Aqsa Ghaffar

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EFFECT OF FOLIAR BORON APPLICATION ON GROWTH, REPRODUCTION, AND OIL

QUALITY OF OLIVE TREES CONDUCTED UNDER A HIGH DENSITY PLANTING


SYSTEM
Abstract

Boron (B) foliar treatments (300 mg L−1 as Solubor DF) were applied at two different dates in
2006 and 2007, prior to flowering and just after fruit set, on olive (Olea europaea L.) trees with
no visual symptoms of B deficiency. Leaf B level increased after the first application as compared
to control (−B). After July treatment, leaf B levels in −B and +B treated trees increased when
compared to the first sampling date. Foliar B application did not significantly affect vegetative
growth in either year. During the first year of study (considered as an “on year”), B application
had no significant effect on several phenological characteristics including fruit set, yield oil
contents and oil quality. In the second year (“off year”), B sprays improved blooming rate, which
increased from 20% in −B to 30% in + B treated trees, and olive yield, which increased by 27% in
response to B.
INTRODUCTION
Mineral fertilization has been reported to influence olive yield and oil quality (Fernández-Escobar
et al., Citation2006). Boron (B) is a micronutrient and adequate B nutrition is critical not only for
high yields but also for high quality of crops (Ateyyeh and Shatat, Citation2006). It is well known
that B deficiency results in low pollen viability, poor pollen germination, and reduced pollen tube
growth (Nyomora et al., Citation1997). Furthermore, low B has been found to restrict the
proportion of flowers setting fruit, to reduce fruitlet retention and development, and to decrease
yield in several tree crop species (Hanson and Breen, Citation1985; Shrestha et al., Citation1987).
Several authors report that B applications increase flower set and yield in fruit and nut crops
(Baron, Citation1973, Chaplin et al., Citation1977; Hanson, Citation1991; Nyomora et al.,
Citation1997). Boron fertilization has exhibited a significant effect on fruit set of almond (Prunis
dulcis) (Nyomora et al., Citation1999), ‘Italian’ prune (Prunus domestica) (Hanson et al.,
Citation1985), sour cherry (Prunus cerasus L.) (Hanson, Citation1991), apple (Malus domestica
Borkh.) (Wojcik et al., Citation1999), highbush blueberry (Vaccinium corymbosum L.) (Hanson,
Citation2000). Conversely, other authors found no crop response to B fertilization (Ferran and
Taous, Citation1997; Silva et al., Citation2003). Reports on the effect of foliar B application on
olive trees are also contradictory. Increase of fruit set in olive after foliar B application was
reported by Perica et al. (Citation2001b), yet others reported that foliar B application did not
affect fruit set of olive trees (Talaie and Taheri, Citation2001; Ateyyeh and Shatat, Citation2006).
Most of these experiments were conducted on trees without obvious vegetative signs of B
deficiency, suggesting that there is a specific requirement for B in the reproductive processes of
fruit tree. Foliar B fertilization is actually widely used in Tunisian and intensive olive orchards
worldwide without knowledge of its effectiveness. Little is known about reproductive
requirements of olive for B, effects of B on flowering and fruit set and effects of foliar B
applications on vegetative growth, yield, oil content, and quality. Furthermore, the common
symptoms of B deficiency described as abnormal fruit development, or “monkey face,” leaf tip
yellowing, shoot die-back, fruit drop and bark abnormalities were not observed in Tunisian olive
orchards (Perica et al., Citation2001b).

The purpose of the present work was to study the effect of foliar applied B fertilization on
vegetative, phenology, yield, and oil quality.

MATERIALS AND METHODS


A two-year experiment was conducted in a commercial olive orchard (Arbequina variety) in the
west-central region of Tunisia (Meknassi, 34°36′ N and 9°35′E). Climate is semi arid. Annual
rainfall averages less than 200 mm, most of which occurs during the cold months. Four-year old
trees were set at 2 × 4 m spacing in a sandy soil with low cation exchange capacity (3.7 meq g−1)
and organic matter content (0.54%). Water and fertilization were supplied by a drip irrigation
system. This olive orchard received 3350 and 2700 m3 of water, 195 and 119 kg of ammonium
nitrate, 85 and 55 kg phosphoric acid and 100 and 127 kg of potassium nitrate during 2006 and
2007, respectively.

Treatments were applied to a block of ten adjacent trees without B deficiency symptoms and
each block was replicated six times. A total of sixty trees per treatment were used [B treated (+B)
and control (−B) trees]. Treatments were arranged in a randomized complete block design. Trees
were selected with comparable morphological features such as canopy volume and trunk
diameter to guarantee a homogeneous sample for −B and +B treatments.

The + B treatment consisted on a foliar application of B, 300 mg L−1 using Solubor DF (78%
Na2B8O13 · 4H2O and 20% Na2B4O7 · 5H2O, US Borax Company, Denver CO, USA) at pre-
flowering stage (March) and at fruit set stage (May). The untreated trees received the same
volume of water without B (−B).

Agronomic characteristics were evaluated on observations from ten trees from each block. At
least five shoots were sampled on each tree. Shoot length was measured monthly from March
to October for 2006 and from March to July for 2007.
The development of buds into inflorescence or vegetative shoot was recorded. The percentage
of staminate flower was determined by observation on 200 inflorescences sampled inside and all
around the trees. Percentage fruit set was also determined 45 days after full bloom by counting
the number of fruitlets on previously counted inflorescence on each marked shoot. The average
number of flowers in each inflorescence was also determined using 200 inflorescences from each
treatment to calculate the fruit set percentage.

Inflorescences of treated and control trees were sampled at the white-button stage. Anthers
were removed and dehisced overnight at room temperature and pollen collected. Pollen viability
was estimated using acetocarmin staining technique (Martoja and Martoja_Pierson,
Citation1967). Pollen grains from (+B) and (−B) flowers were cultured on a sucrose-agar medium
(0.8% agar, 10% sucrose) and 100 mg L−1 boric acid at pH 5 (Mehri et al., Citation2006). Cultures
were held at 25°C for 24 h. Counted pollen grains were at least 500 per Petri dish. Pollen grains
were considered germinated when the length of its pollinic tube was equal to or exceeded its
diameter (Stanley and Linskens, Citation1974). Pollen germination was recorded after 24 h of
incubation, on pollen grain chosen at random from various locations of the Petri dish.
Germination rate was determined using five replicates of approximately 100 pollen grains each
one (Pinney and Polito, Citation1990).

Olive yield (kg), canopy volume (m3) and yield efficiency (kg m-3 of canopy) were determined
separately for each tree at harvesting (Lombard et al., Citation1988). Canopy volume (m3) was
calculated by considering the tree as a parallelepiped since trees were trained to a central leader.
Consequently the formula (1) was used

where h, L and e are the canopy height, width and thickness of the tree, respectively.

Three samples of 3 kg of olives from each block were used to determine fruit characteristics.
Average fruit weight was measured and samples were then dried in a forced-air oven at 100°C
for 48 hours. Dried samples were weighed to determine the moisture content and then the oil
percentage was measured using NMR analyzer (Oxford 4000, Mumbai, India).

Oil was extracted using laboratory oil mill Abencor (MC2 Ingeniería y Sistemas S.L, Seville, Spain)
consisting of a hammer mill, a thermobeater, and a paste centrifuge (Martinez et al.,
Citation1975). Oil fatty acid composition was determined by gas chromatography of the
corresponding methyl esters prepared as described by the EU official method (European Union
Commission, Citation1991). Percentages of the main fatty acids, palmitic (C16:0), oleic (C18:1),
and linoleic (C18:2) were determined. An automated Metrohmn Rancimat 679 apparatus
(Metrohm, Herisau, Switzerland) operating at 120°C was used to determine the oxidative
stability. An oil sample (5.0 g) was weighed into reaction vessel, the conductimetry cells were
filled to 90 mL, and the air was injected through the heated oil at 20 l h−1. Determinations were
made in duplicate and results were expressed as induction time.

Leaf B concentration was measured immediately following the first B application (in March) and
in July of 2006 by sampling 100 leaves from the mid part of shoots all around the tree from each
block. Leaves were washed in a 1% liquid detergent, rinsed three times with deionised water,
and dried at 65° for 72 hours. Dry leaves were ground and sent to Laboratory (AGQ:
Agroalimentaria y Medio Ambiente: ENAC ENSAYOS N° 350: LE 1322/LE 1323; Seville, Spain) for
B analysis.

For each of these parameters analysis of variance (one-way Anova) were performed to test
significant differences between (+B) treated and control (−B) trees using SPSS Base 8.0 software
(SPSS, Chicago, IL, USA).

RESULTS AND DISCUSSION

Effect of Foliar B Application on Leaf B Level


Boron sprays increased foliar B concentration immediately after the first application (Figure 1),
but by the second sampling date in mid July leaf B concentration was similar in +B and −B trees.
Adequate leaf B concentration in olive trees ranges from 19 to 150 mg kg−1 (Freeman et al.,
Citation1994). Our results confirm that trees used in this experiment are not B deficient. Indeed
foliar B concentration of untreated (−B) trees was higher than the currently recognized adequate
level of 19 mg kg−1 (Freeman et al., Citation1994). Previous studies have reported that foliar B
sprays increased leaf and inflorescence B concentrations in olive trees (Perica et al.,
Citation2001b; Ateyyeh and Shatat, Citation2006). Several authors indicated also that B
concentration in inflorescence is higher than in leaves, which suggests that B needs may be higher
for reproductive processes than for vegetative growth (Dell and Huang, Citation1997; Blevins and
Lukzszewski, Citation1998; Perica et al., Citation2001b). Delgado et al. (Citation1994) reported
that B is mobilized from young leaves during the anthesis to satisfy B needs of flowers and fruits.
This may explain the increase of leaf B by July, after fruit set, in (−B) trees, while it was not
significant in +B trees (Figure 1). Indeed, when B demands decrease, B levels in leaves increase.
Similar increases of leaf B concentration in summer were reported by Fernández-Escobar et al.
(Citation1999) and Delgado et al. (Citation1994). However, our results suggest that B application
reduced B mobilization needs.

FIGURE 1 Olive tree leaf B concentrations at two sampling dates in 2006 without (−B) and with
(+B) foliar B applications. NS is not significant at P < 0.05 and * and** is significant at P < 0.05 and
<0.01, respectively.

FIGURE 1 Olive tree leaf B concentrations at two sampling dates in 2006 without (−B) and with
(+B) foliar B applications. NS is not significant at P < 0.05 and * and** is significant at P < 0.05 and
<0.01, respectively.
Display full size

Effect of Foliar B Application on Vegetative Growth


Boron sprays had no effect on vegetative growth of Arbequina olive tree in 2006 while in 2007
vegetative growth was higher in −B than +B trees (Figures 2A and 2B). Shoot length increases
from March to September in 2006 averaged 24.1 cm (0.15 cm day−1) while from March to July
in 2007 vegetative growth was about 22.4 (0.18 cm day−1) and 19.3 cm (0.16 cm day−1) in −B
and +B trees, respectively. Several authors reported that foliar B spray has no effect on vegetative
growth of olive trees (Perica et al., Citation2001b; Ateyyeh and Shatat, Citation2006).

FIGURE 2 Olive tree vegetative growth during growing season in A) 2006 and B) 2007 without
(−B) and with (+B) foliar B applications. NS indicates not statistically significant at P < 0.05
comparing −B and +B at each date within each year.

FIGURE 2 Olive tree vegetative growth during growing season in A) 2006 and B) 2007 without
(−B) and with (+B) foliar B applications. NS indicates not statistically significant at P < 0.05
comparing −B and +B at each date within each year.
Display full size
Effect of Foliar B Application on Flowering

Foliar B application (+B) did not significantly impact development of buds into vegetative, floral
and quiescent buds of Arbequina compared to untreated (−B) trees in 2006 (Figure 3A), but
significantly increased floral buds and decreased vegetative and quiescent buds on the shoots of
+ B trees in 2007 (Figure 3B). Indeed flower bud percentage was 30% in +B trees compared to
20% in −B trees (Figure 3B). Despite this increase of flower bud percentage for B application,
inflorescence length and staminate flower percentage were not affected by foliar B application
(Table 1). Several authors reported that B application improves flower development (Perica et
al., Citation2001b; Dell and Huang, Citation1997; Rawson, Citation1996). The physiological
processes leading to olive flower differentiation in March and bloom in April-May begin in the
preceding summer with flower induction followed by flower initiation in November (Fernández-
Escobar et al., Citation1992). The pre-flowering foliar B application had no effect on the
percentage of floral buds in the first year presumably because the treatment was made after
flower induction and initiation. However, continuous foliar B application lead to a significant
increase of percentage in flower bud in the second year, which points out the importance of
application date. Nyomora et al. (Citation1999) reported that the timing of foliar B application
significantly influenced almond response, where autumnal application were more effective than
pre-flowering one. This was explained by the fact that dormant buds are inefficient at absorbing
applied B (Hanson and Breen, Citation1985; Nyomora et al., Citation1999).

Effect of Foliar B Sprays on Pollen Viability and Germination


Pollen viability was not significantly affected by field applied B treatment (Table 1). Our results
were consistent with other studies carried out with other species (Nyomora et al., Citation1997,
2000). However, Robbertse et al. (Citation1990) reported that B application improved pollen
viability. Loss of viability in pollen has been related to B which is involved in maintaining plasma
membrane integrity (Goldbach et al., Citation1991; Shivanna and Heslop-Harrison, Citation1981).

Foliar-applied B resulted in significantly higher in vitro pollen germination compared to −B trees


(Table 1). Germination percentages ranged from 52% for pollen from untreated (−B) trees to 60%
in pollen collected from trees sprayed with B (+B). This is opposite to results of Perica et al.
(Citation2001b) and Ateyyeh and Shatat. (2006) obtained on olive. However, others reported that
foliar B application improved in vitro pollen germination in almond (Nyomora et al., Citation1997,
2000) and of lowbush blueberry (Chen et al., Citation1998).

Effect of Foliar B Application on Fruit Set


Results obtained here indicate that B application had no positive effect on fruit set of olive trees
(Figure 4). Instead, fruit set percentage was usually higher in untreated (−B) trees (6.3 and 2.6%)
as compared to treated ones (5.7 and 2.4%) during 2006 and 2007, respectively (Figure 4).
Ateyyeh and Shatat (Citation2006) and Talaie and Taheri (Citation2001) indicated that B
application had no effect on fruit set or yield when no symptoms of B deficiency were present on
trees but Perica et al. (Citation2001a) reported that B influence fruit set and yield positively
during years of low production (off year).
Effect of Foliar B Application on Yield
Boron application had no effect on average yield per tree in the first year of experimentation
compared to no application, but did increase average yield per tree in 2007 (Figure 5A). This
translates to an increase of production in B treated trees of about 1012 kg per ha. Our results
indicated that B sprays positively affect production only in the year of lower production (off year).
Yield per ha was 46% lower in 2007 compared to 2006. It has been reported that B sprays increase
yield of olive trees (Perica et al., Citation2001b), almonds (Nyomora et al., Citation1999), and
sweet cherry (Prunus avium L.) (Usenik and Stampor, Citation2007) during years of lower
production (off year) and even when trees have no symptoms of B deficiency. Our results indicate
that no relationship was found between improved yield and fruit set percentage, which was
unaffected by foliar B application. Nyomora et al. (Citation1999) indicated that an increase in set
should increase yield, however yields are not always increased as other unknown factors may
ultimately determine the percentage of fruit that remain on the tree at harvest.
Several authors reported that B is required for fruit retention (Nyomora et al., Citation2000;
Blevins et al., Citation1996; Brown et al., Citation1999). The causes of fruit drop such as hormone
imbalance in favour of abscission and resources limitation particularly of C and N (Nyomora et
al., Citation1999) can generally be related to B status (Bellalouli et al., Citation1994; Perica et al.,
Citation2001a).

Effect of Foliar B Sprays on Oil Content and Quality


Foliar B sprays did not affect the oil content on fresh and dry weight basis in either year (Table
2). Indeed, oil content on fresh weight basis averaged about 27% and 32.5% in 2006 and 2007,
respectively. On dry weight basis, oil content averaged 55% in 2006 and 51% in 2007. Fruit size
and pulp stone ratio also were not affected by foliar B treatment through years of
experimentation (Table 2).

Also fatty acid composition and oxidative stability were not affected by foliar B treatment (Table
3). Oleic, palmitic and Linoleic acid contents averaged 58.9, 18.9 and 16.7%, respectively. Oil
oxidative stability averaged 5.47 h (Table 3). Thus, unlike N, B had no effect on oil quality
(Fernández-Escobar et al., Citation2006).

In conclusion, the response of olive to foliar B application was observed only when fruit set was
low. No relationship was found between improved yield and fruit set percentage. The effect of
foliar B application on yield can be attributed to an increase in flowering development percentage
and better pollination and fertilization. Also B application might influence positively fruit
retention. No negative effect of B on oil quantity or quality was observed. Given the yield increase
observed during the off year of production and the low costs of B fertilizers, the practice of foliar
application of B should be recommended in years following high production (on year).

ACKNOWLEDGMENTS

This work was supported by grants from the Spanish AECID (A/8333/07) to Msallem Monji and
Anunciacion Abadia. The authors gratefully acknowledge Rio Tinto Minerals Company for
providing commercial product and for B analysis. Both Ajmi Larbi and Kamel Gargouri contributed
equally to this work.

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