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Intra-canopy variability of microclimate conditions, plant water relations, and

fruit quality in kiwifruit vines under deficit irrigation and two covering methods

Article in Acta Horticulturae · June 2021

DOI: 10.17660/ActaHortic.2021.1314.5

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1 Intra-canopy variability of microclimate conditions, plant water

2 relations, and fruit quality in kiwifruit vines under deficit irrigation
3 and two covering methods
4
5
6 A. Calderón-Orellana1,a, D.I. Silva1; and R.M. Bastías1
7
8 1Departamento de Producción Vegetal, Facultad de Agronomía, Universidad de Concepción,
9 Chile.
10
11
12 Abstract
13 Chile is one of the main producers of kiwifruit (Actinidia spp) in the world.
14 However, the predicted reduction in water availability and the increase in both air
15 temperature and evaporative demand in many producing areas may jeopardize the
16 Chilean kiwifruit industry. Protected cultivation and deficit irrigation have been
17 progressively gained in popularity among kiwifruit growers as new methods to
18 improve water use efficiency and fruit uniformity. Unfortunately, few studies have
19 addressed the extent of the effects of both cultural practices on microclimate
20 conditions that determine water requirements and fruit variability in commercial
21 kiwifruit orchards. Two irrigation treatments were applied in mature kiwifruit vines
22 (Actinidia deliciosa A. Chev. var. Hayward) under open-field conditions and covered by
23 plastic films. Results showed that plastic-covered vines experienced a lower
24 transmission of blue light (400 - 500 nm), but a higher transmission of red light (600 -
25 700 nm) and far-red light (700-800 nm), which may be related to a lower sensitivity of
26 leaf stomata from plastic-covered vines to moderate water stress (leaf water potential
27 values between -1.3 and -1.0 MPa). Moreover, vines covered by plastic films exhibited
28 20-30% higher stomatal conductance than those under open field conditions, but
29 similar values of chlorophyll fluorescence and temperature of fruits and leaves,
30 regardless of the irrigation strategy. Despite the increase in the proportion of diffusive
31 radiation and lower variability in photosynthetically active radiation, plants beneath
32 plastic covers exhibited lower uniformity in stomata opening, which seemed to be
33 associated with greater variability in soluble solids concentration at harvest in fruits
34 from well-irrigated vines. These results highlighted the importance of spectral light
35 determinations to understand responses of plants to water stress, which may be
36 determinant in a global warming scenario.
37
38
39 Keywords: kiwifruit, protected cultivation, water stress, fruit quality, light quality.
40
41
42 INTRODUCTION
43 Chile is the fourth largest exporter of kiwifruit (Actinidia spp) in the world (FAOSTAT,
44 2018). However, the progressive decline in water availability for irrigation has jeopardized
45 the Chilean kiwifruit industry, as many growers may suffer from severe water stress at the

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46 end of the growing season. This is particularly serious for kiwifruit, a fruit crop that is widely
47 known for its high water requirements (~10.000 m3/ha) (Holzapfel et al., 2000) and low
48 tolerance to water stress (Miller et al., 1998), especially under Mediterranean climate
49 conditions.
50 During the last couple of years, the use of high density polyethylene films have gained in
51 popularity in kiwifruit orchards, as growers need protection from rain, wind, hail, and
52 drought. Protected cultivation has been reported to ameliorate the effects of water scarcity on
53 plant water status in grapevine (Novello and de Palma, 2006). However, there is a lack of
54 information about the impact of this practice on kiwifruit water relations and reproductive
55 growth and development. Furthermore, there is scientific evidence that suggests that the use
56 of protected cultivation can improve uniformity of fruit quality attributes, as microclimate
57 conditions are less variable in covered orchards (Novello and de Palma, 2006).
58 Fruit uniformity is a key quality factor in fresh fruit production, but it has been rarely
59 measured or evaluated in horticultural sciences (Calderón-Orellana et al, 2019). The aim of
60 this study was to determine the effect of a transparent plastic cover on uniformity of
61 microclimate conditions, water relations, and fruit quality in a commercial kiwifruit orchard.
62
63
64 MATERIALS AND METHODS
65 A field study was carried out in a fourteen-year old kiwifruit orchard (Actinidia deliciosa
66 A. Chev. cv. Hayward), grafted onto seedling rootstock in San Nicolás (Ñuble Region, Chile)
67 (36°32'45.8"S, 72°11'18.4"W) for two consecutive growing seasons (2018 to 2019). Kiwifruit
68 vines were planted at 4.0 m x 3.0 m, trained as a pergola system, and drip irrigated (~12,000
69 m3/ha). Soils at the study site are clayey loam and have been locally described as members of
70 the “Talquipen” series. The climate is Mediterranean and the average maximum and minimum
71 temperatures are 32°C (January) and 3°C (July), respectively.
72
73 At the beginning of January (60 DAFF), three groups of 30 vines were covered with a
74 high density, transparent poliethylene film (PC), while other three groups of plants remained
75 under open field conditions (OF). At the onset of the dry matter accumulation period (90
76 DAFF), irrigation was ceased in 15 vines within each covering method (RDI), while the
77 remaining plants were irrigated to satisfy 100%ETc (WET). The experiment was laid out as a
78 split-plot arregement in a randomized complete block design, where covering methods were

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79 replicated three times and represented the mainplots, and irrigation treatments the subplots.
80 Midday values of leaf water potential were weekly measured from 12:00 to 15:00 h in two leaf
81 samples under cloud-free conditions, using a pressure chamber (Model 615, PMS Instruments,
82 Corvallis, USA), according to the protocol described by McCutchan and Shackel (1992).
83 Stomatal conductance was estimated with a steady-state porometer (SC-1, Decagon devices,
84 Pullman, USA) in the same plants where leaf water potential was determined. Intercepted
85 photosynthetically active radiation (PAR) at midday was determined at harvest time with a
86 portable ceptometer (LP-80, Decagon Devices). The ceptometer was placed immediately
87 below the fruiting zone (about 1.8 m above the soil level) and parallel to the vine row. Four
88 PAR measurements were taken per plant, while an outside PAR reading was made at a height
89 of 50 cm above the plant canopy. The spectral transmission of light was determined under
90 laboratory conditions using a dual-channel spectrum-photometer system, which was
91 configured to measure the spectral light transmission in the photosynthetic (400-700 nm),
92 red/far-red (600-800 nm), and infra-red (950-1700 nm) ranges. Measurements were carried
93 out on a 2 x 5 cm sample of material, which was carefully placed in a light integrating sphere
94 (IC2 cube type, StellarNet INC., Tampa FL, USA). A halogen lamp (model SL1, StellarNet INC.,
95 Tampa FL, USA) was used as a light source, which coverede the light range between 350 and
96 2500 nm. Maturity evaluations were weekly made from the onset of fruit ripening to harvest
97 time in randomly taken samples of twenty fruits per experimental unit. Harvest time was
98 defined when the concentration of soluble solids and the percentage of dry matter were 6.5
99 Brix and 16.0%, respectively. Soluble solids concentration (SSC) was evaluated with a hand-
100 held digital refractometer (HI 96801, Hanna Instruments, Rhode Island, USA). At harvest, 360
101 fruits per experimental unit were randomly collected from three contiguous plants and
102 immediately taken to the laboratory for quality evaluations. The data were subjected to
103 analysis of variance (ANOVA). When significant differences were detected, the LSD mean
104 separation test was used on means. The analysis of variability included descripitive statistical
105 analsys, comparing ranges, variances, and box-and-whisker plots of residuals and means. All
106 statistical procedures were carried out using the statistics package SAS/STAT 9.2 (SAS
107 Institute, North Carolina USA).
108
109 RESULTS AND DISCUSSION
110 The plastic film transmited less radiation in the blue light range (74% between 400 and
111 500 nm) than in the red (94% between 600 and 700 nm) and far-red (94% between 700 and

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112 800 nm) ranges (Table 1). The transmission of PAR (400-700 nm) averaged 86%, while the
113 transmission of IR radiation (950-1700 nm) was 95%.
114
Table 1. Percentage of light transmission of the high density polyethylene film used as
covering material for various light ranges under laboratory conditions.
Light range (nm) Percentage of transmission (%)
400-500 (blue) 74%
600-700 (red) 94%
700-800 (far-red) 95%
400-700 (PAR) 86%
950-1700 (IR) 95%
115
116 Irrigation treatments changed plant water status during the growing season each year
117 (Table 2). Before the application of irrigation treatments, midday leaf water potential ranged
118 between -0.7 and -0.5 MPa in WET and RDI plants, indicating well-irrigated conditions for
119 kiwifruit (Chartzoulakis et al., 1993). Once irrigation treatments were applied, midday leaf
120 water potential was reduced in RDI plants to levels representative of severe water stress
121 (between -1.5 and -1.3 MPa). However, severe levels of water stress during stage III occurred
122 twice as fast in uncovered vines (data not shown).
123
Table 2. Average minimum values of midday leaf water potential reached during the first and
third stage of fruit growth in mature ‘Hayward’ kiwifruit plants under open field (OF) and
protected cultivation (PC) conditions and subjected to regulated deficit irrigation (RDI) and
commercial irrigation (WET) in 2018 and 2019.
Fruit growth stage PC-WET OF-WET PC-RDI OF-RDI
(MPa) (MPa) (MPa) (MPa)
Stage I -0.71 a -0.72 a -0.73 a -0.72 a
Stage III -0.81 a -0.83 a -1.46 b -1.43 b
124 Different letters indicates significant differences at 95% confidence level (LSD). N=8.
125
126 There was a weaker response of stomata to water stress in leaves from covered plants
127 (Figure 1). This lower sensitivity of stomata to severe water stress (midday leaf water
128 potential ~-1.5 MPa) may be related to a reduction in transmited PAR, as previous works have

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129 shown higher values of stomatal conductance in water-stressed kiwifruit plants under shaded
130 conditions (Montanaro et al., 2009).
131

132
133 Figure 1. Relationship between midpoints of midday leaf water potential and stomatal
134 conductance of mature ‘Hayward’ kiwifruit plants under open field (OF) and
135 protected cultivation (PC) conditions. Each symbol represents the mean of 12
136 plants.
137
138 There was a higher percentage of transmitted PAR at the fuit zone in RDI plants under open
139 field conditions (~10%), whereas covering showed no influence on transmitted PAR in well-
140 irrigated kiwifruit plants (Figure 2). This may indicate that RDI induced earlier leaf
141 senescence than commercial irrigation practices only in uncovered vines, which means that
142 protected cultivation was effective in reducing the negative effects of water stress on plant
143 physiology. Regardless of the irrigation strategy, plants under open field conditions showed
144 less variability in transmitted PAR at the fruit zone, as indiceted by the length differences in
145 error bars between covering treatments.
146

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147
148

149
150 Figure 2. Seasonal average of transmitted PAR of mature ‘Hayward’ kiwifruit plants under
151 open field (OF) and protected cultivation (PC) conditions and subjected to regulated
152 deficit irrigation (RDI) and commercial irrigation (WET). Each symbol represents the
153 mean of 4 plants. Different letters indicates significant differences at 95% confidence
154 level (LSD). Error bars represent ± 1 S.E.
155
156 Stomatal conductance of covered plants tended to be slightly higher, regardless of leaf
157 position (Figure 3). This is surprising, as the lower transmission of blue light to covered plants
158 may have limited the generation of the inside-negative electric potential nececesary to
159 accumulate K+ in the guard cells and open stomata (Shimazaki et al., 2007). Furthrermore, the
160 use of the plastic canopy seems to have caused a greater impact on stomatal conductance of
161 apical leaves. Box-and-whisker plots were larger in covered plants, which may indicate less
162 uniform stomatal conductance along the shoot. The effect of protected cultivation on the
163 efficiency of photosystem II and fruit temperature was not observed, as fv/fm and IR fruit
164 temperature were similar between cover treatments (data not shown).
165

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166
167 Figure 3. Box-and-whisker plots for stomatal conductance of leaves located at different shoot
168 positions in mature ‘Hayward’ kiwifruit plants under open field (OF) and protected
169 cultivation (PC) conditions at harvest in 2019 (April, 6th). Each boxplot represents the
170 mean of 12 leaves.
171
172 The range of stomatal conductance residuals was wider for covered vines (Figure 4A),
173 which may indicate a more heterogeneus gas exchange capacity in covered kiwifruit vines. On
174 the other hand, the range of fv/fm residuals were similar in covered and uncovered plants
175 (Figure 4B). These results suggest that any potential effect of protected cultivation on the
176 homogeneity of photosynthesis, and hence carbon supply to fruits, is due to greater variability
177 in carbon dioxide diffusion rather than electron transport. These results are unexpected, as
178 one goals of protected cultivation is to improve uniformity of plant physiological parameters.
179

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180
181 Figure 4. Boxplots for the residuals of (A) stomatal conductance in 2018 and 2019 and (B)
182 Fv/Fm in 2019 of leaves from mature ‘Hayward’ kiwifruit plants under open field (OF)
183 and protected cultivation (PC) conditions. Each boxplot represents the mean of 44
184 leaves.
185
186 Protected cultivation reduced Brix variability among fruits in RDI plants, as range
187 (Figure 5A) and variance values (Figure 5B) were higher in fruits from water-stressed vines
188 under open field conditions. It seems the plastic canopy reduces the loss of fruit uniformity
189 that has been previously reported to occur in plants subjected to severe water stress in areas
190 with Mediterranean climate (Calderon-Orellana et al., 2014). Results showed that RDI plants
191 under open field conditions intercepted lower PAR than WET plants (~10%) (Figure 2),
192 probably due to earlier leaf senescence. Earlier defoliation in plants under severe water stress
193 might have either increased dehydration or accelerated ripening rates in a small proportion of
194 fruits, leading to uneven distribution of Brix at harvest. This coincides with a study in
195 winegrape (Calderon-Orellana et al., 2014), in which severely water-stressed Cabernet
196 Sauvignon vines exhibited more heterogeneous Brix and anthocyanin content in clusters due
197 to greater fruit dehydration. On the other hand, the use of plastic covering in WET plants
198 showed no improvement in Brix uniformity. In fact, variance and range values for Brix were
199 slightly higher in fruits from covered plants. The lower uniformity of Brix at harvest found in
200 covered plants under commercial irrigation may be explained by the lower stomatal
201 conductance uniformity observed in leaves from covered vines (Figure 4A).

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202
203 Figure 5. Range and variance for Brix at harvest in fruits from mature ‘Hayward’ kiwifruit
204 plants under open field (OF) and protected cultivation (PC) conditions and subjected
205 to regulated deficit irrigation (RDI) and commercial irrigation (WET). Each bar
206 represents the mean of 200 fruits.
207

208 CONCLUSIONS
209 Protected cultivation has been widely used to protect fruit crops against biotic and
210 abiotic stress. In the present study, the use of a plastic film to cover kiwifruit plants reduced
211 microclimate variability. However, the impact of covering was not the same under different
212 irrigation strategies. While plastic films improved fruit uniformity under water stress, there
213 was no impact of protected cultivation in fruits from well-irrigated plants. In fact, stomatal
214 conductance and fruit maturity was more heterogeneous in well-irrigated covered plants.
215 These results suggest that changes in light transmittance may induce alterations in stomatal
216 response to water stress.

217
218 ACKNOWLEDGEMENTS
219 We gratefully acknowledge the funding provided by the “Comisión Nacional de Ciencia y
220 Tecnología - Conicyt” (Proyecto Fondecyt Iniciación 11160876). We are grateful to Carsol
221 Fruit and Michael Medina for their technical support and assistance.
222
223 Literature Cited
224

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225 Calderon-Orellana, A., Matthews, M. A., Drayton, W. M., and Shackel, K. A. (2014). Uniformity of ripeness and size in
226 cabernet sauvignon berries from vineyards with contrasting crop price. Am. J. Enol. Viticult. 65 (1), 81–88
227 https://doi.org/10.5344/ajev.2013.13084.
228
229 Calderon-Orellana, A., Bambach, N., Aburto, F., and Calderón, M. (2019). Water deficit synchronizes berry color
230 development in Crimson seedless table grapes. Am. J. Enol. Viticult. 70 (1), 60–67
231 https://doi.org/10.5344/ajev.2018.17070.
232
233 Chartzoulakis, K., Noitsakis, B., and Therios, I. (1993). Photosynthesis, plant growth and dry matter distribution in
234 kiwifruit as influenced by water deficits. Irrig. Sci. 14 (1), 1–5 https://doi.org/10.1007/BF00194999.
235
236 Food and Agriculture Organization of the United Nations. (2018). FAOSTAT statistical database. [Rome] :FAO
237 Holzapfel, E. A., Merino, R., Mariño, M. A., and Matta, R. (2000). Water production functions in kiwi. Irrig. Sci. 19 (2), 73–79
238 https://doi.org/10.1007/s002710050003.
239
240 McCutchan, H., and Shackel, K. A. (2019). Stem-water Potential as a Sensitive Indicator of Water Stress in Prune Trees
241 (Prunus domestica L. cv. French). J. Am. Soc. Hortic. Sci. 117 (4), 607–611 https://doi.org/10.21273/jashs.117.4.607
242
243 Miller, S. A., Smith, G. S., Boldingh, H. L., and Johansson, A. (1998). Effects of Water Stress on Fruit Quality Attributes of
244 Kiwifruit. Ann. Bot. 81 (1), 73–81 https://doi.org/10.1006/anbo.1997.0537.
245
246 Montanaro, G., Dichio, B., and Xiloyannis, C. (2007). Response of photosynthetic machinery of field-grown kiwifruit under
247 Mediterranean conditions during drought and re-watering. Photosynthetica 45 (4), 533–540
248 https://doi.org/10.1007/s11099-007-0091-4.
249
250 Novello, V., and de Palma, L. (2006). Growing grapes under cover. Acta Horticulturae 785, 353-362
251 https://doi.org/10.17660/ActaHortic.2008.785.44
252
253 Shimazaki, K., Doi, M., Assmann, S. M., and Kinoshita, T. (2007). Light Regulation of Stomatal Movement. Annu. Rev. Plant
254 Biol. 58 (1), 219–247 https://doi.org/10.1146/annurev.arplant.57.032905.105434.
255
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