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Am. J. Enol. Vitic. 60:1:50-56 (2009)
Copyright © 2009 by the American Society for Enology and Viticulture.
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Response of Container-Grown Girdled Grapevines to Short-Term Water-Deficit Stress

Takayoshi Yamane1,*, Katsutoshi Shibayama2, Yoji Hamana1 and Hiroshi Yakushiji3

1 Fruit Tree Research Division, Agricultural Technology Research Center, Hiroshima Prefectural Technology Research Institute, Akitsu, Higashi-Hiroshima, Hiroshima 739-2402, Japan; 2 Agriculture, Forestry, and Fisheries Department, Hiroshima Prefecture, Hiroshima 730-8511, Japan; and 3 Department of Grape and Persimmon Research Station, National Institute of Fruit Tree Science, Akitsu, Higashi-Hiroshima, Hiroshima 739-2494, Japan.

Acknowledgments: This work was supported by NARO Research Project 166, "Establishment of Agricultural Production Technologies Responding to Global Warming."

* Corresponding author (email: takayoshi.yamane{at}gmail.com)


   Abstract
Top
 Abstract
Materials and Methods
 Results
 Discussion
 Conclusions
 Literature Cited
 
The response of container-grown, girdled vines to water-deficit stress was examined by investigating the leaf net CO2 assimilation rate (A), stomatal conductance (gs), transpiration rate (E), predawn leaf water potential ({Psi}PD), and the fluctuations of stem diameter in vines. At the beginning of withholding irrigation, A, gs, and E were lower in girdled vines than in control vines. A and gs decreased 4 and 5 days after the start of withholding irrigation in control and girdled vines, respectively. {Psi}PD reached –1.13 MPa in the control vines and –0.67 MPa in girdled vines at 6 days after withholding irrigation. The higher soil water potential in water-deficient girdled vines than in control vines showed lower water consumption in girdled vines. Daytime stem contraction was smaller in girdled vines than in control vines while drying. Hence, results indicated that decreased water use as a result of girdling reduced the depletion of soil water in the containers, which caused high {Psi}PD and subsequent low stem contraction in girdled vines. Combined treatment of girdling and short-term strong water-deficit stress did not cause leaf wilt or decrease shoot growth in the subsequent growing season. These results reveal the response of girdled vines to water-deficit stress, especially severe and short-term water-deficit stress, in controlled conditions under which vines were grown in containers in a vinyl greenhouse.

Key words: girdling, stem fluctuation, stomatal conductance, water-deficit stress

Girdling prior to fruit set improves berry set (Brown et al. 1988), which has a positive effect on berry size after set (Carreño et al. 1998) and, if done at the beginning of the ripening phase, enhances skin color and berry ripening in grape (Carreño et al. 1998, Peacock et al. 1977, Yamane and Shibayama 2006a, Yamane et al. 2007). The anthocyanin concentration in grape berry skin is depressed by high temperature (Kataoka et al. 1984, Kliewer and Lider 1970, Mori et al. 2007, Yamane et al. 2006, Yamane and Shibayama 2006b). Poor skin color of grapes grown in warm regions of Japan is one of the most serious problems associated with grape production and girdling is a practical technique used in commercial vineyards to enhance skin coloration (Yamane et al. 2007). However, girdling has been reported to have a debilitating effect on vine growth when combined with water-deficit stress (Winkler 1962), although the actual status of water-deficit stress and vine conditions were not shown. Thus, we have tried to clarify the effects of water-deficit stress on girdled vines.

Girdling has been reported to decrease leaf net CO2 assimilation rate (A) and stomatal conductance (gs) in grape (Düring 1978, Harrell and Williams 1987, Hofacker 1978, Roper and Williams 1989). A and gs also decreased under water-deficit stress conditions (El-Ansary and Okamoto 2007); therefore, comprehensive consideration of the effect of water-deficit stress on girdled vines is needed.

There are several instances in which changes in stem diameter have been used as an indicator for plant water status (Fujita et al. 2003a, 2003b, Imai et al. 1991, Ito et al. 2002, Klepper et al. 1971). Changes in stem diameter reflect changes in stem tissue hydration, and measurement of stem diameter appears to be more sensitive than other plant parameters to changes in net radiation (Klepper et al. 1971). There is a simple technique for measuring micromorphometric shrinkage and expansion of the stem diameter (Iwao and Takano 1988). It is anticipated that measuring the fluctuations of stem diameter in girdled vines can clarify the responses of the girdled vines to water-deficit stress in detail. Our research here investigated the effects of water-deficit stress on leaf A, gs, transpiration rate (E), predawn leaf water potential ({Psi}PD), and the fluctuations of stem diameter in girdled vines. The experiment was conducted in a vinyl greenhouse to avoid rainfall and container-grown vines were used to obtain precise results under severe water-deficit conditions.


   Materials and Methods
Top
 Abstract
 Materials and Methods
Results
 Discussion
 Conclusions
 Literature Cited
 
 Plant material and treatments.   Four-year-old Aki Queen (Vitis labrusca L. x V. vinifera L.) grapevines grafted on 5BB rootstocks in a vinyl greenhouse were used for study. Vines were grown in 60-L plastic containers with a sand and compost mixture (9:1 v/v) at a soil depth of 30 cm in Higashi-Hiroshima, Japan. A compound fertilizer containing 4.5, 3, and 4.5 g of nitrogen (N), phosphorus (P), and potassium (K), respectively, was applied per vine before budbreak. Until the onset of withholding irrigation from vines, 4 L of water was added to each container automatically when the soil water potential decreased to –0.01 MPa, the level at which water-deficit stress is relieved in container-grown vines (Imai et al. 1991). Vines were trained to a flat-top trellis with a single trunk. Each vine developed six to nine shoots, some with one or two clusters. Three weeks after budbreak, shoots were thinned to three per vine to maintain optimal vine vigor. Each primary shoot was pinched before blooming, retaining 12 nodes; after which each lateral shoot was pinched once a week to retain one leaf. Clusters were thinned to three per vine two weeks after blooming. The number of berries was also thinned to 32 berries in each cluster at the time of cluster thinning. Trunks were girdled to a width of 20 mm on 6 July 2005, 35 days after bloom. A utility knife was used for girdling. To avoid girdle closure when irrigation was withheld and to clarify the debilitating effects on vine growth, girdling was conducted at wider distances than 5 mm, which is the usual practice. Therefore, the girdle did not heal until the end of the treatment. Control vines were not girdled. Six vines of uniform growth were selected before girdling, and three replications (vines) were performed for each treatment.

The withholding of irrigation began after full watering (11 L per container) at 5:00 hr on 8 July, which was two days after girdling and before veraison. Withholding irrigation was stopped at 8:50 hr on 15 July with the addition of 11 L of water for each container. After the addition, 4 L of water for each container was automatically added when soil water potential decreased to –0.01 MPa to keep the vines well watered. Soil water potential at 15 cm depth was monitored every 10 min with a tensiometer (DM-8HG; Takemura Denki Seisakusho, Tokyo, Japan), which had a digital output connected to a voltage recorder (VR-71; T&D Co., Nagano, Japan). Photon flux density (PFD) was also monitored every 10 min using a quantum light sensor (LI-190; LI-COR, Lincoln, NE).

 Photosynthesis, stomatal conductance, and transpiration.   Leaf A, gs, and E were measured on mature leaves attached between the sixth and eighth nodes using a portable infrared gas analyzer (LI-6400; LI-COR). Measurements were conducted between 13:00 and 13:30 hr at 0, 2, 3, 4, 5, 6, 7, and 11 days after treatment. The leaf area clipped by the chamber was 6 cm2. Leaf chamber temperature, relative humidity, and CO2 concentration were 26–31°C, 40–90%, and 400 – 490 µmol mol–1, respectively; these were the natural conditions in the vinyl greenhouse. The light source provided by the equipment was adjusted to a PFD of 1000 µmol m–2 s–1. On 12 July, four days after treatment, the diurnal change of A, gs, and E were measured at 7:00, 9:00, 11:00, 13:00, 15:00, and 17:00 hr. In the leaf gas exchange measurement conducted on 12 July, PFD was set to the natural light intensity at each time, which was measured using a light sensor in the portable gas analyzer, except at 13:00 hr, when the PFD was set to 1000 µmol m–2 s–1.

 Stem diameter and predawn leaf water potential.   Stem diameters were recorded at 10-min intervals following a micromorphometric technique with a micro-displacement detector of the shrinkage type (Imai et al. 1990, Iwao and Takano 1988). The stem of a shoot was passed through a clamp screw and Tygon tubing (10 mm diam, 5 mm long). The displacement sensor was placed between the clamp screw and stem. The variation in the stem diameter was transmitted through the Tygon tubing to the displacement sensor, which was connected to a computerized data acquisition system (NEC, Sanei Kogyo, Tokyo). Blank runs were conducted by putting a glass rod in place of the plant sample, and the measurement sensitivity was within the limit of 2 µm. Sensors were placed on each vine for each treatment. Response patterns were similar from vine to vine, and the averages of the three vines in each treatment were recorded.

{Psi}PD was measured at 0, 2, 4, 5, 6, and 11 days after withholding irrigation using a pressure chamber between 3:30 and 4:00 hr each day (DIK-700; Daiki Rika Kogyo, Saitama, Japan). Three leaves were used for each measurement.

 Shoot growth and yield of subsequent growing season.   Shoot length and leaf area of the subsequent growing season were measured on 4 Aug 2006. Subsequent growing seasonal yield was measured on 31 Aug 2006 at harvest. Methods for the cultivation of vines were the same in both treatments.

 Statistical analysis.   The t test was used to evaluate significant differences in {Psi}PD, A, gs, and E between treatments. Excel software (Microsoft, Redmond, WA) and the Excel statistical package (Esumi, Tokyo) were used for statistical calculations.


   Results
Top
 Abstract
 Materials and Methods
 Results
Discussion
 Conclusions
 Literature Cited
 
The weather was cloudy and PFD was low for the first three days of withholding irrigation (Figure 1Go). Soil water potential of each treatment decreased slowly during this time, after which it decreased rapidly until 5 days after withholding irrigation, when it reached –0.08 MPa, the minimum value measurable by the tensiometer. Soil water potential decreased faster in control vines than in girdled vines (Figure 2Go). After adding water 6 days after withholding irrigation, the soil water potential was maintained at over –0.01 MPa by irrigation every 1 or 2 days.


Figure 1
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  		Figure 1 Photon flux density (PFD) in the greenhouse during the experiment. Each point indicates PFD at 10-min intervals.

 

Figure 2
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  		Figure 2 Effects of withholding irrigation and reirrigation on soil water potential in girdled and control vines. Arrows indicate irrigation.

 
{Psi}PD was significantly lower in control vines than in girdled vines during treatment and decreased markedly to –1.13 MPa in control vines and –0.67 MPa in girdled vines at 6 days after withholding irrigation (Figure 3Go). At 6 days after treatment leaf wilting did not appear in girdled vines, but did appear in control vines (Figure 4Go). {Psi}PD in girdled vines recovered after adding water, although {Psi}PD in control vines did not return to the value it had at the beginning of the treatment.


Figure 3
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  		Figure 3 Effects of withholding irrigation and reirrigation treatment on predawn leaf water potential ({Psi}PD) in girdled and control vines. Arrow indicates irrigation; * and NS indicate significance at p < 0.01 and not significant, respectively.

 

Figure 4
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  		Figure 4 Effects of withholding irrigation on leaf wilting at 6 days after treatment in girdled (A) and control (B) vines.

 
A, gs, and E were significantly lower in girdled vines than in control vines at the onset of withholding irrigation, which was 2 days after girdling. A in control vines decreased faster than A in girdled vines, attaining the value of girdled vines at 4 days after withholding irrigation (Figure 5AGo). Between 5 and 6 days of withholding irrigation, A in control vines became lower than A in girdled vines. There were no differences in A in either treatment after adding water. gs was similar to A (Figure 5BGo). For E, the difference between treatments disappeared at 2 and 5 days after withholding irrigation (Figure 5CGo). At 6 days after withholding irrigation, E nearly stopped in control vines but not in girdled vines. The patterns of diurnal changes of A, gs, and E were not different at 4 days after withholding irrigation, except that E at 9:00 hr was significantly higher in control vines than in girdled vines (Figure 6Go).


Figure 5
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  		Figure 5 Effects of withholding irrigation and reirrigation treatment on leaf net CO2 assimilation rate (A), stomatal conductance (B), and transpiration rate (C) in girdled and control vines. Arrows indicate irrigation; *, **, and NS indicate significance at p < 0.05, p < 0.01, and not significant, respectively. Girdling was implemented two days before irrigation was withheld.

 

Figure 6
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  		Figure 6 Effects of withholding irrigation and reirrigation treatment on the diurnal changes of the leaf net CO2 assimilation rate (A), stomatal conductance (B), and transpiration rate (C) at 4 days after treatment in girdled and control vines. * and NS indicate significance at p < 0.01 and not significant, respectively.

 
Stem diameter during the first 3 days did not decrease in either treatment and no fluctuation of daytime contraction and nighttime expansion in the stem was observed because it was cloudy (Figure 7Go). Stem diameter began to decrease within 4 days of withholding irrigation. Clear fluctuation patterns appeared within 4 days of withholding irrigation; diameters started to shrink early in the morning and then reached the minimum value just after noon. Daytime contraction and nighttime swelling of the stem were smaller in girdled than in control vines at between 4 and 6 days of withholding irrigation (Figure 8Go). Daytime contractions were largest at 6 days after withholding irrigation in both treatments. At that time, stem diameter shrank by 2.5 mm in girdled vines and by 3.6 mm in control vines. Stem diameter recovered rapidly in all vines just after irrigation at 6 days. Stem swelling was smaller in girdled vines than in control vines throughout the treatment. Fluctuation patterns in stem diameter after irrigation were similar in both treatments.


Figure 7
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  		Figure 7 Effects of withholding irrigation and reirrigation treatment on changes in the stem diameter in girdled and control vines. Arrows indicate irrigation.

 

Figure 8
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  		Figure 8 Effects of withholding irrigation on the stem diameter in girdled and control vines at 5 days after treatment.

 
There were no significant differences in shoot length (girdling: 100 cm; control: 101 cm), total leaf area (girdling: 5,893 cm2; control: 5,802 cm2), and yield (girdling: 1,136 g; control: 1,137 g) between the two treatments in the subsequent growing season.


   Discussion
Top
 Abstract
 Materials and Methods
 Results
 Discussion
Conclusions
 Literature Cited
 
Girdling has been known to decrease A with stomatal closure (Williams et al. 2000). In this study, gs and E at the beginning of withholding irrigation (2 days after girdling) were lower in girdled vines than in control vines. Girdling has been shown to increase abscisic acid (ABA) concentration in leaves (Düring 1978). The effect of ABA on decreasing the stomatal aperture is well known (Keller 2005). The mechanism of the reduction in A after girdling is assumed to be a reduction in gs induced by an increase of ABA in the leaves (Roper and Williams 1989, Williams et al. 2000). In the present experiment, the reduction of photosynthetic parameters may have occurred immediately after girdling; however, as that was not determined, the additional time (2 days) could contribute to the potential for an increase in ABA.

Feedback inhibition of A is another possible reason for stomatal closure; reduced phloem transport by girdling and subsequent accumulation of photosynthates in the chloroplasts may have reduced A. However, it has been reported that increased carbohydrate concentration in leaves by girdling is too low to induce feedback inhibition (Roper and Williams 1989, Williams et al. 2000).

Stomatal closure is also one of the first responses to soil drying, and a parallel decline in A and gs under progressive water stress has been reported (El-Ansary and Okamoto 2007). A depletion of A and gs with decreasing {Psi}PD has been found (Escalona et al. 1999). In our study, A and gs decreased at 4 and 5 days after withholding irrigation in control and girdled vines, respectively (Figure 5Go), while {Psi}PD values were –0.48 in control vines and –0.44 MPa in girdled vines (Figure 3Go). {Psi}PD has been reported to decrease to –0.4 MPa in nonirrigated vineyards (Escalona et al. 1999). Therefore, the control and girdled vines began to experience water-deficit stress on day 4 and day 5 of withholding irrigation. {Psi}PD reached –0.67 MPa at 6 days after withholding irrigation in girdled vines and was –1.13 MPa in control vines. Although gs in control vines was higher than in girdled vines after 3 days of withholding irrigation, E did not differ between treatments at this time. When the vapor pressure deficit (VPD) was low, E became low despite the high gs (Bunce 1996). The low VPD of 1.37 kPa at 3 days after withholding irrigation, which was due to rainy weather, was considered to decrease E despite the high gs.

Because gs and E were low, water consumption was assumed to be lower in girdled vines during the initial phase of the treatment. This assumption is supported by the findings of Williams and Ayars (2005), who measured water use by mature grapevines with a weighing lysimeter and showed that water use by girdled vines sometimes decreased when the girdle was open. The higher soil water potential in girdled vines than in control vines also indicated lower water consumption in girdled vines (Figure 2Go). Hence, we concluded that decreased water use by girdled vines reduced the depletion of water in their containers, which caused high {Psi}PD in girdled vines. Moreover, girdling and short-term severe water-deficit stress did not cause more leaf wilt or decrease shoot growth the following season.

A, gs, and E at 4 days after withholding irrigation showed almost the same pattern of diurnal change in girdled and control vines (Figure 6Go). The gas exchange parameters at 13:00 hr in control vines decreased to those of girdled vines at 4 days after withholding irrigation (Figure 5Go). Therefore, the differences in diurnal pattern in A and gs between girdled and control vines were assumed to have diminished because of water-deficit stress in control vines. The same pattern in diurnal changes observed in A and gs at the time when nearly identical values were achieved in both treatments suggests that the reduction in the parameters of leaf-gas exchange as a result of girdling and water-deficit stress may follow a similar physiological mechanism.

The initial responses to girdling and withholding irrigation were different, thus perhaps indicating the involvement of other mechanisms. In the present experiment, daytime stem contraction was smaller in girdled than in control vines under water-deficit stress (Figure 7Go). Stem shrinkage occurs almost exclusively in tissues external to the xylem (Kozlowski 1972). As for shrinkage in the stem, xylem shrinkage was minor, while shrinkage in the external tissues to the cambium was substantial (Brough et al. 1986). Biophysical measurements indicate that extensible tissues (i.e., immature xylem, phloem, and cortical cells) behave as a single osmotic entity separated from the matured xylem by a membrane system that shrinks or swells depending on the water status of the stem (Génard et al. 2001). In this study, low stem contraction also indicated that more water was available in girdled vines because of less consumption of water in the soil.

Although A was reduced in girdled vines, girdling enhances stem growth above the girdle because the increased carbon supply acts as a structural material and as a source of metabolic energy (Daudet et al. 2005). This is due to the elimination of the root system as a sink for carbohydrates in girdled vines (Williams et al. 2000). However, in this study, because mature stems were used, there was no apparent enhancement of stem growth in either treatment, and changes in stem diameter were mainly subject to the water status of vines.

The results of the present study indicate that the combination of girdling and short-term severe water-deficit stress did not result in leaf wilt or decrease shoot growth of the vines in the subsequent growing season. High crop load is another factor that debilitates vine growth in girdled vines (Winkler 1962). In general, optimal leaf area per fruit weight is 7 to 14 cm2 g–1 (Howell 2001). In the present study, the crop load was 5.0 cm2 g–1 in girdled vines. Despite the high crop load in girdled vines, the vines did not weaken.

In this study, container-grown vines were used, and the study was conducted in a vinyl greenhouse to clarify the effect of severe water-deficit stress. Therefore, the results are limited and further investigation of field-grown girdled vines that would undergo much slower water-deficit stress is needed.


   Conclusions
Top
 Abstract
 Materials and Methods
 Results
 Discussion
 Conclusions
Literature Cited
 
There are few reports concerning the combined effects of water stress and girdling on vine growth. In this study, physiological parameters associated with the water status of vines were measured to determine the responses of girdled vines to strong water-deficit stress. Specific attention was given to changes in stem diameter, which were measured at 10-min intervals following a micromorphometric technique with a micro-displacement detector of the shrinkage type. As a result, changes in stem diameter, leaf gas exchange parameters, soil water potential, and predawn leaf water potential indicated that decreased water use in girdled vines due to low gs and E reduced the depletion of water in the containers. In addition, short-term water-deficit stress in girdled vines when irrigation was withheld did not induce leaf wilt and depletion of shoot growth in the subsequent season any differently than in nongirdled vines.

Manuscript submitted March 2008; revised June 2008, August 2008

Accepted for publication September 2008


   Literature Cited
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 Abstract
 Materials and Methods
 Results
 Discussion
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 Literature Cited
 
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