Optimal lemongrass (Cymbopogon citratus) supplementation improves growth, feed efficiency, and digestive development without altering hematological status in red tilapia (Oreochromis spp.)

Abstract

The use of phytogenic feed additives in aquaculture has gained attention as a sustainable strategy to enhance fish growth and health; however, optimal inclusion levels and mechanistic responses remain insufficiently defined. This study aimed to evaluate the effects of dietary aqueous Cymbopogon citratus (lemongrass) extract on growth performance, feed utilization, morphometric development, biometric indices, and hematological status of red tilapia (Oreochromis spp.). A 60-day feeding trial was conducted using 360 fingerlings randomly assigned to four dietary treatments (0, 10, 20, and 30 g kg^-¹), with three replicates per treatment. Growth metrics, feed efficiency, morphometric traits, organ indices, and hematological parameters were analyzed using parametric and non-parametric statistical approaches, complemented by dose–response modeling. Lemongrass supplementation significantly improved growth performance, with higher final weight, weight gain, and specific growth rate (p < 0.05), alongside reduced feed conversion ratio (lowest at ~2.4) and increased biomass. Morphometric traits, including total length and fin development, were significantly enhanced at 20 g kg^-¹, while hematological parameters remained stable, indicating physiological safety. Quadratic dose–response modeling identified an optimal inclusion level of ~18–20 g kg^-¹, linking enhanced growth performance to improved digestive organ development. Biometric responses further suggested selective increases in gut and visceral mass without adverse effects on liver indices. In conclusion, dietary lemongrass extract optimizes growth and feed efficiency through enhanced digestive capacity and allometric development, without compromising fish health. These findings highlight its potential as a functional phytogenic additive for sustainable and welfare-oriented tilapia aquaculture.

Introduction

Aquaculture remains one of the fastest-growing food-producing sectors globally and plays a central role in food security, livelihood generation, and economic development. Global fish production from aquaculture continues to increase in response to rising demand for affordable animal protein, with tilapia representing one of the most commercially important cultured fish groups worldwide (Food and Agriculture Organization of the United Nations (FAO), 2024). Among tilapia species, red tilapia (Oreochromis spp.) has gained substantial economic value because of its rapid growth, tolerance to diverse culture conditions, consumer acceptance, and high market demand in many Asian countries, including China, Thailand, and Malaysia (Bardhan et al., 2021; Munguti et al., 2014). Global tilapia production exceeded 6.7 million tonnes in 2023 and is projected to continue expanding, underscoring its importance in sustainable aquaculture development (Global Seafood Alliance, 2023; Food and Agriculture Organization, 2025). However, intensive tilapia production systems frequently expose fish to environmental stressors, including overcrowding, fluctuating water quality, and pathogen outbreaks, which negatively affect growth performance, physiological condition, and farm productivity (Frimpong et al., 2017).

To sustain productivity under intensive farming conditions, aquaculture has traditionally relied on antibiotics and synthetic growth promoters to improve fish health and production efficiency. Nevertheless, the indiscriminate use of these compounds has raised major concerns regarding antimicrobial resistance, environmental contamination, and antibiotic residues in aquatic food products intended for human consumption (Done et al., 2015; Kılıç & Gültekin, 2024). Consequently, there has been increasing interest in environmentally sustainable alternatives capable of enhancing fish performance without compromising animal welfare or ecosystem integrity. Functional feeds supplemented with phytogenic additives have emerged as promising alternatives because of their capacity to improve digestion, nutrient utilization, immune response, antioxidant activity, and disease resistance in cultured fish species (Gabriel, 2019). Recent studies have shown that medicinal plant supplementation can significantly enhance growth performance, hematological profiles, immune activity, and pathogen resistance in fish under culture conditions (Ujan et al., 2024). Similarly, functional phytogenic diets have been reported to improve feed efficiency, antioxidant balance, and physiological resilience in common carp and other freshwater species exposed to intensive production stressors (Habib et al., 2024).

Among medicinal plants evaluated in aquaculture nutrition, lemongrass (Cymbopogon citratus) has attracted growing attention because of its rich content of bioactive compounds, including citral, flavonoids, tannins, and terpenoids, which exhibit antimicrobial, antioxidant, digestive-stimulatory, and anti-inflammatory properties (Oladeji et al., 2019; Shah et al., 2011). In fish nutrition, phytogenic compounds derived from medicinal herbs have been associated with improved digestive enzyme activity, enhanced gut morphology, increased nutrient absorption, and strengthened immune responses (Reverter et al., 2014; Windisch et al., 2008). In Nile tilapia, dietary lemongrass essential oil improved growth performance and immune responses through enhanced physiological efficiency and health status (Al-Sagheer et al., 2018). Similarly, supplementation with medicinal plant extracts such as Moringa oleifera has been shown to improve growth and reproductive performance in Nile tilapia, although excessive inclusion levels may reduce overall biological performance, indicating the importance of determining optimal dietary concentrations (Naz et al., 2025). These findings collectively highlight the growing potential of phytogenic additives as sustainable nutritional strategies in aquaculture systems.

Despite these advances, important knowledge gaps remain regarding the application of aqueous C. citratus extract in red tilapia nutrition. Existing studies have largely focused on isolated performance indicators such as growth or immunity, while integrated assessments linking morphometric development, digestive organ responses, biometric indices, and hematological stability remain limited. In addition, variation in extraction procedures, dietary inclusion levels, and fish species has produced inconsistent findings across studies, making it difficult to establish standardized recommendations for practical aquaculture applications (Soh et al., 2022). Moreover, studies evaluating dose-dependent responses and mechanistic relationships between phytogenic supplementation and digestive development in red tilapia hybrids are still scarce.

Therefore, the present study evaluated the effects of dietary aqueous C. citratus extract on growth performance, feed utilization, morphometric traits, biometric indices, and hematological parameters of red tilapia. In addition, quadratic dose–response modeling was applied to determine the optimal dietary inclusion level associated with improved biological performance. By integrating growth, structural, physiological, and biometric responses, this study provides a more comprehensive understanding of the role of lemongrass as a functional phytogenic additive. It contributes to the development of sustainable, welfare-oriented, and antibiotic-free feeding strategies for tilapia aquaculture.

Materials and methods

Study area and experimental fish

The feeding trial was conducted at the Northeastern Mindanao State University–CFAS Aquaculture Complex from November 2025 to January 2026. Juvenile red tilapia obtained from the Technical Education and Skills Development Authority–Northern Mindanao School of Fisheries hatchery in Buenavista, Agusan del Norte, Philippines, were used in the study. The stock consisted of commercially cultured red tilapia hybrids, which generally originate from crosses involving Nile tilapia and Mozambique tilapia, although the precise genetic lineage was not genetically verified. A total of 360 healthy fingerlings (mean initial length ≈3 cm; mean initial weight ≈ 0.46 g) were transported in oxygenated containers.

Upon arrival, fish were acclimatized for 30 min in aerated tanks and conditioned for 1 week in a controlled culture system. During acclimation, fish were fed a commercial diet containing 35% crude protein twice daily to apparent satiation. Water quality parameters, including temperature, pH, dissolved oxygen (DO), and ammonia, were regularly monitored to maintain optimal conditions for tilapia culture. Partial water exchange (30%–50%) was conducted every other day following standard aquaculture management practices (El-Sayed & Mansour, 2021).

Ethics statement

All experimental procedures were conducted in accordance with institutional guidelines for the care and use of aquatic organisms, and ethical approval was obtained from the North Eastern Mindanao State University – Institutional Animal Care and Use Committee (IACUC).

Preparation of lemongrass aqueous extract

Fresh leaves of Lemongrass (Cymbopogon citratus) were collected locally from Buenavista, Agusan del Norte. Only the above-ground portions were used due to their higher concentration of essential oils and bioactive compounds, which are widely associated with antimicrobial and antioxidant properties (Negrelle and Gomes, 2007; Aćimović et al., 2020). Samples were washed, oven-dried at 38 °C for 48 h, and ground into fine powder.

Measured quantities of powdered lemongrass (10 g, 20 g, and 30 g) were soaked separately in 100 mL distilled water for 24 h at room temperature with intermittent agitation. The mixtures were filtered using sterile muslin cloth followed by Whatman No. 1 filter paper. Filtrates were stored at 5 °C and considered concentrated aqueous extracts before diet preparation (Kigigha and Kalunta, 2017).

Diet preparation

A commercial tilapia feed (TATEH Surfer Tilapia Pre-Starter) containing 36% crude protein, 7% crude fat, 5% crude fiber, 12% crude ash, and 12% moisture was used as the basal diet. Experimental diets were prepared using the coating method by spraying measured volumes of lemongrass aqueous extract onto pre-weighed pellets to obtain supplementation levels of 10, 20, and 30 g kg^-¹ diet. The coated feeds were thoroughly mixed, oven-dried at 37 °C for 1 h, cooled, and stored at 4 °C until use. Control diets received an equivalent volume of distilled water to maintain similar moisture content and palatability (Kro’l and Zakeś, 2016; Kumar et al., 2017).

Nutrient composition was based on manufacturer specifications, and no post-coating proximate analysis was conducted, reflecting practical farm-level application despite limited nutrient standardization. Pellet integrity remained stable throughout the trial. To minimize potential leaching of water-soluble bioactive compounds, coated feeds were dried immediately after spraying and administered promptly during feeding periods.

Experimental design and feeding management

The study followed a Completely Randomized Design with four dietary treatments and three replicates each. Twelve plastic tanks (50-L capacity) were filled with 40 L of dechlorinated freshwater, and thirty fish were randomly stocked per tank. This stocking density is consistent with tilapia nutrition trials evaluating phytogenic feed additives (El-Sayed & Mansour, 2021).

Fish were fed four times daily (06:00, 09:00, 12:00, and 16:00) at a feeding rate equivalent to 8% of body weight per day for 60 days. Feed rations were adjusted weekly following bulk weighing. Uneaten feed and metabolic wastes were siphoned daily prior to the first feeding to maintain water quality. Continuous aeration was provided through a centralized blower system.

Water quality monitoring during culture

Water temperature, pH, and dissolved oxygen were measured twice daily using portable meters, while ammonia concentrations were assessed twice weekly (Table 1). Partial water exchange (30–50%) was conducted every other day or as necessary. Water quality parameters were maintained within acceptable ranges recommended for tilapia culture (El-Sayed, 2006).

Growth monitoring

Fish were batch-weighed and measured at seven-day intervals to monitor growth trends. At the end of the 60-day feeding period, all surviving fish were counted and bulk-weighed per tank. Growth and feed utilization parameters were computed using standard fisheries equations (Hwihy et al., 2021):

• ◼ Weight Gain (WG) = Final Weight − Initial Weight

• ◼ Specific Growth Rate (SGR, % day^-¹) = [(ln FW − ln IW)/days] × 100

• ◼ Survival Rate (SR %) = (final fish ÷ initial fish) × 100

• ◼ Feed Conversion Ratio (FCR) = feed intake ÷ weight gain

• ◼ Protein Efficiency Ratio (PER) = weight gain ÷ protein intake

• ◼ Condition Factor (K) = 100 × body weight/length³

• ◼ Biomass gain per tank

Morphometric and biometric assessment

Twenty-four hours after the final feeding, nine (9) fish per treatment were randomly sampled. Fish were anesthetized using tricaine methanesulfonate (MS-222) prior to handling to minimize stress.

External morphometric measurements including body weight, total length, standard length, snout length, eye diameter, caudal peduncle depth, and fin lengths were measured using a digital caliper and analytical balance.

Fish were dissected through a ventral incision to obtain internal organ measurements. The digestive tract, liver, and visceral organs were carefully excised, rinsed, blotted dry, and weighed. Biometric indices were calculated as follows (Bagenal and Tesch, 1978; Hwihy et al., 2021):

• ◼ Relative Gut Length (RGL) = intestine length ÷ standard length

• ◼ Gut Somatic Index (GSI, %) = 100 × gut weight ÷ body weight

• ◼ Hepatosomatic Index (HIS, %) = 100 × liver weight ÷ body weight

• ◼ Viscerosomatic Index (VSI, %) = 100 × viscera weight ÷ body weight

• ◼ Condition Factor (K) = 100 × body weight ÷ standard length³

Hematological analysis

Blood sampling was conducted 24 h after the final feeding to avoid post-prandial metabolic fluctuations. Blood was drawn from the caudal vein using sterile syringes, and approximately 0.25 mL per fish was transferred into EDTA-coated tubes to prevent coagulation.

Hematological profiling was performed at the Butuan Veterinary Clinic using an automated hematology analyzer (Rayto RT-7600S). Parameters measured included red blood cell count, white blood cell count, hemoglobin concentration, hematocrit, and platelet count following standard clinical laboratory procedures (Fazio, 2019). Units were expressed as ×10⁹ L^-¹ and ×10¹² L^-¹ where appropriate.

Statistical analysis

All data were expressed as mean ± standard deviation (SD). Normality and homogeneity of variance were tested using the Shapiro–Wilk and Levene’s tests, respectively. Statistical significance was set at p < 0.05.

Growth performance, feed utilization, morphometric traits, biometric indices, and hematological parameters were analyzed using one-way ANOVA, followed by Tukey’s HSD test. When assumptions were not met, the Kruskal–Wallis test with Dunn–Bonferroni post hoc comparisons was applied.

Quadratic polynomial regression was used to evaluate dose–response relationships for selected biometric variables. The model was defined as:

where is the response variable and is the dietary inclusion level (g kg^-¹). The optimal inclusion level was estimated using:

Model fit was assessed using the coefficient of determination (R²), and results were visualized with 95% confidence intervals.

Pearson’s correlation analysis was used to examine relationships among morphometric, biometric, and physiological parameters, and results were presented as a heatmap. Statistical analyses were performed using JASP (v0.19.3), while regression modeling and visualization were conducted in R (v4.3.3).

Results

Growth performance

Final weight, total weight gain (TWG), average daily weight gain (ADWG), and weight-specific growth rate (WSGR) were significantly higher in all lemongrass-supplemented groups compared with the control (p < 0.05; Table 2). Similarly, length-based parameters (final length, TLG, ADLG, and LSGR) were significantly improved in supplemented treatments. The 20 g kg^-¹ group consistently exhibited the highest mean values; however, no significant differences were observed among supplemented groups (p > 0.05), indicating a plateau response beyond the lowest inclusion level.

Survival ranged from 76.67% to 85.56%, with numerically higher values in supplemented groups, although differences were not statistically significant (p > 0.05; Table 2).

Feed utilization

Feed conversion ratio (FCR) was significantly lower in all lemongrass-supplemented groups (2.47–2.64) than in the control group (3.04) (p < 0.05; Table 3). Final biomass was significantly higher in all supplemented treatments compared with the control, with no significant differences among supplementation levels (p > 0.05).

Condition factor (K) differed significantly among treatments, with the highest values observed at 10 and 20 g kg^-¹ (1.57 and 1.50, respectively), while the 30 g kg^-¹ group showed the lowest value (1.35) compared with the control (1.46) (p < 0.05).

Protein efficiency ratio (PER) also varied significantly among treatments (p < 0.05). The 30 g kg^-¹ group (1.16) exhibited the highest PER and differed significantly from the 20 g kg^-¹ group (0.62). Meanwhile, the control (0.95) and 10 g kg^-¹ (1.09) groups showed intermediate values and did not differ significantly from either treatment.

Hematological parameters

All measured hematological parameters, including leukocyte profiles, erythrocyte indices, hemoglobin, hematocrit, and platelet parameters, did not differ significantly among treatments (p > 0.05; Table 4), indicating that lemongrass supplementation did not affect physiological homeostasis.

Morphometric traits

Total length, standard length, head length, snout length, and fin dimensions (dorsal, anal, and pectoral) were significantly higher in supplemented groups compared with the control (p < 0.05; Table 5). The highest values were generally observed at 20 g kg^-¹.

In contrast, pelvic fin length, eye diameter, and caudal peduncle depth were not significantly affected (p > 0.05), indicating selective morphological enhancement rather than uniform structural growth.

Biometric indices and dose–response relationships

Visceral weight (VW) and gut weight (GW) were significantly higher in all supplemented groups compared with the control (p < 0.05; Table 6). The highest VW was observed at 20 g kg^-¹, while GW increased progressively across treatments, with no significant differences among supplemented levels, suggesting a plateau response.

Quadratic dose–response models (Figure 1) revealed strong non-linear relationships for VW (R² = 0.975) and GW (R² = 0.952). The estimated optimal inclusion level for VW was approximately 18.99 g kg^-¹, while GW showed a continuous increasing trend within the tested range. Intestinal length (IL) also followed a quadratic pattern (R² = 0.924), with an estimated optimum at ~16.02 g kg^-¹, consistent with the highest observed value at 20 g kg^-¹.

Figure 1

In contrast, liver weight (LW), relative gut length (RGL), gut somatic index (GuSI), hepatosomatic index (HSI), and viscerosomatic index (VSI) were not significantly affected (p > 0.05).

Relationships among morphometric and physiological traits

Correlation analysis (Figure 2) showed moderate negative associations between viscerosomatic index and body size parameters, indicating reduced relative visceral mass with increasing growth. Gut somatic index exhibited weak-to-moderate negative relationships with selected morphometric traits, while relative gut length showed positive correlations with intestinal length and other structural parameters. Hepatosomatic index was strongly associated with liver weight but weakly related to external morphometrics. Condition factor showed no significant correlations.

Figure 2

Discussion

Dietary supplementation with aqueous Cymbopogon citratus improved the growth performance of red tilapia (Oreochromis spp.), as indicated by higher final weight, weight gain, and specific growth rates compared with the control group. However, supplementation levels above 10 g kg^-¹ did not produce additional growth benefits, suggesting an optimal inclusion threshold. Similar growth-enhancing effects of phytogenic feed additives have been reported in tilapia and other cultured fish species (Abdel-Tawwab et al., 2018; El-Dakar et al., 2008; Acar et al., 2015). These responses are likely associated with bioactive compounds such as citral and phenolic constituents in lemongrass, which may improve appetite, digestive activity, and nutrient utilization efficiency (Hashemi and Davoodi, 2011; Al-Sagheer et al., 2018).

The present study did not quantify individual phytochemical constituents such as citral, flavonoids, or total phenolic compounds in the prepared aqueous extracts. Therefore, the precise concentration of active bioactive compounds delivered through the diets remains uncertain. Nevertheless, previous studies have consistently identified citral as the dominant bioactive constituent of C. citratus, together with several phenolic and flavonoid compounds associated with antioxidant and antimicrobial activity (Oladeji et al., 2019; Mishra, 2024). Future studies should include phytochemical profiling and extract standardization to improve reproducibility and dose precision.

Phytogenic compounds have also been associated with improved gut morphology and absorption efficiency in fish (Reverter et al., 2014; Windisch et al., 2008). Although water quality parameters remained within acceptable ranges for tilapia culture, the relatively high stocking density used in the present study may have contributed to physiological stress, reduced growth potential, and moderate survival rates across treatments. Therefore, the observed biological responses should be interpreted within the context of controlled high-density experimental conditions.

Feed utilization also improved in fish receiving lemongrass supplementation, as indicated by lower feed conversion ratios and higher biomass production. These results reflect improved nutrient assimilation and metabolic efficiency, consistent with previous findings on phytogenic additives enhancing digestive processes and intestinal health (Windisch et al., 2008; Hashemi and Davoodi, 2011). The moderate variation in condition factor suggests improved energy allocation without excessive lipid deposition.

Hematological parameters remained stable across treatments, indicating that lemongrass supplementation up to 30 g kg^-¹ did not compromise physiological or immune function. This stability supports the safety of phytogenic supplementation, as similar studies have reported enhanced growth performance without adverse hematological alterations (Fazio et al., 2013). The relatively high variability observed in some hematological parameters, particularly WBC counts, may have reduced statistical sensitivity for detecting treatment-related differences. Such variability may reflect normal inter-individual physiological variation under experimental culture conditions, combined with relatively limited sample size.

Morphometric analysis revealed enhanced axial growth, particularly at 20 g kg^-¹, indicating that moderate supplementation promotes structural development. The slight decline at higher inclusion levels supports a quadratic dose–response pattern, consistent with previous studies demonstrating optimal growth at intermediate phytogenic inclusion levels (Dawood et al., 2020; Hoseinifar et al., 2015; Ahmadifar et al., 2022). Fin development followed a similar trend, while some anatomical traits remained unaffected, suggesting selective rather than uniform morphological responses.

Biometric indices indicated enhanced digestive organ development, particularly increased visceral and gut weights at moderate supplementation levels, suggesting possible physiological adaptations related to nutrient utilization and digestive capacity (Abdel-Latif et al., 2020; Dawood et al., 2021; Alagawany et al., 2024). In contrast, liver weight and other somatic indices remained unchanged, indicating that supplementation selectively affected digestive organs without disrupting overall physiological balance. However, these interpretations were based primarily on biometric measurements, as intestinal histomorphology, digestive enzyme activity, and molecular analyses were not evaluated. Therefore, the proposed digestive-related mechanisms should be considered preliminary and require further confirmation through mechanistic studies.

Correlation analysis revealed allometric growth patterns, with somatic growth inversely associated with some visceral indices, reflecting proportional development between external body structures and internal organs. The strong relationship between intestinal length and relative gut length highlights coordinated digestive development, while the independence of condition factor from most parameters suggests stable physiological status across treatments.

This study was conducted under controlled tank conditions and over a 60-day feeding period; thus, extrapolation to commercial-scale systems and longer production cycles should be interpreted with caution. Environmental variability, stocking density, and management practices in field conditions may influence the magnitude of phytogenic responses.

Future studies should investigate gut histomorphology, digestive enzyme activity, and microbiome modulation to better elucidate the mechanisms underlying phytogenic-induced growth enhancement. Long-term feeding trials and validation under commercial farming conditions are also recommended to confirm the applicability of the optimal inclusion level. As the aqueous extract was applied using a coating/spraying method, partial leaching of water-soluble bioactive compounds into the culture water before feed consumption may have occurred, potentially causing slight differences between nominal and actual dietary exposure levels. Therefore, future research should assess nutrient leaching stability and explore encapsulation or binder-assisted delivery systems to improve dose precision and feed stability.

Taken together, the findings demonstrate that dietary supplementation with aqueous lemongrass extract enhances growth performance, feed efficiency, and digestive organ development in red tilapia, with optimal responses observed at approximately 20 g kg^-¹. These effects are likely mediated by bioactive compounds that improve digestive physiology and nutrient utilization without compromising hematological health, supporting the use of phytogenic additives as sustainable and welfare-oriented strategies in aquaculture systems.

Conclusion

Dietary supplementation with aqueous Cymbopogon citratus extract improved growth performance, feed utilization efficiency, and selected morphometric and biometric traits of Oreochromis spp. without adversely affecting hematological health or survival. Fish fed lemongrass-supplemented diets exhibited enhanced weight gain, feed conversion efficiency, and structural growth, suggesting improved nutrient utilization and physiological performance. Quadratic dose–response modeling indicated that optimal biological responses occurred at intermediate inclusion levels (~18–20 g kg^-¹), where growth performance and digestive organ development were greatest. Increases in visceral weight, gut weight, and intestinal length further suggest potential physiological adaptations associated with digestive development that may contribute to improved feed efficiency and growth. Importantly, the absence of significant hematological alterations indicates that lemongrass supplementation was physiologically safe at the tested inclusion levels. Overall, this study provides empirical evidence supporting the potential use of lemongrass aqueous extract as a phytogenic feed additive in tilapia culture systems. These findings support its application in sustainable and antibiotic-free aquaculture practices. However, because intestinal histomorphology, digestive enzyme activity, and molecular analyses were not evaluated, interpretations regarding digestive mechanisms remain preliminary. Future studies should therefore investigate these mechanisms further and validate the observed responses under commercial-scale production conditions.

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