12 Mineral Nutrition of Higher Plants

minor fraction of the K+

(42
K) is readily exchangeable within this 30-min period, most
of the K+
having already been transported across the membranes into the cytoplasm
and vacuoles ('inner space').
Although the plasma membrane and the tonoplast are the main biomembranes
directly involved in solute uptake and transport in roots, it must be borne in m
ind that
compartmentation by biomembranes is a general prerequisite for living systems (L
eigh
and Wyn Jones, 1986). Solute transport into organelles such as mitochondria and
chloroplasts must also therefore be regulated by membranes which separate these
organelles from the surrounding cytoplasm. An example of solute transport across
the
outer chloroplast membrane is given in Section 8.4 for phosphorus and sugars.
The capability of biomembranes for solute transport and its regulation is closel
y
related to their chemical composition and molecular structure. Before the mechan
isms
of solute transport across membranes are discussed in more detail (Sections 2.4
and
2.5), it is therefore appropriate to consider some fundamental aspects of the co
mposition and structure of biomembranes.
2.3 Structure and Composition of Membranes
The capacity of plant cell membranes to regulate solute uptake has fascinated bo
tanists
since the nineteenth century. At that time the experimental techniques available
limited the investigation of the process. Nevertheless, even by the early years
of the
twentieth century some basic facts of solute permeation across the plasma membra
ne
and tonoplast had been established, as for example of the inverse relationship b
etween
membrane permeation and the diameter of uncharged molecules and the rates at whi
ch
they permeate membranes. These ultrafilter-like properties of membranes have bee
n
confirmed more recently, at least in principle (Table 2.5).
Thus, in addition to the cell walls (Section 2.2.1) cell membranes are effective
barriers to solutes of high molecular weight. Most synthetic chelators such as E
DTA
(see also Table 2.4) and microbial siderophores as specific chelators of iron (S
ection
16.5) are of high molecular weight and their rate of permeation is restricted th
rough the
Table 2.5
Reflection Coefficient (?) of Some Nonelectrolytes
at the Cell Membranes of Valonia utricularisa
Compound db
Molecule radius (nm)
Raffinose 1.00 0.61
Sucrose 1.00 0.53
Glucose 0.95 0.44
Glycerol 0.81 0.27
Urea 0.76 0.20

"Based on Zimmermann and Steudle (1970). 6

1.00 indicates that the membranes are impermeable to the
solute; 0 indicates that the membranes are freely permeable
to the solute.
Ion Uptake Mechanisms of Individual Cells and Roots 13
Fig. 2.4 Model of a biomembrane with polar lipids and with either extrinsic or i
ntrinsic,
integrated proteins. The latter can cross the membrane to form 'protein channels
'.
plasma membrane of root cells. It is possible, therefore, to use high-molecularweight
organic solutes such as polyethyleneglycol at high external concentrations as ef
fective
osmotica in order to induce water deficiency (drought stress) in plants.
Molecules which are highly soluble in organic solvents, i.e. with lipophilic pro
perties,
penetrate membranes much faster than would be predicted on the basis of their si
ze.
The solubility of these molecules in the membrane and their ability to diffuse t
hrough
the lipid core of the membranes are presumably the main factors responsible for
the
faster permeation.
Membranes are typically composed of two main classes of compounds: proteins and
lipids. Carbohydrates comprise only a minor fraction of membranes. The relative
abundance of proteins and lipids can be quite variable depending on whether the
membrane is a plasma, mitochondrial, or chloroplast membrane (Clarkson, 1977).
Membranes also differ in diameter, for example in spinach from 10.5 nm (plasma
membrane) to 8.1 nm (tonoplast) and 6.3 nm (endoplasmic reticulum; Auderset etal
.,
1986). However, all biomembranes have some common basic structure as shown in a
model in Fig. 2.4.
Polar lipids (e.g. phospholipids) with the hydrophilic, charged head regions (ph
osphate, amino, and carboxylic groups) are oriented towards the membrane surface.
Protein molecules can be attached (extrinsic proteins), for example, by electros
tatic
binding to the surfaces as membrane-bound enzymes. Other proteins may be integra
ted
into membranes (intrinsic proteins), or traverse the membranes to form 'protein
channels' (transport proteins) which serve to function in membrane transport of
polar
solutes such as ions (Section 2.4).
Three polar lipids represent the major lipid components of membranes: phospholipids, glycolipids, and less abundant, sulfolipids (except in the thylakoid mem
branes of
chloroplasts, where they occur in substantial amounts). Examples of these polar
lipids
are shown below:
14 Mineral Nutrition of Higher Plants
QLI R1
/\/\/\/\A/\/ v
O-CH 2
-0-P-0-CH2-CH2-N+
(CH3)3
O"
Phosphatidylcholine (lecithin)
H n ^ 2
0 J\ H Oh^c

(Long chain polyunsaturated

fatty acids)
CH 2 R2
/Ny^As/V^NyN^O-CH a
CH2OH
OH/OH
OH H
Monogalactosyl diglyceride
o
CH2 --0"
\H O
OH/OH
OH H
Sulfoquinovosyl diglyceride
Another important group of membrane lipids consists of sterols, for example sistosterol:
-Sistosterol
Through their structural role in membranes sterols may indirectly affect transpo
rt
processes such as the activity of the proton pumping ATPase in the plasma membra
ne
(Sandstrom and Cleland, 1989). In agreement with this assumption the sterol cont
ent is
very low in endomembranes (e.g. endoplasmic reticulum) but may make up more than
30% of the total lipids in the plasma membrane (Brown and DuPont, 1989) and also
in
the tonoplast (Table 2.6). Despite these differences in lipids, the fatty acid c
omposition
of the phospholipids is similar in both membranes. The long-chain fatty acids in
polar
membrane lipids vary in both the length and degree of unsaturation (i.e. number
of
double bounds) which influence the melting point (Table 2.6).
Lipid composition not only differs characteristically between membranes of individual cells but also between cells of different plant species (Stuiver et aL, 1
978), it is
also strongly affected by environmental factors. In leaves, for example, distinc
t annual
variations in the levels of sterols occur (Westerman and Roddick, 1981) and in r
oots
both phospholipid content and the proportion of highly unsaturated fatty acids d
ecrease
Ion Uptake Mechanisms of Individual Cells and Roots 15
Table 2.6
Lipid and Fatty Acid Composition of Plasma Membranes and Tonoplasts from Mung Be
ana
Lipids
Plasma membrane
t??\ mg- 1
protein
Tonoplast
p??? mg- 1
protein
Phospholipids
Sterols
Glycolipids
1.29
1.15

0.20
1.93
1.05
0.80
Fatty acid
Fatty acid composition of the phospholipids
Melting Plasma
Chain point membrane
length (C) (% of total)
Tonoplast
(% of total)
Palmitic acid
Stearic acid
Oleic acid
Linoleic acid
HOW PATHOGENS ATTACK PLANTS
175
MECHANICAL FORCES EXERTED BY PATHOGENS ON HOST TISSUES
177
CHEMICAL WEAPONS OF PATHOGENS
179
ENZYMES IN PLANT DISEASE
180
ENZYMATIC DEGRADATION OF CELL WALL SUBSTANCES
180
CUTIN
PECTIC SUBSTANCES
CELLULOSE
CROSS-LINKING GLYCANS (HEMICELLU
CUTICULAR WAX
LOSES) SUBERIN
LIGNIN
CELL WALL FLAVONOIDS CELL WALL STRUCTURAL PROTEINS
180
ENZYMATIC DEGRADATION OF SUBSTANCES CONTAINED IN PLANT CELLS
189
PROTEINS STARCH
LIPIDS
189
MICROBIAL TOXINS IN PLANT DISEASE
190
TOXINS THAT AFFECT A WIDE RANGE OF HOST PLANTS
190
TABTOXIN PHASEOLOTOXIN
TENTOXIN
CERCOSPORIN
OTHER NON-HOST-SPECIFIC TOXINS
191
The intact, healthy plant is a community of cells built
in a fortress-like fashion. Plant cells consist of cell
wall, cell membranes, and cytoplasm, which con-tains the nucleus and various org
anelles (Fig. 5-1) and
all the substances for which the pathogens attack them.
The cytoplasm and the organelles it contains are sepa-rated from each other by m
embranes that carry various
types of proteins embedded in them (Fig. 5-2). The plant
surfaces that come in contact with the environment
either consist of cellulose, as in the epidermal cells of
roots and in the intercellular spaces of leaf parenchyma
cells, or consist of a cuticle that covers the epidermal cell
walls, as is the case in the aerial parts of plants. Often
an additional layer, consisting of waxes, is deposited
outside the cuticle, especially on younger parts of plants
(Fig. 5-3).
Pathogens attack plants because during their evolu-tionary development they have
acquired the ability to
live off the substances manufactured by the host plants,
and some of the pathogens depend on these substances

for survival. Many substances are contained in the

protoplast of the plant cells, however, and if pathogens
are to gain access to them they must first penetrate the
outer barriers formed by the cuticle and/or cell walls.
Even after the outer cell wall has been penetrated,
further invasion of the plant by the pathogen necessi-tates the penetration of m
ore cell walls. Furthermore,
the plant cell contents are not always found in forms
immediately utilizable by the pathogen and must be
broken down to units that the pathogen can absorb and
assimilate. Moreover, the plant, reacting to the presence
and activities of the pathogen, produces structures and
chemical substances that interfere with the advance or
the existence of the pathogen; if the pathogen is to
survive and to continue living off the plant, it must be
able to overcome such obstacles.
Therefore, for a pathogen to infect a plant it must be
able to make its way into and through the plant, obtain
nutrients from the plant, and neutralize the defense reac-tions of the plant. Pa
thogens accomplish these activities
mostly through secretions of chemical substances that
affect certain components or metabolic mechanisms of
176 5. HOW PATHOGENS ATTACK PLANTS
HOST-SPECIFIC OR HOST-SELECTIVE TOXINS
193
T TOXIN [ RACE T TOXIN]
HC TOXIN
VICTORIN, OR HV TOXIN
TOXINS
OTHER HOST-SPECIFIC TOXINS
194
GROWTH REGULATORS IN PLANT DISEASE
196
POLYSACCHARIDES
201
DETOXIFICATION OF LOW MOLECULAR WEIGHT ANTIMICROBIAL MOLECULES
201
PROMOTION OF BACTERIAL VIRULENCE BY GENES
202
ROLE OF TYPE III SECRETION IN BACTERIAL PATHOGENESIS
202
SUPPRESSORS OF PLANT DEFENSE RESPONSES
202
PATHOGENICITY AND VIRULENCE FACTORS IN VIRUSES AND VIROIDS
203
AVR
ALTERNARIA ALTERNATA
COCHLIOBOLUS (HELMINTHOSPORIUM) HETEROSTROPHUS
Middle lamella
Primary cell wall
Secondary cell wall
Plasma membrane
Cell wall
Middle lamella
Air space
Plasmodesma
Nucleolus
Nucleus
Chloroplast
Golgi body
Peroxysome
Vacuole
Nuclear membrane

Mitochondrion
Endoplasmic
reticulum
FIGURE 5-1 Schematic representation of a plant cell and its main components.
their hosts. Penetration and invasion, however, seem to
be aided by, or in some cases be entirely the result of,
the mechanical force exerted by certain pathogens on
the cell walls of the plant.
MECHANICAL FORCES EXERTED
BY PATHOGENS ON HOST TISSUES
Plant pathogens are, generally, tiny microorganisms that
cannot apply a voluntary force to a plant surface.
Only some fungi, parasitic higher plants, and nematodes
appear to apply mechanical pressure to the plant surface
they are about to penetrate. The amount of pressure,
however, may vary greatly with the degree of presoft-ening of a plant surface by e
nzymatic secretions of the
pathogen.
For fungi and parasitic higher plants to penetrate a
plant surface, they must, generally, first adhere to it.
Hyphae and radicles are usually surrounded by
mucilaginous substances, and their adhesion to the plant
seems to be brought about primarily by the intermolec-ular forces developing bet
ween the surfaces of plant and
pathogen on close contact with the adhesive substances
and with one another. In some cases an adhesion pad
forms from the spore when it comes in contact with a
moist surface, and cutinase and cellulase enzymes
released from the spore surface help the spore adhere to
the plant surface. Spores of some fungi carry adhesive
substances at their tips that, on hydration, allow spores
to become attached to various surfaces.
After contact is established, the diameter of the tip of
the hypha or radicle in contact with the host increases
and forms the flattened, bulb-like structure called the
appressorium (Figs. 2-4 and 2-5). This increases the area
of adherence between the two organisms and securely
fastens the pathogen to the plant. From the appresso-rium, a fine growing point,
called the penetration peg,
HOW PATHOGENS ATTACK PLANTS
175
MECHANICAL FORCES EXERTED BY PATHOGENS ON HOST TISSUES
177
CHEMICAL WEAPONS OF PATHOGENS
179
ENZYMES IN PLANT DISEASE
180
ENZYMATIC DEGRADATION OF CELL WALL SUBSTANCES
180
CUTICULAR WAX
CUTIN
PECTIC SUBSTANCES
CELLULOSE
CROSS-LINKING GLYCANS (HEMICELLU
LOSES) SUBERIN
LIGNIN
CELL WALL FLAVONOIDS CELL WALL STRUCTURAL PROTEINS
180
ENZYMATIC DEGRADATION OF SUBSTANCES CONTAINED IN PLANT CELLS
189
PROTEINS STARCH
LIPIDS
189
MICROBIAL TOXINS IN PLANT DISEASE
190
TOXINS THAT AFFECT A WIDE RANGE OF HOST PLANTS

190
TABTOXIN
PHASEOLOTOXIN
TENTOXIN
CERCOSPORIN
OTHER NON-HOST-SPECIFIC TOXINS
191
The intact, healthy plant is a community of cells built
in a fortress-like fashion. Plant cells consist of cell
wall, cell membranes, and cytoplasm, which con-tains the nucleus and various org
anelles (Fig. 5-1) and
all the substances for which the pathogens attack them.
The cytoplasm and the organelles it contains are sepa-rated from each other by m
embranes that carry various
types of proteins embedded in them (Fig. 5-2). The plant
surfaces that come in contact with the environment
either consist of cellulose, as in the epidermal cells of
roots and in the intercellular spaces of leaf parenchyma
cells, or consist of a cuticle that covers the epidermal cell
walls, as is the case in the aerial parts of plants. Often
an additional layer, consisting of waxes, is deposited
outside the cuticle, especially on younger parts of plants
(Fig. 5-3).
Pathogens attack plants because during their evolu-tionary development they have
acquired the ability to
live off the substances manufactured by the host plants,
and some of the pathogens depend on these substances
for survival. Many substances are contained in the
protoplast of the plant cells, however, and if pathogens
are to gain access to them they must first penetrate the
outer barriers formed by the cuticle and/or cell walls.
Even after the outer cell wall has been penetrated,
further invasion of the plant by the pathogen necessi-tates the penetration of m
ore cell walls. Furthermore,
the plant cell contents are not always found in forms
immediately utilizable by the pathogen and must be
broken down to units that the pathogen can absorb and
assimilate. Moreover, the plant, reacting to the presence
and activities of the pathogen, produces structures and
chemical substances that interfere with the advance or
the existence of the pathogen; if the pathogen is to
survive and to continue living off the plant, it must be
able to overcome such obstacles.
Therefore, for a pathogen to infect a plant it must be
able to make its way into and through the plant, obtain
nutrients from the plant, and neutralize the defense reac-tions of the plant. Pa
thogens accomplish these activities
mostly through secretions of chemical substances that
affect certain components or metabolic mechanisms of
176 5. HOW PATHOGENS ATTACK PLANTS
HOST-SPECIFIC OR HOST-SELECTIVE TOXINS
193
HC TOXIN
VICTORIN, OR HV TOXIN T TOXIN [ RACE T TOXIN]
TOXINS
OTHER HOST-SPECIFIC TOXINS
194
GROWTH REGULATORS IN PLANT DISEASE
196
POLYSACCHARIDES
201
DETOXIFICATION OF LOW MOLECULAR WEIGHT ANTIMICROBIAL MOLECULES
201
PROMOTION OF BACTERIAL VIRULENCE BY GENES
202
ROLE OF TYPE III SECRETION IN BACTERIAL PATHOGENESIS

202
SUPPRESSORS OF PLANT DEFENSE RESPONSES
202
PATHOGENICITY AND VIRULENCE FACTORS IN VIRUSES AND VIROIDS
203
AVR
ALTERNARIA ALTERNATA
COCHLIOBOLUS (HELMINTHOSPORIUM) HETEROSTROPHUS
Middle lamella
Primary cell wall
Secondary cell wall
Plasma membrane
Cell wall
Middle lamella
Air space
Plasmodesma
Nucleolus
Nucleus
Chloroplast
Golgi body
Peroxysome
Vacuole
Nuclear membrane
Mitochondrion
Endoplasmic
reticulum
FIGURE 5-1 Schematic representation of a plant cell and its main components.
their hosts. Penetration and invasion, however, seem to
be aided by, or in some cases be entirely the result of,
the mechanical force exerted by certain pathogens on
the cell walls of the plant.
MECHANICAL FORCES EXERTED
BY PATHOGENS ON HOST TISSUES
Plant pathogens are, generally, tiny microorganisms that
cannot apply a voluntary force to a plant surface.
Only some fungi, parasitic higher plants, and nematodes
appear to apply mechanical pressure to the plant surface
they are about to penetrate. The amount of pressure,
however, may vary greatly with the degree of presoft-ening of a plant surface by e
nzymatic secretions of the
pathogen.
For fungi and parasitic higher plants to penetrate a
plant surface, they must, generally, first adhere to it.
Hyphae and radicles are usually surrounded by
mucilaginous substances, and their adhesion to the plant
seems to be brought about primarily by the intermolec-ular forces developing bet
ween the surfaces of plant and
pathogen on close contact with the adhesive substances
and with one another. In some cases an adhesion pad
forms from the spore when it comes in contact with a
moist surface, and cutinase and cellulase enzymes
released from the spore surface help the spore adhere to
the plant surface. Spores of some fungi carry adhesive
substances at their tips that, on hydration, allow spores
to become attached to various surfaces.
After contact is established, the diameter of the tip of
the hypha or radicle in contact with the host increases
and forms the flattened, bulb-like structure called the
appressorium (Figs. 2-4 and 2-5). This increases the area
of adherence between the two organisms and securely

fastens the pathogen to the plant. From the appresso-rium, a fine growing point,
called the penetration peg,
Linolenic acid
Others
Ci 6
Cl 8
G b
^-18:1
r b
r b
^18:3
+62.8
+70.1
+ 13.0
-5.5
-11.1
35
6
9
21
19
10
39
6
9
22
20
4
"Based on Yoshida and Uemura (1986). Reprinted by permission of the American Soc
iety of Plant
Physiologists.
^Numeral to the right of the colon indicates the number of double bounds.
under zinc deficiency (Cakmak and Marschner, 1988c). In many instances changes i
n
lipid composition reflect adaptation of a plant to its environment through adjus
tment of
membrane properties. Generally, highly unsaturated fatty acids predominate in pl
ants
that grow in cold climates. During acclimatization of plants to low temperatures
an
increase in highly unsaturated fatty acids is also often observed (Bulder et al.
, 1991).
Such a change shifts the freezing point (i.e. the transition temperature) of mem
branes
to a lower temperature and may thus be of importance for maintenance of membrane
functions at low temperatures. It is questionable, however, to generalize about
the
effect of temperature on lipid composition of membranes. In rye, for example, wh
ich is
a cold-tolerant plant species, the proportion of polyunsaturated fatty acids in
the roots
decreased rather than increased as the roots were cooled (White et al., 1990b).
During acclimatization of roots to low temperatures synthesis of new membrane
proteins is also enhanced (Mohapatra et al., 1988) and phospholipids increase co
nsiderably (Kinney et al., 1987). Since phospholipids probably act as receptors for ph
ytohor-

mones such as gibberellic acid, increasing responsiveness of membranes to gibber

ellic
acid at low temperatures may be related to these changes (Singh and Paleg, 1984)
.
The property of membranes in ion selectivity and lipid composition are often hig
hly
correlated as for example between chloride uptake and sterols (Douglas and Walke
r,
i983) and galactolipids (Section 16.6). Also the crop plant species bean, sugar
beet and
barley differ not only in the fatty acid composition of root membranes (Stuiver
et al.,
1978) but also considerably in the uptake of sodium (Section 10.2).
Alterations in the lipid composition of root membranes are also typical response
s to
changes in the mineral nutrient supply or exposure to salinity (Kuiper, 1980). O
f the
12 Mineral Nutrition of Higher Plants
minor fraction of the K+
(42
K) is readily exchangeable within this 30-min period, most
of the K+
having already been transported across the membranes into the cytoplasm
and vacuoles ('inner space').
Although the plasma membrane and the tonoplast are the main biomembranes
directly involved in solute uptake and transport in roots, it must be borne in m
ind that
compartmentation by biomembranes is a general prerequisite for living systems (L
eigh
and Wyn Jones, 1986). Solute transport into organelles such as mitochondria and
chloroplasts must also therefore be regulated by membranes which separate these
organelles from the surrounding cytoplasm. An example of solute transport across
the
outer chloroplast membrane is given in Section 8.4 for phosphorus and sugars.
The capability of biomembranes for solute transport and its regulation is closel
y
related to their chemical composition and molecular structure. Before the mechan
isms
of solute transport across membranes are discussed in more detail (Sections 2.4
and
2.5), it is therefore appropriate to consider some fundamental aspects of the co
mposition and structure of biomembranes.
2.3 Structure and Composition of Membranes
The capacity of plant cell membranes to regulate solute uptake has fascinated bo
tanists
since the nineteenth century. At that time the experimental techniques available
limited the investigation of the process. Nevertheless, even by the early years
of the
twentieth century some basic facts of solute permeation across the plasma membra
ne
and tonoplast had been established, as for example of the inverse relationship b
etween
membrane permeation and the diameter of uncharged molecules and the rates at whi
ch
they permeate membranes. These ultrafilter-like properties of membranes have bee
n
confirmed more recently, at least in principle (Table 2.5).
Thus, in addition to the cell walls (Section 2.2.1) cell membranes are effective

barriers to solutes of high molecular weight. Most synthetic chelators such as E

DTA
(see also Table 2.4) and microbial siderophores as specific chelators of iron (S
ection
16.5) are of high molecular weight and their rate of permeation is restricted th
rough the
Table 2.5
Reflection Coefficient (?) of Some Nonelectrolytes
at the Cell Membranes of Valonia utricularisa
Compound db
Molecule radius (nm)
Raffinose 1.00 0.61
Sucrose 1.00 0.53
Glucose 0.95 0.44
Glycerol 0.81 0.27
Urea 0.76 0.20
"Based on Zimmermann and Steudle (1970). 6
1.00 indicates that the membranes are impermeable to the
solute; 0 indicates that the membranes are freely permeable
to the solute.
Ion Uptake Mechanisms of Individual Cells and Roots 13
Fig. 2.4 Model of a biomembrane with polar lipids and with either extrinsic or i
ntrinsic,
integrated proteins. The latter can cross the membrane to form 'protein channels
'.
plasma membrane of root cells. It is possible, therefore, to use high-molecularweight
organic solutes such as polyethyleneglycol at high external concentrations as ef
fective
osmotica in order to induce water deficiency (drought stress) in plants.
Molecules which are highly soluble in organic solvents, i.e. with lipophilic pro
perties,
penetrate membranes much faster than would be predicted on the basis of their si
ze.
The solubility of these molecules in the membrane and their ability to diffuse t
hrough
the lipid core of the membranes are presumably the main factors responsible for
the
faster permeation.
Membranes are typically composed of two main classes of compounds: proteins and
lipids. Carbohydrates comprise only a minor fraction of membranes. The relative
abundance of proteins and lipids can be quite variable depending on whether the
membrane is a plasma, mitochondrial, or chloroplast membrane (Clarkson, 1977).
Membranes also differ in diameter, for example in spinach from 10.5 nm (plasma
membrane) to 8.1 nm (tonoplast) and 6.3 nm (endoplasmic reticulum; Auderset etal
.,
1986). However, all biomembranes have some common basic structure as shown in a
model in Fig. 2.4.
Polar lipids (e.g. phospholipids) with the hydrophilic, charged head regions (ph
osphate, amino, and carboxylic groups) are oriented towards the membrane surface.
Protein molecules can be attached (extrinsic proteins), for example, by electros
tatic
binding to the surfaces as membrane-bound enzymes. Other proteins may be integra
ted
into membranes (intrinsic proteins), or traverse the membranes to form 'protein
channels' (transport proteins) which serve to function in membrane transport of
polar
solutes such as ions (Section 2.4).

Three polar lipids represent the major lipid components of membranes: phospholipids, glycolipids, and less abundant, sulfolipids (except in the thylakoid mem
branes of
chloroplasts, where they occur in substantial amounts). Examples of these polar
lipids
are shown below:
14 Mineral Nutrition of Higher Plants
QLI R1
/\/\/\/\A/\/ v
O-CH 2
-0-P-0-CH2-CH2-N+
(CH3)3
O"
Phosphatidylcholine (lecithin)
H n ^ 2
0 J\ H Oh^c
(Long chain polyunsaturated
fatty acids)
CH 2 R2
/Ny^As/V^NyN^O-CH a
CH2OH
OH/OH
OH H
Monogalactosyl diglyceride
o
CH2 --0"
\H O
OH/OH
OH H
Sulfoquinovosyl diglyceride
Another important group of membrane lipids consists of sterols, for example sistosterol:
-Sistosterol
Through their structural role in membranes sterols may indirectly affect transpo
rt
processes such as the activity of the proton pumping ATPase in the plasma membra
ne
(Sandstrom and Cleland, 1989). In agreement with this assumption the sterol cont
ent is
very low in endomembranes (e.g. endoplasmic reticulum) but may make up more than
30% of the total lipids in the plasma membrane (Brown and DuPont, 1989) and also
in
the tonoplast (Table 2.6). Despite these differences in lipids, the fatty acid c
omposition
of the phospholipids is similar in both membranes. The long-chain fatty acids in
polar
membrane lipids vary in both the length and degree of unsaturation (i.e. number
of
double bounds) which influence the melting point (Table 2.6).
Lipid composition not only differs characteristically between membranes of individual cells but also between cells of different plant species (Stuiver et aL, 1
978), it is
also strongly affected by environmental factors. In leaves, for example, distinc
t annual
variations in the levels of sterols occur (Westerman and Roddick, 1981) and in r
oots
both phospholipid content and the proportion of highly unsaturated fatty acids d

ecrease
Ion Uptake Mechanisms of Individual Cells and Roots 15
Table 2.6
Lipid and Fatty Acid Composition of Plasma Membranes and Tonoplasts from Mung Be
ana
Lipids
Plasma membrane
t??\ mg- 1
protein
Tonoplast
p??? mg- 1
protein
Phospholipids
Sterols
Glycolipids
1.29
1.15
0.20
1.93
1.05
0.80
Fatty acid
Fatty acid composition of the phospholipids
Melting Plasma
Chain point membrane
length (C) (% of total)
Tonoplast
(% of total)
Palmitic acid
Stearic acid
Oleic acid
Linoleic acid
Linolenic acid
Others
Ci 6
Cl 8
G b
^-18:1
r b
r b
^18:3
+62.8
+70.1
+ 13.0
-5.5
-11.1
35
6
9
21
19
10
39
6
9
22
20
4
"Based on Yoshida and Uemura (1986). Reprinted by permission of the American Soc

iety of Plant
Physiologists.
^Numeral to the right of the colon indicates the number of double bounds.
under zinc deficiency (Cakmak and Marschner, 1988c). In many instances changes i
n
lipid composition reflect adaptation of a plant to its environment through adjus
tment of
membrane properties. Generally, highly unsaturated fatty acids predominate in pl
ants
that grow in cold climates. During acclimatization of plants to low temperatures
an
increase in highly unsaturated fatty acids is also often observed (Bulder et al.
, 1991).
Such a change shifts the freezing point (i.e. the transition temperature) of mem
branes
to a lower temperature and may thus be of importance for maintenance of membrane
functions at low temperatures. It is questionable, however, to generalize about
the
effect of temperature on lipid composition of membranes. In rye, for example, wh
ich is
a cold-tolerant plant species, the proportion of polyunsaturated fatty acids in
the roots
decreased rather than increased as the roots were cooled (White et al., 1990b).
During acclimatization of roots to low temperatures synthesis of new membrane
proteins is also enhanced (Mohapatra et al., 1988) and phospholipids increase co
nsiderably (Kinney et al., 1987). Since phospholipids probably act as receptors for ph
ytohormones such as gibberellic acid, increasing responsiveness of membranes to gibber
ellic
acid at low temperatures may be related to these changes (Singh and Paleg, 1984)
.
The property of membranes in ion selectivity and lipid composition are often hig
hly
correlated as for example between chloride uptake and sterols (Douglas and Walke
r,
i983) and galactolipids (Section 16.6). Also the crop plant species bean, sugar
beet and
barley differ not only in the fatty acid composition of root membranes (Stuiver
et al.,
1978) but also considerably in the uptake of sodium (Section 10.2).
Alterations in the lipid composition of root membranes are also typical response
s to
changes in the mineral nutrient supply or exposure to salinity (Kuiper, 1980). O
f the
//BIMx
//Copyright 2002-2013 Graphisoft SE All rights reserved
//Sat Nov 28 10:55:38 2015
//0.000 InitCore
Program directory is 'C:\Program Files\GRAPHISOFT\ArchiCAD 18\Add-Ons\BIMx\'
Incoming path is 'C:\Program Files\GRAPHISOFT\ArchiCAD 18\Add-Ons\BIMx\BIMx.ap
x'
//0.003 Shutdown
Shutting down core
//BIMx
//Copyright 2002-2013 Graphisoft SE All rights reserved
//Sat Nov 28 10:55:38 2015

//0.000 InitCore
Program directory is 'C:\Program Files\GRAPHISOFT\ArchiCAD 18\Add-Ons\BIMx\'
Incoming path is 'C:\Program Files\GRAPHISOFT\ArchiCAD 18\Add-Ons\BIMx\BIMx.ap
x'
//0.003 Shutdown
Shutting down core
//BIMx
//Copyright 2002-2013 Graphisoft SE All rights reserved
//Sat Nov 28 10:55:38 2015
//0.000 InitCore
Program directory is 'C:\Program Files\GRAPHISOFT\ArchiCAD 18\Add-Ons\BIMx\'
Incoming path is 'C:\Program Files\GRAPHISOFT\ArchiCAD 18\Add-Ons\BIMx\BIMx.ap
x'
//0.003 Shutdown
Shutting down core