AJEV American Journal of

Enology and Viticulture

Research Report
DOI: 10.5344/ajev.2025.25009

Alleviating Water Stress at Veraison May Trigger

Berry Shrivel Disorder in Susceptible Grapevines
Andreas Wenter ,1,3 Carlo Andreotti ,1 Damiano Zanotelli ,1 and Markus Keller 2*

Abstract Introduction
Background and goals Following seed and fruit set, grape (Vitis sp.) berries grow in a double-
Berry shrivel (BS), a grape ripening disorder sigmoid fashion, with two growth phases separated by a short phase of
which is linked to arrested sugar accumulation little growth, called the lag phase (Keller 2020). Fruit ripening starts at the
and metabolic alterations, is thought to be end of the lag phase with berry softening, followed by sugar accumulation,
of physiological origin. Its underlying causes malate depletion, and renewed berry growth (Shahood et al. 2020, Hernán-
remain unknown, but abrupt environmental
dez-Montes et al. 2021). Additionally, anthocyanins begin to accumulate in
changes might be involved. We tested whether
cultivars with dark-skinned berries once the berry sugar concentration,
BS may be triggered by plant water stress lead-
estimated as total soluble solids (TSS), exceeds 9 to 10 Brix (Keller 2020).
ing up to veraison followed by irrigation with
water at different temperatures. Thus, grape berries grow and ripen at the same time. Each berry may ini-
tiate its own ripening program independently of the other berries on the
Methods and key findings same cluster or vine (Keller et al. 2015b, Zhang and Keller 2017, Shahood
In a two-year field trial with deficit-irrigated et al. 2020, Hernández-Montes et al. 2021). In doing so, it pulls phloem sap
Cabernet Sauvignon winegrapes in arid south- into the berry and unloads the dissolved sugar for storage. Under favorable
eastern Washington, irrigation was suspended
conditions (i.e., adequate light, temperature, water, nutrients, and plant
during the lag phase of berry development,
health), the photosynthesizing leaves also push the phloem sap toward all
then resumed at veraison, using either water
berries (Keller et al. 2015b). Thus, phloem flow follows a pressure gradient
at ambient temperature (20 to 24°C) or ice-
cooled water (0°C). Measurements of soil mois-
that is established within the phloem’s sieve tubes by the production and
ture, leaf water potential, and leaf gas exchange loading of sugar in sources and its unloading and use or storage in sinks
showed that interrupting irrigation induced (Knoblauch et al. 2016).
moderate-to-severe plant water stress before Despite the asynchronous ripening onset and progression, grape clus-
rewatering. Compared with ambient-temper- ters can suffer from ripening disorders that are seemingly initiated in-
ature water, cooling the irrigation water used dependent of the maturity status of the individual berries (Griesser et al.
for rewatering reduced the soil temperature by 2024). Among different disorders, berry shrivel (BS; also referred to as sour
4 to 8°C for 1 to 3 days, but had a minor effect shrivel or sugar accumulation disorder) and bunch-stem necrosis (BSN)
on the vines’ response to water stress. Rewa- are especially intractable. Both are thought to be of physiological origin,
tering at 50% veraison, but not at 5% veraison, but their internal or external triggers remain unknown (Griesser et al.
significantly increased BS incidence. Cooling
2024). While the rachis of clusters affected by BSN develops necrotic le-
the irrigation water also tended to increase BS
sions that interrupt phloem flow, it typically remains green in BS-affected
incidence, but not consistently so.
clusters, although cell death has sometimes been observed in their ra-
Conclusions and significance chis phloem (Hall et al. 2011, Bondada and Keller 2012, Zufferey et al. 2015,
This study showed that sudden alterations in Griesser et al. 2024). However, premature loss of mesocarp cell membrane
plant water availability and water temperature integrity during early ripening is a prominent internal symptom of berries
may be involved in triggering BS. Though the
cause-effect relationship remains unclear, this
knowledge may be useful in devising vineyard 1
Faculty of Agricultural, Environmental and Food Sciences, Free University of Bolzano-Bozen,
irrigation or soil management strategies that 39100 Bolzano-Bozen, Italy; 2Department of Viticulture and Enology, Washington State Uni-
versity, Irrigated Agriculture Research and Extension Center, Prosser, WA 99350; and 3present
avoid abrupt changes in soil moisture and soil address, Laimburg Research Centre, 39040 Auer-Ora, Italy.
temperature around veraison. *Corresponding author ([email protected])
Manuscript submitted and accepted Feb 2025, published April 2025
This is an open access article distributed under the CC BY 4.0 license.
Key words: grape berry, ripening disorder, By downloading and/or receiving this article, you agree to the Disclaimer of Warranties and
soil moisture, soil temperature, Vitis Liability. If you do not agree to the Disclaimers, do not download and/or accept this article.

American Journal of Enology and Viticulture — ajevonline.org 1 of 10 2025 Volume 76 Article 0760010
Wenter et al. Berry Shrivel After Water Stress Alleviation

from BS clusters (Krasnow et al. 2008, 2009, Hoff et al. 2021). 2015). Though neither the water temperature nor the soil
Membrane failure, more specifically rupture of the vacuolar temperature were measured in those studies, one might as-
membrane, leads to cell death and might be associated with sume both rainwater and irrigation water in those Alpine re-
oxidative stress as a result of hypoxia in the mesocarp cells gions to be rather cold. Applying cold water to dry soil can
surrounding the seeds (van Doorn and Woltering 2004, Xiao lead to a rapid, transient decline in soil temperature, which
et al. 2018). Affected berries lose the ability to accumulate has been found to inhibit growth of some plant species
sugar, though they still metabolize malate (Keller et al. 2016, (Brockwell and Gault 1976). By decreasing the roots’ hydrau-
Griesser et al. 2024). As phloem inflow stops while evapora- lic conductance, low soil temperature may exacerbate water
tive water loss continues, the berries begin to shrink, and stress responses such as stomatal closure and reduced pho-
their low sugar-to-acid ratio and low potassium content re- tosynthesis (Ameglio et al. 1990, Rogiers and Clarke 2013).
sults in their sour taste (Keller et al. 2016). In contrast, the When severe enough, the ensuing imbalance between root
cellular membranes of healthy berries remain intact until water uptake and leaf water demand may trigger xylem cavi-
late in ripening (Krasnow et al. 2008, Fuentes et al. 2010, tation and leaf wilting (Scheenen et al. 2007).
Keller and Shrestha 2014). To our knowledge, the effect of fluctuations in rootzone
While BSN will affect all berries downstream of necrotic temperature on fruit ripening has not been investigated. In
lesions on the rachis regardless of the ripening status of each addition to the effects of water stress (and its alleviation)
berry, membrane failure leading to BS starts in individual and root chilling on plant responses at the leaf and cano-
berries after their seeds are mature and sugar accumulation py level, it is possible that responses also occur at the fruit
has started (Hall et al. 2011, Keller et al. 2016, Hoff et al. 2021). level, especially during a potentially vulnerable phenologi-
Therefore, rather than appearing simultaneously across en- cal stage such as veraison. Consequently, we speculated that
tire clusters, BS may develop in individual berries but spread alleviating water stress at the beginning of grape ripening
quickly over the cluster as each berry initiates its ripening might trigger processes that culminate in the BS syndrome
process. Compared with normal ripening whose onset can in susceptible cultivars. We further hypothesized that apply-
span 20 to 30 days across the berries of a cluster (Hernán- ing cold irrigation water during periods of high evaporative
dez-Montes et al. 2021), this change might be accelerated in demand might temporarily exacerbate the stress perceived
BS clusters because sugar accumulation stops as each berry by the vines. Taking advantage of eastern Washington's arid
in turn fails to properly execute its ripening program. Never- climate and the relatively frequent appearance of BS in this
theless, it remains unknown why entire clusters rather than region (Keller et al. 2016, Hoff et al. 2021), we tested the in-
individual berries or groups of berries typically develop BS fluence of a period of soil drydown during the lag phase of
symptoms (Griesser et al. 2024). berry development followed by rewatering at veraison with
As grape berries begin to ripen, they undergo a change irrigation water that was either left at ambient temperature
in vascular flow to and from the berries. Green hard ber- or cooled with ice on physiological responses and BS and
ries receive most of their water via the xylem, but as they BSN incidence in Cabernet Sauvignon. The field trial was re-
enter the ripening period, the berries’ rapid rise in sugar de- peated in two growing seasons to test whether any potential
mand leads them to rely on phloem-derived water (Keller et effects were consistent across years.
al. 2015b, Zhang and Keller 2017). While the berries retain a
small portion of that water for renewed growth, the surplus
is discharged via berry transpiration (especially under con- Materials and Methods
ditions of high evaporative demand) and xylem backflow to Site and plant material
the leaves (Zhang and Keller 2015, 2017). At the same time, the A field experiment was conducted in 2017 and 2018 in a
berries’ phloem-associated cells close their plasmodesmata vineyard at Washington State University’s Irrigated Agri-
to switch from symplastic to apoplastic phloem unloading culture Research and Extension Center near Prosser, WA
and facilitate sugar import and storage (Zhang et al. 2006). It (46°17’N; 119°44’W; 365 m asl). The region receives less than
is possible that the direction and magnitude of these chang- 200 mm annual precipitation and has warm and very dry
es, coupled with the shifts in gene expression required for summers and moderately cold winters. The vineyard soil is
ripening initiation, make the berries vulnerable to abrupt a Warden silt loam with a pH ~8 and organic matter content
changes in their environment (Zufferey et al. 2015, Fasoli et of 0.25% (https://websoilsurvey.sc.egov.usda.gov). A cali-
al. 2018, Hewitt et al. 2023, Griesser et al. 2024). Such chang- che layer at 50 to 100 cm depth limits root growth. The soil
es may include temperature fluctuations, as well as fluctua- volumetric water content (θv) is 26% (v/v) at field capacity
tions in water supply. (FC) and 8% at permanent wilting point (PWP) (Groenveld et
Observations in Italy’s South Tyrol region suggested BS al. 2023). Own-rooted Vitis vinifera L. cv. Cabernet Sauvi-
may have been especially prominent in vineyards that ex- gnon (clone FPS 07) vines propagated from certified material
perienced soil water deficit leading up to veraison followed were planted in 2010 at 2.7 m between rows and 1.8 m within
by rainfall at veraison (Raifer et al. 2023). In addition, BS was rows. Rows were oriented north-south on a 2% slope with a
prevalent in a mountain valley in southwestern Switzerland southwestern aspect. Vines were trained to two trunks with
in years with fluctuating temperatures and rainfall around bilateral cordons 95 cm above the ground, spur-pruned in
veraison, and was exacerbated by irrigation (Zufferey et al. winter to 24 buds per vine, and vertically shoot-positioned

American Journal of Enology and Viticulture — ajevonline.org 2 of 10 2025 Volume 76 Article 0760010
Wenter et al. Berry Shrivel After Water Stress Alleviation

between two pairs of foliage wires 30 cm apart and 30 and 75 International) in four access tubes installed equidistant be-
cm above the cordon. A 1.2-m herbicide strip was maintained tween pairs of drip emitters. To permit comparisons across
in the rows, and permanent but summer-dormant volunteer different soil types, soil moisture was normalized relative
vegetation grew between rows. The vineyard was drip-irri- to that at FC and PWP by converting θv to extractable soil
gated using two 2-L/hr emitters per vine. Fertilizer and pest water: ESW = [θv ‒ θPWP]/[θFC ‒ θPWP] (Groenveld et al. 2023).
management practices were applied uniformly across the Because the irrigation water temperature changes gradu-
vineyard according to regional standards. Annual disease ally as water runs through drip tubes and percolates into
testing and rogueing (if necessary) kept the vineyard as free the soil, we measured the resulting soil temperature (Tsoil)
from viral and bacterial diseases as possible. Budbreak oc- at 30-min intervals during the experiment, using iButton
curred on day of year (DOY) 122 and 123, bloom (50% capfall) temperature loggers buried at a depth of 25 cm beneath five
on DOY 163 and 156, fruit set on DOY 179 and 165, and verai- (2017) or three (2018) drip emitters per treatment replicate.
son (50% berry color change) on DOY 230 and 225 in 2017 and
2018, respectively. Physiological measurements
Physiological measurements were taken on three leaves,
Experimental design each on a different vine, per treatment replicate (i.e., nine
The 2017 experiment tested vines that experienced a tem- leaves per treatment) at maximum water stress (day before
porary suspension of water supply during the lag phase of irrigation) and the day after rewatering. A Scholander pres-
berry growth, followed by rewatering at veraison with either sure chamber (model 600, PMS Instrument Company) was
ambient irrigation water or ice-cooled water against deficit- used to measure leaf water potential at predawn (Ψpd) and
irrigated control vines. The 2018 experiments included the midday near solar noon (Ψmd), as described previously (Keller
same treatments but additionally tested the supply of ice- et al. 2015a). Leaf net assimilation rate (A), stomatal conduc-
cooled water to control vines. Treatments (three in 2017 and tance (gs), transpiration rate (E), and leaf temperature (Tleaf )
four in 2018) were applied in a randomized complete block were measured at midmorning (0900 to 1000 hr) on recently
design with three replicates and 25 consecutive vines per mature sun-exposed leaves, using a portable infrared gas an-
replicate. Following an initial drydown period after fruit set alyzer (LCi, ADC BioScientific). Earlier diurnal measurements
to control shoot growth, control vines were irrigated weekly in Merlot and Syrah winegrapes under similar environmental
at 70% of crop evapotranspiration (ETc) calculated from ref- and viticultural conditions as the vines in the present study
erence evapotranspiration (ETo) and a crop coefficient that had indicated that gs reached its daily minimum before 0900
increased from zero at budbreak to 0.8 at veraison. To gener- hr and did not begin to recover until late afternoon at this
ate transient vine water stress during the lag phase, irriga- time of year (Keller et al. 2015a, Romero et al. 2017). Gas ex-
tion was interrupted on DOY 213 (2017) or DOY 205 (2018) change measurements were taken under ambient light (pho-
and resumed at 50% veraison on DOY 230 (2017) or DOY tosynthetic photon flux >1000 µmol/m2sec) and CO2 (~400
225 (2018). In 2018, we additionally tested rewatering at 5% µmol/mol) conditions using a broad-leaf chamber (6.25 cm2)
veraison (DOY 213) in an adjacent block with an otherwise and flow rate of 200 mL/min. The evaporative flux method
identical experimental design. In 2017, the temperature of was used to estimate the leaf area-specific whole-plant hy-
the incoming ambient irrigation water, measured with iBut- draulic conductance (Kl = E/[Ψsoil – Ψmd]), assuming steady-
ton temperature loggers (DS1922L, Maxim Integrated Prod- state conditions (Keller et al. 2015a). We further assumed the
ucts), varied diurnally from 20.1 to 23.6°C, and cooled water soil water potential (Ψsoil) to be equal to Ψpd at ESW ≤ 0.4, but
was supplied to the drip tubes from a 1000-L container with 0.11 MPa higher than Ψpd at ESW > 0.4, based on data present-
an ice-water mixture (i.e., water at 0°C), using a diaphragm ed by Groenveld et al. (2023) for this vineyard site.
pump. In addition, a preliminary experiment conducted with
five vines in one row of the water stress treatment tested Ripening disorder assessment
water cooling by passing it through perforated plastic boxes At harvest time, the incidence of BS and BSN was quanti-
containing 14 kg of ice placed under the drip emitters. This fied by visually assessing and counting all clusters on five
latter approach was used for water cooling in 2018, when the vines per treatment replicate (i.e., 15 vines per treatment).
ambient water temperature ranged from 21.6 to 24.4°C. Following the definition by Seem (1984), incidence was ex-
pressed as the percentage of symptomatic clusters per vine.
Weather and soil data collection Clusters were evaluated on DOY 299 in 2017, and DOY 299
Daily weather data were obtained from the Roza AgWeath- and 303 in 2018. Symptomatic clusters were distinguished
erNet station located ~400 m from the vineyard (https:// from healthy clusters by the typical symptoms of flac-
weather.wsu.edu). Growing degree days (GDD) for the April- cid, shriveled berries for BS, and necrotic rachis portions
October growing season were estimated from daily mini- for BSN (Bondada and Keller 2012, Griesser et al. 2024). To
mum (Tmin) and maximum (Tmax) temperatures, applying a further confirm the presence of these ripening disorders,
base temperature of 10°C. Soil moisture (θv) was measured the TSS concentration of symptomatic and healthy clusters
weekly starting at budbreak to support irrigation manage- from each vine was measured with a handheld refractometer
ment. Measurements were taken at three depths (30, 60, (PAL-1, Atago). Individual clusters were manually crushed in
and 90 cm) using a neutron probe (503 DR Hydroprobe, CPN plastic bags, and the juice was analyzed immediately.

American Journal of Enology and Viticulture — ajevonline.org 3 of 10 2025 Volume 76 Article 0760010
Wenter et al. Berry Shrivel After Water Stress Alleviation

Data analysis irrigation water during the lag phase of berry growth led
Data were analyzed in R (Rx64 3.5.0 Core Team) and Sta- to rapid soil drying. The average ESW in the top 60 cm of
tistica 14.2 (Cloud Software Group). We used one-way anal- the drying soil declined below 0.15, while in the control it
ysis of variance (ANOVA) to test treatment effects in 2017 remained above 0.25 in both 2017 (Table 1) and 2018 (Tables 2
and two-way ANOVA to test main and interactive effects of and 3). Rewatering at veraison rapidly increased ESW but did
water stress and water temperature in 2018. Means were not fully restore soil moisture as it was not possible to apply
separated by F-test at 5% significance. Data are presented differential amounts of water to the different treatments.
as means ± standard error (SE) where appropriate. Pearson The soil temperature at 25-cm depth was primarily driven
correlation analysis was used to test associations between by, and lagged, diurnal changes in air temperature, but was
response variables. modulated by irrigation water cooling. The Tsoil peaked be-
tween dusk and midnight, and the daily minimum occurred
between sunrise and midday (Figure 3). Compared with ir-
Results rigation water at ambient temperature, irrigating with ice-
Weather cooled water in 2017 reduced Tsoil by a maximum of 3.8°C
The two growing seasons were slightly warmer than the over the course of the subsequent day, after which the tem-
long-term (24 yrs) average. The cumulative seasonal GDD was perature recovered to the level of the soil irrigated with
1485°C in 2017 and 1471°C in 2018 (long-term average 1442°C). ambient-temperature water (Figure 3). The Tsoil in the ice-
There were 7 days with Tmax > 35°C (peak 36.6°C) during the box test dropped by 7.8°C during the afternoon (from 25.4 to
17-day preveraison water stress period in early to mid-August 17.6°C within 7 hr) and recovered after sunrise the following
of 2017, and 5 days with Tmax > 35°C (peak 37.3°C) during the morning. In 2018, interrupting the irrigation water supply
20-day stress period in 2018 (Figure 1). At the same time, raised Tsoil by a maximum of 2.9°C, and an average of 1.9°C
there were 5 days with Tmin < 15°C (minimum 9.1°C) in 2017, during the week before rewatering, compared with the con-
and 8 days with Tmin < 15°C (minimum 10.0°C) in 2018. Total trol. Applying ice-cooled water to the previously unirrigat-
annual precipitation was 240 mm in 2017 and 158 mm in 2018 ed, and hence warmer, soil decreased Tsoil by 8.2°C, whereas
(long-term average 175 mm). Only 3.1 mm rain fell during the in the irrigated, and hence cooler, soil the decrease was only
2017 water stress period, and 9.9 mm fell early during the 2018 4.3°C (Figure 3). The cooling effect gradually lessened until
stress period. The daily ETo during the water withholding pe- it disappeared after 3 days.
riod varied from 4.1 to 6.6 mm (cumulative 93 mm) in 2017, and
from 4.1 to 7.4 mm (cumulative 125 mm) in 2018. Physiological traits
Withholding irrigation decreased Ψpd and Ψmd and led to
Soil moisture and soil temperature almost complete stomatal closure (gs < 0.05 mol/m2sec),
Following budbreak, irrigation maintained the ESW across along with large reductions in A and E and an increase in
the soil profile above 0.3 through fruit set, after which it de- Tleaf, in both 2017 (Table 1) and 2018 (Tables 2 and 3). The vines
clined through mid-July to ~0.2 in the top 30 cm and to 0.1 at did not fully recover a day after rewatering at veraison. Wa-
lower depths as a result of deficit irrigation (Figure 2). A fur- ter withholding also strongly decreased K l, but the day after
ther decrease occurred at the 90-cm depth; from veraison water was resupplied, K l recovered to the level of control
onwards, θv at 90-cm depth remained near the PWP through vines. While ice-cooling the irrigation water consistently
harvest in both 2017 and 2018, while irrigation increased θv
at the 30- and 60-cm depths. In both years, withholding
1.0
2017 Soil depth 2018
0.8 30 cm
40 60 cm
2017 2018 0.6 90 cm
ESW
Temperature (°C)

30 0.4

0.2
20
0.0

10 162 184 205 226 253 276 162 184 205 226 253 276
176 197 217 240 261 176 197 217 240 261

0 Day of year Day of year

200 210 220 230 240 200 210 220 230 240

Day of year Day of year Figure 2 Changes in extractable soil water (ESW; varying from 0 at
permanent wilting point to 1 at field capacity) at three depths, from fruit
Figure 1 Daily changes in maximum (red line) and minimum (blue set through harvest, in a Cabernet Sauvignon vineyard in southeastern
line) temperatures during the lag phase and early ripening period in Washington in 2017 and 2018. Days of year were 163 and 156 for bloom,
a Cabernet Sauvignon vineyard in southeastern Washington in 2017 179 and 165 for fruit set, and 230 and 225 for veraison in 2017 and 2018,
and 2018. Blue bars indicate water withholding periods lasting 17 and respectively. Blue bars indicate water withholding periods lasting 17 and
20 days in 2017 and 2018, respectively, and ending at 50% veraison. 20 days in 2017 and 2018, respectively, and ending at 50% veraison.

American Journal of Enology and Viticulture — ajevonline.org 4 of 10 2025 Volume 76 Article 0760010
Wenter et al. Berry Shrivel After Water Stress Alleviation

reduced Ψpd the next morning, the effect on Ψmd, K l, and leaf 28
gas exchange was minor and mostly nonsignificant (Tables 2017 2018
26
1 to 3). Across measurements before and after each irriga-

Temperature (°C)
tion event over the two years, gs correlated with Ψpd (Figure 24

4), Ψmd (r > 0.53), E (r > 0.90), and A (r > 0.95) (all p < 0.001). 22
Stomatal conductance varied widely at high Ψpd as a result
of variable vapor pressure deficit, but gs declined at Ψpd < 20

-0.3 MPa and the stomata were essentially closed (gs < 0.05 18 Stress amb Control amb
mol/m2sec) at Ψpd < -0.8 MPa (Figure 4). Consequently, both Stress cold Control cold
16
E (r > 0.76) and A (r > 0.68) also decreased with decreasing 0 24 48 72 0 24 48 72
Ψpd (p < 0.001). However, unlike gs, K l was driven much more
Time (hr) Time (hr)
strongly by Ψmd (Figure 5) than by Ψpd (r > 0.29, p < 0.025).
Manipulating the irrigation water temperature did not alter Figure 3 Diurnal changes in soil temperature (25-cm depth) during
any of these relationships between physiological variables. and after irrigation following water withholding in a Cabernet Sauvignon
vineyard in southeastern Washington. Water-stressed and control vines
Ripening disorders and TSS were irrigated at 50% veraison on 18 Aug 2017 (n = 5) or 13 Aug 2018
(n = 3) with ambient (amb) or ice-cooled (cold) water. Temperature
Across the two years, both transient preveraison wa- was not monitored in control vines in 2017. Blue bars indicate irrigation
ter stress (p = 0.001) and cold irrigation water (p = 0.021) periods (i.e., rewatering events).

Table 1 Effect of transient water stress (Stress) during the lag phase of berry growth followed by irrigation (Recovery) with ambient or
ice-cooled (Cold) water at 50% veraison on soil moisture and physiological traits, measured the day before and the day after recovery
irrigation in Cabernet Sauvignon vines in a vineyard in southeastern Washington in 2017.
Kl
ESWa Ψpd Ψmd Tleaf A E gs
(mmol/m2sec
(ratio) (MPa) (MPa) (°C) (µmol/m2sec) (mmol/m2sec) (mol/m2sec)
MPa)
Stress
Control 0.32 ndb nd 40.1 16.45 7.12 0.16 nd
Stressed 0.17 -0.47 -1.05 41.1 6.95 2.49 0.04 4.51
p 0.012 nd nd <0.001 <0.001 <0.001 <0.001 nd

Recovery
Ambient 0.21 -0.16 -0.98 39.6 14.18 5.88 0.14 8.77
Cold 0.26 -0.19 -0.87 39.7 12.96 5.41 0.12 6.84
p 0.46 <0.001 0.021 0.33 0.064 0.031 0.016 0.015
a
ESW, relative extractable soil water in top 60 cm; Ψpd, predawn leaf water potential; Ψmd, midday leaf water potential; Tleaf, leaf temperature;
A, net assimilation rate; E, transpiration rate; gs, stomatal conductance; Kl, whole-vine hydraulic conductance.
b
nd, not determined.

Table 2 Effect of transient water stress (Stress) during the lag phase of berry growth followed by irrigation (Recovery) with ambient or
ice-cooled (Cold) water at 5% veraison on soil moisture and physiological traits, measured the day before and the day after recovery
irrigation in Cabernet Sauvignon vines in a vineyard in southeastern Washington in 2018.
Kl
ESWa Ψpd Ψmd Tleaf A E gs
(mmol/m2sec
(ratio) (MPa) (MPa) (°C) (µmol/m2sec) (mmol/m2sec) (mol/m2sec)
MPa)
Stress
Control 0.26 -0.22 -1.04 33.2 18.78 6.08 0.37 7.25
Stressed 0.12 -1.04 -1.63 35.2 5.18 1.44 0.04 2.38
p <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001 <0.001

Recovery
Control 0.42 -0.17 -0.82 34.6 16.04 6.58 0.36 13.14
Stress 0.23 -0.67 -0.92 36.8 6.98 3.06 0.09 10.09
p (water) 0.016 <0.001 0.006 0.95 <0.001 <0.001 <0.001 0.057
Ambient 0.34 -0.39 -0.87 35.7 12.11 5.07 0.23 11.64
Cold 0.31 -0.45 -0.87 35.7 10.91 4.57 0.21 11.59
p (temperature) 0.73 0.002 0.95 0.95 0.051 0.36 0.095 0.97
p (interaction) 0.09 0.031 0.89 0.89 0.061 0.23 0.53 0.71
a
ESW, relative extractable soil water in top 60 cm; Ψpd, predawn leaf water potential; Ψmd, midday leaf water potential; Tleaf, leaf temperature;
A, net assimilation rate; E, transpiration rate; gs, stomatal conductance; Kl, whole-vine hydraulic conductance.

American Journal of Enology and Viticulture — ajevonline.org 5 of 10 2025 Volume 76 Article 0760010
Wenter et al. Berry Shrivel After Water Stress Alleviation

increased BS incidence, with no significant interaction across the water stress experiment varied from 34 to 91 per
(p = 0.41). However, the variation due to water stress was vine and correlated positively with BS incidence (Figure 7).
more than twice that due to water temperature. In 2017, At harvest time, the berries from healthy clusters had 24.8
interrupting irrigation during the lag phase followed by ± 0.4 Brix, whereas those from BS clusters had only 15.0 ±
rewatering at veraison increased BS incidence 21-fold 0.7 Brix (p < 0.001). The preveraison water stress did not
relative to the control (p = 0.023). While only 0.5% of the affect TSS in either the healthy clusters (p = 0.67) or the
clusters on control vines developed BS, on previously wa- BS clusters (p = 0.55). The BSN incidence was minimal in
ter-stressed vines, 10% of the clusters developed BS (Fig- 2017; only two vines, both in the cold-water treatment, had
ure 6). The cold-water treatment was applied only to the a cluster with BSN symptoms.
water-stressed vines and had no significant effect on BS In 2018, there was very little BS (0.2%) and no BSN in the
incidence (p = 0.26). In the ice-box test, however, 50% of block where rewatering occurred at 5% veraison, and nei-
the clusters on cold-water-treated vines developed BS (p ther water stress (p = 0.60) nor water cooling (p = 0.33) in-
< 0.001), and those clusters had a TSS of 14.3 ± 0.5 Brix at creased BS incidence relative to the control. In the adjacent
harvest compared with 23.5 ± 0.5 Brix for the unaffected block that was used to test rewatering at 50% veraison (the
clusters on the same vines (p < 0.001). The cluster number same block as in 2017), BS was much more prevalent (12%).

Table 3 Effect of transient water stress (Stress) during the lag phase of berry growth followed by irrigation (Recovery) with ambient or
ice-cooled (Cold) water at 50% veraison on soil moisture and physiological traits, measured the day before and the day after recovery
irrigation in Cabernet Sauvignon vines in a vineyard in southeastern Washington in 2018.
Kl
ESWa Ψpd Ψmd Tleaf A E gs
(mmol/m2sec
(ratio) (MPa) (MPa) (°C) (µmol/m2sec) (mmol/m2sec) (mol/m2sec)
MPa)
Stress
Control 0.32 -0.42 -1.20 42.7 7.74 4.16 0.07 5.45
Stressed 0.13 -0.87 -1.35 45.5 0.20 0.08 0.01 0.14
p <0.001 <0.001 <0.001 0.014 <0.001 <0.001 <0.001 <0.001

Recovery
Control 0.37 -0.18 -0.67 41.2 14.29 9.05 0.23 20.41
Stressed 0.19 -0.28 -0.66 41.5 10.83 7.58 0.17 20.29
p (water) 0.015 <0.001 0.93 0.56 <0.001 <0.001 <0.001 0.95
Ambient 0.32 -0.22 -0.67 41.6 12.67 8.36 0.20 20.46
Cold 0.24 -0.24 -0.66 41.1 12.07 8.10 0.18 20.21
p (temperature) 0.23 0.033 0.60 0.28 0.67 0.71 0.32 0.32
p (interaction) 0.72 0.87 0.27 0.82 0.37 0.55 0.34 0.34
a
ESW, relative extractable soil water in top 60 cm; Ψpd, predawn leaf water potential; Ψmd, midday leaf water potential; Tleaf, leaf temperature;
A, net assimilation rate; E, transpiration rate; gs, stomatal conductance; Kl, whole-vine hydraulic conductance.

40
0.5
2017, 50% veraison
2017, 50% veraison 2018, 5% veraison
2018, 5% veraison 2018, 50% veraison
Kl (mmol/m2sec MPa)
0.4 30
2018, 50% veraison r = 0.78, p < 0.001
r = 0.61, p < 0.001
gs (mol/m2 sec)

0.3
20

0.2
10

0.1

0
0.0 -1.8 -1.6 -1.4 -1.2 -1.0 -0.8 -0.6 -0.4
-1.4 -1.2 -1.0 -0.8 -0.6 -0.4 -0.2 0.0
Ψmd (MPa)
Ψpd (MPa)
Figure 5 Relationship between midday leaf water potential (Ψmd) and
Figure 4 Relationship between predawn leaf water potential (Ψpd) and leaf area-specific whole-vine hydraulic conductance (Kl) at veraison
stomatal conductance (gs) at veraison in a Cabernet Sauvignon vineyard in a Cabernet Sauvignon vineyard in southeastern Washington over
in southeastern Washington over two years. two years.

American Journal of Enology and Viticulture — ajevonline.org 6 of 10 2025 Volume 76 Article 0760010
Wenter et al. Berry Shrivel After Water Stress Alleviation

There also was some BSN, but its incidence was again very Table 4 Effect of transient water stress during the lag phase
low (Table 4). Here, irrigation of previously water-stressed of berry growth followed by irrigation with ambient or ice-cooled
vines more than doubled the BS incidence (17.3%) com- (Cold) water at 50% veraison on bunch-stem necrosis (BSN)
pared to the control (6.9%; p < 0.001). The main effect of incidence and berry total soluble solids (TSS) at harvest in a
Cabernet Sauvignon vineyard in southeastern Washington in
water cooling was not significant (p = 0.25), but a Tukey 2018.
test across all treatment combinations showed that con-
TSS (Brix)
tinuously irrigating with ambient-temperature water was BSN (%)
BS BSN Healthy
once again associated with the lowest BS incidence (Fig-
ure 6). Neither water stress nor water cooling significantly Control 1.1 16.3 17.4 24.7
altered the BSN incidence, though there was a trend (p = Stress 1.3 18.4 18.3 24.5
0.07) of more BSN in the cold-water treatment (Table 4). p (water) 0.84 0.17 0.68 0.71
The incidence of both BS (Figure 7) and BSN (r = 0.34, p Ambient 0.7 17.7 16.7 24.5
= 0.008) correlated with the number of clusters per vine, Cold 1.7 16.7 18.5 24.8
which ranged from 52 to 123. Moreover, there was a weak p (temperature) 0.07 0.38 0.57 0.58
p (interaction) 0.59 0.91 0.50 0.64
correlation (r = 0.33, p = 0.01) between BS incidence and
BSN incidence. While there was again no treatment effect
on berry TSS at harvest, both BS and BSN reduced TSS to
similar degrees relative to healthy clusters (Table 4).
22
20 2017 2018 a a
Discussion 18 Ambient
a

Berry shrivel (%)

16 Cold
In this two-year field trial in arid southeastern Wash-
14
ington, suspending irrigation water supply during the lag b
12 a
phase of Cabernet Sauvignon berry development followed 10
by rewatering at 50% veraison markedly increased the in- 8
c
cidence of BS compared with vines that did not experience 6
an interruption in water supply. Rewatering at 5% verai- 4
son in one year did not affect BS incidence. Compared with 2 b
0
water at ambient temperature, using ice-cooled water to Control Water stress Control Water stress
release the water stress had a comparatively minor effect
on BS incidence, but the least BS occurred consistently on
Figure 6 Effect of transient water stress during the lag phase of berry
vines that were regularly irrigated with ambient-tempera- growth followed by irrigation with ambient or ice-cooled (Cold) water
ture water. Conversely, the incidence of BSN remained low at 50% veraison on berry shrivel incidence in a Cabernet Sauvignon
and unaltered by manipulation of irrigation water supply vineyard in southeastern Washington over two years. Bars show means
or temperature. Though there was a trend of increasing BS ± SE (n ≥ 9), and letters denote significant differences according to
Tukey’s honest significant difference test (p < 0.05); water cooling was
and BSN incidence in more heavily cropped vines, the vari- not tested in the control in 2017.
ation in cluster number explained only 18% of the variation
in BS and 12% of the variation in BSN. Nevertheless, the
(albeit weak) correlation between BS and BSN incidence,
and the similarly low TSS of BS and BSN clusters, suggests 50

the two disorders were initiated at the same time shortly 2017
r = 0.44, p = 0.002
after the onset of ripening and lends some support to the 40 2018 I
idea that they, or their causal factors, might be related r = 0.42, p < 0.001
(Hall et al. 2011).
Berry shrivel (%)

30
Our results concur with the observation that preverai-
son water stress followed by rainfall at veraison was as-
sociated with the appearance of BS (Raifer et al. 2023), 20
but they seemingly contrast with the results obtained by
Zufferey et al. (2015) in a 13-yr field trial with Humagne
10
rouge winegrapes. Those authors found that irrigation be-
fore or after veraison increased BS incidence relative to
unirrigated vines, with a steep rise in BS incidence when 0

the average Ψpd > -0.3 MPa during veraison ±2 wk. While 20 40 60 80 100 120 140
the deficit-irrigated control vines in our experiment expe- Clusters/vine
rienced mild-to-moderate water stress according to the
Figure 7 Relationship between cluster number per vine and berry
Ψpd, Ψmd, and gs thresholds defined by Rienth and Scho- shrivel incidence in a Cabernet Sauvignon vineyard in southeastern
lasch (2019), the temporary suspension of irrigation water Washington over two years.

American Journal of Enology and Viticulture — ajevonline.org 7 of 10 2025 Volume 76 Article 0760010
Wenter et al. Berry Shrivel After Water Stress Alleviation

supply led to moderate water stress in 2017 and moderate- stress during the lag phase through the “beginning of ve-
to-severe stress in 2018. Since the vineyard was deficit irri- raison” reduced anthocyanin accumulation during berry
gated, the soil was already quite dry at the beginning of the ripening but had no effect on sugar accumulation (Palai et
water withholding period. Thus, the development of vine al. 2022). However, sugar accumulation was reduced in pot-
water stress during water withholding likely resulted from ted Sangiovese and Montepulciano vines that were water-
a combination of relatively dry soil meeting high evapora- stressed from fruit set to veraison (no definition provided)
tive demand, as this period included several days with T max and not thereafter, though BS was not evaluated (Palliotti
> 35°C and had a cumulative ETo of 93 mm in 2017 and 125 et al. 2014). It is possible that cultivars differ in their re-
mm in 2018. sponse to rewatering at the beginning of ripening after a
The incomplete recovery of physiological traits the day period of water stress, but if the impact of minor shifts in
after irrigation that was observed here is common in field- the timing of environmental fluctuations, such as water
grown vines (Romero et al. 2017). Contrary to our expecta- availability or temperature, can be confirmed, this might
tion, however, the decrease in soil temperature following help to explain the seemingly erratic spatial and temporal
application of ice-cooled irrigation water to the dry (and nature of BS appearance in vineyards (cf. Keller et al. 2016).
warm) soil had only a minor influence on the vines’ recov- Oxidative stress and membrane failure in the mesocarp
ery from water stress. In both years, Ψpd was slightly lower cells have been observed in association with both early-
the day after irrigation with cold water compared with wa- ripening BS of different cultivars and late-ripening shrivel-
ter at ambient temperature. Though this effect was more ing of Syrah (Xiao et al. 2018, Griesser et al. 2024). Moreover,
pronounced in 2018 than in 2017, gs and E were affected pressurizing the root system of fully irrigated grapevines
only in 2017, and only mildly so. A study with pot-grown at veraison was found to inhibit xylem backflow and berry
Syrah winegrapes had found positive correlations between sugar accumulation (Zhang and Keller 2017). We speculate
rootzone temperature and gs or E, though the measure- that releasing moderate-to-severe water stress, leading to
ment variability was substantial (Rogiers and Clarke 2013). recovery of leaf photosynthesis and hence phloem export,
Unlike earlier work with cucumber (Scheenen et al. 2007), at a time when most berries are transitioning to ripening,
we observed no leaf wilting when the water-stressed vines or perhaps just after the berries have initiated the process-
were irrigated with cold water. es involved in the mesocarp cell wall modifications that re-
Despite the consistent appearance over two years of BS sult in berry softening, might lead to flooding of the berry
symptoms in vines that experienced transient water stress apoplast from incoming phloem water. In some cases, or in
leading up to veraison, identifying an underlying physiolog- susceptible cultivars, the resulting hypoxia in the interior
ical cause remains challenging. It seems likely that BS was mesocarp might induce oxidative stress through accumu-
triggered at or shortly after the onset of ripening, as TSS lation of reactive oxygen species, which may culminate in
values of 15 to 18 Brix are indicative of red-purple berries, loss of membrane integrity and thus trigger BS. In other
which have not completed their color change (Hernández- cases, or in other cultivars, the buildup of internal pressure
Montes et al. 2021). It is not clear, however, if BS was trig- might result in failure of the cuticle and culminate in berry
gered by the water stress going into veraison, the rewater- cracking or splitting (Keller et al. 2015b, Chang et al. 2019,
ing event at a time when the berries were initiating their Chang and Keller 2021).
ripening program, or a combination of the two. The role of
cold irrigation water in BS induction also remains uncer-
tain, especially considering that water cooling had only a Conclusion
minor influence on the vines’ recovery from water stress. The present study showed that suspending water sup-
While the present results support our first hypothesis that ply during the lag phase of berry development followed
alleviating water stress at veraison promotes BS, they pro- by rewatering at 50% veraison led to BS in field-grown,
vide no conclusive support for our second hypothesis of own-rooted Cabernet Sauvignon vines in arid southeast-
cold water exacerbating BS. Though we tested an earlier ern Washington. Ice-cooling the irrigation water led to
time of stress release only in 2018, it may be significant a comparatively minor increase in BS incidence. Though
that rewatering at 50% veraison, but not at 5% veraison, the underlying causes remain unclear, fluctuations in
seemed to trigger BS. Previously we had found that allevi- vine water status during the transition to berry ripening
ating severe water stress at ripening onset did not result in may be involved in triggering BS. This knowledge may be
BS in potted Merlot and Concord grapevines, but instead useful for developing vineyard management strategies
accelerated both renewed berry growth and sugar accu- that avoid sudden changes in soil moisture, and perhaps
mulation (Keller et al. 2015b). Withholding irrigation water soil temperature, around veraison. In irrigated vineyards,
from field-grown Cabernet Sauvignon vines during the pe- large fluctuations might be avoided by increasing the ir-
riod leading up to veraison followed by irrigation at 100% rigation frequency where possible, especially during the
of ETc in a hot Australian region reduced berry size and period leading up to veraison and through early ripening.
malate content, but neither BS nor any physiological indi- While unirrigated vineyards do not have that option avail-
cators of water stress were evaluated in that study (Cooley able, growers might be able to manage cover crops with the
et al. 2017). In pot-grown Sangiovese vines, transient water same goal in mind.

American Journal of Enology and Viticulture — ajevonline.org 8 of 10 2025 Volume 76 Article 0760010
Wenter et al. Berry Shrivel After Water Stress Alleviation

Hernández-Montes E, Zhang Y, Chang B-M, Shcherbatyuk N and

Keller M. 2021. Soft, sweet, and colorful: Stratified sampling reveals
Acknowledgments sequence of events at the onset of grape ripening. Am J Enol Vitic
This project was funded by a scholarship from Free University of Bozen- 72:137-151. DOI: 10.5344/ajev.2020.20050
Bolzano, the Italian Ministry of University and Research Foundation (TN
2043), the National Institute of Food and Agriculture of the U.S. Depart- Hewitt S, Hernández-Montes E, Dhingra A and Keller M. 2023.
ment of Agriculture (Hatch project 7003737), and the Chateau Ste. Michelle Impact of heat stress, water stress, and their combined effects
Distinguished Professorship in Viticulture. We thank Alan Kawakami and on the metabolism and transcriptome of grape berries. Sci Rep
Lynn Mills for skilled technical assistance. 13:9907. DOI: 10.1038/s41598-023-36160-x
Hoff RT, Bondada BR and Keller M. 2021. Onset and progression of
the berry shrivel ripening disorder in grapes. Aust J Grape Wine
ORCID Res 27:280-289. DOI: 10.1111/ajgw.12471
Andreas Wenter 0009-0009-5480-6396 Keller M. 2020. The Science of Grapevines. 3d ed. Elsevier Academic
Carlo Andreotti 0000-0003-2588-8723 Press, London, UK. DOI: 10.1016/C2017-0-04744-4
Damiano Zanotelli 0000-0002-7700-5761 Keller M and Shrestha PM. 2014. Solute accumulation differs in the
Markus Keller 0000-0003-2144-2388 vacuoles and apoplast of ripening grape berries. Planta 239:633-
642. DOI: 10.1007/s00425-013-2004-z
Keller M, Deyermond LS and Bondada BR. 2015a. Plant hydraulic
Citation conductance adapts to shoot number but limits shoot vigour in
Wenter A, Andreotti C, Zanotelli D and Keller M. 2025. Alleviating water stress grapevines. Funct Plant Biol 42:366-375. DOI: 10.1071/FP14206
at veraison may trigger berry shrivel disorder in susceptible grapevines. Am
J Enol Vitic 76:0760010. DOI: 10.5344/ajev.2025.25009 Keller M, Zhang Y, Shrestha PM, Biondi M and Bondada BR. 2015b.
Sugar demand of ripening grape berries leads to recycling of
surplus phloem water via the xylem. Plant Cell Environ 38:1048-
Data Availability 1059. DOI: 10.1111/pce.12465

The data underlying this study are available on request from the Keller M, Shrestha PM, Hall GE, Bondada BR and Davenport JR. 2016.
corresponding author. Arrested sugar accumulation and altered organic acid metabolism
in grape berries affected by berry shrivel syndrome. Am J Enol
Vitic 67:398-406. DOI: 10.5344/ajev.2016.16048
References Knoblauch M, Knoblauch J, Mullendore DL, Savage JA, Babst BA,
Ameglio T, Morizet J, Cruiziat P, Martignac M, Bodet C and Raynaud Beecher SD et al. 2016. Testing the Münch hypothesis of long
H. 1990. The effects of root temperature on water flux, potential distance phloem transport in plants. eLife 5:e15341. DOI: 10.7554/
and root resistance in sunf lower. Agronomie 10:331-340. DOI: eLife.15341
10.1051/agro:19900407 Krasnow M, Matthews M and Shackel K. 2008. Evidence for sub-
Bondada BR and Keller M. 2012. Not all shrivels are created equal— stantial maintenance of membrane integrity and cell viability in
Morpho-anatomical and compositional characteristics differ normally developing grape (Vitis vinifera L.) berries throughout
among different shrivel types that develop during ripening of development. J Exp Bot 59:849-859. DOI: 10.1093/jxb/erm372
grape (Vitis vinifera L.) berries. Am J Plant Sci 3:879-898. DOI: Krasnow M, Weis N, Smith RJ, Benz MJ, Matthews M and Shackel K.
10.4236/ajps.2012.37105 2009. Inception, progression and compositional consequences of
Brockwell J and Gault RR. 1976. Effects of irrigation water tempera- a berry shrivel disorder. Am J Enol Vitic 60:24-34. DOI: 10.5344/
ture on growth of some legume species in glasshouses. Aust J Exp ajev.2009.60.1.24
Agric Anim Husb 16:500-505. DOI: 10.1071/EA9760500 Palai G, Caruso G, Gucci R and D’Onofrio C. 2022. Berry flavonoids
Chang B-M and Keller M. 2021. Cuticle and skin cell walls have are differently modulated by timing and intensities of water deficit
common and unique roles in grape berry splitting. Hort Res 8:168. in Vitis vinifera L. cv. Sangiovese. Front Plant Sci 13:1040899. DOI:
DOI: 10.1038/s41438-021-00602-2 10.3389/fpls.2022.1040899
Chang B-M, Zhang Y and Keller M. 2019. Softening at the onset of Palliotti A, Tombesi S, Frioni T, Famiani F, Silvestroni O, Zamboni
grape ripening alters fruit rheological properties and decreases M et al. 2014. Morpho-structural and physiological response of
splitting resistance. Planta 250:1293-1305. DOI: 10.1007/s00425- container-grown Sangiovese and Montepulciano cvv. (Vitis vi-
019-03226-y nifera) to re-watering after pre-veraison limiting water deficit.
Cooley NM, Clingeleffer PR and Walker RR. 2017. Effect of water Funct Plant Biol 41:634-647. DOI: 10.1071/FP13271
deficits and season on berry development and composition of Raifer B, Weitgruber T, Schmid A , Haas F and Pertoll G. 2023.
Cabernet Sauvignon (Vitis vinifera L.) grown in a hot climate. Aust Besteht ein Zusammenhang zwischen Extremwetterereignissen
J Grape Wine Res 23:260-272. DOI: 10.1111/ajgw.12274 und dem Auftreten von Traubenwelke? Laimburg J 5:2023.007.
Fasoli M, Richter CL, Zenoni S, Bertini E, Vitulo N, Dal Santo S et DOI: 10.23796/LJ/2023.007
al. 2018. Timing and order of the molecular events marking the Rienth M and Scholasch T. 2019. State-of-the-art tools and methods
onset of berry ripening in grapevine. Plant Physiol 178:1187-1206. to assess vine water status. OENO One 4:619-637. DOI: 10.20870/
DOI: 10.1104/pp.18.00559 oeno-one.2019.53.4.2403
Fuentes S, Sullivan W, Tilbrook J and Tyerman S. 2010. A novel analy- Rogiers SY and Clarke SJ. 2013. Nocturnal and daytime stomatal
sis of grapevine berry tissue demonstrates a variety-dependent conductance respond to root-zone temperature in ‘Shiraz’ grape-
correlation between tissue vitality and berry shrivel. Aust J Grape vines. Ann Bot 111:433-444. DOI: 10.1093/aob/mcs298
Wine Res 16:327-336. DOI: 10.1111/j.1755-0238.2010.00095.x
Romero P, Botía P and Keller M. 2017. Hydraulics and gas exchange
Griesser M, Savoi S, Bondada B, Forneck A and Keller M. 2024. Berry
recover more rapidly from severe drought stress in small pot-grown
shrivel in grapevine: A review considering multiple approaches. J
grapevines than in field-grown plants. J Plant Physiol 216:58-73.
Exp Bot 75:2196-2213. DOI: 10.1093/jxb/erae001
DOI: 10.1016/j.jplph.2017.05.008
Groenveld T, Obiero C, Yu Y, Flury M and Keller M. 2023. Predawn
Scheenen TWJ, Vergeldt FJ, Heemskerk AM and Van As H. 2007.
leaf water potential of grapevines is not necessarily a good proxy
Intact plant magnetic resonance imaging to study dynamics in
for soil moisture. BMC Plant Biol 23:369. DOI: 10.1186/s12870-
long-distance sap flow and flow-conducting surface area. Plant
023-04378-6
Physiol 144:1157-1165. DOI: 10.1104/pp.106.089250
Hall GE, Bondada BR and Keller M. 2011. Loss of rachis cell viability
is associated with ripening disorders in grapes. J Exp Bot 62:1145- Seem RC. 1984. Disease incidence and severity relationships. Ann Rev
1153. DOI: 10.1093/jxb/erq355 Phytopathol 22:133-150. DOI: 10.1146/annurev.py.22.090184.001025

American Journal of Enology and Viticulture — ajevonline.org 9 of 10 2025 Volume 76 Article 0760010
Wenter et al. Berry Shrivel After Water Stress Alleviation

Shahood R, Torregrosa L, Savoi S and Romieu C. 2020. First quan- Zhang Y and Keller M. 2017. Discharge of surplus phloem water may
titative assessment of growth, sugar accumulation and malate be required for normal grape ripening. J Exp Bot 68:585-595. DOI:
breakdown in a single ripening berry. OENO One 54:1077-1092. 10.1093/jxb/erw476
DOI: 10.20870/oeno-one.2020.54.4.3787 Zhang X-Y, Wang X-L, Wang X-F, Xia G-H, Pan Q-H, Fan R-C et al.
van Doorn WG and Woltering EJ. 2004. Senescence and programmed 2006. A shift of phloem unloading from symplasmic to apoplasmic
cell death: Substance or semantics? J Exp Bot 55:2147-2153. DOI: pathway is involved in developmental onset of ripening in grape
10.1093/jxb/erh264 berry. Plant Physiol 142:220-232. DOI: 10.1104/pp.106.081430
Xiao Z, Rogiers SY, Sadras VO and Tyerman SD. 2018. Hypoxia in Zufferey V, Spring J-L, Voinesco F, Viret O and Gindro K. 2015.
grape berries: The role of seed respiration and lenticels on the Physiological and histological approaches to study berry shrivel in
berry pedicel and the possible link to cell death. J Exp Bot 69:2071- grapes. OENO One 49:113-125. DOI: 10.20870/oeno-one.2015.49.2.89
2083. DOI: 10.1093/jxb/ery039
Zhang Y and Keller M. 2015. Grape berry transpiration is determined
by vapor pressure deficit, cuticular conductance, and berry size.
Am J Enol Vitic 66:454-462. DOI: 10.5344/ajev.2015.15038

American Journal of Enology and Viticulture — ajevonline.org 10 of 10 2025 Volume 76 Article 0760010