Chapter 19 PDF

MOLECULAR BIOLOGY OF THE CELL: SIXTH EDITION
CHAPTER 19: CELL JUNCTIONS AND THE

EXTRACELLULAR MATRIX
© Garland Science 2015
1. Indicate whether each of the following descriptions better applies to connective (C) or
epithelial (E) tissues. Your answer would be a four-letter string composed of letters C and E
only, e.g. EEEE.
( ) Cells are usually distributed sparsely in the extracellular matrix.
( ) Gap junctions are rarely found.
( ) Direct cell–cell attachments are common.
( ) Cells are tightly associated into sheets.
2. In the following schematic diagram of a simple columnar epithelium lining the digestive
tract, indicate which position (A to F) along the basal–apical axis better corresponds to each of
the following features. Your answer would be a six-letter string composed of letters A to F only,
e.g. FEDABC.
A B C D E F
Gut Connective
lumen tissue
( ) Basal lamina
( ) Cell apex
( ) Adherens junctions
( ) Gap junctions
( ) Hemidesmosomes
( ) Tight junctions
3. Which of the following cell junctions uses cadherin cell adhesion molecules to anchor the
actin cytoskeleton?
A. Tight junction
B. Adherens junction
C. Desmosome
D. Hemidesmosome
E. Gap junction
4. Which of the following is NOT an anchoring junction?

A. Adherens junction
B. Desmosome
C. Actin-linked cell–matrix junction
D. Hemidesmosome
E. Tight junction
5. Indicate true (T) and false (F) statements below regarding cadherins. Your answer would
be a four-letter string composed of letters T and F only, e.g. TFTT.
( ) Cadherins are present in all multicellular organisms.
( ) Adhesion by cadherins is dependent on Ca2+ ions.
( ) In contrast to nonclassical cadherins, classical cadherins are more closely related in
sequence.
( ) E, N, and P cadherins are mainly found in desmosomes.
6. You have isolated epithelial cells from mouse mammary glands and have grown them in
a three-dimensional gelatinous matrix in the presence or absence of a specific inhibitor that
targets a transmembrane metalloprotease. The protease is capable of cleaving cadherin near the
base of its extracellular domain to release an N-terminal fragment containing the extracellular
cadherin domains. The distribution of the cells in the matrix, as seen under a light microscope,
one day after the transplantation of the isolated tissue is presented in the following schematic
drawings. Which condition (1 or 2) do you think corresponds to the presence of the protease
inhibitor? Write down 1 or 2 as your answer.
1 2
7. Cadherin molecules at the cell surface are often clustered side-to-side to create a
molecular Velcro that attaches the cell to another cell or to the extracellular matrix. In the
following simplified drawings, two cells are initially attached via such a Velcro. When one of the
cells moves in either of three different ways (A to C), it is faced with a mechanical resistance
from the cadherin interactions. The resistance against which type of movement do you think is
the weakest? Write down A, B, or C as your answer.
Two A B C
neighboring The cell moves The cell slides The cell rolls
cells away from its past its over its
neighbor neighbor neighbor
8. Cells expressing either N-cadherin (A), high levels of E-cadherin (B), or low levels of E-
cadherin (C) have sorted themselves out on a substratum in a cadherin-dependent manner as
shown in the schematic drawing below. Which cells (A to C) would you expect to correspond to
white, gray, and black in the drawing, respectively? Your answer would be a three-letter string
composed of letters A to C only, e.g. CAB.
9. Overproduction of cadherins such as E-cadherin …
A. is often found in cancers originating from epithelia.
B. is induced by the transcription regulatory protein Twist.
C. induces epithelial–mesenchymal transition.
D. leads to stronger cell–cell adhesion.
E. All of the above.
10. Consider two cells attached to each other via adherens junctions. The forces generated by
the actin cytoskeleton in one cell …
A. are dependent on myosin activity.
B. are normally balanced by similar forces in the neighboring cell.
C. can strengthen the adherens junction by unfolding catenin proteins.
D. All of the above.
11. Consider a sheet of epithelial tissue that invaginates to form a tube along the anterior–
posterior axis of a developing vertebrate embryo. The tube is then closed concomitant with a
“convergence and extension” process similar to that observed in germ-band extension in
Drosophila melanogaster. During this process, the tube elongates and narrows before pinching
off from the sheet. Which axis (1 or 2) in the following schematic drawing of the tube cells do
you think corresponds to the anterior–posterior axis? Write down 1 or 2 as your answer.
1
2
Cells of the epithelial tube
12. Fill in the blank in the following paragraph. Do not use abbreviations.
“Particularly abundant in cells of heart muscle and the epidermis,
… are structurally similar to adherens junctions and contain a
number of homologous components. However, their cadherin
molecules are linked to intermediate filaments instead of the actin
cytoskeleton.”
13. A researcher has grown monolayers of cells from either of two epithelial cell lines on a
permeable supporting membrane that separates two electrolyte chambers. One of the cell lines
(A) has been derived from the mammalian kidney and forms epithelial sheets resembling those
lining the renal collecting duct and the urinary bladder. The other (B) is derived from the
mammalian small intestine. The researcher has measured the voltage (V) and electrical current
(I) across the monolayer and has plotted the results in the graph below. Which line (1 or 2) in the
graph do you think corresponds to cell line A? Recall that current (I) is related to the applied
voltage (V) by the equation I = V/R, in which R is the electrical resistance. Write down 1 or 2 as
your answer.
100-
V (millivolts)
2
0 -
0 150
I (microamperes)
14. Indicate whether each of the following descriptions better applies to adherens junctions
(A), desmosomes (D), gap junctions (G), or tight junctions (T). Your answer would be a four-
letter string composed of letters A, D, G, and T only, e.g. AAAD.
( ) They play a direct role in structural polarization of epithelial cells.
( ) They couple cells both electrically and metabolically.
( ) They are formed from hexameric hemichannels that can assemble into heterotypic
complexes.
( ) They create semipermeable paracellular pores.
15. In the following schematic drawing of connexons as seen from the cytosol of a cell, what
is the approximate diameter of each of the pores? Which of the indicated connexons (1 or 2) is
more likely to have been incorporated more recently?
1
2
Pore
A. About 20 nm; connexon 1

B. About 4 nm; connexon 1
C. About 1.5 nm; connexon 1
D. About 4 nm; connexon 2
E. About 1.5 nm; connexon 2
16. Plasmodesmata in plant cells are functionally similar to animal cells’ …

A. adherens junctions.
B. desmosomes.
C. gap junctions.
D. hemidesmosomes.
E. tight junctions.
17. In insects such as Drosophila melanogaster, electrical synapses are abundant in the
ventral nerve cord. Unlike a chemical synapse, the cytosols of pre- and postsynaptic cells in an
electrical synapse are connected, allowing action potentials to spread rapidly and without delay.
What junctional proteins in the flies are chiefly responsible for this unique feature of the
electrical synapses?
A. Cadherins
B. Claudins
C. Connexins
D. Innexins
E. Integrins
18. Selectins …
A. mainly mediate cell–matrix attachments.
B. are members of the Ig superfamily of cell adhesion molecules.
C. do NOT require Ca2+ for their adhesive function, unlike integrins.
D. are carbohydrate-binding proteins.
19. Malignant cancer cells that have entered the bloodstream can imitate lymphocytes in
exiting the bloodstream and entering underlying tissues in the process of metastasis. This
“lymphocyte mimicry” can be summarized as …
A. an initial cadherin-dependent rolling followed by an integrin-dependent adhesion and
emigration.
B. an initial cadherin-dependent rolling followed by an I-CAM-dependent adhesion and
emigration.
C. an initial cadherin-dependent rolling followed by a selectin-dependent adhesion and
emigration.
D. an initial selectin-dependent rolling followed by a cadherin-dependent adhesion and
emigration.
E. an initial selectin-dependent rolling followed by an I-CAM-dependent adhesion and
emigration.
20. In the following schematic diagram, the estimated attractive or repulsive force between
two adhesive cells is plotted as a function of distance between the cells. The dashed curve
represents a condition under which the cells express cadherin but not N-CAM. The other two
curves represent conditions under which both cadherin and heavily sialylated N-CAM are
expressed. In one of the conditions, however, the ionic strength of the medium has been
artificially increased by addition of salt. Which curve (1 or 2) do you think represents the high-
salt condition? Write down 1 or 2 as your answer.
Repulsion
2 1
Force
0
Attraction
0
Intercellular distance
21. Indicate whether each of the following descriptions better applies to glycosaminoglycans
(G), fibrous proteins (F), or glycoproteins (P) of the extracellular matrix in animal cells. Your
answer would be a six-letter string composed of letters G, F, and P only, e.g. GFPGFP.
( ) They include laminin and fibronectin.
( ) They primarily include collagens.
( ) They include hyaluronan.
( ) They are found in proteoglycans.
( ) They occupy large volumes and form hydrated gels.
( ) They constitute the major proteins of the matrix.
22. Indicate true (T) and false (F) statements below regarding glycosaminoglycan chains in
the extracellular matrix. Your answer would be a four-letter string composed of letters T and F
only, e.g. TFTT.
( ) They are highly branched polysaccharides.
( ) They are strongly hydrophilic and absorb a large amount of water.
( ) They all contain N-acetylglucosamine or N-acetylgalactosamine.
( ) They are highly positively charged.
23. Fill in the blank in the following paragraph regarding the extracellular matrix. Do not use
abbreviations.
“The extracellular matrix of connective tissues in animals is
primarily made and secreted by the … family of cells, although
they may have more specific names in certain tissues. Being
common in connective tissues, these cells can also migrate and
proliferate if need be.”
24. How is hyaluronic acid different from other glycosaminoglycans of the extracellular
matrix?
A. It lacks sulfate groups.
B. It is generally not covalently linked to proteins.
C. It is not assembled in the Golgi apparatus.
D. It can be several megadaltons in mass.
25. In the following schematic drawings of aggrecan and decorin, which one (1 or 2)
corresponds to aggrecan? Which one (1 or 2) is synthesized primarily by chondrocytes in
cartilage?
2
1
A. 1; 1
B. 1; 2
C. 2; 1
D. 2; 2
26. A giant aggrecan aggregate can have a molecular mass of 108 daltons or more, and can be
as large as a bacterium. Its structure resembles a brush. What is responsible for attaching the
bristles of the brush to its core?
A. Covalent bonds between aggrecan core protein and keratan or chondroitin sulfate.
B. Noncovalent bonds between aggrecan core protein and keratan or chondroitin sulfate.
C. Covalent bonds between aggrecan core protein and hyaluronan.
D. Noncovalent bonds between aggrecan core protein and hyaluronan.
27. Which of the following extracellular matrix components contains a transmembrane
protein?
A. Type I collagen
B. Perlecan
C. Aggrecan
D. Syndecan
E. Decorin
28. Collagens are extremely rich in …

A. alanine and valine.
B. glycine and proline.
C. lysine and arginine.
D. serine and threonine.
E. tyrosine and phenylalanine.
29. Type IV collagen molecules can interact at their C-termini to form dimers and at their N-
termini to form tetramers. What category do you think this type of collagen belongs to?
A. Fibrillar
B. Fibril-associated
C. Network-forming
D. Proteoglycan core
E. Anchorage fibril
30. Which collagens are the most common in our body?

A. Fibrillar
B. Fibril-associated
C. Network-forming
D. Proteoglycan core
E. Anchorage fibril
31. Which components of the extracellular matrix are mostly responsible for its ability to
resist compressive and tensile forces, respectively?
A. Collagens and glycoproteins
B. Glycoproteins and collagens
C. Glycosaminoglycans and glycoproteins
D. Glycosaminoglycans and collagens
E. Collagens and glycosaminoglycans
32. Which of the following is true regarding fibril-associated collagens such as type IX
collagen?
A. They aggregate with one another to form long fibrils in the extracellular space.
B. They lack the triple-stranded helical structure found in other collagens.
C. They are more flexible than collagens of types I to IV.
D. They are the most common among collagens.
33. The size distribution of exons coding for collagen helical domains in various vertebrates
reveals a unique pattern. This pattern provides clues to the evolutionary history of the genes
encoding fibrillar collagen α chains. What is this pattern?
A. The length of most exons is a multiple of 100 nucleotides.
B. Most exons code for 54, or a multiple of 54, Gly-X-Y repeats.
C. The length of most exons is a multiple of 30 nucleotides.
D. Most exons code for 6, or a multiple of 6, Gly-X-Y repeats.
E. None of the above.
34. The elasticity of the components of the extracellular matrix can be visualized in a stress–
strain curve, in which the tensile stress (corresponding to load) applied to a tissue sample is
plotted against the strain (corresponding to deformation) measured in the tissue. In the following
stress–strain graph, the elasticity of an intact connective tissue is represented by a dashed curve.
The other two curves correspond to the same tissue that has been treated with either formic acid
(to digest collagen) or trypsin (to digest elastin). Which curve (1 or 2) do you think corresponds
to trypsin treatment?
1
Stress (Pa) 2
Strain (%)
35. This large glycoprotein of the extracellular matrix is associated with elastin fibers and is
essential for their integrity. Its deposition in the developing connective tissues often precedes the
appearance of elastin. It is part of microfibrils that cover the elastic fibers. This protein is …
A. collagen.
B. fibronectin.
C. fibrillin.
D. filamin.
E. laminin.
36. Multidomain glycoproteins in the extracellular matrix have binding sites for many other
macromolecules. These glycoproteins …
A. can serve as tracks along which cells can migrate.
B. can bind to growth factors and influence cell signaling.
C. can bind to cell-surface receptors.
D. can serve as repellents to forbid cell migration.
37. Indicate true (T) and false (F) statements below regarding fibronectin in the extracellular
matrix. Your answer would be a four-letter string composed of letters T and F only, e.g. TFTT.
( ) Fibronectin carries asparagine-linked oligosaccharides.
( ) Fibronectin often forms homodimers by cross-linking through covalent attachment of
lysine residues.
( ) Fibronectin binds to integrins, collagens, and glycosaminoglycans through its type III
repeats.
( ) Fibronectin molecules can assemble into fibrils only in the vicinity of cells.
38. Neuromuscular junctions of vertebrates are special types of chemical synapses formed
from close association between motor neuron axon terminals and muscle fibers. A thin layer of
basal lamina separates the two cell membranes at the neuromuscular junction and is important
for its organization and maintenance. Which of the following would you NOT expect to find in
the basal lamina in these structures?
A. Nidogen
B. Laminin
C. Type I collagen
D. Type IV collagen
E. Perlecan
39. Indicate true (T) and false (F) statements below regarding the interaction of cells with the
extracellular matrix using matrix receptors. Your answer would be a four-letter string composed
of letters T and F only, e.g. TFTT.
( ) Integrins are the only known matrix receptors in animal cells.
( ) Integrins can transmit signals in both directions across the membrane; that is, both
outside-in and inside-out.
( ) Tension can increase the binding affinity of an integrin for its intracellular and
extracellular ligands.
( ) Integrins can convert molecular signals into mechanical ones.
40. Indicate whether each of the following descriptions better applies to the intracellular (I)
or extracellular (E) domain of integrins. Your answer would be a four-letter string composed of
letters I and E only, e.g. EEEE.
( ) It binds to talin.
( ) It binds to the RGD tripeptide motif.
( ) It is bulkier.
( ) It contains divalent cation-binding domains.
41. A researcher has coated the wells in a multiwell plate with human plasma fibronectin.
She introduces integrin-expressing cells into these wells in the presence of different
concentrations of a synthetic peptide. She then washes the wells to remove unbound cells and
counts the number of cells that remain attached to each well. The results of her experiment are
presented in the following schematic graph. The synthetic peptide sequences used in these
experiments were GRGDSPC (1) or GGDRSLN (2). Which one of these peptides (1 or 2) would
you expect to correspond to curve a in the graph? Which curve (a or b) would you expect if a
synthetic peptide with the sequence GDGRSPC was used? C is cysteine, D is aspartic acid, G is
glycine, L is leucine, N is asparagine, P is proline, R is arginine, and S is serine.
a
Number of remaining cells
Peptide concentration used
A. 1; a
B. 1; b
C. 2; a
D. 2; b
42. EDTA is a metal-ion “chelator” that is commonly used to lower the concentration of free
divalent cations in solution. In the lab, your friend has cultured a human cell line on elastic
membranes that have been coated with fibronectin and attached to a device that can stretch the
membrane. She has quantified the binding of talin to integrin under different conditions, and has
presented the results in the following schematic graph, which shows the binding in the presence
or absence of EDTA in the cell-culture medium. Which experiment (A or B) do you think was
performed in the presence of EDTA? Write down A or B as your answer.
100 - Stretched
No stretch
Bound integrin (%)
0 -
A B
43. The following schematic drawing shows two integrin molecules. Which molecule (a or b)
is depicted in its active configuration? Which side of the plasma membrane (1 or 2) represents
the cytosol?
a b
Plasma membrane
α β
β
α
A. a; 1
B. a; 2
C. b; 1
D. b; 2
44. In the active state of the integrin dimer, …

A. both intracellular and extracellular binding sites are exposed.
B. the intracellular binding sites are inaccessible, while the extracellular binding sites are
exposed.
C. the intracellular binding sites are exposed, while the extracellular binding sites are
inaccessible.
D. both intracellular and extracellular binding sites are inaccessible.
45. Platelets that express a constitutively active Rap1 are able to activate integrins even in the
absence of thrombin stimulation. In contrast, cells expressing a binding-deficient version of talin
are unable to perform integrin activation. In platelets expressing both of these mutant proteins,
what would you expect to observe: the Rap1 gain-of-function phenotype (R) or the talin loss-of-
function phenotype (T), as described above? Write down R or T as your answer.
46. How does talin activate the inside-out signaling through integrin?
A. It binds to both integrin subunits to bring them together in an interlocked
conformation.
B. It is a protein kinase that phosphorylates both integrin subunits and induces activation
by conformational change.
C. It competes with the α subunit for binding to the β subunit, therefore blocking
intimate α-β linkage.
D. It binds to the α subunit, and dissociates the dimer into individual subunits that are
functionally active.
47. Which of the following molecules is capable of tension-sensing by exposing binding sites
for other proteins when mechanically stretched?
A. α-Catenin
B. Talin
C. Fibronectin
D. All of the above
48. The chemical compound PMA is a potent stimulator of the epithelial–mesenchymal

transition in epithelial cells. The activity of two of the following types of proteins is typically up-
regulated in epithelial cells treated with PMA, while that of the other two is mainly down-
regulated. Choose the two proteins whose activity you think is stimulated by PMA. Your answer
would be a two-letter string composed of letters A to D only in alphabetical order, e.g. BC.
(A) Cadherin
(B) Connexin
(C) Matrix metalloprotease
(D) Integrin
49. The focal adhesion kinase …

A. is a cytosolic serine/threonine kinase.
B. binds directly to integrin dimers.
C. can be phosphorylated by Src kinases.
D. prevents the relay of outside-in integrin signaling.
50. Indicate true (T) and false (F) statements below regarding integrin signaling. Your
answer would be a four-letter string composed of letters T and F only, e.g. TFTT.
( ) Integrins usually bind their ligands with a higher affinity compared to other cell-
surface receptors.
( ) Unlike cadherins, integrins usually function as individual dimers and do not cluster.
( ) Anchorage dependence is mainly mediated via integrin signaling.
( ) Integrins can activate the Ras–MAPK pathway.
51. A cell’s response is not the same when placed on a rigid compared to a soft matrix.
Indicate whether each of the following is expected to occur in a cell that is placed on a rigid
matrix (R) or a soft matrix (S). Your answer would be a three-letter string composed of letters R
and S only, e.g. RSR.
( ) More tension is generated in cell–matrix contacts.
( ) More vinculin proteins would bind to the C-terminal tail of talin.
( ) More actin filaments are recruited to the site of cell–matrix adhesion.
52. Indicate whether each of the following descriptions better applies to the plant cell wall
(W) or the extracellular matrix of animal cells (M). Your answer would be a three-letter string
composed of letters W and M only, e.g. WMM.
( ) It is generally thicker and stronger.
( ) It is more rigid.
( ) It has higher nitrogen content.
53. What is the most common additional polymer present in the plant secondary, but not
primary, cell wall?
A. Cellulose
B. Pectin
C. Lignin
D. Starch
54. Indicate whether each of the following descriptions better applies to cellulose microfibrils
(C) or pectins (P) in plant cell walls. Your answer would be a five-letter string composed of
letters C and P only, e.g. CPPPP.
( ) It is made into a network by cross-linking glycans.
( ) It is particularly abundant in the middle lamella.
( ) Divalent ions such as calcium can cross-link it to form gels.
( ) It forms more homogeneous polymers.
( ) It contains many negatively charged galacturonic acid units.
55. In the following schematic diagram of a cubic plant cell that has just left the meristem
zone, the cellulose microfibrils are shown as black lines. In which direction (1 or 2) do you think
the cell will elongate when the cell grows? If the cell wall is momentarily dissolved, in what
direction (3 or 4) would water tend to flow?
A. 1; 3
B. 1; 4
C. 2; 3
D. 2; 4
56. Cellulose is deposited onto the plant cell wall in highly ordered crystalline aggregates by
the cellulose synthase complex embedded in the plasma membrane. Would you expect cell wall
deposition to continue following treatment of plant cells with a drug that depolymerizes
microtubules? Would you expect these cells to be able to switch the orientation of the microfibril
pattern between successive lamellae?
A. Yes; yes
B. Yes; no
C. No; yes
D. No; no
Answers:
1. Answer: CCEE
Difficulty: 1
Section: Cell–Cell Junctions
Feedback: Direct cell–cell attachments are rare in connective tissue compared to
epithelial tissue. Cell–matrix attachments are instead dominant in connective tissues.
2. Answer: FACDEB
Difficulty: 2
Feedback: The adhesion belt (C) is usually formed beneath the strands of tight junctions
(B) near the apex (A). Gap junctions (D) are usually more basally located. Finally,
hemidesmosomes (E) link the cytoskeleton to the underlying basal lamina (F) at the basal
surface of the cells.
3. Answer: B
Difficulty: 1
Feedback: In adherens junctions, classical cadherins from two neighboring cells interact
to link the actin cytoskeletons of the cells together. In these structures, actin filaments
bind cadherin indirectly via intracellular adaptor proteins.
4. Answer: E
Difficulty: 1
Feedback: An anchoring junction links the cytoskeleton of a cell to that of its neighbor
(cell–cell junction) or to the extracellular matrix (cell–matrix junction).
5. Answer: FTTF
Difficulty: 1
Feedback: Cadherins form a diverse family of Ca2+-dependent cell adhesion molecules
found in all animals but not in plants and fungi. The more closely related classical
cadherins include the E, N, and P cadherins and are found in adherens junctions.
6. Answer: 1
Difficulty: 2
Feedback: The inhibitor prevents the shedding of cadherin induced by the protease, and
therefore facilitates cell–cell adhesion. This helps the cells clump together.
7. Answer: C
Difficulty: 2
Feedback: According to the Velcro model, clustered cadherin molecules would provide
cell adhesions with great shear strength (B) as well as tensile strength (A), while
simultaneously allowing “peeling” (C) to occur easily.
8. Answer: CAB
Difficulty: 2
Feedback: Cells expressing N-cadherin (gray) sort out from cells expressing E-cadherin.
Cells expressing high levels of E-cadherin (black) sort out from cells expressing low
levels (white), which end up mostly at the periphery because of their weaker adhesion.
9. Answer: D
Difficulty: 1
Feedback: Loss of cadherin function is often found in malignant cancer cells. Twist
negatively regulates cadherin and induces epithelial–mesenchymal transition. A higher
level of cadherin typically leads to stronger cadherin-dependent adhesions.
10. Answer: D
Difficulty: 1
Feedback: Adherens junctions are subjected to pulling forces generated by the attached
contractile bundles of actin filaments and non-muscle myosin II. The protein α-catenin
can be stretched to an extended conformation by this force, recruiting more actin to the
junction to strengthen it. Through such mechanotransduction mechanisms, pulling on a
junction in one cell increases the contractile force generated in the attached cell.
11. Answer: 2
Difficulty: 2
Feedback: The remodeling of cell–cell adhesions proceeds by contraction of actin–
myosin bundles and formation and expansion of new cadherin-based adhesions along the
tube. This process extends the tube and accompanies its closure.
12. Answer: desmosomes
Difficulty: 1
Feedback: Desmosomes provide epithelial cells with mechanical strength and are
plentiful in tissues that are subject to high levels of mechanical stress.
13. Answer: 1
Difficulty: 3
Feedback: The transepithelial electrical resistance is represented by the slopes of the two
lines and is expected to be higher for cell line A and the epithelial cells lining the urinary
bladder. These cells are orders of magnitude less permeable to small ions compared to
intestinal epithelia.
14. Answer: TGGT
Difficulty: 1
Feedback: Tight junctions form a fence between plasma membrane domains in a
polarized cell, contributing to the difference in apical and basolateral surface proteins.
They also form seals between cells in an epithelial sheet to confer particular permeability
properties on the sheet. Gap Junctions, formed from aligned hexameric hemichannels on
the plasma membranes of two neighboring cells, form molecular sieves that that allow the
cells to share small molecules and ions.
15. Answer: C
Difficulty: 1
Feedback: The pore formed by the intercellular channels in gap junctions is about 1.5 nm
in diameter, allowing the diffusion of only small molecules. Gap-junction plaques are
dynamic structures; new channels are added at the periphery and older channels are
removed from near the center of the plaque.
16. Answer: C
Difficulty: 1
Feedback: Like gap junctions, plasmodesmata connect the cytosols of neighboring cells
in tissues, allowing small metabolites and signaling molecules to diffuse between the
cells.
17. Answer: D
Difficulty: 1
Feedback: Gap junctions at electrical synapses allow action potentials to spread without
delay. In invertebrates, they are made of innexin subunits.
18. Answer: D
Difficulty: 1
Feedback: Selectins are lectins that mediate transient, calcium-dependent cell–cell
adhesions.
19. Answer: E
Difficulty: 2
Feedback: The “rolling” step is accompanied by relatively weak selectin-mediated
adhesions and is followed by stronger adhesions as a result of integrin binding to
immunoglobulin (Ig) superfamily proteins such as I-CAMs.
20. Answer: 2
Difficulty: 3
Feedback: The overall force is the sum of cadherin-dependent attractive forces, N-CAM-
dependent electrostatic repulsion, and steric exclusions. In the presence of N-CAM with
long polysialic acid chains, the repulsive forces are dominant: the chains are negatively
charged and occupy a large volume. Increasing the ionic strength weakens the repulsive
forces.
21. Answer: PFGGGF
Difficulty: 1
Section: The Extracellular Matrix of Animals
Feedback: Glycosaminoglycan chains, such as hyaluronan or those covalently linked to
core proteins in proteoglycans, occupy large amounts of space and form hydrated gels
that resist compression. Fibrous proteins, primarily collagens, constitute a significant
fraction of the total protein mass in the extracellular matrix. Noncollagen glycoproteins,
such as fibronectin and laminin, carry conventional N-linked oligosaccharides.
22. Answer: FTTF
Difficulty: 1
Feedback: Glycosaminoglycans (GAGs) are unbranched and highly anionic.
23. Answer: fibroblast
Difficulty: 1
Feedback: In most connective tissues, the extracellular matrix components are secreted
by fibroblasts.
24. Answer: E
Difficulty: 1
Feedback: Being the simplest of the glycosaminoglycans (GAGs), hyaluronan chains can
be remarkably long. Not typical of GAGs, it contains no sulfated sugars, is not generally
linked to proteins covalently, and is synthesized and spun out directly from the cell
surface by a transmembrane enzyme.
25. Answer: D
Difficulty: 1
Feedback: Aggrecan (2) is a major component of cartilage.
26. Answer: D
Difficulty: 1
Feedback: This brushlike structure is an order of magnitude larger than the bottlebrush
structure of the aggrecan proteoglycan itself. Each bristle represents an aggrecan that is
noncovalently attached to a central hyaluronan molecule.
27. Answer: D
Difficulty: 1
Feedback: Syndecans are found at the plasma membrane in many cell types, where they
regulate cell adhesions and signaling.
28. Answer: B
Difficulty: 1
Feedback: Both glycine (small size) and proline (unique backbone angles) are important
in the formation of the triple-stranded helix in collagen.
29. Answer: C
Difficulty: 2
Feedback: This type of interaction creates sheetlike networks of type IV collagens in
basal lamina.
30. Answer: A
Difficulty: 1
Feedback: Type I collagen, which is a fibrillar collagen, is by far the most common
collagen in the body.
31. Answer: D
Difficulty: 1
Feedback: While glycosaminoglycans of the extracellular matrix provide a tissue with
resistance to compression, collagen fibrils form structures that can confer great tensile
strength to the tissue.
32. Answer: C
Difficulty: 1
Feedback: Fibril-associated collagens bind in a periodic manner to fibrils made of
fibrillar collagens (e.g. type I). The presence of nonhelical interruptions within their
triple-stranded helical structure makes these fibril-associated collagens more flexible than
fibrillar collagen molecules.
33. Answer: D
Difficulty: 2
Feedback: The size of the coding exons for collagen helical domains is often 54 or a
multiple of 54 nucleotides. This suggests that duplications of an ancestral gene (that was
54 nucleotides long and coded for six Gly-X-Y repeats) gave rise to the helical domains
in modern fibrillar collagens.
34. Answer: 1
Difficulty: 3
Feedback: The elastic fibers (composed of elastin) give tissues their elasticity. Under
increasing load, these fibers stretch to a greater extent compared to collagen fibers.
Therefore, the stress-strain curve for elastin fibers (2) has a lower slope compared to that
for collagen fibers (1). When elastin fibers are digested by trypsin, the curve for the
tissue approaches that of the trypsin-resistant collagen.
35. Answer: C
Difficulty: 1
Feedback: The elastin core in elastin fibers is covered with a sheath of microfibrils which
are elastic in their own right and seem to provide scaffolding to guide elastin deposition
in developing tissues. Fibrillin is a glycoprotein in microfibrils that is essential for the
integrity of elastic fibers.
36. Answer: E
Difficulty: 1
Feedback: The extracellular matrix contains a large and varied set of multidomain
glycoproteins such as fibronectin. They help organize the matrix and cell–matrix
attachments. They also guide and control cell migration during development. Their
binding to peptide growth factors and small molecules influences cell signaling within a
tissue.
37. Answer: TFFT
Difficulty: 2
Feedback: Fibronectin is a matrix multidomain glycoprotein and can form fibronectin
fibrils on the surface of cells. This assembly is regulated by the tension exerted by the
cells via surface proteins such as integrins. Type III fibronectin repeats are responsible
for integrin binding, whereas binding to collagen and heparin is mediated by type I and
type II repeats. Fibronectin heterodimers are held together by C-terminal disulfide bonds.
38. Answer: C
Difficulty: 2
Feedback: Laminin and type IV collagen are major and ubiquitous components of the
basal lamina. Glycoproteins such as nidogen and proteoglycans such as perlecan are also
commonly found in basal laminae.
39. Answer: FTTT
Difficulty: 1
Section: Cell–Matrix Junctions
Feedback: Integrins are the principal matrix receptors in animal cells. They can transmit
mechanical and molecular signals in both directions across the plasma membrane.
40. Answer: IEEE
Difficulty: 1
Feedback: The large N-terminal extracellular domain of integrins can bind to proteins
containing the RGD or other sequences. The binding is modulated by the concentration of
divalent ions such as Ca2+. The intracellular domain can engage the actin cytoskeleton or
the intermediate filaments by using adaptor proteins such as talin and plectin,
respectively.
41. Answer: C
Difficulty: 3
Feedback: The synthetic peptide containing the RGD sequence (1) competes with
fibronectin for integrin binding. At higher concentrations, its presence results in fewer
cells attaching to the fibronectin on the well surface, as shown in curve B. A sequence
that lacks the RGD motif would be expected to result in curve A.
42. Answer: B
Difficulty: 2
Feedback: Dependence of the binding of integrin on the presence of divalent cations is
reflected in the inhibition of integrin binding in the presence of the chelating agent.
43. Answer: A
Difficulty: 2
Feedback: Integrins have two major activity states; in the active form, the large
extracellular domains are more extended and further apart from each other.
44. Answer: A
Difficulty: 1
Feedback: In the active state of integrins, the two subunits in each dimer are unhooked at
the membrane to expose the binding sites at both sides of the membrane.
45. Answer: T
Difficulty: 3
Feedback: Since talin functions downstream of Rap1, the talin loss-of-function phenotype
is also observed in the double mutants.
46. Answer: C
Difficulty: 2
Feedback: Talin activates integrin by tilting the transmembrane domains of the two
integrin subunits away from each other, favoring the active state.
47. Answer: D
Difficulty: 1
Feedback: Fibronectin is a mechanical sensor in the extracellular matrix, while α-catenin
and talin are intracellular sensors mainly involved in adherens junctions and focal
adhesions, respectively.
48. Answer: CD
Difficulty: 2
Feedback: Activation of some integrins and matrix metalloproteases occurs in the
epithelial–mesenchymal transition, whereas cadherin-based adherens junctions and
connexin-based gap junctions are down-regulated in this process. This allows the cells to
lose their epithelial character.
49. Answer: C
Difficulty: 2
Feedback: Focal adhesion kinase (FAK) is a tyrosine kinase recruited to active integrins
via adaptor proteins such as talin and paxillin; it is autophosphorylated and also
phosphorylated by recruited Src kinases, helping to relay signals into the cell.
50. Answer: FFTT
Difficulty: 2
Feedback: Integrins usually bind to their ligands weakly; their high concentration in the
membrane and their clustering affect their collective affinity.
51. Answer: RRR
Difficulty: 2
Feedback: When the cell is placed on a rigid substratum that strongly resists pulling
forces applied at the cell-matrix junctions, the increased tension is sensed by the junction
and more proteins are recruited to the junction as a result.
52. Answer: WWM
Difficulty: 2
Section: The Plant Cell Wall
Feedback: Plant cell wall is generally thicker, stronger, and more rigid compared to
extracellular matrix surrounding animal cells. It has a very low protein content and is
instead rich in polymers such as cellulose and lignin which can be made from “cheap”
carbon-based structural building blocks.
53. Answer: C
Difficulty: 2
Feedback: Lignin is a complex network of aromatic polymers found in woody tissues.
54. Answer: CPPCP
Difficulty: 2
Feedback: In plant cells walls, cellulose microfibrils are homogeneous polymers that are
cross-linked into a complex network by a heterogeneous set of cross-linking glycans.
Pectins constitute a heterogeneous group of branched polysaccharides which are rich in
galacturonic acid residues and are abundant in the middle lamella. They can turn into a
semirigid gel by the addition of Ca2+.
55. Answer: C
Difficulty: 3
Feedback: The orientation of cellulose microfibrils in the diagram favors growth,
powered by turgor pressure, in the horizontal direction. Without the cell wall, water
would flow into the cell, potentially bursting it open.
56. Answer: B
Difficulty: 2
Feedback: Synthesis of cellulose and its deposition as microfibrils would continue after
such treatment; however, it is thought that in the absence of cortical microtubules, the
process is not spatially restrained, and therefore the microfibril pattern is locked: new
microfibrils are deposited in the same orientation as the layer put down previously.

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MOLECULAR BIOLOGY OF THE CELL: SIXTH EDITION

CHAPTER 19: CELL JUNCTIONS AND THE

4. Which of the following is NOT an anchoring junction?

Cells of the epithelial tube

A. About 20 nm; connexon 1

16. Plasmodesmata in plant cells are functionally similar to animal cells’ …

28. Collagens are extremely rich in …

30. Which collagens are the most common in our body?

Peptide concentration used

Bound integrin (%)

44. In the active state of the integrin dimer, …

48. The chemical compound PMA is a potent stimulator of the epithelial–mesenchymal

49. The focal adhesion kinase …

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