Benzyladenine plus Gibberellins (GA4+7) Increase Fruit Size and Yield in Greenhouse-Grown Hot Pepper (Capsicum annuum L.)
A two year study was conducted to establish effects
of a commercial preparation (BA plus GA4+7) called accel, on
growth and yield of hot pepper. Five concentrations: 0, 5, 10, 15, 20
and 25 mg L-1 of the preparation were used. Concentrations
of 10, 15, 20 and 25 mg L-1 significantly (p<0.001) increased
yield in a two year pooled data. Accel treatment did not significantly
affect number of fruits, number of branches and plant height. The increase
in yield due to accel treatment was associated with significant increase
in fruit fresh weight (p<0.001) and length (p<0.001). Therefore
the increase in yield was attributed to increase in fruit size. Accel
application has the potential to be used as a management practice in greenhouse
production of hot pepper.
Hot pepper (Capsicum annuum L.) (syn.chili pepper) is among
the species of the genus Capsicum, domesticated for vegetable and industrial
(oleoresin and capsaicin) purposes. Available literature indicates that
hot pepper is widely cultivated throughout the world, mainly in Asia,
South and North America and parts of Africa (Contreras-Padilla and Yahia,
1998; Kumar et al., 2006). The fresh fruits are used in pickles
and sauces, while the red dried and powdered fruits are used as hot spices.
Plant Growth Regulators (PGR) are used extensively in horticulture to
enhance plant growth and improve yield by increasing fruit number, fruit
set and size. Improvement in vegetative growth and yield attributes may
enhance crop productivity. Productivity in horticultural systems is often
dependent on manipulation of physiological activities of the crop by chemicals
means (Yeshitela et al., 2004) and this is modulated by the interaction
of the PGR with plant developmental processes. According to Gianfagna
(1987), plant growth regulators can modify development by interfering
with biosynthesis, metabolism or translocation of endogenous hormones,
or may supplement endogenous hormones when their levels are reduced.
Developmental processes reported to be influenced by PGR application
include; induced flowering in Arabidopsis thaliana by gibberellins
(Richards et al., 2001), increased flowering in Aquilea
species by GA4+7 (Gianfagna and Merrit, 1988), increased fruit
fresh weight in cucumber by BA plus GA4+7 in cucumber
(Batlang et al., 2006) and GA4+7 (Yang et al.,
1992) and stimulation of individual branch growth by BA plus GA4+7
in apple (Voltz et al., 1994). Increase in flowering may
lead to more fruits and coupled with increase in fruit size and yield
in fruit crops. Fruit size may be determined by increased cell layer formation
and cell division and increased sink strength of the fruit (Dyer et
al., 1990). Fruit size increase due to cytokinins and gibberellins
applications to apples, cucumber and grapes was suggested to be
caused by increased cell division and elongation and cell wall extensibility
(Emongor and Murr, 2001; Yu et al., 2001).
In the semi-arid areas such as Botswana, there is low rainfall and low
fertility and as such there is a need to practice controlled environment
horticulture. Intensive production of hot pepper requires the utilization
of high yielding cultivars or improved management of the locally available
genetic materials. This situation validated the evaluation of accel as
a management avenue for intensive production of hot pepper.
MATERIALS AND METHODS
Plant culture: Seeds of a commonly grown landrace were obtained
from a farmer in Bobonong (21° 58' S, 28° 27' E), in the Central
district of Botswana. The experiments were conducted at Botswana College
of Agriculture greenhouses in Gaborone (24° 33' S, 25° 54' E).
The landrace has not been genetically characterized and developed into
a cultivar and will be referred to as Bob-02; where Bob is Bobonong and
02 is collection year of 2002. Fruits were collected and air dried on
a laboratory bench and seeds were removed, washed under a running tap
water and germinated in vermiculite in a growth chamber at 25 °C in
the dark. After five days uniformly emerged seedlings were transplanted
in seedling trays filled with compost soil and grown for four weeks in
The plants were then transplanted into 8 L polyethene bags filled with
soil mixture (4 river sand: 2 top soil and 1 compost) on per volume basis.
When plants showed first sings of nutrient deficiency they were irrigated
with two hydroponics solutions containing 110 g L-1 hydrogrow
(85 g N kg-1, 45 g P kg-1, 240 g K kg-1,
30 g Mg kg-1, 60 g S kg-1, 1.63 g Fe kg-1,
0.4 g Mn kg-1, 0.2 g Zn kg-1, 0.03 g Cu kg-1,
0.05 g Mo kg-1, 0.5 g B kg-1 and 10 g L-1
CaNO3 (26% N) fertilizers. Each fertilizer solution was applied
to field capacity at weekly intervals.
Accel treatment: Accel (Abbot Laboratories, North Chicago, IL)
is a plant growth regulator that contains the cytokinin 6-benzyladenine
(1.8%) and gibberellins (GA4+7) (0.18%). Accel treatment was
applied 56 days after transplanting to the vegetatively growing plants.
Aqueous solutions were prepared at 0 (control), 5, 10, 15, 20 and 25 mg
L-1 of active ingredient (a.i) and applied to run-off with
a hand-held sprayer. The control treatments were sprayed with distilled
water. Five plants were sprayed for each treatment. Plant became dormant
in winter (between May and July) 2003, during which minimal irrigation
was applied to carry them through the dormancy period. There was re-growth
in the summer beginning August, when hydro-grow and CaNO3 fertilizers
regime was resumed, accel application was re-applied at the same rate
as in the first year.
Data collection and analysis: Data such as plant height, branch
number, marketable fruit number, fruit fresh weight and fruit length were
collected between January and May 2003. In this case yield refers to marketable
fruits. Fruits were harvested approximately every fourteen days from fruit
set until plants ceased to bear fruits. Fruit fresh weight at each time
of harvest was divided by the number of fruits to obtain fresh weight
per fruit. Data for each parameter was divided by five plants to obtain
per plant basis for each level of PGR treatment. In the second year (2003
to 2004) the same data was collected. However, for comparison purposes
between years, only data collected between January and May was used for
analysis, to minimize seasonal confounding effects.
The experiment was laid out in a randomized complete block design replicated
four times. Means were calculated across the five plants in each replication
and were subjected to analysis of variance using the general linear models
(Proc GLM) of the Statistical Analysis System program (SAS Institute,
Carey, NC). Treatment means were separated using the Least Significant
Difference (LSD) at p = 0.05. Appropriate linear regression models were
used to determine the yield response to fruit number, fruit length and
fruit fresh weight per fruit.
RESULTS AND DISCUSSION
It was observed that hot pepper plants did not grow in the winter,
even though temperatures were kept at 27 ± 2 °C throughout the
experimental period. This observation can be ascribed to the fact that
there was low irradiance in winter and high irradiance is required for
optimal growth as reported in sweet pepper (Ulvskov et al., 1992).
Accel treatment significantly increased yield, fruit fresh weight and
fruit length, while plant height, number of branches and fruits were not
affected (Table 1). In order to determine how accel might
affect chili pepper due to plant age, repeated application of accel was
performed on the same plants used in the first year and results were compared.
Vegetative growth (plant height and number of branches), fruit number
and yield increased with plant age and the increase of the respective
parameters was at least 60%. This was to be expected, since in the second
year plants had grown more than in the first year and fruiting branches
were increased. However, there were no interactions between accel treatments
and plant age for all parameters that were assessed (Table
1). The effect of accel treatment and plant age were additive, indicating
that the PGR treatment did not affect plant behaviour differently in both
years. As a result data was pooled across two years before mean separations.
The PGR treatment at 10, 15, 20 and 25 mg L-1 concentrations
increased yield, with 20 and 25 mg L-1 giving highest yield
and there were no significant increase in yield between the controls and
5 mg L-1 (Table 2). It was also observed that
the increase in yield
The ANOVA (F-values)
of the effects of accel treatment and plant age and their interaction
on plant growth, yield and yield components
|***Significant at p<0.001. FW: Fruit
Effects of accel
concentration on yield and its respective components (fruit number,
fruit length, fruit fresh weight) in a two year pooled data.
|***significant at p<0.001. NS: Non-Significant
at p = 0.05. FW: Fruit Fresh Weight, Means within columns followed
by the same letter(s) are not significantly different at p = 0.05,
of the relationship
between yield and its respective components (fruit number, fruit
length, fruit fresh weight) in a two year pooled data
in parentheses are standard errors
due to PGR treatment was generally accompanied with increase in fruit
fresh weight per fruit and fruit length, while fruit number was not increased
(Table 2, Fig. 1). This is confirmed
by regression analyses testing the influence of fruit number and size
on yield as a result of PGR treatment (Table 3). It can
therefore be reasoned that PGR treatment increased fruit size, which ultimately
increased yield on per plant basis. This can be attributed to both cytokinins
and gibberellins in their role in cell division and elongation, respectively.
The two hormones, which are the components of accel, are known to promote
cell division and elongation, respectively (Salisbury and Ross, 1996;
Wismer, 1994). During tomato fruit development, there is endogenous gibberellins
accumulation, which coincides with activation of cell division and expansion
(Gillaspy et al., 1993). The increase in cell volume due to expansion
may contribute to the final size of the fruits as observed in this study.
Recently accel was applied to cucumber plants and this caused increased
yield through increase in fruit number and size (Batlang et al.,
2006). In other fruits such as pear (Stern and Flaishman, 2003) and apple
(Wismer et al., 1995; Yuan and Greene, 2000), the increase in fruit
size was accompanied by thinning effect due to benzyladenine. While this
established the effect of thinning on bigger fruit development, in our
case with hot pepper, the increase in fruit size is probably not due to
thinning as there was no significant reduction in fruit number due to
accel treatment (Table 1, 2). This further
implicates the involvement of cytokinins and gibberellins in accel on
fruit development in hot pepper.
Effects of accel
concentration on the increase of yield and its component
Foliar application of both benzyladenine and GA3 to citrus
was reported to enhance assimilate (sucrose) export by the foliage to
the developing fruit and GA3 was especially active in promoting
fruit sink activity (Mauk et al., 1986). Exogenous application
of benzyladenine and GA3 increased expansion of excised leaf
discs in sweet pepper (Nielsen and Ulvskov, 1992; Ulvskov et al.,
1992). According to Yuan and Greene (2000) application of benzyladenine
to apples increased levels of zeatin riboside in fruits, suggesting that
it might promote cell division through the activity of zeatin riboside,
which is one of the dominant endogenous cytokinins found in pepper (Ulvskov
et al., 1992).
Although no attempts have been made to measure cell division in this
experiment, it is hypothesized that accel influenced yield in hot pepper
directly through processes that increased cell division and improved fruit
sink capacity. The next experiments will address direct involvement of
accel on fruit growth and development in hot pepper.
The author thanks Botswana College of Agriculture, Department of
Crop Science and production for financial and material support and Ms.
Biganani Chalegwa for her invaluable technical assistance. Thanks are
due to Motshwari Obopile for his critical review of the Manuscript.
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