© The Authors, 2026, Published by the Universidad del Zulia*Corresponding authorgeorge.cedeno@utm.edu.ecc
Keywords:
Biochar
Compost
Chlorophyll uorescence
Mineral nutrition
Organic matter
Chlorophyll uorescence and leaf nutritional status of passion fruit (Passiora edulis f.
avicarpa) seedlings grown in dierent organic substrates
Fluorescencia de la clorola y estado nutricional foliar de plántulas de maracuyá (Passiora edulis
f. avicarpa) cultivadas en distintos sustratos orgánicos
Fluorescência da clorola e estado nutricional foliar de mudas de maracujá (Passiora edulis f.
avicarpa) cultivadas em diferentes substratos orgânicos
Luís Angel Giler Cool
2
Adriana Celi Soto
1
Galo Cedeño García
1
George Cedeño-García
1*
Rev. Fac. Agron. (LUZ). 2026, 43(1): e264315
ISSN 2477-9407
DOI: https://doi.org/10.47280/RevFacAgron(LUZ).v43.n1.XV
Crop production
Associate editor: Dr. Jorge Vilchez-Perozo
University of Zulia, Faculty of Agronomy
Bolivarian Republic of Venezuela
1
Universidad Técnica de Manabí, Facultad de ingeniería
agroambientales, km 13,5 vía Portoviejo – Santa Ana.
2
Estudiante de posgrado,
Universidad Técnica de Manabí,
Facultad de ingeniería agroambientales, km 13,5 vía
Portoviejo – Santa Ana.
Received: 30-10-2025
Accepted: 26-01-2026
Published: 22-02-2026
ABSTRACT
Passion fruit (Passiora edulis f. avicarpa) has high economic
and nutritional value, and the nursery stage is crucial for its early
development. The objective of the present study was to evaluate
chlorophyll uorescence, foliar nutritional status, and growth of
passion fruit seedlings in response to four organic substrates: S1
(sand + biochar + organic matter), S2 (sand + compost + organic
matter), S3 (sand + compost + agricultural soil), and S4 (agricultural
soil), under a completely randomized design. The physiological
variables F
v
/F
m
, Φ(II), and ETR were measured, as well as the foliar
concentration of macro and micronutrients and growth variables,
whose data were analyzed using ANOVA and Tukey’s test (5 %).
Plants grown in substrate S2 showed the highest photochemical
eciency (F
v
/F
m
≈ 0.79; Φ(II) ≈ 0.44; ETR ≈ 160), whereas those in
substrate S1 exhibited the highest foliar concentrations of K, Fe, and
Zn. The principal component analysis explained 71 % of the total
variability, associating substrate S2 with better physiological yield
and growth. It can be concluded that substrates containing compost
enhance PSII photochemical eciency, while those including
biochar are associated with greater foliar mineral accumulation.
This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.
Rev. Fac. Agron. (LUZ). 2026, 43(1): e264315 January-March ISSN 2477-9409.
2-7 |
Resumen
El maracuyá (Passiora edulis f. avicarpa) presenta un alto
valor económico y nutricional, y la fase de vivero es determinante
para su desarrollo inicial. El presente trabajo tuvo como objetivo
evaluar la uorescencia de la clorola, el estado nutricional foliar
y el crecimiento de plántulas de maracuyá en función de cuatro
sustratos orgánicos: S1 (arena + biocarbón + materia orgánica), S2
(arena + compost + materia orgánica), S3 (arena + compost + tierra
agrícola) y S4 (tierra agrícola), bajo un diseño completamente al
azar. Se midieron las variables siológicas Fv/Fm, Φ(II) y ETR, así
como la concentración foliar de macro y micronutrientes y variables
de crecimiento, cuyos datos fueron analizados mediante ANOVA
y prueba de Tukey (5 %). Las plantas cultivadas en el sustrato S2
presentaron la mayor eciencia fotoquímica (F
v
/F
m
≈ 0,79; Φ(II)
≈ 0,44; ETR ≈ 160), mientras que las cultivadas en el sustrato S1
mostraron las mayores concentraciones foliares de K, Fe y Zn. El
análisis de componentes principales explicó el 71 % de la variabilidad
total, asociando el sustrato S2 con un mejor desempeño siológico
y de crecimiento. Se puede concluir que los sustratos con compost
favorecen la eciencia fotoquímica del PSII, mientras que aquellos
que incluyen biocarbón se asocian con una mayor acumulación foliar
de minerales.
Palabras clave: biocarbon, compost, uorescencia de clorola,
nutrición mineral, materia orgánica.
Resumo
O maracujá (Passiora edulis f. avicarpa) apresenta elevado
valor econômico e nutricional, e a fase de viveiro é determinante para
o seu desenvolvimento inicial. O objetivo deste estudo foi avaliar a
uorescência da clorola, o estado nutricional foliar e o crescimento
de mudas de maracujá em função de quatro substratos orgânicos:
S1 (areia + biocarvão + matéria orgânica), S2 (areia + composto +
matéria orgânica), S3 (areia + composto + solo agrícola) e S4 (solo
agrícola), sob um delineamento inteiramente casualizado. Foram
avaliadas as variáveis siológicas Fv/Fm, Φ(II) e ETR, bem como
as concentrações foliares de macro e micronutrientes e variáveis de
crescimento, cujos dados foram analisados por ANOVA e teste de
Tukey (5 %). As mudas cultivadas em substratos S2 apresentaram
a maior eciência fotoquímica (Fv/Fm ≈ 0,79; Φ(II) ≈ 0,44; ETR ≈
160), enquanto substratos S1 apresentou as maiores concentrações
foliares de K, Fe e Zn. A análise de componentes principais explicou
71 % da variabilidade total, associando o substrato S2 a melhor
desempenho siológico e de crescimento. Conclui-se que substratos
contendo composto favorecem a eciência fotoquímica do PSII,
enquanto aqueles que incluem biocarvão estão associados a maior
acúmulo foliar de minerais
Palavras-chave: biocarvão, composto, uorescência da clorola,
nutrição mineral, matéria orgânica.
Introduction
In Ecuador, passion fruit has high economic relevance, with an
annual production of approximately 48,000 tons according to ocial
reports from the National Institute of Statistics and Censuses (INEC,
2023), and constitutes one of the main fruits destined for export
(MAG, 2020). The quality of the plant material, as well as the vigor
and yield in the eld, are directly associated with nursery management
(Santos et al., 2020).
Substrate selection is decisive for the initial growth and quality of
seedlings (Bonifácio et al., 2025; Lessa et al., 2023). Substrates with
low fertility reduce nutrient availability and root expansion, aecting
the initial vigor (Paixão et al., 2021). On the contrary, the incorporation
of compost and biochar improves porosity, water holding capacity, and
nutrient availability, favoring early development in the nursery (Yang
et al., 2023; Wu et al., 2023). From the physiological approach, the
photosynthetic eciency evaluated through the maximum quantum
eciency of photosystem II (F
v
/Fm), maximum quantum yield of
photosystem II (Φ[II]), electron transport rate (ETR), and chlorophyll
index, is a key indicator of plant yield (Ni et al., 2020). Organic
substrates such as compost and biochar increase these parameters
by improving light use and energy generation, favoring the growth
and productivity of passion fruit (Silva et al., 2020). Chlorophyll
concentration is directly related to photosynthetic eciency and
potential yield of seedlings, making it a key physiological indicator in
the nursery stage (Sun et al., 2025; Su et al., 2024). The application of
organic matter and renewable substrates has been shown to increase
chlorophyll, PSII eciency, and leaf expansion in Passioraceae (Da
Silva et al., 2023; Bonifácio et al., 2025). Despite these advances,
it is unknown how the combinations of organic substrates used
with locally available material and accessible to producers modify
the photochemical responses of chlorophyll uorescence and foliar
nutritional concentration of P. edulis. The objective of the study
was to evaluate chlorophyll uorescence, nutritional status, and the
growth of Passiora edulis seedlings grown in dierent organic
substrates during the nursery stage.
Materials and methods
The research was carried out in the agro-industrial crop nursery
of the Faculty of Agronomic Engineering of the Technical University
of Manabí, located in the canton of Santa Ana, Manabí, Ecuador. The
site is located at an altitude of 60 m.a.s.l, with an average annual
temperature of 26 °C, a rainfall of 625 mm, and a relative humidity that
varies between 74.8 mm (dry season) and 82.48 mm (rainy season).
The experiment was developed under a Completely Randomized
Design (CRD) with four types of substrates as treatments (detailed
in Table 1) and four replications, each consisting of 25 plastic bags
per experimental unit. For the test, 400 black polyethylene plastic
bags of 6 × 8 inches, with an approximate volume of 2.36 L, were
used. The substrates were formulated using a mixture of agricultural
soil, washed river sand, compost, and organic matter in a ratio of
1:1:1:1.5, to which biochar from Ceratonia siligua was added in a
ratio of 1:0.25.
Table 1. Formulation of substrates based on combinations of
organic materials.
Treatments Composition
Substrate 1 Sand + Biochar + Organic matter
Substrate 2 Sand + Compost + Organic matter
Substrate 3 Sand + Compost + Agricultural soil
Substrate 4 Agricultural soil
This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.
Giler et al. Rev. Fac. Agron. (LUZ). 2026, 43(1): e264315
3-7 |
Photochemical evaluations, chlorophyll index, and growth
variables (plant height, stem diameter, and leaf area) were performed
The physicochemical characteristics of the substrates are
presented in Tables 2 and 3. The materials were pre-composted
for three months before sowing. Two certied passion fruit seeds,
Passiora edulis f. avicarpa (INIAP-2009), treated with Thiodicarb
+ Imidacloprid (systemic insecticides; i.a) at a dose of 25 mL.kg
-1
seed, were placed in bags with substrate previously treated with
fungicide Captan 80 WP (180 g.100 L
-1
). After 15 days, thinning was
performed, leaving one seedling per experimental unit. Irrigation was
applied through micro-sprinklers with an 8 mm depth twice a week.
Agronomic management included manual weeding and preventive
applications of an insecticide, fosetyl-aluminum (2 g.L
-1
) and a
fungicide, Spinetoram (0.12 mL.L
-1
).
Table 2. Chemical composition of carob biochar, washed river sand, and compost.
Chemical Composition
Carob biochar % River sand % Compost %
pH 8.7 Silica (SiO
2
/quartz) 90 pH 7.37
EC (dS.m
-1
) 1.49 Metal oxides (Fe
2
O
3
, Al
2
O
3
, etc.) 2 EC (dS.m
-1
) 3.5
CEC cmol
+
.kg 15.7 Humidity 34.45
H/C 0.11 Other minerals (Feldspar, calcite,
clay, etc.) 5
Organic Matter 53.65
O/C 0.13 C/N ratio 12.97
C 71.17 Ashes 18.75
H 2.96 Total Nitrogen 1.2
O 23.86 Total Phosphorus 0.3
N 1.3 Total Potassium 0.79
S 0.37
Humidity 2.7
Ashes 0.32
Fixed carbon 29.64
Volatile matter 67.47
Table 3. Physical-chemical composition of agricultural soil and organic matter used in the mixture of substrates.
Soil chemical composition Agricultural soil Organic matter
Organic Matter (%) 4.2 32.5
Texture (%) Clay loam – silty -
Base saturation (%) 74 62
CIC (meq.100g) 28.9 45.3
EC (mS.cm) 0.15 1.82
pH (H
2
O) 6.6 7.1
Nitrate (NO
3
-N) (mg.kg
-1
) 7.4 215
Ammonium (NH
4
-N) (mg.kg
-1
) 2.1 96
Total N (NO
3
+NH
4
) (mg.kg
-1
) 9.5 311
Phosphorus (P) (mg.kg
-1
) 38.8 412
Potassium (K) (mg.kg
-1
) 228 1450
Magnesium (Mg) (mg.kg
-1
) 259 520
Calcium (Ca) (mg.kg
-1
) 845 1820
Sulfur (S) (mg.kg
-1
) 8.3 112
Iron (Fe) (mg.kg
-1
) 22.2 310
Manganese (Mn) (mg.kg
-1
) 12.8 145
Copper (Cu) (mg.kg
-1
) 3.3 21
Zinc (Zn) (mg.kg
-1
) 2.3 38
Boron (B) (mg.kg
-1
) 0.28 1.9
Sodium (Na) (mg.kg
-1
) 12 89
Chloride (Cl) (mg.kg
-1
) 9 165
eight weeks after sowing. Chlorophyll uorescence (F
v
/F
m
, Φ(II), and
ETR) and chlorophyll index were determined in adult leaves exposed
to full luminosity, using an OS-1p+ uorometer (Opti-Sciences) and
a SPAD 502 chlorophyll meter, respectively. For the determination of
F
v
/F
m
, the leaves were previously adapted to the dark for 30 min. Plant
height was measured with a exometer from the base of the stem to
the insertion of the last fully expanded leaf. The diameter of the stem
was determined with a manual (Vernier) caliper, and the leaf area was
estimated using the gravimetric method. For this purpose, ten leaf
discs of a known area were collected using a punch; these were then
dried to a constant weight and used as a reference to calculate the total
leaf area per plant, with values expressed in cm² per plant. The foliar
concentration of macro and micronutrients was quantied in samples
of 200 g per experimental unit, following INIAP protocols.
This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.
Rev. Fac. Agron. (LUZ). 2026, 43(1): e264315 January-March ISSN 2477-9409.
4-7 |
The data were subjected to analysis of variance (ANOVA; α =
0.05) and comparison of means through Tukey’s test (5 %), using
the InfoStat 8.0 software (2018). Additionally, a principal component
analysis (PCA) was performed to identify multivariate patterns
between physiological, nutritional, and growth variables.
Results and discussion
Nutrient concentration in dierent substrates
The foliar concentration of N was higher in substrates S1 and S3,
while substrate S2 showed the lowest accumulation. This variation
is explained by the ability of biochar to retain N in substrate S1 and
by the gradual release of organic N from the compost in substrate S3,
which coincides with studies that describe the importance of biochar
in ion retention and the tendency of Passiora to present suboptimal
foliar levels of N (Antunes et al., 2022; Hauer-Jákli & Tränkner,
2022; de Lima et al., 2023). Substrates S1 and S2 showed the highest
concentrations of P, mainly associated with the contribution of organic
matter, which has higher concentrations of phosphorus and a higher
fraction available for absorption by the plant, compared to agricultural
soil. This trend is consistent with studies that point to an increase in P
availability and acid phosphatase activity in substrates enriched with
organic materials (Antunes et al., 2022; Yang et al., 2023).
K reached its maximum in substrate S1, which is consistent
with the compost-biochar synergy that improves monovalent cation
retention and reduces leaching; in addition, P. edulis has a high demand
for this nutrient (Antunes et al., 2022; de Lima et al., 2023). Finally, the
accumulation of Ca and Mg was higher in substrates with agricultural
soil and compost (S3 and S4), in accordance with reports that highlight
that organic matter increases the solubility and availability of these
divalent cations in tropical soils (Paixão et al., 2021).
The growth assessment showed that substrate S2 (sand + compost
+ agricultural soil) generated the most vigorous seedlings (higher
height, diameter, and leaf area), as the compost optimized the
nutrition and physical structure of the substrate (Figure 1), a result
that is consistent with recent studies in P. edulis (Da Silva et al., 2023;
Bonifácio et al., 2025). Substrates S1 and S3 oered intermediate
improvements, while substrate S4 (unamended soil) resulted in the
poorest growth due to its nutritional and physical limitations (Paixão
et al., 2021). The best foliar development in substrate S2 aligns
with the reported relationship between organic matter and improved
photosynthetic eciency in Passioraceae (Sun et al., 2025; Su et al.,
2024), making the use of compost key to improving the quality of P.
edulis seedlings in the nursery stage.
On the other hand, sulfur (S) (Figure 2E) presented its highest
concentration in substrate S1 (≈ 490 mg.100 kg⁻¹), attributed to carob
biochar, which contains 0.37 % S and volatile compounds that act
as a slow-release reservoir. Unlike soluble sulfate forms, biochar
improves the retention and availability of the element, showing how
organic amendments modify the dynamics of S and its interaction
with other nutrients (Freire et al., 2022; Antunes et al., 2022).
Figure 2. Foliar nutrient concentration of Passiora edulis f.
avicarpa grown in four substrates. S1 = biochar + river
sand; S2 = river sand + agricultural soil; S3 = compost
+ agricultural soil; S4 = biochar + compost; where. A)
Nitrogen, B) Phosphorus, C) Potassium, D) Calcium, E)
Sulfur, and F) Magnesium. Dierent letters in the variables
dier statistically according to Tukey’s test (p≤0.05).
Figure 1. Growth variables of passion fruit seedlings (Passiora edulis f. avicarpa) grown in four organic substrates during the nursery
stage. A) Seedling height (cm); B) Stem diameter (mm); C) Leaf area (cm²). Values represent means ± standard deviation.
This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.
Giler et al. Rev. Fac. Agron. (LUZ). 2026, 43(1): e264315
5-7 |
Foliar micronutrient concentration varied signicantly between
treatments (Table 4). Substrate S1 presented the highest values of
Zn (0.53 mg.g
-1
), Cu (0.13 mg.g
-1
), and Fe (2.48 mg.g
-1
), reecting
a greater availability and absorption of these elements, favored by
the combined action of biochar and compost. Similar results were
reported by Da Silva et al. (2023) in Passiora and other horticultural
crops. Instead, Mn reached its maximum concentration in substrate
S4 (0, mg.g
-1
), possibly due to the higher acidity of the medium.
Boron (B) showed no signicant dierences between treatments
(~0.04 mg.g
-1
), in agreement with Silva et al. (2020), who highlighted
the stability of this element in substrates with high organic content.
The micronutrient analysis (Table 4) showed a clear inuence of
the type of substrate. The substrate with carob biochar (S1) recorded
the highest concentrations of Zn (≈0.53 mg.g⁻¹) and Fe (≈2.48
mg.g⁻¹), signicantly exceeding S4 (Zn ≈0.29 mg.g⁻¹). This behavior
is related to the higher cation exchange and ion retention capacity
generated by biochar in organic mixtures (Yang et al., 2023).
The rest of the micronutrients presented dierentiated responses:
Mn reached its highest value in substrate S4 (≈0.28 mg.g⁻¹), while
Cu showed its highest accumulation in substrate S1 (≈0.13 mg.g⁻¹).
The lower availability of Mn in biochar-based substrates is consistent
with studies reporting immobilization of Mn and Cu by pH increases
and surface adsorption of biochar (Zhou et al 2023). Regarding B,
no dierences were detected between treatments, indicating that the
eect of biochar is not uniform between micronutrients.
The uorescence of chlorophyll (Figure 3) showed clear dierences
in the photochemical eciency of PSII in the plants grown in the dierent
substrates evaluated. Plants grown on substrate S2 showed the highest
values of F
v
/Fm (~0.79), Φ(II) (~0.44), and ETR (~160), indicating a
stable PSII, with ecient electron transport and low photoinhibition, in
response to the physical and nutritional conditions of the substrate (Da
Silva et al., 2023). In contrast, plants grown in S4 had the lowest values
(F
v
/Fm ~0.70; Φ(II) ~0.23; ETR < 100), reecting marked physiological
stress associated with reduced aeration or water imbalance. The F
v
/Fm
values < 0.75 are associated with damage to reaction centers and reduced
energy dissipation (Faria et al., 2020).
Substrates S1 and S3 promoted intermediate responses,
evidencing adequate photochemical eciency. Together, these
results conrm that the chemical and physical characteristics of the
substrate modulate the photosynthetic eciency of plants. Previous
studies in passion fruit agree that substrates with higher organic
matter content and better water holding capacity favor uorescence
and photochemical eciency (Sun et al., 2025), while abiotic stress
conditions, such as water decit or salinity, signicantly reduce Φ(II)
and ETR (Tomaškinová et al., 2025).
Table 4. Foliar concentration of zinc, copper, iron, manganese, and boron in Passiora edulis f. avicarpa, under the eect of organic
substrates with dierent physical mixtures.
Substrates Zn Cu Fe Mn B
mg.g
-1
M* D.E** M D.E M D.E M D.E M D.E
Substrate 1 0.53 ± 0.02
a
0.13 ± 0.005
a
2.48 ± 0.18
a
0.23 ± 0.005
b
0.36 ± 0.042
ns
Substrate 2 0.35 ± 0.04
b
0.09 ± 0.01
b
1.95 ± 0.11
bc
0.23 ± 0.025
b
0.38 ± 0.062
Substrate 3 0.33 ± 0.005
b
0.11 ± 0.005
b
1.78 ± 0.13
c
0.25 ± 0.015
ab
0.42 ±.047
Substrate 4 0.29 ± 0.009
b
0.11 ± 0.005
b
2.28 ± 0.01
ab
0.28 ± 0.044
a
0.43 ±0.02
*Means; ** Standard deviation
Figure 3. Chlorophyll uorescence parameters in Passiora edulis
f. avicarpa seedlings grown in four substrates. S1 =
biochar + river sand; S2 = river sand + agricultural soil;
S3 = compost + agricultural soil; S4 = biochar + compost,
where Fv/Fm = maximum PSII eciency, Φ(II) = eective
quantum yield of PSII, and ETR = electron transport rate.
Principal component analysis (PCA) explained 71 % of the total
variability (PC1 = 51.06 %; PC2 = 20.39 %), clearly dierentiating the
substrates (Figure 4). Substrate S2 was associated with higher values
of F
v
/F
m
, Φ(II), and ETR, reecting high photosynthetic eciency
and stable electron transport, in agreement with Zhang et al. (2023),
who reported that adequate nutrition and light management increase
photosynthetic yield in P. edulis (Figure 4). In contrast, substrate S4 was
located at the negative end of PC1, linked to nutritional deciencies;
Zn or P deciency reduces F
v
/F
m
and ETR (Zhang et al., 2023), and the
low availability of N under salinity limits pigments and physiological
eciency (Tomaškinová et al., 2025). Substrate S3 was related to Ca, Mg,
and Mn, essential elements for thylakoid stability and enzyme activity, as
observed by Lessa et al. (2023) in passion fruit grown in substrates with
carbonized rice husk. Substrate S1, located in the positive quadrant of
PC1, showed a yield conditioned by its physical properties. Under abiotic
stress, Φ(II) and ETR decrease before F
v
/F
m
in triploid P. edulis (Su et al.,
2024), which conrms that an adequate water and nutritional balance, as
in substrate S2, optimizes photosynthetic eciency.
Correlation analysis (Figure 5) revealed that N, P, and K were
positively associated with photochemical yield, particularly with ETR,
reecting their essential role in the synthesis of PSII components,
energy ow, and stomatal regulation (Zhou et al., 2023). In contrast,
Mg and Mn showed weak or negative correlations: Mg could generate
cationic interferences without reaching toxic levels (Hauer-Jákli &
Tränkner, 2019), while high Mn values are related to oxidative stress
and PSII impairment (Millaleo et al., 2020). Among micronutrients,
Fe, Zn, and S showed positive correlations with photochemical
parameters, consistent with their role in electron transport, enzyme
stability, and maintenance of redox homeostasis (Saleem et al., 2022).
This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.
Rev. Fac. Agron. (LUZ). 2026, 43(1): e264315 January-March ISSN 2477-9409.
6-7 |
Figure 4. Principal component analysis (PCA) of physiological and
nutritional variables of Passiora edulis f. avicarpa
seedlings grown in four substrates. S1 = biochar + river
sand; S2 = river sand + agricultural soil; S3 = compost +
agricultural soil; S4 = biochar + compost—showing the
grouping and relationships among the evaluated variables.
Figure 5. Correlation matrix between foliar nutrients (N, P, K,
Ca, Mg, S, Zn, Cu, Fe, Mn, and B) and chlorophyll
uorescence parameters (F
v
/F
m
, Φ(II), and ETR) in
Passiora edulis seedlings grown in dierent organic
substrates.
Conclusions
The physiological response of Passiora edulis depended on
the nutritional balance provided by the substrates. High contents
of N, P, and K in the substrate improved photochemical eciency
and PSII functionality, whereas Mg and Mn imbalances reduced
F
v
/F
m
, Φ(II), and ETR in seedlings. Overall, the results show that
balanced nutritional management in organic substrates is essential
for maintaining seedlings with high photochemical eciency and
promoting more sustainable passion fruit production systems.
Acknowledgment
The authors thank the Fondo de Investigación de
Agrobiodiversidad, Semillas y Agricultura Sustentable (FIASA)
for fully funding this research through the project ‘‘Generación de
tecnologías climáticamente inteligentes para potenciar la agricultura
de secano en Manabí”, code FIASA-CA-2023-002, conducted in the
province of Manabí, Ecuador.
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