Membrane Structure and Function campbell

omar164646 80 views 83 slides Aug 16, 2024
Slide 1
Slide 1 of 83
Slide 1
1
Slide 2
2
Slide 3
3
Slide 4
4
Slide 5
5
Slide 6
6
Slide 7
7
Slide 8
8
Slide 9
9
Slide 10
10
Slide 11
11
Slide 12
12
Slide 13
13
Slide 14
14
Slide 15
15
Slide 16
16
Slide 17
17
Slide 18
18
Slide 19
19
Slide 20
20
Slide 21
21
Slide 22
22
Slide 23
23
Slide 24
24
Slide 25
25
Slide 26
26
Slide 27
27
Slide 28
28
Slide 29
29
Slide 30
30
Slide 31
31
Slide 32
32
Slide 33
33
Slide 34
34
Slide 35
35
Slide 36
36
Slide 37
37
Slide 38
38
Slide 39
39
Slide 40
40
Slide 41
41
Slide 42
42
Slide 43
43
Slide 44
44
Slide 45
45
Slide 46
46
Slide 47
47
Slide 48
48
Slide 49
49
Slide 50
50
Slide 51
51
Slide 52
52
Slide 53
53
Slide 54
54
Slide 55
55
Slide 56
56
Slide 57
57
Slide 58
58
Slide 59
59
Slide 60
60
Slide 61
61
Slide 62
62
Slide 63
63
Slide 64
64
Slide 65
65
Slide 66
66
Slide 67
67
Slide 68
68
Slide 69
69
Slide 70
70
Slide 71
71
Slide 72
72
Slide 73
73
Slide 74
74
Slide 75
75
Slide 76
76
Slide 77
77
Slide 78
78
Slide 79
79
Slide 80
80
Slide 81
81
Slide 82
82
Slide 83
83

About This Presentation

Membrane Structure and Function : The Fluid Mosaic Model
The cell membrane, also known as the plasma membrane, is a complex structure that surrounds every cell. It's composed primarily of a phospholipid bilayer, which is a double layer of phospholipid molecules. This bilayer ...


Slide Content

Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
PowerPoint
®
Lecture Presentations for
Biology
Eighth Edition
Neil Campbell and Jane Reece
Lectures by Chris Romero, updated by Erin Barley with contributions from Joan Sharp
Chapter 7
Membrane Structure and
Function

Overview: Life at the Edge
•The plasma membrane is the boundary that
separates the living cell from its surroundings
•The plasma membrane exhibits selective
permeability, allowing some substances to
cross it more easily than others
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-1

Concept 7.1: Cellular membranes are fluid mosaics
of lipids and proteins
•Phospholipids are the most abundant lipid in
the plasma membrane
•Phospholipids are amphipathic molecules,
containing hydrophobic and hydrophilic regions
•The fluid mosaic model states that a
membrane is a fluid structure with a “mosaic” of
various proteins embedded in it
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Membrane Models: Scientific Inquiry
•Membranes have been chemically analyzed
and found to be made of proteins and lipids
•Scientists studying the plasma membrane
reasoned that it must be a phospholipid bilayer
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-2
Hydrophilic
head
WATER
Hydrophobic
tail
WATER

•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, 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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-3
Phospholipid
bilayer
Hydrophobic regions
of protein
Hydrophilic
regions of protein

•Freeze-fracture studies of the plasma
membrane supported the fluid mosaic model
•Freeze-fracture is a specialized preparation
technique that splits a membrane along the
middle of the phospholipid bilayer
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-4
TECHNIQUE
Extracellular
layer
Knife
ProteinsInside of extracellular layer
RESULTS
Inside of cytoplasmic layer
Cytoplasmic layerPlasma membrane

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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-5
Lateral movement
(~10
7
times per second)
Flip-flop
(~ once per month)
(a) Movement of phospholipids
(b) Membrane fluidity
Fluid Viscous
Unsaturated hydrocarbon
tails with kinks
Saturated hydro-
carbon tails
(c) Cholesterol within the animal cell membrane
Cholesterol

Fig. 7-5a
(a) Movement of phospholipids
Lateral movement
(10
7
times per
second)
Flip-flop
( once per month)

Fig. 7-6
RESULTS
Membrane proteins
Mouse cell
Human cell
Hybrid cell
Mixed proteins
after 1 hour

•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 that those rich in saturated fatty
acids
•Membranes must be fluid to work properly;
they are usually about as fluid as salad oil
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-5b
(b) Membrane fluidity
Fluid
Unsaturated hydrocarbon
tails with kinks
Viscous
Saturated hydro-
carbon tails

•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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-5c
Cholesterol
(c) Cholesterol within the animal cell membrane

Membrane Proteins and Their Functions
•A membrane is a collage of different proteins
embedded in the fluid matrix of the lipid bilayer
•Proteins determine most of the membrane’s
specific functions
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-7
Fibers of
extracellular
matrix (ECM)
Glyco-
protein
Microfilaments
of cytoskeleton
Cholesterol
Peripheral
proteins
Integral
protein
CYTOPLASMIC SIDE
OF MEMBRANE
Glycolipid
EXTRACELLULAR
SIDE OF
MEMBRANE
Carbohydrate

•Peripheral proteins are bound to the surface
of the membrane
•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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-8
N-terminus
C-terminus
 Helix
CYTOPLASMIC
SIDE
EXTRACELLULAR
SIDE

•Six major functions of membrane proteins:
–Transport
–Enzymatic activity
–Signal transduction
–Cell-cell recognition
–Intercellular joining
–Attachment to the cytoskeleton and
extracellular matrix (ECM)
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-9
(a) Transport
ATP
(b) Enzymatic activity
Enzymes
(c) Signal transduction
Signal transduction
Signaling molecule
Receptor
(d) Cell-cell recognition
Glyco-
protein
(e) Intercellular joining(f) Attachment to
the cytoskeleton
and extracellular
matrix (ECM)

Fig. 7-9ac
(a) Transport (b) Enzymatic activity(c) Signal transduction
ATP
Enzymes
Signal transduction
Signaling molecule
Receptor

Fig. 7-9df
(d) Cell-cell recognition
Glyco-
protein
(e) Intercellular joining
(f) Attachment to
the cytoskeleton
and extracellular
matrix (ECM)

The Role of Membrane Carbohydrates in Cell-Cell
Recognition
•Cells recognize each other by binding to
surface molecules, often carbohydrates, on the
plasma membrane
•Membrane carbohydrates may be covalently
bonded to lipids (forming glycolipids) or more
commonly to proteins (forming glycoproteins)
•Carbohydrates on the external side of the
plasma membrane vary among species,
individuals, and even cell types in an individual
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-10
ER
1
Transmembrane
glycoproteins
Secretory
protein
Glycolipid
2Golgi
apparatus
Vesicle
3
4
Secreted
protein
Transmembrane
glycoprotein
Plasma membrane:
Cytoplasmic face
Extracellular face
Membrane glycolipid

Concept 7.2: Membrane structure results in
selective permeability
•A cell must exchange materials with its
surroundings, a process controlled by the
plasma membrane
•Plasma membranes are selectively permeable,
regulating the cell’s molecular traffic
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

•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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Concept 7.3: Passive transport is diffusion of a
substance across a membrane with no energy
investment
•Diffusion is the tendency for molecules to
spread out evenly into the available space
•Although each molecule moves randomly,
diffusion of a population of molecules may
exhibit a net movement in one direction
•At dynamic equilibrium, as many molecules
cross one way as cross in the other direction
Animation: Membrane SelectivityAnimation: Membrane Selectivity Animation: DiffusionAnimation: Diffusion
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-11
Molecules of dyeMembrane (cross section)
WATER
Net diffusion Net diffusion Equilibrium
(a) Diffusion of one solute
Net diffusion
Net diffusion
Net diffusion
Net diffusion
Equilibrium
Equilibrium
(b) Diffusion of two solutes

Molecules of dye
Fig. 7-11a
Membrane (cross section)
WATER
Net diffusion Net diffusion
(a) Diffusion of one solute
Equilibrium

•Substances diffuse down their concentration
gradient, the difference in concentration of a
substance from one area to another
•No work must be done to move substances
down the concentration gradient
•The diffusion of a substance across a biological
membrane is passive transport because it
requires no energy from the cell to make it
happen
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

(b) Diffusion of two solutes
Fig. 7-11b
Net diffusion
Net diffusion
Net diffusion
Net diffusion
Equilibrium
Equilibrium

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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Lower
concentration
of solute (sugar)
Fig. 7-12
H
2O
Higher
concentration
of sugar
Selectively
permeable
membrane
Same concentration
of sugar
Osmosis

Water Balance of Cells Without Walls
•Tonicity is the ability of a solution to cause a
cell to gain or lose water
•Isotonic solution: Solute concentration is the
same as that inside the cell; no net water
movement across the plasma membrane
•Hypertonic solution: Solute concentration is
greater than that inside the cell; cell loses
water
•Hypotonic solution: Solute concentration is
less than that inside the cell; cell gains water
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-13
Hypotonic solution
(a) Animal
cell
(b) Plant
cell
H
2
O
Lysed
H
2
O
Turgid (normal)
H
2
O
H
2
O
H
2O
H
2
O
Normal
Isotonic solution
Flaccid
H
2O
H
2O
Shriveled
Plasmolyzed
Hypertonic solution

•Hypertonic or hypotonic environments create
osmotic problems for organisms
•Osmoregulation, the control of 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
Video: Video: ChlamydomonasChlamydomonas Video: Video: ParameciumParamecium Vacuole Vacuole
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-14
Filling vacuole
50 µm
(a) A contractile vacuole fills with fluid that enters from
a system of canals radiating throughout the cytoplasm.
Contracting vacuole
(b) When full, the vacuole and canals contract, expelling
fluid from the cell.

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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Video: PlasmolysisVideo: Plasmolysis
Video: Turgid Video: Turgid ElodeaElodea
Animation: OsmosisAnimation: Osmosis
•In a hypertonic environment, plant cells lose
water; eventually, the membrane pulls away
from the wall, a usually lethal effect called
plasmolysis
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Facilitated Diffusion: Passive Transport Aided by
Proteins
•In facilitated diffusion, transport proteins
speed the passive movement of molecules
across the plasma membrane
•Channel proteins provide corridors that allow a
specific molecule or ion to cross the membrane
•Channel proteins include
–Aquaporins, for facilitated diffusion of water
–Ion channels that open or close in response
to a stimulus (gated channels)
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-15
EXTRACELLULAR
FLUID
Channel protein
(a) A channel protein
Solute
CYTOPLASM
Solute
Carrier protein
(b) A carrier protein

•Carrier proteins undergo a subtle change in
shape that translocates the solute-binding site
across the membrane
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

•Some diseases are caused by malfunctions in
specific transport systems, for example the
kidney disease cystinuria
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Concept 7.4: Active transport uses energy to move
solutes against their gradients
•Facilitated diffusion is still passive because the
solute moves down its concentration gradient
•Some transport proteins, however, can move
solutes against their concentration gradients
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

The Need for Energy in Active Transport
•Active transport moves substances against
their concentration gradient
•Active transport requires energy, usually in the
form of ATP
•Active transport is performed by specific
proteins embedded in the membranes
Animation: Active TransportAnimation: Active Transport
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

•Active transport allows cells to maintain
concentration gradients that differ from their
surroundings
•The sodium-potassium pump is one type of
active transport system
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-16-1
EXTRACELLULAR
FLUID
[Na
+
] high
[K
+
] low
Na
+

Na
+

Na
+

[Na
+
] low
[K
+
] high CYTOPLASM
Cytoplasmic Na
+
binds to
the sodium-potassium pump.
1

Na
+
binding stimulates
phosphorylation by ATP.
Fig. 7-16-2
Na
+

Na
+

Na
+

ATP
P
ADP
2

Fig. 7-16-3
Phosphorylation causes
the protein to change its
shape. Na
+
is expelled to
the outside.
Na
+

P
Na
+

Na
+

3

Fig. 7-16-4
K
+
binds on the
extracellular side and
triggers release of the
phosphate group.
P
P

K
+
K
+
4

Fig. 7-16-5
Loss of the phosphate
restores the protein’s original
shape.
K
+
K
+
5

Fig. 7-16-6
K
+
is released, and the
cycle repeats.
K
+
K
+
6

2
EXTRACELLULAR
FLUID
[Na
+
] high
[K
+
] low
[Na
+
] low
[K
+
] high
Na
+
Na
+
Na
+
Na
+
Na
+
Na
+
CYTOPLASM
ATP
ADP
P
Na
+
Na
+
Na
+
P
3
K
+
K
+
6
K
+
K
+
5 4
K
+
K
+
P
P
1
Fig. 7-16-7

Fig. 7-17
Passive transport
Diffusion Facilitated diffusion
Active transport
AT
P

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
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

•Two combined forces, collectively called the
electrochemical gradient, drive the diffusion
of ions across a membrane:
–A chemical force (the ion’s concentration
gradient)
–An electrical force (the effect of the membrane
potential on the ion’s movement)
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

•An electrogenic pump is a transport protein
that generates voltage across a membrane
•The sodium-potassium pump is the major
electrogenic pump of animal cells
•The main electrogenic pump of plants, fungi,
and bacteria is a proton pump
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-18
EXTRACELLULAR
FLUID
H
+

H
+

H
+

H
+

Proton pump
+
+
+
H
+

H
+

+
+
H
+





ATP
CYTOPLASM

Cotransport: Coupled Transport by a Membrane
Protein
•Cotransport occurs when active transport of a
solute indirectly drives transport of another
solute
•Plants commonly use the gradient of hydrogen
ions generated by proton pumps to drive active
transport of nutrients into the cell
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-19
Proton pump






+
+
+
+
+
+
ATP
H
+
H
+
H
+
H
+
H
+
H
+
H
+
H
+
Diffusion
of H
+
Sucrose-H
+
cotransporter
Sucrose
Sucrose

Concept 7.5: Bulk transport across the plasma
membrane occurs by exocytosis and endocytosis
•Small molecules and water enter or leave the
cell through the lipid bilayer or by transport
proteins
•Large molecules, such as polysaccharides and
proteins, cross the membrane in bulk via
vesicles
•Bulk transport requires energy
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Exocytosis
•In exocytosis, transport vesicles migrate to the
membrane, fuse with it, and release their
contents
•Many secretory cells use exocytosis to export
their products
Animation: ExocytosisAnimation: Exocytosis
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

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
Animation: Exocytosis and Endocytosis IntroductionAnimation: Exocytosis and Endocytosis Introduction
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

•In phagocytosis a cell engulfs a particle in a
vacuole
•The vacuole fuses with a lysosome to digest
the particle
Animation: PhagocytosisAnimation: Phagocytosis
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-20
PHAGOCYTOSIS
EXTRACELLULAR
FLUID
CYTOPLASM
Pseudopodium
“Food”or
other particle
Food
vacuole
PINOCYTOSIS
1 µm
Pseudopodium
of amoeba
Bacterium
Food vacuole
An amoeba engulfing a bacterium
via phagocytosis (TEM)
Plasma
membrane
Vesicle
0.5 µm
Pinocytosis vesicles
forming (arrows) in
a cell lining a small
blood vessel (TEM)
RECEPTOR-MEDIATED ENDOCYTOSIS
Receptor
Coat protein
Coated
vesicle
Coated
pit
Ligand
Coat
protein
Plasma
membrane
A coated pit
and a coated
vesicle formed
during
receptor-
mediated
endocytosis
(TEMs)
0.25 µm

Fig. 7-20a
PHAGOCYTOSIS
CYTOPLASM EXTRACELLULAR
FLUID
Pseudopodium
“Food” or
other particle
Food
vacuole
Food vacuole
Bacterium
An amoeba engulfing a bacterium
via phagocytosis (TEM)
Pseudopodium
of amoeba
1 µm

•In pinocytosis, molecules are taken up when
extracellular fluid is “gulped” into tiny vesicles
Animation: PinocytosisAnimation: Pinocytosis
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-20b
PINOCYTOSIS
Plasma
membrane
Vesicle
0.5 µm
Pinocytosis vesicles
forming (arrows) in
a cell lining a small
blood vessel (TEM)

•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
Animation: Receptor-Mediated EndocytosisAnimation: Receptor-Mediated Endocytosis
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Fig. 7-20c
RECEPTOR-MEDIATED ENDOCYTOSIS
Receptor
Coat protein
Coated
pit
Ligand
Coat
protein
Plasma
membrane
0.25 µm
Coated
vesicle
A coated pit
and a coated
vesicle formed
during
receptor-
mediated
endocytosis
(TEMs)

Fig. 7-UN1
Passive transport:
Facilitated diffusion
Channel
protein
Carrier
protein

Fig. 7-UN2
Active transport:
ATP

Fig. 7-UN3
Environment:
0.01 M sucrose
0.01 M glucose
0.01 M fructose
“Cell”
0.03 M sucrose
0.02 M glucose

Fig. 7-UN4

You should now be able to:
1.Define the following terms: amphipathic
molecules, aquaporins, diffusion
2.Explain how membrane fluidity is influenced
by temperature and membrane composition
3.Distinguish between the following pairs or
sets of terms: peripheral and integral
membrane proteins; channel and carrier
proteins; osmosis, facilitated diffusion, and
active transport; hypertonic, hypotonic, and
isotonic solutions
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

4.Explain how transport proteins facilitate
diffusion
5.Explain how an electrogenic pump creates
voltage across a membrane, and name two
electrogenic pumps
6.Explain how large molecules are transported
across a cell membrane
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings