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Review

Aquatic Plants for Blue Protein Innovation: Bridging Nutrition, Sustainability, and Food Security

by
Anil Kumar Anal
1,*,
Abhishek Khadka
1,
Daniel Lee Rice
1,
Nabindra Kumar Shrestha
1,
Johnmel Abrogena Valerozo
1,2,
Khin Nyein Chan Zaw
1 and
Ryunosuke Nagase
1
1
Department of Food Agriculture and Bioresources, Asian Institute of Technology, Pathum Thani 12120, Thailand
2
College of Agriculture, Food and Sustainable Development, Mariano Marcos State University, Batac Campus, City of Batac 2906, Ilocos Norte, Philippines
*
Author to whom correspondence should be addressed.
Resources 2025, 14(12), 192; https://doi.org/10.3390/resources14120192
Submission received: 24 October 2025 / Revised: 12 December 2025 / Accepted: 15 December 2025 / Published: 18 December 2025
Review Reports Versions Notes

Abstract

The global population is rising sharply and is expected to be 10 billion by 2050. Nutrition security, especially protein, is a major concern, as it is one of the essential ingredients for body growth. However, consumption of meat is unsustainable, as the use of natural resources and greenhouse gas (GHG) emissions are relatively high compared to plant-based protein sources. Aquatic plants like duckweed, Azolla, and water spinach, as well as macroalgae and microalgae, contain good amounts of protein, ranging from 25% to 60% dry weight (DW) and comprising major essential amino acids (EAAs). These plants are rich in vitamins and minerals and possess antimicrobial, anti-inflammatory, antidiabetic, and anti-fatigue properties. In addition, green food processing (GFP) technologies minimize the antinutritional factors, which in turn increase the bioaccessibility and biodigestibility of aquatic plants. Fermentation is one of the oldest known GFP methods. Recent advances include high-pressure processing, pulsed electric field, ultrasound-assisted, and microwave-assisted extraction, which are among the most promising techniques. Hence, government initiatives, as well as research and private sector collaboration for cultivation, processing, and advocating for such nutrient-dense food, are necessary. This will ensure sustainable production and consumption.

1. Introduction

Population growth, increasing incomes, and dietary changes toward a greater intake of foods produced from animals are all contributing to the increase in the need for protein worldwide. Recent estimates indicate that by 2050, there will be around 10 billion people on the planet, necessitating a roughly 70% upsurge in food production to keep up with demand [1]. In addition to requiring large amounts of natural resources, traditional sources of animal protein, such as meat, dairy, and fish, are linked to high greenhouse gas emissions, loss of biodiversity, and other environmental stresses. Therefore, relying on traditional animal proteins will become more challenging [2,3].
Considering this, there is rising interest in finding more ecologically friendly, efficient, and sustainable protein sources, such as plant proteins. Plant proteins have long been utilized to supplement protein, replacing animal sources to some extent, but current agricultural production cannot meet the growing demand [4]. Aquatic plants are an underexplored resource and an alternative output to meet the global protein demand. Such aquatic plants and organisms are promising, as they can fix nitrogen, use water or non-arable land, exhibit moderate-to-high protein, and are climate-resilient. These include freshwater species of fast-growing duckweed, leafy vegetables, and ferns, to slow-growing seed crops like lotus and fox nut, as well as commonly marine-sourced microalgae and macroalgae. For example, duckweed has a photosynthetic enzyme called RuBisCo (ribulose biphosphate carboxylase/oxygenase), which catalyzes the initial stages of carbon fixation. Similarly, microalgae synthesize carbohydrates through photosynthesis and carbon fixation, enabling them to capture CO2 from the atmosphere [5,6].
Protein content varies among the groups, with moderate levels of around 15–25% in vegetable crops and ferns, moderate-to-high levels in duckweed and macroalgae, and high protein content (40–70%) in microalgae. This indicates the great potential of these crops as blue protein resources. Also, many plant resources contain high fiber and low fat and exhibit high bioactivities. Additionally, while some of the sources are commercially cultivated, many have yet to be fully explored or industrialized [5,6]. However, like with many plant sources, the presence of antinutrients and other components reduces the bioavailability and digestibility of the products. Therefore, to ensure environmental sustainability by utilizing such plants as the next protein sources, green food processing (GFP) is recommended. GFP ensures that water consumption is minimal and allows byproducts to be recycled, ensuring a safe and high-quality product [7].
Environmentally safe and efficient cultivation, nutrition, and processing are essential to navigating a sustainable global food chain and developing a blue economy from aquatic plant sources. A blue economy refers to the sustainable use of aquatic resources for economic growth [8] and development while maintaining circularity and ecosystem health. Aquatic plants in particular offer hidden potential as protein sources to meet the growing economic demand while also protecting and maintaining aquatic ecosystems.
This review explores the wide range of aquatic plant species that offer great potential as a sustainable blue protein source. It covers the cultivation, nutrition, digestibility, bioactivity, and overall sustainability for several critical moderate-to-high-protein aquatic plant sources. Although most previous reviews highlight individual plant species for aquatic blue protein resources, this review takes a broader view across many potential aquatic plant species, providing insights into protein quality, functional properties, and potential processing avenues. Both conventional and underexplored species are explored to provide a holistic view of aquatic plants as a blue protein resource to meet the global protein demand while also promoting sustainable food systems. The review encompasses the cultivation and nutritional quality of potential aquatic resources in both fresh and salt water, potential sustainable green processing strategies to improve quality, and the overall impact on sustainability and the blue economy that is essential for global food production.

2. Methodology

This review was conducted through a structured and systematic search of the recent scientific literature focusing on alternative protein sources, aquatic plants, microalgae, seaweeds, bioprocessing innovations, nutritional composition, and sustainable food systems. Relevant publications were identified from reputable academic databases, including ScienceDirect, PubMed, SpringerLink, Wiley Online Library, Google Scholar, and additional credible repositories. The search strategy employed targeted keywords such as “alternative protein sources,” “microalgae nutrition,” “seaweed bioactive compounds,” “duckweed protein,” “aquatic plant bioprocessing,” “sustainable food production,” “functional foods,” “agro-industrial valorization,” and “circular bioeconomy.” Only peer-reviewed journal articles, book chapters, and review papers published between 2014 and 2025 were included to ensure comprehensive and contemporary coverage, while older studies were cited only when necessary to provide historical context. All collected materials were screened for methodological robustness, thematic relevance, and scientific contribution. Publications that were duplicated, lacked peer review, or did not align with the scope of this review were excluded. A total of 131 relevant sources were synthesized, encompassing topics such as microalgal and seaweed biorefinery, duckweed nutritional evaluation, phytochemical characterization of aquatic plants, advanced extraction technologies, sustainable aquaculture integration, and global perspectives on future protein production. This systematic approach enabled the development of a comprehensive synthesis highlighting nutritional potential, processing innovations, environmental implications, and industrial applications of emerging sustainable protein and functional food resources.

3. Aquatic Plant Resources

Plant protein production and utilization have grown massively in recent years [9], while many aquatic plants remain underutilized in this context. Aquatic plants offer another sustainably sourced moderate-to-high protein option. These plants grow in or around water, including both freshwater and marine sources, ranging from underexplored to historically produced crops. Including duckweed, aquatic vegetables and ferns, macroalgae, and microalgae, each of these groups showcases plants and species of great potential for nutritious and high-quality blue protein.

3.1. Fresh Water

3.1.1. Duckweed

Duckweeds, belonging to the family Lemnaceae, are recognized as the smallest and most rapidly multiplying angiosperms. They comprise about 36 species across five genera: Spirodela, Landoltia, Lemna, Wolffia, and Wolffiella [10]. Globally distributed across freshwater habitats, duckweeds are notable for their rapid vegetative propagation, high resilience to diverse environmental conditions, and remarkable capacity to store proteins and carbohydrates. Their unique biological traits make them attractive for future food systems, offering high biomass yields under minimal inputs, often in non-arable aquatic environments [11,12]. Among the genera, Wolffia, especially W. globosa, has been traditionally consumed in Asian diets and is highly valued for its nutritional quality [11,13]. Meanwhile, starch-rich species such as Landoltia and Spirodela are increasingly considered for feed and biorefinery purposes [13,14].
The cultivation of Lemnaceae for human food increasingly emphasizes controlled and hygienic systems to guarantee safety, nutritional consistency, and compliance with food-grade standards. While all five genera can naturally thrive in ponds, large-scale food production requires strictly controlled conditions. Lemna spp. show rapid doubling rates, environmental tolerances, and protein content that can surpass 30% [10]. Furthermore, Spirodela polyrhiza or giant duckweed demonstrates 25% protein, fast production, and moderate fiber [15]. W. globosa and W. arrhizal have been traditionally consumed by humans in Southeast Asia [12,13,16]. For modern applications, indoor hydroponic vertical farms with LED lighting are increasingly preferred, allowing precise control of nutrient supply, photoperiod, and water quality to produce consistent, protein-rich biomass with bioavailable Vitamin B12 and a balanced amino acid profile [17].
As shown in Table 1, duckweeds generally contain 20–45% protein, 25–50% carbohydrates, and 3–7% lipids, alongside diverse vitamins and minerals, positioning them as sustainable alternatives to terrestrial vegetables and conventional plant proteins [13]. This protein profile makes duckweeds a good candidate to mitigate amino acid deficiencies in combination with other plants, as cereal grains are often limited in lysine, while methionine is limited in legumes. W. globosa and W. arrhiza are the most protein-dense, reaching up to 45% dry weight, with digestible indispensable amino acid scores (DIAASs) comparable to egg and dairy proteins [18]. These contain a good amount of PUFA (alpha-linolenic acid and linolenic acid), whereas Lemna minor offers a good omega-3-to-omega-6 ratio, supporting cardiovascular and cognitive health [19].
The digestibility and bioavailability of the Lemnaceae family are high because of their simple cell wall structure and presence of low lignin. In vitro assays report digestibility above 80% for several species, with Lemna averaging 72%, Wolffia 69%, and Spirodela substantially lower (39%) [23]. Among the genera, Wolffia globosa and W. arrhiza exhibit the most promising protein quality, with clinical studies demonstrating amino acid bioavailability comparable to dairy and legumes [24]. Micronutrient bioavailability is most validated in Wolffia, where human trials confirm effective absorption of iron, zinc, and the rare plant-derived Vitamin B12 [24,25,26]. By contrast, antinutritional factors such as oxalates and phytates in Lemna and Spirodela can restrict mineral uptake, though blanching or fermentation substantially mitigate these effects [13].
Taken together, Wolffia and Lemna emerge as the most relevant for direct human consumption, supported by robust digestibility, validated nutrient bioavailability, and established food use in Southeast Asia, while Spirodela, Landoltia, and Wolffiella hold potential mainly in processed or blended applications [27,28,29]. Thus, duckweeds are present as a clear blue protein resource for sustainable production, although improvements in processing and scalable production are critical for global utilization.

3.1.2. Floating Vegetables and Ferns

Freshwater vegetables and ferns are another key resource for underutilized plant proteins. This includes Azolla (Azolla spp.), water spinach (Ipomoea aquatica), and watercress (Nasturtium officinale), among others. As shown in Table 2, these plants exhibit a protein content of approximately 20–30%, are rich in micronutrients, and possess important health-promoting bioactivities. These crops are widely available and cultivated in various regions globally and are very common in Asia, with many demonstrating climate resilience and tolerance. To varying degrees, these resources are already produced. Among the floating plants, Azolla are small freshwater floating ferns, well known for their high protein content and symbiotic relationship with cyanobacteria [30,31].
Azolla is most common in Vietnam, China, and the Philippines, but due to its high potential and versatile uses, it has spread to many parts of the world (Europe, Africa, and Iran). Furthermore, it thrives in tropical and subtropical regions and grows in many sources, including rivers, drainages, ponds, rice paddies, and more [32]. Azolla doubles its biomass every 5–6 days, fixes nitrogen due to the symbiotic cyanobacteria, and is a strong phytoremediator. Current cultivation methodology has primarily targeted phytoremediation and highly contaminated or mineral-rich waters. As such, Azolla is minimally consumed by humans, but has been used as animal feed in tilapia, chickens, and ruminants, showcasing strong potential as a novel protein source with cleaner cultivation systems. Azolla’s biomass is approximately 25% crude protein, rich in essential amino acids, vitamins, minerals, and phytohormones [33,34]. Furthermore, Winstead et al. (2024) [35] reported moderate protein biodigestibility of 78.45% in Azolla carliniana, which is similar to levels found in many common legumes and likely linked to antinutritional factors (ANFs) within the plant. Despite the high protein content, rapid growth, and moderate digestibility, Azolla spp. show some risk for human consumption through cyanobacterial neurotoxin production or heavy metal absorption [35]. Thus, strict regulation, cultivation strategies, and policy are critical to investigating Azolla spp. for human consumption [32].
Table 2. Protein and nutrition profile of floating vegetables, ferns, and seed-bearing crops.
Table 2. Protein and nutrition profile of floating vegetables, ferns, and seed-bearing crops.
CropProtein Content (DW)EAA ProfileVitamins/BioactivitiesReferences
Azolla (Azolla spp.)~20–25%High in Lys, MetAntimicrobial, antioxidant, anticancer, anti-inflammatory[36,37]
Ipomoea aquatica (water spinach)~20%All EAAs
Low in Trp, His, Thr		Vitamins B1, C, A
Anticancer, antihyperglycemic		[38,39]
Nasturtium officinale (watercress)6–30%-Lutein and β-carotene; antioxidant, anti-inflammatory, anticancer[40,41,42]
Nymphae spp. (water lilies) 23% (rhizome)High in Leu, good EAA
Limiting: Met		Antimicrobial, anti-inflammatory, anti-depressant[43,44]
Nelumbo nucifera (lotus seed)16–28%36% EAA to TAA, variable studies, Limiting: Phe, Lys, Leu, and Tyr Vitamin B, Zn, Fe, Ca, Mn
Anticancer, antidiabetic, neuroregenerative		[45,46]
Euryale ferox (fox nut)9.7–11.16%Rich in EAA; Leu, Ile, Met, Cys, Arg, and Glu are the highestVitamins C and E, Mg, K, P, Fe, Zn
Antidiabetic, anti-fatigue, anti-aging		[47,48,49]
Leafy vegetables like water spinach (Ipomoea aquatica), watercress (Nasturtium officinale), and water lilies (Nymphaea lotus) are underutilized, nutrient-rich plants commonly consumed in Asia and globally. They are cultivated in a wide range of aquatic systems, including freshwater ponds, wetlands, ditches, marshes, and rice fields throughout the year [50,51]. Water spinach, watercress, and water lilies have demonstrated protein highs of 19.55%, 31.4%, and 23.01% (rhizome) on a dry basis, respectively. However, the range of proteins is variable between studies for these crops, suggesting that cultivation and species are critical. Additionally, all essential amino acids (EAAs) are present in water lilies and water spinach, although water spinach is low in tryptophan, threonine, and histidine, and water lilies are low in methionine [37,42]. These aquatic plants are rich in vitamins, minerals, and bioactivities that increase their potential and relevance as moderate protein sources, as shown in Table 2. Water spinach is high in Vitamin C, E, and riboflavin and exhibits anticancer, antioxidant, and anti-inflammatory characteristics. Alternatively, watercress also exhibited high nutrient content and has shown strong antioxidant, anti-inflammatory, and anticancer potential [52]. However, like many leafy greens, water spinach contains ANFs that can interfere with nutrient absorption, although processing and cooking reduce these [53]. The development of further processing and cultivation strategies is essential for developing these crops as sustainable and functional sources of blue protein. Their rich nutritional composition and use of underutilized land segments will be essential for a sustainable food supply chain.

3.1.3. Seed-Bearing Crops

While many aquatic plants may not present high protein in their leaves or rhizomes, several varieties generate seeds with moderate-to-high protein and overall nutritional value. Beyond the leafy plants, seed- and fruit-bearing plants like lotus (Nelumbo nucifera) and fox nut (Euryale ferox) offer an alternative to aquatic-sourced protein. Seed-bearing crops exhibit longer-term growth, improving ecological niches and demonstrating long-term carbon capture, while producing moderate-to-high protein products with strong bioactive potential [54]. However, these crops have less established cultivation and industrialization.
Among these, Nelumbo nucifera is widely cultivated, and every part of the lotus plant, i.e., seeds, rhizomes, leaves, and flowers, is edible and valued for its nutritional richness, medicinal properties, and industrial applications [55]. The leaves and flowers of the lotus plant are commonly used to make teas and extracts, which provide water-soluble phytochemicals like flavonoids and alkaloids [56]. On the other hand, the seeds in particular exhibit potential as a micronutrient-rich and high-protein resource at 16–28% DW [45]. Lotus is typically cultivated in ponds and lakes, and the seeding varieties take multiple weeks to produce the highly nutritious seeds, but the fresh seeds exhibit rapid quality degradation [57]. However, raw seeds contain antinutritional factors such as phytic acid and tannins, which can hinder nutrient absorption. Thermal treatments like boiling have been shown to effectively reduce these antinutrients [58]. Danhassan et al. (2018) [59] demonstrated that the raw seed had an in vitro protein digestibility of 76.9%, which is comparable to that of many legumes and cereal proteins.
Alternatively, fox nut is less cultivated but is a potential resource exhibiting protein around 10%. Fox nut is produced mostly in India, in stagnant water sources, and is consumed for medicinal and nutritional benefits [60]. The seed is highly nutritious, contains fiber, and is rich in vitamins, minerals, and EAAs [49].

3.2. Marine

3.2.1. Microalgae

Microalgae are microscopic plants with species that grow in both marine and freshwater environments. These include Chlorella vulgaris, Nannochloropsis spp., Tetraselmis spp., and often the cyanobacterium Arthrospira plantensis due to its similarity. These microalgae are high in protein and healthy oils, and rich in vitamins and minerals, as shown in Table 3. Additionally, their rapid growth and minimal nutrient requirements offer strong potential for alternative protein production. In open pond systems, approximately 30 tons of dry matter are produced per hectare per year, but they also have limitations such as low biomass production, high harvesting cost, evaporative water loss, high risk of contamination, and reduced carbon dioxide utilization efficiency [61,62]. The disadvantages of open pond systems are minimized by closed systems. Photobioreactors enable low contamination and single species cultivation, use less land, and combine natural and artificial light to maximize the productivity of microalgae [63]. Hybrid cultivation systems can generate more biomass and less area compared to open pond systems [64].
Cyanobacteria operate in similar systems while having fewer nutrient requirements and fixing atmospheric nitrogen [65]. These species also provide the full spectrum of essential amino acids (EAAs) [66]. Microalgae can yield up to ten times more protein per hectare than soy [61]. Microalgae also provide bioactive compounds such as β-carotene, lutein, and astaxanthin, as well as lipid fractions rich in essential fatty acids, particularly long-chain polyunsaturated fatty acids (PUFAs) belonging to the omega-3 and omega-6 series, which are used in feed industries. These exhibit diverse health-promoting effects such as antioxidants, anti-inflammatory, anticancer, and cardio-protective activities. However, their incorporation into functional foods is complicated by species variability, cultivation and processing challenges, and unresolved concerns regarding the bioavailability and safety of these compounds, in addition to sensory limitations [67].
Another major challenge in utilizing microalgae as food is that certain strains possess robust cell walls, which hinder digestive enzymes from accessing intracellular components [68]. Microalgal digestibility varies by species, with the cyanobacterium Spirulina showing variability in digestibility from as high as 94% to 74% on the lower end, which is less than many traditional plant proteins, while strains like Chlorella require cell disruption [69,70].

3.2.2. Macroalgae

Macroalgae and seagrass are major coastal primary producers that structure nearshore ecosystems and sustain coastal economies. Seaweed offers nutrient-rich biomass, containing essential minerals, fiber, and healthy fatty acids, in addition to various bioactive compounds. Due to these benefits, it is increasingly cultivated as a sustainable resource for both foods and advanced bioproducts, such as packaging, nutraceuticals, and industrial biorefining materials [71].
The annual global aquaculture production of macroalgae now exceeds tens of millions of metric tons, emphasizing their contributions to food security and coastal livelihoods [72]. Although seldom consumed directly, seagrass meadows provide critical ecosystem services, including support of fisheries, sediment stabilization, and substantial blue-carbon sequestration, thereby advancing climate mitigation and biodiversity conservation objectives [73].
Different methods of cultivation are used for macroalgae and seagrass. This includes open water systems, tanks, and integrated aquaculture. Seaweeds are grown using a variety of systems: longlines or rafts in coastal waters (e.g., Saccharina latissima), fixed or floating monolines in the tropics (e.g., Kappaphycus alvarezii), ponds or tanks (e.g., Gracilaria), and land-based RASs (Recirculating Aquaculture Systems) where Ulva acts as a biofilter and biomass crop [74,75,76]. Integrated Multi-Trophic Aquaculture (IMTA) pairs seaweeds with bivalves or finfish to capture dissolved nutrients and improve overall system efficiency [77].
Table 3. Protein and nutritional profile of macro- and microalgae.
Table 3. Protein and nutritional profile of macro- and microalgae.
TypeMacronutrient ProfileEAA Profile HighlightsQuality and DigestibilityReference
Chlorella vulgaris (microalgae)High protein: up to 67%Balanced EAAs, high in Lys
Limiting: Met		Cell-wall disruption/extraction improves bioaccessibility [78,79,80]
Arthrospira platensis (cyanobacterium)Very high protein (50–70% DW) All EAAs, strong in Leu and Val
Limiting: sulfur AA		Generally high digestibility; matrix-dependent [5,81]
Nannochloropsis spp. (microalga)Moderate-to-high proteinRobust Leu and Lys
Limiting: Met		Composition is reasonably stable across media in one study [82,83]
Tetraselmis spp. (microalga)Protein-richBalanced EAAs; good Lys
Limiting: Thr and Met		Digestibility increases with milling/enzymatic pretreatment [82]
Pyropia/Porphyra (nori; red)Moderate-to-high protein; low fatAll EAAs; rich in Leu, Lys, and Val
Limiting: Trp and Met		Vitamin B12 is present and bioavailable in animal and human studies [84,85,86,87]
Ulva spp. (sea lettuce; green)15–30% protein (DW); high fiber; low fatHigh percentage of EAAs; rich in His;
Limiting: Trp and Met		Minerals (Ca, Fe, Mg, and I) and Vitamin B [84,87,88,89,90]
Gracilaria spp. (red)Carbohydrate-rich (agar); moderate protein; low fat All EAAs; rich in Leu, Val, and Thr
Limiting: Trp and Met		Fe and Ca can be appreciable; variable by site/season [85,87,91]
Saccharina/Laminaria (kelp; brown)Low-to-moderate protein and fiber, and low fat Reliable EAA source
Limiting: Met, Cys, and Lys		Iodine is very high; managed via portioning/processing [84,85,92,93]
Zostera marina seeds (eelgrass; seagrass) 9–13% proteinBalanced amino-acid pattern Food use is still niche; composition varies by site/season [94,95]
Macroalgae supply all three macronutrients (carbohydrate, protein, and fat), though amounts vary widely by taxon and environment. As presented in Table 3, protein can be substantial in green and red genera (e.g., Ulva and Pyropia/Porphyra), while browns tend to be lower [84,89]. Predominant carbohydrates are the phycocolloids agar (reds), carrageenan (reds), and alginate (browns) that function as dietary fiber and industrial hydrocolloids [85]. Total lipids are typically low, yet profiles often include nutritionally desirable n-3 PUFA [96]. Micronutrient density is notable: kelps can be exceptionally iodine-rich, useful where iodized salt intake is low, but also a potential excess-intake risk without portion control or processing to reduce iodine. Certain red seaweeds (nori) contain physiologically active Vitamin B12; both mechanistic and intervention evidence indicate bioavailability in humans [86]. Seaweeds also contain antioxidants/bioactives such as phlorotannins and fucoxanthin in browns, and ulvan in Ulva, which contribute to reported functional effects, though magnitudes are species- and processing-dependent [90,93]. Overall composition reflects species identity, season, habitat, and post-harvest handling; thus, data should be interpreted with explicit context [84].
Across macroalgae, EAA distributions and total protein vary by phylum and species. Red seaweeds (Porphyra/Pyropia) and green seaweeds (Ulva) often show higher protein and favorable EAA proportions, with lysine and threonine that can complement cereal-based diets [84,87]. Recent analytical work on Pyropia yezoensis confirmed adequate total EAA content, with methionine as the first limiting amino acid, a pattern comparable to many terrestrial plant proteins [97]. By contrast, evidence for seagrasses remains limited for direct food use; developmental and environmental modulation of free amino-acid pools has been documented in Zostera japonica, underscoring that EAA composition can be dynamic and context-dependent [98]. Relative to terrestrial sources, several seaweeds meet or exceed adult EAA reference patterns when appropriately formulated, but digestibility and cell-wall polysaccharides can depress DIAASs; processing steps are critical to improving bioaccessibility. In human foods, macroalgal proteins and peptides support fortification and the development of alternative-protein matrices [84]. In animal nutrition, systematic and narrative reviews report performance or health benefits at modest inclusion rates, though responses are species- and dose-dependent [99,100].

4. Green Processing

Aquatic plants offer incredible potential as a renewable resource for protein and other nutritionally significant compounds. However, as with other plants, many aquatic plants exhibit high amounts of antinutritional compounds that reduce the bioavailability or digestibility of these components [28,101]. Additionally, as with other plant proteins, the cell wall makes protein extraction more difficult. Thus, pre-processing is critical for extracting and enhancing protein quality, without contributing to unsustainable practices. Green processing strategies refer to environmentally safe, resource-efficient, and energy-efficient technologies that enable the release of proteins from biomass [102]. Therefore, processes that use high heat, fossil fuels, or materials obtained from externally unsustainable processes must be avoided to maintain the sustainability of the whole system for aquatic plants. Processes such as ultrasonication, fermentation, microwave, and high-pressure processing have all exhibited potential as green processing solutions to improve aquatic plant quality [103].

4.1. Fermentation

Fermentation exhibits clear potential as a sustainable technology with low GHG emissions while improving the quality and nutritional content of protein [104]. Most aquatic plants exhibit less lignin or cellulose than terrestrial plants, making them more readily digestible by many microbes without requiring additional pretreatment. Various microbes enhance overall protein quality and digestibility in plants through the enzymatic degradation of ANFs and protein hydrolysis, releasing amino acids and potentially consuming and synthesizing new components [105]. Many studies have exhibited changes in protein patterns, free amino acid content, and changes in amino acid composition utilizing lactic acid bacteria on aquatic plants, including red seaweed, spirulina, duckweed, and mixed silage [106,107,108,109]. There are still limited overall studies on the variety of aquatic resources and microbial species fermentations, indicating significant potential for improving the quality of sustainable aquatic plant products.

4.2. Ultrasound-Assisted Extraction

Ultrasound-assisted extraction (UAE), which uses acoustic cavitation to disrupt cells, is an emerging technology and processing application commonly applied to plant-based proteins. While less studied, the application to aquatic plant resources is beginning to gain traction. The study by Dohaei et al. (2020) [110] exhibited a maximum of 24.4% protein and approximately 12% soluble protein extracted from Pacific mosquitofern (Azolla filiculoides) utilizing ultrasound-assisted alkaline extraction (UAAE) after optimization. Similarly, O’ Connor et al. (2020) [111] showed a yield of approximately 35% in two different seaweed species using UAAE. The efficacy of this process on aquatic resources was further validated by Nitiwuttithorn et al. (2024) [105], who showcased a 35% protein recovery using UAAE on Wolffia. Their study showed a 2.5 times higher yield in fresh protein versus dried, potentially reducing the additional energy input from drying. Additionally, UAE or UAAE exhibit modified AA profiles with higher EAA in aquatic plants as compared to traditional extractions [111,112,113]. UAE exhibits great potential for rapidly extracting and improving proteins, while current advances in industrial probe systems indicate improved viability at scale [114].
Microwave-assisted extraction (MAE) is a low-thermal processing method that has exhibited excellent potential for rapidly extracting plant protein. MAE has often been used to extract alternative metabolites from aquatic plants, ranging from pigments, lipids, phenolic compounds, and more [115,116,117]. However, Chew et al. (2019) [118] observed a 63.2% recovery in microalgae C. vulgaris using optimized microwave-assisted three-phase partitioning for two minutes. This indicates the potential that still exists, requiring greater investment in research to optimize conditions among various species. However, MAE for aquatic plants exhibits some constraints, as it is less effective in dried samples, which is often necessary for preserving samples.

4.3. High-Pressure Processing

High-pressure processing (HPP) or high hydrostatic pressure treatment (HHPT) is typically utilized for preserving food but has recently been explored for extracting proteins from plants by disrupting cell walls and membranes to release proteins without thermal processing. Despite the limited current work on aquatic plants, O’ Connor et al. (2020) [111] successfully recovered protein yields in four seaweed species, with results ranging from a low of 3.2% to 23.7%. This work showcases the variability between species associated with HPP extraction but also highlights the potential for more targeted studies across a wide range of aquatic species.

4.4. Pulsed Electric Field

Pulsed electric field (PEF) utilizes a high voltage pulse between two electrodes to destabilize the cell wall and increase overall permeability [119]. The work on aquatic plants is growing, but recent research in macro- and microalgae has exhibited potential using PEF. In spirulina, research showcased yields of approximately 48 g/100 g while also yielding 85 mg/g in phycocyanin [120]. Additionally, in the green macroalgae Ulva spp., PEF exhibited higher protein yields than osmotic shock and demonstrated high antioxidant ability [121]. Despite the limited research in other aquatic resources, including duckweed or seagrasses, the composition of these cell walls typically exhibits less lignin, indicating potential for PEF. While Pfeifer et al. (2022) [122] found no lignin in the seagrass Zostera, Zhao et al. (2014) [123] identified low lignin content in L. minor. Given these findings, PEF presents significant potential for processing these aquatic resources, though no current studies on this application are available.
Although there are significant research gaps for aquatic plants among green processing methods, many strategies have shown effective protein extraction or enhancement. The majority of the available studies are on macro- and microalgae. However, as these and other aquatic plant resources are given more consideration, there is incredible potential for processing growth and advancements. Furthermore, there is exceptional potential for combining processing strategies to enhance the protein even more, as has previously been demonstrated for land crops [103]. As climate-resilient aquatic crops gain recognition as vital resources, it is essential that investment in processing technologies and innovation advances in parallel.

5. Sustainability and Blue Economy

Meeting the global demands for protein is putting a lot of strain on traditional food systems and often leads to a decrease in sustainable food practices. Aquatic plants demonstrate strong potential to meet the growing protein demand while mitigating or benefiting environmental and nutritional sustainability. Furthermore, the sustainable production and potential of aquatic plant resources promote the development of blue economies to support local communities and sustainable circular systems.

5.1. Sustainability

With their low requirements for land and water, aquatic plants like duckweed, Azolla, water spinach, and microalgae offer a very sustainable substitute for traditional food production [12,124,125]. They greatly lower energy use and greenhouse gas emissions by producing biomass that is high in protein for human consumption and animal feed [126,127]. For instance, Azolla is excellent at sequestering carbon, and microalgae significantly reduce greenhouse gas emissions, efficiently valorize wastewater, and generate biomass for feed and biofuels. When compared to conventional animal-based protein sources, this cultivation model offers a significantly reduced environmental footprint in terms of land use, water consumption, and carbon emissions [128].

5.2. Blue Economy

Breeding for the best strains helps boost sustainability, profitability, and enhance production efficiency. Breeding priorities are mainly focused on areas like accelerated growth, increased protein content, enhanced resilience to climate change (like heat tolerance), and improved digestibility. High-yielding seaweed, duckweed, and microalgae are already produced using technologies such as marker-assisted selection and gene editing [129,130].
The government, as well as academia and the private sector, can play important roles by giving recognition as a top developmental priority, public sector funding, and equity investment for the cultivation, processing, and marketing of aquatic plants which have minimal impact on the environment. Such activities can create supply, whereas boosting consumer preference can create demand among them. Apart from subsidies, the government also needs to assist the capacity building on cultivation, marketing, disease management, and processing. For international trade, FAO and other international organizations are also creating standards to maintain a uniform quality. This would assist in combating transboundary challenges (like climate change) and ensure the benefits of the blue economy [131].

6. Conclusions

The utilization of freshwater aquatic plants, like duckweed, freshwater floating ferns, and vegetables, and marine aquatic organisms, such as macroalgae and microalgae, is foundational to promoting sustainable food systems. These sources offer substantial nutritional benefits, including high protein and mineral content, and properties such as anti-inflammatory, antidiabetic, and anti-fatigue effects. They also foster sustainable farming practices due to low land and water usage, coupled with superior GHG emissions and carbon sequestration relative to other similar nutrient sources. Advanced processing techniques are crucial for maximizing the nutritional value of aquatic resources, as processing methods like GFP can be used to minimize antinutritional properties and increase the bioavailability of minerals in these plants. One technological approach being studied involves high-pressure microfluidization, which is a method shown to improve the nutritional quality of plant-based materials.
Further research and development focusing on microbial optimization combined with green technologies will enhance the digestibility and sensory attributes of aquatic-based foods, building upon existing methods that enhance macro- and micronutrient properties. Ensuring the safety and quality of convenience foods derived from aquatic sources requires the adoption of novel, nonthermal processing methods that preserve the inherent nutritional profile. High-pressure processing has been identified as a novel alternative nonthermal technology to heat pasteurization. Applying such nonthermal technologies is vital for developing RTE aquatic products that are safe, retain the plants’ beneficial properties, and meet consumer demand for minimally processed, nutrient-dense foods. The integration of advanced green and microbial technologies is essential for leveraging the sustainable advantages of aquatic organisms and maximizing their utility within a circular bioeconomy framework. The cultivation techniques, processing techniques, and nutritional benefits of these plants need active promotion by the government and other sectors. By harmonizing the standards of such value-added products, regulatory support can facilitate transboundary trade, ultimately promoting sustainable nutrition, food security, and the blue economy. In essence, if the foundational cultivation of aquatic resources is the fertile ground providing sustainable crops, then green and microbial processing technologies act as the precision tools that unlock the maximum value and digestibility of those crops, ensuring they transition smoothly into high-quality, safe, global food products.

Author Contributions

Conceptualization: A.K.A. and A.K.; Methodology: A.K.A., A.K., D.L.R., N.K.S., K.N.C.Z., J.A.V. and R.N.; Investigation: A.K.A., A.K., D.L.R., N.K.S., K.N.C.Z., J.A.V. and R.N.; Validation: A.K.A., A.K., D.L.R. and N.K.S.; Original Draft Preparation: A.K., D.L.R., N.K.S., K.N.C.Z., J.A.V. and R.N.; Supervision: A.K.A., A.K., D.L.R. and N.K.S.; Review and Editing: A.K.A., A.K., D.L.R. and N.K.S.; Revision: A.K. and N.K.S. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Data Availability Statement

No new data was created.

Conflicts of Interest

The authors declare no conflicts of interest.

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Table 1. Nutritional profile of major duckweed genera.
Table 1. Nutritional profile of major duckweed genera.
GenusProtein (%)Lipid (%)Carbohydrate (%)Fiber (%)Key MicronutrientsReferences
Lemna
(L. minor, L. gibba,
L. trisulca)		25–354–730–4010–15β-carotene, lutein, Vitamin C, Fe, Zn[20]
Wolffia
(W. globosa, W. arrhiza)		35–453–625–358–12Fe, Zn, Vitamin B12, C, E, β-carotene[12]
Spirodela
(S. polyrhiza)		20–303–635–4512–18Iron, Mg, calcium[10]
Landoltia
(L. punctata)		20–283–540–5010–14Polyphenols, starch, trace minerals[21]
Wolffiella
(W. hyalina, W. lingulata)		20–303–530–4015–20 [22]
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Anal, A.K.; Khadka, A.; Rice, D.L.; Shrestha, N.K.; Valerozo, J.A.; Zaw, K.N.C.; Nagase, R. Aquatic Plants for Blue Protein Innovation: Bridging Nutrition, Sustainability, and Food Security. Resources 2025, 14, 192. https://doi.org/10.3390/resources14120192

AMA Style

Anal AK, Khadka A, Rice DL, Shrestha NK, Valerozo JA, Zaw KNC, Nagase R. Aquatic Plants for Blue Protein Innovation: Bridging Nutrition, Sustainability, and Food Security. Resources. 2025; 14(12):192. https://doi.org/10.3390/resources14120192

Chicago/Turabian Style

Anal, Anil Kumar, Abhishek Khadka, Daniel Lee Rice, Nabindra Kumar Shrestha, Johnmel Abrogena Valerozo, Khin Nyein Chan Zaw, and Ryunosuke Nagase. 2025. "Aquatic Plants for Blue Protein Innovation: Bridging Nutrition, Sustainability, and Food Security" Resources 14, no. 12: 192. https://doi.org/10.3390/resources14120192

APA Style

Anal, A. K., Khadka, A., Rice, D. L., Shrestha, N. K., Valerozo, J. A., Zaw, K. N. C., & Nagase, R. (2025). Aquatic Plants for Blue Protein Innovation: Bridging Nutrition, Sustainability, and Food Security. Resources, 14(12), 192. https://doi.org/10.3390/resources14120192

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