Fruits, 2016, vol. 71(4), p.

209-214

c Cirad / EDP Sciences 2016
DOI: 10.1051/fruits/2016010
Available online at:
www.fruits-journal.org

Original article

Pineapple harvest index and fruit quality improvement

by application of gibberellin and cytokinin
Tri Suwandi1, , Kumala Dewi1 and Priyo Cahyono2
1
Faculty of Biology, Gadjah Mada University, Yogyakarta, Indonesia
2
Research and Development Department, Great Giant Pineapple company, Lintas Sumatera street, Km. 77 Terbanggi Besar,
Lampung, Indonesia

Received 23 November 2015 – Accepted 18 April 2016

Abstract – Introduction. Gibberellin application alone in Lampung, Indonesia, has not been successful in overcom-
ing the low harvest index of pineapple. Cytokinin is known to promote the allocation of assimilates to sink organs. The
effect of gibberellin and cytokinin on the harvest index and fruit quality of pineapple was evaluated in this study. Mate-
rials and methods. Three levels of gibberellin (0, 100 and 200 ppm) combined with three levels of cytokinin (0, 24 and
48 ppm) were applied in the 12th and 14th weeks after flowering induction treatment. The experiment was conducted
with the pineapple cv. Smooth Cayenne in a randomized complete block design with three replications. Harvest index
and fruit quality analyses were carried out in fruits harvested during the 23rd week after flowering induction treatment.
Results and discussion. Gibberellin alone (100 or 200 ppm) or combined with cytokinin (24 or 48 ppm) increased the
fruit weight, harvest index and fruit crown length – a positive effect on the quality of this planting material – and de-
layed fruit maturity (5 days) of ‘Smooth Cayenne’ pineapple. Conclusion. Gibberellin and/or cytokinin improved some
properties of the pineapple fruit quality. Treatment with 100 ppm of gibberellin combined with 24 ppm of cytokinin
was the best treatment for improvement of the pineapple harvest index and fruit quality.

Keywords: Indonesia / pineapple / Ananas comosus / plant growth regulators / crop management / fruit maturity

Résumé – Amélioration de l’indice de récolte et de la qualité du fruit de l’ananas par application de gibbérel-
lines et de cytokinines. Introduction. L’application de gibbérellines seules n’a pas réussi à contrecarrer la faiblesse
de l’indice de récolte de l’ananas à Lampung, en Indonésie. Les cytokinines sont connues pour favoriser la répartition
des assimilats qui alimentent organes. Les effets combinés des gibbérellines et des cytokinines sur le point de récolte
et sur la qualité du fruit de l’ananas ont été étudiés. Matériels et méthodes. Trois doses de gibbérellines (0, 100 et
200 ppm) combinées à trois doses de cytokinines (0, 24, et 48 ppm) ont été appliquées durant les semaines 12 et 14
après le traitement d’induction de la floraison. L’expérience a été menée sur le cv. Cayenne lisse en bloc complet ran-
domisé avec trois répétitions. Le calcul de l’indice de récolte et l’analyse qualitative des fruits ont été réalisés sur les
fruits récoltés au cours de la 23e semaine après traitement d’induction de la floraison. Résultats et discussion. Les
gibbérellines seules (100 ou 200 ppm) ou en combinaison avec des cytokinines (24 ou 48 ppm) ont augmenté le poids
des fruits, l’indice de récolte, la longueur de la couronne des fruits – un effet positif pour la qualité de ce matériel de
plantation – et ont retardé (de 5 jours) la maturité des fruits du ‘Smooth Cayenne’. Conclusion. Les gibbérellines et/ou
cytokinines améliorent certains critères de qualité du fruit de l’ananas. La combinaison de 100 ppm de gibbérellines et
de 24 ppm de cytokinines s’est montré le meilleur traitement pour l’amélioration de l’indice de récolte et la qualité des
fruits de l’ananas.

Mots clés : Indonésie / ananas / Ananas comosus / hormones végétales de croissance / gestion des cultures / maturité
des fruits
Abbreviations. GA : gibberellin ; CK : cytokinin ; ABA : abscisic acid ; PGR : plant growth regulator ; ppm : part per
million ; DAF : day after flowering induction treatment ; TSS : total soluble solids ; TA : titratable acidity.


Corresponding author: tri.suwandi@mail.ugm.ac.id
210 Tri Suwandi et al.: Fruits 71 (2016) 209–214

1 Introduction on developmental processes and signaling are exerted by GA

and CK [11,12]. Previous studies have shown that GA and CK
The pineapple fruit is technically called a sorosis – “a fu- have a synergistic relationship to increase the fruit weight of
sion of many fruits together to form one unit”. Each “eye” ‘Hass’ avocado [13] and ‘Golden Delicious’ and ‘Jonagold’
(fruitlet) is a complete fruit. Flowering starts at the bottom apples [14]. To our knowledge, there is no information on CK
of the sorosis and continues up as a spiral to the last eye. or the interaction between GA and CK in pineapple. Moreover,
When the formation of fruitlets stops, the growing point re- no studies have been conducted to evaluate the harvest index,
verts to a vegetative state and the top (crown) is formed [1]. and physical and biochemical quality of the fruit in response
There is probably no direct effect of the crown on the growth to GA and CK application in ‘Smooth Cayenne’ pineapple.
of the fruit and no relationship among the crown, fruit size and The effect of GA and CK on the harvest index, and fruit and
stem starch [2]. Fruit growth from blossoming inflorescence crown quality of ‘Smooth Cayenne’ pineapple is reported in
to mature fruit results in a twenty-fold increase in weight. The the present study.
enlargement of the calyx results from continued growth by cel-
lular division, in the stages up to flowering, and cell enlarge-
ment, in the later stages [3]. Fruit growth is more rapid around 2 Materials and methods
100 days after the flowering induction treatment when there is
a decrease in stem dry matter and probably in stem starch, or 2.1 Experimental design
both, indicating a mobilization of starch from the stem to help
meet the fruit carbon demand. Leaf starch is apparently also
mobilized to meet the increasing demand for the rapidly devel- The experiment was conducted at the research station of
oping fruit [4]. Good cultural care will ensure a large number GGP located in Terbangi Besar, Lampung, Indonesia, from
of fruitlets are formed, which will fill out to give a well-shaped August 2014 until February 2015. The experimental station
and high-yielding fruit. has the following geographic coordinates: 4◦ 49 15.5” S and
105◦ 15 27.4” E, with an average altitude of around 46 m.
In some pineapple plantations of the Great Giant Pineap-
The average annual rainfall is about 148.4 mm, average an-
ple (GGP) company, Indonesia, there has been a decrease in
nual temperature is 27.5 ◦ C, and average relative air humidity
productivity and fruit quality, resulting in small fruits or large
is 88.6%.
fruits with small-sized crowns or with a conical shape. This is
disadvantageous because the crown is one of the main pineap- Ground suckers of ‘Smooth Cayenne’ pineapple (about
ple planting materials. Therefore, the goal is always to ob- 40 cm in length) were planted in August 2013. A plantation of
tain the best fruit quality with vigorous crowns. An effort to 67,340 plants × ha−1 was maintained with a single-row plant-
overcome this problem is by applying plant growth regulators ing system (55 cm × 27 cm). Flower induction was performed
(PGRs). Gibberellin (GA) plays a role in increasing the source- at 12 months after planting (August 2014) by spraying with
sink relation or sink strength by activating enzymes involved 3 kg × ha−1 of ethylene, 25 kg × ha−1 of kaolin and 50 kg ×
in sugar metabolism [5,6]. The partitioning of assimilates from ha−1 of urea dissolved in 4,000 L × ha−1 of water.
source leaves is a key factor for fruit development, as any lim- The experiment was conducted in a randomized complete
itation of assimilate supply affects the final fruit size [7]. Gib- block design with three replications. The experimental plot
berellin is a PGR commonly used to increase fruit weight in (5.5 m × 2.7 m) consisted of at least 110 plants in 10 rows and
various types of plants. Gibberellin increases the fruit number there were border plants between plots. A factorial arrange-
ment of treatments that consisted of ProGibb r 40SG (GA3)
and size of grapes, delays maturity and reduces physiological
abnormalities in citrus peel, maintains the photosynthetic tis- levels of 0, 100 and 200 ppm active ingredient (a.i.) and Nov-
sue of lemon peel, and extends the postharvest shelf life of elgro Alpha (zeatin:kinetin, 3:1 v/v; CK) levels of 0, 24 and
banana [8]; it increases the fruit weight of ‘Comte de Paris’ 48 ppm a.i. (table I) were applied at 12 weeks after forcing
pineapple [9]; it delays the maturity of ‘MD-2’ pineapple for (November 28, 2014) and again at 14 weeks after forcing (De-
7–10 days, and increases its crown length [8]. Previous studies cember 11, 2014). The solution was sprayed until runoff using
conducted at GGP have shown that GA increased the pineap- approximately 10–15 mL of solution per fruit in each applica-
ple fruit size, but the internal fruit quality was poor due to such tion, which was done with a knapsack sprayer in the morning.
aspects as more translucency, bland taste and variable TSS/TA
ratio (personal communication, 2013). These results may have
been affected by inappropriate time of application and concen- 2.2 Analysis of pineapple harvest index
trations of GA. and fruit quality
It is known that plant hormones do not work by themselves
to stimulate or inhibit the growth, but they interact with other Five fruits at the mature green stage, reached at 156 DAF,
hormones or components. Besides GA, cytokinin (CK) is also were harvested per treatment plot for fruit quality analy-
a hormone that stimulates cell division and growth, controls sis. Meanwhile, the remaining fruits were harvested as they
cell division and cell differentiation, and triggers the nutrient reached the half-yellow stage when fruit were designated as
mobilization and allocation from the source to the sink or- mature. The data on days from induction to harvest for all fruits
gans [10]. The interactions between GA and CK include both within each treatment were averaged to assess the effect of the
negative and positive mutual regulation, depending on the tis- PGR treatments on days from induction to fruit maturity. The
sue and signaling context. Also, numerous antagonistic effects fresh weights of the fruit, crown and plant, without roots, were
 Tri Suwandi et al.: Fruits 71 (2016) 209–214 211

Table I. Effect of gibberellin and cytokinin on the fruit weight, crown length and harvest index of pineapple.
Treatments Fruit weight (g) Crown length (cm) Harvest index Days to maturity
GA0 CK0 1165.56b 22.29d 0.50c 161
GA0 CK1 1125.00b 23.78d 0.48d 161
GA0 CK2 1238.33ab 22.09d 0.53a−c 161
GA1 CK0 1414.00a 29.77ab 0.55ab 166
GA1 CK1 1459.78a 27.37bc 0.56a 166
GA1 CK2 1235.56ab 26.81c 0.54a−c 166
GA2 CK0 1235.83ab 29.09a−c 0.52c 166
GA2 CK1 1318.56ab 31.07a 0.55ab 166
GA2 CK2 1458.44a 28.89a−c 0.58a 166
1
Fruit treatments were 0 (GA0 ), 100 (GA1 ) or 200 (GA2 ) ppm of gibberellic acid and 0 (CK0 ), 24 (CK1 ) or 48 (CK2 ) ppm of cytokinin. All
fruits were harvested at the half-yellow stage.
2
Values are means for 45 fruits. Values within a column followed by the same letter are not significantly different (P < 0.05) according to
Duncan’s multiple range test.
3
Days to maturity (DTM) were the number of days from induction on 04-08-2014 to fruit maturity.

measured on 15 fruits from each treatment plot. The harvest in- of NaOH. The flask was boiled again on the condenser for
dex was calculated based on the ratio of fruit weight at harvest 30 min. The suspension was filtered through filter paper while
(without the crown) to the plant weight at harvest [15]. being washed with boiling distilled water until the washing
Five samples of fruits were sliced horizontally at the point water was no longer basic (tested with a PP indicator). The
of the largest diameter. Internal fruit maturity indices were residue was washed with boiling distilled water containing
evaluated using a subjective color chart varying from 0 to 6; 15 mL of alcohol 95%. The residue present in the filter pa-
with 0 being a completely white flesh and 6 a completely yel- per was dried at a temperature of 100–110 ◦ C for 1–3 h until it
low one (available at http://www.verita.cr/internal.jpg). Imma- reached constant mass, cooled in the desiccator, and weighed.
ture fruits have a white flesh color, while mature or ripe fruits The difference in weight of the filter paper and residue before
have a yellowish-white flesh. The flesh also becomes slightly and after drying was the percentage of fibers contained in the
translucent in appearance at maturity. Fruits are over-mature sample.
when more than half of the cross-sectional area of the fruit is The vitamin C content of the fruit flesh was determined
translucent [16]. by titration of the filtrate with 2.6-dichlorophenol indophenol
Fruit firmness was measured on triangle regions of fruit (2.6-D). The filtrate was put into a 250-mL Erlenmeyer flask
slices taken from the middle part using a penetrometer (Instron and distilled water was added up to the mark. Five mL of fil-
5542) and results were expressed as N. trate were put into a 100-mL Erlenmeyer flask and 5 mL of
The fruit shell, core, and the top and bottom 3 cm of flesh HPO3 -acetatic acid were added. The mixture was titrated with
were removed, then the flesh was taken for biochemical fruit a standard solution of 2.6-D until the color changed to pink. At
quality analysis. The flesh was cut into small pieces and the the end point of the titration, the solution volume was recorded.
juice was extracted with a small benchtop juice extractor. To
determine the total soluble solids (TSS) content, the filtrate For fruit potassium content determination, 20 mL of the
was measured by a hand refractometer. Titratable acidity (TA) juice were put into a 25-mL volumetric flask and 40 g L−1 CsCl
was measured using filtrate titration with 0.1 N of NaOH until solution was added until the content in the sample reached
it reached the titration end point (pH 8.3). about 0.1–0.4%. The solution was diluted 5 times. The blank
Water content was analyzed using the thermogravimetric solution was 0.1–0.4% of CsCl solution. Standard solution ab-
method. Samples were heated in an oven at a temperature of sorbance was measured using an atomic absorbance spectrom-
100–105 ◦ C for 3 h to reach constant mass. The difference in eter (AAS) at the wavelength of 769.9 nm. The standard curve
weight before and after drying was the amount of water con- was done afterwards. The absorbance of the sample solution
tained in the sample. was also measured by AAS.
Fruit fiber content was analyzed by alcohol extraction and The sugar content of the pineapple fruits was analyzed us-
the thermogravimetric method. Ten grams of the juice were ing high-performance liquid chromatography (HPLC). Fruc-
put in a boiling flask and 200 mL of boiled 0.255N H2 SO4 tose, glucose and sucrose standards were purchased from
were added. This apparatus was placed on the condenser and Merck. The sugar content in the sample was calculated by
heated for 30 min. Once completed, the suspension was fil- comparing the sample’s peak area with the standards at the
tered and the residue was washed with distilled water until the wavelength of 250 nm. The mobile phase used was acetoni-
washing water was no longer acidic (tested with litmus pa- trile and distilled water (80:20; v/v). The stationary phase used
per). The residue was transferred quantitatively from the fil- was a Purospher Star NH2 250-4 µm column with a RI detector
ter paper back into the boiling flask with a spatula. The re- at a temperature of 40 ◦ C, flow rate of 1 mL min−1 and volume
maining residue was washed with 200 mL of boiled 0.313 N injection of 20 µL.
212 Tri Suwandi et al.: Fruits 71 (2016) 209–214

2.3 Statistical analysis of data function of GA in this mechanism, contributing to increasing

the fiber content in pineapple, as was observed for the treat-
Data were analyzed using analysis of variance (ANOVA) ment GA1 CK1 in this study.
followed by Duncan’s multiple range test (DMRT) at the 5% Pineapple fruit treated with GA, CK, or the combination of
level. GA with CK significantly increased the fruit water content (ta-
ble II). The water content was increased by 7.07% in response
to application of 200 ppm of GA combined with 24 ppm of
3 Results and discussion CK. This may be related to the effect of GA reducing the water
potential in the cell by increasing the plasticity of the cell wall
3.1 Effect of gibberellin and cytokinin on the pineapple followed by hydrolysis of starch into sugar [36]. The entry of
harvest index water into the fruit cells helps cell enlargement, thus increas-
ing the fruit size [24]. In addition, CK also plays a role in the
Pineapple fruit treated with GA alone or in combination absorption of water during fruit growth [10, 20, 21].
with CK slightly increased the fruit weight and harvest in- The fruit potassium content was affected by GA and
dex (table I). The harvest index was increased by 6 and 10% CK. Cytokinin at 24 ppm increased the potassium content
by treatment with 48 ppm of CK and 100 ppm of GA, re- by 4.85%, but it was decreased by higher CK concentration
spectively. However, the treatments of 100 ppm of GA com- (48 ppm) and GA treatments, such as 100 GA, which de-
bined with 24 ppm of CK, and 200 ppm of GA combined with creased it by 23.86% compared with the control. Increasing
48 ppm of CK showed a significant increase in the harvest in- potassium content showed correlation with the water content
dex compared with control (12 and 16%, respectively). Appli- in pineapple fruit when CK was applied (table II), an ef-
cation of GA alone or in combination with CK significantly fect related to the osmotic regulation in the cell. Potassium
increased the crown length, which was a positive effect on the is involved in the charge balance of organic acids in the cell
quality of this planting material. The fruits not treated with GA vacuole [37]. Some phytohormones, such as ABA and GA,
matured five days earlier than those treated with GA, while CK maintain the balance of osmotic pressure by regulating K+ /H+
had no effect on days to maturity (table I). Similar results were exchangers [38]. However, CK has not yet been reported in
obtained by others [8]. this context.
It is known that GA plays a key role in cell expansion [17]. The vitamin C content of fruits treated with GA alone or
Gibberellin may not increase the number of cells, but increases in combination with CK was significantly higher than those
the cell volume of pineapple fruit [9]. Gibberellin is known of the control and CK alone, and there was a significant con-
to be an important phytohormone that regulates source-sink centration response (table II). Treatment of 200 ppm of GA
relations or sink demand by activating enzymes involved in combined with 48 ppm of CK was the best combination, in-
sugar metabolism [18, 19]. CK also plays an important role in creasing the content of vitamin C by 78.5% compared with the
the absorption of water and mobilization of assimilates to the control. These results are consistent with those of a previous
sink organs such as the fruit [10, 20, 21]. Application of GA study which showed that 50 or 100 ppm of GA increased the
increased the fruit weight of ‘MD-2’ [8] and ‘Comte de Paris’ vitamin C content of ‘MD-2’ pineapple fruit [9]. The increase
pineapple [9]. Application of CK increased the fruit weight of in the vitamin C content is favored by the delay in the physi-
Lupinus angustifolius [22] and ‘Duke’ blueberry [23]. The fact ological ripening of the pineapple fruit [39]. Vitamin C is an
that GA combined with CK increases the fruit weight of non- antioxidant compound essential for all parts of a plant, and it
climacteric multiple fruits (sorosis) has not been reported. also plays a role in cell expansion [40, 41]. The increase in the
vitamin C content in pineapple fruits may be one of the reasons
why GA increases the fruit cell area and size [9].
3.2 Effect of gibberellin and cytokinin on pineapple The total soluble solids (TSS) and titratable acidity (TA)
fruit quality values of fruits treated with GA and CK were significantly
lower than the control, and there was a significant concen-
The effect of gibberellin and cytokinin on pineapple fruit tration response (table III). For example, TSS were reduced
quality is shown in tables II and III. As shown in table II, con- by 19.88% and TA by 17.02% in response to application of
trol and fruits treated with CK alone had a more yellowish- 100 ppm of GA combined with 48 ppm of CK compared with
white flesh than fruits treated with GA alone or combined with the control. However, there was no effect of GA and CK on
CK. However, the results indicate that fruit firmness was not the ratio of sugar to acid (TSS/TA) (table III). These results
affected by GA and CK treatments. were different from those obtained in a previous study, where
Gibberellin at 100 ppm combined with 24 ppm of CK GA increased TSS and TA of ‘MD-2’ pineapple fruits, and
was the best combination to increase the fruit fiber content by where TSS/TA was increased in low-GA-concentration treat-
106.06% compared with the control (table II). Changes in fruit ments and decreased in high-level GA treatment [9]. These
fiber contents, as determined by GA and CK, may lead to in- results may be related to the delay in the fruit physiological
hibition of the fruit softening [30, 31]. Gibberellin is known to maturity in response to GA application [8], as changes of TSS
inhibit the reduction of the stability of the cell wall [32] by in- and TA in pineapples are highly correlated with the level of
fluencing the enzymes involved in this process [27,33–35]. On fruit maturity [42–45].
the other hand, there is no information available on the effects Single treatments of GA or CK also significantly af-
of CK on fruit fiber content. However, CK may improve the fected the sugar content of ‘Smooth Cayenne’ pineapple fruits
 Tri Suwandi et al.: Fruits 71 (2016) 209–214 213

Table II. Effect of gibberellin and cytokinin on pineapple fruit quality (part 1). Fruits were collected at 156 DAF (green mature stage). Values
are means for 45 fruits.
Treatments Flesh color index Fruit firmness (N) Fruit fiber (%) Water content (%) Potassium (ppm) Vitamin C (ppm)
GA0 CK0 2 288.70a 0.33b 81.70f 117.02b 9.44e
GA0 CK1 2 281.85a 0.31b 85.33d 122.70a 9.28e
GA0 CK2 2 304.63a 0.45b 84.98e 108.45d 9.59e
GA1 CK0 1 291.67a 0.33b 84.73e 89.31h 14.13bc
GA1 CK1 1 329.07a 0.68a 86.76b 100.56f 12.31cd
GA1 CK2 1 321.67a 0.43b 85.84c 102.03e 10.80de
GA2 CK0 1 282.96a 0.37b 85.86c 95.60g 14.73b
GA2 CK1 1 318.52a 0.31b 87.48a 102.39e 15.34ab
GA2 CK2 1 299.07a 0.47b 85.61cd 109.74c 16.85a
Values within a column followed by the same letter are not significantly different (P < 0.05) according to Duncan’s multiple range test.

Table III. Effect of gibberellin and cytokinin on pineapple fruit quality (part 2). Fruits were collected at 156 DAF (green mature stage). Values
are means for 45 fruits.
Treatments TSS (◦ Brix) TA (%) TSS/TA Sucrose (%) Fructose (%) Glucose (%) Sucrose/Hexose
GA0 CK0 11.92a 0.47a 25.25a 8.17a 1.65b 1.93b 2.28d
GA0 CK1 10.64ab 0.41ab 26.12a 6.22f 1.62c 1.89b 1.77e
GA0 CK2 10.25ab 0.40ab 26.11a 6.80c 1.56d 1.19g 2.47c
GA1 CK0 11.25ab 0.45ab 25.58a 8.14a 0.95i 1.30e 3.62a
GA1 CK1 10.17ab 0.43ab 23.90a 6.54d 1.34f 1.62d 2.21d
GA1 CK2 9.95b 0.39b 25.62a 6.80c 1.03h 1.23f 3.01b
GA2 CK0 10.40ab 0.42ab 25.54a 7.04b 1.38e 1.71c 2.28d
GA2 CK1 9.51b 0.40ab 23.78a 6.31e 1.19g 1.65d 2.23d
GA2 CK2 9.55b 0.41ab 24.10a 4.10g 2.43a 2.71a 0.80f
Values within a column followed by the same letter are not significantly different (P < 0.05) according to Duncan’s multiple range test.

including sucrose and hexose (fructose and glucose). For ex- Jakarta for supplying cytokinin, and Bina Guna Kimia company,
ample, application of 200 ppm of GA combined with 48 ppm Jakarta, for supplying gibberellin.
of CK significantly decreased the sucrose content by 49.78%,
increased the fructose content by 47.27%, increased the glu-
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Cite this article as: Tri Suwandi, Kumala Dewi, Priyo Cahyono. Pineapple harvest index and fruit quality improvement by application of
gibberellin and cytokinin . Fruits 71 (2016) 209–214.