Characterization of Gracilaria salicornia, Turbinaria conoides, and Kappaphycus alvarezii as functional ingredients for companion animal diets

Introduction

Climate change, rapid urbanization, shrinking agricultural land, and increasing competition between food, feed, and biofuel sectors are placing unprecedented pressure on conventional animal feed resources. Simultaneously, the growing companion animal population and increasing demand for functional, health-promoting pet foods have intensified the search for sustainable and nutritionally superior alternative feed ingredients. In this context, marine macroalgae (seaweeds) have emerged as promising biomass resources because of their high nutritional value, rich bioactive composition, and environmentally sustainable production systems (Morais et al., 2020; Tanna and Mishra, 2019) [15,30] . Unlike terrestrial feed crops, seaweeds do not require arable land, freshwater, or intensive fertilizer inputs, making them highly suitable for climate-smart animal nutrition strategies (Wan et al., 2019; Farghali et al., 2022) [8,32] . Global seaweed production currently exceeds 30 million tonnes annually, largely driven by aquaculture, highlighting their growing importance in future food and feed systems (Corino et al., 2019 [3] ; Jönsson and Karlsson, 2023) [13] . India, with an extensive coastline of nearly 8100 km and approximately 700 reported seaweed species, possesses enormous yet underutilized potential for seaweed-based feed applications (Dhingra and Kandiannan, 2021) [6] . However, despite this abundance, seaweeds in India are still predominantly utilized for phycocolloid extraction and agricultural applications, while their direct incorporation into animal nutrition, particularly companion animal feeding systems, remains limited (Pandey et al., 2023) [16] . This underexploited marine biomass therefore offers significant opportunities for developing sustainable, functional, and economically viable pet nutrition systems. Seaweeds are broadly classified into brown algae (Phaeophyceae), red algae (Rhodophyceae), and green algae (Chlorophyceae), each possessing distinct nutritional and biochemical characteristics (Leandro et al., 2020). Among the commercially important tropical seaweeds, Turbinaria conoides (brown seaweed), Gracilaria salicornia (red seaweed), and Kappaphycus alvarezii (red seaweed) have gained increasing attention because of their abundance, adaptability to tropical marine ecosystems, and high nutraceutical potential. Brown seaweeds such as Turbinaria conoides are particularly rich in fucoidan, alginate, polyphenols, and antioxidant metabolites that support gut health, oxidative balance, and immune modulation, whereas red seaweeds including Gracilaria salicornia and Kappaphycus alvarezii are important sources of carrageenans, sulfated polysaccharides, essential minerals, and functional bioactive compounds (Rengasamy et al., 2019; Costa et al., 2021) [4,21] . In general, seaweeds contain appreciable quantities of proteins, essential amino acids, polyunsaturated fatty acids, vitamins, dietary fiber, and minerals such as iodine, selenium, zinc, iron, calcium, and magnesium, along with diverse bioactive compounds possessing antioxidant, antimicrobial, anti-inflammatory, prebiotic, and immunomodulatory properties (Michalak et al., 2022) [14] . These multifunctional properties make seaweeds attractive as nutraceutical feed ingredients capable of improving digestive health, immunity, metabolic stability, and overall well-being in animals. The increasing interest in seaweed supplementation is primarily linked to their role as functional feed additives rather than complete feed replacements. Studies in livestock, poultry, aquaculture, and monogastric nutrition have demonstrated that moderate inclusion of seaweed-derived ingredients can improve gut microbiota, antioxidant status, immune responses, growth performance, and product quality (Ribeiro et al., 2021) [22] . Such benefits are particularly relevant in companion animal nutrition, where modern pet owners increasingly demand natural, health-oriented diets that support immunity, gastrointestinal health, healthy aging, and skin and coat quality. Furthermore, seaweeds can partially substitute conventional feed ingredients such as soybean meal and cereal-based components, thereby reducing dependence on terrestrial agricultural resources and easing pressure on food-feed competition (González-Meza et al., 2023) [10] . Beyond their nutritional importance, seaweed cultivation also contributes to carbon sequestration, nutrient recycling, coastal livelihood generation, and marine bioremediation, further strengthening their role in sustainable animal production systems (Sultana et al., 2022; Waqas et al., 2024) [28,33] . Despite their considerable potential, several challenges limit the broader application of seaweeds in companion animal diets. The nutritional composition of seaweeds varies significantly depending on species, geographical origin, season, environmental conditions, and processing methods, thereby affecting consistency and feed quality (Reddy et al., 2024) [20] . Additionally, some seaweeds may accumulate excessive levels of iodine, arsenic, cadmium, lead, and other potentially toxic compounds, raising concerns regarding safety when incorporated indiscriminately into animal diets (Michalak et al., 2022) [14] . Although processing methods such as washing and thermal treatments can reduce certain contaminants, careful species selection, controlled sourcing, standardized processing, and regulated inclusion levels remain essential to ensure nutritional safety and efficacy (Costa et al., 2021) [4] . Moreover, scientific information regarding the proximate composition, fiber fractions, mineral profile, and antioxidant potential of tropical seaweed species intended for companion animal nutrition remains limited, necessitating detailed species-specific evaluation before dietary incorporation. Therefore, the present study was undertaken to evaluate the proximate composition, crude fiber fractions, mineral profile, and antioxidant potential of Gracilaria salicornia, Turbinaria conoides, and Kappaphycus alvarezii for their potential application as functional ingredients in companion animal nutrition.

Materials and methods

The study was conducted at the Companion Animal Laboratory, Division of Animal Nutrition, ICAR-Indian Veterinary Research Institute (IVRI), Izatnagar, Uttar Pradesh, India. All experimental and laboratory procedures were performed in accordance with the guidelines approved by the IVRI-Institutional Animal Ethics Committee (IAEC) (Approval No. V-137(21)/4/2024-CPCSEA-DADF).

Collection and processing of seaweed samples

Fresh samples of the seaweeds Gracilaria salicornia (red seaweed), Turbinaria conoides (brown seaweed), and Kappaphycus alvarezii (red seaweed) were procured from AquAgri Processing Private Limited, Delhi, India. The laboratory analyses were carried out at the Companion Animal Laboratory, Division of Animal Nutrition, ICAR-Indian Veterinary Research Institute (IVRI), Izatnagar, Uttar Pradesh, India. The seaweed samples were taxonomically authenticated prior to analysis. Samples were thoroughly washed with distilled water to remove adhering salts, sand particles, epiphytes, and other impurities, followed by drying in a hot-air oven at 50-60 °C until constant weight was achieved. The dried samples were ground into fine powder using a laboratory grinder and stored in airtight containers for subsequent proximate, fiber, mineral, and antioxidant analyses.

Proximate composition analysis

Proximate composition of the seaweed samples was determined according to standard AOAC procedures. Parameters analyzed included moisture, crude protein, ether extract, crude fiber, total ash, and nitrogen-free extract. Crude protein content was estimated using the Kjeldahl method, ether extract by Soxhlet extraction, crude fiber through acid-alkali digestion, and total ash by incineration in a muffle furnace at 550 °C. Nitrogen-free extract was calculated by difference.

Fiber fraction analysis

Fiber fractions including neutral detergent fiber (NDF), acid detergent fiber (ADF), and acid detergent lignin (ADL) were analyzed using the Van Soest detergent fiber method.

The analysis was performed to evaluate the structural carbohydrate composition and dietary fiber characteristics of the seaweed samples.

Mineral composition analysis

Mineral composition of the seaweed samples was determined following wet acid digestion of the dried seaweed powders using a nitric acid-perchloric acid digestion mixture. Non-volatile minerals including calcium, phosphorus, sodium, potassium, magnesium, iron, zinc, copper, and manganese were quantified using Atomic Absorption Spectrophotometry (AAS), whereas volatile minerals such as selenium and iodine were analyzed using Inductively Coupled Plasma-Optical Emission Spectroscopy (ICP-OES). The mineral concentrations were expressed on a dry matter basis.

Preparation of seaweed extracts for antioxidant analysis

Seaweed extracts were prepared for antioxidant analysis using methanol/ethanol solvent extraction. Briefly, dried seaweed powder was mixed with the extraction solvent and incubated under continuous shaking for 24-48 h at room temperature. The extracts were filtered through Whatman No.1 filter paper and centrifuged, and the supernatants obtained were stored at 4 °C until further antioxidant assays were performed.

Antioxidant analysis 2.6.1 DPPH radical scavenging activity:

Antioxidant activity of the seaweed extracts was evaluated using the DPPH free radical scavenging assay. The reduction in DPPH absorbance was measured spectrophotometrically at 517 nm, and the radical scavenging activity was expressed as percentage inhibition.

Ferric Reducing Antioxidant Power (FRAP):

Ferric reducing antioxidant power (FRAP) assay was performed to assess the reducing capacity of the seaweed extracts. Absorbance was recorded at 593 nm, and antioxidant activity was expressed as Trolox equivalent antioxidant capacity.

Total Phenolic Content (TPC):

Total phenolic content of the seaweed extracts was estimated using the Folin-Ciocalteu colorimetric method. Absorbance was measured at 765 nm, and the results were expressed as mg gallic acid equivalents per gram of extract.

Total Flavonoid Content (TFC):

Total flavonoid content was determined using the aluminum chloride colorimetric assay. Absorbance was recorded at 415 nm, and the flavonoid concentration was expressed as mg quercetin equivalents per gram of extract.

Statistical analysis

All analyses were carried out in triplicate, and data were expressed as mean ± standard error. Statistical analysis was performed using one-way analysis of variance (ANOVA) with SPSS software. Differences among mean values were considered statistically significant at p<0.05.

Results

Proximate composition and fiber fraction analysis

The proximate and fiber composition of the three seaweed species revealed significant interspecies variation (P < 0.05). Gracilaria salicornia exhibited significantly higher crude protein content (16.8 ± 0.6%) compared with Turbinaria conoides (12.9 ± 0.4%) and Kappaphycus alvarezii (9.8 ± 0.5%). Total ash, crude fiber, NDF, ADF, and insoluble dietary fiber contents were significantly higher in T. conoides (p<0.05), indicating its comparatively greater mineral-rich and structural polysaccharide composition. Both G. salicornia and T. conoides demonstrated superior dietary fiber characteristics compared with K. alvarezii, particularly with respect to soluble dietary fiber, where G. salicornia recorded the highest value (21.4 ± 0.8%), followed closely by T. conoides (19.8 ± 0.7%), while K. alvarezii showed significantly lower levels (11.6 ± 0.5%) (p<0.05). Ether extract was significantly greater in T. conoides, whereas nitrogen-free extract was highest in K. alvarezii (p<0.05). Overall, the findings indicate that G. salicornia and T. conoides possessed comparatively superior nutritional and functional fiber profiles than K. alvarezii, supporting their greater potential as functional ingredients for companion animal nutrition.

Table 1:

Proximate composition and dietary fiber fractions of Gracilaria salicornia, Turbinaria conoides, and Kappaphycus alvarezii (DM basis). Values are expressed as mean ± standard error (SE). Data were analyzed using one-way analysis of variance (ANOVA) followed by Tukey's multiple comparison test to determine differences among seaweed species. Statistical significance was considered at p<0.05, while highly significant differences were considered at p<0. 01

Discussion

The growing demand for sustainable and functional feed resources has intensified interest in marine macroalgae as alternative ingredients for animal nutrition. Seaweeds possess several advantages over conventional feed resources because they are rich in bioactive compounds, minerals, dietary fiber, and antioxidant metabolites while requiring neither arable land nor freshwater for cultivation. Their multifunctional nutritional properties have attracted considerable attention in livestock and companion animal nutrition, particularly for the development of health-oriented and climate-smart feeding strategies. However, substantial interspecies variability exists in nutrient composition and bioactive potential, necessitating detailed characterization of individual seaweed species before their incorporation into animal diets. In the present study, marked differences were observed among Gracilaria salicornia, Turbinaria conoides, and Kappaphycus alvarezii with respect to proximate composition, fiber fractions, mineral profile, and antioxidant properties, indicating distinct functional attributes and nutritional applications for each species.

Proximate composition and fiber fraction analysis

The present study revealed significant species-specific variation in proximate composition and fiber fractions among Gracilaria salicornia, Turbinaria conoides, and Kappaphycus alvarezii, reflecting the influence of taxonomy, environment, and cultivation conditions. Such variability highlights the need for species-wise evaluation before using seaweeds in companion animal feed formulations. G. salicornia showed higher crude protein, consistent with reports describing Gracilaria spp. as moderate-to-high protein seaweeds with functional polysaccharides suitable for feed use (Rosemary et al., 2019; Purwaningsih et al., 2024) [18,23] . In contrast, K. alvarezii exhibited higher nitrogen-free extract, indicating carbohydrate dominance typical of carrageenan-rich red algae (Araújo et al., 2022; Islamy et al., 2025) [2,12] . Turbinaria conoides recorded higher ash, crude fiber, NDF, ADF, and insoluble dietary fiber, confirming the mineralrich and structurally complex nature of brown seaweeds due to high levels of alginate, fucoidan, and related polysaccharides (Diachanty et al., 2017; Rasyid et al., 2025) [7, 19] . In contrast, G. salicornia showed superior soluble dietary fiber, followed by T. conoides, indicating stronger prebiotic and gut-health potential (Sakthivel and Devi, 2015) [24] . T. conoides also exhibited higher ether extract, aligning with reports of diverse lipid fractions and bioactive fatty acids in brown seaweeds (Diachanty et al., 2017) [7] . Overall, G. salicornia and T. conoides demonstrated superior nutritional and functional profiles compared to K. alvarezii. However, reported variability in protein and fiber content of K. alvarezii across studies suggests strong dependence on season, strain, and cultivation conditions (Araújo et al., 2022) [2] . This underscores the need for standardized production systems and species-specific evaluation before large-scale application in functional pet nutrition.

Mineral composition analysis

Significant interspecific variation in mineral composition highlights the importance of species-specific selection of seaweeds for companion animal nutrition. Turbinaria conoides exhibited higher concentrations of essential trace minerals including K, Na, Mg, Fe, Zn, Cu, Mn, and Se, consistent with the mineral-rich nature of brown seaweeds (Siahaan et al., 2018; Yoganandham et al., 2018) [25,34] . This supports its potential as a natural source of trace minerals and electrolytes in functional pet feeds, as also reported in Turbinaria spp. (Rasyid et al., 2025) [19] . In contrast, Gracilaria salicornia showed higher Ca and P, in agreement with reports on red seaweeds as rich sources of structural macro-minerals (Aknaf et al., 2024) [1] . [29,31] . Regulatory assessments further emphasize strong species dependence in contaminant accumulation (Guo et al., 2023) [11] . Overall, T. conoides and G. salicornia demonstrated more balanced and desirable mineral profiles for functional feed applications, whereas K. alvarezii, despite its nutritional value, requires careful sourcing and controlled inclusion due to elevated iodine and heavy metal content.

Antioxidant analysis

Marked interspecific variation in TPC, TFC, and antioxidant activities (DPPH, FRAP, ABTS) reflects the speciesdependent nature of antioxidant potential in seaweeds and its strong association with phenolic and flavonoid content (González-Meza et al., 2023; Golu et al., 2025) [9,10] . Consistent correlations between TPC/TFC and radicalscavenging activity further confirm phenolics and flavonoids as key contributors to antioxidant capacity (Souid et al., 2024) [26] . Gracilaria salicornia exhibited the strongest antioxidant activity, supported by earlier reports on Gracilaria spp. showing high phenolic and flavonoid content with strong DPPH, ABTS, and FRAP activity (Subramanian et al., 2020) [27] . This reinforces its potential as an antioxidant-rich functional ingredient for companion animal nutrition. Turbinaria conoides showed moderate antioxidant activity, consistent with brown seaweed studies attributing bioactivity to phlorotannins, fucoxanthin, and other polyphenols (Morais et al., 2020; Paul et al., 2024) [15,17] . These compounds contribute to oxidative stress reduction, immune modulation, and gut health, supporting its functional feed potential (Michalak et al., 2022) [14] . In contrast, Kappaphycus alvarezii recorded lower [5,20] . However, improved antioxidant potential under optimized systems indicates strong environmental and processing dependence (Islamy et al., 2025) [12] . Overall, G. salicornia and T. conoides demonstrated superior antioxidant and biofunctional potential compared to K. alvarezii, primarily due to higher phenolic and flavonoid content. These findings align with evidence that phenolic-rich seaweeds serve as effective natural antioxidants and sustainable alternatives to synthetic additives in animal feed systems (Morais et al., 2020) [15] .

Conclusion

This study demonstrates clear species-specific variation in the proximate composition, mineral profile, and antioxidant potential of Gracilaria salicornia, Turbinaria conoides, and Kappaphycus alvarezii, highlighting their differential suitability as functional ingredients in companion animal nutrition. G. salicornia exhibited higher protein content, favorable soluble dietary fiber, and superior antioxidant activity, indicating strong potential as a nutraceutical-rich functional feed component. T. conoides showed enhanced mineral density, structural fiber fractions, and appreciable antioxidant capacity, supporting its role as a valuable natural source of trace minerals and bioactive compounds. In contrast, K. alvarezii was characterized by higher carbohydrate, iodine, and cadmium levels, suggesting nutritional potential but also raising concerns regarding safe inclusion without strict quality control. Overall, G. salicornia and T. conoides emerged as more balanced and functionally promising seaweed candidates for companion animal feed formulations due to their combined nutritional and bioactive advantages. However, the marked variability observed across species underscores the necessity for species-specific evaluation, standardized cultivation practices, and contaminant monitoring prior to large-scale utilization. These findings support the strategic use of selected seaweeds as sustainable, functional alternatives to conventional feed additives in animal nutrition systems.

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