Distinct Nutritional Functions of Leptadenia reticulata and Moringa oleifera in Nutrient Digestibility of Lactating Sahiwal Cattle

Introduction

Efficient utilization of dietary nutrients is a fundamental prerequisite for sustainable and profitable dairy production. In ruminants, productive performance depends not only on the quantity of nutrients supplied through feed but also on the efficiency with which these nutrients are digested, absorbed and utilized for productive and physiological functions. Nutrient digestibility directly influences the availability of energy, protein and structural carbohydrates required for maintenance, milk synthesis, reproduction and health (Van Soest, 1994; McDonald et al., 2011) [30] . Since feed expenditure constitutes the largest component of dairy production costs, improving nutrient utilization efficiency remains a major objective of modern ruminant nutrition (NRC, 2001; Owens and Basalan, 2016) [22,23] . The digestive efficiency of ruminants is closely associated with the activity of the ruminal microbial ecosystem. The rumen contains a highly specialized microbial population comprising bacteria, protozoa, fungi and archaea that ferment dietary substrates into volatile fatty acids, microbial proteins and other metabolites utilized by the host animal (Russell and Rychlik, 2001; Jami and Mizrahi, 2012) [28] . Adequate protein availability supports microbial International Journal of Agriculture and Food Science https://www.agriculturaljournals.com growth and microbial protein synthesis, whereas efficient fibre degradation increases the availability of fermentable substrates and energy for ruminal microorganisms (Hoover and Stokes, 1991; McDonald et al., 2011) [11] . Consequently, nutritional interventions that enhance protein and fibre digestion can substantially improve feed utilization efficiency and productive performance in dairy cattle.

In recent years, increasing emphasis on sustainable livestock production has stimulated interest in herbal feed additives as natural alternatives to conventional growth promoters and synthetic supplements. Plant-derived feed resources contain a variety of bioactive compounds, including flavonoids, tannins, saponins, alkaloids and phenolic compounds, which may influence ruminal fermentation, microbial activity and nutrient utilization (Patra and Saxena, 2009; Patra, 2010) [26,25] . Such phytogenic additives have attracted considerable attention because of their potential to improve animal productivity while maintaining animal health and product quality (Windisch et al., 2008; Amad and Zentek, 2023) [31,2] . Among the herbal resources evaluated for livestock feeding, Moringa oleifera has received considerable scientific attention because of its exceptional nutritional composition.

The leaves are rich in crude protein, essential amino acids, vitamins, minerals and antioxidant compounds, with reported crude protein contents ranging from 20 to 30% (Makkar and Becker, 1996; Moyo et al., 2011) [17,20] . In addition, Moringa oleifera contains several biologically active compounds, including flavonoids, phenolic acids and glucosinolates, which may positively influence ruminal fermentation and nutrient metabolism (Anwar et al., 2007; Gopalakrishnan et al., 2016) [3,10] [1,6,7,14,27,29] . These beneficial effects have largely been attributed to its high protein content and ability to support microbial growth and fermentation efficiency within the rumen (Fadiyimu et al., 2010; Liu et al., 2023) [9] . Another important medicinal plant used in traditional livestock production systems is Leptadenia reticulata (Jivanti). The plant is valued for its galactagogue, immunomodulatory and healthpromoting properties and has been widely used in indigenous veterinary practices (Patel et al., 2016; Kumari et al., 2024) [24,15] . Phytochemical investigations have identified the presence of flavonoids, terpenoids, alkaloids, glycosides, steroids and phenolic compounds that contribute to its biological activities (Mohanty et al., 2017; Kumari et al., 2024) [19,15] . Unlike Moringa oleifera, Leptadenia reticulata possesses a comparatively higher fibre content, suggesting its potential role in influencing fibre degradation and structural carbohydrate utilization. Fibre-rich feed resources are known to stimulate fibrolytic microbial populations responsible for the degradation of cellulose and hemicellulose in the rumen, thereby improving the utilization of fibrous feed components (Van Soest, 1994; Russell and Rychlik, 2001) [28,30] . However, information regarding the influence of Leptadenia reticulata on nutrient digestibility in dairy cattle remains limited. The response of nutrient digestibility to herbal supplementation is not necessarily uniform across all nutrient fractions. Variations in nutrient composition and phytochemical characteristics may result in differential effects on protein digestion, fibre degradation and overall feed utilization (Patra and Saxena, 2009; Bashar et al., 2020) [26. 5] . Therefore, understanding the specific nutritional functions of different herbal feed resources is essential for developing targeted feeding strategies aimed at improving nutrient utilization efficiency in dairy cattle.

Although numerous studies have independently investigated the nutritional benefits of Moringa oleifera and the medicinal properties of Leptadenia reticulata, direct comparative evaluations of their effects on nutrient digestibility under identical feeding and management conditions are scarce. Furthermore, information regarding their influence on individual nutrient fractions in indigenous dairy breeds such as Sahiwal cattle remains limited. Based on their contrasting nutrient profiles, it was hypothesized that Moringa oleifera would preferentially improve dry matter and crude protein digestibility, whereas Leptadenia reticulata would exert a greater influence on crude fibre utilization. Therefore, the present study was undertaken to compare the effects of dietary supplementation with Leptadenia reticulata root powder and Moringa oleifera leaf powder, administered individually and in combination, on the digestibility of major nutrient fractions in lactating Sahiwal cattle under farm conditions.

Materials and Methods

Experimental Site

The present investigation was conducted at the Instructional Dairy Farm (IDF), Nagla, College of Veterinary and Animal Sciences, Govind Ballabh Pant University of Agriculture and Technology (GBPUA&T), Pantnagar, Uttarakhand, India. The experimental farm is located at approximately 29.0°N latitude and 79.3°E longitude with an altitude of about 243.8 m above mean sea level. The study was carried out under uniform management and feeding conditions during a 120-day experimental period.

Ethical Approval

The experimental protocol was approved by the Institutional Animal Ethics Committee (IAEC), College of Veterinary and Animal Sciences, GBPUA&T, Pantnagar, Uttarakhand, India. All animal handling and experimental procedures were conducted in accordance with the guidelines of the Committee for the Purpose of Control and Supervision of Experiments on Animals (CPCSEA), Government of India.

Experimental Animals and Design

Twenty-eight clinically healthy lactating Sahiwal cattle maintained at the university dairy farm were selected for the study. The animals were selected on the basis of similar parity, body condition score and stage of lactation to minimize variation among treatment groups and were maintained under similar managemental conditions throughout the experiment. The animals were randomly allocated into four treatment groups consisting of seven animals each following a Completely Randomized Design (CRD). Prior to initiation of the feeding trial, all animals were acclimatized to the experimental conditions for one week.

The experimental treatments were as follows: T₁ Basal diet + Leptadenia reticulata root powder @ 90 g/animal/day T₂ Basal diet + Moringa oleifera leaf powder @ 150 g/animal/day T₃ Basal diet + Leptadenia reticulata root powder @ 45 g + Moringa oleifera leaf powder @ 75 g/animal/day

The feeding trial was conducted for a period of 120 days.

Feeding and Management

All experimental animals were maintained in a loose housing system with free access to clean drinking water throughout the experimental period. Animals were fed according to the prevailing feeding schedule of the dairy farm.

The basal ration consisted of concentrate mixture, wheat straw and seasonal green fodders, including berseem (Trifolium alexandrinum), oat (Avena sativa) and mustard greens. Herbal supplements were individually administered once daily during the afternoon milking session to ensure complete consumption. Routine management practices, including feeding, milking, sanitation and healthcare, were maintained uniformly for all treatment groups throughout the study.

Digestibility Trial

A five-day digestion trial was conducted at the end of the 120-day feeding period to determine nutrient digestibility coefficients. During the digestibility trial, the quantity of feed offered and feed refusals was recorded daily for each experimental animal. Representative fecal samples were collected daily from individual animals throughout the collection period. The daily samples from each animal were pooled proportionately to obtain a representative composite sample, which was subsequently processed, preserved and stored for laboratory analysis. Feed and fecal samples were dried, ground and stored in airtight containers until chemical analysis.

Chemical Analysis

Representative samples of concentrate mixture, roughages, Leptadenia reticulata root powder, Moringa oleifera leaf powder and feces were analyzed for proximate constituents according to the standard procedures of the Association of Official Analytical Chemists (AOAC, 2005) [4] . The analyzed parameters included dry matter (DM), organic matter (OM), crude protein (CP), ether extract (EE), crude fibre (CF) and total ash (TA). Crude protein was estimated using the Micro-Kjeldahl method, whereas ether extract was determined using Soxhlet extraction. Crude fibre and total ash were estimated using standard gravimetric procedures. Nitrogen-free extract (NFE) was calculated by difference.

Calculation of Nutrient Digestibility

Digestibility coefficients of dry matter, organic matter, crude protein, ether extract, crude fibre and nitrogen-free extract were calculated using feed intake and fecal output data according to the following equation:

The calculated digestibility coefficients were expressed as percentages.

Statistical Analysis

The experimental data were analyzed using Statistical Package for Social Sciences (SPSS) software. Treatment effects on nutrient digestibility parameters were evaluated using one-way analysis of variance (ANOVA) under a Completely Randomized Design. The statistical model used was:

where:  Yᵢⱼ= observation on the jᵗʰ animal receiving the i th treatment,  μ= overall mean,  Tᵢ= effect of the iᵗʰ treatment,  eᵢⱼ= random experimental error.

When significant treatment effects were detected, treatment means were separated using Duncan's Multiple Range Test (DMRT). Results are presented as Mean±Standard Error (SE), and differences were considered significant at P < 0.05.

Results and Discussion

Chemical Composition of Feed Ingredients and Herbal Supplements

The proximate composition of the feed ingredients and herbal supplements used during the experimental period is presented in Table 2. Considerable variation was observed among the feed resources with respect to their nutrient composition, reflecting differences in their nutritional quality and potential contribution to nutrient utilization. Such variations are expected to influence ruminal fermentation patterns and digestibility responses in lactating cattle. Among the herbal supplements, Moringa oleifera leaf powder exhibited substantially higher crude protein (27.05%) and ether extract (6.33%) contents than Leptadenia reticulata. The high crude protein concentration highlights the potential of Moringa as a protein-rich feed resource capable of supporting microbial growth and nitrogen metabolism within the rumen. Similar nutrient profiles have been reported by Makkar and Becker (1996) [17] , Moyo et al. (2011) [20] and Gopalakrishnan et al. (2016) [10] , who identified Moringa leaves as an excellent source of protein, minerals and bioactive compounds.

In contrast, Leptadenia reticulata root powder was characterized by markedly higher crude fibre (43.20%) and total ash (14.72%) contents. [19,15] .

The contrasting nutrient composition of the two herbal supplements suggests that they may influence nutrient utilization through different physiological mechanisms. Whereas Moringa oleifera may primarily enhance nitrogen availability and microbial protein synthesis, Leptadenia reticulata may exert greater influence on fibre degradation and utilization. These compositional differences provide a biological basis for the differential digestibility responses observed in the present investigation.

Effect of Herbal Supplementation on Nutrient Digestibility

The effects of dietary supplementation with Leptadenia reticulata and Moringa oleifera on nutrient digestibility are presented in Table 3. Significant treatment effects (P<0.05) were observed for dry matter, crude protein and crude fibre digestibility, whereas organic matter, ether extract and nitrogen-free extract digestibility remained statistically comparable among treatment groups. 1. [29] , Kholif et al. (2015) [14] , Roy et al.

(2016) [27] , Abdel-Raheem and Hassan (2021) [1] and El-Badawi et al. (2023) [7] .

International Journal of Agriculture and Food Science https://www.agriculturaljournals.com

The numerical improvement observed in the combined treatment group suggests that the beneficial effects of Moringa on dry matter utilization were partially retained when administered with Leptadenia reticulata. However, the response was lower than that observed in the Moringaonly treatment, indicating that the protein-rich nature of Moringa may have played a dominant role in improving dry matter digestion.

Crude Protein Digestibility

Crude protein digestibility was significantly influenced by dietary treatment, with the highest value observed in the Moringa oleifera supplemented group (53.32%). This value was significantly higher than those recorded in the control, Leptadenia reticulata and combination treatment groups.

The enhanced crude protein digestibility associated with Moringa supplementation may be attributed to its high crude protein concentration and favorable amino acid profile. Increased availability of degradable nitrogen supports microbial protein synthesis and improves nitrogen utilization efficiency within the rumen. Enhanced microbial activity facilitates greater degradation of dietary protein, resulting in improved protein digestibility and nutrient utilization. Similar findings have been reported by Kholif et al. (2015) [14] , Roy et al. (2016) [27] , Cohen-Zinder et al.

(2016) [6] and El-Hamd et al. (2024) [8] . Furthermore, the presence of antioxidant and phytogenic compounds in Moringa leaves may contribute to stabilization of the ruminal environment, thereby promoting efficient microbial metabolism and nutrient degradation. The substantial improvement in both dry matter and crude protein digestibility observed in the present study supports the hypothesis that Moringa oleifera functions primarily as a protein-oriented nutritional supplement capable of enhancing nitrogen utilization in lactating dairy cattle.

Crude Fibre Digestibility

Crude fibre digestibility was significantly (P<0.05) influenced by dietary supplementation and exhibited a response pattern distinct from that observed for dry matter and crude protein digestibility. The highest crude fibre digestibility was recorded in animals receiving Leptadenia reticulata supplementation (50.48%), followed by the Moringa oleifera supplemented group (45.59%) and the combination treatment (45.12%). The lowest crude fibre digestibility was observed in the control group (43.66%). The superior fibre digestibility observed in the Leptadenia reticulata supplemented group may be attributed to its comparatively higher crude fibre content and unique phytochemical composition. Fibre-rich feed resources are known to stimulate fibrolytic microbial populations responsible for the degradation of cellulose and hemicellulose in the rumen. Enhanced fibrolytic activity increases the breakdown of structural carbohydrates and improves the utilization of fibrous feed components (Van Soest, 1994; Russell and Rychlik, 2001) [28,30] . In addition to its fibre content, Leptadenia reticulata contains several bioactive compounds, including flavonoids, phenolic compounds and glycosides, which may influence ruminal microbial ecology and digestive activity (Mohanty et al., 2017; Kumari et al., 2024) [19,15] . Improved microbial adaptation to fibrous substrates may have contributed to the greater crude fibre digestibility observed in the present study. Enhanced fibre degradation improves the release of fermentable carbohydrates and ultimately contributes to better nutrient utilization efficiency.

Interestingly, the highest crude fibre digestibility was not observed in the Moringa oleifera supplemented group despite its positive effects on dry matter and crude protein digestibility. This finding suggests that the two herbal supplements exerted different nutritional influences on ruminal digestion. While Moringa oleifera primarily enhanced protein utilization and microbial growth through improved nitrogen availability, Leptadenia reticulata appeared to favor fibre degradation and structural carbohydrate utilization. Such differential responses highlight the importance of considering nutrient composition when selecting herbal feed additives for specific nutritional objectives.

The findings of the present investigation support the hypothesis that fibre-rich herbal supplements may preferentially improve fibre utilization, whereas protein-rich supplements may exert greater effects on nitrogen metabolism and crude protein digestion. These results indicate that the digestibility responses observed in the present study were closely associated with the nutrient composition of the herbal supplements.

Organic Matter, Ether Extract and Nitrogen-Free Extract Digestibility

Dietary supplementation with Leptadenia reticulata and Moringa oleifera did not significantly (P>0.05) influence organic matter, ether extract or nitrogen-free extract digestibility. Although numerical differences were observed among treatment groups, the magnitude of variation was insufficient to attain statistical significance. Organic matter digestibility ranged from 38.33% to 44.42% and did not differ significantly (P>0.05) among treatment groups. Although numerical variation was observed, dietary supplementation with Leptadenia reticulata and Moringa oleifera did not produce a measurable effect on overall organic matter utilization. The absence of significant differences suggests that the herbal supplements exerted their primary influence on specific nutrient fractions, particularly crude protein and crude fibre digestibility, rather than on total organic matter digestion. Ether extract digestibility remained consistently high across all treatment groups, varying between 79.16% and 80.80%. The lack of significant differences may indicate that lipid digestion and absorption were already highly efficient under the prevailing feeding conditions and therefore were not substantially influenced by herbal supplementation.

Comparable findings have been reported by Roy et al.

(2016) [27] and Abdel-Raheem and Hassan (2021) [1] , who observed minimal effects of herbal feed additives on ether extract digestibility in ruminants.

Similarly, nitrogen-free extract digestibility remained statistically comparable among treatment groups, with values ranging from 72.99% to 74.10%. Nitrogen-free extract primarily represents readily fermentable carbohydrate fractions such as soluble sugars and starches, which are generally highly digestible under normal ruminal conditions. Consequently, supplementation with the herbal feed additives used in the present study may not have been sufficient to produce measurable alterations in their digestion and utilization. The absence of significant effects on organic matter, ether extract and nitrogen-free extract digestibility indicates that the principal influence of the herbal supplements was directed towards specific nutrient fractions rather than generalized enhancement of all digestive processes. This selective response further supports the concept of differential nutritional functionality between Moringa oleifera and Leptadenia reticulata.

Conclusion

Dietary supplementation with Moringa oleifera and Leptadenia reticulata exerted differential effects on nutrient digestibility in lactating Sahiwal cattle. Significant improvements were observed in dry matter, crude protein and crude fibre digestibility, whereas organic matter, ether extract and nitrogen-free extract digestibility remained unaffected by dietary treatments. Among the evaluated supplements, Moringa oleifera was more effective in improving dry matter and crude protein digestibility, indicating its potential as a protein-rich herbal feed additive for enhancing nutrient utilization and feed efficiency. In contrast, Leptadenia reticulata resulted in the highest crude fibre digestibility, suggesting its suitability for improving the utilization of structural carbohydrates in roughage-based feeding systems.

The findings demonstrate that the two supplements perform distinct nutritional functions rather than exerting uniform effects on all nutrient fractions. While Moringa oleifera primarily enhanced dry matter and protein utilization, Leptadenia reticulata contributed more effectively to fibre digestion. These results suggest that the selection of herbal feed additives should be guided by specific nutritional objectives. Moringa oleifera may be preferred for improving protein utilization and feed efficiency, whereas Leptadenia reticulata may be strategically utilized to enhance fibre digestion in roughagebased diets. Further studies on rumen fermentation characteristics, microbial ecology and long-term production responses are warranted to elucidate the biological mechanisms underlying these differential digestibility responses.

Highlights

 Supplementation with Moringa oleifera significantly improved dry matter and crude protein digestibility in lactating Sahiwal cattle.  Leptadenia reticulata supplementation resulted in the highest crude fibre digestibility among the treatment groups.  The two herbal supplements exhibited distinct nutritional functions, indicating nutrient-specific effects on digestibility.  Organic matter, ether extract and nitrogen-free extract digestibility were not significantly influenced by dietary treatments.  Moringa oleifera favored protein utilization, whereas Leptadenia reticulata enhanced fibre digestion under farm conditions.

Practical Implications

The findings of the present study suggest that Moringa oleifera and Leptadenia reticulata can be strategically utilized as natural herbal feed additives according to specific nutritional objectives in dairy cattle. Supplementation with Moringa oleifera improved dry matter and crude protein digestibility, indicating its potential as a protein-rich feed resource for enhancing nutrient utilization and feed efficiency. In contrast, Leptadenia reticulata improved crude fibre digestibility, suggesting its suitability for roughage-based feeding systems where enhanced fibre utilization is desired. Therefore, both herbal supplements may serve as sustainable and plant-based nutritional interventions for improving feed utilization efficiency in lactating Sahiwal cattle under farm conditions.

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