Journal of Plant Physiology 169 (2012) 1023 1031
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Journal of Plant Physiology
jou rn al h o mepage: www.elsev ie
Berry morphology and composition in irrigated a(Vitis vinifera L.)
Adriano Sofoa,, Vitale Nuzzoa, Giuseppe Tatarannia, Michele M
a Dipartimento asilicab Dipartimento Potenc Dipartimento sano 4
a r t i c l
Article history:Received 7 November 2011Received in revised form 6 March 2012Accepted 7 March 2012
The present study was carried out in a 5-year-old vineyard (Vitis vinifera L., cv. Aglianico) located in South-ern Italy. Half of the plants (IRR) were fully irrigated, whereas the other half were not irrigated (NIRR). Inboth of the treatments, plant water status, gas exchange, photosynthetic efciency and productive per-formance were determined. The arid conditions resulted in signicant decreases in stem water potentialin NIRR (minimum values of 1.34 and 1.52 MPa in IRR and NIRR, respectively). The values of yield perplant, cluster weight and total berry weight were signicantly higher in IRR. Grape berries were sepa-
Among responsibleThey are loskins and l2007; Gam
Abbreviatiospiration; ETo,photochemistrplants; PPFD, pleaf-to-air vapwater potentia
Correspone dellAmbient85100 Potenza
0176-1617/$ http://dx.doi.orated into four weight classes, and morphometric and microscopic analyses were carried out to measureand calculate berry skin characteristics. Irrigation determined a marked shift toward heavier (+23% in theclass 1.25 g) and bigger (336.35 mm3 vs 299.15 mm3) berries, and induced signicant changes in othermorphometric berry parameters. No differences among berry weight classes and irrigation treatmentswere observed for berry skin thickness. In all of the berry weight classes, total anthocyanins extractedfrom berry skins were signicantly higher in NIRR than in IRR (12301.53 and 9585.52 mg kg1 fresh berryskin, respectively), and appeared to be positively related to berry weight, whereas total avonols werenot signicantly different between the two treatments. Qualitative changes in the levels of single antho-cyanin and avonol compounds were detected between IRR and NIRR. In addition, iron, copper and zinc,whose high concentration can negatively affect wine quality, were signicantly higher in the IRR treat-ment. The results highlighted that the absence of irrigation did not determine decreases in grape quality.Such data can be of primary importance in environments where water availability is by far the mostimportant limiting factor for plant growth.
2012 Elsevier GmbH. All rights reserved.
avonoids, anthocyanins are pigmented compounds for the dark blue coloration of ripened grape berries.cated in the vacuoles of the hypodermal cells of berryeaf epidermis (Clarke and Bakker, 2004; Conde et al.,buti et al., 2007; Ristic et al., 2007). Flavonols, another
ns: An, net photosynthesis; E, transpiration; ETc, cultural evapotran- reference evapotranspiration; Fv/Fm, maximum quantum yield of PSIIy; gs, stomatic conductance; IRR, irrigated plants; NIRR, non-irrigatedhotosynthetic photon ux density; SSC, soluble solid content; VPD,or pressure decit; PSII , effective quantum yield of PSII; w, steml.ding author at: Dipartimento di Scienze dei Sistemi Colturali, Forestalie Universit degli Studi della Basilicata Viale dellAteneo Lucano n. 10,, Italy. Tel.: +39 0971 206228; fax: +39 0971 204307.ress: firstname.lastname@example.org (A. Sofo).
class of avonoids, are colorless berry skin constituents generallyconsidered to act as UV protectants and free-radical scavengers, andto contribute to wine color as anthocyanin co-pigments (Downeyet al., 2006; Terrier et al., 2009). The quantity and composition ofanthocyanins and avonols greatly depend on genetic factors (Bosset al., 1996), but their amount can be affected by environmentalfactors and cultural practices (Downey et al., 2006). In grapevine,water availability alters fruit chemical composition both directly,through the activation of specic metabolic pathways (Castellarinet al., 2007a,b), or indirectly through the control of berry size(Roby and Matthews, 2004; Roby et al., 2004) and other plantbiochemical and physiological processes (Conde et al., 2007). Atthe same time, grapevine anthocyanin and avonol content alsodepend on berry growth and berry skin characteristics, such asthickness and total surface per unit of volume, and the relativeproportion with respect to seeds and pulp, which is turn affectedby berry size (Hardie et al., 1996; Roby et al., 2004; Cadot et al.,2011).
see front matter 2012 Elsevier GmbH. All rights reserved.rg/10.1016/j.jplph.2012.03.007 di Scienze dei Sistemi Colturali, Forestali e dell Ambiente, Universit degli Studi della B di Chimica, Universit degli Studi della Basilicata, Viale dellAteneo Lucano 10, I-85100
di Chimica Farmaceutica e Tossicologica, Universit di Napoli Federico II, Via D. Monte
e i n f o a b s t r a c tr .de / jp lph
nd non-irrigated grapevine
anfrab, Mauro De Niscoc, Antonio Scopaa
ta, Via dellAteneo Lucano 10, I-85100 Potenza, Italyza, Italy9, I-80131 Napoli, Italy
1024 A. Sofo et al. / Journal of Plant Physiology 169 (2012) 1023 1031
Other than phenolics, grape and wine chemistry at the elemen-tal level (ionomic), including the content of all mineral nutrientsand trace elements, are poorly studied. All of the inorganic cationsare naturally present (not from winemaking equipment containingalloys nor ftions in graand copper(Ribreau-Gof the grapalso to the compositio
Based ongate anthocberries of Aety of soutworldwide (1) to evaluand physiolpolyphenolthe absencevide useful iprocesses a
The expthe harvestgrafted on 1Marina (42
is a climaticmation of 231 Octoberten rows oferal cordoncharacterizbetween thof 4000 vine
Half of t(from the eaequal to 10per each ofwhereas thwas calculatranspiratiothe culturalfor grapevinirrigation sthan 0.8 Mume was oby two drirological vawithin 50 mature, rainfataken throurecorded atLeaf-to-air to Goudrian
The planimental perstem water
the row, where microclimatic conditions and soil physic-chemicalcharacteristics were similar, were chosen. The values of w weremeasured around midday on 5 fully expanded leaves and well-lightened selected from each plant on fruiting shoots situated in
least et a
eachents cencd frof th
was70 techuoreS1 g moed wting lcencical ttedd lea
lightrk (Flighttry iriablh lea
harveld pd totoot wlantberreigh
ries perryter. Cean e, su
ratiod aserry ted fzed ws, puSC) ual rrom fertilizers and pesticides) at non-toxic concentra-pe berries and wine, and some of them, such as iron, play a major role in winemaking and wine qualityayon et al., 2006). Furthermore, the metal compositione berry is of concern not only to the viticulturist, butoenologist due to the direct impact on juice and mustn, which in turn affect wine quality.
this accepted knowledge, it was of interest to investi-yanin, avonol and metal composition within the grapeglianico, a high-quality and late-ripening grape vari-hern Italy, whose premium red wine is appreciated(Gambuti et al., 2007). The aims of this research wereate the effect of irrigation management on plant yieldogical status, and on berry morphologic characteristics,
and metal composition, and (2) to establish whether of irrigation can affect grape quality. This could pro-nformation for improving vine cultivation, winemakingnd nal product quality.
l site and plant material
eriment was carried out in 2008, from bud break to, in a 5-year-old vineyard (cv. Aglianico clone VCR11103 Paulsen) sited on a clay-lam soil in Montegiordano02N, 1635E; Southern Italy). According to Winkler, it
region 5, classied as very hot, with a thermic sum-603 C above the threshold of 10 C between 1 April and. The experimental plot, of about 0.30 ha, consisted of
spur-pruned vines to a permanent horizontal unilat-. Each vine, decked at 0.60 m above the ground, wased by about 8 spurs of 23 buds each. The distancee vines was of 2.5 m 1.0 m, with a nal plant densitys ha1. Rows were north-south oriented.he plants (IRR) were irrigated from 9 June to 1 Augustrly stages of fruit set to veraison) using a water amount0% of cultural evapotranspiration (ETc) (24 L per plant
ten irrigation turn at approximately 5-day intervals),e other half were not irrigated (NIRR). The value of ETcted using ETo Kc, where ETo is the reference evapo-n calculated according to Hargreaves method, and Kc is
coefcient during the experimental period, equal to 0.6e, according to Allen et al. (1996). In the watered plot,
tarted when the stem water potential ( w) was lowerPa and ended at veraison. The seasonal irrigation vol-
f 960 m3 ha1 (240 L plant1). Each vine was irrigatedp emitter per plant discharging 4 L h1 each. Meteo-riables were monitored by a weather station placed
of the experimental plot. Measurements of temper-ll, and photosynthetic photon ux density (PPFD) wereghout the experimental period. The values of PPFD were
1-h intervals, and daily integrated values were logged.vapor pressure decit (VPD) was calculated according
and Van Laar (1994).
status, gas exchange and chlorophyll uorescence
t water status was determined throughout the exper-iod on ten plants per treatment by measurements of
potential ( w). Plants located in the central part of
the mement C w, leabag at (Chon
Forsuremuoresselectezone oout ontosynt362-cmLED sosourcestant 3(Hansaphyll The FMexcitinequippsaturauoresan optdome adapteactinicthe dawhite chemisthe vaFo. Eacbeforeactual)
At and yiter, anper shsame pof the four wand x ple pop20 bermine bdiameical (mvolumas the(deneEach bseparadeioniof skintent (Sa manItaly). zone of the plant using a pressure chamber (PMS Instru-orvallis, OR, USA, model 600). For the determination ofwere covered with an aluminum foil and a polyethylene
2 h before each measurement for avoiding transpirationl., 2001).
treatment, the same ten plants used for w mea-were chosen to measure gas exchange and chlorophylle on ve fully-expanded and well-lightened leavesm each plant on fruiting shoots situated in the mediane cordon. Gas exchange measurements were carriedugust and 4 September using a portable Li-6400 pho-
system (Li-Cor, Lincoln, NE, USA) equipped with aide leaf chamber. Light was provided by an articial red
emitting at 670 nm, and an external bottled 12-g CO2 used to inltrate the leaf chamber with air at a con-mol mol1 CO2. A pulse modulated FMS1 uorometer
Instruments, Norfolk, UK) was used to carry out chloro-scence measurements along the experimental period.uorometer adopted a pulsed light source as a very weakdulating (amber) light, peaked at 594 nm. The FMS1 wasith a halogen white lamp source to generate a super-ight pulse of 17,000 mol m2 s1 applied for 0.7 s fore induction, and delivered to the leaf sample throughber probe inserted at 45 inclination into a closed black
over the leaf-clip. The basal uorescence yield in dark-ves (Fo) or the steady state uorescence yield under
(Fs) were measured. The maximal uorescence yield inm) and in light conditions (F m) were determined using
pulses. The maximum quantum yield of PSII photo-n dark-adapted leaves was calculated as Fv/Fm, wheree uorescence (Fv) is the difference between Fm andf was dark-adapted for 30 min by means of a leaf clipmeasurement. In light-adapted leaves, the effective (or
ntum yield of PSII (PSII) was calculated as (F m F s)/F m.
est, on 27 September 2008, the number of clusterser plant, cluster weight, number of berries per clus-al berry weight per cluster, and the number of leavesere determined on 30 plants per treatment. From the
s, three clusters per plant were randomly collected. Allies of these clusters were picked and separated intot classes: x < 0.60 g, 0.60 g x < 0.90 g, 0.90 g x < 1.25 g,
g. The weight class frequency distribution (% of sam-ion) per each treatment was calculated. For each plant,er each weight class were randomly chosen to deter-
fresh weight, berry equatorial diameter and berry polaronsidering that berry shape was approximately spher-equatorial/polar diameter ratio = 0.99 0.08), surface,rface/volume ratio, and skin specic surface (dened
berry surface/skin weight), and skin specic weight the ratio skin weight/berry weight) were calculated.was cut into two halves and the seeds and skins wererom the pulp through a small metal spatula, washed inater, dried with absorbent paper, and the fresh weightslps and seeds were determined. The soluble solid con-of each berry was obtained by squeezing his pulp onefractometer (model MT-032 ATC; Turoni & C., Forl,
A. Sofo et al. / Journal of Plant Physiology 169 (2012) 1023 1031 1025
Anthocyanin and avonol extraction and determination
At harvest, three clusters per plant were randomly sampledin the central and well-irradiated area of the canopy from threeplants locatsoil differenthe four wewith a scalstored for twere discaranalysis in berries wer(w/w) soluout for 24 hMinisart SFtingen, Germwere analythe anthocyextracts weraphy (HP berry skin eThe HPLCMeach peak utor coupledan ESI sour5 -Phenyl-Torrance, Csolution cocolumn wacratic gradi1 min, then1 min, and 1ow rate ofup to m/z 8tive ionizatavonols w(Extrasynth
Berry skcyanin andsolution (5:unit (MLS-1copper, zincinductivelyDRC II, Perkas follows: Meinhard n(L min1): p50 ms; intemaximum Iwere used, unidentiedwavelengthwith the otplastic contgrade HNO3pure waterfrom multi-the calibrattion solutioalso analyzetory equipmtreatment a
Berries deriving from the same clusters used for anthocyanin,avonol and metal analysis were used to measure skin thickness.
clasquatladetion. rente 80iraphkin wg So
datepreferen). Thre rediffereme
as tnt plyaniroscisticA) wfferest.
exp andn 15
fromdencn 12ring alue ust. Dboveher
SeptTc) dradiat valuis d
e anden de
envier reues oed in the central part of the rows, in order to minimizeces between the two treatments. Again, the berries ofight classes were detached from each cluster, peeledpel and the skin, rapidly frozen at 80 C, and thenhe analytical determinations. The berries of x < 0.60 gded, as they were not always sufcient for a completeIRR and NIRR plants. Five grams of skin of frozen grapee collected and placed in a 100 mL methanolHCl 0.75%tion at 20 C in the dark. The extraction was carried. The resulting extracts were ltered through 0.20 mCA sterile lters (Sartorius Stedim Biotech GmbG, Goet-
any), and immediately frozen at 80 C. Berry extractszed by high-perf...