MEMBRANES
2
Membranes function to organize biological
processes by
compartmentalizing them. Indeed, the cell, the
basic unit of
life, is essentially defined by its enveloping
plasma membrane.
3
Moreover, in eukaryotes, many subcellular
organelles,
such as nuclei, mitochondria, chloroplasts,
the endoplasmic
reticulum, and the Golgi apparatus are
likewise
membrane bounded.
4
FUNCTIONS OF MEMBRANE
4
 Protective barrier
 Regulate transport in & out of cell
(selectively permeable)
 Allow cell recognition
 Provide anchoring sites for filaments
of cytoskeleton
5
 Provide a binding site for
enzymes
 Interlocking surfaces bind cells
together (junctions)
Contains the cytoplasm (fluid in
cell)
6
Membrane Structure and Function
Membrane Structure
 Lipids and proteins are the main
components of the membranes,
although carbohydrates are also
important.
7
 The most abundant lipids in most
membranes are phospholipids
 Phospholipids and most of
proteins of membrane are
amphipathic molecules.
8
Amphipathic molecules : A molecule
that has both hydrophilic region and a
hydrophilic regions.
.
HISTORY OF MEMBRANE STUDIES
11
 The molecules in the bilayer are
arranged such that the
hydrophobic fatty acid tails are
sheltered from water
while the hydrophilic
phosphate groups
interact with water.
14
 Further investigation revealed two
problems.
First, not all membranes were alike, but
differed in thickness, appearance when
stained, and percentage of proteins to
lipids.
15
Second, measurements showed
that membrane proteins are actually
not very soluble in water.
• Membrane proteins are amphipathic,
with hydrophobic and hydrophilic
regions.
• If at the surface, the hydrophobic
regions would be in contact with water.
17
THE FLUID MOSAIC MODEL
1818
Unit membrane model
1919
Cell membrane and its surface structure
LIPID RAFTS MODEL
20
PROTEIN, LIPID AND CARBOHYDRATE
COMPOSITIONS OF SOME MEMBRANES
21
Depending on the precise conditions and the
nature
of the lipids, three types of lipid aggregates can
form
when amphipathic lipids are mixed with water
22
• a) Micelle
• (b) Bilayer
• (c) Liposome
23
MICELLES
Micelles are spherical structures that contain
anywhere from a few dozen to a few thousand
amphipathic
molecules. Their hydrophobic regions aggregated in the
interior,
where water is excluded, and their hydrophilic head
groups at the surface, in contact with water.
24
BILAYER
in which two lipid monolayers (leaflets) form
a two-dimensional sheet. Bilayer formation occurs
most
readily when the cross-sectional areas of the head
group
and acyl side chain(s) are similar
25
The hydrophobic portions
in each monolayer, excluded from water, interact
with
each other. The hydrophilic head groups interact
with
water at each surface of the bilayer.
26
LIPOSOME
the bilayer sheet is relatively
unstable and spontaneously forms a third
type of
aggregate: it folds back on itself to form a
hollow sphere,
a vesicle or liposome.
27
By forming vesicles,
bilayers lose their hydrophobic edge regions, achieving
maximal stability in their aqueous environment. These
bilayer vesicles enclose water, creating a separate
aqueous
compartment.
28
BIOLOGICAL MEMBRANES
Biological membranes are composed of proteins
associated with a lipid bilayer matrix. Their lipid
fractions consist of complex mixtures that vary
according to the membrane
source and, to some extent, with the diet and
environment of the organism that produced the
membrane
29
.
Membrane proteins carry out the dynamic processes
associated with membranes, and therefore specific
proteins
occur only in particular membranes.
30
MEMBRANE STRUCTURE AND COMPOSITION
MEMBRANE IS A COLLAGE OF PROTEINS & OTHER MOLECULES
EMBEDDED IN THE FLUID MATRIX OF THE LIPID BILAYER
Extracellular fluid
Cholesterol
Cytoplasm
Glycolipid
Transmembrane
proteins
Filaments of
cytoskeleton
Peripheral
protein
Glycoprotein
Phospholipids
32
PHOSPHOLIPIDS
• Fatty acid tails
• hydrophobic
• Phosphate group head
• hydrophilic
• Arranged as a bilayer
Fatty acid
Phosphate
33
PHOSPHOLIPID BILAYER
polar
hydrophilic
heads
nonpolar
hydrophobic
tails
polar
hydrophilic
heads
34
MORE THAN LIPIDS…
• In 1972, S.J. Singer & G. Nicolson proposed that membrane
proteins are inserted into the phospholipid bilayer
It’s like a fluid…
It’s like a mosaic…
It’s the
Fluid Mosaic Model!
35
MEMBRANE FAT COMPOSITION VARIES
• Fat composition affects flexibility
• membrane must be fluid & flexible
• about as fluid as thick salad oil
36
MEMBRANE PROTEINS
• Proteins determine membrane’s specific functions
• cell membrane & organelle membranes each
have unique collections of proteins
37
PERIPHERAL PROTEINS
•loosely bound to surface of
membrane
•cell surface identity marker
(antigens)
38
• INTEGRAL PROTEINS
•penetrate lipid bilayer, usually across
whole membrane
•transmembrane protein
•transport proteins
• channels, permeases (pumps)
39
Integral protein
(hydrophobic domain
coated with detergent)
Peripheral protein
40
PROTEINS DOMAINS ANCHOR MOLECULE
• Within membrane
• nonpolar amino acids
•hydrophobic
•anchors protein
into membrane
Polar areas
of protein
Nonpolar areas of protein
41
• On outer surfaces of membrane
• polar amino acids
•hydrophilic
•extend into extracellular fluid & into
cytosol
42
NH2
H+
COOH
Cytoplasm
Retinal
chromophore
Nonpolar
(hydrophobic)
a-helices in the
cell membrane H+
Porin monomer
b-pleated sheets
Bacterial
outer
membrane
proton pump channel
in photosynthetic bacteria
water channel
in bacteria
function through
conformational change =
shape change
EXAMPLES
43
MANY FUNCTIONS OF MEMBRANE PROTEINS
Outside
Plasma
membrane
Inside
Transporter Cell surface
receptor
Enzyme
activity
Cell surface
identity marker
Attachment to the
cytoskeleton
Cell adhesion
44
MEMBRANE CARBOHYDRATES
• Play a key role in cell-cell recognition
• ability of a cell to distinguish one cell from
another
• antigens
• important in organ &
tissue development
• basis for rejection of
foreign cells by
immune system
45
Any Questions??
2007-2008
MOVEMENT ACROSS THE
CELL MEMBRANE
47
DIFFUSION
• 2nd Law of Thermodynamics
governs biological systems
• universe tends towards disorder (entropy)
 Diffusion
 movement from high  low concentration
48
DIFFUSION
• Move from HIGH to LOW concentration
• “passive transport”
• no energy needed
diffusion osmosis
movement of water
49
DIFFUSION ACROSS CELL MEMBRANE
• Cell membrane is the boundary between inside &
outside…
• separates cell from its environment
IN
food
carbohydrates
sugars, proteins
amino acids
lipids
salts, O2, H2O
OUT
waste
ammonia
salts
CO2
H2O
products
cell needs materials in & products or waste out
IN
OUT
Can it be an impenetrable boundary? NO!
50
DIFFUSION THROUGH PHOSPHOLIPID BILAYER
• What molecules can get through directly?
• fats & other lipids
inside cell
outside cell
lipid
salt
aa H2Osugar
NH3
 What molecules can
NOT get through
directly?
 polar molecules
 H2O
 ions
 salts, ammonia
 large molecules
 starches, proteins
51
CHANNELS THROUGH CELL MEMBRANE
• Membrane becomes semi-permeable with protein
channels
• specific channels allow specific material across cell
membrane
inside cell
outside cell
sugaraaH2O
saltNH3
52
FACILITATED DIFFUSION
• Diffusion through protein channels
• channels move specific molecules across
cell membrane
• no energy needed
“The Bouncer”
open channel = fast transport
facilitated = with help
high
low
53
ACTIVE TRANSPORT
• Cells may need to move molecules against concentration
gradient
• shape change transports solute from
one side of membrane to other
• protein “pump”
• “costs” energy = ATP
“The Doorman”
conformational change
ATP
low
high
54
symportantiport
ACTIVE TRANSPORT
ATP ATP
55
TRANSPORT SUMMARY
simple
diffusion
facilitated
diffusion
active
transport
ATP
56
HOW ABOUT LARGE MOLECULES?
• Moving large molecules into & out of cell
• through vesicles & vacuoles
• endocytosis
• phagocytosis = “cellular eating”
• pinocytosis = “cellular drinking”
• exocytosis
exocytosis
57
ENDOCYTOSIS
phagocytosis
pinocytosis
receptor-mediated
endocytosis
fuse with
lysosome for
digestion
non-specific
process
triggered by
molecular
signal
2007-2008
THE SPECIAL CASE OF WATER
MOVEMENT OF WATER ACROSS
THE CELL MEMBRANE
59
• Diffusion of water from
high concentration of water to
low concentration of water
• across a
semi-permeable
membrane
60
CONCENTRATION OF WATER
• Direction of osmosis is determined by comparing total
solute concentrations
• Hypertonic - more solute, less water
• Hypotonic - less solute, more water
• Isotonic - equal solute, equal water
hypotonic hypertonic
water
net movement of water
61freshwater balanced saltwater
MANAGING WATER BALANCE
• Cell survival depends on balancing water uptake & loss
62
AQUAPORINS
• Water moves rapidly into & out of cells
• evidence that there were water channels
1991 | 2003
Peter Agre
John Hopkins
Roderick MacKinnon
Rockefeller
63
Any Questions??
64
65
SUMMARY
68
THE COMPOSITION AND ARCHITECTURE
OF MEMBRANES
69
Biological membranes define cellular boundaries,
divide cells into discrete compartments, organize
complex reaction sequences, and act in signal
reception and energy transformations.
70
Membranes are composed of lipids and proteins
in varying combinations particular to each
species, cell type, and organelle. The fluid
mosaic model describes features common to all
biological membranes. The lipid bilayer is the
basic structural unit.
71
Fatty acyl chains of phospholipids and the
steroid nucleus of sterols are oriented toward the
interior of the bilayer;
their hydrophobic interactions stabilize the
bilayer but give it flexibility.
72
Peripheral proteins are loosely associated with
the membrane through electrostatic interactions
and hydrogen bonds or by covalently attached
lipid anchors..
73
Integral proteins associate firmly
with membranes by hydrophobic interactions
between the lipid bilayer and their nonpolar
amino acid side chains, which are oriented
toward the outside of the protein molecule
74
75
76
Membrane Dynamics
77
Lipids in a biological membrane can exist in
liquid-ordered or liquid-disordered states; in
the latter state, thermal motion of acyl chains
makes the interior of the bilayer fluid. Fluidity
is affected by temperature, fatty acid
composition, and sterol content.
78
Flip-flop diffusion of lipids between the
inner
and outer leaflets of a membrane is very
slow
except when specifically catalyzed by
flippases.
79
Lipids and proteins can diffuse laterally within
the plane of the membrane, but this mobility is
limited by interactions of membrane proteins
with internal cytoskeletal structures and
interactions of lipids with lipid rafts.
80
One class
of lipid rafts consists of sphingolipids and
cholesterol with a subset of membrane proteins
that are GPI-linked or attached to several
long-chain fatty acyl moieties.
81
Caveolin is an integral membrane protein that
associates with the inner leaflet of the plasma
membrane, forcing it to curve inward to form
caveolae, probably involved in membrane
transport and signaling.
82
Integrins are transmembrane proteins of the
plasma membrane that act both to attach cells to
each other and to carry messages between
the extracellular matrix and the cytoplasm.
■ Specific proteins mediate the fusion of two
membranes, which accompanies processes such
as viral invasion and endocytosis and
exocytosis.
83
SOLUTE TRANSPORT ACROSS
MEMBRANES
Movement of polar compounds and ions across
biological membranes requires protein
transporters. Some transporters simply
facilitate passive diffusion across the membrane
from the side with higher concentration to the
side with lower.
84
Others bring about active
movement of solutes against an electrochemical
gradient; such transport must be coupled to a
source of metabolic energy.
85
Carriers, like enzymes, show saturation and
stereospecificity for their substrates.
Transport via these systems may be passive
or active.
■
86
Primary active transport is driven by ATP
or
electron-transfer reactions; secondary
active
transport, by coupled flow of two solutes,
one of which (often H or Na) flows down
its
electrochemical gradient as the other is
pulled up its gradient.
87
The GLUT transporters, such as GLUT1 of
erythrocytes, carry glucose into cells by
facilitated diffusion. These transporters are
uniporters, carrying only one substrate.
88
Symporters permit simultaneous passage of two
substances in the same direction; examples are
the lactose transporter of E. coli, driven by the
energy of a proton gradient (lactose-H
symport), and the glucose transporter of
intestinal epithelial cells, driven by a Na
gradient (glucose-Na symport)
89
Antiporters mediate simultaneous passage of two
substances in opposite directions; examples are
the chloride-bicarbonate exchanger of
erythrocytes and the ubiquitous NaK
ATPase.
90

Membranes

  • 1.
  • 2.
    2 Membranes function toorganize biological processes by compartmentalizing them. Indeed, the cell, the basic unit of life, is essentially defined by its enveloping plasma membrane.
  • 3.
    3 Moreover, in eukaryotes,many subcellular organelles, such as nuclei, mitochondria, chloroplasts, the endoplasmic reticulum, and the Golgi apparatus are likewise membrane bounded.
  • 4.
    4 FUNCTIONS OF MEMBRANE 4 Protective barrier  Regulate transport in & out of cell (selectively permeable)  Allow cell recognition  Provide anchoring sites for filaments of cytoskeleton
  • 5.
    5  Provide abinding site for enzymes  Interlocking surfaces bind cells together (junctions) Contains the cytoplasm (fluid in cell)
  • 6.
    6 Membrane Structure andFunction Membrane Structure  Lipids and proteins are the main components of the membranes, although carbohydrates are also important.
  • 7.
    7  The mostabundant lipids in most membranes are phospholipids  Phospholipids and most of proteins of membrane are amphipathic molecules.
  • 8.
    8 Amphipathic molecules :A molecule that has both hydrophilic region and a hydrophilic regions. .
  • 9.
  • 10.
    11  The moleculesin the bilayer are arranged such that the hydrophobic fatty acid tails are sheltered from water while the hydrophilic phosphate groups interact with water.
  • 11.
    14  Further investigationrevealed two problems. First, not all membranes were alike, but differed in thickness, appearance when stained, and percentage of proteins to lipids.
  • 12.
    15 Second, measurements showed thatmembrane proteins are actually not very soluble in water. • Membrane proteins are amphipathic, with hydrophobic and hydrophilic regions. • If at the surface, the hydrophobic regions would be in contact with water.
  • 13.
  • 14.
  • 15.
    1919 Cell membrane andits surface structure LIPID RAFTS MODEL
  • 16.
    20 PROTEIN, LIPID ANDCARBOHYDRATE COMPOSITIONS OF SOME MEMBRANES
  • 17.
    21 Depending on theprecise conditions and the nature of the lipids, three types of lipid aggregates can form when amphipathic lipids are mixed with water
  • 18.
    22 • a) Micelle •(b) Bilayer • (c) Liposome
  • 19.
    23 MICELLES Micelles are sphericalstructures that contain anywhere from a few dozen to a few thousand amphipathic molecules. Their hydrophobic regions aggregated in the interior, where water is excluded, and their hydrophilic head groups at the surface, in contact with water.
  • 20.
    24 BILAYER in which twolipid monolayers (leaflets) form a two-dimensional sheet. Bilayer formation occurs most readily when the cross-sectional areas of the head group and acyl side chain(s) are similar
  • 21.
    25 The hydrophobic portions ineach monolayer, excluded from water, interact with each other. The hydrophilic head groups interact with water at each surface of the bilayer.
  • 22.
    26 LIPOSOME the bilayer sheetis relatively unstable and spontaneously forms a third type of aggregate: it folds back on itself to form a hollow sphere, a vesicle or liposome.
  • 23.
    27 By forming vesicles, bilayerslose their hydrophobic edge regions, achieving maximal stability in their aqueous environment. These bilayer vesicles enclose water, creating a separate aqueous compartment.
  • 24.
    28 BIOLOGICAL MEMBRANES Biological membranesare composed of proteins associated with a lipid bilayer matrix. Their lipid fractions consist of complex mixtures that vary according to the membrane source and, to some extent, with the diet and environment of the organism that produced the membrane
  • 25.
    29 . Membrane proteins carryout the dynamic processes associated with membranes, and therefore specific proteins occur only in particular membranes.
  • 26.
  • 27.
    MEMBRANE IS ACOLLAGE OF PROTEINS & OTHER MOLECULES EMBEDDED IN THE FLUID MATRIX OF THE LIPID BILAYER Extracellular fluid Cholesterol Cytoplasm Glycolipid Transmembrane proteins Filaments of cytoskeleton Peripheral protein Glycoprotein Phospholipids
  • 28.
    32 PHOSPHOLIPIDS • Fatty acidtails • hydrophobic • Phosphate group head • hydrophilic • Arranged as a bilayer Fatty acid Phosphate
  • 29.
  • 30.
    34 MORE THAN LIPIDS… •In 1972, S.J. Singer & G. Nicolson proposed that membrane proteins are inserted into the phospholipid bilayer It’s like a fluid… It’s like a mosaic… It’s the Fluid Mosaic Model!
  • 31.
    35 MEMBRANE FAT COMPOSITIONVARIES • Fat composition affects flexibility • membrane must be fluid & flexible • about as fluid as thick salad oil
  • 32.
    36 MEMBRANE PROTEINS • Proteinsdetermine membrane’s specific functions • cell membrane & organelle membranes each have unique collections of proteins
  • 33.
    37 PERIPHERAL PROTEINS •loosely boundto surface of membrane •cell surface identity marker (antigens)
  • 34.
    38 • INTEGRAL PROTEINS •penetratelipid bilayer, usually across whole membrane •transmembrane protein •transport proteins • channels, permeases (pumps)
  • 35.
    39 Integral protein (hydrophobic domain coatedwith detergent) Peripheral protein
  • 36.
    40 PROTEINS DOMAINS ANCHORMOLECULE • Within membrane • nonpolar amino acids •hydrophobic •anchors protein into membrane Polar areas of protein Nonpolar areas of protein
  • 37.
    41 • On outersurfaces of membrane • polar amino acids •hydrophilic •extend into extracellular fluid & into cytosol
  • 38.
    42 NH2 H+ COOH Cytoplasm Retinal chromophore Nonpolar (hydrophobic) a-helices in the cellmembrane H+ Porin monomer b-pleated sheets Bacterial outer membrane proton pump channel in photosynthetic bacteria water channel in bacteria function through conformational change = shape change EXAMPLES
  • 39.
    43 MANY FUNCTIONS OFMEMBRANE PROTEINS Outside Plasma membrane Inside Transporter Cell surface receptor Enzyme activity Cell surface identity marker Attachment to the cytoskeleton Cell adhesion
  • 40.
    44 MEMBRANE CARBOHYDRATES • Playa key role in cell-cell recognition • ability of a cell to distinguish one cell from another • antigens • important in organ & tissue development • basis for rejection of foreign cells by immune system
  • 41.
  • 42.
  • 43.
    47 DIFFUSION • 2nd Lawof Thermodynamics governs biological systems • universe tends towards disorder (entropy)  Diffusion  movement from high  low concentration
  • 44.
    48 DIFFUSION • Move fromHIGH to LOW concentration • “passive transport” • no energy needed diffusion osmosis movement of water
  • 45.
    49 DIFFUSION ACROSS CELLMEMBRANE • Cell membrane is the boundary between inside & outside… • separates cell from its environment IN food carbohydrates sugars, proteins amino acids lipids salts, O2, H2O OUT waste ammonia salts CO2 H2O products cell needs materials in & products or waste out IN OUT Can it be an impenetrable boundary? NO!
  • 46.
    50 DIFFUSION THROUGH PHOSPHOLIPIDBILAYER • What molecules can get through directly? • fats & other lipids inside cell outside cell lipid salt aa H2Osugar NH3  What molecules can NOT get through directly?  polar molecules  H2O  ions  salts, ammonia  large molecules  starches, proteins
  • 47.
    51 CHANNELS THROUGH CELLMEMBRANE • Membrane becomes semi-permeable with protein channels • specific channels allow specific material across cell membrane inside cell outside cell sugaraaH2O saltNH3
  • 48.
    52 FACILITATED DIFFUSION • Diffusionthrough protein channels • channels move specific molecules across cell membrane • no energy needed “The Bouncer” open channel = fast transport facilitated = with help high low
  • 49.
    53 ACTIVE TRANSPORT • Cellsmay need to move molecules against concentration gradient • shape change transports solute from one side of membrane to other • protein “pump” • “costs” energy = ATP “The Doorman” conformational change ATP low high
  • 50.
  • 51.
  • 52.
    56 HOW ABOUT LARGEMOLECULES? • Moving large molecules into & out of cell • through vesicles & vacuoles • endocytosis • phagocytosis = “cellular eating” • pinocytosis = “cellular drinking” • exocytosis exocytosis
  • 53.
  • 54.
    2007-2008 THE SPECIAL CASEOF WATER MOVEMENT OF WATER ACROSS THE CELL MEMBRANE
  • 55.
    59 • Diffusion ofwater from high concentration of water to low concentration of water • across a semi-permeable membrane
  • 56.
    60 CONCENTRATION OF WATER •Direction of osmosis is determined by comparing total solute concentrations • Hypertonic - more solute, less water • Hypotonic - less solute, more water • Isotonic - equal solute, equal water hypotonic hypertonic water net movement of water
  • 57.
    61freshwater balanced saltwater MANAGINGWATER BALANCE • Cell survival depends on balancing water uptake & loss
  • 58.
    62 AQUAPORINS • Water movesrapidly into & out of cells • evidence that there were water channels 1991 | 2003 Peter Agre John Hopkins Roderick MacKinnon Rockefeller
  • 59.
  • 60.
  • 61.
  • 62.
  • 63.
    68 THE COMPOSITION ANDARCHITECTURE OF MEMBRANES
  • 64.
    69 Biological membranes definecellular boundaries, divide cells into discrete compartments, organize complex reaction sequences, and act in signal reception and energy transformations.
  • 65.
    70 Membranes are composedof lipids and proteins in varying combinations particular to each species, cell type, and organelle. The fluid mosaic model describes features common to all biological membranes. The lipid bilayer is the basic structural unit.
  • 66.
    71 Fatty acyl chainsof phospholipids and the steroid nucleus of sterols are oriented toward the interior of the bilayer; their hydrophobic interactions stabilize the bilayer but give it flexibility.
  • 67.
    72 Peripheral proteins areloosely associated with the membrane through electrostatic interactions and hydrogen bonds or by covalently attached lipid anchors..
  • 68.
    73 Integral proteins associatefirmly with membranes by hydrophobic interactions between the lipid bilayer and their nonpolar amino acid side chains, which are oriented toward the outside of the protein molecule
  • 69.
  • 70.
  • 71.
  • 72.
    77 Lipids in abiological membrane can exist in liquid-ordered or liquid-disordered states; in the latter state, thermal motion of acyl chains makes the interior of the bilayer fluid. Fluidity is affected by temperature, fatty acid composition, and sterol content.
  • 73.
    78 Flip-flop diffusion oflipids between the inner and outer leaflets of a membrane is very slow except when specifically catalyzed by flippases.
  • 74.
    79 Lipids and proteinscan diffuse laterally within the plane of the membrane, but this mobility is limited by interactions of membrane proteins with internal cytoskeletal structures and interactions of lipids with lipid rafts.
  • 75.
    80 One class of lipidrafts consists of sphingolipids and cholesterol with a subset of membrane proteins that are GPI-linked or attached to several long-chain fatty acyl moieties.
  • 76.
    81 Caveolin is anintegral membrane protein that associates with the inner leaflet of the plasma membrane, forcing it to curve inward to form caveolae, probably involved in membrane transport and signaling.
  • 77.
    82 Integrins are transmembraneproteins of the plasma membrane that act both to attach cells to each other and to carry messages between the extracellular matrix and the cytoplasm. ■ Specific proteins mediate the fusion of two membranes, which accompanies processes such as viral invasion and endocytosis and exocytosis.
  • 78.
    83 SOLUTE TRANSPORT ACROSS MEMBRANES Movementof polar compounds and ions across biological membranes requires protein transporters. Some transporters simply facilitate passive diffusion across the membrane from the side with higher concentration to the side with lower.
  • 79.
    84 Others bring aboutactive movement of solutes against an electrochemical gradient; such transport must be coupled to a source of metabolic energy.
  • 80.
    85 Carriers, like enzymes,show saturation and stereospecificity for their substrates. Transport via these systems may be passive or active. ■
  • 81.
    86 Primary active transportis driven by ATP or electron-transfer reactions; secondary active transport, by coupled flow of two solutes, one of which (often H or Na) flows down its electrochemical gradient as the other is pulled up its gradient.
  • 82.
    87 The GLUT transporters,such as GLUT1 of erythrocytes, carry glucose into cells by facilitated diffusion. These transporters are uniporters, carrying only one substrate.
  • 83.
    88 Symporters permit simultaneouspassage of two substances in the same direction; examples are the lactose transporter of E. coli, driven by the energy of a proton gradient (lactose-H symport), and the glucose transporter of intestinal epithelial cells, driven by a Na gradient (glucose-Na symport)
  • 84.
    89 Antiporters mediate simultaneouspassage of two substances in opposite directions; examples are the chloride-bicarbonate exchanger of erythrocytes and the ubiquitous NaK ATPase.
  • 85.