Scientia Horticulturae 126 (2010) 3741

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Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti

Parthenocarpic fruit production in loquat (Eriobotrya japonica Lindl.) by using

gibberellic acid
C. Mesejo, C. Reig, A. Martnez-Fuentes, M. Agust
Instituto Agroforestal Mediterrneo, Universidad Politcnica de Valencia, Camino de Vera s/n, 46022 Valencia, Spain

a r t i c l e

i n f o

Article history:
Received 3 March 2010
Received in revised form 1 June 2010
Accepted 10 June 2010
Keywords:
Algerie
Ringing
Seedless
Thinning

a b s t r a c t
This study evaluates the effect of gibberellic acid (GA3 ) in inducing parthenocarpy in Algerie loquat,
as well as the optimum treatment conditions and associated techniques, hand thinning and ringing, to
produce seedless fruit with high enough quality for fresh consumption. GA3 applied in the course of the
phenological growth stages 504508 of the BBCH-scale produced seedless fruits, with the magnitude of
the response depending on the concentration applied and number of treatments. Percentage of panicles
bearing seedless fruitlets signicantly increased with increasing GA3 concentrations up to 100 mg l1 and
signicantly and positively correlated with the number of treatments applied. Trees treated three times
with 100 mg l1 developed more than 90% of panicles bearing almost 7 seedless fruits per panicle, which
were smaller in size, drier and slightly acid but similar in TSS concentration and skin colour than seeded
fruits from untreated trees. Fruit thinning to 3 fruits per panicle did not increase seedless fruit size, but
ringing performed at the onset of cell enlargement stage, growth stage 702 of the BBCH-scale, signicantly
increased fruit size by 1215%, depending on the year. Trees treated three times with 100 mg l1 of GA3
and ringed produced 26 kg, on average, of seeded fruit of suitable commercial quality.
2010 Elsevier B.V. All rights reserved.

1. Introduction
Loquat (Eriobotrya japonica Lindl.) belongs to the Rosaceae family, subfamily Maloideae. It is a subtropical evergreen fruit tree
species native to Southern China that, under Mediterranean climatic conditions, owers in autumn and early winter, sets in winter
and ripens in spring. By 2006 world loquat crop area exceeded
130 103 ha and fruit production was about 550 103 t, China
(450 103 t) and Spain (36 103 t) being the main producers (Lin,
2007). At present, Spain is the most important exporting country for fresh consumption and, thus, producing high quality fruit
is of prime importance. Algerie loquat leads the production with
1.1 103 ha which yield 18 103 t.
The ovary of loquat fruit contains 5 carpels with 2 ovules
each. Under open pollination conditions the fruit bears among 15
seeds. Seeds are relatively large and occupy 2030% of fruit volume (Sadamatsu et al., 2004), making the fruit somewhat difcult
for fresh consumption. Producing seedless loquat is of commercial
importance and represents a valuable horticultural objective.
Guo et al. (2007) have selected natural triploids from open populations of various cultivars. Some of these clones have vegetative

Corresponding author at: Instituto Agroforestal Mediterrneo, Universidad

Politcnica de Valencia, Campus de Vera s/n, 46022 Valencia, Spain.
Tel.: +34 963879330; fax: +34 963877331.
E-mail address: magusti@prv.upv.es (M. Agust).
0304-4238/$ see front matter 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.scienta.2010.06.009

parthenocarpy, whereas others have stimulative parthenocarpy

and require pollinizers as fruit set inducers. Production of these
triploid cultivars is still a distant goal and although they could have
a profound effect on the loquat industry in the future, fruit quality
for fresh consumption is far removed from that of current varieties
selected according to consumer taste. Therefore, parthenocarpic
fruit production from growing cultivars may be a good approach
to improve fruit quality.
In loquat, parthenocarpy is induced by means of gibberellic acid
(GA3 ). In short, concentrations between 25 and 1000 mg l1 applied
before full bloom result in parthenocarpic fruits, the proportion
depending on the concentration applied and on the cultivar (Kumar,
1976; Goubran and El-Zeftawi, 1986; Fan, 1989; Takagi et al., 1994;
Yahata et al., 2006). However, some cultivars, such as Fusahikari,
do not develop parthenocarpic fruits even when applying GA3 concentrations as high as 500 or 1000 mg l1 (Sadamatsu et al., 2004).
Therefore, horticultural factors regulating effectiveness of GA3 to
produce seedless loquat fruit remain unclear.
On the other hand, seedless fruit is generally smaller and more
elongated than seeded fruit, and differences in total soluble sugars, acidity and fruit colour at harvest have been reported among
cultivars (Goubran and El-Zeftawi, 1986; Takagi et al., 1994). The
application of 1-(2-Cl-4-pyridyl)-3-phenylurea (CPPU) during the
cell division stage increased nal fruit size of parthenocarpic loquat
fruit (Zhang et al., 1998; Matsui et al., 2004); however, CPPU is not
registered at present in the EU for use as a fruit enhancer in loquat.
In Spain, farmers thin fruits and ring branches to increase loquat

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C. Mesejo et al. / Scientia Horticulturae 126 (2010) 3741

fruit size; thinning reduces competition among developing fruit,

ringing increases fruit growth rate (Agust et al., 2000, 2005). The
effect of these techniques on increasing seedless loquat fruit size
remains unknown.
The objectives of this study were to evaluate the effectiveness
of GA3 in inducing parthenocarpy in Algerie loquat (1) determining the optimum date of treatment and concentration applied and
(2) establishing associated techniques, such us hand thinning and
ringing, to produce fruit of enough quality for commercialization.

(Facchini S.R.L., Alfonsine, Italy) with a 1.5 mm diameter at cylinder probe. Total soluble solids (TSS) concentration of juice ( Brix)
was measured using a digital refractometer (Atago, Tokyo, Japan)
and total acidity was determined by titration with 0.1N NaOH, using
phenolphthalein as the indicator. Fruit colour was established by
determining the a and b Hunter coordinates; three measurements
were made per fruit around the equator using a Minolta Chroma
Meter CR-300 (Minolta Camera Co. Ltd., Osaka, Japan).
2.5. Statistical analysis

2. Materials and methods

2.1. Plant material
Experiments were carried out in commercial orchards of adult
trees (20-year-old) of Algerie loquat trees (E. japonica Lindl.),
grafted onto loquat seedlings, and located in Callosa dEn Sarri,
Spain (38 39 N; 00 06 W; 150 m a.s.l.). Trees were pruned to the
vase shape, planted 4 m 3 m apart, grown on loamy-clay soil, with
drip irrigation. The experiments were carried out in three consecutive developmental periods (2006/072008/09).
2.2. Treatments
To determine the optimum concentration of GA3 inducing parthenocarpy, 0, 50, 100 and 150 mg l1 were applied
(2006/072007/08) at the phenological growth stage 504 of the
BBCH-scale (Martnez-Calvo et al., 1999). To determine the appropriate number of treatments, GA3 100 mg l1 was applied twice
(stages 504 and 506 of the BBCH-scale) or three times (stages 504,
506 and 508) (2007/082008/09). The loquat BBCH-scale consists
of three numbers (i.e. 504) the rst one expressing the major stage,
i.e. 5 refers to inorescence development, the second one expressing the mesostadium (03) referring to the generative growth (0)
or the vegetative growth (13), and the third one expressing the
secondary stage within the course of the specic major stage, i.e. 4
means inorescence at about 40% of nal size.
To establish associated techniques to improve fruit size, at the
onset of the cell enlargement stage (phenological fruit growth
stage 702 of the BBCH-scale) (2007/082008/09) trees were hand
thinned to 3 fruits per panicle, only keeping seedless fruits in
treated trees, and/or principal branches ringed as described by
Agust et al. (2005). Untreated and not ringed trees were used as
control. Only trees treated three times were used to study fruit
development and to determine commercial fruit characteristics at
harvest.

A randomized complete block design with single tree plots

and ve replications was used in all the experiments. Data were
analyzed by analysis of variance, and comparison of means carried out using Duncans multiple range test. Percentages were
analyzed after arc-sin transformation of the data. The effect on
frequency distribution of diameters at harvest was evaluated by
KolmogorovSmirnov test. The experimental data were analyzed
with Statgraphics 4.1 software (Statistical Graphics Corp., MD,
USA).
3. Results
GA3 proved to be effective in inducing parthenocarpy in Algerie
loquat trees. This effect was consistent in the 3 years of the experiment, while the magnitude of the response depended on GA3
concentration and on the number of treatments applied. The percentage of panicles bearing seedless fruitlets signicantly increased
with increasing GA3 concentrations up to 100 mg l1 (Fig. 1) and
signicantly correlated with the number of treatments applied
(y = 30.9x + 1.8; R2 = 0.997; P 0.01). On trees treated three times
more than 90% of the panicles bore seedless fruits, signicantly differing from trees treated twice (64%), once (34%), and untreated
trees (Table 1). The total number of fruitlets set per panicle, seedless plus seeded fruitlets, remained unchanged by treatments (7
fruitlets per panicle) (Table 1). On the other hand, the number of
seedless and seeded fruitlets set per panicle was signicantly modied by treatments, the former increasing and the latter decreasing
with increasing number of GA3 treatments (Table 1). In trees treated
once 4.4 seedless fruitlets per panicle were set, whereas 6.6 seedless
fruitlets per panicle were set in trees treated three times. Control
trees did not set seedless fruitlets (Table 1). Conversely, control

2.3. Fruit set, fruit development and yield

When seeded and seedless fruits were clearly distinguishable,
the total number of panicles per tree bearing seedless fruits and
number of seedless fruits per panicle were recorded. From owering to harvest, fruit development was followed by periodically
measuring fresh (FW) and dry (DW) weight of 10 fruits per tree
randomly selected around the canopy at a height of 1.52.0 m. The
fruit was harvested when they reached appropriate commercial
standards (MAPA, 1990). The yield and the number of fruits per
tree were recorded at harvest. Fruit size was evaluated at harvest
date by measuring the equatorial diameter and weight of 50 fruits
per tree randomly selected from among harvested fruits.
2.4. Sampling and fruit characteristic analysis
At harvest, 10 fruit samples from harvested fruits were collected
from each replication to determine commercial fruit characteristics. Fruit rmness was assessed using a fruit pressure tester FT-011

Fig. 1. The inuence of gibberellic acid (GA3 ) concentration applied on the percentage of Algerie loquat panicles bearing seedless fruitlets. Treatment was applied at
growth stage 504 of the BBCH-scale. Data for growing season 2006/07. Values are
the average of 10 panicles per tree and 5 trees. Standard errors are given as vertical
bars. Different letters indicate signicant differences (P 0.05).

C. Mesejo et al. / Scientia Horticulturae 126 (2010) 3741

39

Table 1
Effects of number of gibberellic acid treatments on seedless and seeded Algerie loquat fruit set. Gibberellic acid concentration applied 100 mg l1 . Data for growing season
2007/08. Dates of treatment given as BBCH growth stage. Date of evaluation was 702 BBCH fruit growth stage.
BBCH-scale growth stage at treatment

Panicles bearing seedless fruita (%)

Control
504
504 + 506
504 + 506 + 508

0.0a
34.9 5.1b
64.2 2.7c
93.1 2.6d

Fruit setb (fruits panicle1 )

Seedless

Seeded

Total

0.0a
4.4 0.3b
nd
6.6 0.3c

6.9 0.3c
2.6 0.3b
nd
0.6 0.1a

6.9 0.3
6.9 0.4
nd
7.2 0.3

Means followed by different letters in the same column differ signicantly (P 0.05); nd: not determined.
a
Values are the average of 5 trees.
b
Values are the average of 10 panicles per tree and 5 trees.

Fig. 2. The time-course of seedless Algerie loquat fruit weight as affected by 100 mg l1 of gibberellic acid compared to seeded loquat fruit from untreated trees. Gibberellic
acid applied three times (504, 506 and 508 BBCH-scale). Panicles were hand thinned and branches ringed at the growth stage 702. Data for growing season 2008/09. Values
are the average of 10 fruits per tree and 5 trees. Standard errors are given as vertical bars. Different letters indicate signicant differences (P 0.05); ns: not signicant.

trees only set seeded fruitlets, 6.9 fruitlets per panicle, whereas
trees treated once or three times set 2.6 and 0.6 seeded fruitlets
per panicle, respectively (Table 1).
Fresh and dry weight of seeded fruits was signicantly higher
than those of seedless fruits, in the course of fruit development
(702709 BBCH; Fig. 2). Differences in fresh and dry weight of pulp
between seedless and seeded fruits became signicant (P 0.05)
at growth stage 704 and 705 BBCH-scale, respectively, and reached
their highest value at growth stage 709 BBCH-scale (2.33- and 1.38fold higher for seeded fruit) (Fig. 2).
The effects of fruit thinning and ringing on seedless fruit development are shown in Table 2 for two growing seasons. Fruit
thinning did not signicantly improve the seedless fruit weight
but ringing signicantly increased it by 12% on average (Table 2).
Both fruit thinning and ringing signicantly increased seeded fruit
weight by 78% on average, respectively.
During fruit growth and ripening pulp acidity was signicantly
higher in seedless fruit than in seeded fruit (Fig. 3) whereas TSS

Table 2
Seedless Algerie loquat fruit weight (g) at harvest as affected by 100 mg l1 of gibberellic acid, fruit thinning and ringing. Trees were treated with gibberellic acid
three times, at the growth stages 504, 506 and 508 of the BBCH-scale. Thinning and
ringing were performed at the growth stage 702 of the BBCH-scale. Data for two
growing seasons. Values are the average of 10 fruits per tree and 5 trees.
2007/08

2008/09

Ringing

Ringing

Fruit thinning

18.7Aa
+
18.8Aa
Average
18.8a

Average

Average

22.0Ab
21.4Ab
21.7b

18.1A
17.9A

19.9Aa
20.8Aa
20.4a

22.3Ab
23.0Ab
22.7b

21.2A
21.9A

Means followed by different letters in the same line (lower case) or column (upper
case) differ signicantly (P 0.05).

Fig. 3. The time-course of seedless Algerie loquat fruit TSS and total acidity as
affected by 100 mg l1 of gibberellic acid compared to seeded loquat fruit from
untreated trees. Gibberellic acid applied three times (504, 506 and 508 BBCH-scale).
Panicles were hand thinned and branches ringed at the growth stage 702 of the
BBCH-scale. Data for growing season 2007/08. Values are the average of 10 fruits
per tree and 5 trees. *Signicant at P 0.05.

concentration signicantly differed between seedless and seeded

fruit during fruit growth stage (703801 BBCH-scale) but not during
fruit ripening (805809 BBCH-scale; Fig. 3). At harvest, seedless
fruit was smaller and more elongated and had lower water content
but higher acidity than seeded fruit (Table 3). No differences in TSS
concentration and skin colour were found between seedless and
seeded fruit (Table 3). Although seedless fruit showed signicantly
higher resistance to puncturing, their senescence appeared faster
than seeded fruit senescence.
In our experiments, since thinning did not improve seedless
fruit size (Table 2), the best results were those from trees treated
three times and ringed. Yield of these trees averaged 26 kg tree1
of seedless fruits, whereas untreated and ringed trees averaged
60 kg tree1 of seeded fruit. At harvest, the frequency distribution
of seedless fruit according to diameter, shifted toward lower size

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C. Mesejo et al. / Scientia Horticulturae 126 (2010) 3741

Table 3
Fruit characteristics at harvest of seedless fruits from Algerie loquat trees treated
three times with 100 mg l1 of gibberellic acid, compared to seeded loquat fruits
from untreated trees. Trees were treated at the growth stages 504, 506 and 508 of
the BBCH-scale. All trees were ringed at the growth stage 702 of the BBCH-scale.
Data for growing season 2008/09. Values are the average of 10 fruits per tree and 5
trees.
Seedless

Seeded

(mm)
Height (mm)
/heigth
Fruit colour (a/b)

28.7 0.3a
49.1 1.2a
0.60 0.02a
0.30 0.01

44.8 1.0b
52.5 1.2b
0.85 0.04b
0.30 0.01

Flesh
Fresh weight (g)
Dry weight (g)
Water content (%)
TSS ( Brix)
Acidity (%)
Resistance (kg cm2 )

22.5 1.0a
3.1 0.3a
86.5 0.4a
14.9 0.6
1.2 0.1b
11.7 0.8b

54.5 2.7b
4.1 0.3b
92.2 0.4b
14.5 0.7
0.9 0.1a
7.6 0.6a

Means followed by different letters in the same line differ signicantly (P 0.05).

Fig. 4. Effect of 100 mg l1 of gibberellic acid on the frequency distribution of seedless Algerie loquat fruit diameter compared to untreated seeded loquat fruit at
harvest. Gibberellic acid applied three times (504, 506 and 508 BBCH-scale). All
trees were ringed at the growth stage 702 of the BBCH-scale. Data for growing
season 2008/09. Values are the average of 10 fruits per tree and 5 trees. Standard errors are given as vertical bars. Distributions differ signicantly according
to KolmogorovSmirnov test (KS = 3.66; DN = 0.725; P 0.05).

categories (Fig. 4). Fruits ranged 2531 mm amounted to about

70% of the yield from trees treated three times with GA3 and ringed,
whereas untreated and ringed trees achieved only 7%, and fruits
ranged 3238 mm amounted to 20% and 32%, respectively.
4. Discussion
Pre-blossom application of 100 mg l1 of GA3 proved to be
effective in inducing parthenocarpy in Algerie loquat fruit. Our
results are in accordance with Kumar (1976) using Pale Yellow and Thames Pride loquat, and differ from others in which
2001000 mg l1 of GA3 were necessary for parthenocarpic fruit
set in Tanaka, Mogi and Nawasaki-Wase loquat (Kumar, 1976;
Goubran and El-Zeftawi, 1986; Fan, 1989; Takagi et al., 1994; Yahata
et al., 2006). Sadamatsu et al. (2004) reported that seedless fruit
set was not accomplished using 500 or 1000 mg l1 of GA3 in
Fusahikari loquat.
In our experiments, the magnitude of the response positively
correlated with the number of treatments applied. This is because
owering in loquat lasts for more than 4 weeks, different phenological oral stages overlapping and, consequently, leading to owers
receptive to GA3 for about one month, making repeated treatment

necessary. The mechanism of GA3 inducing parthenocarpy has been

studied in grape (Kimura et al., 1996), sweet cherry (Stsser and
Anvari, 1982) and citrus (Mesejo et al., 2008), but how it works in
loquat remains unknown. It depends critically on the oral growth
stage, being preferably accomplished prior to anthesis, suggesting
that GA3 may operate on ovule development prior to fertilization,
in a similar manner to that observed in Citrus (Mesejo et al., 2008).
In fact, in our experiments seedless fruit set was preferably accomplished in the panicles from the most immature shoots and in the
most immature owers of each panicle (data not shown).
On the other hand, the larger the seedless fruit number the
lower the seeded fruit number per panicle. This phenomenon
modies competition among developing fruits with regard to the
controls, and explains why our best results were those from trees
treated three times. Successive GA3 treatments increase fruit sink
strength and consequently the more treatments, up to three, the
more parthenocarpic fruits harvested per tree. Competition for
carbohydrates among developing fruitlets has been studied in
loquat (Agust et al., 2000) and is the reason for thinning. Furthermore, ringing signicantly increases fruit size in loquat by means
of translocation and accumulation of carbohydrates in the fruit
(Wallerstein et al., 1974; Goldschmidt et al., 1985) reinforcing the
role of carbohydrates content in determining fruit size. Gibberellins
produced by seeds are responsible for sink strength of pome fruit
(Zhang et al., 2007), thus allowing taking up of carbohydrates and,
as a result, increasing water uptake, i.e., seeded fruit is heavier
than seedless fruit, both in fresh and dry weight. Therefore, lower
fruit weight and higher total acid concentrations of seedless fruit,
compared to seeded fruit, are due to seed absence.
It is important to note that thinning did not increase average
seedless fruit size. It is also due to seed absence, which deprives
fruit of sink strength and, thus, incapacitates it for competing for
carbohydrates, all fruits reaching almost the same size. However,
ringing increased signicantly seedless fruit size. In Citrus, Rivas et
al. (2007) reported that ringing increases quantum yield efciency
of photosystem II in young leaves of fruiting shoots, altering their
carbohydrate concentrations and photosynthetic performance and,
thus enhancing fruit size of Loretina mandarin, a parthenocarpic
Clementine cultivar. According to these results, ringing enhances
Clementine fruit size regardless to seed, in a similar manner to
our results in seedless loquat fruits which develop in leafy fruiting
shoots.
In conclusion, 100 mg l1 of GA3 applied three times between
504 and 508 growth stages of the BBCH phenological growth scale
induce parthenocarpy in Algerie loquat trees. Seedless fruits are
smaller in size than seeded fruits, but ringing performed at the
onset of the cell enlargement stage produce fruits of enough quality
for fresh consumption.

Acknowledgements
This research was supported by grants from Universidad Politcnica de Valencia (Spain), project PAID-06-08 no. 3026. Thanks are
due to Mr. E. Soler and Mr. V. Martnez for technical assistance, to
Cooperativa Agrcola de Callosa dEn Sarri S.C.V. (Alicante, Spain)
for the orchard facilities provided, and to the Centro de LenguasUPV for proof reading the English.

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