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Cape Chemistry Unit 1 Paper 1 2010
Cape Chemistry Unit 1 Paper 1 2010
Journal of Experimental Botany, Vol. 48, No. 306, pp. 45-58, January 1997 Experimental
Botany
1
Fax: +46 18 673037. E-mail: Wieland.Fncke©emc.s/u.se
solvent drag effect on solutes (Zimmermann et al., 1992). CaSO4. Eight seedlings were then transferred to 5 1 of aerated,
An alternative explanation for the direction of gradients modified Hoagland solution containing 1 mM NaCl (Fricke
et al., 1994) and grown at a PAR of 300-400 jumol photons m~ 2
has been provided by the results of Meshcheryakov et al. s~' and a 16 h day/8 h dark period. Plants were grown in the
(1992) which suggest that in the mature region of the same room where measurements were carried out. Plants were
hypocotyl of Ricinus communis, gradients in turgor (and analysed when they were 12-22-d-old. Second, third or fourth
77) are related to the proximity of cells to the phloem leaves were analysed at the time they represented the youngest,
representing the major source of organic osmotica. A fully-, or nearly fully-expanded leaf. Leaves were analysed half-
way along the expanded blade at the adaxial (upper) epidermis,
similar explanation has been given by Frensch and Hsiao within a small region covering the first large lateral vein (LV),
(1994) for the observation that cells located in the inner counted from the midrib, the adjacent (outward) intermediate
cortex of osmotically-stressed maize roots recovered faster vein (IV) and the second large lateral vein. For details of the
in turgor than cells located in the outer cortex. NaCl treatments see Figure legends and Tables.
Pressure-probe studies on leaves (mature tissue of the
Location of epidermal cells
leaf epidermis of Tradescantia virginiana, Tomos et al.,
RIDGE
Fig. 1. Cross-section (A) and view of the upper epidermis (B) of a barley leaf. The cross-section was cut fresh by hand from the third leaf of
barley, 1 d following full expansion. The same leaf was used to obtain a view of the upper leaf surface using a double-reprint technique (Fncke
el al., 1995). Ridges that cover large lateral veins (LV), with a bundle sheath extension stretching from the bundle to the centre top of the ridge-
epidermal surface are referred to as LV-ridges; ridges that cover intermediate veins (IV), which lack such a far-stretching bundle sheath extension,
are referred to as IV-ridges. At the transition from a ndge (R-cells) to trough region (TR-cells) is typically one file of stomata, and interstomatal
cells (IS) are located in between stomata. RT, RM and RB refers to ridge cells located at the top-, mid- or base-position analysed within ndges; UE,
upper epidermis; LE, lower epidermis; MS, mesophyll; SC, substomatal cavity. The bar represents 50 ^m.
(Bangor, North Wales, UK). To minimize vibration during Clark Electromedical Instruments, Pangbourne, UK) using a
turgor measurements, all pieces of equipment were tightened capillary puller (type 50-2013, Harvard Apparatus Ltd,
to a metal plate that rested on inflated rubber (bicycle) Edenbridge, UK) and broken with a microforge (type MF-83,
tubes. Probe capillaries were pulled from borosilicate glass Narashige Scientific Laboratories, Tokyo, Japan). Non-silanized
capillaries (outer diameter, 1.0 mm; inner diameter, 0.58 mm; capillaries were used and filled, as was the rest of the probe,
48 Fricke
with silicone oil (type AS4, Wacker Chemie, MOnchen, (Malone et al., 1989). The extracted sap was transferred within
Germany). The underestimation of cell turgor caused by the seconds under liquid paraffin placed on to the osmometer stage
volume of cell solution outside the cell (in the capillary) during and analysed for osmolality within the following 70 min.
turgor measurements should be highest for the smallest cells,
i.e. here interstomatal cells (Fig. IB). The volume of inter-
stomatal cells was generally between 31-50 pi (calculated from
leaf cross-sections and reprints of the epidermal surface; trough- Results
cell volume generally between 400-520 pi and ridge-cell volume
Turgor profile within IV- and LV-ridges
between 750-980 pi). The volume of cell solution outside the
cell was less than 0.078 pi, i.e. less than 0.16-0.25% of the The turgor relationship of the first and second LV
interstomatal cell volume. Assuming a volumetric elastic
modulus of 5 MPa (see Malone and Tomos [1990] for leaf (counted from the midrib) was similar and results from
epidermal cells of wheat) this would result in an underestimation these two LVs were therefore pooled. Since IV- and
of interstomatal cell turgor of 0.008-0.013 MPa. This is about LV-ridges were analysed from the same leaves, their mean
an order of magnitude smaller than the differences in turgor cell turgor could be compared directly.
found here between cell types and, therefore, does not interfere Both IV- and LV-ridges showed even distribution pro-
0.6 OJ
o 0J • OJ
0.0 0.0
15 B Large lateral vein D Large latara vein
is
o. 4T
-+—t
1.2 1 ?
1 1 1 I
0.9 r •— T ~~ OJ
0.6 o.e
o OJ
0.0 0.0
Bate Middle Top Top Middle Bate Bate Middle Top Top Middle Bate
Fig. 2. Turgor pressure profile within IV- and LV-ndges of the upper epidermis of barley leaves. Cells located at adjacent positions (R B , RM> ^ T )
within ridge regions overlying an IV (IV-ridge) or LV (LV-ridge) were analysed for turgor. The centre top of a ridge is indicated by a dashed line.
Analyses were carried out under low (A, B) or high PAR (C, D). Second, third and fourth leaves were analysed between 2 d preceding and
following full expansion. Cells were probed in such a way that first every other cell was analysed and thereafter the cells in between. Cells analysed
under low PAR conditions were located at direct neighbouring positions, whereas under high PAR conditions cells were displaced from each other
by one to two cell lengths up- or downstream along the vein. Results in (A) are expressed as means ± S D of 5-8 leaf analyses since not every leaf
analysed was composed of 6 ridge cells or a cell was lost during turgor measurement. Results in (B). (C) and (D) are expressed as means ± S D of
7, 5 and 7 complete leaf analyses, respectively.
Turgor in leaf epidermal cells 49
cells had turgors that were different by up to 0.15 MPa in four leaves significantly higher than trough-cell turgor
(not shown). The latter was not an artefact caused by the (indicated in Fig. 3A by '4/15'). The difference in turgor
probing of neighbouring cells, i.e. there was no such was more pronounced and more often significant when
tendency that the turgor of a cell probed second was leaves were analysed under high PAR (Fig. 3B). On
lower than the turgor of its neighbour cell probed first. average, ridge-cell turgor was 0.09 MPa higher than
The mean (±SD) ridge-cell turgor, calculated from trough- and 0.08 MPa higher than interstomatal-cell
the values shown in Fig. 2, for IV and LV-ridges were: turgor. The differences was in 6 of 8 plants significant
low PAR: IV-ridge 1.03 ±0.02 MPa, LV-ridge (mostly at 1% level). In some leaves, ridge-cell turgor
1.04±0.03MPa; high PAR: IV-ridge 1.21 +0.03 MPa, was up to 0.19 MPa higher than trough- or interstomatal-
LV-ridge 1.19 ±0.03 MPa. Thus, under both measurement cell turgor (not shown). The difference between
conditions, IV- and LV-ridge cell turgor was almost interstomatal- and trough-cell turgor was on average
identical. minimal (0.01 MPa) and in none of the leaves statistically
significant. Plants were treated for 3-9 d before analysis
Table 1. Combined measurement of turgor and osmotic pressure, and inferred water potential (t/i) in cells of the upper epidermis of
barley leaves
The leaf region analysed was exposed during measurements to a by-passing stream of dry air and brightly illuminated with the
cold light source. The leaf number (L3, 4) is given with leaf developmental stage (d) expressed in days before (minus numbers) or
days following (plus numbers) full expansion. Turgor and osmotic pressure were measured from the same cell, and <p was calculated
as the difference between. Values are given as means ± S D of (n) cell analyses per cell type and plant. The absolute differences
between trough- and ridge-cell values are given in parentheses and their significance is shown at the 5% (*) and 1% (**) level.
'Means ±SD of the difference between the trough and ridge cell value (n = 9 leaf analyses).
50 Fricke
IV
1.8 Control
1.5
QL
1.2
'gor,
0.9
a 0.6
03
0.0
TR TR IS
Cell Type
Fig. 3. Turgor pressure in leaf epidermal cells of barley. Plants were analysed while being viewed horizontally, either under low (A, C) or high
PAR (B), or while being viewed vertically, with the leaf area analysed exposed to high local illumination and stream of dry air (D, E). In control
plants (A, B, D), second, third and fourth leaves were analysed between 1 d preceding and 4 d following full expansion. NaCl-treated plants (C,
E) were treated for 3-9 d prior to analysis with 100 mM NaCl and third and fourth leaves were analysed between 2 d preceding and following full
expansion. NaCl-treatments were started at the time the third leaf emerged from the surrounding second-leaf sheath by two daily additions of
50 mM NaC1. Results are presented as means of 15 (A), 8 (B), 13 (C), 9 (D), and 5 (E) leaf analyses, with 4-11 cells analysed per cell type and
leaf. The error bars shown do nol represent the SD of the means of the n leaf analyses This would represent the leaf-to-leaf variation in cell turgor.
Instead, the error bars represent the mean cell-to-cell variation in turgor for a particular cell type ( = means of the SDs of the individual leaf
analyses). The number of leaves where turgor differences between ndge and trough, and between ridge and interstomatal cells were significant is
given in relation to the total number of leaves analysed fa/b').
turgor was on average 0.15 MPa higher than trough-cell stream, i.e. evaporative-demand conditions comparable
turgor (Fig. 3D). The difference was in 8 out of 9 plants to the low PAR set-up. Ridge-cell n was on average only
significant (mostly at 1% level). In NaCl-treated 0.04 MPa higher than trough-cell n and in none of the
plants analysed under the same conditions, ridge-cell 17 leaves analysed significantly higher (Fig. 4A). In NaCl-
turgor was on average 0.28 MPa higher than trough- and treated plants analysed under the same conditions, ridge-
0.17 MPa higher than interstomatal-cell turgor (Fig. 3E). cell 7T was on average 0.19 MPa higher than trough-, and
Differences were in 4, and in 5 out of 5 plants significant 0.03 MPa higher than interstomatal-cell -n (Fig. 4B). The
(1% level), respectively. Interstomatal-cell turgor was on difference in n between ridge and trough cells and between
average 0.11 MPa higher than trough-cell turgor. This ridge and interstomatal cells was in 7 out of 8, and in 2
difference was in 4 out of 5 leaves significant (1% level). out of 8 plants significant (mostly at 1%, and 5% level),
Epidermal-cell -n: Leaves were analysed under condi- respectively.
tions of low local illumination and no by-passing air Leaves were analysed under conditions of high local
Turgor in leaf epidermal cells 51
| Low local illumination and no ilr itraim| High local Illumination and air itream
2/8
Q.
0/17
s/g
_I_
O
a.
o
o
Call Type
Fig. 4. Osmotic pressure (n) in leaf epidermal cells of barley. Plants were viewed vertically and the leaf area analysed was exposed to low local
illumination and no air stream (A, B) or high local illumination and dry-air stream (C, D) In control plants (A, C), second, third and fourth
leaves were analysed between 2 d preceding and 3 d following full expansion. NaCl-treated plants (B, D) were treated for 3-8 d prior to analysis
with 100 mM NaCI and third and fourth leaves were analysed between 1 d preceding and 4 d following fulJ expansion NaCl-treatments were
started at the time the third leaf emerged from the surrounding second-leaf sheath, by two daily additions of 50 mM NaCI. Results are presented
as means of 17 (A), 8 (B), 9 (C), and 5 (D) leaf analyses, with 4-8 cells analysed per cell type and leaf. The error bars represent the mean cell-to-
cell variation in -n for a particular cell type ( = means of the SDs of the individual leaf analyses). The number of leaves where TT differences between
ridge and trough, and between ridge and interstomatal cells were significant is given in relation to the total number of leaves analysed ('a/b').
illumination and by-passing stream of dry air. Ridge-cell treated plants (-0.70 MPa; Fig. 5B). As for control plants,
TT was on average 0.12 MPa higher than trough-cell n differences in turgor between ridge cells (highest turgor)
(Fig. 4C) and in 5 out of 9 leaves significantly higher and trough and interstomatal cells were almost entirely
(mostly at 1% level). In NaCl-treated plants analysed due to differences in TT.
under the same conditions, ridge-cell n was on average
0.24 MPa higher than trough- and 0.16 MPa higher than Short-term changes in turgor in ridge and trough cells
interstomatal-cell n (Fig. 4D). The difference in TT between
ridge and troughs cells, and between ridge and inter- Plants were kept for 24 h in a plastic bag at saturated
stomatal cells was in 5 out of 5, and in 4 out of 5 plants humidity. Upon removal of the plastic bag, i.e. sudden
significant (mostly at 1% level), respectively. drop in atmospheric RH (room RH c. 25%), both ridge
Epidermal-cell ip: Water potentials of cells were obtained and trough cell turgor dropped within minutes by about
in two ways. Either by using turgor and n obtained from 0.7 MPa (Fig. 6A). Thereafter, turgor in ridge and trough
different leaf analyses and under set-ups likely to be cells followed the same time-course of recovery and
comparable in evaporative demand (Fig. 5A) or by using reached a final value after 40-90 min, with a Tlj2 of 918 s
turgor and -n obtained from identical leaves and cells for the experiment shown (Fig. 6A). The T1/2 of the
(Fig. 5B). Calculation of </r from separate determinations immediate, exponential, decline in cell turgor following
of turgor and TT gave identical mean tfi (-0.41 MPa) for the removal of the plastic bag was 78 s (Fig. 6B, separate
ridge and trough cells in control plants (Fig. 5A); in experiment). The following recovery in turgor was accom-
NaCl-treated plants, mean ridge cell ifi (-0.76 MPa) was panied by a decrease in stomatal conductance (Fig. 6C;
0.07 MPa more positive than trough cell </. (-0.83 MPa) TU2 of 702 s; separate experiments).
and 0.12 MPa more positive than interstomatal cell </« To lower the external </i of the root suddenly, 100 mM
(-0.88 MPa; Fig. 5A). Calculation of 0 from combined NaCI was added to the medium while the plant was
determinations of turgor and TT gave nearly identical mean analysed (room RH c. 34%). Turgor started to decrease
ip for ridge (-0.31 MPa) and trough cells (-0.34 MPa) in within 10 s, in both ridge (Fig. 6D) and trough cells
control plants (Fig. 5B; see also Table 1 for data of (Fig. 6E); 10 s were needed to add all the salt in the form
individual leaf analyses) and for ridge (-0.69 MPa), of a stock solution and to readjust the meniscus in the
trough (-0.73 MPa) and interstomatal cells in NaCl- probe to its initial position. Within the following 10-12
52 Fricke
Saparat* d*ttrmtnatk>n of turgor and otmotlc p r t u u r * Combined dtUrminttton of turgor and oimotk praitur*
0.9
1M 0.9
0.6
|1 |M| 0.6
0 1 II
R TR IS R TR IS
R TR
I HI I I
IS
1
0
R TR IS R TR IS
R TR IS
-0J
III1 -OJ
-0.6
-0J
•II• -0.6
-0.9
-1.5 -13
-1.8 -1.8
-2.1 -2.1
Fig. 5. Turgor pressure, osmotic pressure (IT) and inferred water potential (<j>) in leaf epidermal cells of barley. Turgor and n data were taken from
Figs 3 and 4 (in (A) from Fig. 3A, C and Fig. 4A, B; in (B) from Fig. 3D, E and Fig. 4C, D) Water potentials were inferred in two ways Either
(A) by comparing turgor data with n data obtained from different plant batches and cells (i.e separate determinations), or (B) by comparing turgor
data with n data obtained from the identical plants and cells (combined determinations). For details of NaCl-treatments, leaf number and leaf
developmental stage see legends to Figs 3 and 4. Water potentials (and turgor and n) in B represent the means of 9 (control) or 5 (NaCl-treated)
leaf analyses, and error bars give for each cell type the means of the SDs of the individual leaf analyses.
min, turgor in ridge and trough cells decreased by about The RH was suddenly increased from 36% to >94%
0.5 MPa and finally stabilized (Tm of 226 and 170 s for RH by use of an ultrasonic humidifier. Epidermal (here:
the two experiments shown). Upon removal of NaCl, ridge) cell turgor started to increase within 2 s and
turgor started to increase within 30 s in both ridge attained within minutes a new stable value (Tl/2 of 62 s
(Fig. 6D) and trough cells (Fig. 6E); 30 s was needed to for the experiment shown; Fig. 6F). Subsequent addition
exchange media and probe a new cell. Turgor continued of NaCl caused a similar response in cell turgor as under
to increase and stabilized after about 10 min at a value 34% RH (compare Fig. 6D, E). However, T{/2 was faster
comparable to that previous to the experiment (T1/2 of (50 s) and the absolute decrease in turgor smaller than at
153 s and 215 s for the two experiments shown). 34% RH.
Fig. 6. Short-term response in epidermal-cell turgor and stomatal conductance to a sudden change in external ui. Plants were subjected under low
PAR to sudden changes in either the root-surrounding or atmospheric ip. This was achieved in three ways. Either by keeping the plant for 24 h
under a plastic bag at saturated humidity (bag with small holes for gas exchange) and then suddenly removing the bag (A, B, C; room RH c.
25%); or by adding 100 mM NaCl to, or removing it from the medium (D, E, F); or by suddenly increasing the leaf-surrounding RH from 36 to
^ 9 4 % RH through use of an ultrasonic humidifier (F). Each square represents the turgor of a separate cell, except the first 2-3 squares following
addition of NaCl in (D) and (E) and all the squares following increase in RH or addition of NaCl in (F), which were obtained from continuous
turgor measurement in the same cells. Stomatal conductance was measured with a portable photosynthesis system (LI-6200, Li-Cor, Lincoln, ME,
USA); results from four leaf analyses are plotted together, and each symbol represents the means of six determinations per leaf. Turgor relaxations
were fitted with curves (dotted lines) for a mono-exponential change plus residual to calculate 7", 2. Second or third leaves were analysed between
2 d preceding and following full expansion.
Turgor in leaf epidermal cells 53
Change in atmospheric
0.6
i- 0.4 ri T
t/i~
T t / I - 918 . Q.p
• Ridge e e l
o 0.2
• Trough ceB
0.0
•
1.5
1.0
6\ Ti/i- 702 *
0.5
o.
b- • -Q. •o- ...<P
0.0
CO -20 20 40 60 80 100
Time, min
D
+ 100 mM
1.0 I NaCI -
0.8
o ^T 1 / t " 226 i
at 0.6 153 t .
3
0.4
•i Naa
o -
NaCI
1.0 + 100 mM 1.4
a. I T 1 / I -82
•I
r
2
0.8 ODD 1.2
o \ , T 1 r t - 170 t 215 t
0.6 1.0
•ifj Q
\- 0.8 94 X
0.4 T,/ 2-50 t-
RH
Cell
0.2 t- 0.6 36 1
RH
0.0 0.4 -
0.6
c(T,/i- 109 . ©-• £ I a
0.6
- "^
S •.
0.4 0.4 E —
o, • rldga call M
a, • trough call
0.2 •
o
0^
0.0 1
Q.
*
-20 -10 10 20 30 40 50
Time, min
Fig. 7. Short-term response in turgor, osmotic pressure and water potential in leaf epidermal cells of barley to the addition of 100 mM NaCI to the
medium. Each symbol represents the turgor (A), osmotic pressure (A) and inferred water potential (B) for a separate ridge or trough cell. During
each cell analysis, turgor was measured immediately before cell sap was extracted for determination of osmotic pressure and calculation of i/>. For
better visual comparsion, curves were fitted by first order (osmotic pressure) or third order (turgor, water potential) polynoms (dotted lines); 7*1/2
was calculated from curve fits for mono-exponential decline plus residual.
et al., 1981; Steudle and Jeschke, 1983; Malone and ences in </i between cell types are transpiration-induced,
Tomos, 1992; Frensch and Hsiao, 1994). The proximity then the finding that >p is lowest in trough cells indicates
of ridge cells to stomata or bundle sheath extension did that interstomatal cells are lined at their inner walls with
not affect turgor. Similarity, trough-cell turgor showed a cuticle as observed in Tradescantia (Nonami et al., 1990)
no obvious dependency on the proximity to stomata (not and that the transpiration tension in the apoplast is highest
shown). Thus, there may be at least three 'units' of turgor in trough cells. The difference between barley and
relations within the upper epidermis of barley leaves: Tradescantia leaves concerning << / gradients may partly
the ridge-, the trough- and the interstomatal-cell unit result from the generally slower Tl/2 of water exchange in
(other peristomatal cells and guard cells were not epidermal cells of Tradescantia (Tomos et al., 1981;
considered). Tyerman and Steudle, 1982) compared to barley (Table 2).
Epidermal-cell i/i: Differences in if> between and within
non-growing leaf tissues have been reported for Short-term regulation of epidermal-cell turgor, and T1/2 of
Tradescantia and explained by the existence of transpir- water flux equilibrium
ation-induced </r gradients (Frensch and Schulze, 1988;
Nonami et al., 1990). However, in the present study there The response of epidermal cell turgor to a sudden change
were no systematic differences in 0 between epidermal cell in external </r (Fig. 6) suggested that changes in the cell's
types and regions. The difference in i/> between cells (vacuolar) solute content do not contribute to the turgor
obtained from calculation of I/I from combined analyses of response and that differences in I/I between ridge and
turgor and n ranged between 0.01 and 0.04 MPa (Fig. 5B) trough cells are at any time negligibly small. These
and was statistically non-significant. A too low transpir- assumptions were proved correct by an experiment in
ation rate as a reason for missing epidermal <p gradients is which turgor and -n (and inferred </«) were determined in
very unlikely. Even under low PAR where plants were the same leaf and cells following addition of NaCI
likely to transpire the least in the present study, the (Fig. 7). While turgor and <> / decreased by about 0.4 MPa,
'remaining' transpiration rate was still high enough to and with a similar TU2, -n remained almost constant
lower epidermal cell turgor by 0.2 MPa (compared to (<0.015 MPa increase). This also indicates that cell volu-
turgor at no transpiration; Fig. 6F). If the small differ- metric elastic modulus remained almost unchanged.
56 Fricke
The Tl/2 for local water flux equilibrium in the cells plants was only 46 s (Table 2). This shows that under the
studied was between 0.7-1.7 s (Table 2) and, thus, one conditions tested, control of stomatal aperture (water
to two orders of magnitudes smaller (faster) than the T1/2 loss) is the time-limiting process for water flux- and, thus,
for epidermal turgor relaxations at plant level (Table 2). turgor- and i/i-equilibration in the leaf epidermis of barley
This shows that water flux equilibration in the leaf (see also Fig. 6A, C). This may be a direct consequence
epidermis of barley is not time-limited by the water of the initial, non-controllable and 'counteracting', hydro-
exchange properties of epidermal cells. Falk (1966) passive movement of guard cells. The Tl/2 of 46 s is 2-17
observed that following addition of 100 mM NaCl to the times smaller than the Tl/2 obtained for excised barley
root medium of wheat, water uptake dropped within roots using the root pressure probe (Steudle and Jeschke,
seconds and recovered within the following 15 min; 1983). This points to a difference in hydraulic properties
transpirational water loss also responded within seconds, between roots of intact plants and excised roots.
slowly increased during the following 5 min and decreased The turgor of a ridge cell next (axially or laterally) to
thereafter such that, after 12-20 min, it balanced water a previously punctured ridge cell was not obviously lower
Table 2. Half times (T 1/2 ) of water flux equilibrium of (A) individual epidermal cells and (B) the hydraulic continuum between root,
xylem, leaf epidermis, and stomatal pore
Half-times of water flux equilibrium of individual cells were obtained by pressure relaxations using the pressure probe. For each
cell two Tl/2 determinations were carried out, one from endosmotic and one from exosmotic water flows. Results are presented as
means ± S D of 22-32 determinations per cell type. Third or fourth leaves were analysed, between 2 d preceding and 3 d following
full expansion, and NaCl plants were exposed for 3-9 d prior to analysis to 100 mM NaCl. Half-times of water flux equilibrium
of the hydraulic continuum between root, xylem, leaf epidermis, and stomatal pore were calculated from curves (mono-exponential
change plus residual) fitted to epidermal turgor relaxations obtained during short-term imposed changes in external 0; results are
presented as means ± S D of (n) experiments and the range of Tl/2 is given in '[ ]'.
Treatment/Experiment TUi
(B) Tu2 of hydraulic continuum between root, xylem, leaf epidermis, and stomata
Addition of NaCl to medium (c. 34% RH) 162 ±42 (6) [96; 228]
(£94% RH) 47±6(3) [45. 50]
(leaf greased)' 46(1)
Removal of NaCl from medium (c. 34% RH) 198±24 (3) [174; 216]
Increase in RH from 34 to 2: 94% 54±6(3) [48; 57]
Sudden drop in RH from 100% to 34%
Initial turgor drop 78(1)
Subsequent turgor recovery 858±156(6) [618; 1008]
'Leaf covered in stopcock grease prior to addition of 100 mM NaCl
Turgor in leaf epidermal cells 57
an increase in epidermal -n since turgor was generally at Dewar RC. 1995. Interpretation of an empirical model for
the control level (Fig. 5B; no significant difference in stomatal conductance in terms of guard cell function. Plant.
Cell and Environment 18, 365-72.
turgor between control and NaCl treatment). The latter Falk SO. 1966. Effect on transpiration and water uptake by
might be indicative of turgor regulation in NaCl-treated rapid changes in the osmotic potential of the nutrient
barley (osmotic adjustment). During short-term exposure solution. Plivsiologia Plantarum 19, 602-17.
to 100 mM NaCl, changes in epidermal turgor seemed Franks PJ, Cowan IR, Tyerman SD, Cleary AL, Lloyd J,
also to be equivalent to changes in external </f (Fig. 6D, Farquhar GD. 1995. Guard cell pressure/aperture character-
istics measured with the pressure probe. Plant, Cell and
E). However, this was rather coincidental since the recov-
Environment 18, 795-800.
ery decreased with increasing RH to 48+17% (means Frensch J, Hsiao TC. 1994. Transient responses of cell turgor
± S D of eight experiments at 94% RH; see also Fig. 6F, and growth of maize roots affected by changes in water
and Thiel et al., 1988). Assuming that the pressure change potential. Plant Physiology 104, 246-54.
in root-xylem transmitted quantitatively into a change in Frensch J, Schulze E-D. 1988. The effect of humidity and light
epidermal I/I, then the results give a radial root reflection on cellular water relations and diffusion conductance of