The influence of watering intervals and types of biochar on the agrophysiological of Kejibeling (Strobilanthes crispus (L.) Blume)

A influência dos intervalos de irrigação e tipos de biochar na agrofisiologia da Kejibeling (Strobilanthes crispus (L.) Blume)

Abstract

Kejibeling (Strobilanthes crispus (L.) Blume) is a medicinal plant with various health benefits, including antioxidant, antidiabetic, anti-inflammatory, and anticancer properties, but its cultivation in Indonesia remains suboptimal. Proper watering intervals and growing medium composition, particularly biochar application, may enhance plant growth and physiological performance. This study employed an experimental quantitative approach using a factorial randomized completely block design. The objective of this study was to assess the impacts of watering intervals and biochar types on the growth, physiological aspects, and yield of S. crispus. The experiment was conducted in a screenhouse at the Faculty of Agriculture, Universitas Sebelas Maret, from April-October 2025, using a randomized completely block design (RCBD) with two factors: watering intervals (2, 4, and 6 days) and biochar types (without biochar, water hyacinth, coconut shell, and rice husk), resulting in 12 treatment combinations with three replications. The growth parameters evaluated included plant height, leaf number, leaf area, root volume, fresh and dry mass of roots, stems, and leaves, while the physiological parameters assessed included chlorophyll a, chlorophyll b, total chlorophyll, leaf greenness (SPAD), stomatal density, stomatal aperture width, stomatal conductance, transpiration rate, photosynthetic rate, and water use efficiency. The results showed significant interaction effects between watering interval and biochar type on chlorophyll b content and leaf fresh mass. The 4-day interval with water hyacinth biochar produced the highest leaf fresh mass (57.20 g), while the 6-day interval with coconut shell biochar produced the highest chlorophyll b content (1.83 mg g^-1). A 4-day watering interval resulted in the highest leaf number (145.96), stem fresh and dry mass (26.69 and 8.50 g), plant fresh and dry mass (125.36 and 30.23 g), and leaf dry mass (12.67 g). Water hyacinth biochar produced the highest leaf area (14.22 cm²), stem fresh mass (28.05 g), plant fresh mass (128.84 g), and leaf dry mass (12.41 g). The 4‑day interval optimizes moisture and nutrients, water hyacinth biochar improves leaf biomass, and coconut shell biochar enhances chlorophyll b via magnesium and oxidative stress mitigation. In conclusion, both watering interval and biochar type significantly influence the growth and physiological aspects of S. crispus. A 4-day watering interval combined with either water hyacinth or coconut shell biochar is recommended for optimal cultivation.

Keywords:
chlorophyll; growth; medicinal plant; organic matter; soil amendment

Resumo

A kejibeling (Strobilanthes crispus (L.) Blume) é uma planta medicinal com diversos benefícios para a saúde, incluindo propriedades antioxidantes, antidiabéticas, anti-inflamatórias e anticancerígenas, mas seu cultivo na Indonésia ainda é limitado. Intervalos de irrigação adequados e a composição do meio de cultivo, particularmente a aplicação de biochar, podem melhorar o crescimento e o desempenho fisiológico das plantas. Este estudo empregou uma abordagem quantitativa experimental, utilizando um delineamento fatorial em blocos casualizados. O objetivo foi avaliar os impactos dos intervalos de irrigação e tipos de biochar no crescimento, nos aspectos fisiológicos e na produção de S. crispus. O experimento foi conduzido em casa de vegetação na Faculdade de Agricultura da Universidade Sebelas Maret, de abril a outubro de 2025, utilizando delineamento em blocos casualizados com dois fatores: intervalos de irrigação (2, 4 e 6 dias) e tipos de biochar (sem biochar, biochar de jacinto-d’'água, casca de coco e casca de arroz), resultando em 12 combinações de tratamentos com três repetições. Os parâmetros de crescimento avaliados incluíram altura da planta, número de folhas, área foliar, volume radicular, massas fresca e seca de raízes, caules e folhas, enquanto os parâmetros fisiológicos avaliados incluíram clorofila a, clorofila b, clorofila total, teor de clorofila (SPAD), densidade estomática, largura da abertura estomática, condutância estomática, taxa de transpiração, taxa fotossintética e eficiência do uso da água. Os resultados mostraram efeitos de interação significativos entre o intervalo de irrigação e o tipo de biochar no teor de clorofila b e na massa fresca das folhas. O intervalo de 4 dias com biochar de jacinto-d’água produziu a maior massa fresca das folhas (57,20 g), enquanto o intervalo de 6 dias com biochar de casca de coco produziu o maior teor de clorofila b (1,83 mg g^-1). O intervalo de irrigação de 4 dias resultou no maior número de folhas (145,96), massa fresca do caule (26,69 g e 8,50 g), massas fresca e seca da planta (125,36 g e 30,23 g) e massa seca das folhas (12,67 g). O biochar de jacinto-d’água produziu a maior área foliar (14,22 cm²), massa fresca do caule (28,05 g), massa fresca da planta (128,84 g) e massa seca das folhas (12,41 g). O intervalo de 4 dias otimiza a umidade e os nutrientes; o biochar de jacinto-d’água melhora a biomassa foliar; e o biochar de casca de coco aumenta a clorofila b por meio do magnésio e da mitigação do estresse oxidativo. Portanto, tanto o intervalo de irrigação quanto o tipo de biochar influenciam significativamente o crescimento e a atividade fisiológica deS. crispus. Recomenda-se o intervalo de irrigação de 4 dias combinado com biochar de jacinto-d’água ou de casca de coco para um cultivo otimizado.

Palavras-chave:
clorofila; crescimento; planta medicinal; matéria orgânica; condicionador de solo

1. Introduction

Compared to previous generations, society today is more concerned with adopting a healthy lifestyle, and public awareness of the adverse effects of chemical therapies has coincided with a sharp rise in the use of herbal medicine. Medicinal herbs are highly valued in traditional healthcare systems and international markets, where they are greatly in demand (Aftab and Hakeem, 2021; Singh et al., 2024). In addition to acting as supplements or alternatives to modern medical treatments, medicinal plants improve human health and are considered safer than pharmaceuticals because they are natural and free from harmful chemicals (Agidew, 2022). Kejibeling (Strobilanthes crispus), a horticultural plant in the Acanthaceae family, it ranks among the most widely used medicinal plants because of its various health benefits. Native to Madagascar, Indonesia, and Malaysia, this plant is also known as 'pecah beling' or yellow strobilanthus, and its primary medicinal component is its leaves (Tajudin et al., 2022). Traditionally, S. crispus leaves have been used to treat various ailments, including kidney stones, gallstones, hypercholesterolemia, diabetes, and cancer (Chua et al., 2019; Rubiyanti et al., 2026). Ng et al. (2021)demonstrated that S. crispus leaves contain active compounds such as flavonoids (3.98%), alkaloids (3.2%), tannins (1%), and catechins (1.18%). However, its cultivation in Indonesia remains suboptimal, which negatively impacts yields. This issue can be addressed by ensuring proper watering intervals and an appropriate growing medium composition, as these are important factors that can enhance plant growth.

The recent rise in global temperatures indirectly affects crop agriculture, as elevated temperatures increase plant water consumption. Plant growth and development are significantly influenced by watering practices, which should be adjusted based on soil moisture and ambient temperature. Water availability affects the success of plant production, both vegetative and generative, because water is a basic requirement for plants (Idaryani et al., 2021). While water is essential for irrigation, excessive consumption may result in water scarcity, as well as harm to crops and deterioration of soil quality (Gudisa et al., 2021). Regular watering creates conditions in which water remains available in the growing medium, allowing plants to absorb water more effectively (Alharbi et al., 2024). Previous research indicates that proper watering intervals play a key role in water use efficiency, thereby supporting optimal plant growth without causing water stress or wastage. Research by (Efretuei et al., 2022) showed that watering at a 7-day interval on Talinum fruticosum produced a plant fresh mass of 8.68 g plant^-1. Research by Islam (2023) showed that watering every 3 days on tomato plants significantly increased unit fruit mass, yield per plant, yield per plot, and gross yield. InMesembryanthemum crystallinumplants, watering every 4 days resulted in higher leaf number, plant fresh mass, plant dry mass, and chlorophyll content (Mndi et al., 2023).

Strategies for optimal water use are closely linked to the growing medium employed, given that the ideal watering interval depends heavily on how well the planting medium functions. According to Wasis and Fitriani (2022), good growing media are those that can supply plants with sufficient water and nutrients to support their growth. Consequently, adding biochar to the growth medium is one effective way to achieve such improvement. Biochar refers to a carbon compound created when biomass is subjected to high heat with minimal oxygen present, a process known as pyrolysis (Adhikari et al., 2022; Amalina et al., 2022; Rahayu et al., 2025). Biochar incorporation into the soil enhances its physical, chemical, and biological characteristics, along with enhancing the accessibility of cations, phosphorus, nitrogen, and cation exchange capacity, thereby improving soil fertility and increasing available water content by 45% in coarse-textured soil and 14% in fine-textured soil; moreover, due to the abundance of macropores within its structure, biochar causes both immediate and long-lasting changes in soil water retention capacity, retaining water and increasing soil nutrients to facilitate enhanced plant growth (Fischer et al., 2019; Meng et al., 2024; Rahayu et al., 2022). Biochar production can utilize various agricultural wastes, making it an environmentally friendly solution, such as water hyacinth, coconut shell, and rice husk. Research by Gezahegn et al. (2024) indicated that applying water hyacinth biochar at a rate of 20 t ha^-1 raised the dry biomass of maize shoots and roots from 47.7% to 17.6% and from 78.4% to 54.1%, in that order. Research by Vinolina et al. (2024) showed that the application of coconut shell biochar at 10 t ha^-1 on shallots significantly enhanced plant height, leaf count, wet bulb mass per sample, dry bulb mass per sample, and wet bulb mass per plot. Research by Walianggen (2022) showed that rice husk biochar at a dose of 7.5 t ha^-1 on long bean plants enhanced the height of plants, the fresh mass of long beans, and the fruit mass produced per plant.

To date, no study has investigated the interactive effects of watering intervals and biochar types on the growth, physiological aspects, and yield of S. crispus. The research questions addressed in this study were: (1) Is there an interactive effect between watering interval and biochar type on the growth and physiological aspects of S. crispus? (2) Do different watering intervals affect the vegetative growth and physiological aspects of S. crispus? (3) Do different biochar types affect the vegetative growth and physiological aspects of S. crispus? Therefore, the objective of this study was to assess the impacts of watering intervals and biochar types on the growth, physiological aspects, and yield of S. crispus.

2. Materials and Methods

2.1. Experimental site

The research was conducted over the period of April through October 2025 within the screenhouse belonging to the Faculty of Agriculture at Universitas Sebelas Maret, in Jumantono District, Karanganyar Regency, Central Java Province, Indonesia (07^o 37' 831" S, 110^o 56' 905" E; 148 m a.s.l.). The site is characterized by a tropical climate, with a mean air temperature of 32^oC., a mean relative humidity of 58.81%, and a mean light intensity of 76.69% during the experimental period. Daily temperature, humidity, and light intensity data are presented in Figure 1.

Figure 1
The weekly fluctuations in temperature, relative humidity, and light intensity at the research location over the course of the observation period.

2.2. Biochar material preparation

The biochar employed in this research originated from agricultural waste materials, namely water hyacinth (Pontederia crassipes Mart.), coconut shell (Cocos nucifera L.), and rice husk (Oryza sativa L.), obtained from the Surakarta region, Central Java, Indonesia. The biochar was produced using a simple pyrolysis method. The feedstocks were air-dried, chopped into small pieces (2-5 cm), and placed in a pyrolysis drum. The pyrolysis process was conducted under limited oxygen conditions for 2-3 hours. After cooling, the biochar was crushed, and homogenized prior to application. The biochar production process is illustrated in Figure 2. The physicochemical properties of the biochar are shown in Table 1.

Figure 2
Biochar production process from agricultural waste.

Table 1
Physicochemical properties of water hyacinth, coconut shell, and rice husk biochar.

Type of Biochar	pH	Organic C (%)	N (%)	P2O5 (%)	K2O (%)	Organic matter (%)	C/N
Water hyacinth biochar	9.92	22.50	0.15	0.79	3.29	38.79	150
Coconut shell biochar	9.36	24.15	0.30	0.13	1.36	41.64	80.50
Rice husk biochar	6.54	26.36	0.76	0.39	1.02	45.44	34.68

2.3. Experimental design

The experiment was arranged using a randomized completely block design (RCBD) with two factors. The first factor was watering interval, consisting of three levels: every 2 days, every 4 days, and every 6 days. The second factor was biochar type, applied at a rate of 15 t ha^-1 which was selected as the optimum biochar dosage as recommended by Sakya et al. (2023), consisting of four levels: without biochar (control), water hyacinth biochar, coconut shell biochar, and rice husk biochar. The experiment was arranged in a 3 × 4 factorial scheme, totaling 12 treatment combinations (Table 2), with three replications (blocks), each consisting of three polybags.

Table 2
Treatment combination of watering intervals and biochar types.

Treatment code	Watering interval (days)	Biochar type
W2B0	2	Without biochar (control)
W2B1	2	Water hyacinth biochar
W2B2	2	Coconut shell biochar
W2B3	2	Rice husk biochar
W4B0	4	Without biochar (control)
W4B1	4	Water hyacinth biochar
W4B2	4	Coconut shell biochar
W4B3	4	Rice husk biochar
W6B0	6	Without biochar (control)
W6B1	6	Water hyacinth biochar
W6B2	6	Coconut shell biochar
W6B3	6	Rice husk biochar

2.4. Plant material preparation and planting

S. crispusseedlings were propagated from stem cuttings. The stem cuttings, approximately 10 cm in length with 6-8 nodes, were obtained from healthy mother plants. The cuttings were planted in a nursery medium consisting of a mixture of soil and chicken manure (2:1, v/v) and maintained under screenhouse conditions for four weeks until they were ready for transplanting. During this period, the cuttings were watered daily. After six weeks, the cuttings had developed 6-8 fully expanded leaves and a well-established root system, indicating they were suitable for transplanting. Each polybag (20 cm diameter × 26 cm height) was filled with 6.4 kg of Latosol soil. Biochar was applied at a rate of 15 t ha^-1 (45.9 g polybag^-1) and incorporated into the growing medium during soil preparation. Planting holes were made, and one seedling was placed into each hole at a depth of 3-5 cm, leaving at least two nodes above the soil surface. Each polybag contained one plant. Planting was carried out in the late afternoon to minimize transplant shock caused by excessive evapotranspiration.

2.5. Plant maintenance

Maintenance consisted of watering, weeding, pest and disease control, fertilization, and harvesting. During the first two weeks after planting (WAP), all plants were watered daily with a volume of 500 ml polybag^-1 to support establishment and root development. Watering was applied at intervals of 2, 4, or 6 days according to treatment, starting at two WAP, with a volume of 500 ml polybag^-1, increased to 800 m polybag^-1 after 45 days after planting (DAP). This increase was due to the advancing plant age, which led to higher water requirements as the plants grew larger and developed more extensive canopy and root systems. Weeding was performed manually by uprooting any weeds growing in the polybags. Pest control was conducted weekly; pests were removed manually, and chemical pesticides were used only when infestations caused severe damage. The main pests observed during the experiment were aphids (Aphis sp.), which were manually removed when present. No severe disease symptoms were observed throughout the experiment. Fertilization was applied in two splits: basal fertilization at 2 t ha^-1 (6.1 g polybag^-1) one week before transplanting, and a second application at 4 t ha^-1 (12.2 g polybag^-1) at 45 DAP. The fertilizer used was chicken manure, applied by broadcasting evenly around the plant base and lightly incorporated into the soil surface. Harvesting was carried out at 12 WAP, when plants attained a minimum height of 50 cm and at least eight leaves per plant, by uprooting the entire plant including the roots.

2.6. Agronomic variables

The agronomic characteristics assessed at 12 WAP comprised plant height, leaf number, leaf area, root volume, and both fresh and dry mass of the roots, stems, and leaves. Plant height was recorded by measuring from the stem's base to the highest point of growth with a measuring tape (Liew et al., 2021). Leaf number was determined by manually counting fully expanded leaves per plant. Leaf area was calculated using the formula length × width × correction factor (0.654) for all leaves. Root volume was assessed by cutting the root system at the stem base, cleaning the roots of adhering soil, and submerging them in a graduated cylinder filled with 500 mL of water; root volume was calculated as the water volume displacement (De Miguel-Díez et al., 2022). Fresh mass was obtained by separating roots, stems, and leaves and weighing them on a digital balance. Dry mass was measured after oven, drying each plant part (roots, stems, and leaves) at 60°C until constant mass (3 × 24 hours) using a digital balance.

2.7. Physiological variables

Physiological parameters, including leaf greenness, chlorophyll a, b, and total chlorophyll content, stomatal density, stomatal aperture width, stomatal conductance, transpiration rate, photosynthetic rate, and water use efficiency, were measured at 9 WAP. A SPAD meter was used to assess leaf greenness, with measurements performed on three leaves per sample: upper, middle, and lower canopy leaves. Chlorophyll content was determined following the method of Arnon (Asgari-Ravandi et al., 2025). Fresh leaves (1 g) were weighed using an analytical balance and ground into a fine powder using a mortar. Acetone (20 mL) was added using a pipette, and the mixture was stirred until homogeneous. The solution was filtered through filter paper and a funnel into a small bottle. The supernatant was transferred into a cuvette up to the marked line, while another cuvette was filled with acetone as a blank. Absorbance readings were taken with a spectrophotometer set to wavelengths of 654 nm and 663 nm. Chlorophyll concentrations were calculated using the following equations (Equations 1, 2 and 3):

$$\begin{array}{l}\text{Chlorophyll a}\left(mg{\text{g}}^{-1}\right)=\\ \left(12.7\times {\text{A}}_{663}-2.69\times {\text{A}}_{645}\right)\times (20mL)/(1000\times 1\text{g})\end{array}$$ (1)

$$\begin{array}{l}\text{Chlorophyll b}\left(mg{\text{g}}^{-1}\right)=\\ \left(22.9\times {\text{A}}_{645}-4.68\times {\text{A}}_{663}\right)\times (20mL)/(1000\times 1\text{g})\end{array}$$ (2)

$$\begin{array}{l}\text{Total chlorophyll}\left(mg{\text{g}}^{-1}\right)=\\ \left(20.2\times {\text{A}}_{645}+8.02\times {\text{A}}_{663}\right)\times (20mL)/(1000\times 1\text{g})\end{array}$$ (3)

Stomatal density and aperture width were assessed using the imprint method. Stomatal imprints were collected in the morning (09:00-12:00) (Anggraini et al., 2025). Leaves that were healthy and fully developed, taken from the middle section of the plant shoot, were chosen. A small amount of transparent nail polish was spread onto the underside (abaxial side) of each leaf. Once the nail polish had hardened, it was peeled off with adhesive tape. The tape with the stomatal imprint was then attached to a plastic sheet and transported to the laboratory. Stomatal density was observed under a light microscope at 10× magnification, and the number of stomata was counted using Image Raster software. Stomatal aperture width was measured from the same imprints at 40× magnification. Stomatal conductance, transpiration rate, and photosynthetic rate were measured using a portable photosynthesis analyser between 09:00-12.00 (Hu et al., 2023). Measurements were performed on fully expanded leaves from the middle canopy of each plant. The instrument was set to maintain a leaf temperature of 25 °C, and a relative humidity of 60-70% inside the chamber. For each experimental unit (polybag), five consecutive readings were recorded from the same leaf, and the average value was used for statistical analysis. Thus, a total of five readings were obtained per replication. Water use efficiency (WUE) was calculated using Equation 4:

$\text{WUE}=\frac{\text{A}}{\text{E}}$ (4)

Notes:

WUE = water use efficiency

$\left(\mu molC{O}_2{\left(mmol{\text{H}}_2\text{O}\right)}^{-1}\right)$

A = photosynthetic rate

$\left(\mu molC{O}_2{\text{m}}^{-2}{\text{s}}^{-1}\right)$

E = transpiration rate

$\left(mmol{\text{H}}_2{\text{Om}}^{-2}{\text{s}}^{-1}\right)$

(Jin et al., 2024).

2.8. Statistical analysis

The experiment was arranged using a randomized completely block design (RCBD) with a two-factor factorial arrangement. This design was chosen because it effectively controls variability caused by environmental heterogeneity in light intensity, by grouping experimental units into homogeneous blocks. Each block represented one replication, and treatments were randomly assigned within each block to ensure unbiased estimation of treatment effects.The entire dataset was analyzed using analysis of variance (ANOVA) with a two-factor factorial arrangement (3 watering intervals × 4 biochar types), testing the main effects of each factor as well as their interaction. The two-way ANOVA was employed to determine whether each factor independently influenced the measured variables and whether the combination of both factors produced synergistic or antagonistic effects through significant interactions. When significant differences were found, Duncan's multiple range test (DMRT) was applied to compare means, with a significance threshold of α = 0.05. The Duncan test was used because it allows for the identification of homogeneous subsets among treatment means, providing a clear distinction between treatments that are significantly different from one another while grouping those that are statistically similar. All statistical analyses were performed using SPSS software (version 26.0, IBM Corp., Armonk, NY, USA). Additionally, correlation analyses (Pearson's correlation coefficient) were conducted to examine the associations between growth variables and physiological variables. This analysis was conducted to identify which physiological traits are most strongly linked to growth performance, thereby providing insights into the underlying mechanisms driving treatment responses.

3. Result

3.1. Growth parameter

3.1.1. Plant height

The effects of watering intervals and biochar types on S. crispus plant height are presented in Figure 3. At 12 WAP, no significant interaction was detected between watering interval and biochar type (Figure 3a and 3b). The main effects of watering interval and biochar type were also not significant. The mean plant height for watering intervals of 2, 4, and 6 days ranged from 39.92 to 40.79 cm, with no significant differences among treatments. Similarly, the mean plant height for biochar ranged from 39.61 to 41.72 cm, with no significant differences observed.

Figure 3
Effects of watering interval and biochar type on growth parameters ofS. crispus: (a) Effect of watering interval on plant height and leaf number; (b) Effect of biochar type on plant height and leaf number; (c) Effect of watering interval on leaf area and root volume; (d) Effect of biochar type on leaf area and root volume. Notes: Values followed by the same letter are not significantly different according to Duncan's Multiple Range Test (α = 0.05).

3.1.2. Number of leaves

The application of watering intervals and biochar types significantly affected the number of S. crispus leaves at 12 WAP, whereas no significant interaction was detected between the two factors (Figure 3a and 3b). The 4-day watering interval produced the highest number of leaves (145.96 leaves), which was significantly higher than the 2-day and 6-day intervals. Among biochar types, the control (without biochar) produced the highest number of leaves (143.94 leaves), followed by coconut shell biochar, water hyacinth biochar, and rice husk biochar. The control was significantly higher than rice husk biochar. Coconut shell biochar also significantly increased leaf number compared to rice husk biochar. However, no significant difference was observed between the control and coconut shell biochar, or between water hyacinth biochar and any other treatment

3.1.3. Leaf area

The effects of watering intervals and biochar types on S. crispus leaf area at 12 WAP are presented in Figure 3c and 3d. No significant interaction was detected between watering interval and biochar type. The main effect of watering interval was not significant, whereas the main effect of biochar type was significant. The mean leaf area for watering intervals of 2, 4, and 6 days ranged from 12.10 to 12.63 cm², with no significant differences among them. Among biochar types, water hyacinth biochar produced the highest leaf area, which was significantly higher than the control (without biochar), coconut shell biochar, and rice husk biochar. However, no significant difference in leaf area was observed among the control, coconut shell biochar, and rice husk biochar.

3.1.4. Root volume

The effects of watering intervals and biochar types on S. crispus root volume at 12 WAP are presented in Figure 3c and 3d. No significant interaction was detected between watering interval and biochar type. Furthermore, the main effects of watering interval and biochar type were also not significant. The mean root volume for watering intervals of 2, 4, and 6 days ranged from 43.75 to 47.21 mL, while the mean root volume for biochar types ranged from 43.33 to 47.39 mL, with no significant differences among treatments within each factor.

3.1.5. Fresh and dry leaves mass

The interaction between watering intervals and biochar types significantly affected the fresh leaf mass of S. crispus at 12 WAP (Figure 4a, 4b, and 4c). The combination of a 4-day watering interval with water hyacinth biochar produced the highest fresh leaf mass (57.20 g), which was significantly higher than most other treatment combinations. Compared to the control (4-day interval without biochar), this treatment increased fresh leaf mass by 12.1%. The same watering interval with coconut shell biochar (51.77 g) and the 6-day interval with water hyacinth biochar (54.33 g) showed no significant difference from the highest treatment. However, the 2-day and 6-day intervals with coconut shell biochar produced the lowest fresh leaf mass (34.03 g and 33.90 g, respectively).

Figure 4
Effects of watering interval and biochar type on growth parameters ofS. crispus: (a) Effect of watering interval on fresh and dry leaves mass; (b) Effect of biochar type on fresh and dry leaves mass; (c) Interaction between watering interval and type of biochar on fresh leaves mass; (d) Effect of watering interval on fresh and dry stem mass; (e) Effect of biochar type on fresh and dry stem mass. Notes: Any values marked with the same letter are considered not significantly different from one another, as determined by Duncan's Multiple Range Test (p ≤ 0.05).

The application of watering intervals and biochar types significantly affected the leaf dry mass of S. crispus at 12 WAP, whereas no significant interaction was detected between the two factors (Figure 4a and 4b). The 4-day watering interval produced the highest leaf dry mass (12.67 g), which was significantly higher than the 2-day (11.01 g) and 6-day (9.50 g) intervals. Among biochar types, water hyacinth biochar produced the highest leaf dry mass (12.41 g), representing a 13.3% increase compared to the control without biochar (10.95 g). However, no significant differences were observed between these treatments.

3.1.6. Fresh and dry stem mass

The application of watering intervals and biochar types significantly affected the fresh stem mass of S. crispus at 12 WAP, whereas no significant interaction was detected between the two factors (Figure 4d and 4e). The 4-day watering interval produced the highest fresh stem mass (26.69 g), which was significantly higher than the 2-day (20.75 g) and 6-day (22.21 g) intervals. Among biochar types, water hyacinth biochar produced the highest fresh stem mass (28.05 g), representing increases of 21.9%, 39.3%, and 29.4%, respectively, compared to without biochar (23.01 g), coconut shell biochar (20.13 g), and rice husk biochar (21.68 g). Coconut shell biochar produced the lowest fresh stem mass and was significantly lower than without biochar and water hyacinth biochar.

The application of watering intervals significantly affected the dry stem mass of S. crispus at 12 WAP, whereas biochar type had no significant effect, and no significant interaction was detected between the two factors (Figure 4d and 4e). The 4-day watering interval produced the highest dry stem mass (8.50 g), which was significantly higher than the 6-day interval (7.33 g) by 16.0%, but not significantly different from the 2-day interval (7.45 g). No significant differences were observed among biochar types (Figure 4e).

3.1.7. Fresh and dry root mass

The application of watering intervals and biochar types did not significantly affect the fresh root mass of S. crispus at 12 WAP, and no significant interaction was detected between the two factors (Figure 5a and 5b). The mean fresh root mass for watering intervals of 2, 4, and 6 days ranged from 43.67 to 47.63 g, while the mean fresh root mass for biochar types ranged from 42.28 to 47.44 g, with no significant differences among treatments within each factor.

Figure 5
Effects of watering interval and biochar type on growth parameters ofS. crispus: (a) Effect of watering interval on fresh and dry root mass; (b) Effect of biochar type on fresh and dry root mass; (c) Effect of watering interval and type of biochar on fresh plant mass; (d) Effect of watering interval and type of biochar on dry plant mass. Notes: Any values marked with the same letter are considered not significantly different from one another, as determined by Duncan's Multiple Range Test (p ≤ 0.05).

The application of watering intervals and biochar types did not significantly affect the dry root mass of S. crispus at 12 WAP, and no significant interaction was detected between the two factors (Figure 5a and 5b). The mean dry root mass for watering intervals of 2, 4, and 6 days ranged from 8.25 to 9.07 g, while the mean dry root mass for biochar types ranged from 8.40 to 8.83 g, with no significant differences among treatments within each factor.

3.1.8. Fresh and dry plant mass

The application of watering intervals and biochar types significantly affected the plant fresh mass of S. crispus at 12 WAP, whereas no significant interaction was detected between the two factors (Figure 5c and 5d). The 4-day watering interval produced the highest plant fresh mass (125.36 g), which was significantly higher than the 2-day (105.48 g) and 6-day (108.30 g) intervals. Among biochar types, water hyacinth biochar produced the highest plant fresh mass (128.84 g), representing increases of 14.1%, 22.8%, and 22.2%, respectively, compared to without biochar (112.96 g), coconut shell biochar (104.92 g), and rice husk biochar (105.47 g). No significant differences were observed among without biochar, coconut shell biochar, and rice husk biochar.

The application of watering intervals significantly affected the plant dry mass of S. crispus at 12 WAP, whereas biochar type had no significant effect, and no significant interaction was detected between the two factors (Figure 5c and 5d). The 4-day watering interval produced the highest plant dry mass (30.23 g), which was significantly higher than the 2-day (26.58 g) and 6-day (26.90 g) intervals. No significant differences were observed among biochar types, with plant dry mass ranging from 26.62 g (coconut shell biochar) to 29.71 g (water hyacinth biochar).

3.2. Physiological parameter

3.2.1. Chlorophyll content

The effects of watering intervals and biochar types on the chlorophyll content of S. crispus at 9 WAP are presented in Figure 6a, 6b, and 6c. A significant interaction was detected between the two factors for chlorophyll b content, whereas no significant interaction or main effects were observed for chlorophyll a and total chlorophyll content (Figure 6a and 6b). The combination of a 6-day watering interval with coconut shell biochar produced the highest chlorophyll b content (1.83 mg g^-1), significantly increasing chlorophyll b content by 22.8%, 79.4%, and 103.3%, respectively, compared to the control (6-day interval without biochar; 1.49 mg g^-1), the 6-day interval with rice husk biochar (0.90 mg g^-1), and the 2-day interval with coconut shell biochar (1.02 mg g^-1) (Figure 6c). The lowest chlorophyll b content (0.90 mg g^-1) was recorded in the 6-day interval with rice husk biochar.

Figure 6
Effects of watering interval and biochar type on physiological aspects ofS. crispus: (a) Effect of watering interval on chlorophyll a, b, and total chlorophyll; (b) Effect of biochar type on chlorophyll a, b, and total chlorophyll; (c) Interaction between watering interval and type of biochar on chlorophyll b content; (d) Effect of watering interval on stomatal conductance, transpiration rate, and photosynthetic rate; (e) Effect of biochar type stomatal conductance, transpiration rate, and photosynthetic rate. Notes: Any values marked with the same letter are considered not significantly different from one another, as determined by Duncan's Multiple Range Test (p ≤ 0.05).

3.2.2. Stomatal conductance, transpiration rate, photosynthetic rate

The application of watering intervals and biochar types did not significantly affect the stomatal conductance, transpiration rate, and photosynthetic rate of S. crispus at 9 WAP, and no significant interaction was detected between the two factors (Figure 6d and 6e).

3.2.3. Water use efficiency, leaf greenness, stomatal density, and stomatal aperture width

The application of watering intervals and biochar types did not significantly affect the water use efficiency (μmol CO₂ (mmol H₂O)^-1), leaf greenness (SPAD value), stomatal density, and stomatal aperture width of S. crispus at 9 WAP, and no significant interaction was detected between the two factors (Table 3).

Table 3
Effect of watering interval and type of biochar on water use efficiency, leaf greenness, stomatal density, and stomatal opening width.

Treatment	Water use efficiency (μmol CO2 (mmol H2O)-1)	Leaf greennes (SPAD unit)	Stomatal density (unit mm-2)	Stomatal opening width (μm)
Watering interval
2 Days	55.88±48.70a	43.87±5.66a	146.00±18.98a	2.55±0.60a
4 Days	59.32±101.43a	46.67±3.32a	148.33±21.17a	2.55±0.46a
6 Days	31.01±42.27a	46.16±4.31a	171.33±46.36a	2.55±0.57a
Type of biochar
Without biochar	34.50±44.86	44.11±5.36a	164.66±33.08a	2.60±0.65a
Water hyacinth biochar	34.02±48.12	46.33±5.36a	147.56±28.00a	2.42±0.51a
Coconut shell biochar	67.54±103.33	46.94±4.17a	152.22±23.90a	2.35±0.54a
Rice husk biochar	58.88±67.74	44.87±3.31a	157.44±44.89a	2.43±0.55a
Interaction	-	-	-	-
p-value of watering interval	0.152 ns	0.273ns	0.161ns	0.300ns
p-value of types of biochar	0.889 ns	0.514ns	0.735ns	0.764ns
p-value of interaction	0.277 ns	0.270ns	0.673ns	0.921ns
CV (%)	51.48	9.67	22.18	21.57

• Note: Any values marked with the same letter are considered not significantly different from one another, as determined by Duncan's Multiple Range Test (p ≤ 0.05).
• A plus symbol (+) denotes the presence of a positive interaction, whereas a minus symbol (-) signifies the absence of any interaction.
• ^ns
  Non significance.
• CV: coefficient of variate.

3.2.4. Stomatal conductance, transpiration rate, photosynthetic rate

The application of watering intervals and biochar types did not significantly affect the stomatal conductance, transpiration rate, and photosynthetic rate of S. crispus at 9 WAP, and no significant interaction was detected between the two factors (Figure 6d and 6e).

3.3. Correlation analysis

Pearson's correlation revealed strong associations among the measured traits, underscoring the importance of above-ground biomass and root development in enhancing S. crispus growth under varying watering frequencies and biochar types (Figure 7). Fresh stem mass showed positive associations with most growth and biomass traits, particularly with fresh leaf mass (r = 0.972) and fresh plant mass (r = 0.926). A very strong correlation (0.972) between dry stem mass and dry plant mass also implied the significant role of stem biomass in determining overall plant productivity. Moreover, root development had a substantial impact on total plant biomass, showing strong correlations (>0.83) between dry root mass and dry plant mass (r = 0.837), as well as between fresh root mass and dry plant mass (r = 0.784).

Figure 7
Pearson correlation analysis between observation variables. Notes: PH: plant height, NL: number of leaves, LA: leaf area, RV: root volume, FLM: fresh leaves mass, DLM: dry leaves mass, FSM: fresh stem mass, DSM: dry stem mass, FRM: fresh root mass, DRM: dry root mass, FPM: fresh plant mass, DPM: dry plant mass, LG: leaves greenness, CA: chlorophyll a, CB: chlorophyll b, CT: chlorophyll total, SD: stomatal density, SOW: stomatal opening width, SC: stomatal conductance, TR: transpiration rate, PR: photosynthetic rate, WUE: water use efficiency. * The asterisk (*) indicates that the correlation is significant at the 0.05 level.

Fresh and dry root mass were nearly perfectly correlated with each other (r = 0.940), indicating consistent root tissue hydration across treatments. Leaf number exhibited strong positive correlations with dry leaf mass (r = 0.748) and dry plant mass (r = 0.721), suggesting that leaf count can serve as a reliable proxy for overall biomass accumulation. Plant height showed modest positive correlations with dry stem mass (r = 0.782) and dry plant mass (r = 0.720), but demonstrated a negative correlation with stomatal conductance (r = -0.431). Among photosynthetic pigments, chlorophyll b was very strongly correlated with total chlorophyll (r = 0.958), implying that variations in total chlorophyll content were primarily driven by chlorophyll b. Stomatal conductance showed an extremely strong positive correlation with transpiration rate (r = 0.974), confirming the physiological coupling between these two parameters. Fresh leaf mass was also strongly correlated with dry leaf mass (r = 0.814).

4. Discussion

4.1. Effect of watering interval and type of biochar on growth performance of S. crispus

The present study assessed the mutual influences of watering intervals and biochar types on the agronomic characteristics of S. crispus cultivated under screenhouse conditions. A significant interaction between watering interval and biochar type was detected for fresh leaf mass, suggesting a synergistic effect between optimal watering frequency and high-quality biochar in promoting leaf biomass production. The highest fresh leaf mass was obtained from the combination of a 4-day watering interval with water hyacinth biochar. The 4-day interval maintains soil moisture at an optimal level, thereby maximizing the water-holding capacity and nutrient release efficiency of water hyacinth biochar. Consequently, this optimal moisture condition enhances root activity and nutrient absorption without causing waterlogging or drought stress, as water hyacinth biochar also improves soil porosity and aeration (Hammam et al., 2022). The gradual and steady release of nutrients from the biochar under consistent soil moisture supports sustained leaf cell division and expansion, ultimately leading to higher fresh leaf biomass. This finding aligns with Sepehri et al. (2023) who reported that a 4-day irrigation interval improves water use efficiency, and with Ntoulas et al. (2025) who stated that the 4-day interval is an effective strategy for maintaining soil water status. Moreover,Yaebiyo et al. (2024) reported that the combined application of biochar and deficit irrigation significantly increased biomass production. The benefit of biochar is maximized under adequate water supply, as biochar application increases soil porosity and improves soil hydrology when soil moisture is well-maintained (Wu et al., 2022; Yang et al., 2022). In contrast, suboptimal watering intervals (2-day and 6-day) negated the potential benefits of biochar application, further emphasizing that the positive interaction between watering frequency and biochar type is critical for optimizing plant growth.

Watering intervals significantly affected the number of leaves, leaf dry mass, fresh and total plant mass, with the 4-day watering interval consistently producing the highest values. This finding indicates that a 4-day watering frequency provides optimal soil moisture conditions for vegetative growth of S. crispus. The intermediate watering interval (4 days) thus represents the most balanced regime for maintaining soil moisture without causing either waterlogging or drought stress. Watering every 2 days may lead to waterlogged conditions that reduce root zone aeration, inhibit nutrient uptake, and potentially induce hypoxia stress, which suppresses leaf initiation and biomass accumulation. This result aligns with Islam and Pervin (2023) who reported that a 4-day irrigation interval produced the highest leaf number in cauliflower, and withZhang et al. (2025) who stated that watering every 4 days enhances nitrogen uptake in tomatoes. Conversely, watering every 6 days likely induces mild to moderate water deficit that limits cell division, leaf expansion, and overall photosynthetic capacity, resulting in reduced dry matter production (Jarin et al., 2024).

Among biochar types, water hyacinth biochar consistently produced the highest leaf area, leaf dry mass, fresh stem mass, and fresh plant mass, demonstrating its superiority over coconut shell and rice husk biochar. This positive effect can be attributed to its high cation exchange capacity (CEC), nutrient content, and water-holding capacity, which collectively enhance soil fertility and plant nutrient availability (Canning, 2025; Gezahegn et al., 2024; Lewoyehu et al., 2024). Additionally, water hyacinth as a feedstock produces biochar with favorable physicochemical properties, including high surface area and porosity (Ullah and Rahman, 2024), which improve soil aeration, microbial activity, and nutrient retention. According to Lewoyehu et al. (2024), the porous structure and high carbon content of water hyacinth biochar can promote microbial activity by providing habitat and energy, thereby accelerating the decomposition of organic waste. Similarly, Rahayu et al. (2026) noted that biochar acts as a habitat for soil microorganisms and aids in the formation of stable soil aggregates, which enhances water retention. The high fresh stem mass observed with water hyacinth biochar is thought to be due to its high potassium (K) content, which aids in the translocation of photosynthetic products. This finding is consistent with Hossain (2020) who reported that water hyacinth biochar has the highest K content compared to other forms of biochar. In contrast, rice husk biochar often resulted in the lowest values across several parameters, possibly due to its high ash content (Hamidu et al., 2025), which can increase electrical conductivity and negatively affect plant development (De La Rosa et al., 2023). Coconut shell biochar showed intermediate effects, performing similarly to the control for most parameters.

The absence of significant effects on plant height, root volume, and fresh and dry root mass suggests that S. crispus exhibits a considerable degree of tolerance to variations in water availability and biochar application for these specific parameters. However, the correlation analysis reveals that plant height remains a meaningful indicator of overall biomass despite its lack of treatment response. The consistent relationship between height and dry matter accumulation across treatments suggests that taller plants inherently produce more biomass, even when height itself is not significantly altered by watering intervals or biochar types. Furthermore, the strong linkage between root biomass and total plant productivity underscores the functional importance of root system development. Even though root parameters did not respond significantly to treatments, the close association between root and shoot biomass implies that any improvement in root growth, whether through genetic or management practices, would directly translate into enhanced above ground productivity.

4.2. Effect of watering interval and type of biochar on physiological aspects of S. crispus

The present study revealed that watering intervals and biochar types had a significant interactive effect only on chlorophyll b content, while other physiological parameters remained largely unaffected. The highest chlorophyll b content was observed under the combination of a 6-day watering interval and coconut shell biochar. Chlorophyll b plays an essential role in light energy capture and photosynthetic adaptation to variable light conditions (Li et al., 2024; Simkin et al., 2022). The enhancement of chlorophyll b under mild water deficit combined with coconut shell biochar suggests an adaptive strategy to optimize light-harvesting efficiency when water availability is moderately restricted. Shahid et al. (2022) reported that increased chlorophyll b under water deficit in certain wheat genotypes serves as an adaptive strategy to optimize light harvesting and protect the photosynthetic apparatus from oxidative stress, thus sustaining photosynthesis and yield. In contrast, chlorophyll a content did not show significant treatment effects, which is consistent with findings by Pompelli et al. (2022) who reported that chlorophyll a is generally more stable than chlorophyll b under moderate stress conditions because it is centrally located in the reaction centers of photosystems and is structurally more protected. The positive effect of coconut shell biochar is likely attributable to its physicochemical properties. Coconut shell biochar increases total pore volume and water-holding capacity (Wang et al., 2023) and exhibits a high cation exchange capacity along with substantial nutrient content, particularly potassium, phosphorus, and magnesium (Lita et al., 2022; Manikandan and Nair 2025). Magnesium, in particular, regulates energy transfer between photosystem I and photosystem II and maintains chlorophyll structure integrity (Zhu et al., 2025). Thus, the combined supply of suitable nitrogen and magnesium from the biochar soil matrix may have facilitated more efficient chlorophyll b biosynthesis under the 6-day watering interval. The dominant role of chlorophyll b in determining total chlorophyll content is clearly reflected in the correlation structure. The very strong relationship between chlorophyll b and total chlorophyll indicates that variations in overall chlorophyll levels are almost entirely driven by changes in chlorophyll b rather than chlorophyll a.

No significant treatment effects were detected for leaf greenness (SPAD), stomatal density, stomatal aperture width, stomatal conductance, transpiration rate, photosynthetic rate, and water use efficiency. This overall stability suggests that S. crispus maintains robust physiological homeostasis across watering frequencies of 2 to 6 days and across different biochar types. A plausible explanation is that water availability remained sufficient even at the 6-day interval, meaning that the plants did not experience physiologically meaningful water stress. Aminah et al. (2020) reported that leaf greenness is influenced by accessible water; therefore, the lack of SPAD variation implies that water supply across treatments remained within the threshold required for normal chlorophyll biosynthesis. This interpretation is confirmed by the unchanged chlorophyll a and total chlorophyll levels, confirming that photosynthetic pigmentation was not compromised.

Stomatal density is known to increase under high relative humidity (RH) environments (Driesen et al., 2020). The comparable stomatal density observed across treatments therefore suggests that the microclimate around the leaf canopy was relatively uniform, with no treatment induced differences in RH. Furthermore, water scarcity typically reduces stomatal aperture (Driesen et al., 2023); however, the absence of aperture variation in this study indicates that even a 6-day watering interval did not impose substantial water deficit. From a biochemical perspective, stomatal density is determined by epidermal cell division and differentiation during leaf development, which are regulated by systemic signals such as plant hormones and nutrient availability (Anggraini et al., 2025). The stable stomatal density across treatments implies that nutrient absorption and hormonal signaling related to stomatal development remained unaffected. Pérez-Bueno et al. (2022) noted that stomatal width decreases with RH variation, which in turn affects stomatal conductance and transpiration. The stable aperture width across treatments implies that neither watering interval nor biochar type created differential humidity gradients at the leaf surface. From a biochemical perspective, stomatal opening is regulated by potassium ion (K⁺) influx and efflux in guard cells, which is influenced by plant water status and abscisic acid (ABA) signaling (Liu et al., 2022). The lack of significant change in stomatal aperture suggests that water availability across all treatments remained sufficient to maintain guard cell turgor and normal stomatal functioning. Furthermore, the stable K⁺ content in the growing medium, particularly from water hyacinth biochar which has high K₂O content (3.29%), may have contributed to maintaining normal stomatal regulation despite variations in watering frequency.

Stomatal conductance typically declines under increasing groundwater stress, whether from drought or flooding (Goto et al., 2021). The lack of significant change in stomatal conductance in the present study further supports the conclusion that the plants remained within a well-watered physiological state. The physiological coupling between stomatal conductance and transpiration rate is strongly evident in this study. The nearly inseparable relationship between these two parameters confirms that transpiration is primarily regulated by stomatal opening in S. crispus. Consequently, the absence of treatment effects on stomatal conductance directly explains the lack of effects on transpiration rate. Biochar application did not alter transpiration, photosynthesis rates, and water use efficiency. One possible reason is that biochar decomposes slowly and releases nutrients over an extended period (Rahayu et al., 2025); thus, within the relatively short experimental duration, the effects of biochar on plant water relations may not have fully manifested. These findings are consistent with (Purwanto et al., 2025) observed that 3- and 6-day watering intervals did not significantly influence transpiration rate. The absence of treatment effects on photosynthetic rate can be attributed to adequate soil water supply across all treatments, as photosynthetic aspects is impaired only when water scarcity co-occurs with other stressors (Xu et al., 2020). Moreover, the effect of biochar on photosynthetic rate depends on soil type and application dosage (Piao et al., 2023). From a biochemical standpoint, the photosynthetic rate is determined by both stomatal (CO₂ diffusion) and non-stomatal (biochemical) limitations (Bellasio, 2025). The stable photosynthetic rate across treatments indicates that neither stomatal conductance nor biochemical processes such as Rubisco activity, electron transport, and Calvin cycle enzymes were impaired under the tested conditions. Collectively, these results indicate that S. crispus exhibits considerable physiological tolerance to moderate variations in watering frequency, and that biochar addition offers selective benefits under the tested conditions.

5. Conclusion

The interaction between watering interval and biochar type significantly affected fresh leaf mass and chlorophyll b content, with the combination of a 4-day watering interval and water hyacinth biochar producing the highest leaf mass, and the 6-day interval with coconut shell biochar enhancing chlorophyll b. A 4-day watering interval consistently enhanced most agronomic traits, including leaf number, stem mass, and total plant mass, indicating that moderate watering frequency provides optimal soil moisture for S. crispus cultivation. Among biochar types, water hyacinth biochar produced the highest leaf mass compared to coconut shell and rice husk biochar. Physiological parameters remained largely stable across treatments, demonstrating that S. crispus possesses strong physiological tolerance to watering intervals of 2 to 6 days. For optimal cultivation ofS. crispus, a 4-day watering interval combined with either water hyacinth or coconut shell biochar is recommended, depending on whether the goal is to maximize leaf biomass or enhance specific physiological traits like chlorophyll b content.

Acknowledgements

This research was funded by the Annual Work Plan and Budget (RKAT) of Universitas Sebelas Maret for the 2025 Fiscal Year, under the Research Group Capacity Strengthening Research (PKGR-UNS) A scheme, with research assignment agreement number: 371/UN27.22/PT.01.03/2025. The authors would like to thank the Faculty of Agriculture, Universitas Sebelas Maret, for providing the screenhouse and research facilities. The authors also acknowledge the technical assistance provided during the experimental period. The authors declare that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

Data Availability Statement

The complete dataset generated in this study is included here. Detailed methodologies and unprocessed data can be obtained by contacting the corresponding author with a justifiable request.

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*

e-mail: [e-mail]

Editor:

Takako Matsumura Tundisi

Publication Dates

• Publication in this collection
  09 Oct 2026
• Date of issue
  2026

History

• Received
  17 June 2026
• Accepted
  24 Aug 2026