6

nds of subunits:

olipid molecule.
Membranes

B101 Lecture
Concept Outline
the phospholipid molecule.

6.1 Biological membranes are fluid layers of lipid.

The Phospholipid Bilayer. Cells are encased by

Lipids &
membranes composed of a bilayer of phospholipid.
The Lipid Bilayer Is Fluid. Because individual
phospholipid molecules do not bind to one another, the lipid
bilayer of membranes is a fluid.

6.2 Proteins embedded within the plasma membrane

determine its character.
The Fluid Mosaic Model. A varied collection of proteins
float within the lipid bilayer.
Examining Cell Membranes. Visualizing a plasma
ids, long chains of C—H bonds (hydrocarbon

ids are attached to the glycerol backbone in a

ending in a carboxyl (—COOH) group. Two
l phospholipid is a composite molecule, made

ol, a three-carbon alcohol, with each carbon

a hydroxyl group. Glycerol forms the back-

membrane requires a powerful electron microscope.

Kinds of Membrane Proteins. The proteins in a

Carbohydrates
membrane function in support, transport, recognition, and
reactions.
Structure of Membrane Proteins. Membrane proteins are
anchored into the lipid bilayer by their nonpolar regions.
FIGURE 6.1

4
6.3 Passive transport across membranes moves down Membranes within a human cell. Sheets of endoplasmic

HO
the concentration gradient. reticulum weave through the cell interior. The large oval is a
H
H

3
H 5
mitochondrion, itself filled with extensive internal membranes.
sugar

Diffusion. Random molecular motion results in a net

1

Water
Water
Water

H
H
movement of molecules to regions of lower concentration.
H C
C
Oil C

OH
3-carbon

Facilitated Diffusion. Passive movement across a

membrane is often through specific carrier proteins. mong a cell’s most important activities are its interac-
O

Osmosis. Polar solutes interact with water and can affect tions with the environment, a give and take that never

6 CH2OH

Glucose
A
OH
OH

the movement of water across semipermeable membranes.

H
ceases. Without it, life could not persist. While living cells

HO
O
Glyceraldehyde
carbohydrates

OH
2
and eukaryotic organelles (figure 6.1) are encased within a
6-carbon sugars
6.4 Bulk transport utilizes endocytosis. lipid membrane through which few water-soluble sub-

H
Bulk Passage Into and Out of the Cell. To transport large stances can pass, the membrane contains protein passage-

1
OH
particles, membranes form vesicles. ways that permit specific substances to move in and out of
the cell and allow the cell to exchange information with its
4

5
6.5 Active transport across membranes is powered by environment. We call this delicate skin of protein mole-
H

HO
energy from ATP. cules embedded in a thin sheet of lipid a plasma mem-
biological membranes.

brane. This chapter will examine the structure and func-

3

Active Transport. Cells transport molecules up a

4
H concentration gradient using ATP-powered carrier proteins. tion of this remarkable membrane.
H

OH
Coupled Transport. Active transport of ions drives coupled
OH
cules that contain carbon,

6 CH2OH
5 CH 2OH
|
|

uptake of other molecules up their concentration gradients.

atoms in the molecule with H

O
O

Fructose
H

H
HO
Ribose
5-carbon sugars

3
OH
2
molar ratio 1:2:1. Their empirical

1
H

103
2
1
OH
H—C—Fatty acid

CH 2OH
4

4
H

H
OH
H—C—Phosphate group

3
are well suited for energy storage.

H
H
5
OH

OH
5 CH 2OH

O
Because the C—H bonds in lipids are very nonpolar,
they are not water-soluble, and aggregate together in

6 CH2OH
H

H
water. This kind of aggregation by phospholipids forms

Galactose
2
OH
Deoxyribose
H

H
1
1
bon atoms. Because they contain many carbon-hydrogen
there are of each) is (CH2O)n, where n is the number of car-
The carbohydrates are a loosely defined group of mole-

subscripts to indicate how many

hydrogen, and oxygen in the
formula (which lists the

(C—H) bonds, which release energy when they are broken,

OH

O OH
 gether
CH3 water, forming O
two CHlayers of tails pointed toward each ot
O C (CH2)14 CH3
Biological
ds is of er—a lipid
membranes arebilayer
f luid (figure
layers of 3.18).
lipid. Lipid bilayers are the bas
CH2 O C (CH2)7 CH CH (CH2)7 CH3
framework of biological membranes, discussed in detail
ructure
Phospholipid Bilayer
chapter 6. The Core
O
mbranes that encase all living cells are sheets of
G
Fatty acid

(a) Phospholipid (phosphatidyl

L choline)
y two molecules thick; more than 10,000 of these Y
led on one another would just equal the thickness C
H lipid layer ❖
heet of paper. The O H
that forms
Glycerol H H H H
the foun- H E
R Fatty acid
a cell membrane is composed of molecules called
lipids (figure
H 6.2).
C O C C C C C C H
| O
L
H—C—Fatty
Phosphorylated
acid
ules
olipidsof H H H H H
alcohol
|
branes.
fat molecules you studied in chapter
O 3,Ha phos-
H H
(a)
H HH—C—Fatty acid
, made
has a backbone derived from a three-carbon
C C H |
e called glycerol. Attached to this backbone are
H C O
ds, long chains of carbon atoms endingC Cin a C car- C
CH 3
—COOH) group. A fat molecule has three such H—C—Phosphate group
H Hbe- H
ne attached to each carbon in the backbone; H H |
carbon
ese chains are nonpolar, they do not form hydro-
ds with water, and the fat moleculeOis not H H
H water- H H H
back-
A phospholipid, by contrast, has only two fatty Polar
HO
H to its
ns attached C backbone.
O C carbon
The third H
C Con C C C(hydrophilic) region
Nonpolar (hydrophobic) region
bone is attached instead to a highly polar organic
carbon H hydrogen
that readily forms Because
bonds withthe HC—H
H water. H HbondsH in lipids are very nonpolar,
this alcohol is attached they are
by a phosphatenot water-soluble,
group, and aggregate together in
p. Two
cule is called a phospholipid.
(b)water.
Triacylglycerol molecule
is,This kind of aggregation by phospholipids forms
ne in a
end of a phospholipid molecule therefore,
nonpolar (water-insoluble), while the other end is
biological membranes.
polar (water-soluble). The two nonpolar fatty
(b)
 H—C—Fatty acid
An individual phospholipid is a composite molecule, made |
up of three kinds of subunits: H—C—Phosphate group
1. Glycerol, a three-carbon alcohol, with each carbon |
H
bearing a hydroxyl group. Glycerol forms the back-

The Membrane
bone of the phospholipid molecule.
2. Fatty acids, long chains of C—H bonds (hydrocarbon
chains) ending in a carboxyl (—COOH) group. Two
Because the C—H bonds in lipids are very nonpolar,
they are not water-soluble, and aggregate together in
water. This kind of aggregation by phospholipids forms
fatty acids are attached to the glycerol backbone in a
biological membranes.
phospholipid molecule.

Oil
Hydrophobic
"tails"
Hydrophilic
"heads" Water
(a)

Water
Hydrophilic
region

Hydrophobic
region

Hydrophilic
region
Water
(b)

FIGURE 3.18
Phospholipids. (a) At an oil-water interface, phospholipid molecules will orient so that their polar (hydrophilic) heads are in the polar
medium, water, and their nonpolar (hydrophobic) tails are in the nonpolar medium, oil. (b) When surrounded by water, phospholipid
molecules arrange themselves into two layers with their heads extending outward and their tails inward.
 A Phospholipid

CH3 O–

CH3 N+ CH2 CH2 O P O CH2 O

CH3 O CH O C (CH2)14 CH3

CH2 O C (CH2)7 CH CH (CH2)7 CH3

❖ Phosphatidyl Choline
O C

(a) Phospholipid (phosphatidyl choline)

H O H H H H H

H C O C C C C C C H H3C

H H H H H
CH
O H H H H H

H C O C C C C C C H CH3
 3 2 2 2

CH3 O CH O C (CH2)14 CH3

Storage
CH2 O C (CH2)7 CH CH (CH2)7 CH3

O
(a) Phospholipid (phosphatidyl choline)

H O H H H H H ❖ Triglycerides OR
H C O C C C C C C H Triacylglycerol
O H H H H H H H H H H H H H H H
HO C C C C C C C C C C C C C C C C H
O H
HO C C
H H H H H H H H H H H H H H H H

H H H H H No double bonds between carbon atoms; fatty acid

Saturated Fatty acids
chains fit close together
Dou
cha

O H H H H H

H C O C C C C C C H C CH3 C
O H H H H H H H H H H H H H H H O H H H H H H H H H H H H H H
HO C C C C C C C C C C C C C C C C H HO C C C C C C C C C C C C C C C C C C H
H H H H
H H
H H H H H
H H
H H H H H
H
H H Saturated
(a) H H H fat H H H H H H H H H H
No double bonds between carbon atoms; fatty acid Double bonds present between carbon atoms; fatty acid

O H
chains fit close together
H H H H Unsaturated Fatty acids
chains do not fit close together

H C O C C C
H H
(a) Saturated fat
(b) Triacylglycerol molecule
HO
C C C H FIGURE 3.20
Saturated and unsaturated fats. (a) Palmitic acid, with no double bonds and, thu
carbon chain, is a saturated fatty acid. Many animal triacylglycerols (fats) are satur
together, these triacylglycerols form immobile arrays called hard fat. (b) Linoleic a
H H H H maximum number of hydrogen atoms bonded to the carbon chain, is an unsaturat
C C
many kinks the double bonds introduce into the fatty acid chains prevent the triac
are liquid at room temperature.
(b) Unsaturated fat

Fats as Food bonds.

FIGURE 3.19 from hy
 Other Lipids
CH3 CH3 OH
2)14 CH3 CH3 CH3 OH

32)7 CH CH (CH2)C
7 CH3
CH2 C CH
CH2 CHCH2 CH2 Terpenes - long chain fatty acids - pigments, rubber
CH2
CH2 CH2 CHCH
2
2 CH2
CH 2
CH2

dyl choline) (c) Terpene (citronellol)

(c) Terpene (citronellol)

H3C CH2 CH2 CH3

CH CH2 CH
H3C CH2 CH3 CH2 CH3 CH3

CH CH3 CH2 CH Steroids - 4 carbon rings - animal membranes, harmones

CH3
CH3
HO

(d) Steroid (cholesterol)

iologically important lipids: (a) phospholipids, (b) triacylglycerols

(d) Steroid (cholesterol)

tween one or more pairs of successive carbon atoms, the

fatty acid is said to be unsaturated. If a given fatty acid has
pids, fat more than one double bond, it is said to be polyunsaturat-
glycerol ed. Fats made from polyunsaturated fatty acids have low
a)tophospholipids,
each (b) triacylglycerols
melting points because their fatty acid chains bend at the
ns three double bonds, preventing the fat molecules from aligning
or, more closely with one another. Consequently, a polyunsaturated
ee fatty fat such as corn oil is usually liquid at room temperature and
ten they is called an oil. In contrast, most saturated fats such as those
the en- in butter are solid at room temperature.
e many Organisms contain many other kinds of lipids besides
 A plasma membrane is composed of both lipids and glob-
ular proteins. For many years, biologists thought the pro-
tein covered the inner and outer surfaces of the phospho-
lipid bilayer like a coat of paint. The widely accepted
Davson-Danielli model, proposed in 1935, portrayed the
Membranes are Fluid
membrane as a sandwich: a phospholipid bilayer between
two layers of globular protein. This model, however, was
not consistent with what researchers were learning in the
1960s about the structure of membrane proteins. Unlike
most proteins found within cells, membrane proteins are
not very soluble in water—they possess long stretches of
nonpolar hydrophobic amino acids. If such proteins in-
deed coated the surface of the lipid bilayer, as the
Davson-Danielli model suggests, then their nonpolar por-
tions would separate the polar portions of the phospho-
lipids from water, causing the bilayer to dissolve! Because
this doesn’t happen, there is clearly something wrong
with the model.
Extracellular fluid
Glycolipid
Singer & Nicolson
Carbohydrate
Cholesterol
Filaments of
Cytoplasm
cytoskeleton
proteins are inserted into the lipid bilayer, with their nonpo-
lar segments in contact with the nonpolar interior of the
bilayer and their polar portions protruding out from the
membrane surface. In this model, called the fluid mosaic
model, a mosaic of proteins float in the fluid lipid bilayer
like boats on a pond (figure 6.5).
Components of the Cell Membrane
A eukaryotic cell contains many membranes. While they
are not all identical, they share the same fundamental ar-
chitecture. Cell membranes are assembled from four com-
ponents (table 6.1):
1. Lipid bilayer. Every cell membrane is composed of
a phospholipid bilayer. The other components of the
membrane are enmeshed within the bilayer, which
provides a flexible matrix and, at the same time, im-
poses a barrier to permeability.
Glycoprotein
Transmembrane
protein
Peripheral
protein
The Fluid Mosaic Model
 onsix
n sixkey
keyclasses
classes 5.5.Cell
Celladhesion
adhesionproteins.
proteins. Cells CellsuseuseAttachments
specificproteins
specific proteins
6.6.
6.Attachments
Attachments toto the
tothe cytoskeleton.
thecytoskeleton.
cytoskeleton.Outside Surface
Surface pro-
Surfacepro-
pro-
Transporters.
Transporters.
ollowing
llowing chapterchapter Membranes
Membranes are
are
totoglue very
very selective,
selective,
gluethemselves
themselvestotoone al-
al-
oneanother.
another.SomeSome act
act like
like Vel-
Vel-
teins that
teinsthat
teins interact
thatinteract with
interactwith other
withother cells
othercells are
cellsare often
areoften anchored
oftenanchored
anchored
lowingonly
owing certainsubstances
onlycertain substances cro,
cro, towhile
enter
towhile
enter orleave
or
others
others leave
form
form the
the
a amore
morepermanent
permanent bond.
bond.
to the
totothe cytoskeleton
thecytoskeleton by linking
cytoskeletonbybylinking proteins.
linkingproteins.
proteins.
cell, eitherthrough
ell, either throughchannels
channels orcarriers.
or carriers.In
6.6.Attachments
Attachments Intosome
some
tothe in-
in-
the cytoskeleton. Surface
cytoskeleton. Surfacepro- pro-
eryselective,
ry
tances,selective,
they al-up
al-
take upmolecules
molecules already presentwithinthe
the
stances, theytake already
teins
teins that present
thatinteract
interact inwith othercells
other cellsareareoften
oftenanchored
anchored Plasma membrane
nter
ter oror leave
leave the
the

Functions on the membrane

ell ininhigh
cell concentration. totothe
highconcentration. The
The
The many
many
many proteins
proteins
proteins embedded
embedded
embedded within
within a
within a membrane
membrane
a membrane carry
carry
carry
thecytoskeleton
cytoskeletonbybylinking
linkingproteins.
proteins.
riers.
ers. InInsome
Enzymes. some in-carry
in-
Cells
Cells carryout
outmany
manychemical
chemicalreactions
reactions out
outaaahost
out host
hostofof functions,
offunctions, many
manyofof
functions,many which
ofwhich
whichare are associated
areassociated
associated
Enzymes.
adypresent
dy presentininthethesurfaceofofthe with
with transport
transportofof
withtransport materials
materialsoror
ofmaterials information
orinformation across
informationacross the
the
across the
on
on the
the interior
interiorsurface The
The the
many plasma
plasma
many proteinsmembrane,
membrane,
proteins embeddedwithin
embedded within a a membrane
membrane carry
carry
membrane.
membrane.
membrane.
using
usingenzymes attachedtotoout
enzymesattached theamembrane.
the
out membrane.
ahost
hostofoffunctions,
functions,manymanyofofwhich
whichare areassociated
associated Inside
hemical reactions
emical reactions
withtransport
with transportofofmaterials
materialsororinformation
informationacrossacrossthethe
asma membrane,
ma membrane,
membrane.
membrane.
brane.
ane. Transporter Enzyme

de
e

ma membrane
a membrane
ma membrane

e
e

Transporter
Transporter
Transporter Enzyme
Enzyme
Enzyme Cell
Cell
Cell surface
surface receptor
receptor
surface receptor Cell surface identity Cell adhesion
Reaction marker

Transport Signalling
Figure 6.7 Identifiers
Enzyme
Enzyme Cell
Catalysis
Cellsurface
surfacereceptor
receptor
Functions of plasma membrane proteins. Membrane proteins act as transporters, en
markers, as well as aiding in cell-to-cell adhesion and securing the cytoskeleton.

Cell
Cellsurface
Cell surfaceidentity
surface identity
identity Cell
Celladhesion
Cell adhesion
adhesion Attachment
Attachment
Attachmenttotothe
to the
the
marker
marker
marker cytoskeleton
cytoskeleton
cytoskeleton

ee6.7
6.7
6.7 Celladhesion
Cell adhesion Cytoskeletal
Attachmenttotothe
Attachment the
ons
ionsofof
ions
s,
rs,as
as
ofplasma
well
plasmamembrane
plasma
wellas
as
membrane
aiding
aidingin
in Adhesion
membraneproteins.
proteins.Membrane
proteins.
cell-to-cell
Membraneproteins
Membrane
adhesion
cell-to-cell adhesionand
and
proteinsact
proteins
securing
securingthe
the
act
actasastransporters,
as transporters,enzymes,
transporters,
cytoskeleton
cytoskeleton
cytoskeleton.
cytoskeleton.
enzymes,cell
enzymes, cellsurface
cell surfacereceptors,
surface receptors,and
receptors, andcell
and cellsurface
cell surface
surface
Attachment
rs, as well as aiding in cell-to-cell adhesion and securing the cytoskeleton.

embraneproteins
mbrane proteinsact
actasastransporters,
transporters,enzymes,
enzymes,cell
cellsurface
surfacereceptors,
receptors,and
andcell
cellsurface
surface
 layer of the membrane. immersed in a beaker of distilled water, the salt cannot cross the
Even water molecules, which are very polar, cannot
Glycophorin membrane, but water can. The water entering the tube causes the
in cross a lipid bilayer. Water flows through aquaporins, salt solution to rise in the tube. (c) Water will continue to enter the
which are specialized channels for water. A simple experi-
Cytoskeletal tube from the beaker until the weight of the column of water in the

Osmosis
r ment demonstrates this. If proteins
you place an amphibian egg in tube exerts a downward force equal to the force drawing water
n Junctional
hypotonic spring water, it does not swell. If you then inject molecules upward into the tube. This force is referred to as
complex
aquaporin mRNA into the egg, the channel proteins are ex- osmotic pressure.
100 nm
pressed and the egg then swells.
Dissolved solutes interact with water molecules, which
form hydration shells about the charged solute. When there Hyperosmotic Isosmotic Hypoosmotic
is a concentration gradient of solutes, the solutes will move solution solution solution
from a high to a low concentration, dragging with them their
hydration shells of water molecules. When a membrane sepa-
Porin monomer
rates two solutions, hydration shell water molecules move
with the diffusing ions, creating a!-pleated
net movement
sheets
of water to-
wards the low solute. This net water movement across a
membrane by diffusion is called osmosisBacterial
(figure 6.14). Shriveled cells Normal cells Cells swell and
The concentration of all solutes in a outer
solution determines eventually burst
the osmotic concentration of the solution. membrane If two solu-
tions have unequal osmotic concentrations, the solution
with the higher concentration is hyperosmotic (Greek
hyper, “more than”), and the solution with the lower con-
centration is hypoosmotic (Greek hypo, “less than”). If the
osmotic concentrations of two solutions are equal, the solu-
tions are isosmotic (Greek iso, “the same”).
In cells, a plasma membrane separates two aqueous solu-
tions, one inside the cell (the cytoplasm) and one outside Human red blood cells

FIGURE 6.15
Osmosis. In a hyperosmotic solution water moves out of the cell
toward the higher concentration of solutes, causing the cell to
shrivel. In an isosmotic solution, the concentration of solutes on
either side of the membrane is the same. Osmosis still occurs, but
water diffuses into and out of the cell at the same rate, and the cell Cell body shrinks
doesn’t change size. In a hypoosmotic solution the concentration of Flaccid cell Normal turgid cell
from cell wall
FIGURE
solutes is 6.11
higher within the cell than without, so the net movement
is in Aofpore protein. Thecell.
bacterial transmembrane protein Plant cells
water is into the
 Some Transports need Fuel
Extracellular

P
P
P
P
ATP A
P ATP P
P
P P
Na+ A A
ADP
Intracellular

1. Protein in membrane binds intracellular 2. ATP phosphorylates protein with bound 3. Phosphorylation causes conformational
sodium. sodium. change in protein, allowing sodium to
leave.

K+

P P ATP
P P P
P A
A ADP+Pi P
A
ADP

4. Extracellular potassium binds to exposed 5. Binding of potassium causes dephos- 6. Dephosphorylation of protein triggers
sites. phorylation of protein. change back to original conformation,
potassium moves into cell, and the cycle
repeats.
 Meeting the Energy Requirement
Outside of cell

entration
Sugar
ctly. The
Na+
r protons
This type
ponents:
s used to
Coupled
gradient, Na/K
transport
molecule pump
protein

proteins
the mol-
together
K+
proton is Inside of cell
be trans-
 H C OH O OH O OH
2
4 1 4 1
H
3
C OH Carbohydrates
H
3 2
H
H H
3
H H H
2
H
H
OH OH OH H
C : H : O
Glyceraldehyde Ribose
1 : 2 : 1 Deoxyribose

(C H2 O)n
6-carbon sugars

6 CH2OH 6 CH2OH 6 CH2OH

5
H 5 O H O H OH O OH
H H
4 1 5 2 4 1
OH HO
H H HO OH H
HO OH HO CH 2OH H H
3 2 4 3 1 2
3
H OH OH H H OH
Glucose Fructose Galactose
Glucose Fructose Galactose
FIGURE 3.21
 es. Many familiar sugars like sucrose are CH2OH CH2OH

Carbohydrates
rs,” two monosaccharides joined by a covalent O O
H H H
3.23). Called disaccharides, they often play a H
ansport of sugars, as we will discuss shortly. OH H O H HO
HO CH2OH

ides. Polysaccharides are macromolecules H OH OH H

monosaccharide subunits. Starch is a polysac- Disaccharides Glucose Fructose
by plants to store energy. It consists entirely of Sucrose
cules, linked one after another in long chains.
Glucosethat
a polysaccharide + Fructose
serves as a structural Glucose + Galactose
erial in plants. It too consists entirely of glucose
CH2chains,
OH CH2OH CH2OH
ked together into and special enzymes O
o break the O O CH OH H OH
H links. H H 2 H
H OH O OH HO H
H O H
OH H O H HO
ers HO CH2OH H
OH H
H H OH
ot the only sugar
H withOH the formulaOH C6H12H O6. H OH Glucose
on six-carbon sugars such as fructose and ga- Galactose
Glucose Fructose
Sucrose
have this same empirical formula (figure 3.24). Lactose
Lactose
Sucrose
are isomers, or alternative forms, of glucose.
h isomers have the same empirical formula, FIGURE 3.23
are arranged in different ways; that is, theirPolysaccharides
Disaccharides. Sugars like sucrose and lactose are disaccharides,
ional structures are different. These structural composed of two monosaccharides linked by a covalent bond.
ften account for substantialStarch
functional differ-
CH2OH Cellulose
n the isomers. Glucose and fructose, for exam-O
CH 2OH
tural isomers. In fructose, H
the double-bonded OH galactose and glucose are called stereoisomers. Again, this
H
ached to OH O
an internal carbon ratherOHthanHOto
H a seemingly slight difference has important consequences, as
H O H
 Plants

-ssorc era sniahc eseht

-losni na etaerc sknil-sso
-ed eb nac hcihw ,esocu
fo dnik rehtona yb
sreffid hsem eht fo ezis
001 tuoba ecir ni ;tnalp
owt ro eno htiw hcae ,
.hsem eht sm
-ylg si hcrats fo noisrev
na si n e g o c y l g ,nitcepolym
gniniatnoc edirahccas
Amylose

,negocylg nI .sniahc eso

Amylopectin
hcum si htgnel niahc
sehcnarb erom era ere
nitcepolymA )b( esolymA )a( -uH .) c62.3 erugif( hcra
-xe erots setarbetrev re
-vil eht ni negocylg sa y
eht nehw ;sllec elcsu
-saercni eussit a ni ygre
esaeler ot dezylordyh s

steewS g
taht ragus elbat fo dn
Storage

laer eht ekil skooc dna

lufmrah ro seirolac on
sniatnuom tae dluoc uo
sreneteews hcus morf e
-saP siuoL sA .thgiew g
tsom ,s0081 etal eht ni d
,selucelom ”dednah-thg
a sdnib taht puorg lyxor
thgir eht no si mota n
,sragus ”dednah-tfel“ ,
Glycogen

eht no si puorg lyxordy

eht ni ylidaer edam eb
era sragus citehtnys ese
eht fo sniwt lacimehc
negocylG )c(
taht semyzne eht tub
62.3 ERUGIF -id namuh eht ni sragu
.stnalp ni ygrene erots taht sremylop esoculg gnol era sehcratS .sedirahccasylop egarotS
Animals .ecnereffid eht llet nac
pu lioc ot dnet hcihw ,esolyma dellac esotlam fo sniahc gnol era sehcrats tselpmis ehT )a( emyzne na ,elucelom rag
era hcihw ,snitcepolyma dellac sehcrats xelpmoc erom niatnoc stnalp tsoM )b( .retaw ni eohs a ekil hcum ,ti ps
naht dehcnarb ylevisnetxe erom si hcihw ,negocylg ni esoculg erots slaminA )c( .dehcnarb eht fo lla dna ,toof a
.esolyma fo sniahc regnol sniatnoc dna nitcepolyma A !dednah-thgir era se
 Protection

FIGURE 3.27
A journey into wood. This jumble of cellulose

Animals
fibers (a) is from a yellow pine (Pinus ponderosa)
Plants

(20×). (b) While starch chains consist of alpha-

glucose subunits, (c) cellulose chains consist of
beta-glucose subunits. Cellulose fibers can be
very strong and are quite resistant to metabolic
breakdown, which is one reason why wood is such
a good building material.

O O O O
1 1 4
O O O
H

e Starch: chain of α-glucose subunits

O
 8.3
Life’s Currency
ATP is the energy currency of life.

What Is ATP? ATP

The chief energy currency all cells use is a molecule called
adenosine triphosphate (ATP). Cells use their supply of Triphosphate
ATP to power almost every energy-requiring process they group
carry out, from making sugars, to supplying activation en- O
–

ergy for chemical reactions, to actively transporting sub-

—
stances across membranes, to moving through their envi- O == P — O–
ronment and growing. (a)

O
Structure of the ATP Molecule
Each ATP molecule is a nucleotide composed of three High-energy
bonds
smaller components (figure 8.14). The first component is a
O == P — O– NH2
five-carbon sugar, ribose, which serves as the backbone to Adenine
which the other two subunits are attached. The second N
N
component is adenine, an organic molecule composed of
two carbon-nitrogen rings. Each of the nitrogen atoms in O
N
the ring has an unshared pair of electrons and weakly at- N
tracts hydrogen ions. Adenine, therefore, acts chemically as AMP
a base and is usually referred to as a nitrogenous base (it is O == P — O
— CH2 O core
one of the four nitrogenous bases found in DNA and
ATPRNA). The third component ADPof + ATPPiis a triphosphate O–
H
H H
H
group (a chain of three phosphates).
7.3 kcal / mole OH OH
(b) Ribose
How ATP Stores Energy
er what happens when the trans- to be channeled into more useful paths. In the combustion
te. of gasoline, the same amount of energy is released whether
all of the gasoline in a car’s gas tank explodes at once, or
whether the gasoline burns in a series of very small explo-

Energy for ATP

ation-Reduction sions inside the cylinders. By releasing the energy in gaso-
line a little at a time, the harvesting efficiency is greater and
is an oxidation-reduction reac-
ns in the C—H bonds of glucose
equally between the C and H
hydrogen nuclei have about the Electrons from food
ectrons (that is, they exhibit sim-
ever, when the carbon atoms of
to form carbon dioxide, the elec- e– High energy
bonds take a different position. Energy for
ually, the electrons that were as- synthesis of
toms in glucose shift far toward
ATP
ecause oxygen is very electroneg-
are pulled farther from the car-
ms of glucose have been oxidized
oxygen atoms reduced (gain of
the hydrogen atoms of glucose
oms to form water, the oxygen
ectrons strongly toward them;
Electron
nd glucose is oxidized. In this re- transport
zing (electron-attracting) agent chain
ms of glucose.

Low energy
g the oxidation of glucose is to
e–
e shared electrons. In a covalent
d to remove an electron from an
be used to roll a boulder up a hill. Formation of water
the atom, the steeper the energy
quence of 10 reactions that convert glucose into 2 three-
carbon molecules of pyruvate (figure 9.7). For each mole-
cule of glucose that passes through this transformation,
the cell nets two ATP molecules by substrate-level phos-
phorylation.
glucose by changing it into a compound that can be
cleaved readily into 2 three-carbon phosphorylated mole-
cules. Two of these reactions require the cleavage of ATP,
so this step requires the cell to use two ATP molecules.
Step B: Cleavage and rearrangement. In the first of

Energy from Carbohydrates

the remaining pair of reactions, the six-carbon product
of step A is split into 2 three-carbon molecules. One is
G3P, and the other is then converted to G3P by the sec-
ond reaction (figure 9.8).

OVERVIEW OF GLYCOLYSIS
1 6-carbon glucose
2 3
(Starting material)
2 ATP

P P P P

6-carbon sugar diphosphate 6-carbon sugar diphosphate

P P P P

3-carbon sugar 3-carbon sugar 3-carbon sugar 3-carbon sugar

phosphate phosphate phosphate phosphate

NADH NADH

2 ATP 2 ATP

3-carbon 3-carbon
pyruvate pyruvate

Priming reactions. Glycolysis begins
with the addition of energy. Two high-
energy phosphates from two molecules
of ATP are added to the six-carbon
molecule glucose, producing a six-carbon
molecule with two phosphates.
Cleavage reactions. Then, the six-carbon
molecule with two phosphates is split in
two, forming two three-carbon sugar
phosphates.
Energy-harvesting reactions. Finally, in
a series of reactions, each of the two three-
carbon sugar phosphates is converted to
pyruvate. In the process, an energy-rich
hydrogen is harvested as NADH, and two
ATP molecules are formed.
 Glucose CH2OH
O
1
ATP
1. Phosphorylation of Hexokinase
glucose by ATP. ADP CH2 O P
O
Glucose 6-phosphate

2
2–3. Rearrangement, Phosphoglucoisomerase
followed by a second CH2 O P
ATP phosphorylation. O CH2OH
Fructose 6-phosphate

3
ATP
Phosphofructokinase
4–5. The six-carbon P O CH2 CH2 O P
molecule is split into ADP O
two three-carbon
molecules—one G3P, Fructose 1,6-bisphosphate
another that is Aldolase
converted into G3P in 4,5
H
another reaction. P O CH2 Isomerase
C O
C O Dihydroxyacetone Glyceraldehyde 3-
6. Oxidation followed by phosphate phosphate (G3P) CHOH
CH2OH
phosphorylation produces
CH2 O P
two NADH molecules and NAD+ Pi Pi NAD+
two molecules of BPG, 6
each with one high-energy NADH Triose phosphate NADH
phosphate bond. dehydrogenase P O C O
1,3-Bisphosphoglycerate 1,3-Bisphosphoglycerate CHOH
(BPG) (BPG)
7. Removal of high-energy CH2 O P
phosphate by two ADP ADP 7 ADP
–
molecules produces two Phosphoglycerokinase O
ATP molecules and leaves ATP
ATP C O
two 3PG molecules.
3-Phosphoglycerate 3-Phosphoglycerate CHOH
(3PG) (3PG) CH2 O P

8 –
O
Phosphoglyceromutase
C O
8–9. Removal of water yields
two PEP molecules, H C O P
2-Phosphoglycerate 2-Phosphoglycerate
each with a high-energy
(2PG) (2PG) CH2OH
phosphate bond.
9 O
–

H 2O Enolase H 2O
C O

C O P
Phosphoenolpyruvate Phosphoenolpyruvate
CH2
(PEP) (PEP)
10. Removal of high-energy O
–
phosphate by two ADP ADP 10 ADP
molecules produces two Pyruvate kinase C O
ATP molecules and two ATP ATP C O
pyruvate molecules.
Pyruvate Pyruvate CH3
 Income from Aerobic Respiration
mmarizing Aerobic
spiration
Glucose
much metabolic energy does a cell
ally gain from the electrons har-
d from a molecule of glucose, using 2 ATP
lectron transport chain to produce 2 ATP Glycolysis
by chemiosmosis?
2 NADH 4 ATP
Pyruvate
oretical Yield
chemiosmotic model suggests that
ATP molecule is generated for 2 NADH 6 ATP
proton pump activated by the
ron transport chain. Since the elec- Acetyl-CoA
s from NADH activate three
ps and those from FADH2 activate
we would expect each molecule of 2 ATP
DH and FADH2 to generate three
two ATP molecules, respectively.
ever, because eukaryotic cells carry 6 NADH 18 ATP
Krebs
glycolysis in their cytoplasm and cycle
Krebs cycle within their mitochon-
they must transport the two mole-
of NADH produced during gly- 2 FADH2 4 ATP
sis across the mitochondrial
branes, which requires one ATP
molecule of NADH. Thus, the net
production is decreased by two. Total net ATP yield = 36 ATP
efore, the overall ATP production
ting from aerobic respiration theo-
lly should be 4 (from substrate-
KEGG Database