Plant Physiol.
(1985) 79, 891-895
0032-0889/85/79/0891 /05/$01.00/0
Measurement of the Water Potential of Stored Potato Tubers'
Received for publication June 14, 1985 and in revised form August 6, 1985
WILLIAM L. BLAND -AND CHAMP B. TANNER*
Department of Soil Science, University of Wisconsin-Madison, Madison, Wisconsin 53706 (C.B.T.)
ABSTRACT
A method of measuring the water potential of stored potato tubers
(Solanum tuberosum L.) was needed to investigate the relationship of
bacterial soft rot in tubers to water potential. Pressure chamber meas-
urements, while useful for tubers with functional stolons, cannot be made
on stored tubers. Measurements could be made on excised tissue pieces
in a hygrometer chamber and with hygrometers implanted into tubers.
We report here our evaluation of these hygrometric methods using a
comparison with the pressure chamber on tubers harvested with stolons
intact.
In tubers of high water potential, measurements on excised tissue were
as much as 0.5 megapascals lower than the pressure chamber, probably
due to turgor-driven expansion of the sample when freed from constraints
imposed by surrounding tissue. Good agreement (0.05 megapascals)
was found between the implanted hygrometer and the pressure chamber
at potentials higher than -0.5 megapascals. At lower water potentials,
both hygrometer measurements were higher than the pressure chamber.
Respirational heating of the tissue contributed to the increase in the
excised tissue samples, but not with the implanted hygrometers because
of the hygrometer design. The osmotic pressure balaced the pressure
chamber measurement of potential at -0.7 megapascals, but was too
small to do so at lower potentials. At most, 25% of this discrepancy can
be accounted for by dilution by apoplastic water. We believe that the
pressure chamber measurement is too low at low water potentials and
that the error is associated with air bubbles in the xylem. At low potentials
air emerged from xylem vessels along w-ith sap, and fewer xylem emitted
sap as potentials decreased.
An unambiguous measurement of the water potential of stored
potato tuber tissue is needed to investigate further an apparent
relationship between water potential and the susceptibility of
stored tubers to attack by soft rot erwinias (15). Methodology
developed should also be of value in studying the water relations
of other fleshy plant storage tissues, particularly for studies of
mechanical damage and postharvest pathology (6). Such storage
organs are an important part of the world food supply and
measurements of their water relations have received inadequate
attention.
Early measurements of potato tuber water potential were made
by liquid-phase equilibrium (18). Later, soil psychrometers were
inserted into tubers attached to plants in the field (5) and the
pressure chamber was used with freshly harvested tubers having
intact stolons (1 1). Gandar and Tanner ( 11) compared pressure
chamber measurements with liquid-phase equilibrium and with
soil psychrometers inserted into tubers attached to plants and
' Research supported by the College of Agricultural and Life Sciences,
University of Wisconsin-Madison, and by United States Department of the pressure chamber, in situ hygrometry, and excised tissue hygrometry,
Agriculture Hatch formula funds (Project 142-5140).
891
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found poor agreement with liquid-phase equilibrium and fair
agreement with psychrometers at potentials > -0.6 MPa, al-
though poor thermal equilibration between psychrometer and
tissue degraded their measurements. The validity of the pressure
chamber measurement was indicated by the agreement with
psychrometers and also by comparable leaf and tuber water
potentials down to -0.7 MPa provided transpiration was low
(10).
The pressure chamber cannot be used to measure the water
potential of stored tubers because these tubers lack a conductive
stolon. This article reports on two methods suitable for stored
tubers which were evaluated by comparison to the pressure
chamber on fresh tubers. The first measures the potential of
excised tissue in a hygrometer chamber, as is frequently done
with leaves. With the second, specially designed in situ hy-
grometers are inserted into the tuber.
MATERIALS AND METHODS
Tuber Material. Potato plants (Solanum tuberosum L. cv
Russet Burbank) growing in the field were sheltered from water
inputs to obtain a range of water potential. Also, entire plants
were harvested and spread on tables under continuous light and
circulating air, with the tubers covered with black plastic. Water
loss through the foliage lowered the tuber water potential. Tubers
also were measured hygrometrically immediately after field har-
vest (2-3 weeks after vine-kill) and after storage from 1 to 18
months in several storage environments.
Pressure Chamber. The pressure chamber and procedure de-
scribed in Gandar and Tanner (10) were used except that a lower
rate of pressurization was found necessary when 4,2 < -(.7 MPa.
Pressurization rates of 2 kPa s' or less gave the same endpoint
with repeated pressurization. Tubers were held for at least 16 h
in a plastic bag or a chamber lined with damp blotters before
measurements were made. This delay was to allow for dissipation
of water potential gradients near the xylem system (9, 10); we
verified that the delay was adequate to bring the tuber to uniform
potential by repeated pressurizations following removal of tubers
from the vine. To separate xylem and phloem exudation (10) for
measurement of the osmotic pressure of xylem sap and to
confirm xylem blockage and air entrainment in xylem sap, we
picked away the cortex, phloem, and pith at the cut end of the
stolon so that the xylem vessels extended beyond other tissue.
Hygrometry. Hygrometric measurements were made with a
Wescor HR-33T microvoltmeter (Wescor, Logan, UT) operated
in the dewpoint mode. The meter was modified with an auxiliary
rechargeable battery pack and an integrating digital voltmeter
attached to the recorder outputs. The mode switch was replaced
with a six-position, four-pole (two poles in parallel), gold-plated
contact rotary switch (Centralab, Fort Dodge, IA). A sixth mode,
'read reverse', was added so the zero offset, zo, could be read in
2 Abbreviations: n, 4{i, 4*,, tuber tissue water potential as measured by
respectively; w, osmotic pressure; zo, zero offset, dry-bulb voltage.
892 BLAND AND TANNER Plant Physiol. Vol. 79, 1985
the reverse from normal polarity. -_ _FLEX::*** FLEXANE 94
The zo arises from two sources: the bias current of the meter .25mm
amplifier (40-50 namps) through the hygrometer junction and EVANOHM LEAD
leads and from temperature differences between the hygrometer COPPER HEAT SINK
reference and measuring junctions. This measured temperature
difference reflects a temperature difference between the reference SUPPORT PIN
junction and the vapor source. Reversing the polarity when
reading zo changes the sign of the thermal component but not
that of the bias current contribution, so by reading zo in both
polarities the thermal portion can be determined. All zo values
reported here represent thermal effects only. I
Excised tissue samples were measured in a 10-sample hygro- 0

meter (model SC-10 with a Peltier thermocouple; Decagon De-

vices, Pullman, WA). We polished the inside surfaces of the
unpolished stainless steel sample holders to speed equilibrium
(present Decagon holders are polished). We also washed the
sample holders with 20% NH4OH and cotton swabs, rinsed with
distilled H20, and dried them with lint-free wipes prior to each
use. Cylinders of tissue (13 mm diameter x10 mm long) were
excised at the xylem ring, parallel to the major axis of the tuber
with a sharp, thin-wall, guided sampling tube. The cores were
lightly blotted with lint-free tissue immediately after excision.
Rinsing the samples with distilled water immediately after exci-
sion and blotting dry had no effect on the potential measured.
Two samples were taken from each tuber, either both from the
tuber midsection or one each from the apical and basal thirds of
the tuber, and their values averaged. The mean difference be-
tween duplicate samples was 0.04 MPa, believed to be random
error. Calibration solutions (usually -0.45 and -0.90 MPa) equal
in volume to the tissue cores were included with each set of
samples. The SC-10 was kept under an insulated box in the
laboratory (about 22C) when in use except during collection of
the 1983 fresh tuber data, when it was mounted inside of a
constant temperature air bath (25C). Knobs needed to operate
the hygrometer were extended outside of the bath with phenolic
rods. Thermal equilibrium in the hygrometer body was checked
by measuring zo over a calibration solution. In order to ensure
vapor and thermal equilibrium, 4e was recorded 2 h after the
samples were excised and placed in the hygrometer. At this time
the zo of fresh tuber samples was 0.20 0.08 MV (tissue warmer
than reference junctions); between 2 and 5 h after excision zo
increased by about 0.16 MV. Samples measured at field harvest
and during storage had average zo's of 0.10 and 0.03 MV, respec-
tively. No dependence on water potential was observed in the zo
values. Determinationsof 4.,e also were made using Wescor C-52
hygrometers; tissue cores 8 mm in diameter and 4 mm thick
were measured.
Measurements of water potential were also made with hygro-
meters placed in close-fitting holes drilled axially in tubers. Two
hygrometers were designed for implantation into tubers, one
with a polyethylene body and the other with a stainless steel
body. The stainless steel unit (Fig. 1) was designed to lock the
hygrometer and the tissue together thermally, minimizing error
caused by temperature gradients. To minimize heat flow along
the leadwires, 12 cm of 0.25-mm (30 gauge) Evanohm3 wire with
12 Am thick polyurethane varnish was used for leads, crimped
into holes drilled in the heat sinks. The wires were extended by
crimping them to stranded copper wire. The hygrometer meas-
uring junction was welded from 25 Mm chromel and constantan
wire and crimped into notches at the ends of the heat sinks. A
0.076 mm Evanohm/constantan thermocouple glued into a
small cavity in the brass core was used to measure the hygrometer
temperature. Snug-fitting holes for the hygrometers were cut in
3 Evanohm wire is available from Pelican Wire Co., Naples, FL. onto a disc of
Relative to copper, it has an electromotive force of only 0.2 ,V/'C, but plicate samples were measured with different hy-
lower thermal conductivity and higher tensile strength.
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0 0 THERMOCOUPLE
lO '--VAPOR PORT
-r-6
16 mm_
,-TEFLON PLUG
FIG. 1. Cross-section of hygrometer designed for implantation into
potato tubers.
the tuber with a twist drill followed by a corkborer-like cutter
with its cutting edge to the outside. After inserting the hygro-
meter, the juncture of the periderm and the hygrometer was
covered with petroleum jelly, the Evanohm leads were taped to
the tuber surface, and about 15 cm of lead was coiled against the
tuber. The exposed end of the hygrometer and adjacent leads
then were covered with 3-mm thick polyethylene foam and
wrapped in aluminum foil. Measurements were madedata in a con-
stant temperature air bath (25C).The stored tuber
collected with both polyethylene and stainless steel units; the
were
fresh and just-harvested tuber data were collected
with the stain-
less steel hygrometers. With fresh tubers, pressure chamber meas-
urements were made both before hygrometers were inserted and
after they were removed to test for shifts in water potential; 14
changed -0.04 0.05 MPa between the two determinations. In
comparisons {i of and 0, the second pressure chamber value
was used because it was made immediately following {i,
the
measurement. Steady values of 4i (0.01
MPa) obtained 8 to 12
h after implanting the hygrometers. Readings at 8 h were within
0.03 MPa ofthe final value. Adjustment ofthe tuber temperature
to that of the constant temperature box was the limiting process
in equilibration.
Once the output was steady it remained so for at least 30 h.
To test whether the cut surface was still exchanging vapor with
the hygrometer, a hygrometer was removed and replaced with
another after the first had been implanted 22 h. The second
hygrometer was read 4 h after implanting and indicated the same
water potential (within 0.02 MPa) as the first hygrometer.
The osmotic pressure of the tubers used in the comparison
also was determined. Duplicate tissue cores (8 mm diameter, 12
mm length) were cut from the xylem ring and put in a 3-ml
polyethylene syringe. To prevent the tissue from plugging the
syringe orifice, a disc of stainless steel screen (IO00-m
pores) had
been placed at the needle-end of the syringe. The plunger was
replaced, the needle inserted in a rubber stopper, and the syringe
was frozen in dry ice and kept in a freezer until measured.
Measurement of r was made by thawing the tissue in the sy-
1
ringe at room temperature for h, then expressing the The exudate
filterpaper in a Wescor C-52 hygrometer.
du-
grometers; duplicates differed, on average, by 0.0 16 MPa. In-
 WATER POTENTIAL OF STORED POTATO TUBERS 893
creases in r due to enzymic action following thawing (3) were NV (MPb)
not observed, whether the cell sap was separated quickly after
rapid thawing or allowed to remain in contact with structural
tissue components. We tested for possible enzymic changes in r
during storage of samples in the freezer for as long as 16 weeks
(7). Decreases in r averaging 0.013 MPa week-' occurred; ac-
cordingly, samples reported here were typically stored no more
than 4 d.
Frequently, the ir of the cell sap freed when tissue samples
were excised was measured to compare with that of frozen tissue.
Sap was collected by touching a filter paper disc to a freshly cut
tissue surface, which was then placed in a Wescor C-52 hygro-
meter. To reduce evaporation, these operations were done in a
box lined with damp blotters. The osmotic pressure measured
on frozen and thawed tissue was the same as that of the sap freed
when tissue was cut. For 51 samples of both fresh and stored
tubers with water potentials from -0.58 to -1.53 MPa, r (frozen/
thawed) averaged only 0.005 MPa higher (drier) than ir (cut cell
sap); this was not significant at P < 0.40.

RESULTS AND DISCUSSION

FIG. 3. Comparisons of the water potential of freshly harvested potato
Comparisons among the pressure chamber, excised tissue, and tubers as determined by the pressure chamber (4,6) and in situ hygrometers
implanted hygrometer measurements of tuber water potential (4's). The sy, is the standard error of the estimate.
are presented in Figures 2 to 4. Osmotic pressure as a function
of A1 is presented in Figure 5. ejz (MPG)
The pressure chamber values presented in this paper have not
been corrected for the osmotic pressure of the xylem sap, as is
thought necessary for pressure chamber values to yield total
water potential (1). We found the osmotic pressure of the xylem
sap by collecting xylem exudate on filter paper discs and meas-
uring these in Wescor C-52's psychrometrically. The mean of
several tubers was 0.10 MPa, so correcting the chamber meas-
urements would shift all data points in Figures 2, 3, and 5 to the
left, worsening the agreement.
Excised Tissue Measurements. For pressure chamber meas-
urements ranging from 0 to -0.3 MPa, the excised tissue water
potentials were -0.25 to -0.35 MPa, independent of 0, (Fig. 2).
The best-fit straight line describing l, as a function of 46^ for
pressure chamber values less than -0.3 MPa (Fig. 2) had a slope
%(MPO)

FIG. 4. Comparisons of the water potential of potato tubers as deter-

mined by excised tissue hygrometry (4e,) and in situ hygrometers (4,). For
{i < -0.3 MPa, the standard error of the estimate of 4' is 0.06 MPa.
of 0.7 (not equal to 1.0 at P < 0.001).
Figure 2 also shows the straight line fit (slope = 0.42) through
21 comparisons (individual points not shown) of 4,, and excised
tissue samples determined with the C-52 hygrometer, data were
evenly distributed along the line. In samples with high turgor, C-
52 measurements were as much as 0.5 MPa below pressure
chamber measurements. Small zo's were recorded in the C-52.
The C-52's design (thermocouple wires only about 1 mm long
[4], and the measuring junction recessed in the metal hygro-
meter body) reduces the zo response to thermal gradients between
the vapor source and the reference junction. However, this does
FIG. 2. Comparisons ofthe water potential of freshly harvested potato not reduce associated errors.
tubers as determined by the pressure chamber (4) and excised tissue We believe the unduly low 4,e at high potentials (Figs. 2, 4)
hygrometry (he). Points and regression line are shown for data collected was caused by turgor-driven expansion of the samples following
with the SC-j0 hygrometer, only the regression line for the C-52 data is their excision. Excision removed constraints imposed on the
shown. The s,, is the standard error of the estimate. sample by surrounding tissue in the intact tuber, allowing the
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894 BLAND AND TANNER
0.5-
-9.0 -0.5
Wc (MPo)
FIG. 5. Osmotic pressure (w) of freshly harvested potato tuber tissue
measured by hygrometry of sap from frozen and thawed tissue as a
function of the tuber water potential measured by the pressure chamber
(14). The line indicates where 4 =
expansion. This expansion occurred at the expense of turgor,
lowering e,. In a plot such as Figure 2 this would tend to push
the data points downward from the 1:1 line where turgor exists
(4c > -0.7 MPa; Fig. 5). The smaller C-52 samples yielded lower
41e values than did the SC-10 samples at all potentials where there
was turgor, as expected for smaller samples on release of con-
straints.
Turgor-driven strain does not affect in situ measurements
because of the large bulk of tissue surrounding the implanted
hygrometer. The small cylindrical cavity cut for the hygrometer
would minimize strain. Also, radial geometry assists water flow
and dissipation of potential gradients near the cavity. Finally,
the tuber tissue volume provides a large capacitance for water,
minimizing the impact of the coring on the tissue.
During the measurements reported here, the excised tissue
samples were isothermal with or warmer than the hygrometer
body, so thermal errors were in the opposite direction to expan-
sion errors. Tissue warming, as indicated by the zo in the SC-10,
ordered fresh > harvest > stored as expected for respiratory
heating (20). The size of respirational errors in y6 can be esti-
mated from zo in the SC-10. The zo underestimates the temper-
ature gradient between the reference and the tissue because the
measuring junction is 0.43 of the distance between them and
because of divergent heat flow geometry. Correcting for the
distance of the measuring junction from the tissue and recogniz-
ing that the measured zo was removed electronically before
making the dewpoint measurement, a minimum error of 0.04
MPa obtains for the fresh tubers. This value is in good agreement
with the displacement of the fresh tuber data from the 1:1 line
in Figure 4; the lower respiration rate of stored tubers did not
cause a similar displacement. This fresh tuber respirational error
is also present in Figure 2. As reviewed by Laties (16), respiration
increased 3- to 5-fold immediately after cutting the tissue and at
least another 2-fold within 8 h. The long-term increase in respi-
ration was inversely related to slice thickness so the error in the
C-52 measurements should have been greater than with the SC-
10 samples.
In the in situ measurement, the hygrometer body makes good
thermal contact with the tissue, whereas the excised samples
make poor contact with the holder walls to avoid tissue com-
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Copyright 1985 American Society of Plant Biologists. All rights reserved.
Plant Physiol. Vol. 79, 1985
pression. Accordingly, errors from respiratory heating are re-
duced with implanted hygrometers as compared to measure-
ments on excised tissue. Additionally, the implanted hygrometer
is surrounded by tissue into which wound-stimulated respiratory
heat could dissipate.
The possible degradation of membranes following wounding
ofpotato tuber tissue (16) could impact #e and fj. This apparently
did not occur to a great extent during our measurements since
the cells maintained appreciable turgor at high water potentials.
This and other physiological reactions to wounding should have
had more effect on the excised tissue measurement than on the
in situ because of the large bulk of intact tissue surrounding the
implanted hygrometer. It is known that when excised potato
tuber discs are reimplanted into intact tubers, some wound
responses of the discs are attenuated (21).
In comparisons with the pressure chamber, the excised tissue
measurement (Fig. 2) was more scattered than the implanted
hygrometer measurement (Fig. 3). This was probably due to
variable effects of turgor-driven expansion, respiration, and
wound responses.
Implanted Hygrometer Measurements. The implanted hy-
grometer measurements agreed well with the pressure chamber
at high potentials (Fig. 3). At low potentials, 4#i was higher than
A; the second-order term of the regression in Figure 3 is signifi-
cant at P < 0.01. This departure from the pressure chamber
measurement at low potentials also occurred with the excised
tissue measurement (Fig. 2) and when osmotic pressure was
determined (Fig. 5). Respiratory heating of the tissue explains
only a small amount of the discrepancy between Ac and 4JJ and
is negligible with Oi. The negative turgor calculable from the data
in Figure 5 seems unlikely in potato tuber tissue at the level of
hydration of these tubers.
Dilution of the symplasm solutes by apoplastic water in the
samples may have contributed to apparent negative turgor (2,
23). In potato tuber tissue, however, there does not appear to be
enough apoplastic water to provide the necesary dilution. To
cause a decrease in r of 0.1 MPa at 0, = -1.0 MPa (Fig. 5), the
apoplastic water must be 10% of the total tissue water. The
apoplastic water present can be estimated in two ways: first by
the amount of water in the wall and the amount of wall in the
tissue and secondly by the fraction of cell volume that is com-
prised by wall. We measured the water retained by isolated potato
tuber cell walls (14) at various water potentials with a soil pressure
plate apparatus. The weight fractions of water (dry wall weight
base) at various potentials were: 0.90 at -1.2 MPa, 1.08 at -0.8
MPa, 1.25 at -0.4 MPa, and 1.60 at -0.2 MPa. Since these are
overestimates due to water retained in menisci between the wall
particles, we can use 1.00 as the upper limit for water weight
fraction and 0.40 as a lower limit (that of never-ried wood;
[22]). Weight fractions can be converted to volumetric fractions
given the specific gravity of dry cell wall, about 1.55 (8, 22).
Measurements of the dry wall weight fraction in fiesh tissue (0.01
to 0.02; [13, 14, 17]) and the water content of the tubers (0.78-
0.80 of fresh weight) lead to apoplastic water fractions between
0.005 and 0.025. The volume fraction of wall in tissue can be
estimated from wall thickness and cell size observed in cut tissue.
Data of Reeve et al. (19) give wall volume fractions between 0.02
to 0.05 for stained tissue and we measured values of 0.02 to 0.04
with unstained, fresh tissue. Data reported by Hayden et al. (12)
yield a value close to 0.05. Since a tuber is about 0.86 water by
volume, apoplastic water fractions of 0.009 to 0.035 obtain.
Dilution by apoplastic water can account for 35% of the discrep-
ancy at most, and 20 to 25% is more likely.
Contrary to pressure chamber measurements, Oi and -r meas-
urements gave reasonable turgors for fresh and stored tubers:
about 0.1 MPa at Ai = -0.7 MPa decreasing to 0.0 at -1.0 MPa.
Correction for the dilution error in xwould increase these turgor
 WATER POTENTLAL OF STORED POTATO TUBERS
estimates slightly. This protracted decrease in turgor at low
potential may be due to variation in water potential components
or bulk modulus of elasticity within the tissue, as evidenced by
the development of sponginess at the basal end of tubers during
water loss before the entire tuber is spongy.
From the above osmotic pressure arguments we believe the
difference between the pressure chamber and the hygrometer
measurements at potentials < -0.7 MPa mainly was due to error
in the pressure chamber measurement. Errors due to rapid
pressurization and to lack of water potential equilibrium within
the tuber were guarded against, as described earlier. We believe
that the low pressure chamber measurements at low potentials
were in some way associated with air entering the xylem of
stressed tubers. When the xylem at the end of the stolon was
separated from the surrounding tissue to permit observation, the
number of xylem exuding sap decreased as the potential de-
creased below about 0.7 MPa. We believe, as observed by Tyree
et al. (24), that as the potential decreased, the water in an
increased number of xylem elements fractured under tension
and air diffused into the xylem. This air embolism resting against
perforation plates would block return of sap. In making duplicate
determinations of the chamber end point, the pressure never was
decreased rapidly nor more than about 0.1 MPa; accordingly,
the embolism would have been created prior to the chamber
measurement. We also observed air bubbles entrained in the sap
and the number of vessels with entrained air increased as the
potentials decreased.
Good measurements of tuber water potential appear to be
provided by the in situ hygrometers over the range of water
potentials encountered under field or storage conditions. Meas-
urement with excised tissue from stored tubers of water potential
< -0.4 MPa are acceptable provided the SC-10 hygrometer, with
its large sample holders, is used. Pressure chamber measurements
are satisfactory with fresh tubers at potentials > -0.7 MPa, but
error obtains at lower potentials. Dilution by apoplastic water
affects measurements of wr little, since only about 0.025 of the
water in potato tubers is apoplastic. Sap collected from cut
surfaces yields the same value of X as obtained by killing the
tissue by freezing.
LITERATURE CITED
1. BOYER JS 1969 Water status measurements in plants. Annu Rev Plant Physiol
20: 351-364
Downloaded from on September 28, 2017 - Published by www.plantphysiol.org
Copyright 1985 American Society of Plant Biologists. All rights reserved.
895
2. BOYER JS, JR POTTER 1973 Chloroplast response to low leaf water potentials.
I. Role of turgor. Plant Physiol 51: 989-992
3. BROWN PW, CB TANNER 1983 Alfalfa osmotic potential: A comparison of the
water-release curve and frozen-tissue methods. Agron J 75: 91-93
4. CAMPBELL GS 1979 Improved thermocouple psychrometer for measurement
of soil water potential in a temperature gradient. J Phys E Sci Instrum 12:
739-743
5. CAMPBELL MD 1972 The lower limit of soil water potential for potato growth.
PhD thesis. Washington State University, Pullman
6. COOK RJ, RI PAPENDICK 1978 Role of water potential in microbial growth
and development of plant disease, with special reference to postharvest
pathology. HortScience 13: 11-16
7. FENNEMA 0 1975 Activity of enzymes in partially frozen aqueous systems. In
R Duckworth, ed, Water Relations of Foods, Proc. Int. Symp. in Glasgow,
1974. Academic Press, London
8. FREY-WYSUNG A 1976 The Plant Cell Wall. Gebruder Borntraeger, Berlin
9. GANDAR PW 1975 Growth and water relations in potatoes (Solanum tubero-
sum L.). PhD thesis. University of Wisconsin-Madison
10. GANDAR PW, CB TANNER 1976 Potato leaf and tuber water potential meas-
urements with a pressure chamber. Am Potato J 53: 1-14
11. GANDAR PW, CB TANNER 1975 Comparison of methods for measuring leaf
and tuber water potentials in potatoes. Am Potato J 52: 387-397
12. HAYDEN RI, CA MoYsE, FW CALDER, DP CRAWFORD, DS FENsoM 1969
Electrical impedance studies on potato and alfalfa tissue. J Exp Bot 20: 177-
200
13. HOFF JE, MD CASTRo 1969 Chemical composition of potato cell walls. J Agric
Food Chem 17: 1328-1331
14. JARVIS MC, MA HALL, DR THREFALL, J FRIEND 1981 The polysaccharide
structure of potato cell walls: Chemical fractionation. Planta 152: 93-100
15. KELMAN A, JW BAUGHN, EA MAHER 1978 The relationship of bacterial soft
rot susceptibility to water status of potato tuber. Phytopathol News 12: 178
Abstr 236
16. LATIES GG 1978 The development and control of respiratory pathways in
slices of plant storage organs. In G Kahl, ed, Biochemistry of Wounded Plant
Tissues. W DeGruyter, Berlin
17. MCGUIRE R 1983 Reduced severity of Erwinia soft rot in potato tubers with
increased calcium content. PhD thesis. University of Wisconsin-Madison
18. MEYER BS, AM WALLACE 1941 A comparison of two methods of determining
the diffusion pressure deficit of potato tuber tissuei Am J Bot 28: 838-843
19. REEVE RM, H TIMM, ML WEAVER 1973 Cell wall thickness during growth of
domestic and foreign potato cultivars. Am Potato J 50: 204-211
20. SCHIPPERS PA 1977 The rate of respiration of potato tubers during storage. 1.
Review of the literature. Potato Res 20: 173-188
21. SMrrH BG, PH RUBERY 1981 The effects ofinfection by Phytophihora infestans
on the control of phenylpropanoid metabolism in wounded potato tissue.
Planta 151: 535-540
22. STAMM AJ 1971 Review of nine methods for determining the fiber saturation
points of wood and wood products. Wood Sci 4: 114-128
23. TYREE MT 1976 Negative turgor pressure in plants: Fact or fallacy? Can J Bot
54: 2738-2746
24. TYREE MT, MA DIXON, EL TYREE, R JOHNSON 1984 Ultrasonic acoustic
emissions from the sapwood of cedar and hemlock. Plant Physiol 75: 988-
992