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Membrane Structure and Function презентация
Содержание
- 1. Membrane Structure and Function
- 2. Overview: Life at the Edge The plasma
- 3. Fig. 7-1
- 4. Concept 7.1: Cellular membranes are fluid mosaics
- 5. Membrane Models: Scientific Inquiry Membranes have been
- 6. Fig. 7-2 Hydrophilic head WATER Hydrophobic tail WATER
- 7. In 1935, Hugh Davson and James Danielli
- 8. Fig. 7-3 Phospholipid bilayer Hydrophobic regions of protein Hydrophilic regions of protein
- 9. Freeze-fracture studies of the plasma membrane supported
- 10. Fig. 7-4 TECHNIQUE Extracellular layer
- 11. The Fluidity of Membranes Phospholipids in the
- 12. Fig. 7-5 Lateral movement (~107 times per
- 13. Fig. 7-5a (a) Movement of phospholipids Lateral
- 14. Fig. 7-6 RESULTS Membrane proteins Mouse
- 15. As temperatures cool, membranes switch from a
- 16. Fig. 7-5b (b) Membrane fluidity Fluid Unsaturated
- 17. The steroid cholesterol has different effects on
- 18. Fig. 7-5c Cholesterol (c) Cholesterol within the
- 19. Membrane Proteins and Their Functions A membrane
- 20. Fig. 7-7 Fibers of extracellular matrix (ECM)
- 21. Peripheral proteins are bound to the surface
- 22. Fig. 7-8 N-terminus C-terminus α Helix CYTOPLASMIC SIDE EXTRACELLULAR SIDE
- 23. Six major functions of membrane proteins: Transport
- 24. Fig. 7-9 (a) Transport ATP (b) Enzymatic
- 25. Fig. 7-9ac (a) Transport (b) Enzymatic activity
- 26. Fig. 7-9df (d) Cell-cell recognition Glyco- protein
- 27. The Role of Membrane Carbohydrates in Cell-Cell
- 28. Synthesis and Sidedness of Membranes Membranes have
- 29. Fig. 7-10 ER 1 Transmembrane glycoproteins
- 30. Concept 7.2: Membrane structure results in selective
- 31. The Permeability of the Lipid Bilayer Hydrophobic
- 32. Transport Proteins Transport proteins allow passage of
- 33. Other transport proteins, called carrier proteins, bind
- 34. Concept 7.3: Passive transport is diffusion of
- 35. Fig. 7-11 Molecules of dye Membrane (cross
- 36. Molecules of dye Fig. 7-11a Membrane (cross
- 37. Substances diffuse down their concentration gradient, the
- 38. (b) Diffusion of two solutes Fig. 7-11b
- 39. Effects of Osmosis on Water Balance Osmosis
- 40. Lower concentration of solute (sugar) Fig. 7-12
- 41. Water Balance of Cells Without Walls Tonicity
- 42. Fig. 7-13 Hypotonic solution (a) Animal
- 43. Hypertonic or hypotonic environments create osmotic problems
- 44. Fig. 7-14 Filling vacuole 50 µm
- 45. Water Balance of Cells with Walls Cell
- 46. Video: Plasmolysis Video: Turgid Elodea Animation: Osmosis
- 47. Facilitated Diffusion: Passive Transport Aided by Proteins
- 48. Fig. 7-15 EXTRACELLULAR FLUID Channel protein
- 49. Carrier proteins undergo a subtle change in
- 50. Some diseases are caused by malfunctions in
- 51. Concept 7.4: Active transport uses energy to
- 52. The Need for Energy in Active Transport
- 53. Active transport allows cells to maintain concentration
- 54. Fig. 7-16-1 EXTRACELLULAR FLUID [Na+]
- 55. Na+ binding stimulates phosphorylation
- 56. Fig. 7-16-3 Phosphorylation
- 57. Fig. 7-16-4 K+
- 58. Fig. 7-16-5 Loss
- 59. Fig. 7-16-6 K+
- 60. 2 EXTRACELLULAR FLUID [Na+]
- 61. Fig. 7-17 Passive transport
- 62. How Ion Pumps Maintain Membrane Potential Membrane
- 63. Two combined forces, collectively called the electrochemical
- 64. An electrogenic pump is a transport protein
- 65. Fig. 7-18 EXTRACELLULAR FLUID H+
- 66. Cotransport: Coupled Transport by a Membrane Protein
- 67. Fig. 7-19 Proton pump –
- 68. Concept 7.5: Bulk transport across the plasma
- 69. Exocytosis In exocytosis, transport vesicles migrate to
- 70. Endocytosis In endocytosis, the cell takes in
- 71. In phagocytosis a cell engulfs a particle
- 72. Fig. 7-20 PHAGOCYTOSIS EXTRACELLULAR FLUID
- 73. Fig. 7-20a PHAGOCYTOSIS CYTOPLASM EXTRACELLULAR
- 74. In pinocytosis, molecules are taken up when
- 75. Fig. 7-20b PINOCYTOSIS Plasma membrane
- 76. In receptor-mediated endocytosis, binding of
- 77. Fig. 7-20c RECEPTOR-MEDIATED ENDOCYTOSIS Receptor
- 78. Fig. 7-UN1 Passive transport: Facilitated diffusion Channel protein Carrier protein
- 79. Fig. 7-UN2 Active transport: ATP
- 80. Fig. 7-UN3 Environment: 0.01 M sucrose 0.01
- 81. Fig. 7-UN4
- 82. You should now be able to: Define
- 83. Explain how transport proteins facilitate diffusion Explain
Слайд 2Overview: Life at the Edge
The plasma membrane is the boundary that
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
Слайд 4Concept 7.1: Cellular membranes are fluid mosaics of lipids and proteins
Phospholipids
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
Слайд 5Membrane Models: Scientific Inquiry
Membranes have been chemically analyzed and found to
Scientists studying the plasma membrane reasoned that it must be a phospholipid bilayer
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 7In 1935, Hugh Davson and James Danielli proposed a sandwich model
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
Слайд 9Freeze-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
Слайд 10
Fig. 7-4
TECHNIQUE
Extracellular
layer
Knife
Proteins
Inside of extracellular layer
RESULTS
Inside of cytoplasmic layer
Cytoplasmic layer
Plasma membrane
Слайд 11The Fluidity of Membranes
Phospholipids in the plasma membrane can move within
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
Слайд 12Fig. 7-5
Lateral movement
(~107 times per second)
Flip-flop
(~ once per month)
(a) Movement of
(b) Membrane fluidity
Fluid
Viscous
Unsaturated hydrocarbon
tails with kinks
Saturated hydro-
carbon tails
(c) Cholesterol within the animal cell membrane
Cholesterol
Слайд 13Fig. 7-5a
(a) Movement of phospholipids
Lateral movement
(~107 times per second)
Flip-flop
(~ once per
Слайд 15As temperatures cool, membranes switch from a fluid state to a
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
Слайд 16Fig. 7-5b
(b) Membrane fluidity
Fluid
Unsaturated hydrocarbon
tails with kinks
Viscous
Saturated hydro-
carbon tails
Слайд 17The steroid cholesterol has different effects on membrane fluidity at different
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
Слайд 19Membrane Proteins and Their Functions
A membrane is a collage of different
Proteins determine most of the membrane’s specific functions
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 20Fig. 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
Слайд 21Peripheral proteins are bound to the surface of the membrane
Integral proteins
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
Слайд 23Six major functions of membrane proteins:
Transport
Enzymatic activity
Signal transduction
Cell-cell recognition
Intercellular joining
Attachment to
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 24Fig. 7-9
(a) Transport
ATP
(b) Enzymatic activity
Enzymes
(c) Signal transduction
Signal transduction
Signaling molecule
Receptor
(d) Cell-cell recognition
Glyco-
protein
(e)
(f) Attachment to
the cytoskeleton
and extracellular
matrix (ECM)
Слайд 25Fig. 7-9ac
(a) Transport
(b) Enzymatic activity
(c) Signal transduction
ATP
Enzymes
Signal transduction
Signaling molecule
Receptor
Слайд 26Fig. 7-9df
(d) Cell-cell recognition
Glyco-
protein
(e) Intercellular joining
(f) Attachment to
the
and extracellular
matrix (ECM)
Слайд 27The Role of Membrane Carbohydrates in Cell-Cell Recognition
Cells recognize each other
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
Слайд 28Synthesis and Sidedness of Membranes
Membranes have distinct inside and outside faces
The
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 29Fig. 7-10
ER
1
Transmembrane
glycoproteins
Secretory
protein
Glycolipid
2
Golgi
apparatus
Vesicle
3
4
Secreted
protein
Transmembrane
glycoprotein
Plasma membrane:
Cytoplasmic face
Extracellular face
Membrane glycolipid
Слайд 30Concept 7.2: Membrane structure results in selective permeability
A cell must exchange
Plasma membranes are selectively permeable, regulating the cell’s molecular traffic
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 31The Permeability of the Lipid Bilayer
Hydrophobic (nonpolar) molecules, such as hydrocarbons,
Polar molecules, such as sugars, do not cross the membrane easily
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 32Transport Proteins
Transport proteins allow passage of hydrophilic substances across the membrane
Some
Channel proteins called aquaporins facilitate the passage of water
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 33Other transport proteins, called carrier proteins, bind to molecules and change
A transport protein is specific for the substance it moves
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 34Concept 7.3: Passive transport is diffusion of a substance across a
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 Selectivity
Animation: Diffusion
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 35Fig. 7-11
Molecules of dye
Membrane (cross section)
WATER
Net diffusion
Net diffusion
Equilibrium
(a) Diffusion of one
Net diffusion
Net diffusion
Net diffusion
Net diffusion
Equilibrium
Equilibrium
(b) Diffusion of two solutes
Слайд 36Molecules of dye
Fig. 7-11a
Membrane (cross section)
WATER
Net diffusion
Net diffusion
(a) Diffusion of one
Equilibrium
Слайд 37Substances diffuse down their concentration gradient, the difference in concentration of
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
Слайд 38(b) Diffusion of two solutes
Fig. 7-11b
Net diffusion
Net diffusion
Net diffusion
Net diffusion
Equilibrium
Equilibrium
Слайд 39Effects of Osmosis on Water Balance
Osmosis is the diffusion of water
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
Слайд 40Lower
concentration
of solute (sugar)
Fig. 7-12
H2O
Higher concentration
of sugar
Selectively
permeable
membrane
Same concentration
of sugar
Osmosis
Слайд 41Water Balance of Cells Without Walls
Tonicity is the ability of a
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
Слайд 42Fig. 7-13
Hypotonic solution
(a) Animal
cell
(b) Plant
cell
H2O
Lysed
H2O
Turgid
H2O
H2O
H2O
H2O
Normal
Isotonic solution
Flaccid
H2O
H2O
Shriveled
Plasmolyzed
Hypertonic solution
Слайд 43Hypertonic or hypotonic environments create osmotic problems for organisms
Osmoregulation, the control
The protist Paramecium, which is hypertonic to its pond water environment, has a contractile vacuole that acts as a pump
Video: Chlamydomonas
Video: Paramecium Vacuole
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 44Fig. 7-14
Filling vacuole
50 µm
(a) A contractile vacuole fills with fluid
a system of canals radiating throughout the cytoplasm.
Contracting vacuole
(b) When full, the vacuole and canals contract, expelling
fluid from the cell.
Слайд 45Water Balance of Cells with Walls
Cell walls help maintain water balance
A
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
Слайд 46Video: Plasmolysis
Video: Turgid Elodea
Animation: Osmosis
In a hypertonic environment, plant cells lose
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 47Facilitated Diffusion: Passive Transport Aided by Proteins
In facilitated diffusion, transport proteins
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
Слайд 48Fig. 7-15
EXTRACELLULAR FLUID
Channel protein
(a) A channel protein
Solute
CYTOPLASM
Solute
Carrier protein
(b) A carrier protein
Слайд 49Carrier proteins undergo a subtle change in shape that translocates the
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 50Some diseases are caused by malfunctions in specific transport systems, for
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 51Concept 7.4: Active transport uses energy to move solutes against their
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
Слайд 52The Need for Energy in Active Transport
Active transport moves substances against
Active transport requires energy, usually in the form of ATP
Active transport is performed by specific proteins embedded in the membranes
Animation: Active Transport
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 53Active transport allows cells to maintain concentration gradients that differ from
The sodium-potassium pump is one type of active transport system
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 54
Fig. 7-16-1
EXTRACELLULAR
FLUID
[Na+] high
[K+] low
Na+
Na+
Na+
[Na+] low
[K+]
CYTOPLASM
Cytoplasmic Na+ binds to
the sodium-potassium pump.
1
Слайд 56
Fig. 7-16-3
Phosphorylation causes
the protein to change its
shape. Na+
the outside.
Na+
P
Na+
Na+
3
Слайд 57
Fig. 7-16-4
K+ binds on the
extracellular side and
triggers release
phosphate group.
P
P
K+
K+
4
Слайд 60
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
Слайд 62How Ion Pumps Maintain Membrane Potential
Membrane potential is the voltage difference
Voltage is created by differences in the distribution of positive and negative ions
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 63Two combined forces, collectively called the electrochemical gradient, drive the diffusion
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
Слайд 64An electrogenic pump is a transport protein that generates voltage across
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
Слайд 65Fig. 7-18
EXTRACELLULAR
FLUID
H+
H+
H+
H+
Proton pump
+
+
+
H+
H+
+
+
H+
–
–
–
–
ATP
CYTOPLASM
–
Слайд 66Cotransport: Coupled Transport by a Membrane Protein
Cotransport occurs when active transport
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
Слайд 67Fig. 7-19
Proton pump
–
–
–
–
–
–
+
+
+
+
+
+
ATP
H+
H+
H+
H+
H+
H+
H+
H+
Diffusion
of H+
Sucrose-H+
cotransporter
Sucrose
Sucrose
Слайд 68Concept 7.5: Bulk transport across the plasma membrane occurs by exocytosis
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
Слайд 69Exocytosis
In exocytosis, transport vesicles migrate to the membrane, fuse with it,
Many secretory cells use exocytosis to export their products
Animation: Exocytosis
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 70Endocytosis
In endocytosis, the cell takes in macromolecules by forming vesicles from
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 Introduction
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 71In phagocytosis a cell engulfs a particle in a vacuole
The vacuole
Animation: Phagocytosis
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 72Fig. 7-20
PHAGOCYTOSIS
EXTRACELLULAR
FLUID
CYTOPLASM
Pseudopodium
“Food”or
other particle
Food
vacuole
PINOCYTOSIS
1 µm
Pseudopodium
of
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
Слайд 73Fig. 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
Слайд 74In pinocytosis, molecules are taken up when extracellular fluid is “gulped”
Animation: Pinocytosis
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 75Fig. 7-20b
PINOCYTOSIS
Plasma
membrane
Vesicle
0.5 µm
Pinocytosis vesicles
forming (arrows) in
a cell
blood vessel (TEM)
Слайд 76
In receptor-mediated endocytosis, binding of ligands to receptors triggers vesicle
A ligand is any molecule that binds specifically to a receptor site of another molecule
Animation: Receptor-Mediated Endocytosis
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Слайд 77Fig. 7-20c
RECEPTOR-MEDIATED ENDOCYTOSIS
Receptor
Coat protein
Coated
pit
Ligand
Coat
protein
Plasma
membrane
0.25 µm
Coated
vesicle
A coated pit
and
vesicle formed
during
receptor-
mediated
endocytosis
(TEMs)
Слайд 80Fig. 7-UN3
Environment:
0.01 M sucrose
0.01 M glucose
0.01 M fructose
“Cell”
0.03 M
0.02 M glucose
Слайд 82You should now be able to:
Define the following terms: amphipathic molecules,
Explain how membrane fluidity is influenced by temperature and membrane composition
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
Слайд 83Explain how transport proteins facilitate diffusion
Explain how an electrogenic pump creates
Explain how large molecules are transported across a cell membrane
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
