INFLUENCE OF VINE WATER STATUS ON ABOVEGROUND
BIOMASS PRODUCTION AND PARTITIONING
INFLUENCE DE L’ÉTAT HYDRIQUE DE LA VIGNE SUR
LA PRODUCTION ET LA RÉPARTITION DE LA BIOMASSE AÉRIENNE
Carlos MIRANDA1,*, Luis G. SANTESTEBAN1, José M. ESCALONA2, Hipolito MEDRANO2,
Felicidad DE HERRALDE3, Xavier ARANDA3 , Monserrat NADAL4,
Diego S. INTRIGLIOLO5, Juan R. CASTEL6, José B. ROYO1
1
Departamento de Producción Agraria, Universidad Pública de Navarra, 31006 Pamplona (Spain)
2
Departament de Biologia, Universitat de les Illes Balears, 07122 Palma de Mallorca (Spain)
3
IRTA. Ecophysiology. Torre Marimon. 08140 Caldes de Montbui (Spain)
4
Facultat d’Enologia, Universitat Rovira I Virgili, 43007 Tarragona (Spain)
5
Centro de Edafología y Biología Aplicada del Segura/CSIC. 30100 Espinardo. Murcia (Spain)
6
IVIA, Centro Desarrollo Agricultura Sostenible, 46113 Moncada (Spain)
*
Corresponding author : C. Miranda, 34948169850, 34948169732, Email : [email protected]
Abstract
Water availability is considered a determinant factor for grape-growing sustainability since vineyard water relations critically
impacts on vine performance and grape and wine composition. Improving water use efficiency (WUE) is therefore of crucial
importance for a sustainable viticulture in Mediterranean and semi-arid regions. In a previous work, we defined ranges and
relationships for biomass production and partitioning among vines of several cultivars at vineyard scale, using an extensive
database of vegetative growth and yield data. In this study those ranges and relationships are used as a tool for evaluating the
influence of seasonal vine water status on biomass production and partitioning in vineyards throughout
semi-arid regions in Spain (Navarra, Cataluña, Baleares and Valencia). Aboveground vegetative growth and yield data were
transformed to biomass values using allometric relationships and partitioning among vegetative and reproductive structures was
calculated. Biomass production and partitioning in the vineyards fell within the confidence ranges of the reference relationships.
Water availability positively influenced biomass production, increasing it up to a 50% respect rain-fed vineyards. Moreover,
seasonal stem water potential differences of 0.1 MPa were needed to modify aboveground biomass production in 100-150 g·vine-1.
Partitioning was also affected, as lower water availability favored up to a 9-12% higher biomass allocation to bunches in rain-fed
vineyards. Overall, the results obtained provide a tool to better evaluate water use efficiency in vineyards.
Keywords : Water use efficiency, partitioning, biomass.
Résumé
La disponibilité de l'eau est considérée comme un facteur déterminant pour la durabilité de la viticulture depuis que les relations de
l'eau de la vigne impacts gravement sur les performances de la vigne et le raisin et la composition du vin. Améliorer l'utilisation
efficace de l'eau (WUE) est donc d'une importance cruciale pour une viticulture durable dans les régions semi-arides et
méditerranéenne. Dans un travail précédent, nous avons défini la gamme de production de biomasse et le partitionnement entre les
vignes de plusieurs cultivars à l'échelle du vignoble, en utilisant une base de données étendue de croissance et de rendement
végétative. L'objectif de ce travail était de d'évaluer l'influence du statut hydrique de la vigne de saison sur la production de biomasse
et le partitionnement des vignobles dans plusieurs régions semi-arides en Espagne (Navarre, Catalogne, Iles Baléares et de Valence)
utilisant des relations obtenues dans le étude précédente. Les données brutes de croissance aérienne végétative et de rendement ont
été transformées en valeurs de biomasse suivant des relations allométriques ainsi que leur répartition. La production de biomasse et le
partitionnement dans les vignobles se situent dans les gammes de confiance des relations de référence. La disponibilité en eau la
production de biomasse positivement influencé, l'augmenter jusqu'à 25% respect vignobles non irriguées. En outre, variations du
potentiel hidrique tige de 0.1 MPa ont été nécessaires pour modifier la production de la biomasse aérienne dans 100-150 g·vigne-1.
Partitionnement a été affecté outre, comme la disponibilité de l'eau réduite favorisée jusqu'à une allocation de la biomasse de 9-12%
supérieur aux grappes dans les vignobles non irriguée. Dans l'ensemble, les résultats obtenus fournissent un outil pour l'évaluer
l'efficacité d'utilisation de l'eau dans les vignobles.
Mots-clés : Efficacité de l'Usage de l'eau, partitionnement, biomasse
1. Introduction
For viticulture, sustainability is becoming a serious concern due to the high extension of the crop in many different climatic
conditions and high inputs required. Water availability is considered a determinant factor for grape-growing sustainability,
particularly in arid and semi-arid areas, since vineyard water relations critically impacts on vine performance and grape and wine
composition (Santesteban and Royo, 2006; Flexas et al, 2010). Moreover, climate change scenarios suggest an increasing
dependence of grape production on irrigation because of higher global air temperatures, heat waves, droughts and uneven rainfall
patterns (Jones et al. 2005, IPCC 2013). Improving water use efficiency (WUE) is therefore of crucial importance for a sustainable
viticulture in Mediterranean and semi-arid regions. Stem water potential (!s) measured at mid-morning or noon can be effectively
used in semi-arid regions to determine to what extent a vineyard is under water deficit, because it integrates all the potential daily
factors that can impact water status (Santesteban et al, 2010). In previous works, Miranda et al. (2013, 2014) estimated the annual
production of biomass for vineyards of several cultivars, using an extensive database of vegetative growth and yield data that allowed
to define the ranges of biomass production and partitioning among vines. Moreover, significant relationships explaining vine
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performance were found between vine vigour and biomass production, and between yield and vegetative/reproductive biomass
allocation. In this study those ranges and relationships are used as a tool for evaluating the influence of water status on biomass
production and partitioning in vineyards throughout semi-arid regions in Spain (Navarra, Cataluña, Baleares and Valencia).
2. Materials and methods
Model building for vine biomass production and partitioning. A detailed database including vegetative and reproductive
development data, yield and weather for 622 vineyards of Tempranillo, Cabernet Sauvignon, Grenache (syn ‘Garnacha’) and
Graciano was used. Data was obtained between 2008 and 2010 as a part of the vineyard-modelling project VitMod (Santesteban et al,
2012). All vineyards were commercially grown in Navarra and La Rioja regions in Spain and encompassed a relatively wide range of
climatic conditions that mainly resulted from differences in altitude (from 307 to 556 m above sea level). Aboveground vegetative
growth and yield data were transformed to biomass values using the allometric relationships detailed in Miranda et al. (2013, 2014).
Regressions using R v3.1.2 (R core team, 2014) with General Linear Model tool in R commander package (Fox, 2005) were fitted
among vine vigour (defined as trunk size per row meter) and biomass production, and among yield and vegetative/reproductive
biomass allocation, pooling the data of all cultivars and years.
Influence of seasonal vine water status on biomass production and partitioning. A database including vegetative and reproductive
development data, yield, and stem water potentials measured from fruit set to harvest for 225 vineyards of Tempranillo, Cabernet
Sauvignon, Grenache, Marselan and Muscat was used (Table 1).
Table 1. Mean values, and ranges for vine characteristics and seasonal water potentials in the vineyards studied.
Tableau 1. Valeurs moyennes, et les potentielles tiges saison obtenus dans les vignobles étudiés.
cv.z
Locationsy
Number
of
vineyards
Vine vigour
(Vm, cm2 trunk section · m-1
row)
Mean
Min
Max
75
5.8
1.1
15.1
GN
IRTA,
URV
TE
UIB, UPNA
62
12.4
3.9
23.2
CS
IRTA, URV
20
6.8
2.8
12.0
MS
IRTA, URV
36
6.3
4.1
9.6
32
29.1
23.4
33.6
MU IVIA
UiB,
Yield
(kg · m-1 row
distance)
Mea Mi
n
n
Max
0.4
2.59
5
5.70
0.6
3.18
5
9.33
0.9
1.70
2
2.86
1.4
2.77
6
4.14
2.9
8.12
0
13.25
!s-noon
(fruit set-harvest, MPa)
Mean
Min
Max
-1.09
-1.46
-0.64
-0.95
-1.44
-0.47
-1.21
-1.38
-1.00
-1.27
-1.59
-0.83
-0.74
-0.98
-0.53
y
Location of the vineyards: IRTA,Barcelona; IVIA Valencia; UiB, Balearic Is.; UPNA, Navarra; URV, Tarragona
z
GN: Grenache, TE: Tempranillo, CS: Cabernet Sauvignon, MS: Marselan, MU: Muscat
Data was obtained between 2010 and 2014 as a part of the Spanish government funded BACAVID project
(Ref: AGL2011-30408-C04-03) on water and carbon balances in vineyard. The vineyards included commercial and experimental
plots, and were grown in Cataluña, Baleares, Valencia and Navarra regions in Spain, under warm climate conditions. Vegetative
growth and yield data were transformed to biomass values with the allometric relationships used in the model building section.
Biomass production and partitioning values were compared to the ranges defined by the confidence intervals of the regressions
obtained using VitMod data. The effect that vine vigour and seasonal water status from fruit set to harvest (!s-FSH) measured at noon
had on biomass production, and that yield and !s-FSH had on biomass allocation, respectively, were evaluated through multiple
regression analysis using R v3.1.2.
3. Results and discussion
The set of vineyards included in the VitMod database covers a broad range of vegetative and productive situations (Miranda et al,
2013, 2014) and they allow obtaining generalizable results (Fig 1a,b). As previously reported by Miranda et al. (2013), strong
relationships (R2=0.630, p<0.0001) were found (Fig 1a) between biomass production expressed on a per trunk area basis (kg·cm-2)
and vine size was expressed on a row meter basis (cm2 trunk cross sectional area · m-1 vine spacing).
The range of situations covered by the vineyards in the BACAVID dataset was very similar to VitMod ones (Fig 1a), and the
biomass production values fell within the confidence limits defined for VitMod data for most of the vineyards, the exceptions being
mostly young ("4 years-old) “Grenache” plots. Biomass production was also strongly related to vine vigour (R2=0.551, p<0.0001).
Viticultural research has seldom taken trunk size and growth into account. However, estimation of trunk cross-sectional area allows
an integration of vineyard carbohydrate balance (Santesteban et al., 2010) and is well related to those factors limiting vine
photosynthetic activity.
!
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Figure 1. Influence of (a), (c) vine vigour on biomass produced by the vines and (b), (d) of yield on allocation in vegetative
organs. Cultivars and institutions are indicated in (a) and (b), in (c) and (d) vineyards are classified in seasonal water stress
levels.
Figure 1. Influence de (a), (c) la vigueur de la vigne sur la biomasse aérienne produite par la vigne et du rendement sur la
répartition de la biomasse aérienne. Cultivars et les institutions sont indiqués en (a) et (b), alors que dans (c) et (d) les vignobles
sont classés dans le niveau de stress de l'eau saisonniers.
Strong relationships (R2>0.67, p<0.001) were also found for VitMod vineyards between yield, measured as dry weight of bunches·m1
of vine spacing in the row (Fig 1b) and the proportion of biomass allocated to vegetative structures (trunk, leaves and canes). The
vineyards from BACAVID dataset fell also within the confidence limits of that relationship (except for the above-mentioned young
“Grenache” fields), and a strong relationship yield/allocation for BACAVID data was also found (R2=0.57, p<0.001). The slopes of
both relationships were similar (p>0.05), but the intercepts indicated that, for any given yield, BACAVID vineyards consistently
allocated around 10% less biomass to their vegetative structures.
This pattern could be due to BACAVID vineyards were grown under warmer conditions than most of VitMod ones. In fact, similar
effects on sink : source ratios were found in field-grown Shiraz by Sadras and Moran (2013) and Miranda et al (2014).
Partitioning to bunches, as fruit/pruning weight ratio, is a way to evaluate sink : source ratio in vines, and their increase with
warming is important for viticulture. Empirical ranges of the fruit/pruning weight ratio are commonly used to guide management
practices such as pruning and training to achieve specific viticultural targets. The effect of zone and climate found in this study
indicates that those indices could not be considered as constant across regions or years, and also that long-term carbon balances of
vines might be affected by global warming.
The influence of seasonal water status on the abovementioned relationships was analysed classifying BACAVID vineyards in four
levels according to the water deficit thresholds suggested by Van Leeuwen et al (2009), from weak to severe water deficit (Fig 1c,d).
When all cultivars were pooled, water availability positively influenced biomass production, so that vineyards grown under weak
seasonal water deficit measured at noon (!s-FSH> -0.9 MPa) produced up to 50% more biomass compared with vineyards under
severe water deficit (!s-FSH< -1.1 MPa). Vine vigour and !s-FSH showed a close relationship to biomass production (R2=0.575,
p<0.001, Table 2), although the relative weight of !s-FSH, estimated with the standardized coefficient (#) was smaller than that of
vine vigour.
The relationship showed that seasonal !s-FSH increases of 0.1 MPa were needed to decrease aboveground biomass production in 100150 g · vine-1. Allocation patterns were also affected by water availability throughout the season, as vineyards grown under weak
deficit consistently allocated 9-12% more biomass in their vegetative structures, and a strong relationship of yield and !s-FSH with
allocation (R2=0.683, p<0.001, Fig 1d) was found, with water deficit also showing smaller relative weight. The range of situations
available for “Tempranillo” and “Grenache” was broader than in the rest of cultivars, allowing analysing the effect of water deficit on
each one separately (Table 2).
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Table 2. Linear regression coefficients between a) Vine vigour (Vm) and seasonal water deficit (!S-FSH)
and biomass production (DWT) and b) biomass allocated in vegetative structures (BAV)
and yield expressed on dry weight basis (DWB) and !S-FSH.
Tableau 2. Coefficients de régression linéaire entre a) la vigueur de la vigne (Vm) et le déficit d'eau saisonnier (!S-FSH) et la
production de biomasse (DWT) et b) la répartition de la biomasse dans les structures végétatives (BAV),
le rendement du poids sec (DWB) et !S- FSH.
Regression variables
Coefficientsz
Rel. Weight (")
cv
R2
R2adj
df
b0
b1
b2
b1
b2
•
Log (DWT)=b0+b1*Vm+b2*!S-FSH
All
GN
TE
0.575
0.385
0.656
0.571
0.367
0.645
226
72
63
6.29
6.22
6.77
-0.05
-0.07
-0.09
0.63
0.46
0.79
-0.89
-0.64
-0.82
0.32
0.23
0.30
•
BAV=bo+b1*Log(DWB)+!S-FSH
All
GN
TE
0.683
0.319
0.707
0.680
0.299
0.698
229
69
63
170.30
124.00
179.62
-17.38
-10.64
-16.71
10.82
10.32
20.19
0.87
0.60
0.83
-0.19
-0.25
-0.35
Z
Regression and coefficients are significant at p<0.001 level except for the underlined coefficient (ns).
Significant relationships were obtained for both cultivars, but for “Tempranillo” the response was stronger
(better relationships as indicated by their R2), and also only in this cultivar a significant effect of !s-FSH could be observed. These
results match with the fact that “Tempranillo” is traditionally grown in cooler areas, where water deficit is unusual, and anisohydric
water management mechanisms prevail in this cultivar (Medrano et al, 2003; Santesteban et al, 2009). Results also match with the
fact that “Grenache” is autochthonous from drier areas, and also with is considered to behave as nearly isohydric (Schultz, 2003;
Santesteban et al, 2009).
4. Conclusion
Allometric equations applied on an extensive database of vegetative growth and yield data for vineyards have allowed us to confirm
that the biomass production and allocation patterns and relationships found in VitMod data can be also observed in different cultivars
grown under warmer conditions and, therefore, be generalizable to vineyards grown in semi-arid conditions.
The influence of seasonal water status seems to agree with the isohydric/anisohydric response of the cultivars. In vineyards grown
under severe seasonal water deficits vineyards could decrease their biomass production up to a 35-40% respect to irrigated vineyards
under weak or no seasonal deficit.
Moreover, seasonal stem water potential differences of 0.1 MPa were needed to modify aboveground biomass production in 100-150
g · vine-1. Partitioning was also affected, as higher water availability favored up to a 9-12% higher biomass allocation to the
vegetative structures in irrigated vineyards. Overall, the results obtained provide a tool to better evaluate water use efficiency in
vineyards.
5. Acknowledgements
This work was developed with funding provided by the Spanish Ministry of Economy and Competitiveness (BACAVID, Ref:
AGL2011-30408-C04-00) and by Dpt. Innovación, Industria & Empleo of the Government of Navarra (MODELVID,
Ref:IIM11879.RI.1).
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