Plant Growth, Fruit Production and Total Terpenoid Compounds in Bitter Gourd (Momordica charantia L.) at Various Levels of Phosphorus Fertilization

Abstract

Bitter gourd (Momordica charantia L.) fruits have been reported to have pharmacological functions such as anti-bacterial, lowering blood sugar, and preventing cardiovascular disease. Terpenoids are the bioactive compounds that play a role in those functions. The phosphorus is essential in the biosynthesis of terpenoids. This research aimed to determine plant growth characteristics, fruit production, and terpenoid production in bitter gourds fertilized with various phosphorus. The experiment was conducted at the IPB experimental station in Cikarawang, 6o32’58.3” S south latitude and 106o43’54.8” E east longitude, Bogor, Indonesia, from July to October 2023. The experiment used a completely randomized block design with a single factor: fertilizer treatments and three replications. The treatments were without fertilizer, chicken manure only, and varying dosages of SP-36 (0, 20, 40, 60 g per plant). Plants treated with 40 g SP-36 per plant significantly had longer stems at four weeks after planting (WAP) (153.5 cm), substantially more female flowers at 5 WAP (6.4 flowers), heavier fresh weight per fruit (243.98 g), and higher fruit carotene level (86 μg.g-1). Plants treated with 60 g SP-36 per plant had the heaviest fresh fruit weight per plant (2,820.9 g). On the other hand, chicken manure fertilizer resulted in an elevated number of female flowers at 7 WAP (6.8 flowers), leaf potassium content (3.41%), chlorophyll a (250 μg.g-1), chlorophyll b (114 μg.g-1), and total chlorophyll concentration (363 μg.g-1). There were no significant differences in IC50, terpenoid content, and terpenoid production; however, plants treated with 60 g SP-36 per plant tended to have lower IC50 (1,347.67 ppm) and terpenoid content (95,227 μmol NE.g-1 dry fruit). In contrast, plants treated with 40 g SP-36 per plant tended to have higher terpenoid production (15,995 mmol NE per plant) than other treatments.

Full text

Journal of Tropical Crop Science Vol. 11 No. 2, June 2024
www.j-tropical-crops.com
184 Dian Novira Rizva, Maya Melati, Sandra Arin Aziz
Received 09/04/2024; Revised 25/06/2024; Accepted 27/06/2024
https://doi.org/10.29244/jtcs.11.02.184-194
Plant Growth, Fruit Production and Total Terpenoid
Compounds in Bitter Gourd (Momordica charantia L.) at
Various Levels of Phosphorus Fertilization
Dian Novira RizvaA, Maya Melati*BC, Sandra Arin AzizBC
A Agronomy and Horticulture Study Program, Graduate School, Bogor Agricultural University, Bogor,
Indonesia,16680
B Department of Agronomy and Horticulture, Faculty of Agriculture, Bogor Agricultural University, Bogor,
Indonesia, 16680
C Tropical Biopharmaca Research Center, Bogor Agricultural University, Bogor, Indonesia
*Corresponding author; email: maya_melati@apps.ipb.ac.id
Abstract
Bitter gourd (Momordica charantia L.) fruits have been
reported to have pharmacological functions such as
anti-bacterial, lowering blood sugar, and preventing
cardiovascular disease. Terpenoids are the bioactive
compounds that play a role in those functions.
The phosphorus is essential in the biosynthesis of
terpenoids. This research aimed to determine plant
growth characteristics, fruit production, and terpenoid
production in bitter gourds fertilized with various
phosphorus. The experiment was conducted at the
IPB experimental station in Cikarawang, 6o32’58.3”
S south latitude and 106o43’54.8” E east longitude,
Bogor, Indonesia, from July to October 2023. The
experiment used a completely randomized block
design with a single factor: fertilizer treatments and
three replications. The treatments were without
fertilizer, chicken manure only, and varying dosages
of SP-36 (0, 20, 40, 60 g per plant). Plants treated
with 40 g SP-36 per plant signicantly had longer
stems at four weeks after planting (WAP) (153.5
cm), substantially more female owers at 5 WAP (6.4
owers), heavier fresh weight per fruit (243.98 g), and
higher fruit carotene level (86 µg.g-1). Plants treated
with 60 g SP-36 per plant had the heaviest fresh fruit
weight per plant (2,820.9 g). On the other hand,
chicken manure fertilizer resulted in an elevated
number of female owers at 7 WAP (6.8 owers), leaf
potassium content (3.41%), chlorophyll a (250 µg.
g-1), chlorophyll b (114 µg.g-1), and total chlorophyll
concentration (363 µg.g-1). There were no signicant
di󰀨erences in IC50, terpenoid content, and terpenoid
production; however, plants treated with 60 g SP-36
per plant tended to have lower IC50 (1,347.67 ppm)
and terpenoid content (95,227 µmol NE.g-1 dry fruit).
In contrast, plants treated with 40 g SP-36 per plant
tended to have higher terpenoid production (15,995
mmol NE per plant) than other treatments.
Keywords: antioxidant, chicken manure, chlorophyll,
precursors
Introduction
Bitter gourd (Momordica charantia L.) is an annual
plant widely grown in tropical and subtropical regions
including East Africa, Asia, the Caribbean, and South
America (Paul et al., 2009). Bitter gourd is commonly
utilized as a vegetable and a medicinal potential plant
(Yaldiz et al. 2015). Bitter gourd contains various
secondary metabolite compounds (Slamet., 2020),
one of the secondary metabolite compounds is a
group of terpenoids. Bitter gourd-derived terpenoid
compounds include cucurbitanes (Huang et al.,
2020), charantin (Weng et al., 2013), and momordicin
(Zhao et al., 2005). These compounds have been
shown to possess pharmacological benets, such
as anti-cancer and anti-bacterial properties (Cuong
et al., 2017), suppressing cardiovascular disease
disorders (Tuan et al. 2017), also as anti-diabetic, at
a level of 300 mg.kg-1 bitter gourd can reduce blood
sugar levels by 31.64% and increase insulin levels by
27.35% (Mahwish et al., 2021).
Due to its numerous properties and benets, bitter
gourd has attained signicant economic value as
a commercial commodity in the market (Lee et al.,
2021). However, the current requirements of bitter
gourd, both in terms of quantity and quality, are
not yet optimal. Therefore, e󰀨orts are necessary
to enhance bitter gourd production, particularly by
increasing the yield of secondary metabolites such
as terpenoid compounds, which can be achieved
through fertilization techniques. Fertilization has
been demonstrated to enhance plant production

Plant Growth, Fruit Production and Total Terpenoid Compounds in Bitter Gourd ..........
Journal of Tropical Crop Science Vol. 11 No. 2, June 2024
www.j-tropical-crops.com
185
Received 09/04/2024; Revised 25/06/2024; Accepted 27/06/2024
https://doi.org/10.29244/jtcs.11.02.184-194
(Purba et al., 2021) and can signicantly inuence
the production of secondary metabolite compounds
(Yang et al., 2018).
Among the fertilizers, phosphorus is a crucial factor
inuencing secondary metabolite compounds.
Phosphorus plays a pivotal role in the production of
terpenoid compounds within the plant, as terpenoid
precursors, including IPP (Isopentenyl diphosphate),
DMAPP (Dimethylallyl pyrophosphate), GDP (Geranyl
diphosphate), and FDP (Farnesyl diphosphate),
contain high-energy phosphorus bonds. Additionally,
phosphorus constitutes an essential component
of ATP and NADPH molecules, which are vital for
synthesizing terpenoids through both the mevalonic
acid (MVA) and methylerythritol phosphate (MEP)
pathways. Thus, phosphorus serves as a fundamental
element in the production of terpenoid compounds
(Bustamante et al., 2020).
Several research results reveal that applying P
fertilizer has a signicant role in the biosynthesis of
secondary metabolite compounds (Nell et al., 2009).
Increasing the amount of P fertilizer can increase
terpenoid biosynthesis (Peng and Teik, 2022). Based
on the research results of Naorem et al., (2019), the
application of P fertilizer (90 kg.ha-1) gave the highest
results in the fruit number per plant, fruit weight, fruit
length, fruit diameter, fruit weight per plant, and fruit
weight per hectare on bitter gourd plants. This is
because the absorption of macro and micronutrients
increases with the addition of P and this is also
related to improved root growth due to the addition
of P (Fageria et al., 2016). P utilization e󰀩ciency can
be increased when applied along with other nutrients,
primarily N and K (Duan et al., 2004).
Phosphorus can be sourced from both inorganic
and organic fertilizers. Inorganic fertilizers like SP-
36 contain phosphorus, while organic fertilizers such
as chicken manure also provide a high phosphorus
level. Considering the role of phosphorus in
increasing the yield and production of secondary
metabolite compounds, especially terpenoid
compounds, research is necessary to determine
the best phosphorus fertilizer treatment that can
increase growth, fruit production, and total terpenoid
production in bitter gourd.
Materials and Methods
The experiment was conducted from July to October
2023 at the Cikarawang experimental station, IPB,
Bogor, West Java, Indonesia. The materials used
are bitter gourd seeds of the Opal F1 variety, laying
chicken manure, urea fertilizer, SP-36 fertilizer, KCl
fertilizer, and silver-black plastic mulch. The tools
used include a UV-vis spectrophotometer, centrifuge,
sonication, and micropipette.
The experiment used a randomized complete block
design with one factor consisting of six fertilizer
treatments: without fertilization, chicken manure,
and four SP-36 fertilizer doses consisting of 0, 20,
40, and 60 g SP-36 per plant (equivalent to 0, 50,
100, and 150% of the recommended dose from PT.
East-West). Each treatment was replicated thrice, so
there were 18 experimental units, each consisting of
32 plants.
Experimental Procedures
Bitter gourd seeds were sown in trays for 14 days,
then transplanted to the experimental eld with a plot
size of 1 m x 11.75 m, planting distance of 0.5 m x
0.75 m, distance between plots of 0.5 m, and distance
between replicates of 1 m. The seedlings with the
criteria of 2-4 leaves fully opened were transplanted.
Chicken manure as a treatment (0.73 kg per plant)
was applied two weeks before transplanting. The SP-
36 fertilizer (Ca(H2PO4)) was applied one week after
transplanting. SP-36 fertilizer with doses of 20, 40,
and 60 g SP-36 per plant (equivalent to 35.5 kg.ha-1;
71 kg.ha-1; and 106.5 kg.ha-1) was given in 6 times-
applications. The rst application of each treatment
was 10, 20, and 30 g SP-36 per plant, and the rest
was applied ve times with doses of 2, 4, and 6 g SP-
36 per plant every two weeks.
For the four SP-36 treatments, N and K fertilizers
were also given with a dose of 45 g urea per plant and
30 g KCl per plant, respectively. The urea and KCl
were also applied six times with the rst application
being 20 g urea per plant and 10 g KCl per plant.
The next application of N and K fertilizer was 5 g
urea per plant urea and 4 g KCl per plant every two
weeks. The rst fruit harvest was done 43 days after
transplanting. The fruit harvest can be done 10 times
with specic criteria, including shiny dark green color,
fruit length of approximately 26 cm, and fruit weight of
approximately 305 g.
Plant Growth and Development
Stem height and leaf number were measured from
2 weeks after planting (WAP) to 5 WAP at 1-week
intervals. The number of stems was counted at 4 and
5 WAP.
The number of male and female owers was counted
on each plant sample. Days to 50% owering are
recorded based on the age at which owers are rst
initiated.

Journal of Tropical Crop Science Vol. 11 No. 2, June 2024
www.j-tropical-crops.com
186 Dian Novira Rizva, Maya Melati, Sandra Arin Aziz
Received 09/04/2024; Revised 25/06/2024; Accepted 27/06/2024
https://doi.org/10.29244/jtcs.11.02.184-194
Relative growth rate (RGR) was calculated based on
plant dry weight per unit time (g.g-1 per day), whereas
net assimilation rate (NAR) is calculated based on
plant weight per unit leaf area within a specic time
(g.cm-2 per day). RGR and NAR are calculated at 3-5
and 5-7 WAP, respectively.
Leaf N, P, and K Content
The mature leaves’ N, P, and K contents were
measured at 4 WAP. The leaves are air-dried and
then dried in the oven at 60oC until reaching a
constant weight. Leaf N analysis was conducted
using the Kjeldahl method. P and K were extracted
using the wet ash method using HNO3 and HClO4.
P levels were calculated using a spectrophotometer,
while K was calculated using an atomic absorption
Spectropotometer (AAS) (Balitttanah, 2005).
Yield
Fruit number, fresh weight per fruit (g), fresh weight
of fruit per plant (g), and dry weight of fruit per plant
(g) were measured as yield components. The fresh
weight per fruit was calculated by weighing each unit
of bitter gourd fruit sample that had been harvested.
The fresh fruit weight per plant was calculated by
adding up all the weights of fruit per plant that had
been harvested. Fruit dry weight per plant was
calculated using the following formula:
Dry weight of fruit per plant = fresh weight of fruit per
plant x (100% - fruit water %)
Fruit Pigment Concentration
The fruit pigment was determined from the fourth-
harvested fresh fruit using the Sims and Gamon
(2002) method. The analysis procedure was initiated
by grinding the fruit samples and then adding 2 mL
acetone Tris. The mixture was then centrifuged at
14000 rpm for 10 minutes. Next, 1 mL of supernatant
was added with 3 mL of acetone Tris and homogenized.
The absorbance of the mixture was measured using
a Shimadzu UV-1201 UV-VIS spectrophotometer at
wavelengths of 663, 647, 537, and 470 nm.
Inhibiting Concentration (IC50)
The IC50 in bitter gourd fruit was determined by using
the DPPH (2,2-Diphenyl-1-Picrylhydrazyl) method
(Salazar et al., 2011) The procedure for this analysis
begins by preparing 100 mg of concentrated bitter
gourd fruit extract, dissolving it with 1 ml of DMSO,
and then diluting it to various concentrations (125 -
8000 ug.ml-1). 100uL of sample extract at di󰀨erent
concentrations was added with 100uL of DPPH
125 Um into a 96-well microplate. Next, the extract
solution was incubated in the dark for 30 minutes.
The absorbance was measured at a wavelength of
517 nm. Ascorbic acid was used as a standard at a
concentration of (0.3125 – 10 ug ascorbic acid per
ml of methanol). The capacity value is expressed in
percent IC50 (e󰀨ective concentration of the sample
capable of inhibiting 50% of DPPH radicals). The
IC50 value is calculated based on the line equation
obtained by entering the value 50 as the y variable
and then determining the x variable value as the
concentration of the sample.
Total Terpenoid Production
The principle of determining total terpenoids using
spectrophotometry is based on the colorimetric
method (Łukowski et al., 2022). A total of 0.2 g of
dry fruit was weighed, then cold 95% methanol
was added. The mixture was incubated for 48
hours at room temperature. The samples were
then centrifuged at 4000 g for 15 minutes at room
temperature. The resulting supernatant was
transferred to a test tube then 1.5 mL of chloroform
was added and homogenized using a vortex. Once
homogeneous, 100 µL of H2SO4 was added to the
sample and incubated at room temperature for 1.5–2
hours. After incubation, 1.5 mL of 95% methanol was
added to the sample and homogenized using a vortex
until the precipitate dissolved again. The samples
were then analyzed with a UV-Vis spectrophotometer
at 520 nm. Total terpenoids were determined based
on the Nerol standard curve. Terpenoid contents were
quantied in Nerol equivalents (NE). Determination
of terpenoid productions (mmol NE per plant) was
obtained by multiplying the dry weight (weight of dry
fruit per plant) by the terpenoid content (μmol NE. g-1
of dry fruit).
Data Analysis
Data were analyzed using ANOVA at α=0.05 and
continued with the Duncan multiple range test (DMRT)
if the means between treatments were signicant.
Statistical analysis was performed using R Studio
software version 4.3.1.
Result and Discussion
Plant Growth and Development
The application of 40 g SP-36 per plant led to a
signicant 17.09% increase in stem length at four
weeks after planting (WAP) (P<0.05) compared to
without fertilizer (Table 1), but leaf numbers were not
di󰀨erent among all fertilizer treatments.

Plant Growth, Fruit Production and Total Terpenoid Compounds in Bitter Gourd ..........
Journal of Tropical Crop Science Vol. 11 No. 2, June 2024
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187
Received 09/04/2024; Revised 25/06/2024; Accepted 27/06/2024
https://doi.org/10.29244/jtcs.11.02.184-194
Leaf N, P and K Content
Various phosphorus fertilizations resulted in
signicant di󰀨erences in the leaf potassium content
(P<0.05) (Table 2). The highest potassium content
was found in chicken manure, with an increase of
39.75% compared to 0 g SP-36 per plant. On the
other hand, various phosphorus fertilizations did not
a󰀨ect leaf nitrogen and phosphorus content.
The di󰀨erences in the number of female owers due
to various fertilizer treatments occurred at 5 and 7
WAP (Table 3). At 5 WAP, the highest ower number
was obtained from the 40 g SP-36 per plant with a
45.45% increase compared to 0 g SP-36 per plant
and 68.42% increase compared to without fertilizer
(P<0.05). On the other hand, 60 g SP-36 per plant led
to a 44.44% increase in 7 WAP (P<0.05) compared
to without fertilizer. The number of male owers at
5, 7 WAP, and the days to 50% owering were not
signicantly di󰀨erent among fertilizer treatments.
Fertilization treatment did not signicantly a󰀨ect the
relative growth and net assimilation rates at 3-5 and
5-7 WAP (Table 4).
Yield
The application of 40 g SP-36 per plant, resulted
in a signicantly 10.84% higher fresh weight per
fruit compared to 20 g SP-36 per plant and 29.38%
higher than without fertilizer (P<0.01, Table 5). The
application of 60 g SP-36 per plant produced 29.43%
higher fresh weight of fruit per plant compared to
without fertilizer (P<0.05, Table 5). Various levels of
phosphorus fertilization did not a󰀨ect the fruit number
and dry weight of fruit per plant.
Fruit Pigment Concentration
Applying 40 g of SP-36 caused a higher concentration
of chlorophyll a, chlorophyll b, carotene, and total
chlorophyll (P<0.05) by 200%, 166.66%, 207.14%,
and 185.83%, respectively, compared to those
without fertilizer (Table 6.).
Table 1. Stem length and leaf number at 3, 4, and 5 WAP with various levels of phosphorus fertilization
Treatment
3 WAP 4 WAP 5 WAP
Stem
length
(cm)
Leaf
number
Stem
length
(cm)
Leaf
number
Stem
length
(cm)
Leaf
number
Without fertilizer 68.6 17.3 131.1b 51.6 172.2 105.2
Chicken manure 73.7 19.2 133.3b 55.7 174.7 105.8
0 g SP-36 per plant 79.3 21.9 151.6a 69.2 176.6 111. 4
20 g SP-36 per plant 75.0 23.2 144.5ab 62.4 178.2 106.9
40 g SP-36 per plant 81.2 24.1 153.5a 66.1 177.3 114.4
60 g SP-36 per plant 71.5 22.8 143.9ab 55.9 177.8 122.9
P-Value 0.305 0.366 0.034 0.415 0.965 0.666
Sig. ns ns * ns ns ns
Note: values followed by di󰀨erent letters in the same column are signicantly di󰀨erent in the Duncan test; ns=non-signicant;
* = signicant at α=0.05
Table 2. Leaf N, P, and K content at di󰀨erent levels of phosphorus fertilization
Treatment Nitrogen (%) Phosphorus (%) Potassium (%)
Without fertilizer 4.54 0.28 2.73abc
Chicken manure 4.89 0.31 3.41a
0 g SP-36 per plant 4.35 0.22 2.44bc
20 g SP-36 per plant 4.66 0.27 2.96abc
40 g SP-36 per plant 4.39 0.23 2.08c
60 g SP-36 per plant 4.59 0.28 3.27ab
P-Value 0.202 0.199 0.040
Sig. ns ns *
Note: values followed by di󰀨erent letters in the same column are signicantly di󰀨erent in the Duncan test; ns=non-signicant;
* = signicant at α=0.05

Journal of Tropical Crop Science Vol. 11 No. 2, June 2024
www.j-tropical-crops.com
188 Dian Novira Rizva, Maya Melati, Sandra Arin Aziz
Received 09/04/2024; Revised 25/06/2024; Accepted 27/06/2024
https://doi.org/10.29244/jtcs.11.02.184-194
Table 3. Number of male and female owers per plant at 5 and 7 WAP, and days to 50% at di󰀨erent levels of
phosphorus fertilizations.
Treatment
Number of males
owers
Number of female
owers Days to 50% owering
5 WAP 7 WAP 5 WAP 7 WAP
Without fertilizer 3.6 5.6 3.8b 4.5b 28.0
Chicken manure 3.2 6.6 5.2ab 6.8a 27.7
0 g SP-36 per plant 4.5 5.8 4.4b 5.6ab 27.3
20 g SP-36 per plant 3.7 6.5 4.8b 6.2a 27.3
40 g SP-36 per plant 3.7 6.3 6.4a 6.4a 27.0
60 g SP-36 per plant 3.7 7.1 4.2b 6.5a 27.3
P-Value 0.380 0.171 0.017 0.017 0.775
Sig. ns ns * * ns
Note: values followed by di󰀨erent letters in the same column are signicantly di󰀨erent according to DMRT; ns=non-
signicant; * = signicant at α=0.05
Table 4. Plant relative growth rate (RGR) and net assimilation rate (NAR) at 3-5 WAP and 5-7 WAP with
various phosphorus fertilization
Treatment
Relative growth rate
(g.g-1 per day)
Net assimilation rate
(g.cm-2 per day)
3-5 WAP 5-7 WAP 3-5 WAP1) 5-7 WAP1)
Without fertilizer 3.46 4.12 0.08 0.13
Chicken manure 3.66 4.22 0.10 0.10
0 g SP-36 per plant 3.53 4.46 0.07 0.18
20 g SP-36 per plant 3.64 4.35 0.09 0.14
40 g SP-36 per plant 3.78 4.47 0.10 0.14
60 g SP-36 per plant 3.84 4.44 0.11 0.13
P-Value 0.486 0.569 0.855 0.877
Sig. ns ns ns ns
Note: ns=non-signicant; 1) Data was transformed using the formula
Note: ns=non-significant; 1) Data was transformed using the formula √ + 0,5
Table 5. Fruit number, fresh weight per fruit (g), fresh weight of fruit per plant (g), and dry weight of fruit per
plant with various phosphorus fertilization
Treatment
Harvest
Fruit number
per plant
Fresh weight
per fruit (g)
Fresh weight of fruit
per plant (g)
Dry weight of fruit
per plant (g)
Without fertilizer 11.7 188.54d 2,179.5b 90.74
Chicken manure 12.1 237.32ab 2,828.4a 118.79
0 g SP-36 per plant 12.2 225.12bc 2,714.9a 114.63
20 g SP-36 per plant 12.5 220.07c 2,725.8a 134.85
40 g SP-36 per plant 11.5 243.93a 2,820.4a 163.75
60 g SP-36 per plant 11.8 240.05ab 2,820.9a 163.90
P-Value 0.737 0.000182 0.00194 0.314
Sig. ns ** ** ns
Note: values followed by di󰀨erent letters in the same column are signicantly di󰀨erent according to the DMRT; ns= non-
signicant; *= signicant at α=0.05; ** = signicant at α=0.01.

Plant Growth, Fruit Production and Total Terpenoid Compounds in Bitter Gourd ..........
Journal of Tropical Crop Science Vol. 11 No. 2, June 2024
www.j-tropical-crops.com
189
Received 09/04/2024; Revised 25/06/2024; Accepted 27/06/2024
https://doi.org/10.29244/jtcs.11.02.184-194
IC50, Fruit Terpenoid Content, and Plant Terpenoid
Production
Various levels of phosphorus fertilization did not
signicantly a󰀨ect IC50, but plants that received
60 g SP-36 tended to have the lowest IC50 (Table
7). Terpenoid content tended to be higher in plants
treated with 60 g SP-36 compared to control (Table
7).
Discussion
Bitter gourd applied with 40 g SP-36 per plant at
4 WAP showed signicantly a longer stem than
without fertilizer. This positive e󰀨ect of fertilizer
shows that fertilization is irreplaceable in increasing
growth, yields and the quality of plant growth and
development (Li and Shang., 2021). Fertilization is
one of the main inputs for providing nutrients. Bitter
gourd treated with 60 g SP-36 per plant exhibited a
higher leaf number at 5 weeks after planting (WAP)
compared to 0 g SP-36 per plant, although this
di󰀨erence was not statistically signicant. Consistent
with prior studies, phosphorus has been observed
to enhance plant vegetative growth, increasing leaf
number (Martin et al., 2018) and stem length (Razaq
et al., 2017). Phosphorus, as a macronutrient, plays
a crucial role in the growth and development of plants
(Wang et al., 2021), particularly in owering (Dey et
al., 2021). Higher phosphorus doses have been linked
to increased ower production in cucumber (Cucumis
sativus L.) (Vaudo et al., 2022), attributed to its
ability to enhance the assimilation, translocation, and
partitioning of oral components (Sahu et al., 2021).
In this study, the application of various phosphorus
fertilizers had a signicant impact on female owers
at 5 and 7 WAP, in line with the same family, the use
of phosphorus signicantly can increase the ratio of
female owers to watermelon (Maluki et al., 2016).
This is because phosphorus is crucial for enzymatic
activity and hormone synthesis that support female
ower development (Choudhary et al., 2013).
Table 6. Fruit pigment concentration with various levels of phosphorus fertilization.
Treatment Chlorophyll a
(µg.g-1)
Chlorophyll b
(µg.g-1)
Carotene
(µg.g-1)
Chlorophyll total
(µg.g-1)
Without fertilizer 80b 39b 28b 120b
Chicken manure 250a 114a 83a 363a
0 g SP-36 per plant 240a 106a 84a 344a
20 g SP-36 per plant 190a 85a 67a 279a
40 g SP-36 per plant 240a 104a 86a 343a
60 g SP-36 per plant 220a 101a 75a 320a
P-Value 0.018 0.019 0.023 0.017
Sig. * * * *
Note: values followed by di󰀨erent letters in the same column are signicantly di󰀨erent according to the DMRT; ns=non-
signicant; * = signicant at α=0.05.
Table 7. Antioxidant activity (IC50), fruit terpenoid content, and terpenoid production with various levels of
phosphorus fertilization.
Treatment IC50
(ppm)
Fruit terpenoid content
(µmol NE.g-1 of dry fruit)1)
Terpenoid production
(mmol NE per plant)1)
Without fertilizer 2,214.5 80,830 6,791
Chicken manure 1,448.6 79,821 8,162
0 g SP-36 per plant 1,602.8 85,266 11,630
20 g SP-36 per plant 1,658.6 94,292 12,888
40 g SP-36 per plant 1,669.0 92,950 15,995
60 g SP-36 per plant 1,347.7 95,227 14,494
P-Value 0.326 0.985 0.624
Sig. ns ns ns
Note: values followed by di󰀨erent letters in the same column are signicantly di󰀨erent in the Duncan test; ns=non-signicant;
1) Data was transformed using the formula
Note: values followed by different letters in the same column are significantly different in the Duncan test; ns=non-
significant; 1) Data was transformed using the formula √ + 0,5; NE = Nerol equivalent
; NE = Nerol equivalent

Journal of Tropical Crop Science Vol. 11 No. 2, June 2024
www.j-tropical-crops.com
190 Dian Novira Rizva, Maya Melati, Sandra Arin Aziz
Received 09/04/2024; Revised 25/06/2024; Accepted 27/06/2024
https://doi.org/10.29244/jtcs.11.02.184-194
Applying 40 g SP-36 per bitter gourd plant can
increase fruit weight compared to plants treated
with 0 g SP-36 per plant. This is due to phosphorus,
with a signicant portion being translocated to the
fruit area. This translocation is driven by the high
energy demand during seed and fruit production
(Johan et al., 2021). Furthermore, phosphorus is an
essential element in cell energy transfer because
it is part of adenosine triphosphate (ATP), cytidine
triphosphate (CTP), guanosine triphosphate (GTP),
uridine triphosphate (UTP), phosphoenolpyruvate,
and other phosphorylated intermediates (Malhotra
et al., 2018). Increasing the dose of phosphorus
fertilizer to 100 kg ha-1 can increase the weight of
bitter gourd fruit, signicantly di󰀨erent from doses
of 80, 60, 40, 20, and 0 ha-1 (Ashraf et al. 2019).
Increasing phosphorus concentration in the same
Cucurbitaceae family can produce larger cucumber
fruit (Lee et al., 2024). The application of phosphorus
can also increase Citrus yields compared to without
phosphorus (Li et al., 2020). Citrus yields increased
with phosphorus doses from 0.1 to 0.3 kg per plant (Li
et al., 2019). In line with this research, increasing the
dose of phosphorus fertilizer can increase fruit weight
but was not signicantly di󰀨erent from the chicken
manure treatment. Chicken manure can improve soil’s
physical, chemical, and biological properties (Lima et
al., 2021). In addition, chicken manure contains two
to four times more phosphorus than other manures,
ranging from 13.6 to 25.4 g P2O5 kg-1dm (Kacprzak
et al., 2022). Chicken manure had the best e󰀨ect on
the weight of bitter gourd fruit per plot, weighing 464
grams (Wardana et al., 2020). Moreover, chicken
manure can increase the weight of bitter gourd up to
54.68% per plant (Miguel et al., 2023).
Bitter gourd without fertilizer signicantly resulted in
a lower concentration of chlorophyll a, chlorophyll b,
carotene, and total chlorophyll compared to those
treated with fertilizer (chicken manure only and
various SP-36 dosages treatments), this is because
those treatments have macronutrients such nitrogen,
phosphorus, and potassium which have an essential
role to increase photosynthetic capacity by increasing
chlorophyll content (Deng et al., 2020). Nitrogen is
used in chloroplasts with thylakoids and photosynthetic
enzymes (Mu et al., 2016). Phosphorus is involved
in cellular processes, including energy conservation,
metabolic regulation, and signal transduction
(Carstensen et al., 2018). Phosphorus deciency
can reduce stomata opening. If stomata opening is
reduced, the lower CO2 can be captured, reducing
triose phosphate, signicantly reducing the recycling
of ATP and NADPH, inhibiting photosynthetic capacity
(Neocleous and Savvas., 2018). Potassium can
increase photosynthetic assimilation and improve
nutrient absorption (Sustr et al., 2019). Potassium
deciency can signicantly inhibit the biosynthesis of
chlorophyll a and b and total chlorophyll (Thornburg
et al., 2020). The maximum value of total chlorophyll
content was obtained at a dose (N:P:K: 300:120:100
kg-1ha), signicantly di󰀨erent from the treatment
without fertilization (Ashraf et al., 2019).
Based on the results of this research, bitter gourd
treated with 60 g SP-36 tend to have a lower IC50
value; namely, 1,347.67 ppm sample concentration is
needed to inhibit 50% of free radicals. On the other
hand, plants treated with 0 g SP-36 tend to have
higher IC50 values, namely 1,602.8 ppm sample
concentration is needed to inhibit 50% of free radicals,
although it was not signicantly di󰀨erent. It can be
inferred that bitter gourds treated with 0 g SP-36 per
plant had lower antioxidant activity because the higher
the IC50 value, the lower the antioxidant activity
(Cruz et al., 2020). This value is calculated based on
the antioxidant concentration required to reduce the
DPPH concentration by 50% (Moreno et al., 1999).
However, antioxidant enzyme activity increases when
phosphate fertilizer is applied (Hekmati et al., 2023).
Conversely, low phosphorus availability will reduce
antioxidant enzyme activity (Kayoumu et al., 2023).
Increasing the dose of phosphorus to 60 g SP-
36 per plant tends to increase the total terpenoid
content compared to 0 g SP-36 per plant, although
it did not have a signicant e󰀨ect. Terpenoid
content and dry fruit weight per plant are the main
factors determining terpenoid production per plant.
Therefore, in this study, various SP-36 treatments
tended to have the potential to increase terpenoid
production because they had higher total terpenoid
and heavier fruit dry weight than 0 g SP-36 treatment.
However, it did not have a signicant e󰀨ect. This is
because phosphorus plays a vital role in secondary
metabolites because it is a constituent of nucleic
acids and phospholipids and plays an important role
in cell metabolism (Marschner 2002). One group
of secondary metabolites that require phosphorus
is terpenoids, terpenoid precursors including IPP
(Isopentenyl diphosphate), DMAPP (Dimethylallyl
pyrophosphate), GDP (Geranyl diphosphate), and
FDP (Farnesyl diphosphate) contain high-energy
phosphate bonds so that phosphate becomes an
element for producing terpenoid compounds and
expect to inuence terpenoid production (Li et al.,
2023). Terpenoid production in Rosmarinus ocinalis
is inuenced by phosphorus availability (Bustamante
et al., 2020). There is a positive inuence between
phosphorus fertilization on the terpenoid content in
Pinus halepensis under drought conditions, although
it is not signicantly di󰀨erent (Branch et al., 2009).
In addition, in the same species, there is a positive
and signicant e󰀨ect on the terpenoid content in
phosphorus fertilization (Ormeno et al., 2008).

Plant Growth, Fruit Production and Total Terpenoid Compounds in Bitter Gourd ..........
Journal of Tropical Crop Science Vol. 11 No. 2, June 2024
www.j-tropical-crops.com
191
Received 09/04/2024; Revised 25/06/2024; Accepted 27/06/2024
https://doi.org/10.29244/jtcs.11.02.184-194
Phosphorus is also involved in the synthesis of ATP
and NADPH where later these molecules are needed
to synthesize terpenoids via the mevalonic acid (MVA)
pathway and the methylerythritol phosphate (MEP)
pathway (Stirbet et al., 2020). Therefore, phosphorus
is an essential component in producing secondary
metabolites, especially terpenoid compounds.
Conclusion
The application of 40 g SP-36 per plant, in addition
to the basic fertilizer N and K, increased stem height
at 4 WAP, number of female owers, fresh weight
per fruit, fresh weight of fruit production per plant,
and leaf chlorophyll a, chlorophyll b, carotene, and
total chlorophyll. Bitter gourd plants without fertilizer,
with chicken manure, and fertilized with 20, 40, and
60 g SP-36 per plant produced similar amounts of
terpenoid, i.e., 6,791, 8,162, 11,630, 12,888, 15,995,
and 14,494 mmol NE per plant, respectively.
Acknowledgment
The authors thanked the Indonesia Endowment Fund
for Education Agency (LPDP) for funding the research
project “Plant and Production of Total Terpenoid
Compounds in Bitter Gourd (Momordica charantia L.)
at Various Phosphorus fertilization” in the 2023-2024
academic year.
References
Ashraf, I., Zain, M., Sarwar, M., Abbasi, K.Y., and
Amin, M. (2019). Role of di󰀨erent levels of
nitrogen, phosphorus, and potassium on
growth, yield and quality attributes of Bitter
gourd (Momordica charantia L.). Food Science
and Technology 7, 16-21. doi:10.13189/
fst.2019.070202
Bustamante, M.A., Michelozzi, M., Caracciolo, A.B.,
Grenni, P., Verbokkem, J., Geerdink, P., Sa,
C., and Nogues, I. (2020). E󰀨ects of soil
fertilization on terpenoid and other carbon-
based secondary metabolites in Rosamarinus
ocinalis plants: A comparative study. Plants
9, 1-19. https://doi.org/10.3390/plants9070830
[Balitttanah] Balai Penelitian Tanah. (2005). “Petunjuk
Teknis Analisis Kimia Tanah, Tanaman, Air dan
Pupuk”. http:/?balittanah.litbang.deptan.go.id
[March 13, 2024].
Carstensen, A., Herdean, A., Schmidt, S.B., Sharma,
A., Spetea, C., Pribil, Mathias., and Husted, S.
(2018). The impacts of phosphorus deciency
on the photosynthetic electron transport chain.
Plant Physiology 177, 271-284. https://doi.
org/10.1104/pp.17.01624
Choudhary, B.R., Fageria, M.S., and Dhaka, R.S.
(2013). Role of macro and micronutrients in
fruit production. Indian Journal of Horticulture
70, 474-478.
Cruz, J.F.R., Pineda, J.G., Chaverri, J.P., Rojas,
J.M.P., Passari, A.K., Ruiz, G.D., and Cruz,
B. E. R. (2020). Phytochemical constituents,
antioxidant, Cytotoxic, and antimicrobial
activities of the ethanolic extract of Mexican
brown propolis. Antioxidants 9, 1-11.
doi:10.3390/antiox9010070
Cuong, D.M., Jeon, J., Morgan, A.M.A., Kim, C.,
Kim, J.K., Lee, S.Y., and Park, S.U. (2017).
Accumulation of charantin and expression of
Tritepernoid Biosynthesis genes in bitter melon
(Momordica charantia L.). Journal Agriculture
and Food Chemistry 65, 7240-7249. https://
doi.org/10.1021/acs.jafc.7b01948
Deng, F., Li, W., Wang, Li., Hu, H., Liao, S., Pu,
S.L., Tao, Y.F., Li, G.H., and Ren, W.J. (2020).
E󰀨ect of controlled-release fertilizers on leaf
characteristics, grain yield, and nitrogen use
e󰀩ciency of machine transplanted rice in
southwest China. Archives of Agronomy and
Soil Science 67, 1739-1753. https://doi.org/1
0.1080/03650340.2020.1807519
Dey, G., Banerjee, P., Sharma, R.K., Maity, J. P.,
Etesami, H., Shaw, A. K., Huang, Y.H., Huang,
H.B., and Chen, C.Y. (2021). Management of
phosphorus in salinity-stressed agriculture
for sustainable crop production by salt-
tolerant phosphate-solubilizing bacteria.
Agronomy 11, 1-27. https://doi.org/10.3390/
agronomy11081552
Duan, Z., Xiao, H., Dong, Z., Li, X., and Wang, G.
(2004). The combined e󰀨ect of nitrogen,
phosphorus, potassium fertilizers and water
on spring wheat yield in an arid desert
region. Communication in Soil Science and
Plant Analysis 35, 161-175. http://dx.doi.
org/10.1081/CSS-120027641
Fageria, N.K., Gheyi, H.R., Carvalho, M.C.S., and
Moreira, A. (2016). Root growth, nutrient
uptake, and use e󰀩ciency by roots of

Journal of Tropical Crop Science Vol. 11 No. 2, June 2024
www.j-tropical-crops.com
192 Dian Novira Rizva, Maya Melati, Sandra Arin Aziz
Received 09/04/2024; Revised 25/06/2024; Accepted 27/06/2024
https://doi.org/10.29244/jtcs.11.02.184-194
tropical legume cover crops as inuenced by
phosphorus fertilization. Journal of Johan Plant
Nutrition 39, 781-792. https://doi.org/10.1080/0
1904167.2015.1088020
Hekmati, J., Hamidoghli, Y., Esmaielpour, B., and
Ghasemnezhad, M. (2023). The e󰀨ect of bio-
fertilizers and phosphorus on physiological
characteristics, antioxidant enzyme activity and
the essential oil content of green mint (Mentha
spicata L.) under arsenic stress conditions.
Agriculturae Conspectus Scienticus 88, 307-
315. https://hrcak.srce.hr/311919
Huang, H., Chen, F., Long, R., and Huang, G. (2020).
The antioxidant activities in vivo of bitter
gourd polysaccharide. International Journal
of Biological Macromolecules 145, 141-144.
https://doi.org/10.1016/j.ijbiomac.2019.12.165
Johan, P.D., Ahmed, O.H., Omar, Latifah., and
Hasbullah, N.A. (2021). Phosphorus
transformation in soil following Co-Application
of charcoal and wood ash. Agronomy 11, 1-25.
https://doi.org/10.3390/agronomy11102010
Kacprzak, M., Malinska, K., Grosser, A., Szoltysek,
J.S., Wystalska,K., Drozdz, D., Jasinka, A.,
and Meers, E. (2022). Cycles of carbon,
nitrogen and phosphorus in poultry manure
management technologies-environmental
aspects. Critical Reviews in Environmental
Science and Technology 8, 914-938. https://
doi.org/10.1080/10643389.2022.2096983
Kayoumu, M., Iqbal, A., Muhammad, N., Li, X.,
Li, L., Wang, X., Gui, H., Qi, Q., Ruan, S.,
Guo, R., Zhang, X., Song, M., and Dong, Q.
(2023). Phosphorus availability a󰀨ects the
photosynthesis and antioxidant system of
controlling low-P-tolerant cotton genotypes.
Antioxidants 12, 1-23. https://doi.org/10.3390/
antiox12020466
Kumari, S., Kumar P., Kiran, S., Kumari, S., and
Singh, A. (2023). Characterization of culture
condition dependent, growth responses of
phosphate solubilizing bacteria (Bacillus
subtilis DR2) on plant growth promotion of
Hordeum vulgare. Vegetos 37, 266-276. doi:
http://dx.doi.org/10.1007/s42535-023-00589-2
Lee, J.J., Yoon, K.Y. (2021). Optimization of ultrasound-
assisted extraction of phenolic compounds
from bitter melon (Momordica charantia) using
response surface methodology. Journal of
Food 19, 721-728. https://doi.org/10.1080/194
76337.2021.1973110
Lee, N.R., Kim, Y.X., Lee, Y., Lee, Chanwook., Song,
Y., Park, H., Lee, C.H., and Lee, Y. (2024).
Metabolomics reveals the e󰀨ects of nitrogen/
phosphorus/potassium (NPK) Fertilizer
levels on cucumber fruit raised in di󰀨erent
nutrient soils. Metabolites 14, 1-12. https://doi.
org/10.3390/metabo14020102
Li, C., Zha, W., Li, W., Wang, J., and You, A. (2023).
Advances in the biosynthesis of terpenoids and
their ecological functions in plant resistance.
International Journal of Molecular Sciences 24,
1-16. https://doi.org/10.3390/ijms241411561
Li, D., Li, X.,Han, Q., Zhou, Y., Dong, J., and Duan, Z.
(2020). Phosphorus application improved the
yield of citrus plants grown for three years in
acid soil in the Three Gorges Reservoir area.
Scientia Horticulturae 273, 1-7. https://doi.
org/10.1016/j.scienta.2020.109596
Li, X., Shang, J. (2021). Decision-making
behavior of fertilizer application of grain
growers in Heilongjiang Province from the
perspective of risk preference and risk
perception. Hindawi 2021, 1-8. https://doi.
org/10.1155/2021/6667558
Li, Z., Zhang, R., Xia, S., Wang, L., Liu, C., Zhang, R.,
Fan, Z., Chen, F., and Liu, Y. (2019). Interaction
between N, P, and K fertilizers a󰀨ects the
environment and the yield and quality of
satsumas. Global Ecology and Conservation
19, 1-13. https://doi.org/10.1016/j.gecco.2019.
e00663
Lima, J.R.D.S., Goes, M.D.C.C.D., Hammecker, C.,
Antonino, A.C.D., Medeiros, E.V., Sampaio,
E.V.D.S.B., Leite, M.C.D.B.S., Silva, V.P.D.S.,
Souza, E.S.D., and Souza, R. (2021). E󰀨ect
of poultry manure and biochar on acrisol soil
properties and yield of common bean. A short-
term eld experiment. Agriculture 11, 1-11.
https://doi.org/10.3390/agriculture11040290
Lukowski, A., Jagiello, R., Robakowski, P., Adamczyk,
D., and Karolewski, P. (2022). Adaptation of
simple method to determine the total terpenoid
content in needles of coniferous trees. Plant
Science 314, 1-9. Https://doi.org/10.1016/j/
plantsci.2021.111090
Mahwish., Saeed, F., Sultan, M.T., Riaz, A., Ahmed, S.,
Bigiu, N., Amarowicz., and Manea, R. (2021).
Bitter melon (Momordica charantia L.) fruit
bio-actives charantin and vicine potential for
diabetes prophylaxis and treatment. Plants 10,
1-13. https://doi.org/10.3390/plants10040730

Plant Growth, Fruit Production and Total Terpenoid Compounds in Bitter Gourd ..........
Journal of Tropical Crop Science Vol. 11 No. 2, June 2024
www.j-tropical-crops.com
193
Received 09/04/2024; Revised 25/06/2024; Accepted 27/06/2024
https://doi.org/10.29244/jtcs.11.02.184-194
Malhotra, H., Sharma, S., and Pandey, R. (2018).
Phosphorus nutrition: plant growth in response
to deciency and excess. In Plant Nutrients and
Abiotic Stress Tolerance pp. 171-190. Springer,
Singapore https://doi.org/10.1007/978-981-10-
9044-8_7
Maluki, M., Gesimba, R.M., and Ogweno, J.O. (2016).
The e󰀨ect of di󰀨erent phosphorous levels
on yield and quality of Watermelon (Citrullus
lanatus (Thunb.) Matsumura and Nakai grew in
the Kenyan Coastal region. Annals of Biological
Research 7, 12-17.
Martin, J. D.L., Soratto, R.P., Fernandes, A.M.,
and Dias P.H.M. (2018). Phosphorus
fertilization and soil texture a󰀨ect potato yield.
Universidade Federal Rural do Semi-Arido
31, 541-550. https://doi.org/10.1590/1983-
21252018v31n302rc
Marschner, H. (2012). Mineral Nutrition of Higher
Plants. Academic Press is an imprint of
Elsevier. http://dx.doi.org/10.1016/B978-0-12-
384905-2.00006-6
Miguel, J., Melati, M., and Aziz, S.A. (2023).
Determination of chicken manure plant for
bitter gourd production organically. Prosiding
Seminar Nasional PERHORTI 37-46.
Moreno, C.S., Larrauri, J.A., and Calixto, F. S.
(1999). A procedure to measure the antiradical
e󰀩ciency of polyphenols. Journal Science Food
Chemistry 76, 270-276. https://doi.org/10.1002/
(SICI)1097-0010(199802)76:2<270AID-
JSFA945>3.0.CO;2-9
Mu, X., Chen, Q., Chen, F., Yuan, L., and Mi, G. (2016).
Within-leaf nitrogen allocation in adaptation to
low nitrogen supply in maize during the grain-
lling stage. Frontiers in Plant Science 7, 1-11.
doi: 10.3389/fpls.2016.00699
Naorem, J., Dakho, J. (2019). E󰀨ect of phosphorus
and potassium on yield and quality
characters of bitter gourd (Momordica
charantia L.) ecotype Mithipagal. Journal
of Pharmacognosy and Phytochemistry
8, 1686-1688. https://doi.org/10.15740/
HAS%2FTAJH%2F10.2%2F207-211
Nell, M., Votsch, M., Vierheilig., Steinkellner, S.,
Zitterl-Eglseer., Franz, C., and Novak, J.
(2009). E󰀨ect of phosphorus uptake on growth
and secondary metabolites of garden sage
(Salvia ocinalis L.). Journal of the Science of
Food and Agriculture 89, 1090-1096. https://
doi.org/10.1002/jsfa.3561
Neocleous, D., Savvas, D. (2019). The e󰀨ect
of phosphorus supply limitation on
photosynthesis, biomass production,
nutritional quality, and mineral nutrition in
lettuce grown in recirculating nutrient solution.
Scientia Horticulturae 252, 379-387. https://
doi.org/10.1016/j.scienta.2019.04.007
Ormeno, E., Baldy, V., Ballini, C., and Fernandez,
C. (2008). Production and diversity of volatile
terpenes from plants on calcareous and
siliceous soils: e󰀨ect of soil nutrients. Journal of
Chemical Ecology 34, 1219-1229. doi:10.1007/
s10886-008-9515-2
Paul, A., Mitter, K., Sen., and Raychaudhuri, S.
(2009). E󰀨ect of polyamines in in vitro somatic
embryogenesis in Momordica charantia L.
Plant cell tissue organ culture 97, 303-311.
http://dx.doi.org/10.1007/s11240-009-9529-7
Peng, C.L., Teik, L. (2022). Impacts of nitrogen and
phosphorus fertilization on biomass polyphenol
contents and essential oil yield and composition
of Vitex negundo Linn. Agriculture 12, 1-13.
https://doi.org/10.3390/agriculture12060859
Purba, T., Situmeang, R., Mahyati. H.F.R., Arsi,
Firgiyanto. R., Saadah, A.S.J.T.T., Herawati,
J.J., and Suhastyo, A.A. (2021). Pupuk dan
Teknologi Pemupukan. Yayasan Kita Menulis.
Sumatera Utara.
Razaq, M., Zhang, P., Shen, H.L., and Salahuddin.
(2017). Inuence of nitrogen and phosphorous
on the growth and root morphology of
Acer mono. Plos One 12,1-13. https://doi.
org/10.1371/journal.pone.0171321
Sahu, J.K., Tiwari, A., and Lakpale R. (2021). E󰀨ect
of nitrogen, phosphorus, and potassium on
growth and owering of chrysanthemum. The
Pharma Innovation Journal 10, 1289-1292.
Salazar, A.R., Alejandro, R.L.L., Lopez, A.J.,
Alicia, A.G.B., and Waksman, T.N. (2011).
Antimicrobial and antioxidant activities
of plants from northeast of Mexico. Evid
Based Complement Altern Med 2011, 1-6.
Doi:10.1093/ecam/nep127
Sims, D.A., Gamon, J. (2002). Relationships between
leaf pigment content and spectral reectance
across a wide range of species, leaf structures

Journal of Tropical Crop Science Vol. 11 No. 2, June 2024
www.j-tropical-crops.com
194 Dian Novira Rizva, Maya Melati, Sandra Arin Aziz
Received 09/04/2024; Revised 25/06/2024; Accepted 27/06/2024
https://doi.org/10.29244/jtcs.11.02.184-194
and developmental stages. Remote Sensing
of Environment 81, 337-354. Doi: http://dx.doi.
org/10.1016/S0034-4257(02)00010-X
Slamet. (2020). The e󰀨ect of Bitter melon leaves
(Momordica charantia L.) ethanol extract on the
Trichophyton rubrum fungal inhibiting zone of
di󰀨usion method. Jurnal Teknologi Kesehatan
Borneo 1, 1-8. https://doi.org/10.30602/jtkb.
v1i1.4
Stirbet, A., Lazar, D., Guo, Y., and Govindjee, G.
(2020). Part of a special issue on functional
structural plant growth modeling. Annals
of Botany 126, 511-537. doi: 10.1093/aob/
mcz171
Sustr, M., Soukup, A., and Tylova, E. 2019. Pottasium
in Root Growth and Development. Plants 8,
1-16. doi:10.3390/plants8100435
Thornburg, T.E., Liu, J., Li, Q., Xue, H., Wang, G.,
LI, L., Fontana, J.E., Davis, K.E., Liu, W.,
Zhang, B., Zhang, Z., Liu, M., and Pan, X.
(2020). Potassium deciency signicantly
a󰀨ected plant growth and development as
well as a microRNA-mediates mechanism in
wheat (Triticum aestivum L.). Frontiers in Plant
Science 11, 1-10. https://doi.org/10.3389/
fpls.2020.010219
Tuan, N.Q., Lee, D.H., Oh, J., Kim, C.H., Heo, K.S.,
Myung, C.S., and Na, M. (2017). Inhibition
of proliferation of vascular muscle cells by
cucurbitanes from Momordica charantia.
Journal of Natural Products 80, 2018-2025.
https://doi.org/10.1021/acs.jnatprod.7b00151
Vaudo, A.D., Erickson, E., Patch, H.M., Grozinger,
C.M., and Mu, J. (2022). Impacts of soil nutrition
on oral traits, pollinator attraction, and tness
in cucumbers (Cucumis sativus L.). Scientic
Reports 12,1-12. doi: 10.1038/s41598-022-
26164-4
Wang, Y., Wang, F., Lu, H., Liu, Y., and Mao, C.
(2021). Phosphate uptake and transport in
plants: an elaborate regulatory system. Plant
Cell Physiology 62, 564-572. doi: 10.1093/pcp/
pcab011
Wardana, A., Boceng, A., Haris, A., Ashar, J.R.,
and Gani, M.S. (2020). Pengaruh pemberian
berbagai jenis pupuk kendang terhadap
pertumbuhan dan produksi tanaman pare
(Momordica charantia L.). Jurnal Agrotek
Mahasiswa 1, 1-8
Weng, J.R., Bai, L. Y., Ciu, C.F., Hu, J. L., Chiu, S.J.,
and Wu, C. Y. (2013). Curcubitane triterpenoid
from Momordica charantia induces apoptosis
and autophagy in breast cancer cells, in part,
through peroxisome proliferator-activated
receptor y activation. Evidence Based
Complementary and Alternative Medicine
2013, 1-12
Xia, Z., Zhang, S., Wang, Q., Zhang, G., Fu, Y., and
Lu, H. (2021). E󰀨ect of root zone warming on
maize seedling growth and photosynthetic
characteristics under di󰀨erent phosphorus
levels. Frontiers in Plant Science 12, 1-11.
doi:10.3389/fpls.2021.746152
Yaldiz, G., Sekeroglu, N., Kulak, M., and Demirkol, G.
(2015). Antimicrobial activity and agricultural
properties of Bitter melon (Momordica charantia
L.) grown in northern parts of Turkey: a case
study for adaptation. Journal Natural Product
Research 29, 543-545. http://dx.doi.org/10.10
80/14786419.2014.949706
Yang, Li., Wen, K.S., Ruan, X., Zhao, Y.X., Wei,
F., and Wang, Q. (2018). Response of plant
secondary metabolites to environmental
factors. Molecules 23, 1-26. doi:10.3390/
molecules23040762
Zhao, J., Davis, L.C., and Verpoorte, R. (2005).
Elicitor signal transduction leads to the
production of plant secondary metabolites.
Biotechnology Advances 23, 283-333. https://
doi.org/10.1016/j.biotechadv.2005.01.003