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GANDARAND TANNER: MEASURING WATER POTENTIALS

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COMPARISON OF METHODS FOR MEASURING L E A F AND

T U B E R WATER P O T E N T I A L S IN POTATOES 1
P. W. Gandar and C. B. Tanner 2
Abstract

Comparative methods for measuring water potential in leaves and

tubers of potatoes (Solanum tuberosum L., var. Russet Burbank) are
described.
For leaves drier than -3 bars, the pressure chamber gave estimates of
water potential which were zero to three bars drier than potentials measured using thermocouple psychrometers. Pressure chamber readings
ranged _+ 2.5 bars from psychrometer value for leaves wetter than -3 bars;
the psychrometer measurement usually was drier than the pressure
chamber when leaves were sampled in the evening.
With t u b e r s , w a t e r p o t e n t i a l m e a s u r e m e n t s using in situ
psychrometers and the pressure chamber agreed to within one bar, except
in tubers drier than -7 bars, where there were discrepancies o f _+ 2.5 bars.
However, if the interval between psychrometer insertion and water potential measurement were longer than 24 hours, serious errors arose in the
psychrometer measurements, apparently from suberization of tissues surrounding the psychrometers which prevented vapor equilibrium.
Pressure chamber measurements of tubers also were compared with
tuber water potentials determined from the weight changes of tuber samples immersed in graded sucrose osmotica. For tubers wetter than -5 bars,
the osmotica method gave drier estimates than the pressure chamber; for
tubers drier than -5 bars the reverse occurred. Causes for these discrepancies are discussed.
Resumen

Se describen m6todos comparativos para medir potenciales de agua en

hojas y tub6rculos de papa (Solanum tuberosum L., var. Russet Burbank).
Para hojas contenidos de humedad menores que -3 bars, la cfimara de
presi6n di6 estimados de potenciales de agua que estuvieron entre cero y
tres bars por debajo de los potenciales medidos usando psic6metros de
termocupla. Las lecturas de la cfimara de presi6n se desviaron+ 2.5 bars de
los valores del psic6metro para hojas mils h6medas que -3 bars; las medidas
del psic6metro di6 contenidos de humedad mils bajos que la cfimara de
presi6n cuando las muestras de hojas se tomaron al anochecer.
1Contribution from the Department of Soil Science, University of Wisconsin. Research
supported by the Collegeof Agricultural and Life Sciences, Universityof Wisconsin, Madison, Wisconsin, 53706, and by USDA Hatch funds.
2Research Assistant (presently Scientist, D.S.I.R., Palmerston North, New Zealand), and
Professorof Soil Science,Universityof Wisconsin, Madison,Wisconsin,53706.Receivedfor
publication April 14, 1975.

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En tubrrculos, las demidas del potencial de agua usando psicdmetros

en situ y la c/tmara de presirn concordaron dentro del limite de 1 bar,
excepto en tubrrculos con humedad menor que -7 bars, donde hubieron
discrepancias de ___2.5 bars.
Sin embargo, si el intervalo entre la inserci6n del psic6metro y la
medida del potencial de agua fue mayor de 24 horas, serios errores surg i e r o n en las medidas del psic6metro, a p a r e n t e m e n t e debido a la
suberizaci6n de tejidos alrededor del psic6metro, impidiendo asi al equilibrio del vapor.
Las medidas de la cfimara de presi6n para tubrrculos tambirn se
compararon con potenciales de agua determinados por el cambio de peso
de las muestras de tub4rculos sumergidos en soluciones osmdticas de
sucrosa. Para tubrrculos con contenidos de humedad mayores que -5 bars,
el mrtico dio estimados de humedad mils bajos que la cfimara de presirn; 1o
opuesto ocurri6 en tubrrculos menos ht~medos que -5 bars. Se discuten las
causas de estas discrepancias.

Introduction

Studies of crop response and growth during periods of water deficits

require measurement of'plant water status. Few measurements of water
status in leaves or tubers of potato have been reported. In a few experiments on potato, leaf relative water contents have been used as an index of
response to changes in soil water supply or atmospheric evaporative demand (16, 27, 29, 32). In other cases, pressure chambers (19, 26), thermocouple psychrometers (11,27), and in situ hygrometers (3, 10) have been
used to measure leaf water potentials (qJl). The only comparisons among
any of these methods for potato leaves were made by Baughn and Tanner
(3, 4). In their experiments, made indoors under low transpiration conditions, the pressure chamber gave estimates of qJ~ which were within +_ 1.5
bars of values measured using an in situ hygrometer, and psychrometer
measurements which were 0 to -2 bars drier than the hygrometer. Similar
comparisons, under field conditions have not been reported for potatoes.
Tuber water status has been measured in only a few experiments.
Epstein and Grant (16), following the method of Meyer and Wallace (23),
determined tuber water potential (Wt) from the weight changes of plugs of
tuber tissue immersed in sucrose solutions with different osmotic potentials. Tuber water potentials have also been measured with soil psychrometers sealed in holes drilled in tubers (11), and with a pressure
chamber (19). No comparisons have been made between these methods.
The purpose of this paper is to provide comparisons of different
methods for measuring qJ~ and Wt in potatoes. Comparative measurements
of water potential in leaves were made using a pressure chamber and

1975)

GANDARAND TANNER: MEASURING WATER POTENTIALS

389

thermocouple psychrometer; in the case of tubers, we compare the pressure chamber method with graded osmotica determinations, and with in
situ soil psychrometers.

Materials and Methods

Plant material--Leaves and tubers from Russet Burbank potatoes

grown in a glasshouse and in the field at Hancock, Wisconsin (18) were
used in these experiments.
L e a f measurements--Pressure chamber and psychrometer measurements were made on leaves from field-grown plants. Young (third to fifth
from top) and old (tenth to fifteenth) leaves were washed and dried on
-plants, prior to excision, to remove contaminants which might affect psy=
chrometer readings (15). Measurements of leaf xylem water potential, qJ~x,
were made with a pressure chamber as described by Gandar and Tanner
(19). Once qJ~xhad been measured, leaves were removed from the chamber
and sampled for thermocouple psychrometer determinations of qJ~, as
described by Baughn and Tanner (4).
Tuber m e a s u r e m e n t s - - T h e c o m p a r i s o n b e t w e e n the p r e s s u r e
chamber, graded osmotica, and in situ psychrometer methods were made
using glasshouse-grown plants. The tubers used in these experiments were
50 to 90 days old (from initiation). Their lengths ranged from five to ten
centimeters.
Soil psychrometers (Wescor, Inc., Logan, Utah, 84321) were inserted
into slightly undersized holes drilled in the apices or sides of tubers without
detaching them from plants. Holes usually were rinsed with distilled water
and swabbed dry before psychrometers were inserted. Psychrometers
were sealed into tubers with a low melting-point wax. To minimize errors
due to heat conduction, leads were coiled against the tubers. Tubers were
then buried under sphagnum moss. Tuber water potentials were measured
18 hours to 10 days after psychrometers had been inserted, usually in the
early afternoon. Once q6 had been determined with the psychrometers,
tubers were excavated and their xylem water potentials, qJ~x, were measured in the pressure chamber (19). Tubers were then sampled for graded
osmotica determination of tuber water potential.
We followed the method of Meyer and Wallace (23) for the graded
osmotic measurements. Plugs 20 mm in length and 5 mm in diameter were
cut from a tuber, rinsed in distilled water, and dried by gentle blotting
between filter papers. After weighing, plugs were equilibrated in 0. I to 0.6
M sucrose solutions for four hours at 5 C. Plugs were then rinsed in distilled
water, dried, and reweighed. Tuber water potential was found by interpolating to the sucrose solution osmotic potential at which the weight of
immersed tissue samples remained constant.

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Results and Discussion

Pressure chamber vs. psychrometers for leaves~The comparison of

~, measured with the pressure chamber, and qJl, obtained using psychrometers, is shown in Fig. 1. Data are presented as the difference A ~ ,
between tlJixand W~. Positive A ~ indicate that the pressure chamber estimate, W~x,is drier than the psychrometer value, W~. Data are plotted against
W~xon the abscissa; since both qJ~xand qJ~ are subject to error, tlh might
equally well have been used. Data have been separated according to the
time of day in which samples were collected. We shall discuss the daytime
results first, and then the morning and evening data.
The pressure chamber gave drier readings than the psychrometer for
leaves collected during the day. This discrepancy would appear even larger
if chamber values were corrected for the 1-bar osmotic potential of the
xylem sap (3). Pressure chamber measurements which are drier than those
of psychrometers have been observed in a number ofother species (2, 7).
However, comparisons of psychrometers and hygrometers, and of hygrometers and the pressure chamber by Baughn and Tanner (3, 4) suggest
that psychrometer readings should be drier than pressure chamber readings for dry potato leaves. The difference between their data and ours may
reflect differences between glasshouse-grown and field-grown plants.
However, it is equally likely that psychrometer-pressure chamber differences resulted from our measurement procedure. Psychrometer and pressure chamber measurements were made on the same leaves. From excision
until insertion of samples in psychrometers, leaves were covered with a
damp cloth to prevent evaporation. This procedure did not affect the
pressure chamber readings (19). However, if moisture from the cloth
permeated the cuticle or remained on the surfaces of leaves, psychrometer
readings would have been wetter than Baughn and Tanner's results. Apparently this occurred, for the psychrometers approached equilibrium
from the wet side rather than the dry, as is usually the case. This systematic
wetting appeared in all our psychrometer measurements, including those
for morning and evening leaves. However, had we eliminated this error by
discarding the damp cloth, the pressure chamber-psychrometer comparison might not have been improved, for vapor sorption by psychrometer
walls and cuticle causes measurements to be too dry (4, 24). All other
errors, such as evaporation and sap loss, which would cause dry pressure
chamber readings, should produce similar shifts in the psychrometer
measurement.
In contrast to the daytime data, the high water potential data during
morning and evening shows both positive and negative AtlJ (Fig. 1). Some
of this scatter arises because pressure chamber endpoints are less clear,
and psychrometer measurements less reliable, at high water potentials.
Part of the scatter may also arise from growth effects in the psychrometers.

1975)

391

GANDAR AND TANNER: MEASURING WATER POTENTIALS

*4

*2

#,

Hancock, Wis.
1973

<I
o
o

-2

tx

-4

I
0

-2

-4

PRESSURE-CHAMBER

Morning, old leaves

Morning, young leaves
Daytime leaves
Evening, old leaves
Evening, young leaves
I

-6
POTENTIAL

-8

-IO

(bars)

F I G . 1. C o m p a r i s o n of p r e s s u r e c h a m b e r and t h e r m o c o u p l e p s y c h r o m e t e r m e a s u r e m e n t s of
leaf w a t e r potential. A q j = qJ~-W~x= W ( p s y c h r o m e t e r ) _tp ( p r e s s u r e c h a m b e r ) . Positive AlP
indicate a drier p r e s s u r e c h a m b e r estimate.

In separate experiments (18), we have established that nearly all the expansion growth of well-watered, field-grown potato leaves occurs in the late
afternoon and evening. We believe that the tendency for negative AW
(psychrometer drier than pressure chamber), evident in the young-leaf
evening data of Fig. I is attributable to the expansion of leaf samples within
the psychrometers, as described by Bayer (8) and Tinklin (31). Such
expansion would occur at the expense of sample turgor, with the result that
W~ is lowered. Negative AqJ were not obtained for the young morning, or
the old evening and daytime leaves, because these leaves were not expanding.
Pressure chamber vs. in situ psychrometer f o r tubers---The relationship between pressure chamber measurements of Wtx, and in situ
psychrometer measurements of Wt, is shown in Fig. 2. Data have been
separated according to the length of time between insertion ofpsychrometers and measurement ofq/t. The 18- to 24-hour, and the 24- to 72-hour data
provide comparisons of measurements made on the same tuber. The 5- to

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10-day data were obtained by comparing psychrometer Wt values with

measurements of W~xrather than Wtx. The implicit assumption that Wtx WJx
was tested in a separate experiment in which we measured qs]x and qJtxof
drying plants. We found (18) that under the low transpiration conditions in
our glasshouse, that ~ x were 0.7 __+0.1 bar drier than qJtx. This difference
was small enough to justify treating the W~x-Wt comparison as if it were
tlJtx-~t. We have not corrected the pressure chamber readings in Fig. 2 for
the osmotic potential of the tuber xylem sap. For a comparable set of
tubers, we found that the average xylem sap osmotic potential was 1.5___0.1
bar (19). If this amount were added to the pressure chamber readings in Fig.
2, all except one of the AW values would be positive (chamber drier than
psychrometer).
*6

*4

0
0

<l

o
0

-2

5 - I0 doys
2 4 - 7 2 hours

18-24 hours
-4

-2

-4

PRESSURE-CHAMBER

-6
POTENTIAL

-8

-I0

(bors)

FIG, 2. Comparison of pressure chamber and in sitt~ psychrometer measurements of tuber

water potential. AKu = q/t-I.IJtx = it/(psychrometer) - ~ (pressure chamber). Positive Akv
indicate a drier pressure chamber estimate. Data are separated according to the length of time
between insertion of psychrometers and measurement of tlJt.

1975)

GANDAR AND TANNER: MEASURING WATER POTENTIALS

393

In the 18- to 24-hour data of Fig. 2, there is good agreement between

the chamber and psychrometer, except for tubers drier than -7 bars. The
agreement between chamber and psychrometer for the 24- to 72-hour data,
and particularly, the 5- to 10-day data, is poor. Positive AtIt values show
that psychrometers gave much wetter readings than the pressure chamber.
We believe that this is caused by gradual suberization of the tissues surrounding the psychrometer cup which slows, and eventually prevents, the
attainment of water equilibrium between tuber and psychrometer. The
efficacy of suberized tissue in reducing water vapor loss from tuber disks
has been demonstrated (20). Wet psychrometer readings in drying tubers
arise because the imposition of a suberized barrier reduces water transfer
from the psychrometer cup to the surrounding tissue, leaving the vapor
pressure in the cup erroneously high.
We examined the development of suberin in tissue surrounding psychrometers by staining hand-sections with 0.05% toluidine blue, a dye
which colors suberized cell walls blue (6). We found that about 50% of the
surface of the hole was suberized 24 hours after insertion of psychrometers; by 48 hours, suberization of the surface was almost complete and
transverse walls in the top cell layer were partly suberized; after eight days,
suberization of most of the cells in the first two layers of tissue surrounding
the psychrometers had occurred. Comparable rates of suberization have
been reported (5, 20). This rate of suberization is in keeping with the decline
in psychrometer performance with time.
In addition to suberization, there are other sources of error in our
pressure chamber-psychrometer comparison. The main error in the pressure chamber method is overestimation of the tuber endpoint caused by
high rates of pressurization or by lack of equilibrium within the tuber (19).
In both cases, ~tx would appear too dry. We minimized the first error by
using rates of pressurization of less than 0.05 bar sec _1 as endpoints were
approached. The second error should not be important under the low
transpiration conditions of our glasshouse experiments.
There are a number of difficulties in using soil thermocouple psychrometers to measure tuber water potentialin situ. Spurious readings will
be obtained if the temperature of the thermocouple differs from that of the
tissue whose water potential is being measured. Temperature errors could
arise from heat conduction down the lead wires, from thermal gradients
along the axis of the psychrometer (22), and from the increased respiratory
heat production in the wound healing process (6, 21), which follows insertion of psychrometers. We attempted to reduce the first two errors by
wrapping the psychrometer leads against the tuber and by the use of
insulation. Nothing could be done about respiratory heat production within
the tuber. When psychrometers were in the dry-bulb condition we observed offsets of from 5 to 15~tV. These suggest that temperature increases,
due to wound-induced respiration, were of the order of 0.2 C (25). We

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compensated for this heat production by nulling the psychrometer output

before each reading.
Errors caused by air gaps between the ceramic cup and surrounding
tissue could also affect in situ psychrometer measurements. We ensured a
tight fit between the psychrometer and the tuber by using an undersized
hole initially, but with time, gaps could have developed. In a limited trial,
we inserted psychrometers, from which the ceramic cups had been removed, into tubers. No significant improvement in the performance of
these psychrometers was apparent.
Pressure c h a m b e r vs. graded osmotica method---The comparison
between tuber water potentials obtained using the pressure chamber, and
from weight changes in graded osmotica, is shown in Fig. 3. There is a clear
trend: in tubers in which Wtx is wetter than about -5 bars, the graded
osmotica estimate is drier than the pressure chamber measurement; for
tubers drier than -5 bars, the converse holds.
Discrepancies shown could arise from errors in either, or both
methods. However, in view of the agreement between the pressure

,4

+2

t,.

,',

,.>,
<1

-2

-4

m.

-2

-4

-6

PRESSURE- CHAMBER POTENTIAL

-8

- 0

(bars)

FIG. 3. Comparison of pressure chamber and graded osmotica determinations of tuber water
potential. AtlJ =tlJ (graded osmotica) - t14 (pressure chamber). Positive AW indicate a drier
pressure chamber estimate.

1975)

GANDAR AND TANNER: MEASURING WATER POTENTIALS

395

chamber and psychrometer in the 18- to 24-hour data of Fig. 2, we have

some confidence that there are no major systematic errors in the pressure
chamber method. Therefore, the graded osmotica method appears to give
unreliable measurements of tuber water potential.
The basic assumption in the graded osmotica method is that changes in
the weights of tissue samples are caused by the uptake or loss of water and
not by the exchange of solutes. Uptake of solutes would cause overestimation of the final weights of tissue samples so that the tuber water potential
would appear too dry. Experiments have shown that about 25% of the
volume of disks of tuber tissue is occupied by external solution following
equilibration in osmotica (17, 30). This volume includes intercellular
spaces, in which external solution replaces air, and cell wails, in which
external solution replaces water (1, 23, 28). In our experiments, penetration of sucrose solution into intercellular spaces and cell walls could, at the
outside, cause a one bar underestimate (more negative) of tuber water
potential. Thus, solute uptake can explain only part of the negative Attt
shown in Fig. 3.
Overestimates of final weights (and underestimates of water potentials) in the graded osmotica method could also be caused by uptake of
external solution associated with the expansion of tissue samples. It has
been established that disks of tuber tissue can show weight gains associated
with cell enlargement when immersed in osmotica (W >-5 bars) and water
(12, 30). These weight gains depend upon auxin-induced changes in the
extensibility of cell walls, and upon the oxidative metabolism of the disks.
Cell enlargement also could occur for physical reasons. In an intact tuber,
cells are constrained by surrounding cells. In a tissue sample, these constraints are partially removed and expansion is possible (9). Because of the
short time and low temperatures used for tissue equilibration in our experiments, removal of physical constraints seems a more likely cause for cell
enlargement than metabolically-linked uptake. In our experiments, a 5%
increase in tissue volume would cause a 2.5-bar underestimate of tuber
water potential.
Although solute uptake and cell enlargement could explain discrepancies between the pressure chamber and the graded osmotica method for
tubers wetter than -5 bars, these mechanisms cannot account for the
positive AW shown for drier tubers. Here, other mechanisms must have an
overriding effect. One possibility is loss of solutes from tissue plugs to the
external solution (13), which could cause overestimates of tuber water
potential. However, although the loss of electrolytes from tuber disks drier
than -6 bars increases (14), this effect is probably more than offset by
sucrose uptake. Another possibility is that dry (but not plasmolysed) tubers
lose part of their ability to rehydrate on immersion in water in the same
manner as disks plasmolysed in osmotica (17). If this happened to tubers
drier than -5 bars, smaller weight gains in hypotonic solutions would be

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AMERICAN POTATO JOURNAL

(Vol. 52

expected. This would cause overestimates of graded osmotica water potentials, and hence, the positive AW of Fig. 3. However, we doubt that this is a
complete explanation for the discrepancies between the pressure chamber
and graded osmotica determinations of tuber water potential for dry tubers.
Both the graded osmotica and pressure chamber methods require
destructive sampling. In view of the ambiguities in the graded osmotica
method we recommend the faster, simpler pressure chamber method for
routine measurements of tuber water potentials.
Acknowledgment
Psychrometer measurements of leaf water potential were made by
J. W. Baughn.

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