• Plant cell walls

• Complex structures involved in

– Structure and strength
– Water relations
– Development and morphogenesis
– Defense
• Figure 15.1 Cross-section of a
stem of a buttercup (Ranuculus
repens)
• Figure 15.3 Diversity of cell wall
structure

• Cell walls
• Primary cell wall
– Laid down by growing cells
– Relatively Simple
– Extensible
• Secondary cell wall
– Laid down largely after growth
ceases
– Internal to 1° cell wall
– May be highly complex,
multilayered
• 15.2 Two views of primary cell
walls
• 15.3 Outer epidermal cell wall
from the growing region of a bean
hypocotyl
• Primary cell walls can also be heavily thickened, and
may differ in morphology across a single cell
• Cell walls
• Primarily consist of sugar
polymers
– Cellulose
– Matrix polysaccharides
 • Pectins (extractable with boiling water
or Ca2+ chelators)
• Hemicelluloses (extractable with hot
NaOH)
• Proteins
• Phenolic lignins and other
components
• Table 14.1 Structural
components of plant cell walls
• 15.5 Conformational structures
of sugars commonly found in
plant cell walls (Part 1)

• 15.5 Conformational structures

of sugars commonly found in
plant cell walls (Part 2)
• (1→4)β-D-glucans
• 15.6 A structural model of a
cellulose microfibril (Part 1)
• Cellulose microfibrils
•
• -Semi-crystalline with abundant H-bonding

-Extremely strong
-inaccessible to
enzymatic attack
-high tensile
strength
• -Crosslinked by matrix polysaccharides and proteins
• 15.6 A structural model of a
cellulose microfibril (Part 2)

• 15.7 Cellulose synthesis by the

cell (Part 1)
• Cellulose microfibrils are constructed on
“rosettes” at the plasma membrane
• -Cellulose synthase (CesA) family
complexes arranged into trimeric to
 hexameric subunits, which combine to make
a rosette
• -Construct β-(1-4)-glucans by addition of
UDP-glucoses
• -Associated with microtubules inside cell
membrane
• Figure 14.7 Cellulose
microfibrils are synthesized at the
cell surface by membrane-bound
complexes containing cellulose
synthase (CESA) proteins
• 15.8 Model of cellulose synthesis
by a multisubunit complex
containing cellulose synthase
• Sugar-nucleotide donor
(UDP-Glucose) provides sequential addition of glucoses to
cellulose synthase

• Sterol glucoside may act as “primer” for the glucan chains of

cellulose
• Major structural components of
the primary cell wall and their
likely arrangement

• Matrix polysaccharides
• Produced by glycosyltransferases
in Golgi
• Delivered by exocytosis to the
cell wall

• Hemicelluloses and Pectins

• 15.11 Partial structures of the
most common pectins (Part 1)
• Pectins
• -From fairly simple to very complex and
variable
• -Form soluble hydrophilic gel in cell wall
• -Porosity limits movement of some
macromolecules in the apoplast
• -May bind borate covalently or form ionic
bonds with Ca2+ ions
• 15.11 Partial structures of the
most common pectins (Part 2)

• 15.11 Partial structures of the

most common pectins (Part 3)

• Figure 15.13 Linear arrangement

of various pectin domains to each
other
• Pectins of different types may be linked into long
chains…
• Hemicelluloses
• Linear matrix polysaccharides
with short branches
– Often very long backbones
 – Associate tightly with cellulose
• may bind microfibrils together or
• may prevent binding to add
flexibility
• 15.10 Partial structures of
common hemicelluloses (Part 1)
• Xyloglucans are the most abundant hemicelluloses in most
plant groups’ primary cell walls
• -Glucan backbone with attached xylose sidechains
• -Link celluloses together and to other matrix components
• Figure 14.11 Partial structures of
the major hemicelluloses (Part 3)
• Figure 14.11 Partial structures of
the major hemicelluloses (Part 4)
• 15.14 A repeated
hydroxyproline-rich motif from a
molecule of HRGP from tomato
• Cell wall matrix proteins include structural
repeat motifs
• -Hydroxyproline-rich glycoproteins (HRGP)
• -proline-rich proteins (PRP)
• -glycine-rich proteins (GRP)
• 15.15 A highly branched
arabinogalactan molecule
• Arabinogalactan proteins (AGP) may be involved in
developmental signaling and embryogenesis
• New cell walls form
at the cell plate
• Directed by the phragmoplast
(microtubules and membrane
vesicle aggregates)
• Cell walls may form by self-
assembly or by enzyme-mediated
assembly
– self aggregation does occur
– form incomplete cell walls without
additional enzymes such as
Xyloglucan endotransglycosylases
(XETs)
 – XETs cut and rejoin xyloglucan
backbones
• Xyloglucan
endotransglucosylases (XET) cut
and join xyloglucan polymers into
new configurations
• Cell wall growth
• As cells grow, cell walls thin and
new material must be added to
retain structural integrity
• Enzymes (such as XET, pectin
methylesterases, glycosylases)
allow integration of new material
into the existing wall;
• expansins allow cellulose
microfibrils to “slide” with
respect to one another
• Cell growth
• Cells can grow by tip growth (e.g.
root hairs, pollen tubes) or by
diffuse growth (most plant cells)
• Expansion of cells depends on
microfibril orientation and cell
wall characteristics
– Isotropic (random) vs. anisotropic
orientation
– Cortical microtubules control where
cellulose microfibrils are deposited

• 15.19 The cell surface expands

differently during tip growth and
diffuse growth
• 15.20 The orientation of newly
deposited cellulose microfibrils

• Cell growth
• Direction and location of cortical
microtubules guiding cellulose
deposition
– Rho-like GTPases (ROPs) activated
by RICs initiate microtubule
formation in specific locations to
orient growth
– Allows complex cellular growth
patterns

• 15.21 Interdigitating cell growth

of leaf pavement cells (Part 1)
• 15.21 Interdigitating cell growth
of leaf pavement cells (Part 2)
• When activated by RIC1, ROP forms “neck”
oriented microtubules; when activated by
RIC4, it orients “lobe” actin filaments
• 15.22 The orientation of
microtubules in the cortical
cytoplasm
• Cellular cortical microtubules (green) coincide with
microfibril orientation (blue)
• 15.23 The disruption of cortical
microtubules

• Figure 15.23 Cortical

microtubules; (C) CesA proteins
and microtubules

• Cell elongation and expansion

• Cell walls must yield for growth
to occur
– Yielding properties (e.g. threshold,
rate)
– Expansion produces stress
relaxation
• Internal pressure (turgor) drives
expansion
– Stress (force per unit area)
• In growing cells, as water enters
the cell the turgor increases and
causes the wall to yield,
increasing the cell volume and
relieving the ΨP .

• Cell elongation and expansion

• Rate of water uptake = cell area
(A) x membrane permeability (LP)
x (ΔΨW)
• In growing cells, as water enters
the cell
– turgor increases
– above threshold, causes the wall to
yield,
– increases the cell volume and
relieves the ΨP .
– As water enters, it dilutes the
solutes and makes ΨS less negative
– Thus, the cell must take up or
generate more solutes to drive water
uptake for expansion (turgor stress)
to continue.

• Cell wall loosening

• Changes in Yield threshold (Y) or
cell wall extensibility (m) are
used to change growth rate for a
given ψp
– Loosening enzymes
– Cell wall acidification (e.g. via
auxin)
• Acid growth hypothesis

• 15.25 Reduction of cell turgor

pressure (water potential) by
stress relaxation
• Auxin induces cell wall acidification, allowing for
rapid stress relaxation and thus loss of turgor
corresponding to increased cell size
--if continued H2O uptake is allowed (control), no loss
of turgor occurs
• 15.26 Acid-induced extension of
isolated cell walls, measured in an
extensometer
• Artificial cell wall acidification increases
extensibility of the cell walls (but only those
cut from growing tissues)
• Acid growth hypothesis
• Acidification of the cell wall (via
auxin or fusicoccin activation of
H+ ATPase) stimulates expansins
and other cell wall enzymes
– Expansins act in catalytic quantities
but do not break covalent bonds
– α-expansins act on cellulose-
cellulose junctions in eudicots,
while β-expansins loosen
glucuranoglycan (GAX) in
gymnosperm primary walls
• Hydrolysis of matrix
polysaccharides
• Cell wall “creep” is irreversible
 – extensibility decreases over time:
rigidification

• 15.27 Scheme for the

reconstitution of extensibility of
isolated cell walls (Part 1)

• 15.27 Scheme for the

reconstitution of extensibility of
isolated cell walls (Part 2)
• Freeze-thaw kills cells without destroying
cell-wall enzymes.
• Heat inactivation of tissue denatures cell
wall-loosening enzymes

• Purified expansins added

back to boiled tissue
cell walls allows
elongation again,
 but only at
acidic pH
• Cell wall growth cessation
• Older cells lose the ability for
continued expansion
– Incorporation or modification of
matrix components that are less
subject to loosening
– Covalent crosslinking (e.g. lignins,
proteins)
– Newer (2°) cell walls control
growth more than older walls

• Figure 14.20 Alternative

concepts of the structural role of
xyloglucan
• Figure 14.21 Secondary cell
walls
• Figure 15.18 Secondary cell
walls; (D) Structure of poplar
lignin
• Lignins are complex polyphenolics that crosslink
cellulose microfibrils and other cell wall
components, particularly in secondary cell walls
• Figure 14.23 Schematic
representation of Casparian strip
deposition
• Casparian strip is produced by lignification and
suberin deposition in specific regions of endodermal
cell walls
• Cell walls in the Casparian strip membrane
domain are loaded with monolignols via plasma
membrane transporters
• Organized by CASP1 protein
• NADPH oxidase, peroxidase produce reactive
peroxides and radical monolignol intermediates for
polymerization
• ESB1 (Enhanced Suberin 1) may help direct
polymerization

• Another change in the calendar!

• This Friday (3/27) – Wednesday,
April 1 “Recalibration”
• Tuesday, April 7 follows
Wednesday schedule
• “Spring Break” April 8-10