Plant and Soil (2006) 279:229–242
Springer 2006

DOI 10.1007/s11104-005-1302-z

Relationships between climatic variables and sap flow, stem water potential
and maximum daily trunk shrinkage in lemon trees

M.F. Ortuño1, Y. Garcı́a-Orellana2, W. Conejero1, M.C. Ruiz-Sánchez1,3, O. Mounzer1,

J.J. Alarcón1,3 & A. Torrecillas1,3,4
1
Dpto. Riego y Salinidad, Centro de Edafologı´a y Biologı´a Aplicada del Segura (CSIC), E-30100 Espinardo,
(Murcia), Spain. 2Dpto. Ingenierı´a Agrı´cola, Universidad Centro Occidental Lisandro Alvarado (UCLA),
Barquisimeto, Venezuela. 3Unidad Asociada al CSIC de Horticultura Sostenible en Zonas Áridas (UPCT-
CEBAS), E-30203 Cartagena, (Murcia), Spain. 4Corresponding author*

Received 24 May 2005. Accepted in revised form 21 July 2005

Key words: irrigation scheduling, lemon, plant–water relations, sap flow, trunk diameter fluctuations

Abstract
The feasibility of obtaining sap flow (SF), maximum daily trunk shrinkage (MDS) and midday stem water
potential (Ystem) baselines or reference values for use in irrigation scheduling was studied in adult Fino
lemon trees (Citrus limon (L.) Burm. fil.) grafted on sour orange (C. aurantium L.) rootstocks. Plants were
irrigated daily above their water requirements in order to obtain non-limiting soil water conditions. The
results indicated that baselines for plant-based water status indicators (MDS, SF and Ystem) can be ob-
tained, even though there was a certain scattering of the data points representing the relations between the
plant-based measurements and the environmental variables (reference evapotranspiration, solar radiation,
vapour pressure deficit and temperature). SF was more closely associated with changes in the studied
evaporative demand variables than were MDS and Ystem. SF and Ystem were more closely correlated with
changes in reference evapotranspiration (ETo) (r2 = 0.93 and 0.79, respectively), while MDS behaviour
was best correlated with mean daily air temperature (Tm) (r2 = 0.76). Increases in the evaporative demand
induced more negative Ystem values and, as a consequence, SF increased, which, in turn, was translated into
an increase in MDS. This confirmed that SF and MDS were very good predictors of the plant water status
during the observation period and their continuous recording offers the promising possibility of their use in
automatic irrigation scheduling in lemon trees.

Abbreviations: Ystem – midday stem water potential; ETc – crop evapotranspiration; ETo – reference
evapotranspiration; MDS – maximum daily trunk shrinkage; Rs – solar radiation; SF – sap flow; TDF –
trunk diameter fluctuations; Tm – mean daily air temperature; Tmd – midday air temperature; VPDm – daily
mean vapour pressure deficit; VPDmd – midday vapour pressure deficit

Introduction tus may be useful for irrigation scheduling

In recent years the use of plant-based water sta-
tus indicators have become very popular to study
plant–water relations and for planning irrigation
programs. The measurement of plant water sta-
* FAX No: +34-968-396213.
E-mail: atorreci@cebas.csic.es
because of its dynamic nature, which is directly
related with climatic and soil conditions, as well
as with crop productivity (Goldhamer et al.,
2003; Remorini and Massai, 2003). The most
widely used approach for evaluating tree water
status has been to determine leaf water potential
(Améglio et al., 1999; Hsiao, 1990). However,
Choné et al. (2001), Naor (2000) and Shackel
230
et al. (1997) showed that midday stem water
potential (Ystem) is a significant and more reliable
plant water status indicator for scheduling the
irrigation of woody crops. However, the main
disadvantage of Ystem is the relatively cumber-
some measurement procedure, the necessity of
frequent trips to the field and a significant input
of labour.
The robustness of the sensors used to measure
sap flow (SF) and trunk diameter fluctuations
have renewed interest in using these parameters
as plant water status indicators (Cohen et al.,
2001; Fernández et al., 2001; Goldhamer and
Fereres, 2001; Moreno et al., 1996). These tech-
niques permit continuous and automated regis-
ters of the plant water status, and an immediate,
consistent and reliable response to water deficit
(Goldhamer et al., 1999; Ortuño et al., 2004a, b),
with a reduction in the labour needed for the
measurement procedure. Goldhamer and Fereres
(2004) obtained promising results using only
maximum daily trunk shrinkage (MDS) for
scheduling almond trees irrigation.
To correctly interpret the values of each
plant-based water status indicator and their use
for irrigation scheduling, it is necessary to devel-
op baselines or reference relationships, which are
obtained with their values in plants under non-
limiting soil water conditions versus the evapora-
tive demand of the atmosphere (Goldhamer and
Fereres, 2001; Martin et al., 1990; Shackel et al.,
1997).
For the above reasons, the objective of this
study was to evaluate the feasibility of obtaining
SF, MDS and Ystem measurement baselines in
adult lemon trees to be used in irrigation schedul-
ing. This approach involved characterizing the
behaviour of SF, MDS and Ystem under non-lim-
iting soil water conditions with respect to changes
in several parameters related with the evaporative
demand of the atmosphere. The relations between
SF, MDS and Ystem were also investigated.
Materials and methods
Plant material and experimental conditions
Experiments were conducted at the CEBAS-
CSIC experimental station in Santomera (Mur-
cia) from 2 April 2004, day of year (DOY) 93, to
10 January 2005, DOY 10. The soil is a Paralith-
ic mollic-calciorthid, and the profile shows only a
slight differentiation between horizons (an ochric
epipedon on a C horizon). The clay loam soil of
the experimental site (1.28% organic matter,
4.92 mmol kg)1 available potassium, 1.81 mmol
kg)1 available phosphorus, 51% lime content
and pH of 7.8) was characterized by volumetric
water content of 0.19 cm3 cm)3 at field capacity
and 0.08 cm3 cm)3 at permanent wilting point,
1.5 g cm)3 bulk density, high stone content (43%
w/w), and 8.0 cm h)1 saturated hydraulic con-
ductivity, which provided an excellent internal
drainage (Trout et al., 1982).
The experiment was performed on 24-year-old
lemon trees (Citrus limon (L.) Burm. fil.) cv. Fino
grafted on sour orange (C. aurantium L.) root-
stocks. Tree spacing followed a 6
6 m square
pattern, with an average ground cover of about
65%.
Micrometeorological hourly data, namely air
temperature, precipitation, solar radiation, air
relative humidity and wind speed at 2 m above
soil surface were collected by a weather station
of the Servicio de Información Agraria de Mur-
cia (SIAM), located 8.4 km from the experimen-
tal site. Daily reference evapotranspiration (ETo)
was calculated using the Penman–Monteith equa-
tion (Allen et al., 1998). Daily mean vapour pres-
sure deficit (VPDm) was calculated from mean
daily vapour pressure and relative humidity
(Goldhamer and Fereres, 2001).
No weeds were allowed to develop within the
orchard, resulting in a clean orchard floor for the
duration of the experiment. Pest control and fer-
tilization practises were those commonly used by
the growers.
Irrigation was carried out during the night by
drip using two lateral pipes per tree row and 12
emitters per plant, six of which delivered 2 L h)1
(0.06 mm h)1) and six 4 L h)1 (0.11 mm h)1).
Plants irrigation requirements were determined
according to ETo and a crop factor based on the
time of the year and the percent of ground area
shaded by the tree canopy (Domingo et al.,
1996). During the experimental period, total crop
evapotranspiration (ETc) was 566.5 mm.
From the beginning of the experimental peri-
od, lemon trees were irrigated daily above crop
water requirements in order to obtain non-limit-
ing soil water conditions. A total water amount

of 843.5 mm, measured with in-line water meters,
was applied during the experiment. Even though
the amount of irrigation water applied to plants
during the experimental period was clearly above
crop water requirements, the absence of plant
symptoms and the lemon tree water relations
indicated the absence of any waterlogging situa-
tion (Garcı́a-Orellana, unpublished data).
Measurements
Trunk diameter fluctuations (TDF) were mea-
sured throughout the experimental period in four
trees, using a set of linear variable displacement
transducers (LVDT) (model DF ± 2.5 mm, accu-
racy ± 10 lm, Solartron Metrology, Bognor
Regis, UK) attached to the trunk, with a special
bracket made of Invar, an alloy of Ni and Fe with
a thermal expansion coefficient close to zero (Kat-
erji et al., 1994), and aluminium. Sensors were
placed on the north side and were covered with
silver thermoprotected foil to prevent heating and
wetting of the device. Measurements were taken
every 10 s and the datalogger (model CR10
with
AM 416 multiplexer, Campbell Scientific Ltd.,
Logan, USA) was programmed to report 30 min
means. Maximum daily trunk shrinkage (MDS)
was calculated as the difference between maxi-
mum and minimum daily trunk diameter.
Sap flow was measured from 2 June 2004
(DOY 154) using the compensation heat-pulse
technique (Swanson and Whitfield, 1981) in the
same trees as used for TDF measurements. One
set of heat-pulse probes was located above the
LVDT sensors on each tree. Each set consisted
of two temperature probes inserted 10 mm
downstream and 5 mm upstream from a heater
needle. Heater and thermocouples had a diame-
ter of 1.8 mm and were installed in parallel holes
drilled radially in the trunks. The temperature
probes comprised four copper–constantan ther-
mocouples to measure the sap velocity at four
radial depths of 10, 20, 30 and 40 mm below the
cambium to monitor the sap velocity profile over
a radial depth. For the experiments, pulse dura-
tion of 1 s was used. Sap velocity was measured
following the procedure of Green and Clothier
(1988) and taking into account the Swanson cor-
rection factors for a wound size of 2.4 mm
(Swanson and Whitfield, 1981).
231
Once the heat-pulse velocity (V) was deter-
mined, the next step was to relate it to the actual
SF. In our analysis, the sap flow density (J) was
related with the heat-pulse velocity (V) using the
equation developed by Edwards and Warwick
(1984):
J ¼ ð0:505Fm þ Fl ÞV
where Fm and Fl are the volume fractions of
wood and water, respectively. These were calcu-
lated at the end of the experiment by taking
5 mm diameter wood samples from all trees with
a special auger and measuring the fresh mass,
mass of displaced water by immersion in a vessel
of distilled water on a balance, and oven dried
mass. The factor 0.505 is related to the thermal
properties of the woody matrix, and is assumed
to be constant within and between species. This
equation provides an estimate of the values of J
at any point in the conducting sapwood. It is
widely recognized that sap flux density is not
uniform throughout the sapwood, for which rea-
son our probes measured J at four radial depths.
The volumetric measurement of total sap flux
(Q) was obtained by the integration of these
point estimates over the sapwood conducting
area, which was determined at the end of the
experiment by cutting the trunks and immersing
them in a safranin solution overnight and mea-
suring the radius of sapwood at the point where
the heat-pulse sensors were installed. The temper-
ature signals and the corresponding heat-pulse
velocities were recorded at a 30 min intervals
using compensation heat-pulse instrumentation
(MITRA-3.1, Polytechnic University of Cartage-
na, Spain) controlled by a data logger (CR10
,
Campbell Scientific Ltd., Logan, USA).
Midday (12.00 h solar time) stem water
potential (Ystem) was measured from 9 July 2004
(DOY 191), every 2–3 days, except when rainfall
prevented measurements, in two mature leaves
per plant, taken from close to the trunk. Leaves
were covered using a small black plastic bag cov-
ered with aluminium foil for at least 2 h before
measurement with a pressure chamber.
Statistical design and analysis
The design of the experiment was completely ran-
domized with four replications, each replication
232
consisting of three adjacent rows of five trees.
Measurements were taken in the inner tree of the
central row of each replicate, the other trees
serving as borders.
All the measurements were taken in the same
tree in each replicate. Ystem values for each day
and replicate were averaged before the mean and
the standard error were calculated.
Results
All the parameters concerning the evaporative
demand of the atmosphere increased from the
beginning of the experiment, reaching maximum
values in summer and then decreasing until the
end of the experiment (Figure 1). During the
experimental period, rainfall was 250.2 mm,
mainly occurring during spring and autumn, the
usual rainy periods in the experimental site area
(Figure 1a). Mean daily air temperature (Tm) and
midday air temperature (Tmd) presented a similar
seasonal trend, reaching maximum values in late
June and early August (Figure 1a). During the
experiment, average Tm and average daily mini-
mum temperatures were 25 and 13 C, respec-
tively (Figure 1a), and average mean relative
humidity was 66% (data not shown).
During the experiment, daily ETo reached
maximum values in June and July and decreased
afterwards. Total ETo was 1009.9 mm (Fig-
ure 1b). Solar radiation (Rs) fluctuated widely
during the experimental period, showing maxi-
mum values in mid June and minimum values in
late November (Figure 1b). Midday VPD
(VPDmd) and VPDm values presented similar sea-
sonal trends, and maximum values were obtained
in late June and early August (Figure 1c).
Ystem values varied between )1.5 and
)0.6 MPa, increasing during the observation per-
iod (Figure 2a). MDS values ranged between
0.03 and 0.49 mm, increasing from the beginning
of the experiment to early July and decreasing
afterwards (Figure 2b). Daily SF values were be-
tween 64 and 227 L day)1, presenting nearly con-
stant values from the beginning of the
measurements to mid August and decreasing
afterwards (Figure 2c).
Figures 3–6 represent the plant-based water
status indicators (MDS, Ystem and SF) as a func-
tion of ETo, Rs, VPDm, VPDmd, Tmd, and Tm.
The overall increases in any of the selected envi-
ronmental variables were associated with in-
creases in MDS and SF and decreases in Ystem
(Figures 3–6). The r2 values of the data around
the regression lines varied from 0.39 to 0.93. In
all cases, r2 was higher for SF than for MDS and
Ystem (Figures 3–6).
Figure 3 presents the MDS, Ystem and SF as
a function of ETo. Of the relationships between
the water status indicators and ETo, the correla-
tion between MDS and ETo (Figure 3a) pre-
sented the weakest r2 value. A cubic fit of the
regression between SF and ETo (Figure 3c) yiel-
ded an equation with a determination coefficient
higher than those found for the first and second
order regressions (data not shown). Increases in
ETo were associated with increases in SF until
ETo reached values of around 3.5 mm, after
which SF levelled off as ETo increased
(Figure 3c). Ystem showed a better correlation
with ETo than with the other selected evapora-
tive demand parameters (Figures 3–6).
The first order regression of Ystem values
against Rs was characterized by a tighter correla-
tion than that of MDS against Rs (Figure 4a and
b). Moreover, in the case of the relationship
between SF and Rs a departure from linearity
was evident (Figure 4c) and a second degree
regression significantly improved the goodness of
the first order fit (data not shown). In this case,
increases in Rs values were associated with
increases in SF until Rs reached values of around
230 W m)2, after which increases in Rs values
were associated with near constant SF values
(Figure 4c).
The MDS and Ystem regressions against
VPDm and VPDmd (Figure 5a and b) showed
relatively low correlations. The quadratic fit
regression between SF and VPDm and VPDmd
(Figure 5c) rendered coefficients of determination
higher than that found for the linear fit (data not
shown). The relations between SF and VPDm and
VPDmd departed from linearity at VPDm and
VPDmd values above 2 and 3 kPa, respectively.
The relations between MDS, and Tm and Tmd
presented tighter correlations than that consider-
ing the other environmental variables (Figure 6a),
while those of SF and Ystem against both
temperatures (Tm and Tmd) correlated well but
more weakly than those obtained against ETo
(Figures 3b, c and 6b, c).
 233

(a)

35 35

30
Air temperature (ºC)

30

Rainfall (mm)
25
25
20
20
15
15 Tm
10
Tmd
10 5

(b)
350
9

8 300
RS
7
ETo

Rs(Wm-2 )
250
ETo (mm)

6
200
5

4 150

3
100
2
50
1

(c)

4 Días Julianos vs Eto (mm/d)

VPDm
Días vs Rad solar
VPDmd
VPD (kPa)

100 120 140 160 180 200 220 240 260 280 300 320 340 360

DOY

Figure 1. Daily mean (Tm, dotted line) and midday (Tmd, solid line) air temperature, and daily rainfall (vertical bars) (a), solar
radiation (Rs, dotted line) and reference evapotranspiration (ETo, solid line) (b), daily mean (VPDm, dotted line) and midday
(VPDmd, solid line) air vapour pressure deficit (VPD) (c) values during the experimental period.
234

-0.6 (a)

Ψstem (MPa) -0.8

-1.0

-1.2

-1.4

(b)
0.5

0.4
MDS (mm)

0.3

0.2

0.1

(c)
225

200
SF (L day )
-1

175

150

125

100

75

100 120 140 160 180 200 220 240 260 280 300 320 340 360

DOY

Figure 2. Midday stem water potential (Ystem) (a), maximum daily trunk shrinkage (MDS) (b) and daily sap flow (SF) (c) values
during the measurements period. Vertical bars are twice the overall mean SE.
 235
(a)
MDS = 0.06 + 0.05 ETo
2
r = 0.6561
0.5

MDS (mm) 0.4

0.3

0.2

0.1

(b)

-0.50

-0.75
Ψstem (MPa)

-1.00

-1.25

-1.50 Ψ stem = -0.60 - 0.13 ETo

r 2 = 0.7921

(c)

200
SF (L day-1 )

150

100

50 SF = 33.28 + 78.75 ETo - 13.56 ETo2 + 0.83 ETo3

r2 = 0.9266

1 2 3 4 5 6

ETo (mm)

Figure 3. Relationships between ETo and maximum daily trunk shrinkage (MDS) (a), midday stem water potential (Ystem) (b) and
daily sap flow (SF) (c) values during the measurement period.
236
(a)
MDS = 0.06 + 0.001 Rs
2
r =0.4666
0.5

0.4
MDS (mm)

0.3

0.2

0.1

(b)
Ψstem = -0.49 - 0.01 Rs
r2 =0.6613
-0.50

-0.75
Ψstem (kPa)

-1.00

-1.25

-1.50

(c)
2
SF = 21.49 + 1.08 Rs - 0.01 Rs
r2 =0.7840

200
SF (L day )
-1

150

100

50

50 100 150 200 250 300 350

Rs (W m-2)

Figure 4. Relationships between solar radiation (Rs) and maximum daily trunk shrinkage (MDS) (a), midday stem water potential
(Ystem) (b) and daily sap flow (SF) (c) values during the measurement period.
 237

(a)
VPDm
VPDmd
0.5

0.4
MDS (mm)

0.3

0.2
MDS = 0.07 + 0.15 VPDm
r 2 =0.5560
0.1 MDS = 0.09 + 0.08 VPD md
2
r =0.5293

(b)

-0.50
Ψst em = -0.70 - 0.18 VPD md
2
r = 0.394
-0.75 Ψstem = -0.68 - 0.31 VPDm
Ψstem (MPa)

r 2 = 0.4268

-1.00

-1.25

-1.50
VPD m
VPD md

(c) VPDm
VPDmd

200
SF (L day-1)

150

100 2
SF = 26.43 + 163.99 VPD m - 36.61 VPDm
2
r = 0.8464
SF = 37.25 + 91.47 VPDmd - 12.09 VPDmd2
50 2
r = 0.7883

1 2 3 4 5
VPD (kPa)

Figure 5. Relationships between daily mean (VPDm, open symbols) and midday (VPDmd, closed symbols) air vapour pressure defi-
cit, and maximum daily trunk shrinkage (MDS) (a), midday stem water potential (Ystem) (b) and daily sap flow (SF) (c) values
during the measurement period.
238

(a)
MDS = -0.07 + 0.02 Tm
2
0.5 r = 0.7591

MDS = -0.13 + 0.02 Tmd

2
0.4 r = 0.7460
MDS (mm)

0.3

0.2
Tm
0.1 Tmd

(b)
Tm
-0.50 Tmd

-0.75
Ψstem (MPa)

-1.00

-1.25
Ψstem = -0.37 - 0.03 Tm
r 2 = 0.5979
-1.50
Ψstem = -0.22 - 0.03 Tmd
2
r = 0.5492

(c)
SF = 31.97 + 6.30 Tm
2
r = 0.8613
200
SF = 1.38 + 6.11 Tmd
2
r = 0.8609
SF (L day )
-1

150

100
Tm
Tmd

50

10 15 20 25 30 35
Air temperature (ºC)

Figure 6. Relationships between daily mean (Tm, open symbols) and midday (Tmd, closed symbols) air temperature, and maximum
daily trunk shrinkage (MDS) (a), midday stem water potential (Ystem) (b) and daily sap flow (SF) (c) values during the measure-
ment period.
 239

(a) (b)
-0.50

-0.75
Ψstem (MPa)

-1.00

-1.25

-1.50 Ψ stem = -0.48 - 2.20 MDS Ψstem = -0.18 - 0.01 SF

r 2 =0.7796 r 2 =0.6453

0.1 0.2 0.3 0.4 0.5 50 100 150 200

MDS (mm) SF (L day-1)

Figure 7. Relationships between midday stem water potential (Ystem), and maximum daily trunk shrinkage (MDS) (a) and daily
sap flow (SF) (b) values during the measurement period.

MDS = - 0.11 + 0.01 SF

2
r =0.7586
0.5

0.4
MDS (mm)

0.3

0.2

0.1

50 100 150 200

SF (L day-1)
Figure 8. Relationship between maximum daily trunk shrinkage (MDS) and daily sap flow (SF) values during the measurement
period.

The regressions analysis between Ystem, and
MDS and SF, pooling data across the observation
period, indicated clear associations (Figure 7).
Also, the MDS versus SF relationship showed a
high coefficient of determination (Figure 8).
Discussion
Since the three plant-based water stress indicators
studied (MDS, SF and Ystem) encompass different
dimensions and time scales, it was to be expected
that MDS and SF would correlate better with the
evaporative demand indicators measured on a
whole-day basis, while Ystem would best fit the
parameters measured at midday. The regression
analysis indicated that the highest coefficients of
determination were obtained for the regressions
of SF, MDS and Ystem against ETo, Tm and ETo,
respectively (Figures 3–6). The fact that SF was
more closely associated with changes in ETo can
be explained by taking into consideration that SF
240
is directly related with daily transpiration
(Alarcón et al., 2005; Ortuño et al., 2004a). Even
though temperature is not an accurate indicator
of the evaporative demand of the atmosphere
(Hatfield and Fuchs, 1990), MDS was closely
related with Tm. In this sense, Fereres and Goldh-
amer (2003) showed that MDS correlated well
with both Tmd and Tm in almond trees. However,
Vélez (2004) indicated that MDS was more
directly related with changes in Rs and ETo than
VPD and temperature in Citrus clementina trees.
According to the characteristics of the data
for the different studied regression models (Fig-
ures 3–6), it can be concluded that the behaviour
of SF can be more adequately predicted (higher
r2) by changes in the evaporative demand
variables than can MDS and Ystem. Also, the
MDS was more adequately described by changes
in the environmental variables than was Ystem,
except when ETo or Rs were considered.
At first sight, the fact that SF levelled off
after ETo, Rs, VPDmd and VPDm reached
threshold values (Figures 3c, 4c and 5c) would
suggests that spatial variability of soil moisture
availability and/or the high hydraulic conductiv-
ity of the soil constraints plant transpiration even
with a high volume of irrigation. However, other
logical explanation could be that maximum and
constant values of SF correspond to physical
constraint of plant transpiration. In this sense,
Pataki and Oren (2003) studied factors control-
ling transpiration of some trees species, and
showed that Quercus rubra, Q. alba, Carya
tormentosa and Fraxinus americana plants were
responsive to vapour pressure deficit and pre-
sented saturation in increases in daily sap flux
density with vapour pressure deficit, these re-
sponses being similar at different soil moisture
levels. Moreover, considering that MDS respond
sooner to water stress than other continuous and
discrete plant and soil water status indicators
(Goldhamer et al., 1999) and that MDS is a ver-
satile indicator of transpiration stream intensity
when the soil water content is not strongly de-
pleted (Huguet et al., 1992; Ortuño et al., 2004a,
b), the fact that when SF presented high and
constant values (Figure 2c) MDS values progres-
sively decreased (Figure 2b) indicated that lemon
trees were not under a water stress situation.
In contrast with the observed behaviour in
the SF versus ETo, Rs and VPD relationships,
the fact that the relationships between MDS and
the environmental variables (Figures 3–6) were
linear even at high values of evaporative demand
was probably the result of the efficient water
recruitment from additional stem tissue capaci-
tances when evaporative demand increased and
Ystem decreased below a threshold. In this sense,
Zweifel et al. (2001) showed that internally
stored water contributed to daily transpiration
even in well-watered trees, indicating that stored
water plays an important role not only during
periods of drought, but whenever water transport
occurs within the tree.
Previous authors have shown weak correla-
tions between MDS and Ystem on a seasonal
basis in almond trees (Fereres and Goldhamer,
2003), in plum trees (Intrigliolo and Castel, 2004)
and in peach trees (Marsal et al., 2002), but
when they were broken into different time peri-
ods the correlation for individual periods clearly
improved, suggesting that this may be a general
trend for deciduous fruit trees due to trunk
growth rate and tissue elasticity changes during
the season (Intrigliolo and Castel, 2004). Our
results, on the other hand, pointed to a constant
relationship between MDS and Ystem (Figure 7a),
which means neither postulated cause was impor-
tant in adult lemon trees during the observation
period.
Molz and Klepper (1973) indicated that the
main factor controlling MDS is the leaf water
potential, which determines the driving force for
water transport between the bark and the xylem.
The goodness of the correlation found between
Ystem and MDS (Figure 7a) confirmed this view.
Moreover, the fact that there were direct relation-
ships between Ystem and ETo (Figure 3b), Ystem
and SF (Figure 7b) and SF and MDS (Figure 8)
indicated that the control of MDS by Ystem could
be mediated through SF. The mechanism, under
non-limiting soil water conditions, would be
described as follow: increases in the evaporative
demand of the atmosphere induce decreases in
Ystem (Figure 3b), which increases the soil–plant
water potential gradient, and as a consequence
SF increases (Figure 7b) turning into an increase
in MDS (Figure 8).
The above mentioned results indicated that
baselines or reference values for plant-based wa-
ter status indicators (MDS, SF and Ystem) can be
obtained for adult lemon trees, even though there

was a certain scattering in the relations between
the plant-based measurements and the environ-
mental variables. According to the r2 values of
the data around the different regression models,
it can be concluded that the SF behaviour more
adequately reflected changes in the evaporative
demand variables than those of MDS and Ystem.
The respective behaviour of SF and Ystem corre-
lated more closely with changes in ETo
(r2 = 0.93 and 0.79, respectively), while MDS
behaviour was best correlated with Tm
(r2 = 0.76). Whatever the case, these correlations
must be used within their confidence levels.
Increases in the evaporative demand induced
decreases in Ystem and, as a consequence, SF in-
creased, which, in turn, was translated into an in-
crease in MDS. This confirmed that SF and
MDS were very good predictors of the plant wa-
ter status during the observation period and their
continuous recording offers the promising possi-
bility of their use in automatic irrigation schedul-
ing in lemon trees.
Acknowledgements
This research was supported by Ministerio de
Educación y Ciencia (MEC), (CICYT/FEDER
AGL2003-9387-C05-02 and AGL2004-0794-C03-
02) and PETRI (PTR1995-0693-OP-02-01) grants
to the authors. M.F. Ortuño and O. Mounzer
were research fellowships from CSIC (Program
I3P) and MEC (FPI), respectively.
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