the membrane
If you add molecules to water they will disperse
until it is equally distributed in the water =
diffusion
Equilibrium is reached when the molecules are
equally distributed
Substances diffuse down their concentration
gradient, the region along which the density of a
chemical substance increases or decreases
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 no energy
is expended by the cell to make it happen
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Movement through the membrane
Concentration gradient
Molecules will go from higher concentration
to lower concentration.
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Movement through the membraneOsmosis
The cell membrane is semipermeable
The cytoplasm is mainly water (solvent) and
molecules and ions are dissolved in it
(solutes)
Water will travel across the membrane to try
to restore the balance of solutes = Osmosis
Effects of Osmosis on Water Balance
Osmosis is the diffusion of water across a
selectively permeable membrane
Water diffuses across a membrane from the
region of lower solute concentration to the
region of higher solute concentration until the
solute concentration is equal on both sides
© 2011 Pearson Education, Inc.
Animation: OsmosisFigure 7.14 Lower
concentrationof solute (sugar)
Higher concentrationof solute
Sugarmolecule
H2O
Same concentrationof solute
Selectivelypermeablemembrane
Osmosis
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Water Balance of Cells Without Walls
Tonicity is the ability of a surrounding 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
© 2011 Pearson Education, Inc.
Figure 7.15
Hypotonicsolution
Osmosis
Isotonicsolution
Hypertonicsolution
(a) Animal cell
(b) Plant cell
H2O H2O H2O H2O
H2O H2O H2O H2OCell wall
Lysed Normal Shriveled
Turgid (normal) Flaccid Plasmolyzed
Hypertonic or hypotonic environments create
osmotic problems for organisms
Osmoregulation, the control of solute
concentrations and water balance, is a necessary
adaptation for life in such environments
The protist Paramecium, which is hypertonic to its
pond water environment, has a contractile vacuole
that acts as a pump
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Figure 7.16
Contractile vacuole50 m
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
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In a hypertonic environment, plant cells lose
water; eventually, the membrane pulls away from
the wall, a usually lethal effect called plasmolysis
© 2011 Pearson Education, Inc.
13
73
If you put a cell in very salty water, would water enter or
leave the cell?
1. Enter
2. Leave
Ente
r
Lea
ve
50%50%
Can Calcium (Ca2+) pass freely through the membrane?
1. Yes
2. No
Yes N
o
50%50%
Can glucose pass freely through the membrane?
1. Yes
2. No
Yes N
o
50%50%
BioFlix: Membrane Transport Transporting molecules across the membrane
Passive transport – molecules travel across
using a concentration gradient
1. Simple diffusion
2. Facilitated diffusion
Active Transport – requires energy, usually
in the form of ATP, to move molecules
against a concentration gradient
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Passive Transport: 1. Simple diffusion
Simple diffusion: molecules that can freely passthrough the membrane are controlled by concentration gradient
They will travel from a high concentration to a low concentration
Gases like oxygen and CO2
Very small molecules that are not charged (H2O, glycerol)
hydrophobic (non-polar) molecules (steroids)
Passive Transport: Facilitated diffusion
Passive Transport: 2. Facilitated diffusion
Facilitated diffusion: Aided by a transport protein, still controlled by concentration gradient, does not require energy
Molecules use this type of transport if they can’t pass freely through the membrane, but travels from high concentration to low concentration
Hydrophilic molecules like glucose and polar amino acids
Ions or charged compounds
Passive Transport: Facilitated diffusion
Figure 7.17
EXTRACELLULARFLUID
CYTOPLASM
Channel proteinSolute
SoluteCarrier protein
(a) A channelprotein
(b) A carrier protein
Passive Transport
15
Active Transport
Sometimes our cells want to move a molecule
across the membrane but against the
concentration gradient. The cell wants more of
the solute on one side of the membrane.
The cell will use energy to maintain the higher
concentration. The energy is usually in the
form of ATP
Example: Used to transport ions
Types of Active Transport Proteins
1. Uniporters – transport a single type of
molecule
2. Symporter – transport two types of
molecules in the same direction
3. Antiporter – transport two molecules in the
opposite direction
Active Transport - Example
Sodium-potassium pump
Energy is used to transport sodium ions to the
outside of a cell in exchange for potassium ion
which enter the cell.
Transport proteins are used
Two potassium ions enter in exchange for three
sodium ions that leave the cell
Used in nerve cells = neurons
How Ion Pumps Maintain Membrane Potential
Membrane potential is the voltage difference across a membrane
Voltage is created by differences in the distribution of positive and negative ions across a membrane
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© 2011 Pearson Education, Inc.
Animation: Active Transport
Right-click slide / select “Play”
Figure 7.18-1
EXTRACELLULAR
FLUID[Na] high
[K] low
[Na] low
[K] highCYTOPLASM
Na
Na
Na
1
16
Figure 7.18-2
EXTRACELLULAR
FLUID[Na] high
[K] low
[Na] low
[K] highCYTOPLASM
Na
Na
Na
1 2
Na
Na
Na
PATP
ADP
Figure 7.18-3
EXTRACELLULAR
FLUID[Na] high
[K] low
[Na] low
[K] highCYTOPLASM
Na
Na
Na
1 2 3
Na
Na
Na
Na
Na
Na
PP
ATP
ADP
Figure 7.18-4
EXTRACELLULAR
FLUID[Na] high
[K] low
[Na] low
[K] highCYTOPLASM
Na
Na
Na
1 2 3
4
Na
Na
Na
Na
Na
Na
K
K
PP
PP i
ATP
ADP
Figure 7.18-5
EXTRACELLULAR
FLUID[Na] high
[K] low
[Na] low
[K] highCYTOPLASM
Na
Na
Na
1 2 3
45
Na
Na
Na
Na
Na
Na
K
K
K
K
PP
PP i
ATP
ADP
Figure 7.18-6
EXTRACELLULAR
FLUID[Na] high
[K] low
[Na] low
[K] highCYTOPLASM
Na
Na
Na
1 2 3
456
Na
Na
Na
Na
Na
Na
K
K
K
K
K
K
PP
PP i
ATP
ADP
Figure 7.19Passive transport Active transport
Diffusion Facilitated diffusion ATP
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If a transport protein is used to move glucose using
a concentration gradient this is called:
1. Simple diffusion
2. Facilitated diffusion
3. Active transport
Sim
ple d
iffusi
on
Fac
ilita
ted
diffus
ion
Act
ive
tran
spor
t
33% 33%33%
Bulk Transport
When the cell needs to transport larger things they can use vesicles to transport things in and out of the cell.
Exocytosis: moving things out of the cell
Endocytosis: moving things into the cell
Used to transport macromolecules including: whole cells (bacteria), cholesterol, and proteins
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Animation: Exocytosis
Right-click slide / select “Play”
Endocytosis
In endocytosis, the cell takes in macromolecules
by forming vesicles from the plasma membrane
Endocytosis is a reversal of exocytosis, involving
different proteins
There are three types of endocytosis
Phagocytosis (“cellular eating”)
Pinocytosis (“cellular drinking”)
Receptor-mediated endocytosis
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Endocytosis
Phagocytosis – when cells transport large particles and cells (bacteria) into the cell using vesicles
Pinocytosis – when cells transport fluid into the cell using vesicles
Receptor-mediated endocytosis – molecules attach to a receptor, and are then transported into the cell. The molecules that specifically bind to a receptor are called ligands
© 2011 Pearson Education, Inc.
Animation: Exocytosis and Endocytosis Introduction
Right-click slide / select “Play”
18
In phagocytosis a cell engulfs a particle in a
vacuole
The vacuole fuses with a lysosome to digest
the particle
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Animation: Phagocytosis
Right-click slide / select “Play”
In pinocytosis, molecules are taken up when
extracellular fluid is “gulped” into tiny vesicles
© 2011 Pearson Education, Inc.
Animation: Pinocytosis
Right-click slide / select “Play”
receptor-mediated endocytosis
In receptor-mediated endocytosis, binding of
ligands to receptors triggers vesicle formation
A ligand is any molecule that binds specifically to
a receptor site of another molecule
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Animation: Receptor-Mediated Endocytosis
Right-click slide / select “Play”
19
Figure 7.22
Solutes
Pseudopodium
“Food” or
other particle
Food
vacuole
CYTOPLASM
Plasma
membrane
Vesicle
Receptor
Ligand
Coat proteins
Coated
pit
Coated
vesicle
EXTRACELLULAR
FLUID
Phagocytosis Pinocytosis Receptor-Mediated Endocytosis
Figure 7.22a
Pseudopodium
Solutes
“Food”
or other
particle
Food
vacuole
CYTOPLASM
EXTRACELLULAR
FLUID
Pseudopodium
of amoeba
Bacterium
Food vacuole
An amoeba engulfing a bacterium
via phagocytosis (TEM).
Phagocytosis
1
m
Figure 7.22b
Pinocytosis vesicles forming
in a cell lining a small blood
vessel (TEM).
Plasma
membrane
Vesicle
0.5
m
Pinocytosis
Figure 7.22c
Top: A coated pit. Bottom: Acoated vesicle forming duringreceptor-mediated endocytosis(TEMs).
Receptor
0.2
5
m
Receptor-Mediated Endocytosis
Ligand
Coat proteins
Coated
pit
Coated
vesicle
Coat
proteins
Plasma
membrane
113
Receptor-mediated endocytosis
HIV (Human Immunodeficiency Virus) entering a cell. HIV infects a number of cells in the
body, though the main target is the lymphocyte cell, which is a type of T-cell. Once the HIV
virus has infected the host cell it then replicates itself with thousands of copies. Image 1 of 3
in series. TEM X120,000.
Credit: © Dr. Hans Gelderblom/Visuals Unlimited
229674
20
HIV (Human Immunodeficiency Virus) being surrounded by a cell. HIV infects a number of
cells in the body, though the main target is the lymphocyte cell, which is a type of T-cell. Once
the HIV virus has infected the host cell it then replicates itself with thousands of copies.
Image 2 of 3 in series. TEM X120,000.
Credit: © Dr. Hans Gelderblom/Visuals Unlimited
229675HIV (Human Immunodeficiency Virus) in a cell. HIV infects a number of cells in the body,
though the main target is the lymphocyte cell, which is a type of T-cell. Once the HIV virus
has infected the host cell it then replicates itself with thousands of copies. Image 3 of 3 in
series. TEM X120,000.
Credit: © Dr. Hans Gelderblom/Visuals Unlimited
229676
when cells transport fluid into the cell using
vesicles this is called:
1. Phagocytosis
2. Pinocytosis
3. Exocytosis
Phag
ocyt
osis
Pin
ocyto
sis
Exo
cyto
sis
33% 33%33%
Important Concepts
Know the vocabulary from this lecture
What are the functions of the plasma membrane
Identify the main components in membrane and
their functions. Be able to draw a membrane.
Be able to identify what can pass freely through a
membrane and what can’t pass freely
What happens to cells in isotonic, hypertonic, and
hypertonic situations
Important Concepts
Be able to describe how things are transported
across membranes – know all the mechanisms
What are the types of membrane proteins, what
are the secondary structures of intergral
membrane proteins