Revista de la Facultad de Ciencias
Agrarias. Universidad Nacional de Cuyo. En prensa. ISSN (en línea) 1853-8665.
Original article
Growth
and Foliar Alkaloid Content of Clitoria ternatea under Different
Irrigation Frequencies
Desarrollo
y contenido foliar de alcaloides de la conchita azul (Clitoria ternatea)
con diferentes frecuencias de riego
Juan Pio
Salazar-Arias2,
Jorge Luis Ramirez
de la Rivera3
1Universidad Técnica de Cotopaxi. Facultad de Ciencias
Agropecuarias y Recursos Naturales. Calle los Almendros y Pujilí. La Maná.
Cotopaxi. C. P. 050202. Ecuador.
2Universidad Técnica de Cotopaxi. Facultad de Ciencias
Agropecuarias y Recursos Naturales. Vía Salache. km 7,53 sector
La Florida. Latacunga. C. P. 050107. Ecuador.
3Universidad de Granma. Facultad de Ciencias Agropecuarias.
Carretera a manzanillo Km 171/2. Bayamo. Granma. Código Postal: 85 100. Cuba.
*tatiana.gavilanez@utc.edu.ec
Abstract
Clitoria ternatea is widely used in
tropical livestock farming. This study evaluates the agronomic performance and
alkaloid content of C. ternatea under different irrigation frequencies
in protected cultivation. A randomized block design with controlled manual
irrigation was implemented with five treatments, four replicates, and five
experimental units per treatment. Treatments included daily irrigation (T0),
irrigation every two days (T1), every three days (T2), every four days (T3),
and every five days (T4). We evaluated plant height, leaf and internode number,
leaf area, green and dry matter yield, and alkaloid content. Irrigation every
three or four days generally resulted in the highest values for the measured
variables compared to the control. Reduced irrigation frequency affects the
agronomic performance of C. ternatea in protected cultivation. Agronomic
indicators were highest under irrigation every three or four days. Alkaloid
content did not vary with irrigation frequency.
Keywords: alkaloids, height,
leaf area, number of leaves, yield
Resumen
La Clitoria
ternatea constituye una de las especies de mayor uso en la ganadería del
trópico. El objetivo de este trabajo es evaluar el comportamiento agronómico y
los alcaloides de la C. ternatea al aplicar diferentes frecuencias de
riego en cultivo protegido. Se empleó un diseño en bloques al azar. Se
emplearon cinco tratamientos con cuatro repeticiones cada uno, con cinco
unidades experimentales cada una. Los tratamientos consistieron en diferentes
frecuencias de riego, sin riego (T0), cada dos días (T1), cada tres (T2), cuatro
(T3) y cada cinco (T4). Se evaluaron las variables altura de la planta, número
de hojas y entrenudos, área de la hoja, rendimiento de materia verde y seca de
la planta, así como los alcaloides. Las frecuencias de riego después de tres y
cuatro días mostraron los mayores valores para las diferentes variables
evaluadas, con respecto al control y resto de los tratamientos. La disminución
de la frecuencia de riego no afectó el comportamiento agronómico de la C.
ternatea en cultivo protegido. Los indicadores agronómicos fueron mayores
para las frecuencias de riego de tres y cuatro días. Por otra parte, el
contenido de alcaloides no varió en dependencia de la frecuencia de riego.
Palabras clave: alcaloides, altura,
área de la hoja, número de hojas, rendimiento
Originales: Recepción: 05/03/2025 - Aceptación: 25/09/2025
Introduction
In Latin America
and the Caribbean, legumes are grown with economically important crops to
promote sustainable agricultural and livestock production systems (14, 20). C. ternatea (Fabaceae) is widely
used in tropical livestock farming due to its adaptability, productivity, and
nutritional value (22). These benefits
improve animal diets and help meet feed demand during the dry season. They also
promote the use of sustainable animal production techniques compatible with
environmental protection.
However, in the
tropics, the agronomic performance, yield, and chemical composition of this
species are highly variable. This variability is influenced by plant age,
management practices (especially irrigation and fertilization), and
edaphoclimatic conditions (3, 23).
A previous study in
Ecuador showed that plant quality decreases with age (10). However, as this research was conducted
during the rainy season, it did not address irrigation effects. Furthermore,
scarce information describes how irrigation affects the development and foliar
alkaloid content of C. ternatea in Ecuador’s various climatic
conditions. Thus, this study aims to evaluate the agronomic performance and
alkaloid content of C. ternatea under different irrigation frequencies
in protected cultivation.
Materials
and methods
Site
and Experiment
The study was
conducted at the Technical University of Cotopaxi’s experimental area, El
Triunfo sector, La Maná Cantón. This site has a maximum temperature of 30°C, a
minimum of 17°C, relative humidity of 86.83%, average annual rainfall of
3029.30 mm, 735.70 light hours per year, and sandy loam soil (7). The experiment lasted 70 days, and irrigation
was applied manually.
Seed
Collection and Storage
Seeds were
collected at the Sacha Wiwa Experimental Center, treated, and stored in the
Germplasm Bank of the Technical University of Cotopaxi Extension, La Maná.
Seeds were stored at 6.50% humidity and 4°C until sowing.
Soil
Preparation, Sewing and Transplanting
Soil in the
research area contains 6.5% organic matter and has a pH of 6.5; therefore, no
amendments were necessary. Seeds were sown during the dry season (July) in
polystyrene trays with 200 cells filled with common substrate from the experimental
center. One seed was placed in each cell. Twenty days after sowing, seedlings
with two to three true leaves were transplanted into polyethylene bags, one
seedling per bag. These were immediately arranged according to the experimental
design with guides for their climbing nature. All this work was carried out at
the final study site. Weeds were controlled manually, and the area was
disinfected by solarization (19).
The crop was
protected with transparent plastic to prevent rain from interfering with treatments.
Sides were covered with dark saran. Treatments were arranged on tables, one
meter above the ground. Each table was divided into five sections, each
representing a replicate. Variables were measured 10 days after transplanting.
Experimental
Design and Treatments
The structured
design with controlled manual irrigation had five treatments, four replicates
and five experimental units, totalizing 100 plants. Treatments included daily
irrigation (T0), irrigation every two days (T1), every three days (T2), every
four days (T3), and every five days (T4).
Plant
Height, Number of Leaves, and Leaf Area
Plant height was
measured from the stem base to the tip of the terminal bud using a measuring
tape, expressed in cm. Leaf number was averaged from 10 plants randomly
selected per treatment. Leaf area was determined using the Smart Petiole
application, calculating leaf area from scanned images. These variables were
evaluated at 10, 20, 30, 40, and 50 days.
Node
Number, Green and Brown Matter, and Alkaloid Content
Internode number
was determined at 40 and 50 days after transplanting. Green and dry matter
yields were measured by weighing 10 randomly selected plants per treatment at
50 days after emergence. Fresh weights were measured using a semi-analytical
balance (precision: 0.01 mg). Dry matter was determined in an air-circulation
oven (1). Results were expressed in tons (2).
Alkaloid content
was determined using Dragendorff’s reagent by mixing (a) 0.8 g pentahydroxy
bismuth nitroxide in 40 ml distilled water and 10 ml acetic acid, and (b) 0.8 g
potassium iodide in distilled water. The bismuth nitrate solution was prepared
by dissolving 10 mg Bi(NO3)3·5H2O in 5 mL concentrated
nitric acid, then diluting to 100 mL with distilled water. Additionally, 3%
thiourea and 1% disodium sulfide were required.
Alkaloid solutions
were diluted in acidic, acid-alcoholic, and basic-alcoholic media (21). All solutions were centrifuged, and the
precipitate was washed with alcohol and re-centrifuged. The resulting
precipitate was mixed with 5 mL thiourea solution, and the compound formed was
measured spectrophotometrically. Results were expressed as moles of alkaloids
per gram (21).
Statistical
Analysis
Analysis of
variance (ANOVA) was performed, and means were compared using Tukey’s test (p ≤
0.05). Normality was assessed using the Kolmogorov-Smirnov test, and
homogeneity of variances was tested using Bartlett’s test.
Results
Plant
Height
At 10 days, treatment T4 showed the greatest plant height
(p<0.05), with significant differences compared to other treatments. T1, T2,
and T3 did not differ significantly, with values ranging from 10.36 to 10.60
cm. The control treatment had the lowest height, significantly different from
the others (table 1).
Table 1. Height
of C. ternatea (cm) according to evaluation days.
Tabla
1. Altura de la C. ternatea expresada
en cm según días de evaluación.
Different
letters in the same column indicate statistical differences for p<0.05.
T0=
daily irrigation, T1= irrigation every two days, T2= irrigation every three
days, T3= irrigation every four days, T4= irrigation every five days.
Letras
desiguales en una misma columna difieren significativamente para p<0,05.
T0=
irrigación diaria, T1= irrigación cada dos días, T2= irrigación cada tres días,
T3= irrigación cada cuatro días, T4= irrigación cada cinco días.
At 20 days, T2 and
T4 reflected the highest values (p<0.05). T1 and T3 showed no differences
between while T0 presented the lowest value (10.60 cm), significantly different
from the rest (table 1). At 30 days, T3 reached 21.54 cm
(highest, significant value). Treatments T1, T2 and T4, ranging between 17.0
and 17.89 cm, did not differ from each other. The lowest value was shown by T0
(table 1).
Treatment T1
yielded the highest value at 40 days (38.50 cm), significantly differing from
the others, while T3 and T4 did not differ from each other, and T2 showed the
lowest height (17.30 cm). However, at 50 days, T4 reached the highest value, T0
and T3 ranged between 40.25 and 40.40 without significant differences, and T2
was the shortest (table 1).
Number
of Leaves
At 10 days, treatment T3 had the greatest number of leaves
(p<0.05) with 11.60, not significantly different from T2. T1, T2, and T4 did
not differ significantly. The control (T0) had the lowest number of leaves,
although this was not significantly different from T1 (table 2). After 20 days, the highest value was reached by T3 with
significant differences compared to the rest. Treatments T0, T1, T2 and T4
showed no differences between themselves, with values between 11 and 12.40.
Table 2. Number
of leaves of C. ternatea according to evaluation days.
Tabla
2. Número de hojas de C. ternatea según
días de evaluación.
Different
letters in the same column indicate statistical differences for p<0.05
T0=
daily irrigation, T1= irrigation every two days, T2= irrigation every three
days, T3= irrigation every four days, T4= irrigation every five days.
Letras
desiguales en una misma columna difieren significativamente para p<0,05.
T0=
irrigación diaria, T1= irrigación cada dos días, T2= irrigación cada tres días,
T3= irrigación cada cuatro días, T4= irrigación cada cinco días.
Treatment T3
reached the highest value (p<0.05) at 30 days, with significant differences
in contrast to the rest. However, T2 and T4 did not show differences between
themselves, the control reflected the lowest height and was differentiated from
the rest. At 40 days, T1 and T4 reached the highest numbers of leaves with
significant differences compared to the rest. Consequently, T2 and T3 were the
lowest, without significant differences between them. Thus, at 50 days, T3
reached the highest value (48), with significant
differences from the rest. However, T0, T1 and T4 showed no differences, while
T2 showed the lowest number of leaves (table 2).
Leaf
Area
At 10 days, T2 and
T3 showed the greatest leaf area (p<0.05), although T2 did not differ from
T4. The lowest leaf area values were observed in T0 and T1, ranging between
3.28 and 3.64 cm2. At 20 days, T2 and T3 did not show differences
showed the highest values, while T4 did not present significant differences
compared to T3 (table 3). The lowest value was from T0 with
3.35 cm2.
Treatment T4
reflected the highest value leaf area at 30 days (8.13 cm2)
with significant differences with respect to T0, T1 and T3. T2 did not show
differences with respect to T3, showing values between 6.99 and 7.79 cm2. The smallest area
was reflected by T0. At 40 days, T2 and T3 did not show significant
differences, ranging between 14.90 cm2 and 16.02.
At 50 days, control and T4 reflected the lowest values, 8.90 and
9.74 cm2, respectively, while T3
showed the significantly highest leaf area (table 3).
Table 3. Leaf area according to evaluation days.
Tabla
3. Área de las hojas según días de
evaluación.
Different
letters in the same column indicate statistical differences for p<0.05.
T0=
daily irrigation, T1= irrigation every two days, T2= irrigation every three
days, T3= irrigation every four days, T4= irrigation every five days.
Letras
desiguales en una misma columna difieren significativamente para p<0,05.
T0=
irrigación diaria, T1= irrigación cada dos días, T2= irrigación cada tres días,
T3= irrigación cada cuatro días, T4= irrigación cada cinco días.
Internode
Number
At 40 days, T4 had
the maximum internodes (6.70, p<0.05), significantly different from the
other treatments, which ranged from 4.70 to 5.26. At 50 days, internode number
was not significantly different among treatments, ranging from 5.58 to 6.68 (figure 1). At 50 days, the number of knots did not reflect
differences between treatments, with values between 5.58 and 6.68 (figure
1).
Different
letters in bars differ significantly for p<0.05.
Letras
distintas en columnas difieren significativamente para p<0,05.
Figure
1. Internode number of C. ternatea at 40 and 50
Days according to Experimental Treatments.
Figura
1. Número de internudos de C.
ternatea a los 40 y 50 días según los tratamientos experimentales.
Yield
and Alkaloid Content
At 50 days, treatment T2 achieved the highest green matter
yield, although not significantly different from T0 and T4. For dry matter,
treatments T0, T1, and T2 did not differ significantly. The lowest yields were
observed in T3 and T4 (figure 2).
Different
letters in bars differ significantly for p<0.05.
Letras
distintasen columnas difieren significativamenbte poara p<0,05.
Figure
2. Fresh (FM) and Dry Matter (DM) of C. ternatea after
50 days of evaluation
Figura
2. Rendimiento de materia fresa y seca
de C. ternatea después de 50 días de evaluación.
Alkaloids reached their highest value for T2 (watering every
three days) with 14.2 g/kg-1 DM. Treatments T0 and T4
did not show differences between them. The lowest value was reached by T1 with
2 g/kg-1 DM (figure
3).
Different
letters differ significantly for p<0.05.
Letras
distintas en las barras difieren significativamente para p<0,05.
Figure
3. Alkaloid content of C. ternatea after 50 days
of evaluation.
Figura
3. Alcaloides de C. ternatea después
de 50 días de evaluación.
Discussion
Maximum plant
height was achieved when Clitoria ternatea received irrigation every
four or five days, with T4 reaching 48.50 cm at 50 days. These results differ
from those reported by other authors (17),
who observed plant heights ranging from 10.37 to 18.67 cm at 30 days and from
18.86 to 31.8 cm at 45 days under tropical conditions. Such discrepancies
underscore the critical role of both irrigation management and environmental
variables in modulating vegetative growth. Optimal water supply and climate
conditions directly affect physiological processes, promoting stem elongation
and overall development in C. ternatea (4,
13).
Leaf number
increased as plants matured, with the highest value recorded in T3 (48 leaves
at 50 days). According to previous studies (11),
before 75 days, C. ternatea predominantly allocates photoassimilates to
roots and young leaves, supporting the observed increase in leaf production
and, consequently, biomass accumulation. This pattern also aligns with the
trends noted for leaf area and internode development in this study.
Leaf area reached
its maximum when irrigation occurred every three or four days and plant age
exceeded 40 days, as similarly documented in previous studies (10). However, leaf area expansion and biomass
production in C. ternatea depend on age, environmental factors, and
agronomic practices. Irrigation management is a decisive factor in cell
expansion, while excessive moisture induces root anoxia (17). Our findings highlight the necessity of
balancing irrigation to optimize physiological performance and yield of C.
ternatea.
Distinct values for
leaf area in C. ternatea have been documented (18). An expanded leaf area enhances light
interception and carbon assimilation (24, 25).
Nevertheless, several agronomic and environmental factors-including irrigation
regime, precipitation patterns, and organic fertilization-modulate the
agronomic performance of C. ternatea.
In our
investigation, optimal vegetative vigor and yield were measured when irrigation
intervals ranged from three to four days. Environmental conditions and crop
management practices, particularly irrigation frequency, affect organ
differentiation and elongation (8, 24).
Yields exceeded
3.26 t DM ha-¹ at 50 days,
demonstrating the productive potential of C. ternatea under optimized
irrigation. Literature indicates that, under adequate irrigation, this species
may achieve up to 30 t ha-1 year-1;
however, reduced rainfall and limited soil moisture substantially decrease
productivity (10). The highest yields in
this study resulted from irrigation every three days, underscoring the
importance of precise water management. Productivity in C. ternatea varies
with seasonality, genotype, agronomic practices, and environmental variables
like humidity and soil properties (9).
These findings highlight the necessity of conducting field trials under diverse
irrigation regimes to validate and potentially improve productivity.
Yield results
obtained in this study exceed those documented by various authors for C.
ternatea under irrigated conditions. Previous reports indicate yields
ranging from 1.6 to 2.5 t ha-1,
as well as 1.3 t ha-1 at 60 days, and 0.86 t ha-1
of dry matter in grass-legume associations involving three rows
of signal grass and two rows of C. ternatea (12,
25, 26). Several factors may explain these discrepancies. Low
planting densities and full sunlight exposure enhance photosynthesis. This,
along with an expanded root system, promotes higher growth rates and biomass
production under adequate soil moisture (8, 15).
Alkaloid
concentrations measured align with those reported for other legume species and
tree crops. Literature cites values between 0.28 and 11 g kg-1,
noting that secondary metabolite content depends on plant age, defense
mechanisms, nutrient status, and edaphoclimatic factors (27). In this study, alkaloid levels ranged from
12 to 14 g kg-1,
with the highest value recorded in plants irrigated every three days. These
findings suggest that, under optimal agronomic management, C. ternatea can
accumulate substantial quantities of alkaloids, enhancing its functional and
nutritional value.
Environmental
contexts strongly affect the synthesis of secondary metabolites. Climate and
soil properties modulate the presence and concentration of bioactive compounds.
Stressful deficiencies or excesses in edaphoclimatic factors often upregulate
the biosynthesis of compounds like alkaloids. For instance, increased wind
speed accelerates evaporation of essential oils, while higher transpiration
rates facilitate the upward movement of tropane alkaloids from roots to
canopies (5, 6). This research
corroborates these observations and emphasizes the importance of integrated
environmental and agronomic management for optimizing yield and quality in C.
ternatea (16).
Conclusions
Reducing irrigation
frequency affects the agronomic performance of Clitoria ternatea under
protected conditions. In fact, most agronomic variables-including plant height,
leaf number, leaf area, and biomass yield-reached their highest values when
irrigation occurred every three or four days. These findings indicate that
watering within this interval optimizes vegetative growth and productivity by
avoiding drought and excessive soil moisture.
Given that daily
irrigation didn’t enhance growth parameters compared to less frequent regimes
under protected conditions, future research should assess whether such
intensive watering is justified under field conditions. Alkaloid stability
across all irrigation treatments further suggests that secondary metabolite
accumulation in C. ternatea remains largely unaffected by variations in
watering frequency within the tested range.
The evidence presented here supports the strategic management of
irrigation schedules according to crop developmental stage and physiological
indicators. This approach enables producers to optimize yield and resource use
efficiency without compromising quality.
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