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ORIGINAL PAPER
Isohydrodynamic behavior in deficit-irrigated CabernetSauvignon and Malbec and its relationship between yieldand berry composition
Krista C. Shellie • Pat Bowen
Received: 4 May 2012 / Accepted: 16 September 2013 / Published online: 2 October 2013
� Springer-Verlag Berlin Heidelberg (outside the USA) 2013
Abstract Cabernet Sauvignon and Malbec grapevines
were irrigated at 70 or 23 % of estimated crop evapo-
transpiration throughout berry development over four
growing seasons. Stomatal behavior was characterized by
relating predawn leaf water potential and mid-morning
stomatal conductance to mid-morning leaf water potential.
Seasonal average weekly midday leaf water potential was
lower in Cabernet Sauvignon than Malbec despite similar
irrigation amounts. Both cultivars exhibited anisohydric
behavior with midday leaf water potential decreasing lin-
early with declining predawn leaf water potential
(r2 = 0.51) and stomatal conductance (r2 = 0.42). How-
ever, both cultivars utilized hydrodynamic mechanisms to
maintain a soil-to-leaf water potential gradient of -0.62
(±0.03) MPa under standard irrigation and -0.75
(±0.04) MPa under reduced irrigation. Berry fresh weight
and titratable acidity decreased, and the concentration of
total anthocyanins increased in both cultivars in response to
decreases in midday leaf water potential. The slope of
regression equations for seasonal mean midday leaf water
potential was used to estimate cultivar-specific levels of
water stress associated with changes in berry weight and
berry composition at fruit maturity.
Introduction
Wine grapes (Vitis vinifera L.) have traditionally been
grown without irrigation in Europe because supplemental
water has been associated with reduced quality for wine
production (Cifre et al. 2005). Irrigation is now used
extensively in semiarid production regions but compromise
between fruit quality and yield remains a major issue of
importance and an active area of research (Cifre et al.
2005; Chaves et al. 2010; Lovisolo et al. 2010; Romero
et al. 2013). Delineation of desirable severities of water
deficit has been complicated by inherent cultivar differ-
ences in drought sensitivity, interactive effects with abiotic
conditions, and limited definitive information relating vine
water status parameters with berry composition at maturity
(De Souza et al. 2005; Romero et al. 2010, 2013; Bowen
et al. 2011).
Wine grape cultivars differ in their response to drought
(Bota et al. 2001; Chaves et al. 2010; Lovisolo et al. 2010),
and the underlying mechanisms responsible for these dif-
ferences remain poorly understood (Schultz 2003; Van-
deleur et al. 2009). Many wine grape cultivars have been
categorized as isohydric or anisohydric according to the
changes in leaf water potential in relation to soil moisture
content; however, there is discrepancy among studies in
cultivar classification, and individual grapevines have been
observed to change from isohydric-like behavior when
transpiration is low to anisohydric-like behavior with
increasing water demand (Chaves et al. 2010; Lovisolo
et al. 2010; Domec and Johnson 2013). Stomatal behavior
has been found to differ in field-grown versus potted vines
(Schultz 2003; Chaves et al. 2010).
Leaf water potential, measured predawn or midday, and
stomatal conductance have been proposed as indicators of
water status for monitoring water-deficit severity and
Communicated by S. Ortega-Farias .
K. C. Shellie (&)
Horticultural Crops Research Unit, USDA-ARS, 29603 U of I
Lane, Parma, ID 83660, USA
e-mail: [email protected]
P. Bowen
Pacific Agri-Food Research Centre, Summerland, BC VOH 1Z0,
Canada
123
Irrig Sci (2014) 32:87–97
DOI 10.1007/s00271-013-0416-y
scheduling irrigation events (Cifre et al. 2005; Intrigliolo
and Castel 2006; Shellie 2006; Bowen et al. 2011). Pre-
dawn measurements of water potential have been used in
many grape studies as the standard to which other measures
of vine performance are compared; however, differences
detected predawn have not always been apparent at midday
(Correia et al. 1995). Strong correlations have also been
observed between leaf water potential, leaf gas exchange,
and soil moisture content (Williams and Araujo 2002;
Williams and Trout 2005; Williams 2012; Williams et al.
2012). Optimum thresholds for stomatal conductance and
midday leaf water potential have been proposed but there
has been limited comparative evaluation of different cul-
tivars under identical field conditions (Cifre et al. 2005;
Chaves et al. 2010; Lovisolo et al. 2010).
Cabernet Sauvignon is widely grown in arid regions
where irrigation is essential for wine grape production. Its
drought response has been categorized as both isohydric
and anisohydric (Bota et al. 2001; Williams and Baeza
2007; Chaves et al. 2010; Hochberg et al. 2013). World
production of Malbec is one-tenth the amount of Cabernet
Sauvignon but its production acreage is steadily increasing.
There is limited available information describing the sto-
matal behavior of Malbec in response to drought. The
objective of this research was to compare the drought
response of Cabernet Sauvignon and Malbec under iden-
tical field conditions when irrigated with similar amounts
of water and to relate their drought response with changes
in yield components and berry composition at fruit matu-
rity. A practical aim was to generate information about
these cultivars that could be used to customize irrigation
management practices to meet particular production goals.
Materials and methods
The trial was conducted over four growing seasons
(2007–2010) in an experimental vineyard located at the
University of Idaho Parma Research and Extension Center
in Parma, ID (43.78�N; 116.94�W; elevation 750 m asl).
Climate at this site has a Koeppen classification of BSK,
which is midlatitude steppe climate. Precipitation during
the growing season (April–October) accounts for *44 %
of annual. Soil at the site was a fine sandy loam (Turbyfill
series, Xeric Torriorthent Entisol) with an available water-
holding capacity of 0.14 cm per cm of soil (US Department
of Agriculture Soil Conservation Service 1972) over a
hardpan at a variable depth to 1 m.
Planting material was obtained from Foundation Plant
Services (University of California, Davis, CA, USA) in
1997 as ungrafted, dormant rooted cuttings. Vines were
planted in eight-vine panels in four replicate blocks, each
block having three rows of 56 vines. The row by vine
spacing was 2.7 by 2.1 m. Row orientation was north to
south. The vines had bilateral cordons with each cordon
supported by a separate trunk at a height of 102 cm above
the soil surface. Shoots were positioned vertically, spur-
pruned to about 30 buds (14–16 buds per meter of cordon)
per vine, and thinned around bloom to *16 shoots per
meter. Nutrient, pest, and disease management practices
were uniformly applied each year according to standard
commercial practice. Chemical and mechanical methods
were used to keep alley and in-vine rows weed-free
throughout the growing season.
The experimental design was a split-plot with each
cultivar in an eight-vine main plot randomly located within
each block. Two irrigation amounts, standard or reduced,
were applied to 4-vine subplots within each main plot. The
interior two vines of each subplot were used for data col-
lection, and the same vines were evaluated each growing
season. The vineyard was equipped to irrigate with two
lines of aboveground drip tubing (Bowsmith, 16 mm i.d.)
suspended 30 cm above the soil surface. One drip line
contained in-line emitters with a flow rate of 1.9 L h-1
spaced 1 m apart (two emitters per vine). A second line of
drip tubing without emitters was located adjacent to the
other line, and only contained emitters (flow rate of
3.8 L h-1) in subplots that received the standard amount of
irrigation. Manually inserted emitters were located midway
between the in-line emitters of the other drip line. Thus,
when both drip lines were employed during an irrigation
event, threefold more water was delivered to standard
versus reduced irrigated subplots.
All vines received equal amounts of water prior to fruit
set and after harvest by employing only the in-line emitter
drip line. Both drip lines were employed in all irrigation
events between fruit set and harvest. Irrigation amount was
calculated weekly using the Penman–Monteith model
(Allen et al. 1998), with well-watered alfalfa as the refer-
ence crop (ETr), and a crop coefficient that increased pre-
veraison from 0.2 to 0.7 and decreased post-veraison to 0.4,
similar to Keller et al. (2008). Values for ETr were
obtained from a weather station located in close proximity
to the trial site (http://www.usbr.gov/pn/agrimet/wxdata.
html). Vines under the standard irrigation treatment were
supplied 70 % of estimated crop evapotranspiration (ETc),
which represented the regional industry standard (Keller
et al. 2008). The crop coefficient was adjusted during the
growing season to maintain the midday leaf water potential
(mdWL) of vines under standard irrigation at *-1.0 MPa
(Williams et al. 2010). The amount of water delivered to
each block was measured by flow meters during each
irrigation event. The irrigation treatments were first applied
in 2006.
Soil moisture was measured during three growing sea-
sons (2007–2009) using a data-logging, time domain
88 Irrig Sci (2014) 32:87–97
123
reflectometry (TDR) system and segmented probes
(Moisturepoint, ESI, Victoria, BC, Canada). Each probe
measured average volumetric moisture content at five
depths: 0–15, 16–30, 31–45, 46–60, and 61–90 cm. Two
probes were installed midway between an emitter and vine,
18 cm toward the alley away from the vine row in the
standard and the reduced irrigation subplots within each
cultivar main plot in a block. Soil moisture at each probe
segment was measured and logged hourly.
The day preceding an irrigation event, the mdWL of two,
fully expanded, exposed leaves was measured in each
subplot using a pressure chamber (PMS Instruments model
610 Corvallis, OR, USA) as described by Turner (1988).
Leaf blades were covered with a plastic bag prior to petiole
excision and remained in the bag during measurement.
Elapsed time between excision and chamber pressurization
was *15 s. Leaf water potential was measured predawn
(pdWL), and stomatal conductance (gs) was measured mid-
morning (L1-1600 steady-state porometer; Ll-COR, Lin-
coln, NE, USA) on cloudless days prior to and after ver-
aison in 2007 and on three sampling dates post-veraison in
2009.
Fruit was harvested when a composite juice sample of
ten clusters collected from non-data vines in each repli-
cate block had a soluble solids concentration (SS) of
*24 % and a titratable acidity (TA) of *6 g L-1. On the
day of harvest, ten clusters were collected from the east-
and west-facing canopy of the data vines in each cultivar
subplot. One hundred berries were subsampled from these
clusters and used to calculate average berry fresh weight
and to measure total monomeric anthocyanins (Iland et al.
2004). The remaining berries on the 10 cluster sample
were crushed, left overnight on the skins at 21 �C, and
then analyzed the following day for SS (refractive index
detector RE40, Mettler-Toledo, Columbus, OH, USA), pH
and TA (Metrohm 716 DMS Titrino, Brinkmann, Herisau,
Switzerland) following methods described by Shellie
(2006).
Seasonal cumulative growing degree days (GDD)
were calculated from daily maximum (no upper limit)
and minimum temperatures measured at the Parma
Experiment Station weather station (US Department of
Interior, Bureau of Reclamation, Pacific Northwest
Cooperative Agricultural weather network) using a base
threshold of 10 �C. Data for measured variables were
analyzed using a mixed model analysis of variance with
irrigation, cultivar, and their interaction as fixed effects.
Differences between mean values for significant
(P B 0.05) main effects were detected using Tukey–
Kramer adjusted t test (SAS version 8.02; SAS Institute,
Cary, NC, USA). Graphs and regression analyses were
generated using SigmaPlot 11.2 (Systat Software, Inc.
San Jose, CA, USA).
Results
Seasonal GDD accumulation was lowest, and harvest was
latest in 2010 relative to the other study years (Table 1;
Fig. 1). The 4-year average seasonal amount of precipita-
tion was 8 % of seasonal ETr and precipitation was lowest
in 2008 (Table 1). Over the four-year study, standard irri-
gated vines were provided *27 % of seasonal ETr. Vines
under reduced irrigation were provided 33 % less water
than vines under standard irrigation in each of the last three
study years, and 46 % less water than standard in the first
year of the study. Irrigation amount in relation to ETr
reached a maximum at veraison in all years except 2009,
when irrigation system malfunctions resulted in reduced
amounts of water supplied to all plots for two consecutive
irrigations (Fig. 1a–d). Cabernet Sauvignon had lower
weekly values of mdWL than Malbec throughout most of
each growing season (Fig. 1e–h). Vines under standard
irrigation had at least one weekly value of mdWL that was
less than -1.0 MPa in each year, with the exception of
Malbec in 2010.
Soil moisture in the top 60 cm increased in response to
weekly irrigation events and then declined to pre-irrigation
levels with little accumulation between irrigation events
(Fig. 2). The one exception to this trend was in 2007 where
soil moisture increased gradually over time in standard
irrigated plots. Soil moisture at the 60–90 cm depth
increased gradually over time in both irrigation plots in
2007 and was similar in standard and reduced irrigated
plots in 2008 and 2009.
Vines under reduced irrigation had lower seasonal
average weekly mdWL than vines under standard irrigation
in each year of the study (Table 2). Cultivar differences in
seasonal average weekly mdWL were detected in 2007,
2008, and 2010. In 2007, seasonal average weekly mdWL
was lower in Cabernet Sauvignon (-1.23 MPa) than
Table 1 Seasonal growing degree days (GDD), precipitation (Pcp),
reference evapotranspiration (ETr), and amount of water supplied to
vines of Cabernet Sauvignon and Malbec grown in Parma ID under
standard or reduced irrigation treatments
Year GDDa (�C) Pcpa (mm) ETra (mm) Irrigation amount
(mm)
Standard Reduced
2007 1,750 97 1,294 366 170
2008 1,659 63 1,276 243 81
2009 1,689 111 1,220 426 142
2010 1,569 141 1,197 321 107
4-year 1,667 103 1,247 339 125
a Accumulated Pcp, Kimberly–Penman alfalfa-based ETr, and GDD
April 1 to October 31 from Agrimet weather station at Parma ID.
Cumulative GDD is daily average temperature above 10 �C
Irrig Sci (2014) 32:87–97 89
123
180 200 220 240 260-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4
Mid
day
leaf
wat
er p
oten
tial(
MPa
)
-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4
180 200 220 240 260-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4
Mid
day
leaf
wat
er p
oten
tial
-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4CS, reducedCS, standardMB, reducedMB, standard
180 200 220 240 260-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4
Mid
day
leaf
wat
er p
oten
tial
-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4
Day of year
180 200 220 240 260 280-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4
Mid
day
leaf
wat
er p
oten
tial
-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4
180 200 220 240 260 280
Irri
gate
d am
ount
: E
Tr
0.0
0.2
0.4
0.6
0.8
1.0
Gro
win
g de
gree
day
s (0
C)
200
400
600
800
1000
1200
1400
1600
180 200 220 240 260 280
Irri
gate
d am
ount
: ET
r
0.0
0.2
0.4
0.6
0.8
1.0
Gro
win
g de
gree
day
s (0 C
)
200
400
600
800
1000
1200
1400
1600
180 200 220 240 260 280
Irri
gate
d am
ount
: E
Tr
0.0
0.2
0.4
0.6
0.8
1.0
Gro
win
g de
gree
day
s (0
C)
200
400
600
800
1000
1200
1400
1600
Day of year
180 200 220 240 260 280
Irri
gate
d am
ount
: E
Tr
0.0
0.2
0.4
0.6
0.8
1.0
Gro
win
g de
gree
day
s (0
C)
200
400
600
800
1000
1200
1400
1600
(e)
(d)
(f)
(g)
(h)
V H
V
V
V
H
H
H
(b)
(c)
(a)
2010
2007
2008
2009
Fig. 1 Ratio of irrigation amount to reference evapotranspiration
(ETr) under reduced (solid circle) or standard (open circle) irrigation
and accumulated GDD (solid square) 2007–2010 (a–d). Day of year
is indicated for veraison (V) and harvest (H). Weekly midday leaf
water potential of cultivars Cabernet Sauvignon (CS) (circle) and
Malbec (MB) (squares) 2007–2010 (e–h). Error bars for midday leaf
water potential depict standard error for weekly mean values
90 Irrig Sci (2014) 32:87–97
123
Malbec (-1.01 MPa) and was lower under reduced
(-1.28 MPa) than standard (-0.96 MPa) irrigation. In
2008, seasonal average mdWL was similar for the two cul-
tivars under standard irrigation but, under reduced irriga-
tion, were lower in Cabernet Sauvignon than Malbec. In
2009, both cultivars had higher seasonal average weekly
mdWL under standard (-1.0 MPa) than reduced irrigation
(-1.37 MPa). In 2010, the seasonal average weekly mdWL
of Cabernet Sauvignon under standard irrigation was sim-
ilar to that of Malbec under reduced irrigation. The 4-year
seasonal average weekly mdWL was -0.98 MPa under
standard irrigation and -1.31 MPa under reduced
irrigation.
The soil-to-leaf water potential gradient at mid-morning
(DWplant), calculated as the difference between predawn
and mid-morning leaf water potential, was similar in both
cultivars in 2007 and 2009 (Table 2). In 2007, vines under
either irrigation amount had similar DWplant; however, in
2009, DWplant was 0.13 MPa lower in reduced relative to
standard irrigated vines in both cultivars (Table 2). Sto-
matal conductance (gs) was about 45 % lower in vines of
both cultivars under reduced relative to standard irrigation
(Table 2). Predawn leaf water potential was linearly related
to mid-morning measurements of leaf water potential
(r2 = 0.51, P \ 0.01) and gs (r2 = 0.42, P \ 0.01) despite
different ambient temperature and evaporative demand on
Vol
umet
ric m
oist
ure
(%)
5
10
15
20
25
30210 215 220 225 230 235 210 215 220 225 230 235
Vol
umet
ric m
oist
ure
(%)
5
10
15
20
25
30
Vol
umet
ric m
oist
ure
cont
ent (
%)
5
10
15
20
25
30
Vol
umet
ric m
oist
ure
(%)
5
10
15
20
25
30
0-60 cm61-90 cm
Day of year
Vol
umet
ric m
oist
ure
cont
ent (
%)
5
10
15
20
25
30
35
Day of year
210 215 220 225 230 235 210 215 220 225 230 235
Vol
umet
ric m
oist
ure
(%)
5
10
15
20
25
30
35
(a)
(b)
(c)
(d)
(e)
(f)
Standard irrigation Reduced irrigation
Fig. 2 Volumetric soil
moisture content during four
typical irrigation events in 2007
(top row), 2008 (middle row),
and 2009 (bottom row) under
standard (a–c) or reduced (d–
f) amounts of irrigation
Irrig Sci (2014) 32:87–97 91
123
each of five sampling dates (Fig. 3). Ambient temperature
and air vapor pressure deficit at mid-morning (10:00 MDT)
on the two sampling dates in 2007 and three sampling dates
in 2009 were as follows: 28.6, 23.8, 24.3, 18.9, and 18.3 �C
and 2.8, 2.0, 1.7, 1.3, and 0.7 kPa, respectively. Mid-
morning leaf water potential declined in association with
pdWL and mid-morning gs over a range of pdWL from -0.10
to -0.9 MPa. Across all sampling dates, mean pdWL was
about 0.2 MPa higher under standard than reduced irriga-
tion, with a mean value of -0.49 MPa in Cabernet Sau-
vignon and -0.46 MPa in Malbec. Mean mid-morning leaf
water potential was about 0.3 MPa higher under standard
than reduced irrigation, and mean values across sampling
dates were about 0.10 MPa lower in Cabernet Sauvignon
than Malbec (-1.21 vs. -1.11 MPa). DWplant across sam-
pling dates and cultivars was 0.13 MPa higher under
standard than reduced irrigation. DWplant was -0.62
(±0.03) MPa under standard irrigation when pdWL ranged
from -0.10 to -0.65 MPa and gs ranged from 50 to
500 mmol m-2s-1. DWplant was -0.75 (±0.04) MPa under
reduced irrigation when pdWL ranged from -0.20 to -
0.90 MPa and gs ranged from\25–370 mmol m-2s-1. The
linear relationships between pdWL or mid-morning gs and
DWplant were nearly horizontal, indicating that, although
DWplant differed by irrigation amount, it was similar in both
cultivars and was unrelated to mid-morning gs or pdWL.
Pronounced variability in leaf water potential under
differing amounts of soil moisture was exhibited in both
cultivars, and the strength of the relationship was identical
(r2 = 0.51). However, mid-morning leaf water potential
was lower in Cabernet Sauvignon than Malbec (Fig. 3a).
Table 2 Seasonal average weekly midday leaf water potential
(mdWL), soil-to-leaf water potential gradient at mid-morning
(DWplant), and mid-morning stomatal conductance (gs) in Cabernet
Sauvignon and Malbec vines irrigated with a standard or reduced
amount of water between fruit set and harvest grown in Parma ID
Year Seasonal average weekly mdWL (MPa) Mid-morning DWplant (MPa) Mid-morning gs (mmol m-2 s-1)
2007 2008 2009 2010 Four years 2007 2009 Two years 2007 2009 Two years
Cabernet Sauvignon
Standard -1.06 -1.05 -1.03 -1.03 -1.04 -0.60 -0.67 267.66 307.13 289.23
Reduced -1.40 -1.47 -1.42 -1.27 -1.39 -0.79 -0.80 147.53 171.15 159.59
Malbec
Standard -0.86 -1.04 -0.97 -0.80 -0.92 -0.59 -0.60 359.95 300.15 321.96
Reduced -1.16 -1.30 -1.31 -1.09 -1.22 -0.65 -0.74 127.56 208.57 174.06
Irrigation * ** ** ** ns * ** **
Cultivar ** * ns ** ns ns ns ns
I 9 C ns * ns * ns ns ns ns
Least square mean values from a mixed model analysis of variance with irrigation (I), cultivar (C), and their interaction as fixed effects
*, **, ns indicates P B 0.05, 0.01, not significant, respectively
Predawn leaf water potential (MPa)
Mid
-mor
ning
leaf
wat
er p
oten
tial (
MP
a)
-2.0
-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4
Leaf
wat
er p
oten
tial g
radi
ent (
MP
a)
-2.0
-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4
CS regressionMB regression
Mid-morning stomatal conductance (mmol m-2 s-1)
-1.0 -0.9 -0.8 -0.7 -0.6 -0.5 -0.4 -0.3 -0.2 -0.1 0.0
0 100 200 300 400 500 600Mid
-mor
ning
leaf
wat
er p
oten
tial (
MP
a)
-2.0
-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4
Leaf
wat
er p
oten
tial g
radi
ent (
MP
a)
-2.0
-1.8
-1.6
-1.4
-1.2
-1.0
-0.8
-0.6
-0.4
Leaf water potential CSLeaf water potential MBGradient CSGradient MB
(b)
(a)
Fig. 3 Relationships between predawn leaf water potential (a), and
mid-morning stomatal conductance (b), mid-morning leaf water
potential (open symbols) and the soil-to-leaf water potential gradient
(closed symbols) in Cabernet Sauvignon (CS) and Malbec (MB) that
were irrigated with a standard or reduced amount of water and grown
in Parma ID in 2007 and 2009
92 Irrig Sci (2014) 32:87–97
123
Mid-morning gs was more strongly related to mid-morning
leaf water potential in Cabernet Sauvignon than Malbec
(r2 = 0.48 vs. 0.37) and was higher or lower in Cabernet
Sauvignon than Malbec depending on whether mid-morn-
ing leaf water potential was above or below -1.0 MPa,
respectively (Fig. 3b).
Crop load, measured as the ratio of yield to pruning
weight, was highest in 2007 in both cultivars due to low
vine vigor (Table 3). Crop load declined between 2007 and
2009 in both cultivars as vine vigor increased threefold. In
Cabernet Sauvignon, the decline in crop load was due to
smaller seasonal increases in yield and cluster number per
vine relative to vine vigor. In Malbec, seasonal declines in
crop load were due to sequential declines in yield and
cluster number per vine. Berry fresh weight was lower in
Cabernet Sauvignon relative to Malbec in each year of the
study.
Reduced irrigation decreased yield, berry fresh weight,
and pruning weight in at least three out of 4 years in each
cultivar (Table 3). Reduced irrigation decreased yield per
vine each year in both cultivars, with the exception of
Malbec in 2010. The average yield reduction was 37 % for
Cabernet Sauvignon and 21 % for Malbec. The number of
clusters per vine was lower under reduced irrigation in both
cultivars in the last 2 years of the study. Reduced irrigation
decreased berry fresh weight by *18 % in both cultivars
and decreased the 4-year average pruning weight by 50 %
in Cabernet Sauvignon and 31 % in Malbec. Average shoot
length under standard irrigation was 166 cm in Cabernet
Sauvignon and 180 cm in Malbec, and was 30 and 25 %
shorter, respectively, under reduced irrigation (data not
shown).
Fruit was harvested at similar levels of SS and pH in
both cultivars in three out of 4 years (Table 4). In 2008,
Malbec had *2 % higher SS concentration than Cabernet
Sauvignon. In 2007, Malbec had 0.1 unit lower pH than
Cabernet Sauvignon. Vines under reduced irrigation pro-
duced fruit with higher pH, lower TA, and higher antho-
cyanins in at least three out of 4 years (Tables 4, 5).
Reduced irrigation had no effect on SS in three out of
4 years but increased SS by 3 % in 2010 (Table 4).
Reduced irrigation increased must pH by *4 % and
decreased TA by *25 %. Berries from vines under
reduced irrigation had *11 % higher concentration of total
anthocyanins (Table 5).
Berries of Cabernet Sauvignon had lower TA and total
anthocyanins than berries of Malbec in three out of 4 years
(Tables 4, 5). In Cabernet Sauvignon, anthocyanin con-
centrations did not change over years as vine vigor and
yield increased. In Malbec, the concentration of anthocy-
anins was lowest in 2007 when crop load was highest.
Berry fresh weight, TA, and total anthocyanins were
related to average weekly mdWL during berry development,
but the strength and nature of the relationships varied by
cultivar (Fig. 4). A sequential increase in deficit severity
from a mdWL of -1.2 MPa to -1.4 MPa to -1.6 MPa was
associated in Cabernet Sauvignon with a 6 % decrease in
berry fresh weight at each sequential decrease in mdWL. In
Malbec, berry fresh weight decreased 8, 9, and 10 % for
each increase in deficit severity. The percent increase in
total anthocyanin concentration associated with each
sequential increase in water stress severity was 3, 8, and
13 % in Cabernet Sauvignon and 8, 7, and 7 % in Malbec,
respectively. The ratio of percent increase in total
Table 3 Yield components in Cabernet Sauvignon and Malbec irrigated with a standard or reduced amount of water over four growing seasons
(2007–2010) in Parma ID
Year Yield (kg/vine) Pruning weight (kg/vine) Berry fresh weight (g) Cluster number/vine
07 08 09 10 Three
years
07 08 09 10 Four
years
07 08 09 10 Four
years
07 08 09 10 Three
years
Cabernet Sauvignon
Standard 3.9 na 5.5 5.4 4.9 0.5 1.0 1.6 1.7 1.2 1.1 1.0 1.2 1.0 1.1 32 na 40 41 38
Reduced 2.0 na 3.6 3.6 3.1 0.3 0.5 0.8 0.9 0.6 0.8 0.9 1.1 1.0 0.9 38 na 35 29 34
Malbec
Standard 8.6 na 6.0 2.8 5.8 0.6 1.0 1.9 1.7 1.3 1.6 1.7 1.7 1.5 1.6 70 na 46 37 51
Reduced 5.6 na 5.3 2.9 4.6 0.5 0.7 1.2 1.3 0.9 1.1 1.4 1.4 1.3 1.3 64 na 43 36 48
Irrigation
(I)
* na * * * * ** * * ** ** ns ns na * *
Cultivar
(C)
** ns ns ** * ns ns ns * ** ** ** ** na ** ns
I 9 C ns ns ns * ns ns ns ns ns ns ns ns ns na ns ns
Least square mean values from a mixed model analysis of variance with irrigation (I), cultivar (C), and their interaction as fixed effects
*, **, ns indicates P B 0.05, 0.01, not significant, respectively
Irrig Sci (2014) 32:87–97 93
123
anthocyanins relative to decrease in berry fresh weight for
each sequential increase in deficit severity was 0.5, 1.3, and
2.2 in Cabernet Sauvignon and 1, 0.8, and 0.7 in Malbec. A
beneficial increase in total anthocyanin concentration
without a concomitant decrease in berry weight was
attained at a -1.2 MPa in Malbec and -1.4 MPa water
stress in Cabernet Sauvignon.
Discussion
Irrigation could be used more effectively to conserve water
and optimize vine productivity and fruit quality if its
management could be customized to accommodate for
different drought response mechanisms among cultivars of
wine grape. Based upon generic stress severity thresholds
for gs and mdWL (Cifre et al. 2005, Lovisolo et al. 2010),
vines in this study under standard irrigation experienced
little if any water deficit (mean gs was 315 mmol m-2 s-1
and mdWL was -0.98 MPa) and vines under reduced irri-
gation experienced a moderate level of water deficit (mean
gs was 153 mmol m-2 s-1 and mdWL was -1.31 MPa).
The mean mdWL of Malbec under standard or reduced
irrigation in this study was similar to the mean mdWL of
field-grown Thompson Seedless grapevines provided 60 or
20 % ETc (Williams et al. 2010), but the mean mdWL of
Cabernet Sauvignon was lower than Thompson Seedless
under both irrigation amounts (Table 1).
Cabernet Sauvignon and Malbec exhibited the classic
anisohydric behavior. Their mdWL varied widely in
response to varying levels of pdWL when irrigated with
standard or reduced amounts of water (Fig. 3). Strong
correlations between midday and predawn leaf water
potential have also been observed in field-grown Thomp-
son Seedless, Chardonnay, Cabernet Sauvignon, and
Tempranillo (Williams and Araujo 2002; Williams and
Trout 2005; Intrigliolo and Castel 2006). Williams and
Araujo (2002) reported a stronger correlation between
predawn and midday leaf water potential (r2 = 0.88) than
we observed in this study (Fig. 3). Our data were most
likely more variable because it included five rather than
two sampling dates, and midday measurements of leaf
water potential are influenced by ambient conditions.
An interesting finding from this study was that DWplant
was maintained at a similar level by both cultivars over low
and high values of predawn leaf water potential and rates of
stomatal conductance (Table 2; Fig. 3). Franks et al.
(2007) found that the transpiration-induced water potential
gradient from roots to shoots, measured as the difference
Table 4 Berry maturity indices at harvest for Cabernet Sauvignon and Malbec irrigated with a standard or reduced amount of water over four
growing seasons (2007–2010) in Parma ID
Year Soluble solids (%) pH Titratable acidity (g L-1)
2007 2008 2009 2010 Four years 2007 2008 2009 2010 Four years 2007 2008 2009 2010 Four years
Cabernet Sauvignon
Standard 22.9 22.8 23.9 23.1 23.2 3.7 3.4 4.0 3.6 3.7 5.25 7.16 3.92 4.92 5.31
Reduced 23.5 24.0 24.1 23.8 23.8 3.9 3.6 4.2 3.8 3.9 3.93 4.74 3.11 3.63 3.85
Malbec
Standard 22.6 24.8 23.5 23.2 23.5 3.5 3.5 4.1 3.6 3.7 7.03 6.75 4.51 7.56 6.46
Reduced 23.8 25.4 24.1 24.3 24.4 3.8 3.6 4.1 3.8 3.8 4.80 5.63 3.81 5.44 4.92
Irrigation ns ns ns * ** ** * ** ** ** ** **
Cultivar ns ** ns ns * ns ns ns * ns ** **
I 9 C ns ns ns ns ns ns ns ns ns ns ns ns
Least square mean values from a mixed model analysis of variance with irrigation (I), cultivar (C), and their interaction as fixed effects
*, **, ns indicates P B 0.05, 0.01, not significant, respectively
Table 5 Total monomeric anthocyanin concentration in Cabernet
Sauvignon and Malbec berries harvested from vines irrigated with a
standard or reduced amount of water for four growing seasons
(2007–2010) in Parma ID
Year Total monomeric anthocyanins
(mg/g fresh weight)
2007 2008 2009 2010 Four years
Cabernet Sauvignon
Standard 1.10 1.22 1.09 1.08 1.12
Reduced 1.07 1.43 1.25 1.21 1.24
Malbec
Standard 1.13 2.67 1.80 2.48 2.02
Reduced 1.46 2.81 1.96 2.70 2.23
Irrigation ns * * *
Cultivar ns ** ** **
I 9 C ns ns ns ns
Least square mean values from a mixed model analysis of variance
with irrigation (I), cultivar (C), and their interaction as fixed effects
*, **, ns indicates P B 0.05, 0.01, not significant, respectively
94 Irrig Sci (2014) 32:87–97
123
between predawn and midday leaf water potential, in
Eucalyptus gomphocephala was maintained at -0.67 MPa
across seasons despite a strong correlation between pre-
dawn and midday leaf water potential and strong stomatal
down-regulation under increasing evaporative demand.
They referred to this pattern of hydraulic regulation as
isohydodynamic. Similar isohydrodynamic hydraulic reg-
ulation has been recently reported in the wine grape cul-
tivar Merlot (Zhang et al. 2013). Embolism formation and
recovery in xylem tissue may be one mechanism involved
with dynamic changes in hydraulic conductance (Franks
et al. 2007). The mean mdWL of Cabernet Sauvignon in this
study under reduced irrigation was at a level that others
have found to be associated with up to 50 % loss in
hydraulic conductivity and induction of xylem embolisms
(Alsina et al. 2007). The consistently lower mdWL of
Cabernet Sauvignon relative to Malbec at similar levels of
pdWL suggests that these two cultivars may differ in their
sensitivity to or type of hydrodynamic mechanisms utilized
to achieve an isohydrodynamic state.
The low pdWL and mdWL in Cabernet Sauvignon relative
to Malbec observed in this study support field observations
of Williams and Araujo (2002) where Cabernet Sauvignon,
irrigated at 0 or 50 % ETc, had lower predawn and midday
leaf water potential than Chardonnay even though both
cultivars were grafted to 5C rootstock. Cabernet Sauvignon
also had the lowest mdWL of three field-grown cultivars that
were irrigated at fractional percentages of ETc (Williams
and Baeza 2007). Understanding the factors responsible for
low leaf water potential in Cabernet Sauvignon could
enhance our understanding of cultivar differences in
drought response.
The influence of water deficit on yield components,
vegetative growth, leaf gas exchange, and berry composi-
tion followed similar trends in both cultivars, but water
deficit was more severe in Cabernet Sauvignon than Mal-
bec. The water deficit imposed by reduced irrigation (23 %
ETc) inhibited vegetative and reproductive growth in each
cultivar, as evidenced by decreased pruning weight and
berry fresh weight under reduced irrigation (Table 3).
Water deficit has been reported to reduce berry fresh
weight in some (Roby and Matthews 2004; Shellie 2006;
Keller et al. 2008), but not all studies (Bowen et al. 2011).
In the 3 years that we observed a reduction in berry fresh
weight under reduced relative to standard irrigation,
the average difference in seasonal weekly mdWL was
0.3 MPa in Malbec and 0.4 MPa in Cabernet Sauvignon
when water deficit was mild in Malbec (3-year mean
-1.25 MPa) and moderate in Cabernet Sauvignon (3-year
mean -1.43 MPa).
Berry SS has been reported to increase, decrease, or not
respond to water deficit (Roby and Matthews 2004; Sivi-
lotti et al. 2005; Shellie 2006; Cramer et al. 2007; Sadras
et al. 2007; Keller et al. 2008; Romero et al. 2010; Bowen
et al. 2011). The increase in SS we observed under water
deficit was smaller than the percent decrease we observed
in berry fresh weight, suggesting that total sugar production
per vine was lower under reduced than standard irrigation.
Roby et al. (2004) collected similar sized berries from
vines under deficit-irrigated and well-watered vines and
found that SS was consistently lower in berries from well-
watered vines. They also showed that total soluble solids
per berry increased approximately linearly with berry size
but deviations from proportionality suggested that SS
Midday leaf water potential (MPa)
Ber
ry fr
esh
wei
ght (
g)
0.60.70.80.91.01.11.21.31.41.51.61.71.81.92.0
CS: r=0.41, p=0.002, y=1.40+0.303xMalbec: r=0.40, p< 0.01, y=2.13+0.619x
Midday leaf water potential (MPa)
Ber
ry a
ntho
cyan
in (
mg
g fr
wt)
0.5
1.0
1.5
2.0
2.5
3.0
3.5
CS: r=0.51, p<0.01, y=1.95+1.633x+0.810x2
Malbec: r=0.33, p=0.01, y=1.31 - 0.880xCS reducedCS standardMB reducedMB standard
(a)
Midday leaf water potential (MPa)
-1.6 -1.4 -1.2 -1.0 -0.8 -0.6 -1.6 -1.4 -1.2 -1.0 -0.8 -0.6-1.6 -1.4 -1.2 -1.0 -0.8 -0.6
Titr
atab
le a
cidi
ty (
g L
-1)
2
4
6
8
10
CS: r=0.48, p< 0.01, y=8.69+3.372xMalbec: r=0.55, p< 0.01, y=10.67+4.67x
(b) (c)
Fig. 4 Relationships between seasonal average weekly midday leaf
water potential and berry fresh weight (a), titratable acidity (b), and
total anthocyanins (c) in Cabernet Sauvignon (circle) and Malbec
(square) grown under reduced (solid symbols) or standard (open
symbols) irrigation in Parma ID over four growing seasons
(2007–2010)
Irrig Sci (2014) 32:87–97 95
123
concentration increased with decreasing berry size. Basile
et al. (2011) reported a bell-shaped quadratic relationship
between SS and leaf water potential and attributed
increases in SS to a concentration effect rather than
increased accumulation.
The reduction in TA we observed at harvest under
reduced irrigation has also been reported by others (Shellie
2006; Keller et al. 2008; Romero et al. 2010; Basile et al.
2011; Bowen et al. 2011). A decrease in organic acids
under water stress has been attributed to increased respi-
ration of malic acid induced by a deficit-related increase in
cluster microclimate temperature and a shift in carbon
skeletons for the production of amino acids from glycolysis
rather than assimilates directly from the chloroplast
(Lawlor and Cornic 2002; Romero et al. 2010). Deficit-
related decreases in vigor and gs have been associated with
increased leaf and berry temperature due to increased
canopy light transmission and decreased transpirational
cooling (Shellie 2006; Shellie and King 2013).
Medrano et al. (2003) observed an increase in berry
phenolics in Manto Negro (near-isohydric) but not in
Tempranillo (near-anisohydric) and concluded that
drought-induced changes in grape quality were cultivar-
dependent. We also observed cultivar differences in berry
composition in response to mdWL (Fig. 4). Water potential
has recently been used to define cultivar-specific thresholds
for irrigation management (Romero et al. 2010; Romero
et al. 2013). The strong association between water potential
and irrigation amount and the differences in leaf water
potential between cultivars observed in this study lend
support for the usefulness of leaf water potential to define
cultivar-specific stress thresholds.
Conclusions
An important finding of this study was that both cultivars
maintained a similar DWplant under widely varying, water
stress-induced changes in pdWL and gs. The implication of
this finding is that both cultivars were able to maintain an
isohydric soil-to-leaf water potential gradient through
dynamic changes in hydraulic conductivity. The lower leaf
water potential of Cabernet Sauvignon relative to Malbec
under identical amounts of irrigation suggests that the
mechanisms used to achieve this isohydrodynamic state
may differ among cultivars. Understanding how wide-
spread isohydrodynamic behavior is among cultivars and
the diversity of mechanisms involved could lead to future
advances in water conservation and irrigation management.
In this study, we used mdWL as an indicator to characterize
stress-induced changes in yield components and berry
composition and showed that their regression equations can
be used to predict response. The cultivar-specific
relationships presented in this study provide useful infor-
mation for customizing irrigation amounts to induce levels
of water deficit that meet production goals.
Acknowledgments This work was conducted under ARS Project
No. 5358-21000-034-00D entitled ‘‘Production Systems to Promote
Yield and Quality of Grapes in the Pacific Northwest.’’ The authors
thank Alan Muir, Monte Shields, and Cheryl Franklin-Miller for
technical assistance and the University of Idaho Parma Research and
Extension Center for the use of their field resources and materials.
References
Allen RG, Pereira LS, Raes D, Smith M (1998) Crop evapotranspira-
tion-guidelines for computing crop water requirements-FAO
irrigation and drainage paper 56. Food Agricultural Organization
(FAO), Rome
Alsina MM, de Herralde F, Aranda X, Save R, Biel C (2007) Water
relations and vulnerability to embolism are not related: exper-
iments with eight grapevine cultivars. VITIS 46:1–6
Basile B, Marsal J, Mata M, Vallverdu’ X, Bellvert J, Girona J (2011)
Phenological sensitivity of Cabernet Sauvignon to water stress:
vine physiology and berry composition. Am J Enol Vitic
62:452–461
Bota J, Flexas J, Medrano H (2001) Genetic variability of photosyn-
thesis and water use in Balearic grapevine cultivars. Ann Appl
Biol 138:353–361
Bowen P, Bogdanoff C, Usher K, Estergaard B, Watson M (2011)
Effects of irrigation and crop load on leaf gas exchange and fruit
composition in red winegrapes grown on a loamy sand. Am J
Enol Vitic 62:9–22
Chaves MM, Zarrouk O, Francisco R, Costa JM, Santos T, Regalado
AP, Rodrigues ML, Lopes CM (2010) Grapevine under deficit
irrigation: hints from physiological and molecular data. Ann Bot
105:661–676
Cifre J, Bota J, Escalona JM, Medrano H, Flexas J (2005)
Physiological tools for irrigation scheduling in grapevine (Vitis
vinifera L.): an open gate to improve water-use efficiency? Agr
Ecosyst Environ 106:159–170
Correia MJ, Pereira JS, Chaves MM, Rodrigues ML, Pacheco CA
(1995) ABA xylem concentrations determine maximum daily
leaf conductance of field-grown Vitis vinifera L. plants. Plant,
Cell Environ 18:511–521
Cramer G, Ergu A et al (2007) Water and salinity stress in grapevines:
early and late changes in transcript and metabolite profiles. Funct
Integr Genomics 7:111–134
De Souza CR, Maroco JP, dos Santos TP, Rodrigues ML, Lopes CM,
Pereira JS, Chaves MM (2005) Impact of deficit irrigation on
water use efficiency and carbon isotope composition (d13C) of
field-grown grapevines under Mediterranean climate. J Exp Bot
56:2163–2172
Domec J, Johnson DM (2013) Does homeostasis or disturbance of
homeostasis in minimum leaf water potential explain the
isohydric versus anisohydric behavior of Vitis vinifera L.
cultivars? Tree Physiol 32:245–248
Franks PJ, Drake PL, Froend RH (2007) Anisohydric but isohydro-
dynamic: seasonally constant plant water potential gradient
explained by a stomatal control mechanism incorporating
variable plant hydraulic conductance. Plant, Cell Environ
30:19–30
Hochberg U, Degu A, Fait A, Rachmilevitch S (2013) Near isohydric
grapevine cultivar displays higher photosynthetic efficiency and
photorespiration rates under drought stress as compared with
96 Irrig Sci (2014) 32:87–97
123
near anisohydric grapevine cultivar. Physiol Plantarum
147:443–452
Iland P, Bruer N, Edwards G, Weeks S, Wilkes S (2004) Chemical
analysis of grapes and wine: techniques and concepts. Patrick
Iland Wine Promotions PTY LTD, Campbelltown, pp 44–48
Intrigliolo DS, Castel JR (2006) Vine and soil-based measures of
water status in a Tempranillo vineyard. VITIS 45:157–163
Keller M, Smithyman RP, Mills LJ (2008) Interactive effects of
deficit irrigation and crop load on Cabernet Sauvignon in an arid
climate. Am J Enol Vitic 59:221–234
Lawlor DW, Cornic G (2002) Photosynthetic carbon assimilation and
associated metabolism in relation to water deficits in higher
plants. Plant, Cell Environ 25:275–294
Lovisolo C, Perrone I, Carra A, Ferrandino A, Flexas J, Medrano H,
Schubert A (2010) Drought-induced changes in development and
function of grapevine (Vitis spp.) organs and in their hydraulic
and non-hydraulic interactions at the whole-plant level: a
physiological and molecular update. Funct Plant Biol
105:661–676
Medrano H, Escalona JM, Cifre J, Bota J, Flexas J (2003) A ten-year
study on the physiology of two Spanish grapevine cultivars
under field conditions: effects of water availability from leaf
photosynthesis to grape yield and quality. Func Plant Bio
30:607–619
Roby G, Matthews MA (2004) Relative proportions of seed, skin and
flesh, in ripe berries from Cabernet Sauvignon grapevines grown
in a vineyard either well irrigated or under water deficit. Aust J
Grape Wine R 10:74–82
Roby G, Harbertson JF, Adams DA, Matthews MA (2004) Berry size
and vine water deficits as factors in winegrape composition:
anthocyanins and tannins. Aust J Grape Wine R 10:100–107
Romero P, Fernandez-Fernandez JI, Martinez-Cutillas A (2010)
Physiological thresholds for efficient regulated deficit-irrigation
management in winegrapes grown under semiarid conditions.
Am J Enol Vitic 61:300–312
Romero P, Gil-Munoz R, del Amor FM, Valdes E, Fernandez JI,
Martinez-Cutillas A (2013) Regulated deficit irrigation based
upon optimum water status improves phenolic composition in
Monastrell grapes and wines. Agric Water Manag 121:85–101
Sadras VO, Stevens RM, Pech JM, Taylor EJ, Nicholas PR, McCarthy
MG (2007) Quantifying phenotypic plasticity of berry traits
using an allometric-type approach: a case study on anthocyanins
and sugars in berries of Cabernet Sauvignon. Aust J Grape Wine
R 13:72–80
Schultz HR (2003) Differences in hydraulic architecture account for
near-isohydric and anisohydric behavior of two field-grown Vitis
vinifera L. cultivars during drought. Plant Cell Environ
26:1393–1405
Shellie K (2006) Vine and berry response of Merlot (Vitis vinifera L.)
to differential water stress. Am J Enol Vitic 57:514–518
Shellie KC, King B (2013) Kaolin-based foliar reflectant and water
deficit influence Malbec leaf and berry temperature, pigments,
and photosynthesis. Am J Enol Vitic. doi:10.5344/ajev.2012.
12115
Sivilotti P, Bonetto C, Paladin M, Peterlunger E (2005) Effect of soil
moisture availability on Merlot: from leaf water potential to
grape composition. Am J Enol Vitic 56:9–18
Turner NC (1988) Measurement of plant water status by the pressure
chamber technique. Irrigation Sci 9:289–308
US Department of Agriculture (1972) Soil survey of Canyon Area,
Idaho. US Department of Agriculture, Soil Conservation Service.
Superintendent of Documents, US Government Printing Office,
Washington, DC
Vandeleur RK, Mayo G, Shelden MC, Gilliham M, Kaiser BN,
Tyerman SS (2009) The role of plasma membrane intrinsic
protein aquaporins in water transport through roots: diurnal and
drought stress responses reveal different strategies between
isohydric and anisohydric cultivars of grapevine. Plant Physiol
149:445–460
Williams LE (2012) Effects of applied water amounts at various
fractions of evapotranspiration (ETc) on leaf gas exchange of
Thompson Seedless grapevines. Aust J Grape Wine R 18:100–108
Williams LE, Araujo FU (2002) Correlations among predawn leaf,
midday leaf, and midday stem water potential and their
correlations with other measures of soil and plant water status
in Vitis vinifera. J Am Soc Hortic Sci 127:448–454
Williams LE, Baeza P (2007) Relationships among ambient temper-
ature and vapor pressure deficit and leaf and stem water
potentials of fully irrigated, field-grown grapevines. Am J Enol
Vitic 58:173–181
Williams LE, Trout TJ (2005) Relationships among vine-and soil-
based measures of water status in a Thompson seedless vineyard
in response to high-frequency drip irrigation. Am J Enol Vitic
56:357–366
Williams LE, Grimes DW, Phene CJ (2010) The effects of applied
water at various fractions of measured evapotranspiration on
water relations and vegetative growth of Thompson Seedless
grapevines. Irrig Sci 28:221–232
Williams LE, Baeza P, Vaughn P (2012) Midday measurements of
leaf water potential and stomatal conductance are highly
correlated with daily water use of Thompson Seedless grape-
vines. Irrig Sci 30:201–212
Zhang Y, Oren R, Kang S (2013) Spatiotemporal variation of crown-
scale stomatal conductance in an arid Vitis vinifera L. Merlot
vineyard: direct effects of hydraulic properties and indirect
effects of canopy leaf area. Tree Physiol 32:262–279
Irrig Sci (2014) 32:87–97 97
123
