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Plant, Soil and Environment, 69, 2023 (7): 303–313 Original Paper

https://doi.org/10.17221/160/2023-PSE

Stem water potential, stomatal conductance and yield


in irrigated apple trees

Lenka Plavcová1*, Radek Jupa1,3, Martin Mészáros2, Klára Scháňková2,


Zuzana Kovalíková1, Jan Náměstek2
1Faculty of Science, University of Hradec Králové, Hradec Králové, Czech Republic
2Research and Breeding Institute of Pomology, Holovousy, Hořice, Czech Republic
3Faculty of Science, Masaryk University, Brno, Czech Republic

*Corresponding author: lenka.plavcova@uhk.cz

Citation: Plavcová L., Jupa R., Mészáros M., Scháňková K., Kovalíková Z., Náměstek J. (2023): Stem water potential, stomatal
conductance and yield in irrigated apple trees. Plant, Soil and Environ., 69: 303–313.

Abstract: Plant-based indicators of water status, such as midday stem water potential (Ψstem) and leaf stomatal con-
ductance (gs), are used to optimise irrigation scheduling in horticultural crops because they integrate the effect of soil
and climatic conditions and the internal physiological constraints. The use of these indicators relies on experimentally
acquired thresholds that relate the value of the indicator to negative effects on yield. In five irrigation treatments, we
monitored yield, fruit size and the courses of Ψstem and gs throughout four consecutive growing seasons. We found
that Ψstem was more sensitive to irrigation treatment than gs. Both indicators increased with available soil water
content (ASWC) and decreased with evaporative demands of the atmosphere (ETC). On a seasonal basis, crop load
had a stronger impact on gs than Ψstem. In summary, our study explored the effect of environmental conditions and
crop load on plant-based indicators of tree water status and can be useful for establishing thresholds for irrigation
scheduling in apple tree orchards.

Keywords: drought; water dynamic; fruit trees; gas exchange; deficit; abiotic stress

In many arid and semi-arid regions, the sustain- the tree’s rooting zone (Dodd 2007, Bauerle et al.
able yield of horticultural crops largely depends 2008). In addition, there are inherent physiological
on irrigation water. Yet, irrigation is becoming an differences in water use between crop species and
important agricultural practice even in temperate cultivars (Levin et al. 2020, Plavcová et al. 2023).
regions due to ongoing climate change (Cancela et Therefore, plant-based indicators that utilise plants
al. 2006). Irrigation scheduling has been traditionally as biosensors of water deficit appear as promising
based on the estimation of orchard water balance tools for more efficient irrigation scheduling (Jones
from measured climatic conditions and estimated 2004, Fernández 2017).
crop coefficients (Allen et al. 1998, Allen and Pereira Several plant-based indicators have been identified
2009) or on soil moisture-based criteria (Campbell as useful descriptors of plant water status. Among
et al. 1982, Hanson et al. 2000). However, these ap- them, midday stems water potential (Ψstem) measured
proaches have limitations as they do not account with a Scholander-type pressure chamber is arguably
for the spatial and temporal heterogeneity which the most widely used plant-based water status indica-
is common under field conditions. For instance, tor (Naor et al. 1995, Shackel et al. 1997), although
soil moisture measured by point sensors does not its measurements are labour intense and restricted to
capture well the complex soil water dynamics within discrete sampling dates. Ψstem responds dynamically

Supported by the Ministry of Agriculture of the Czech Republic, Project No. QK1910165. _________________________

© The authors. This work is licensed under a Creative Commons Attribution-NonCommercial 4.0 International (CC BY-NC 4.0).

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Original Paper Plant, Soil and Environment, 69, 2023 (7): 303–313

https://doi.org/10.17221/160/2023-PSE

to changing soil and atmospheric conditions and the experimental plots of Research and Breeding
integrates the effect of plant water uptake, transport Institute of Pomology Holovousy Ltd., Czech Republic
and loss. Values of Ψ stem provide a useful indication (15°34'48''E, 50°22'24''N) were used for the measure-
of the level of drought stress experienced by the plant ments. The scions were grafted on semi-dwarfing M9
and should be related to hydraulic thresholds such rootstock and spaced 3.5 m between rows and 1.2 m
as turgor loss point or the onset of run-away xylem between trees. The trees were trained as slender
cavitation (Meinzer et al. 2009, Plavcová et al. 2023). spindles combined with "Klik" pruning. The final
Stomatal conductance (gs) has also been suggested as height of the trees was 3.2 m. The measurements
a suitable plant-based indicator because stomatal closure occurred during four consecutive growing seasons
belongs to the first response to water stress, at least from 2019 to 2022. The orchard site experiences
in isohydric species. Apple trees, which are the focal a temperate mild climate with a mean annual tem-
tree species of this study, were considered as isohydric perature of 8.4 °C and mean precipitation of 664 mm.
species with a stringent stomatal control (Lakso 1994, The soil texture composition on the site was 22.2%
Lauri et al. 2016), although more recent study reported clay, 69.1% silt and 8.7% sand based on laboratory
anizohydric behaviour and very narrow hydraulic safety analyses using Kopeckeho o-rings. Hence, the soil was
margins in young high-yield apple trees (Beikircher et classified as silty loam Luvisol soil of medium fertility
al. 2013). While stomata close in response to low soil according to WRB classification (IUSS Working Group
water and high vapour pressure deficit (Fernández WRB 2015). Based on the soil texture, soil water con-
et al. 2011, Ahumada-Orellana et al. 2019), stomatal tent at field capacity was established at 34.2 vol. %,
conductance was found to be a less sensitive water and the permanent wilting point was at 15.9 vol. %.
status indicator than Ψ stem and maximal diurnal trunk To regulate fruit set to optimal crop density, hand
shrinkage (MDS) in apple trees (Doltra et al. 2007). thinning at BBCH 72 (fruit diameter of 20 mm ac-
In horticultural crops, tree water use is also strongly cording to Meier 2001) was carried out with the aid
affected by crop load, with higher yield being asso- of Equilifruit disc (Kon and Schupp 2013) for the
ciated with greater water demands (Wünsche et al. cultivar Red Jonaprince the maintained number of
2000, Naor et al. 2008). Therefore, trees with higher fruit per branch cross-section area was estimated as
crop loads typically have lower water potential val- δ + 1, where δ is the recommended optimal bearing.
ues (Naor et al. 2008), although leaf water potential The plant protection and fertilisation followed the
was not affected by crop load in adult lemon trees rules of integrated fruit production. Herbicides were
(Ortuño et al. 2009). The explanation for the lower applied to maintain the row weed free. The inter
water potential values in heavily cropping trees is rows were covered with grass and periodically mown.
sought in their higher leaf gas exchange rates because Irrigation treatments. Five different irrigation
ample fruits cause high demands for assimilates treatments were applied to sectors consisting of
(Wünsche et al. 2000). 17 adjacent trees grown in five neighbouring north-
This study aimed to evaluate the effect of five ir- south oriented rows. The sectors were not rotated
rigation treatments on fruit yield and size calibre and during the four consecutive seasons of the experi-
relate these yield parameters to midday stem water ment; hence, the effect of irrigation compounded
potential and leaf gas exchange during four consecu- over the four years. The trees were drip-irrigated with
tive years. We expected that non-irrigated trees will the dripline placed within the tree row at the height
bear less fruits and/or smaller fruits compared to the of 0.5 m above the soil surface. The drippers had
irrigated ones. We also expected Ψ stem and g s to be a flow capacity of 2.3 L/h and were placed in 0.5 m
lower in non-irrigated than irrigated trees. By linking spacing. Crop evapotranspiration was estimated
yield parameters with plant-based indicators of water according to the Penman-Monteith equation (Allen
stress, we will be able to refine irrigation scheduling et al. 1998), with later updates by Allen and Pereira
in apple orchards and, by extension, utilise limited (2009). First, reference evapotranspiration (ET 0 )
water irrigation reservoirs more efficiently. was calculated using the Penman-Monteith equa-
tion. ET0 was then multiplied by the crop coefficient
MATERIAL AND METHODS that varied from 0.5 to 1.2 throughout the season,
reflecting the canopy development. The calculation
Plant material. Apple trees (Malus × domestica used micrometeorological data measured within the
Borkh.) c v. Red Jonaprince planted in 2013 at orchard at 2 m above ground. The measured micro-

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Plant, Soil and Environment, 69, 2023 (7): 303–313 Original Paper

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meteorological variables included air temperature lected at the beginning of the experiment in 2019 and
(DS18B20, Dallas Semiconductor, Dallas, USA), air used throughout the whole 4-year period. At the end
humidity (HIH-4000, Honeywell, Charlotte, USA), of each growing season, fruits were manually picked,
wind direction and speed (W2, Tlusťák, Prague, sorted and weighed using a portable scale with an
Czech Republic), solar radiation (SG002, Tlusťák, accuracy of 0.01 kg. Total annual fruit yield (kg/tree),
Prague, Czech Republic) and rainfall totals (Small fruit count and fruit distribution within three size
Rain Gauge 100.053, Pronamic, Skjern, Denmark). categories (small: < 65 mm, medium: 65–75 mm,
The data were further validated against the official large: > 75 mm) were determined.
meteorological data by Czech Hydrometeorological Midday stems water potential. Midday stems wa-
Institute measured in the station situated 500 m from ter potential was measured on sunny days in 2-week
the orchard. The orchard data agreed well with the intervals from June to September. The measurements
official meteorological data, and we used the orchard were done on four tree individuals per irrigation
data for all our analyses. Soil moisture was measured treatment that were selected at the beginning of each
within the root zone of the trees using three sensors growing season to have homogeneous flower density
(VIRRIB, Fiedler AMS, České Budějovice, Czech and hence expected bearing. For each tree, two fully
Republic) per plot placed in soil depths of 10, 30 and expanded healthy leaves on the current year extension
60 cm. The VIRRIB sensors measure soil moisture shoots that faced the sun were selected and covered
based on soil electric conductivity. They were placed with an aluminium bag to prevent transpiration.
in the middle between two neighbouring trees in the Ψ stem was measured after at least 30 min of equili-
row directly under the drip line, thus measuring the bration. The covered leaves were excised between
wetted soil volume. Measured soil volumetric water 11:00–13:00 h and measured immediately using
content was converted to the saturation proportion a portable Scholander pressure chamber (1505D-EXP,
of available soil water content (ASWC) calculated PMS Instrument Company, Albany, USA). We used
as the difference in soil water content between the the average calculated from the two measurements
field capacity and the wilting point. per tree for statistical analyses.
In total, five irrigation treatments were adminis- Leaf gas exchange. Leaf gas exchange parameters
tered. These included ET-100, ET-50 and non-irrigated were measured on the same days, and trees were used
control ET-0, in which 100% or 50% or none of the for Ψ stem measurements. The measurements were
estimated evapotranspiration was supplied after ac- carried out during 11:00–13:00 h using a portable
counting for natural rainfall. These treatments applied infra-red gas analyser (LI-6800P, Li-Cor Inc., Lincoln,
irrigation during the whole growing season from USA). A healthy mature leaf was inserted into the
1 st April to 30 th September. The other two irrigation measuring cuvette, and the leaf gas exchange reading
treatments were regulated deficit irrigation (RDI) was taken after reaching a steady state, which typically
RDI-50 and RDI-50a, in which 50% of the estimated took 2–3 min. The irradiance was set to a constant of
evapotranspiration was replaced during the rapid 1 800 μmol/m 2/s, close to the ambient irradiance on
phase of fruit growth (BBCH 72–77 according to sunny days when the measurements were done and
Meier 2001), while full replacement of evapotranspi- corresponded to light-saturated photosynthetic condi-
ration was done outside of these phenological stages. tions. Reference CO 2 concentrations were 400 ppm,
RDI-50 had an optimal crop load as determined using the flow rate was 300 μmol/s, and the fan speed was
a hand-thinning gauge (Equilifruit). In the RDI-50a 10 000 rpm. The chamber’s air temperature and relative
treatment, the fruit load was regulated to 60% of the air humidity were set to follow the ambient condi-
optimal crop load of the cultivar. Irrigation scheduling tions. Matching of reference and sample infrared gas
was based on soil moisture criterium, and the goal analysers was done after each measurement. Out of all
was to prevent ASWC to decline below 0.7 in fully measured leaf gas exchange parameters, leaf stomatal
irrigated ET-100 treatment. Thus, irrigation volumes conductance was selected and used for analyses as it
and frequencies varied depending on soil moisture affects leaf water loss and leaf carbon gain.
and were between 4 to 11 mm per dose applied Statistical analyses. The difference in flower den-
2–3 times per week during the dry periods. sity and yield was analysed by fitting linear regression
Fruit yield and size calibre. For these measure- models, which a one-way ANOVA followed up to test
ments, 10 trees in each irrigation treatment with for the effect of irrigation treatment. Poisson regression
similar flowering intensity and growth vigour were se- models were fitted and followed with ANOVA-type

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Original Paper Plant, Soil and Environment, 69, 2023 (7): 303–313

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500

400
Cummulative ETc, precipitation or irrigation (mm)

300

200
Variable
100 ETc
0 precipatation
irrigation ET-100
irrigation ET-50
500 irrigation RDI-50
400 and RDI-50a

300

200

100

0
Apr May Jun Jul Aug Sep Oct Apr May Jun Jul Aug Sep Oct
Time (months)
Figure 1. Cumulative reference evapotranspiration (ET C), precipitation and irrigation in ET-100, ET-50, RDI-50
and RDI-50a treatments during four consecutive growing seasons. In 2020, the irrigation was the same for in
RDI-50, RDI-50a and ET-100

1.25

1.00

0.75

0.50 Treatment
ET-0
ET-50
ASWC

ET-100
RDI-50
1.25
RDI-50a
1.00

0.75

0.50

Apr May Jun Jul Aug Sep Oct Apr May Jun Jul Aug Sep Oct
Time (months)

Figure 2. Available soil water content (ASWC) in four irrigation treatments (ET-50, ET-100, RDI-50, RDI-50a) and
a non-irrigated control (ET-0) during four consecutive years

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Plant, Soil and Environment, 69, 2023 (7): 303–313 Original Paper

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λ2-tests for the number of fruits in three size catego- Cumulative evapotranspiration reached 492 mm,
ries. Linear regression models with treatment and date while precipitation during May–September period
as factors followed by ANOVA F-tests were used to was only 228 mm, with very little rain during the peak
analyse differences in Ψstem and gs. The models were of vegetation season from June to August (Figure 1A).
fitted separately for each of the four studied years. Hence, an irrigation water supplement of 294 mm
At each measuring date within the season, individual was applied in fully irrigated treatment ET-100. In
treatment means were separated from the non-irrigated contrast, 2020 and 2021 were relatively wet, and pre-
control (ET-0) by pairwise contrasts using the means cipitation was the main water input, with irrigation
function from the means package (Lenth et al. 2019). only 54 mm and 189 mm in fully irrigated treatment
Bonferroni correction was used to adjust the P-values ET-100 in 2020 and 2021, respectively (Figure 1B, C).
for multiple comparisons. Simple linear regression as- The year 2022 was relatively dry, with an exceptional
sessed the relationship between Ψstem, gs and climatic rainfall event of more than 40 mm at the end of June.
conditions and yield. All analyses were carried out In total, 273 mm of irrigation water was applied in
using R (R Development Core Team 2010). fully irrigated treatment (Figure 1D).
Available soil water content was generally the low-
RESULTS est in non-irrigated control ET-0, the highest in fully
irrigated treatment ET-100, and medium in ET-50
Microclimatic conditions. The year 2019 was and RDI treatments, although the measurements were
the driest and hottest of the four studied years. variable in response to precipitation events (Figure 2).

(A)
10.0
Flower density

7.5

5.0

2.5

0
(B) 300
Treatment
Number of fruits

200 ET-0
(1/tree)

ET-50

100 ET-100
RDI-50
RDI-50a
0
(C)
40
Fruit yield

30
(kg/tree)

20
10
0
Irrigation treatment
Figure 3. (A) Flower density; (B) number of fruits and (C) fruit yield of apple trees subjected to four irrigation
treatments (ET-50, ET-100, RDI-50, RDI-50a) and a non-irrigated control (ET-0) during four consecutive years.
The bars are means, and the error bars are standard errors (n = 10). P-values of ANOVA-type F-tests (flower
density, yield) or λ 2-tests (number of fruits) are shown. Significant differences between individual treatments
and ET-0 are indicated with an asterisk

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Original Paper Plant, Soil and Environment, 69, 2023 (7): 303–313

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(A)
50
Number of fruits

40
< 65 mm

30
20
10
0
(B)
Treatment
Number of fruits

100 ET-0
65–75 mm

ET-50
ET-100
50
RDI-50
RDI-50a
0
(C)
Number of fruits

150
> 75 mm

100

50

0
Irrigation treatment
Figure 4. Number of fruits per tree in three size classes (smaller than 65 mm; between 65 and 75 mm; and
larger than 75 mm) in apple trees subjected to four irrigation treatments (ET-50, ET-100, RDI-50, RDI-50a) and
a non-irrigated control (ET-0) during four consecutive years. The bars are means, and the error bars are standard
errors (n = 10). P-values of ANOVA-type λ 2-tests are shown. Significant differences between individual treat-
ments and ET-0 are indicated with an asterisk

In 2020, the rather wet conditions during the period there were statistically significant differences in the
from June to August were evidenced by the small number of fruits among the five treatments, although
difference between ET-0 and ET-100 (Figure 2B). there was no clear pattern of one irrigation treatment
In contrast, the difference between these treat- being consistently better or worse in each of the four
ments was rather high during most of the seasons studied years (Figure 3B). A notable pattern is the
of 2019 and 2022 (Figure 2A, D) and parts of the lack of alternate bearing in the irrigation treatment
season 2021 (Figure 2C). In 2021 and 2022, there was combined with the fruit thinning (RDI-50a), which
a notable divergence of values between RDI-50 and is evident from a relatively high number of fruits and
RDI-50a, irrigated in the same regime and with the high fruit yield in 2020 (Figure 3B, C).
same amount of water (Figure 2C, D). The distribution of fruits in three size classes was
Fruit yield. Flower density showed signs of alternate also quite variable among years and irrigation treat-
bearing behaviour, with 2019 and 2021 being years ments (Figure 4). In low crop years (2020 and 2022),
with high flowering, whereas 2020 and 2022 were there was a low number of small fruits (< 65 mm)
years with low flowering (Figure 3A). The biennial and a high number of large fruits (> 75 mm) per
pattern in flower density was also reflected in the tree in all five irrigation treatments (Figure 4). The
number of fruits at harvest and annual fruit yield differences in fruit size due to irrigation treatment
(Figure 3B, C). Within years, there were no statisti- were variable, with no clear, consistent pattern across
cally significant differences in flower density among years. No clear pattern among treatments was ob-
the five irrigation treatments (Figure 3A). In contrast, served either when the number of fruits in each size

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Plant, Soil and Environment, 69, 2023 (7): 303–313 Original Paper

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–0.75

–1.00

–1.25
Treatment
–1.50
ET-0
Ψ stem (MPa)

ET-50
ET-100
RDI-50
–0.75 RDI-50a

–1.00

–1.25

–1.50

Jun Jul Aug Sep Jun Jul Aug Sep


Time (months)
Figure 5. Seasonal course of midday stems water potential (Ψ stem) in apple trees subjected to four irrigation
treatments (ET-50, ET-100, RDI-50, RDI-50a) and a non-irrigated control (ET-0) during four consecutive years.
The points are means, and the error bars are standard errors (n = 4). P-values of ANOVA-type F-tests are shown.
Significantly different treatment vs control (ET-0) pairwise comparisons are indicated with an asterisk and
colour-coded according to the treatment that was significantly different from the non-irrigated control (ET-0)

category was expressed as relative fractions of the highest in the middle of the growing season. In 2022,
total number of fruits (data not shown). the fully irrigated treatment (ET-100) had slightly
Midday stems water potential. The measured Ψstem higher values of g s compared to the non-irrigated
values were between –0.6 and –1.7 MPa across dates control (ET-0). This difference was the highest at
and irrigation treatments (Figure 5). In three out of the end of the growing season.
the four years of measurements, the non-irrigated Relationships between physiological param-
control (ET-0) had significantly lower values of Ψstem eters, microclimatic parameters and fruit yield.
than fully irrigated trees on most of the measured Both physiological parameters, Ψ stem and g s, were
days (Figure 5). The difference in Ψ stem between significantly and positively related to ASWC across
ET-0 and the fully irrigated treatment (ETC-100) all measured days (Figure 7A, C). R2 was 0.125 for the
was –0.32 MPa at the maximum and –0.16 MPa on ASWC vs Ψstem relationship and 0.061 for the ASWC
average. The differences among treatments were the vs g s . In contrast, both physiological parameters
lowest in season 2020, which was also particular in were significantly negatively related to daily crop
that the RDI-50a treatment had the lowest Ψ stem evapotranspiration (ET C) across all measured days,
values out of all five treatments. with the R 2 being 0.341 for ET C vs Ψ stem and 0.106
Leaf stomatal conductance. The gs values ranged for ET C vs g s relationship (Figure 7B, D). There was
from 16.4 to 672.5 mmol/m2/s across dates and irriga- a significant negative relationship between minimal
tion treatments (Figure 6). There were no significant seasonal Ψstem and annual fruit yield when excluding
differences among treatments in seasons 2019 and 2021 (Figure 8A), implying that more negative Ψstem
2021. In 2020, there was a clear pattern of higher g s was associated with a higher fruit yield (R 2 = 0.734).
in RDI‑50a treatment, with the difference being the There was also a significant positive association

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Original Paper Plant, Soil and Environment, 69, 2023 (7): 303–313

https://doi.org/10.17221/160/2023-PSE

600

400

200
Treatment
g s (mmol/m 2/s)

ET-0
ET-50
ET-100
600 RDI-50
RDI-50a

400

200

Jun Jul Aug Sep Jun Jul Aug Sep


Time (months)
Figure 6. Seasonal course of leaf stomatal conductance (g s ) in apple trees subjected to four irrigation treat-
ments (ET-50, ET-100, RDI-50, RDI-50a) and a non-irrigated control (ET-0) during four consecutive years. The
points are means, and the error bars are standard errors (n = 4). P-values of ANOVA-type F-tests are shown.
Significantly different treatment vs control (ET-0) pairwise comparisons are indicated with an asterisk and
colour-coded according to the treatment that was significantly different from the non-irrigated control (ET-0)

between minimal seasonal g s and annual fruit yield, pressed as alternate bearing behaviour in a biennial
with R 2 being 0.678 (Figure 8B). cycle. Thus, producing small fruits was associated
with higher crop loads during "on" years rather than
DISCUSSION reduced soil water availability.
Irrigation resulted in less negative values of midday
This study monitored fruit yield, midday stem water stem water potential compared to non-irrigated trees
potential and stomatal conductance for four consecu- (Figure 5). The mean observed difference in Ψstem be-
tive years in apple trees cv. Red Jonaprince was subjected tween fully irrigated and non-irrigated trees of 0.16 MPa
to five irrigation treatments (Figures 1 and 2). Despite was not large but sustained in three out of four growing
the significantly lower Ψstem in non-irrigated trees seasons monitored (i.e., in 2019, 2021, 2022). Similar
throughout most of the growing seasons, there was nei- values of Ψstem and similar differences between irrigated
ther a reduction in the overall fruit yield (Figure 3) nor and non-irrigated treatments were reported for apple
a consistently significant effect on fruit size (Figure 4). trees grown in North East Spain (Doltra et al. 2007) and
These results suggest that the yield parameters of in the Golan Heights in Israel (Naor et al. 1995), which
cv. Red Jonaprince apple trees were robust against are both characterised by much drier climate compared
cumulative but mild drought stress. Our results con- to our study site. The seasonal minima of Ψstem ranged
trast with some other studies that found a significant from –1.0 to –1.6 MPa, which can be considered no
fruit size reduction in apple trees that received no or or mild drought stress (De Swaef et al. 2009, Robinson
low irrigation (Naor et al. 1995, 2008, Robinson et et al. 2019). Our observation that non-irrigated trees
al. 2019). In our study, yield parameters were more experienced only mild drought stress agrees with the
affected by the intrinsic interannual variation ex- non-significant effect on yield components.

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(A) (B)

–0.75
–0.8
Ψ stem (MPa)

–1.00

–1.2 –1.25
Treatment
–1.50 ET-0
–1.6
ET-50
–1.75
ET-100
(C) (D) RDI-50
600 600 RDI-50a
Year
g s (mmol/m 2/s)

2019
400 400
2020
2021
200 200 2022

0 0
0.50 0.75 1.00 2 3 4 5
ASWC ETc (mm)
Figure 7. Relationship between (A) available soil water content (ASWC) and midday stem water potential (Ψstem);
(B) crop evapotranspiration (ET C) and Ψ stem; (C) ASWC and leaf stomatal conductance (g s); (D) ETC and g s
across all measuring dates and irrigation treatments. The points represent irrigation treatment means per each
date, and the error bars represent standard error (SE). n = 4 for Ψ stem and g s. The lines are linear regressions to
the data. Corresponding R 2 and P-values are provided. n = 129 for ASWC; n = 150 for ET C

Leaf stomatal conductance did not differ much tivity of gs to irrigation means that gs is a less suitable
among the irrigation treatments, with the exception plant-based water stress indicator than Ψstem, at least
of a clearly higher gs in RDI-50a treatment in season for cv. Red Jonaprince. However, the extrapolation
2020 (Figure 6). The higher values of g s in RDI-50a of these results to other apple tree cultivars should
can be linked with its higher crop load compared be done with caution because the stomatal response
with the other four treatments. Due to fruit thinning and the degree of izo/anizohydry may differ among
in RDI-50a, the trees did not over-cropped in 2019 cultivars (Beikircher et al. 2013, Levin et al. 2020).
and did not show as strong signs of alternate bearing Both physiological parameters Ψ stem and g s were
as the other four treatments. The high sensitivity responsive to atmospheric water demands (Figure 7),
of g s to crop load have been previously reported in which makes the separation of soil- and atmospheric-
apple trees (Palmer et al. 1997, Wünsche et al. 2000) drought difficult for irrigation practice. While the soil
and demonstrates that trees upregulate their leaf water availability and atmospheric water demands
gas exchange to match the higher carbon demands are typically tightly coupled on monthly to seasonal
of growing and ripening fruits (White et al. 2016). scales (Torres et al. 2013, Jupa et al. 2022), atmos-
Similar g s values between irrigated and non-irri- pheric water demands and high air temperatures may
gated trees, while Ψ stem differed, suggest that the have negative impacts on plant hydraulic integrity
apple trees exhibit anizohydric rather than isohydric even under the conditions of non-limiting soil water
behaviour. During anizohydric response, trees do (Schönbeck et al. 2022). This means that there are
not close their stomates during drought stress and situations in which water stress on trees cannot be
tolerate certain declines in water potential (Klein relieved by supplying irrigation.
2014). Our results are in agreement with Beikircher Seasonal minimal Ψ stem was negatively correlated
et al. (2013), who found late stomatal closure and, with annual fruit yield when one year (2021) was
consequently, negative hydraulic safety margins in excluded (Figure 8A). 2021 was wet in the early
three high-yield apple cultivars. The relative insensi- growing season, and flower density in spring was

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(A) 50 Our data shed more light on two commonly meas-


ured plant-based indicators of tree water status. It was
40 found that Ψstem was more sensitive to differences in
Fruit yield (kg/tree)

irrigation than gs, which suggests that Ψstem is a better


30 plant-based indicator for irrigation scheduling than gs.
Differential irrigation and the resulting differences
20 in tree water status did not affect yield parameters,
and hence, irrigation necessity cannot be proved by
10 yield data under mild-humid climatic conditions.
Such non-significant differences are frequently not
0 reported in the scientific literature, which can lead
–1.7 –1.5 –1.3 –1.1 –0.9 to an unwanted publication bias (Dwan et al. 2008).
Seasonal minimal Ψ stem (MPa)
Acknowledgement. We thank Jana Fenclová for
(B) 50
her assistance during field measurements.
40
Fruit yield (kg/tree)

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Received: April 17, 2023
Accepted: June 5, 2023
Published online: July 10, 2023

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