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<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Mar. Sci.</journal-id>
<journal-title>Frontiers in Marine Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Mar. Sci.</abbrev-journal-title>
<issn pub-type="epub">2296-7745</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmars.2025.1661042</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Marine Science</subject>
<subj-group>
<subject>Review</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Harnessing microalgae for sustainable nutrition and ecosystem services in aquaponic systems: a blue&#x2013;green approach to ecosystem health</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Manoharan</surname>
<given-names>Ramya</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2542573/overview"/>
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<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/visualization/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Somanathan Nair</surname>
<given-names>Chythra</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2542787/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Nishanth</surname>
<given-names>Drishya</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/3145698/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Subramanian</surname>
<given-names>Radhakrishnan</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/3163432/bio"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ahmed</surname>
<given-names>Zienab</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1018458/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Rastrelli</surname>
<given-names>Luca</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/112235/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Xie</surname>
<given-names>Xiu-Lan</given-names>
</name>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1781439/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Ren</surname>
<given-names>Mao-Zhi</given-names>
</name>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
<xref ref-type="aff" rid="aff7">
<sup>7</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/229135/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Jaleel</surname>
<given-names>Abdul</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/436653/overview"/>
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<aff id="aff1">
<sup>1</sup>
<institution>Department of Integrative Agriculture, College of Agriculture and Veterinary Medicine, United Arab Emirates University</institution>, <addr-line>Al Ain</addr-line>,&#xa0;<country>United Arab Emirates</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>ASPIRE Research Institute for Food Security in the Drylands (ARIFSID), United Arab Emirates University</institution>, <addr-line>Al Ain</addr-line>,&#xa0;<country>United Arab Emirates</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>National Biodiversity Future Center (NBFC)</institution>, <addr-line>Palermo</addr-line>,&#xa0;<country>Italy</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Department of Pharmacy, University of Salerno</institution>, <addr-line>Salerno</addr-line>,&#xa0;<country>Italy</country>
</aff>
<aff id="aff5">
<sup>5</sup>
<institution>Laboratory of Space Biology, Institute of Urban Agriculture, Chinese Academy of Agricultural Sciences</institution>, <addr-line>Chengdu</addr-line>,&#xa0;<country>China</country>
</aff>
<aff id="aff6">
<sup>6</sup>
<institution>Zhengzhou Research Base, State Key Laboratory of Cotton Biology, School of Agricultural Sciences, Zhengzhou University</institution>, <addr-line>Zhengzhou</addr-line>,&#xa0;<country>China</country>
</aff>
<aff id="aff7">
<sup>7</sup>
<institution>Hainan Yazhou Bay Seed Laboratory</institution>, <addr-line>Sanya</addr-line>,&#xa0;<country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/720896/overview">Devendra Singh</ext-link>, Central Arid Zone Research Institute (ICAR), India</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/204426/overview">Angel Llamas</ext-link>, University of Cordoba, Spain</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/1961330/overview">Atif Khurshid Wani</ext-link>, Lovely Professional University, India</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Abdul Jaleel, <email xlink:href="mailto:abdul.jaleel@uaeu.ac.ae">abdul.jaleel@uaeu.ac.ae</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>01</day>
<month>09</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>12</volume>
<elocation-id>1661042</elocation-id>
<history>
<date date-type="received">
<day>07</day>
<month>07</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>07</day>
<month>08</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Manoharan, Somanathan Nair, Nishanth, Subramanian, Ahmed, Rastrelli, Xie, Ren and Jaleel.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Manoharan, Somanathan Nair, Nishanth, Subramanian, Ahmed, Rastrelli, Xie, Ren and Jaleel</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Aquaponics is a soilless farming approach that integrates aquaculture with hydroponics to produce food. In regions with limited arable land, aquaponics can help address food insecurity challenges. Both fish and plants are produced using aquaponic systems. The aquafeeds used to feed the fish in aquaponic systems are also the main source of nutrition for the plants. Currently, commercial aquafeeds such as fishmeal and fish oil are used in aquaponics, but they do not completely meet the nutritional requirements of plants. Additionally, commercial aquafeeds are expensive, and their production is unsustainable. This review focuses on the suitability of microalgae as a replacement for commercial aquafeeds and its role in meeting the nutritional requirements of plants growing in aquaponic systems. Microalgae production is sustainable and cost effective compared to commercial aquafeed production. Many studies have been conducted on the impact of microalgae-based feed on fish growth and its role as a biofertilizer and biostimulants for plant growth. However, using microalgae as aquafeed for the development of both fish and plants in aquaponic systems remains underexplored. This review aims to provide insights into the dual role of microalgae in aquaponics&#x2014;enhancing fish nutrition while supplementing plant nutrient requirements. Although some micronutrient gaps may persist, further optimisation could help make aquaponic systems more efficient and sustainable.</p>
</abstract>
<kwd-group>
<kwd>microalgae</kwd>
<kwd>aquafeed</kwd>
<kwd>nutrient cycle</kwd>
<kwd>plant nutrition</kwd>
<kwd>sustainability</kwd>
<kwd>aquaponics</kwd>
</kwd-group>
<contract-sponsor id="cn001">United Arab Emirates University<named-content content-type="fundref-id">10.13039/501100006013</named-content>
</contract-sponsor>
<counts>
<fig-count count="3"/>
<table-count count="2"/>
<equation-count count="0"/>
<ref-count count="186"/>
<page-count count="18"/>
<word-count count="8347"/>
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<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Microbial Symbioses</meta-value>
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</front>
<body>
<sec id="s1" sec-type="intro">
<title>Introduction</title>
<p>Aquaponics is an innovative and sustainable food production system that integrates recirculating aquaculture with hydroponics (<xref ref-type="bibr" rid="B82">K&#xf6;nig et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B116">Okomoda et&#xa0;al., 2023</xref>). Aquaponic plants perform comparably to hydroponic plants, despite lower nutrient concentrations (<xref ref-type="bibr" rid="B157">Sreekumar et&#xa0;al., 2023</xref>). This system offers numerous benefits, including water conservation, a reduced environmental impact, and the simultaneous production of fish and plants (<xref ref-type="bibr" rid="B104">Mishra et&#xa0;al., 2020</xref>). Aquaponics can be implemented in various settings, ranging from urban to rural areas, and at different scales, ranging from small-scale farms to industrial production units (<xref ref-type="bibr" rid="B82">K&#xf6;nig et&#xa0;al., 2016</xref>). This system is particularly valuable for addressing food security challenges as it can be utilised on non-arable land and in areas with limited water resources (<xref ref-type="bibr" rid="B153">Shreejana et&#xa0;al., 2022</xref>). Furthermore, as aquaponics is a closed-loop system that includes both hydroponic and aquaculture systems, it presents potential solutions for food production in the face of climate change-related challenges such as global warming, desertification, water scarcity, famine, and increased pests and diseases. Aquaponics is an eco-friendly cultivation system that has gained interest in various fields and industries including ecology, agriculture, and fisheries (<xref ref-type="bibr" rid="B69">Hao et&#xa0;al., 2020</xref>). Several areas of aquaponic systems need to be explored further to fully understand their potential. These areas include nitrogen cycling, nutrient recovery from fish waste, plant nutrition, plant pathogens, pest control strategies, and sustainable aquafeed (<xref ref-type="bibr" rid="B61">Goddek et&#xa0;al., 2019</xref>). This review focuses on microalgae as a sustainable alternative to commercial aquafeeds in aquaponic systems. Additionally, this review examines how microalgae can fulfil the nutritional requirements of both plants and fish in aquaponic systems., In aquaponic systems, residual fish feed and fish waste provide nutrients for plant growth. Current aquaponic systems that use commercial aquafeeds such as fish oil, fishmeal, and fish waste do not completely meet the nutritional requirements for plant growth (<xref ref-type="bibr" rid="B47">Eck et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B180">Yep and Zheng, 2019</xref>). Macronutrients such as potassium, phosphorus, sulphur, and calcium, along with micronutrients such as manganese, iron, zinc, copper, boron and molybdenum, often accumulate in inadequate amounts or disproportionate ratios in the water. Consequently, plants do not receive sufficient nutrients for their growth (<xref ref-type="bibr" rid="B39">Delaide et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B160">Suhl et&#xa0;al., 2016</xref>). Furthermore, commercial aquafeeds are expensive and unsustainable. I The water footprint of commercial aquafeeds was estimated to be between31&#x2013;35 km<sup>3</sup> in 2008, with the top five species alone accounting for 18.2 km<sup>3</sup>. For example, fisheries that provide commercial aquafeeds, such as fish oil and fishmeal, emitted 4.6 million tons of carbon dioxide-equivalent greenhouse gases in 2014 (<xref ref-type="bibr" rid="B25">Cashion et&#xa0;al., 2017</xref>). The carbon footprints of extruded and pelleted grass carp aquafeeds were 1334 and 1071 kg CO<sub>2</sub> eq/t, respectively, with raw material production being the largest contributor. To overcome these challenges, sustainable sourcing of raw materials and the utilisation of renewable energy in aquafeed production can substantially reduce the environmental impact (<xref ref-type="bibr" rid="B170">Wang et&#xa0;al., 2022</xref>). Additionally, exploring alternative aquafeed sources, such as single-cell proteins and insects, can help address sustainability concerns (<xref ref-type="bibr" rid="B34">D&#x2019;Abramo, 2021</xref>). Microalgae have emerged as promising alternative aquafeed ingredients because of their nutritional profile and sustainability benefits. They contain essential amino acids, fatty acids, vitamins, pigments, and bioactive compounds that enhance fish health, survival, and product quality (<xref ref-type="bibr" rid="B107">Nagappan et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B139">Sagaram et&#xa0;al., 2021</xref>). Compared to traditional feed sources, such as fishmeal and soymeal, microalgae offer a more diverse set of fatty acids, pigments, sterols, and vitamins (<xref ref-type="bibr" rid="B45">Dixit et&#xa0;al., 2022</xref>). Microalgae production has a lower environmental footprint than terrestrial crops in terms of water use and land requirements (<xref ref-type="bibr" rid="B107">Nagappan et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B96">Mahata et&#xa0;al., 2022</xref>). Additionally, microalgae can positively influence gut microbiota and immune responses in aquatic species (<xref ref-type="bibr" rid="B139">Sagaram et&#xa0;al., 2021</xref>). Microalgae, small but powerful photosynthetic organisms, are emerging as a sustainable solution to a range of global challenges&#x2014;from clean energy and environmental protection to food security and human health. Rich in nutrients and bioactive compounds, microalgae are being explored for use in biofuels (<xref ref-type="bibr" rid="B7">Akhtar et&#xa0;al., 2023</xref>), functional foods (<xref ref-type="bibr" rid="B12">Andrade-Bustamante et&#xa0;al., 2025</xref>), and health supplements (<xref ref-type="bibr" rid="B14">Ayub et&#xa0;al., 2025</xref>), offer benefits like antioxidant, anti-inflammatory, and heart-protective effects. Their ability to treat wastewater, capture CO<sub>2</sub>, and support circular practices makes them valuable in aquaculture (<xref ref-type="bibr" rid="B38">Dasari et&#xa0;al., 2025</xref>) and environmental cleanup, including antibiotic pollution removal (<xref ref-type="bibr" rid="B171">Wani et&#xa0;al., 2024</xref>). They use minimal land and water, making them eco-friendly alternatives to traditional crops, and contribute significantly to achieving climate goals and UN Sustainable Development Goals (<xref ref-type="bibr" rid="B5">Ahmad and Ashraf, 2024</xref>). Despite challenges like high production costs, regulatory hurdles, and taste issues in food applications, advances in biotechnology, AI, and strain development are paving the way for large-scale, sustainable use of microalgae across industries. This review examines the potential of microalgae as sustainable components of aquafeed formulations. It also provides valuable insights into the utilisation of microalgae aquafeeds as replacements for commercial aquafeeds such as fish oil and fishmeal. Microalgae may provide complete nutrition for plant growth in aquaponic systems, and its use will enhance the efficiency and sustainability of these systems and contribute to global food security.</p>
</sec>
<sec id="s2">
<title>Aquaponics: an overview</title>
<p>Aquaponics is a climate-smart technology used for sustainable food production (<xref ref-type="bibr" rid="B112">Nishanth et&#xa0;al., 2024</xref>). Aquaponic systems use less than 90% of the water used in conventional fish and plant farming which support sustainable food production and facilitate complete biological processes between fish, plants, and microbes. Aquaponic systems are composed of three main components that work together: the growing bed (hydroponic unit) for plant growth, biofilter for microbes to perform nitrification, and aquaculture tank to rear fish. All three components must function in coordination to support fish and plant growth. Fish waste is the primary nutrient component for plant growth in aquaponics. Fish waste acts as a primary nutrient source undergoing microbial nitrification to convert ammonia into plant-available nitrates. Based on the designs of the hydroponic and aquaculture units, aquaponic systems are classified into coupled and decoupled systems. Common hydroponic designs include Nutrient Film Technique (NFT), floating raft or deep-water culture, and media-based grow beds. peat moss and perlite are used as plant growth media. Media-based systems use substrates like peat moss and perlite and are ideal for vegetables and fruits due to their capacity to support high root density. NFT systems are typically used for smaller vegetables, while floating raft systems are most common, allowing roots to freely absorb nutrients. To maintain a stable environment, it is essential to monitor water quality, pH, temperature, water-use efficiency, waste management, and nutrient cycling (<xref ref-type="bibr" rid="B61">Goddek et&#xa0;al., 2019</xref>).</p>
<p>Nutrient cycling in an aquaponic system has many advantages because it is a recirculating system combining hydroponics and aquaculture. Because no effluent is discharged, it prevents environmental pollution. Additionally, the nutrient-rich aquaculture water can be reused as an organic fertiliser for plants in the hydroponic units. Some studies have indicated that aquaponics produces plant growth and yields comparable to or even exceeding those of soil-grown plants (<xref ref-type="bibr" rid="B181">Yogev et&#xa0;al., 2016</xref>). Nutrient cycling in aquaponic systems is influenced by multiple factors such as the aquafeed type, fish species, fish density, plant type, and microbial community. The main nutrient sources in aquaponic systems are aquafeed and aquaculture water, which contribute essential elements such as magnesium, calcium, and sulphur (<xref ref-type="bibr" rid="B39">Delaide et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B147">Schmautz et&#xa0;al., 2016</xref>). The two main types of aquafeed are plant-based and fishmeal-based feeds. After being introduced into the system, aquafeed is partly consumed and excreted by the fish, while some residual feed remains in the tank. Fish excreta and uneaten feed dissolve in the water, releasing nutrients that are absorbed by plants. To support optimal plant growth, additional supplements like potassium and iron may be introduced&#x2014;without harming the fish (<xref ref-type="bibr" rid="B147">Schmautz et&#xa0;al., 2016</xref>). Residual aquafeed, which accounts for less than 5%, and fish excreta also contribute to carbon dioxide and ammonia production, increasing the nutrient load of the water and influencing plant development (<xref ref-type="bibr" rid="B181">Yogev et&#xa0;al., 2016</xref>). Water quality and fish biomass are strongly influenced by aquafeed type, highlighting the importance of selecting feed that meets the nutritional needs of both fish and plants (<xref ref-type="bibr" rid="B147">Schmautz et&#xa0;al., 2016</xref>).</p>
<p>Microbial communities in aquaponic systems undergo many biological processes that convert fish waste and residual aquafeed into nutrient rich solutions for plant growth. One such process is solubilisation carried out by bacteria that break down complex organic compounds into ionic forms absorbable by plants. Heterotrophic bacteria such as <italic>Pseudomonas</italic> sp., <italic>Flavobacterium</italic> sp., <italic>Rhizobium</italic> sp., <italic>Aeromonas</italic> sp., and <italic>Sphingobacterium</italic> sp. are involved in this solubilisation process. Additionally, some &#x3b3;-proteobacteria can solubilise phytates making phosphorus available to plants. The primary nitrogen source in aquaponic systems is the proteins present in aquafeeds. However, fish utilise only about30% of the nitrogen present in aquafeed, and the remaining is excreted in the form of ammonia (<xref ref-type="bibr" rid="B137">Ru et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B174">Wongkiew et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B178">Yavuzcan Yildiz et&#xa0;al., 2017</xref>). This ammonia is oxidised to nitrite by ammonia-oxidising bacteria such as <italic>Nitrosococcus</italic>, <italic>Nitrosospira</italic>, <italic>Nitrosomonas</italic>, <italic>Nitrosolobus</italic>, <italic>and Nitrosovibrio</italic>, and subsequently converted to nitrate by nitrite-oxidising bacteria such as <italic>Nitrobacter</italic>, <italic>Nitrococcus</italic>, <italic>Nitrospina</italic>, <italic>and Nitrospira</italic> (<xref ref-type="bibr" rid="B174">Wongkiew et&#xa0;al., 2017</xref>). Understanding the nutrient cycles is essential for the effective operation of aquaponic systems, as plants require different nutrients at various growth stages. Some of these nutrients can be supplemented either as foliar application or by adding nutrients directly to the water. Macronutrients, such as carbon, are supplied through the organic compounds in the aquafeed then metabolised by both fish and microbes, releasing carbon dioxide(CO<sub>2</sub>) as a byproduct. This CO<sub>2</sub> is then absorbed by plants and used in photosynthesis via carbon fixation. Plants uptake nitrogen as nitrate or ammonium ions (<xref ref-type="bibr" rid="B174">Wongkiew et&#xa0;al., 2017</xref>), Phosphorus as orthophosphate (<xref ref-type="bibr" rid="B133">Resh, 2022</xref>), and potassium which is important for growth and accumulates especially in fruit (<xref ref-type="bibr" rid="B147">Schmautz et&#xa0;al., 2016</xref>). Other essential elements such as calcium, magnesium, and sulphur typically present in tap water, while micronutrients like manganese, iron, and zinc, are derived from aquafeed., Copper and boron are also present in tap water (<xref ref-type="bibr" rid="B39">Delaide et&#xa0;al., 2017</xref>). Overall, aquaponic systems foster a symbiotic relationship among fish, plants, and microbes in a recirculating sustainable food production process.</p>
</sec>
<sec id="s3">
<title>Nutrient imbalance and aquafeed unsustainability in aquaponics</title>
<p>In aquaponic systems, nutrients are transferred from fish waste to plants through biological processes; however, an imbalance often exists between the nutrient content in fish waste and the nutrient requirements for optimal plant growth. Factors such as fish tank size, biofilter capacity, and system design influence the nutrient availability Therefore, the nutrient composition of aquafeed, and the specific requirements of each plant species must be carefully considered (<xref ref-type="bibr" rid="B133">Resh, 2022</xref>). Monitoring nutrient availability is challenging, as nutrients originate primarily from fish waste and residual aquafeed. Processes such as fish waste removal, water renewal, and denitrification contribute to nutrient loss in the system. Research studies has shown that fish waste and residual aquafeed contain 86% manganese, 22% copper, 89% magnesium, 24% iron, 16% calcium, 6% potassium, 6% nitrogen, and 18% phosphorus. However, not all these nutrients are efficiently utilised by plants, particularly macronutrients like potassium, phosphorus, iron, manganese, and sulphur. Nitrogen released from fish protein metabolism enters nitrogen cycle and is transformed into usable forms usable by plants. Since aquafeed and fish waste are the main nutrient sources their selection and utilisation are crucial for supporting both fish and plant growth in integrated aquaponic systems (<xref ref-type="bibr" rid="B186">Zhanga et&#xa0;al., 2021</xref>).</p>
<p>Some studies have shown that, minerals added as supplements to aquafeed, can be utilised by plants in aquaponic systems. Soluble minerals are not absorbed by fish may be taken up by plants, enhancing nutrient recovery. However, the mineral requirements and metabolism in aquaculture species has not been extensively investigated. The addition of anions and their accompanying cations to aquafeed has been shown to improve nutrient availability for plants (<xref ref-type="bibr" rid="B110">Ng and Koh, 2017</xref>). Plant-based minerals in aquafeed may contain phosphorus in phytate form, which is not readily metabolised by plants. The exogenous addition of enzymes to aquafeed can help release phosphorus from improving bioavailability. However, this approach has some limitations, such as potential release of undesirable compounds that may affect fish health. Further research is needed to evaluate the safe and effective use of these enzymes. Moreover, adding supplements to aquafeed or directly into hydroponic systems is expensive. To reduce this cost it is essential to understand the amount of aquafeed required to meet both fish and plant nutritional requirements. In aquaponic systems, plant physiological processes such as photosynthesis, flowering, defence, and seed germination are regulated in a circadian rhythm pattern which ideally should work in coordination with circadian rhythm patterns of fish. However, when commercial aquafeed is used, this rhythm is not always well-coordinated and plants nutritional requirements may not be fully met. Some microalgae like <italic>Chlamydomonas reinhardtii</italic> has been widely studied for its role in research on photosynthesis, metabolism, and cilia function. Beyond its laboratory significance, it is increasingly recognised for its biotechnological potential due to its fast growth, metabolic flexibility, and low-cost cultivation. It has been applied in biofuel production, nutraceutical development, and wastewater treatment, where it contributes to contaminant removal and resource recovery. Recent studies have also explored the synergistic benefits of co-cultivating <italic>Chlamydomonas</italic> with bacteria to enhance detoxification and bioproduction processes. Although challenges such as genome editing remain, ongoing technological progress continues to expand the industrial and environmental applications of this versatile alga (<xref ref-type="bibr" rid="B141">Salom&#xe9; and Merchant, 2019</xref>; <xref ref-type="bibr" rid="B149">Scranton et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B19">Bellido-Pedraza et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B19">Bellido-Pedraza, Torres and Llamas, 2024</xref>). In addition to its utility in biofuels and bioremediation, microalgae are increasingly explored as sustainable alternatives to commercial aquafeeds.</p>
<p>Moreover, commercial aquafeed production relies heavily on wild fisheries, making it environmentally unsustainable. Some studies have shown that plant-based alternatives like soybean meal and corn gluten meal commonly used to replace fishmeal, contain anti-nutritional factors that limit their effectiveness as aquafeeds (<xref ref-type="bibr" rid="B58">Gerile and Pirhonen, 2017</xref>). Additionally, most plant-based aquafeeds contain phosphorus in phytate form, which is unavailable to plants, necessitating the supplementation of nutrients like phosphorus and zinc in aquaponic systems. Selecting appropriate aquafeed is critical, and supplements should be added carefully to avoid harming both fish and plant health in aquaponics. While plant-based aquafeeds are often promoted as eco-friendly option, they are not fully sustainable due to their negative ecological impacts such as destruction of plant communities for feed production. Animal-based aquafeeds, such as animal proteins sourced from slaughterhouses that are free from anti-nutritional factors, can serve as viable fishmeal substitutes. Additionally, insect-based feeds such as those derived from black soldier flies, have emerged as promising alternatives due to their high protein content, low land and water requirements, reduced greenhouse gas emissions, and superior feed conversion efficiency. However, further research is necessary to evaluate the quality, efficacy, and safety of using insects as aquafeed in aquaponic systems. Recently, the use of microalgae such as <italic>Arthrospira platensis</italic>, <italic>Chlorella vulgaris</italic>, <italic>Schizochytrium</italic> sp., <italic>Nannochloropsis</italic> sp.<italic>, Dunaliella salina., Haematococcus pluvialis.</italic>, and <italic>Isochrysis galbana</italic> as a replacement for commercial aquafeeds has gained increasing attention because microalgae can produce higher biomass than plants. As shown in <xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>, the global average water footprint varies significantly among different aquafeed ingredients, highlighting the need for more sustainable alternatives such as microalgae (<xref ref-type="bibr" rid="B123">Pugazhendhi et&#xa0;al., 2020</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Global average water footprint of different aquafeed ingredients. Adapted from <xref ref-type="bibr" rid="B107">Nagappan et al., 2021</xref>, licensed <uri xlink:href="https://creativecommons.org/licenses/by/4.0/">CC-BY-4.0</uri>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1661042-g001.tif">
<alt-text content-type="machine-generated">Bar chart showing the global average water footprint of various feed ingredients. Water footprint values range from near 0 to approximately 5000 L/kg. Microalgae produced using seawater recycling has the lowest water footprint, while wheat gluten, millet, and insect meal have the highest. X-axis lists feed ingredients; Y-axis shows water footprint in L/kg.</alt-text>
</graphic>
</fig>
<p>Microalgae exhibit remarkable adaptability and rapid growth rates, which make them valuable for various applications. They can thrive in extreme conditions, such as highly alkaline environments, with growth rates of 1.10&#x2013;1.30/d (<xref ref-type="bibr" rid="B122">Praveen et&#xa0;al., 2023</xref>). Thermally tolerant mutant species of <italic>Nitzschia inconspicua</italic> microalgae have shown 1.4- to 6.7-fold higher growth rates than wild types at different temperatures. Adaptive laboratory evolution has been used to enhance the growth rate, stress tolerance, and product yield of microalgae (<xref ref-type="bibr" rid="B88">LaPanse, 2024</xref>). These fast-growing organisms have diverse applications as functional foods and in biofuel production, greenhouse gas mitigation, and wastewater treatment. Their ability to efficiently remove carbon (70&#x2013;80%) and other nutrients (80&#x2013;90%) from wastewater demonstrates their potential in environmental remediation (<xref ref-type="bibr" rid="B122">Praveen et&#xa0;al., 2023</xref>). The high adaptability and rapid growth rate of microalgae make them promising candidates for sustainable biotechnology and industrial innovation across various sectors. Their diverse nutritional compositions makes them valuable in aquaculture, food and other industries. The protein, carbohydrate, and lipid contents of microalgae typically range from 18 to 52%, 18 to 46%, and 12 to 48%, respectively (<xref ref-type="bibr" rid="B184">Zhang et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B166">Tibbetts et&#xa0;al., 2017</xref>). Most species also contain abundant essential amino acids with high digestibility (&gt;80%) (<xref ref-type="bibr" rid="B166">Tibbetts et&#xa0;al., 2017</xref>). Their fatty acid profiles vary, with marine species being rich in monounsaturated fatty acids and freshwater species being rich in polyunsaturated fatty acids. Microalgae are also source of various vitamins, particularly B2 and B3, and pigments such as chlorophyll-a and carotenoids (<xref ref-type="bibr" rid="B184">Zhang et&#xa0;al., 2023</xref>). Cultivation conditions, including irradiance and residence time, strongly influence the nutritional composition. Species such as <italic>Isochrysis galbana</italic>, <italic>Dunaliella tertiolecta</italic>, and <italic>Tetraselmis gracilis</italic> have shown promising nutritional profiles (<xref ref-type="bibr" rid="B184">Zhang et&#xa0;al., 2023</xref>)that meet the United Nations Food and Agriculture Organization nutritional requirements for adults and children, highlighting their potential for food applications.</p>
</sec>
<sec id="s4">
<title>Impact of microalgae on fish growth</title>
<p>Microalgae have great potential as sustainable aquafeed ingredients, offering high nutritional value and environmental benefits. Microalgal species, such as <italic>Nannochloropsis salina</italic> and <italic>Dunaliella salina</italic> can accumulate substantial lipid and protein contents while fixing carbon dioxide (CO<sub>2</sub>) (<xref ref-type="bibr" rid="B29">Chen et&#xa0;al., 2019</xref>). While microalgae are often highlighted for their potential to lower the carbon footprint of aquafeed production, current evidence remains limited. Specifically, comprehensive life-cycle assessment (LCA) studies comparing microalgae with conventional feed ingredients such as fishmeal and fish oil are scarce. Further system-level evaluations are needed to substantiate these environmental claims. Nonetheless, the ability of microalgae to fix CO<sub>2</sub> and reduce reliance on fish stocks presents a promising opportunity for developing more sustainable aquafeed strategies. Microalgae are emerging as a sustainable and nutritious alternative to traditional aquafeed ingredients such as fishmeal and fish oil (<xref ref-type="bibr" rid="B107">Nagappan et al., 2021</xref>; <xref ref-type="bibr" rid="B94">Ma and Hu, 2024</xref>). They offer a high protein content, essential amino acids, omega-3 fatty acids, and bioactive compounds that enhance the growth, colouration, immunity, and survival rates of aquatic species (<xref ref-type="bibr" rid="B43">Dineshbabu et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B72">Idenyi et&#xa0;al., 2022</xref>). Microalgae play a crucial role in aquaculture by supplying essential nutrients that support the health and growth of fish and shellfish (<xref ref-type="bibr" rid="B74">Kapara, 2018</xref>). These microscopic organisms are rich in amino acids, long-chain polyunsaturated fatty acids, vitamins, proteins, and minerals, which are particularly important for enhancing the larval survival, growth, and overall well-being of aquatic species (<xref ref-type="bibr" rid="B154">Siddik et&#xa0;al., 2024</xref>). Microalgae are used as live feed for various growth stages of molluscs, crustaceans, and some fish species. Additionally, certain microalgae contain bioactive compounds with antioxidant, anti-inflammatory, and immunomodulatory properties that can improve immunity and disease resistance in farmed aquatic animals (<xref ref-type="bibr" rid="B1">Abdel-Latif et&#xa0;al., 2022</xref>). Although microalgae are typically cultivated in-house in hatcheries, commercial concentrates are becoming more widely used. However, the high cost of algal biomass limits its widespread use in commercial aquafeeds (<xref ref-type="bibr" rid="B154">Siddik et&#xa0;al., 2024</xref>). Co-cultivation of microalgae with nitrogen-fixing bacteria that release ammonium can significantly reduce the costs of algal biomass. Microalgae are sustainable sources of omega-3 fatty acids, particularly docosahexaenoic acid (DHA) and eicosapentaenoic acid (EPA), which are essential for human and animal nutrition (<xref ref-type="bibr" rid="B113">Norzagaray-Valenzuela et&#xa0;al., 2017</xref>). These fatty acids provide significant health benefits especially for cardiovascular health and brain function in humans. Since fish oil is the traditional source of these compounds concerns over depletion of global fish stocks has led to an increased interest in alternative sources (<xref ref-type="bibr" rid="B167">Topuz, 2016</xref>). Microalgae as primary producers of omega-3 fatty acids can be cultivated at industrial scale and processed into various food products and animal feeds (<xref ref-type="bibr" rid="B113">Norzagaray-Valenzuela et al., 2017</xref>). Recent advancements in microalgal biotechnology, such as metabolic engineering and selective breeding, have further enhanced the potential of omega-3 fatty acid production in autotrophic microalgae. As research progresses, microalgal oil is expected to become a viable replacement for fish oil (<xref ref-type="bibr" rid="B167">Topuz, 2016</xref>).Among microalgae, <italic>Nannochloropsis</italic> spp., show strong potential as sustainable aquafeed ingredients, capable of replacing fishmeal and fish oil. These species can accumulate high lipid (20&#x2013;46%) and protein (30&#x2013;57%) contents while also efficiently fixing carbon dioxide CO<sub>2</sub> (<xref ref-type="bibr" rid="B31">Chen et&#xa0;al., 2020b</xref>). They also produce valuable EPA and can be cost-effectively cultivated in solar-powered open ponds (<xref ref-type="bibr" rid="B91">Li et&#xa0;al., 2020</xref>). Although <italic>Nannochloropsis</italic> spp. have lower digestibility than <italic>Isochrysis</italic> sp. in rainbow trout, they remain a promising fishmeal substitute (<xref ref-type="bibr" rid="B144">Sarker et&#xa0;al., 2020</xref>). Additionally, <italic>Schizochytrium</italic> have shown high digestibility of macronutrients, energy, and fatty acids, particularly DHA, in rainbow trout at both 8&#xb0;C and 15&#xb0;C, further supporting its potential as a fish oil substitute in aquafeeds (<xref ref-type="bibr" rid="B18">B&#xe9;langer et&#xa0;al., 2021</xref>). In Nile tilapia diets, the complete replacement of fish oil with <italic>Schizochytrium</italic> resulted in improved growth, increased feed efficiency, and a higher DHA content in fillets (<xref ref-type="bibr" rid="B143">Sarker et&#xa0;al., 2016</xref>).</p>
<p>
<italic>Schizochytrium</italic> supplemented diets have demonstrated higher phosphorus digestibility and lower solid phosphorus discharge in tilapia, indicating potential environmental benefits (<xref ref-type="bibr" rid="B56">Gamble et&#xa0;al., 2021</xref>). The optimal inclusion level of <italic>Schizochytrium</italic> in fish feed varies by species, ranging from 20 to 80% fish oil replacement, and has been associated with improved growth, survival, and feed intake across various fish species (<xref ref-type="bibr" rid="B121">Pratiwi and Zidni, 2023</xref>). In addition to <italic>Schizochytrium</italic>, <italic>Tetraselmis</italic> spp. have also emerged as promising microalgal ingredients in aquafeeds. For example dietary supplements with <italic>Tetraselmis suecica</italic> has been shown to improve growth performance, feed utilisation, and gene expression in Pacific white shrimp (<italic>Litopenaeus vannamei</italic>) (<xref ref-type="bibr" rid="B150">Sharawy et&#xa0;al., 2020</xref>). Furthermore, cultivation of <italic>Tetraselmis striata</italic> has been optimised at both laboratory and pilot scales, yielding biomass rich in proteins, lipids, carbohydrates, pigments and notable high EPA content (<xref ref-type="bibr" rid="B118">Patrinou et&#xa0;al., 2023</xref>). <italic>Spirulina</italic>, a cyanobacterium also referred to as blue-green algae, has gained attention as a potential aquafeed ingredient due to its high nutritional value and sustainability (<xref ref-type="bibr" rid="B51">El-Sheekh et&#xa0;al., 2023</xref>). It offers a promising alternative to fishmeal, addressing the growing demand for aquaculture feed while reducing environmental impacts (<xref ref-type="bibr" rid="B107">Nagappan et&#xa0;al., 2021</xref>)<italic>. Spirulina</italic> is rich in proteins, essential amino acids, fatty acids, vitamins, and minerals, making it suitable for use by various aquatic species (<xref ref-type="bibr" rid="B128">Ragaza et&#xa0;al., 2020</xref>). In addition, it enhances the innate immunity and disease resistance of fish and shrimp (<xref ref-type="bibr" rid="B129">Rakocy, 2012</xref>). Different species of microalgae, such as <italic>Nannochloropsis</italic>, <italic>Schizochytrium</italic>, and <italic>Isochrysis</italic>, have been incorporated into fish diets. In some studies, microalgae-based diets (e.g. <italic>Nannochloropsis</italic> spp. and <italic>Nanochloropsis salina</italic>) resulted in comparable or slightly lower weight gain than the reference diets but showed acceptable growth rates and feed efficiency. Notable exceptions include defatted <italic>Nannochloropsis oculata</italic> and <italic>Schizochytrium</italic> in juvenile Nile tilapia, where the microalgae-based diet resulted in a higher specific growth rate (SGR) and weight gain than the reference diet. <italic>Schizochytrium</italic> spp. fed to Atlantic salmon and Pacific white shrimp resulted in increased weight gain compared with the reference diets, highlighting the high lipid content of microalgae as an effective substitute for fish oil.</p>
<p>
<italic>Schizochytrium</italic> resulted in a higher weight gain (426 g) than fish oil (326 g) in Atlantic salmon (<italic>Salmo salar</italic>) (<xref ref-type="bibr" rid="B172">Wei et&#xa0;al., 2021</xref>). Defatted <italic>N. oculata</italic> and <italic>Schizochytrium</italic> sp. diets outperformed the reference diet in terms of weight gain and SGR in juvenile Nile tilapia (<italic>Oreochromis niloticus</italic>) (<xref ref-type="bibr" rid="B73">Ju et&#xa0;al., 2017</xref>). Diets with microalgae such as <italic>Schizochytrium</italic> (<xref ref-type="bibr" rid="B9">Allen et&#xa0;al., 2019</xref>) have demonstrated competitive growth performance in shrimp (<italic>Litopenaeus vannamei</italic>). Microalgal diets generally maintain feed conversion ratio (FCR) values that are similar to or slightly higher than those of the reference diets. In some cases, a lower FCR (e.g. <italic>Schizochytrium</italic> spp.) for Atlantic salmon indicates efficient feed utilisation. Thus, microalgae show promise as sustainable feed ingredients with growth performance comparable to that of traditional feeds, especially for specific species. Variations in performance suggest that diet formulations need to be species specific and that some microalgae might not completely replace conventional ingredients without compromising efficiency. Data from previous studies support the viability of microalgae as a sustainable alternative to traditional aquafeed, provided that they are tailored to the nutritional needs of the target fish species in aquaponic systems. <xref ref-type="table" rid="T1"><bold>Table 1</bold></xref> shows the list of microalgae and its effect on fish growth.</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Effect of microalgae on fish growth.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Microalgae</th>
<th valign="middle" align="left">Fish species</th>
<th valign="middle" align="left">Effect</th>
<th valign="middle" align="left">Reference</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">
<italic>Schizochytrium</italic> sp.</td>
<td valign="middle" align="left">Nile tilapia (<italic>Oreochromis niloticus</italic>)</td>
<td valign="middle" align="left">Improved gut health</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B156">Souza et&#xa0;al., 2020</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Euglena</italic> sp.</td>
<td valign="middle" align="left">Atlantic salmon</td>
<td valign="middle" align="left">Immunostimulant</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B80">Kiron et&#xa0;al., 2016a</xref>, <xref ref-type="bibr" rid="B81">b</xref>; <xref ref-type="bibr" rid="B106">Montoya et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B176">Yamamoto et&#xa0;al., 2018</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Schizochytrium</italic> sp.</td>
<td valign="middle" align="left">Atlantic salmon</td>
<td valign="middle" align="left">Enhanced fillet firmness</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B83">Kousoulaki et&#xa0;al., 2016</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Spirulina</italic> sp.</td>
<td valign="middle" align="left">Red tilapia, Koi, Striped jack, Black tiger prawn, and yellow catfish</td>
<td valign="middle" align="left">Enhanced coloration</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B13">Ansarifard et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B43">Dineshbabu et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B92">Liu et&#xa0;al., 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">2.5% <italic>Phaeodactylum tricornutum</italic>
</td>
<td valign="middle" align="left">Gilthead seabream</td>
<td valign="middle" align="left">High fucoxanthin content</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B135">Ribeiro et&#xa0;al., 2017</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Arthrospira platensis</italic>
</td>
<td valign="middle" align="left">Freshwater prawns (<italic>Macrobrachium rosenbergii</italic>)</td>
<td valign="middle" align="left">Enhanced growth performance</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B127">Radhakrishnan et&#xa0;al., 2016</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">5% <italic>Schizochytrium</italic> sp. oil</td>
<td valign="middle" align="left">Atlantic salmon (<italic>Salmo salar L.</italic>)</td>
<td valign="middle" align="left">Weight gain</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B172">Wei et&#xa0;al., 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">0.75% <italic>Tetraselmis suecica</italic>
</td>
<td valign="middle" align="left">Post larvae pacific white shrimp (<italic>Litopenaeus vannamei</italic>)</td>
<td valign="middle" align="left">30% weight gain</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B150">Sharawy et&#xa0;al., 2020</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">15% <italic>Chlorella</italic> sp.</td>
<td valign="middle" align="left">Nile tilapia (<italic>Oreochromis niloticus</italic>)</td>
<td valign="middle" align="left">30% reduction in FCR (feed conversion ratio)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B53">Fadl et&#xa0;al., 2020</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Nannochloropsis gaditana</italic>
</td>
<td valign="middle" align="left">African catfish and Nile tilapia</td>
<td valign="middle" align="left">Improved weight gain and FCR</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B4">Agboola et&#xa0;al., 2019</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Spirulina</italic>-based fish feed</td>
<td valign="middle" align="left">Mozambique tilapia fingerlings (<italic>Oreochromis mossambicus)</italic>
</td>
<td valign="middle" align="left">Improved digestibility</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B151">Sharma et&#xa0;al., 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Nannochloropsis</italic> sp.extruded feed</td>
<td valign="middle" align="left">Gibel carp</td>
<td valign="middle" align="left">Improved digestibility</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B152">Shi et&#xa0;al., 2016</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Pavlova</italic> sp., <italic>Chaetoceros</italic> sp., <italic>Nannochloropsis ocu lata</italic>, and <italic>Isochrysis</italic> sp., in feed,</td>
<td valign="middle" align="left">Seahorses <italic>(Hippocampus reidi)</italic> and Oysters <italic>(Pinctada margaritifera)</italic>
</td>
<td valign="middle" align="left">Increased survivability</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B99">Mart&#xed;nez-Fern&#xe1;ndez and Southgate, 2007</xref>; <xref ref-type="bibr" rid="B102">M&#xe9;lo et&#xa0;al., 2016</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">1-2% <italic>Dunaliella salina</italic> supplemented feed</td>
<td valign="middle" align="left">
<italic>Litopenaeus vannamei</italic>
</td>
<td valign="middle" align="left">Increased survival rate</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B101">Medina-F&#xe9;lix et&#xa0;al., 2014</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Tetraselmis suecica</italic> live cells</td>
<td valign="middle" align="left">White shrimp (F<italic>enneropenaeus indicus</italic>
</td>
<td valign="middle" align="left">Reduced gut pathogenic bacterial load</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B131">Regunathan and Wesley, 2004</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Microencapsulated <italic>Chaetoceros</italic> sp.</td>
<td valign="middle" align="left">Pacific white shrimp <italic>(Lito penaeus vannamei)</italic>
</td>
<td valign="middle" align="left">Survivability at larval stage increased</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B111">Nimrat et&#xa0;al., 2011</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Paramylon in <italic>Euglena</italic> sp. cell wall</td>
<td valign="middle" align="left">Atlantic salmon, mussels, red drum, and matrinxa</td>
<td valign="middle" align="left">Immunostimulant</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B20">Bianchi et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B80">Kiron et&#xa0;al., 2016a</xref>, <xref ref-type="bibr" rid="B81">b</xref>; <xref ref-type="bibr" rid="B106">Montoya et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B176">Yamamoto et&#xa0;al., 2018</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">6-8% of <italic>Chlorella vulgaris</italic>
</td>
<td valign="middle" align="left">Post larvae of <italic>Macrobrachium rosenbergii</italic>
</td>
<td valign="middle" align="left">Improved immune response and survivability agianst <italic>Aeromonas hydrophila</italic> infection</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B97">Maliwat et&#xa0;al., 2017</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Tetraselmis chuii, Nanno chloropsis gaditana</italic>, and <italic>P. tricornutum</italic>
</td>
<td valign="middle" align="left">Gilthead seabream (<italic>Sparus. aurata)</italic>
</td>
<td valign="middle" align="left">Enhanced defence activity</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B26">Cerezuela et&#xa0;al., 2012</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Euglena viridis</italic>
</td>
<td valign="middle" align="left">Rohu fish <italic>(Labeo rohita)</italic>
</td>
<td valign="middle" align="left">Increased immunostimulatory effects</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B35">Das et&#xa0;al., 2009</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Dunaliella salina</italic>
</td>
<td valign="middle" align="left">
<italic>Penaeus monodon</italic>
</td>
<td valign="middle" align="left">Increased antioxidant factors and survival rate</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B95">Madhumathi and Rengasamy, 2011</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Fish diet with <italic>Lactobacillus sakei</italic> and <italic>Navicula</italic> sp.</td>
<td valign="middle" align="left">Pacific red snapper <italic>(Lutjanus peru)</italic>
</td>
<td valign="middle" align="left">Improved humoral response</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B134">Reyes-Becerril et&#xa0;al., 2013</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Feed with <italic>Prunus incisa</italic>
</td>
<td valign="middle" align="left">Guppy fish (<italic>Poecilia reticulata)</italic>
</td>
<td valign="middle" align="left">Increased survival rate</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B109">Nath et&#xa0;al., 2012</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">10% <italic>A. Platensis</italic> diet</td>
<td valign="middle" align="left">Rainbow trout <italic>(Oncorhynchus mykiss)</italic>
</td>
<td valign="middle" align="left">Increased in total proteins level</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B179">Yeganeh et&#xa0;al., 2015</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">5% <italic>Schizochytrium</italic> sp.</td>
<td valign="middle" align="left">Atlantic salmon</td>
<td valign="middle" align="left">Improved fillets quality</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B83">Kousoulaki et&#xa0;al., 2016</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Schizochytrium limacinum</italic>
</td>
<td valign="middle" align="left">Atlantic salmon</td>
<td valign="middle" align="left">Improved fillets taste and odour</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B77">Katerina et&#xa0;al., 2020</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Schizo chytrium</italic> sp.</td>
<td valign="middle" align="left">Atlantic salmon</td>
<td valign="middle" align="left">Rich in PUFA (polyunsaturated fatty acids)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B132">Ren et&#xa0;al., 2010</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">4% defatted-<italic>Spirulina</italic> and 0.4% <italic>Spirulina</italic>-lipid-ex tract</td>
<td valign="middle" align="left">Yellow catfish <italic>(Pelteobagrus fulvidraco)</italic>
</td>
<td valign="middle" align="left">Improved skin colour</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B92">Liu et&#xa0;al., 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">7.5% <italic>Spirulina platensis</italic>
</td>
<td valign="middle" align="left">Showa koi</td>
<td valign="middle" align="left">Improved pigmentation</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B161">Sun et&#xa0;al., 2012</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s5">
<title>Impact of microalgae on plant growth</title>
<p>Studies have shown that the integration of microalgae into aquaponic systems can improve the physicochemical properties of aquaculture water (<xref ref-type="bibr" rid="B3">Addy et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B162">Tejido-Nu&#xf1;ez, 2020</xref>). Residual fish feed and fish excreta that accumulate in aquaculture water can be used by microalgae to support its growth and biomass production (<xref ref-type="bibr" rid="B41">Delrue et&#xa0;al., 2016</xref>). Microalgae cultivation in aquaponics helps to improve water quality by decreasing the pH. Microalgae interactions with bacteria could be the reason for the conversion of fish waste into nutrients, thereby increasing water quality. However, studies on the mechanism of fish waste-to-nutrient conversion through interactions between algae and bacteria are scarce. It has been hypothesised that microalgae facilitate the proliferation of beneficial bacteria and reduce the risk of pathogenic bacteria that could otherwise cause diseases in fish and plants in aquaponic systems. A study evaluating the effects of three microalgal species (<italic>Chlorella vulgaris</italic>, <italic>Scenedesmus</italic> spp., and <italic>Spirulina platensis</italic>), cultivated in an aquaponic system along with Nile tilapia and garlic plants, showed growth similar to that of the control in terms of plant biomass, leaf number, and shoot length. Water quality parameters such as dissolved oxygen, pH, temperature, ammonia, nitrate, and nitrite were maintained at ideal levels for aquaponic systems (<xref ref-type="bibr" rid="B3">Addy et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B163">Tejido-Nu&#xf1;ez et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B30">Chen et&#xa0;al., 2020a</xref>).</p>
<p>A previous study revealed that diverse populations of beneficial microorganisms were significantly higher in fish tanks and biofilters when microalgae were co-cultivated in an aquaponic system. This was a positive outcome owing to the mutual interaction between microalgae and bacteria, which could play an important role in nutrient cycling. Studies have shown that microalgae play a crucial role in atmospheric nitrogen recycling and soil fertility. They can fix atmospheric nitrogen into bioavailable forms like ammonia, particularly through specialized cells called heterocysts in cyanobacteria (<xref ref-type="bibr" rid="B155">Singh, 2021</xref>). Many studies have highlighted that bacterial richness is higher in the presence of <italic>Chlorella vulgaris</italic>, which helps in the removal of nitrogen and phosphorus. A stable association between <italic>C. vulgaris</italic> and specific bacterial species such as <italic>Flavobacterium</italic> sp., <italic>Terrimonas</italic> sp., <italic>Sphingobacterium</italic> sp., <italic>Rhizobium</italic> sp., and <italic>Hyphomonas</italic> sp. has been observed (<xref ref-type="bibr" rid="B130">Ramanan et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B66">Han et&#xa0;al., 2019</xref>). In aquaponic systems, compared with fish tanks, biofilters had a higher bacterial population because they act as a growth substrate for microalgae and bacteria that form biofilm known as a &#x2018;phycosphere&#x2019;. Diverse beneficial bacterial species are attracted to this phycosphere, and this algae&#x2013;bacteria interaction plays a major role in regulating water quality in aquatic environments. Nitrogen cycling in aquaponic systems occurs because of the presence of the phyla Proteobacteria and Bacteroidetes, which are indicators of the good health status of the system (<xref ref-type="bibr" rid="B146">Schmautz et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B175">Wongkiew et&#xa0;al., 2018</xref>). Bacteroidetes convert nitrates into various nitrogen compounds that are essential for degrading complex organic matter (<xref ref-type="bibr" rid="B175">Wongkiew et&#xa0;al., 2018</xref>). In one study, it was found that Bacteroidetes were more abundant when microalgae were present. Bacteroidetes play a vital role in nutrient cycling and support optimal plant growth in aquaponics (<xref ref-type="bibr" rid="B76">Kasozi et&#xa0;al., 2021</xref>). In addition, fish fed microalgae showed resistance to bacterial infection. In a previous study, Nile tilapia fish fed <italic>Spirulina platensis</italic> in an aquaponic system showed lower mortality than the control group. Some studies have shown that microalgae such as <italic>Chlorella vulgaris</italic> can produce the antibacterial compound &#x201c;chlorellin&#x201d;. Another study showed that when <italic>Spirulina platensis</italic> was given as a feed supplement, the antibacterial compound &#x201c;phycocyanin&#x201d; it produces decreased the mortality of Nile tilapia. <italic>Nannochloropsis oculata</italic>, <italic>Schizochytrium</italic> sp., and <italic>Spirulina</italic> sp., the microalgal mix used in Nile tilapia feed, increased immunity against <italic>Vibrio</italic> and <italic>Staphylococcus</italic> bacterial species and enhanced its antioxidant enzyme activity (<xref ref-type="bibr" rid="B54">Falaise et&#xa0;al., 2016</xref>). Studies have indicated that purple sulphur bacteria such as those from the genus <italic>Thiobaca</italic> play a key role in the sulphur cycle and were observed to be more abundant in aquaponics water treated with <italic>Chlorella vulgaris</italic>. Studies on iron-reducing bacterial species belonging to the genus <italic>Geothrix</italic> have shown that they oxidise organic compounds by reducing iron (III) to iron (III) oxide, manganese (IV) oxide, and nitrate, thereby preventing the production of environmentally harmful compounds in aquaponic systems. Another study revealed that the bacterial genus <italic>Fusibacter</italic> contributes to the reduction of elemental sulphur, or thiosulfate, to sulphides during the sulphur cycle. Previous studies on the bacterial genus <italic>Treponema</italic> showed that it plays a crucial role in scavenging nutrients through fermentation processes (<xref ref-type="bibr" rid="B22">Buyuktimkin et&#xa0;al., 2019</xref>). These biological processes are important for converting fish waste into nutrient solutions in aquaponic systems. In aquaponics, the co-cultivation of microalgae (e.g. <italic>C. vulgaris</italic>, <italic>Scenedesmus</italic> sp., and <italic>Spirulina platensis</italic>) showed better performance of the bacterial genera <italic>Thiobaca</italic>, <italic>Geothrix</italic>, <italic>Fusibacter</italic>, and <italic>Treponema</italic> in terms of nutrient cycling. Further studies on microalgae&#x2013;bacteria interactions will provide insights into the effects on fish growth and plant development in aquaponics (<xref ref-type="bibr" rid="B145">Schmautz et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B76">Kasozi et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B17">Bartelme et&#xa0;al., 2019</xref>). Microalgae enhance plant growth by acting as biostimulant, biofertilizers, and biopesticides. These properties are due to the presence of bioactive compounds such as phenols, phytohormones, amino acids, polysaccharides, and terpenoids (<xref ref-type="bibr" rid="B90">Lee and Ryu, 2021</xref>). Although microalgae play an important role in nutrient uptake and cycling, it&#x2019;s the cyanobacteria often referred as blue green algae that are responsible for nitrogen fixation. Microalgae can mobilize nutrients like phosphate, potassium, and copper <xref ref-type="bibr" rid="B173">Win et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B63">Gon&#xe7;alves, 2021</xref>). Another essential micronutrient molybdenum (MO) is a key cofactor for two enzymes: nitrogenase (in nitrogen-fixing microbes like diazotrophs or cyanobacteria) and nitrate reductase (in microalgae and plants). These enzymes are essential for converting atmospheric nitrogen (N<sub>2</sub>) into ammonium and nitrate (NO<sub>3</sub>&#x207b;) into usable nitrogen forms, respectively. In integrated aquaponics systems where microalgae or nitrogen-fixing bacteria are involved, the presence of trace levels of Mo ensures these microbes can effectively perform biological nitrogen fixation and nitrate assimilation (<xref ref-type="bibr" rid="B60">Glass et&#xa0;al., 2012</xref>).</p>
<p>Microalgae, particularly cyanobacteria, have a specific mechanism for fixing nitrogen. Cyanobacteria, such as <italic>Cyanothece</italic> spp., <italic>Lyngbya</italic> spp., and <italic>Trichodesmium</italic> spp., colonise the leaves and roots of plants, penetrate cell tissues, and colonise internally with plant host specificity. Microalgae produce enzymes such as alkaline phosphatases, 5&#x2019;nucleotidases, phytases, and phosphodiesterases that help to release bound phosphorus from organic sources such as phytate. Some species such as <italic>Tetraselmis suecia</italic>, <italic>Nannochloropsis gaditana</italic>, and <italic>Nanochloropsis oceania</italic> adopt a luxury uptake mechanism to store excess or relocate phosphorus by remodelling polar lipids (<xref ref-type="bibr" rid="B24">Ca&#xf1;avate et&#xa0;al., 2017</xref>). Microalgal species, such as <italic>Spirulina platensis</italic>, <italic>Chlorella</italic> spp., <italic>Scenedesmus</italic> spp., <italic>Acutodesmus</italic> spp., <italic>Calothrix elenkini</italic>, and <italic>Dunaliella</italic> spp (<xref ref-type="bibr" rid="B136">Ronga et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B33">Colla and Rouphael, 2020</xref>), enhance crop production by improving nutrient uptake, enhancing resistance to both abiotic and biotic stress, and maintaining essential functions such as respiration, photosynthesis, nucleic acid synthesis, and iron uptake (<xref ref-type="bibr" rid="B90">Lee and Ryu, 2021</xref>; <xref ref-type="bibr" rid="B84">Kumar et&#xa0;al., 2022</xref>).</p>
<p>Microalgae synthesise phytohormones that play important roles in shoot and root development, plant tissue differentiation, aging, and defence against biotic and abiotic stressors. Studies on microalgal species, such as <italic>Coenochloris</italic> spp., <italic>Chlorella</italic> spp., <italic>Scenedesmus</italic> spp., <italic>Chlorococcum</italic> spp., and <italic>Acutodesmus</italic> spp., have shown that they can synthesise auxin hormones, such as indol-3-acetamide and indole 3-acetic acid, which play roles in the formation and elongation of plant roots (<xref ref-type="bibr" rid="B75">Kapoore et&#xa0;al., 2021</xref>). Microalgae that synthesise auxins form colonies with cyanobacteria, which has been observed in wheat and rice plants (<xref ref-type="bibr" rid="B71">Hussain et&#xa0;al., 2017</xref>). Recent studies have revealed that the green alga <italic>Chlamydomonas reinhardtii</italic> can synthesize auxin (indole-3-acetic acid, IAA) through an extracellular L-amino acid oxidase (LAO1) under nitrogen-limited conditions (<xref ref-type="bibr" rid="B23">Calatrava et&#xa0;al., 2022</xref>). This auxin production plays a role in algal-bacterial mutualism, particularly with Methylobacterium species. <italic>Nannochloropsis</italic> spp. synthesise cytokinin phytohormones that enhance resistance to nitrogen and water stress in tomato plants. Studies on <italic>Chlorella vulgaris</italic> extracts containing gibberellic acid phytohormones suggest that the extract could mitigate the harmful effects of heavy metals, such as lead and cadmium, on plant growth. Microalgae belonging to the genera <italic>Chlorella</italic>, <italic>Chlamydomonas</italic>, and <italic>Scenedesmus</italic>, and cyanobacteria including <italic>Anabaena</italic> spp., <italic>Synechococcus</italic> spp., <italic>Calothrix</italic> spp., <italic>Nostoc</italic> spp., <italic>Cylindrospermum</italic> spp., and <italic>Scytonema</italic> spp., have been reported to synthesise ethylene phytohormones that regulate cell division, fruit ripening, aging, and biotic and abiotic stress tolerance (<xref ref-type="bibr" rid="B67">Han et&#xa0;al., 2018</xref>). Microalgae can synthesise signalling molecules, such as jasmonic acid, polyamines, brassinosteroids, and salicylic acid, which are associated with stress-tolerance mechanisms that enhance enzymatic and non-enzymatic defence responses in plants (<xref ref-type="bibr" rid="B75">Kapoore et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B90">Lee and Ryu, 2021</xref>). Some studies have reported that <italic>Spirulina</italic> can produce polyamines which promote the growth of lettuce seedlings (<xref ref-type="bibr" rid="B105">M&#xf3;gor et&#xa0;al., 2018</xref>). Studies on microalgae, such as <italic>Chlorella stigmatophora</italic>, <italic>Chlorella vulgaris</italic>, <italic>Tetraselmis</italic> spp., <italic>Dunaliella salina</italic>, and <italic>Porphyridium cruentum</italic>, have shown that microalgae can synthesise exopolysaccharides (EPS) which stimulate plant growth and metabolism (<xref ref-type="bibr" rid="B28">Chanda et&#xa0;al., 2019</xref>). <xref ref-type="bibr" rid="B50">El Arroussi et&#xa0;al. (2018)</xref> reported that EPS from <italic>Dunaliella saline</italic> microalgal species enhanced salinity stress tolerance in tomato plants. Studies on protein-rich extracts of <italic>Spirulina platensis</italic> have revealed increased flower number, freshness, and dry weight in <italic>Petunia x hybrida</italic> plants (<xref ref-type="bibr" rid="B120">Plaza et&#xa0;al., 2018</xref>). Green algal extracts rich in amino acids enhanced the total solid and organic contents of three hot pepper varieties (<xref ref-type="bibr" rid="B182">Zamljen et&#xa0;al., 2021</xref>). Some microalgal species also synthesise phenolic compounds and carotenoids, which support photoprotection and defence responses in plants (<xref ref-type="bibr" rid="B168">Vidyashankar et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B27">Cezare-Gomes et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B40">Del Mondo et&#xa0;al., 2021</xref>). Some microalgae species are a good source of micronutrients, such as calcium,, iron, zinc, a and magnesium (<xref ref-type="bibr" rid="B142">Sandgruber et&#xa0;al., 2021</xref>). <italic>Tetraselmis chuii</italic> is rich in total calcium and phosphorus, <italic>Chlorella</italic> has high phosphorus and iron contents, and <italic>Spirulina</italic> is rich in potassium. Microalgae synthesize vitamins that act as plant growth promoting factors. Vitamin C and nicotinic acid are abundant in <italic>Tetraselmis suecia</italic>. Freshwater microalgae, such as <italic>Spirulina platensis</italic> and <italic>Chlorella</italic> spp., are rich in vitamins such as niacin, riboflavin, cyanocobalamin, and folic acid (<xref ref-type="bibr" rid="B48">Edelmann et&#xa0;al., 2019</xref>). Studies have shown that plants like soybean, barley, and spinach absorb vitamin B complex when microalgal biomass is applied as biofertilizer. In addition to vitamins and minerals, microalgae produce terpenoids, betaines, humic substances, and peptides that function as biopesticides (<xref ref-type="bibr" rid="B75">Kapoore et&#xa0;al., 2021</xref>). <xref ref-type="table" rid="T2"><bold>Table 2</bold></xref> shows the list of microalgae and its effect on plant growth.</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Effect of microalgae on plant growth.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Microalgae</th>
<th valign="middle" align="center">Plant species</th>
<th valign="middle" align="center">Effect</th>
<th valign="middle" align="center">Reference</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">
<italic>Spirulina platensis</italic>
</td>
<td valign="middle" align="left">
<italic>Raphanus sativus</italic>
</td>
<td valign="middle" align="left">Enhanced germination rate and seedling vigour</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B62">Godlewska et&#xa0;al., 2019</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Chlorella vulgaris</italic>
</td>
<td valign="middle" align="left">
<italic>Solanum lycopersicum</italic> L., <italic>Cucumus sativus</italic>
</td>
<td valign="middle" align="left">Improved root parameters, increased biomass yield</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B21">Bumandalai and Tserennadmid, 2019</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Scenedesmus quadricauda</italic>, <italic>Chlorella vulgaris</italic>, <italic>Arthrospira</italic> spp.</td>
<td valign="middle" align="left">
<italic>Beta vulgaris</italic> L.</td>
<td valign="middle" align="left">Improved root parameters, enhanced biomass and nutritional quality</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B16">Barone et&#xa0;al., 2018</xref>;</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Navicula</italic> spp.</td>
<td valign="middle" align="left">
<italic>Solanum lycopersicum</italic> L., <italic>Capsicum annuum</italic> L., <italic>Solanum melongena</italic>
</td>
<td valign="middle" align="left">Enhanced biomass</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B10">Alshehrei et&#xa0;al., 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Oscillatoria agardhii</italic>
</td>
<td valign="middle" align="left">
<italic>Triticum</italic> spp.</td>
<td valign="middle" align="left">Drought tolerance</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B64">Haggag et&#xa0;al., 2018</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Chlorella vulgaris</italic>, <italic>Nannochloropsis salina</italic>
</td>
<td valign="middle" align="left">
<italic>Moringa oleifera</italic>
</td>
<td valign="middle" align="left">Salinity tolerance</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B8">Al Dayel and El Sherif, 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Chlorella vulgaris</italic>
</td>
<td valign="middle" align="left">
<italic>Vigna mungo</italic> L.</td>
<td valign="middle" align="left">Enhanced growth (acts as a biostimulant)</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B44">Dineshkumar et&#xa0;al., 2019</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Spirulina</italic> extract</td>
<td valign="middle" align="left">
<italic>Triticum aestivum, Hordeum vulgare</italic>
</td>
<td valign="middle" align="left">Enhanced germination and biomass yield</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B6">Akg&#xfc;l, 2019</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Spirulina platensis</italic> extract</td>
<td valign="middle" align="left">
<italic>Calotropis procera</italic> Ait</td>
<td valign="middle" align="left">Improved root growth and germination rate</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B15">Bahmani Jafarlou et&#xa0;al., 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Spirulina platensis</italic> extract</td>
<td valign="middle" align="left">
<italic>Vigna mungo</italic> L.</td>
<td valign="middle" align="left">Enhanced germination, nutritional content, root growth, biomass and stress tolerance</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B165">Thinh, 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Spirulina platensis</italic> Phycocyanin extract</td>
<td valign="middle" align="left">
<italic>Solanum lycopersicum</italic> L.</td>
<td valign="middle" align="left">Increased biomass, nutritional content, and germination</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B103">Metwally et&#xa0;al., 2022</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Chlorella</italic> spp. Cell suspension</td>
<td valign="middle" align="left">
<italic>Triticum aestivum, Hordeum vulgare</italic>
</td>
<td valign="middle" align="left">Enhanced root development, biomass and germination rate</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B115">Odgerel and Tserendulam, 2016</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Nostoc commune</italic> aqueous extracts</td>
<td valign="middle" align="left">
<italic>Oryza sativa</italic> L.</td>
<td valign="middle" align="left">Enhanced root development, biomass and germination rate</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B2">Abedi Firoozjaei et&#xa0;al., 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Scenedesmus quadricauda</italic> and <italic>Chlorella vulagaris</italic> extract</td>
<td valign="middle" align="left">
<italic>Beta vulgaris</italic>
</td>
<td valign="middle" align="left">Improved seed vigour and root growth</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B124">Puglisi et&#xa0;al., 2020</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Consortia of<break/>
<italic>Chlorococcum</italic> spp. <italic>Micractinium</italic> spp.<break/>
<italic>Scenedesmus</italic> spp.<break/>
<italic>Chlorella</italic> spp.</td>
<td valign="middle" align="left">
<italic>Spinacia oleraceae</italic>
</td>
<td valign="middle" align="left">Enhanced biomass, nutritional content and germination rate</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B138">Rupawalla et&#xa0;al., 2022</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Scenedesmus subspicatus</italic>
</td>
<td valign="middle" align="left">
<italic>Allium cepa</italic> L</td>
<td valign="middle" align="left">Improved root development</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B57">Gemin et&#xa0;al., 2022</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Chlorella vulgaris</italic> biomass with cow dung</td>
<td valign="middle" align="left">
<italic>Solanum lycopersicum</italic> L.</td>
<td valign="middle" align="left">Enhanced root growth, leaf phytochemical content, soil, enzyme activity and stress tolerance</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B159">Suchithra et&#xa0;al., 2022</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Chlorella vulgaris</italic> extract</td>
<td valign="middle" align="left">
<italic>Lactuca sativa</italic>
</td>
<td valign="middle" align="left">Increased crop yield, leaf pigment content, fruits, flowers numbers and nutritional quality</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B87">La Bella et&#xa0;al., 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Chlorella vulgaris</italic>
</td>
<td valign="middle" align="left">
<italic>Brassica oleracea</italic> var. italica</td>
<td valign="middle" align="left">Enhanced leaf pigments, stress tolerance, enzymatic activity, early flowering</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B85">Kusvuran, 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Chlorella vulgaris</italic> extract</td>
<td valign="middle" align="left">
<italic>Latuca sativa</italic> L.</td>
<td valign="middle" align="left">Increased enzymatic activity, early flowering, nutritional quality</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B125">Puglisi et&#xa0;al., 2022</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Cell lysates of <italic>Chlamydomonas reinhardtii</italic> CC124<break/>
<italic>Chlorella</italic> sp. MACC360</td>
<td valign="middle" align="left">
<italic>Solanum lycopersicum</italic> L.</td>
<td valign="middle" align="left">Improved crop yield, enzymatic activity, number of fruits, early flowering</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B59">Gitau et&#xa0;al., 2022</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Polysaccharides extract of <italic>Dunaliella salina</italic> MS002 and MS067<break/>
<italic>Phaeodactylum tricornotum</italic> MS023<break/>
<italic>Porphyridium</italic> spp.MS081, <italic>Desmodesmus</italic> spp.<break/>
<italic>Spirulina platensis</italic> MS001</td>
<td valign="middle" align="left">
<italic>Solanum lycopersicum</italic> L.</td>
<td valign="middle" align="left">Enhanced nutritional quality and stress tolerance</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B126">Rachidi et&#xa0;al., 2021</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Extracts of microalgae consortium<break/>
<italic>Chlorella</italic> spp., <italic>Scenedesmus</italic> spp., <italic>Spirulina</italic> spp., <italic>Synechocystis</italic> spp</td>
<td valign="middle" align="left">
<italic>Solanum lycopersicum</italic> L.</td>
<td valign="middle" align="left">Increased biomass, leaf pigment content, nutritional quality</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B68">Hans et&#xa0;al., 2020</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Chlorella vulgaris</italic>
</td>
<td valign="middle" align="left">
<italic>Cyamopsis tetragonoloba</italic> (L.) Taub.</td>
<td valign="middle" align="left">Early flowering, improved nutritional quality and stress tolerance</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B86">Kusvuran and Can, 2020</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">Polysaccharide extracts of <italic>Chlorella vulgaris, Chlorella Sorokiniana</italic>
</td>
<td valign="middle" align="left">
<italic>Solanum lycopersicum</italic> L.</td>
<td valign="middle" align="left">Early flowering, increased enzymatic activity and stress tolerance</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B55">Farid et&#xa0;al., 2019</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Scenedesmus</italic> spp.extract, <italic>Arthrospira platensis</italic> cell hydrolysate</td>
<td valign="middle" align="left">
<italic>Petunia x hybrida</italic>
</td>
<td valign="middle" align="left">Enhanced crop yield, nutritional quality</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B120">Plaza et&#xa0;al., 2018</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Scenedesmus obliquus Chlorella vulgaris</italic> and A<italic>nabaena oryzae</italic> biomass</td>
<td valign="middle" align="left">
<italic>Musa</italic> spp.</td>
<td valign="middle" align="left">Improved root growth, leaf phytochemical content, soil quality, stress tolerance</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B65">Hamouda and El-Ansary, 2017</xref>
</td>
</tr>
<tr>
<td valign="middle" align="left">
<italic>Chlorella fusca</italic>
</td>
<td valign="middle" align="left">
<italic>Cucumis sativus Arabidopsis thaliana</italic>
</td>
<td valign="middle" align="left">Stress tolerance</td>
<td valign="middle" align="left">
<xref ref-type="bibr" rid="B78">Kim et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B89">Lee et&#xa0;al., 2020</xref>
</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s6">
<title>Integration of microalgae in aquaponics</title>
<p>Microalgae serve as excellent nutrient sources for aquatic organisms, providing proteins, omega-3 fatty acids, vitamins, and minerals. They also play crucial roles in water quality management, larviculture, and Integrated Multi-Trophic Aquaculture (IMTA) systems (<xref ref-type="bibr" rid="B70">Hashmi et&#xa0;al., 2023</xref>). Algaeponics is a recent innovation in the field of aquaponics (<xref ref-type="bibr" rid="B108">Nair et&#xa0;al., 2025</xref>) and it is a novel extension of conventional aquaponics that incorporates microalgae as an integral biological component within the system. Unlike standard aquaponics&#x2014;where fish waste provides nutrients for higher plants&#x2014;algaeponics uses microalgae to recycle nutrients, improve water quality, and serve as a supplementary or primary feed source for fish (<xref ref-type="bibr" rid="B185">Zhang et&#xa0;al., 2022</xref>). Microalgae enhance aquaponic systems by supporting nutrient removal, improving water quality, and serving as feed for fish like tilapia (<xref ref-type="bibr" rid="B49">Edwards et&#xa0;al., 1981</xref>; <xref ref-type="bibr" rid="B79">Kinh et&#xa0;al., 2024</xref>). Factors such as fish density, food-to-microorganism ratio (F/M), and hydraulic retention time (HRT) influence algal integration and system stability (<xref ref-type="bibr" rid="B100">Medina and Neis, 2007</xref>). While species like <italic>Chlorella</italic> sp. aid in ammonia control and pH balance, their growth may be limited in systems optimized for fish and plant productivity (<xref ref-type="bibr" rid="B3">Addy et&#xa0;al., 2017</xref>). Microalgae can interact with nitrogen fixing bacteria called diazotrophs that could possess combined biotechnological applications in a sustainable production system. In aquaponics systems, integrating microalgae with nitrogen-fixing bacteria (diazotrophs) offers a promising, sustainable way to enhance nutrient cycling, water quality, and productivity. While microalgae contribute to carbon fixation, oxygenation, and biomass production, diazotrophs help convert atmospheric nitrogen into plant-available forms like ammonium. Together, they can naturally supplement nitrogen when fish waste is insufficient, reduce the need for synthetic inputs, and support plant growth through biofertilization. Additionally, the protein-rich algal biomass can be harvested and reused as fish feed, creating a closed-loop system that improves efficiency, reduces operational costs, and boosts environmental resilience (<xref ref-type="bibr" rid="B93">Llamas et&#xa0;al., 2023</xref>).</p>
<p>Microalgae added to aquaponic systems in the form of aquafeeds is shown in <xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>, as they can act as an essential food source for fish, but in-depth research is necessary to determine their potential benefits for plant growth in aquaponic systems. Microalgae can effectively remediate aquaculture water acting as nutrient recyclers while producing valuable biomass (<xref ref-type="bibr" rid="B46">Dourou et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B66">Han et&#xa0;al., 2019</xref>). This integration reduces environmental impacts, improves water quality, and provides a sustainable source of aquafeed (<xref ref-type="bibr" rid="B66">Han et&#xa0;al., 2019</xref>). Upscaling the production of microalgae can lead to improved resource efficiency and a reduced carbon footprint. Recent advances in recirculating aquaculture systems (RAS) have focused on incorporating microalgae to close the system loop, thereby enhancing performance and deriving value from waste streams. Microalgae in RAS facilitate oxygenation, carbon dioxide sequestration, and nutrient recovery (<xref ref-type="bibr" rid="B52">Ende et&#xa0;al., 2024</xref>). Various cultivation systems, harvesting technologies, and species selection strategies have been explored to optimise microalgae-assisted aquaculture (<xref ref-type="bibr" rid="B66">Han et&#xa0;al., 2019</xref>). Microalgal biomass production has a water footprint of 2857 L/kg when using freshwater. This footprint can be reduced considerably by employing wastewater or seawater or recycling growth media, with recycling potentially lowering the footprint by 90% (<xref ref-type="bibr" rid="B123">Pugazhendhi et&#xa0;al., 2020</xref>). Compared with plant and insect production, microalgae production has a lower water footprint. In open cultivation systems, evaporation is a major contributor to water loss, with evaporation rates reaching up to 2 cm/d in 20 cm deep raceway ponds (<xref ref-type="bibr" rid="B37">Das et&#xa0;al., 2016</xref>). Cultivating microalgae using wastewater for human consumption raises legitimate food safety concerns. These concerns stem from the potential accumulation of harmful substances, including heavy metals, pathogens, and emerging contaminants (<xref ref-type="bibr" rid="B98">Markou et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B11">&#xc1;lvarez-Gonz&#xe1;lez et&#xa0;al., 2023</xref>). While treatment processes such as anaerobic digestion can significantly reduce biological and chemical risks, the persistence of certain xenobiotics remains a challenge. Currently, the legal frameworks in most regions&#x2014;including the European Union&#x2014;do not support the use of such biomass in food products. However, some studies indicate that treated microalgal biomass may be suitable for non-food applications, such as fertilizers and aquafeed, although elements like cadmium can still exceed allowable limits (<xref ref-type="bibr" rid="B11">&#xc1;lvarez-Gonz&#xe1;lez et&#xa0;al., 2023</xref>). Continued research and clearer regulatory guidance are essential as the industry evolves (<xref ref-type="bibr" rid="B42">de Oliveira and Bragotto, 2022</xref>; <xref ref-type="bibr" rid="B140">Salehipour-Bavarsad et&#xa0;al., 2024</xref>).</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Schematic representation of microalgae contributions to Sustainable Aquaponics: From Biomass Cultivation to Aquafeed.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1661042-g002.tif">
<alt-text content-type="machine-generated">Flowchart depicting a balanced aquaponic system integrating microalgae culture and fish tank. The process begins with microalgae culture using CO2 and sunlight, producing algal biomass. This biomass is pelleted into aquafeed for fish. The fish tank connects to a plant trough within the aquaponic system, facilitated by microalgae bacteria consortia. Benefits include improved water quality, efficient nutrient cycling, and enhanced plant and fish growth.</alt-text>
</graphic>
</fig>
<p>Major challenges in microalgae cultivation is its biomass productivity which is highly variable due to numerous cultivation factors such as light intensity and spectrum, nutrient availability, temperature, and strain-specific physiological differences. Light is a key determinant, with suboptimal intensity, poor spectral quality, and inefficient distribution significantly reducing photosynthetic efficiency, especially in dense cultures and closed photobioreactors (<xref ref-type="bibr" rid="B117">Ooms et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B114">Nwoba et&#xa0;al., 2019</xref>). Nutrient limitations, particularly of nitrogen and phosphorus, can both constrain growth and stimulate desired metabolite accumulation, but must be precisely managed to balance productivity and product quality (<xref ref-type="bibr" rid="B32">Chu, 2017</xref>). Additionally, different microalgal strains respond uniquely to environmental conditions, making strain selection critical for consistent biomass yield and target compound production (<xref ref-type="bibr" rid="B158">&#x160;t&#x11b;rbov&#xe1; et&#xa0;al., 2023</xref>). Innovations such as spectral conversion, temperature control strategies, genetically modified strains, and advanced photobioreactor designs aim to mitigate these inconsistencies and improve biomass uniformity and scalability (<xref ref-type="bibr" rid="B114">Nwoba et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B183">Zhang et&#xa0;al., 2024</xref>).Freshwater is often required to counteract evaporation and maintain salinity for marine microalgae. However, certain halotolerant microalgal strains (e.g. <italic>Dunaliella</italic> sp., <italic>Tetraselmis</italic> sp., and <italic>Picochlorum</italic> sp.) can adapt to salinity changes, thereby reducing freshwater use and lowering the overall water footprint (<xref ref-type="bibr" rid="B36">Das et&#xa0;al., 2019</xref>). Microalgae show potential as sustainable alternatives to fish-based aquafeed in addressing the growing demand for high-quality proteins (<xref ref-type="bibr" rid="B164">Tham et&#xa0;al., 2023</xref>; <xref ref-type="bibr" rid="B177">Yarnold et&#xa0;al., 2019</xref>). Integrating microalgae cultivation with aquaculture, agriculture, aquaponics, and livestock farming could create a circular bioeconomy based on recycling nutrients and wastewater. This approach offers environmental benefits, resource recovery, and potential socioeconomic improvements in rural areas. However, challenges remain, including developing large-scale production methods and addressing energy-intensive harvesting and processing methods. Some studies have shown that freshwater microalgae like spirulina might contain contaminants like microcystins (MCs) which have raised increasing concern due to their potential health risks. Spirulina, a cyanobacterial supplement&#x2014;has been examined for safety, especially in France where over 180 small-scale farms contribute to local production. A review of data from 95 producers between 2013 and 2021, showed that MCs levels generally remained within safe limits. These findings support the relative safety of French spirulina and other microalgae while emphasizing the importance of refining cultivation practices to prevent contamination (<xref ref-type="bibr" rid="B148">Scoglio, 2018</xref>; <xref ref-type="bibr" rid="B119">Pinchart et&#xa0;al., 2023</xref>). Further research on life cycle assessment and pilot-scale demonstrations is needed to establish the feasibility and sustainability (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>) of integrating algae-based systems into aquaculture, aquaponics, and related sectors (<xref ref-type="bibr" rid="B169">Vishwakarma et&#xa0;al., 2022</xref>).</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>SWOT analysis of microalgae-based aquafeed in aquaponic systems.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1661042-g003.tif">
<alt-text content-type="machine-generated">SWOT analysis diagram for microalgae aquafeed. Strengths: high in nutrients, renewable, improves water quality. Weaknesses: high cultivation cost, requires optimal conditions, perishable. Opportunities: rising demand, reduces carbon footprint. Threats: environmental risks, market competition, strict regulations.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s7" sec-type="conclusions">
<title>Conclusion</title>
<p>Aquaponics holds immense potential to address global food and nutrition security challenges by integrating fish and plant production in a sustainable manner. However, one of the key limitations of current systems lies in the inefficient conversion of fish effluent into complete nutrient solutions for plant growth, often necessitating external fertiliser inputs. Recent studies suggest that microalgae could offer a promising solution to this bottleneck by serving dual roles as functional aquafeed for fish and as biostimulants or biofertilizers for plants. Certain species, such as <italic>Spirulina</italic> and <italic>Chlorella</italic>, have demonstrated benefits in nutrient recycling, water purification, and enhancement of fish health and plant biomass. While some microalgae species needs to be optimised for its application as aquafeed. Despite these promising insights, the research on microalgae as sustainable aquafeed in aquaponics system remains fragmented and limited in scope. Most existing studies are either species-specific or focused on isolated benefits rather than on integrated system-wide performance. Additionally, the long-term stability, scalability, and economic viability of incorporating microalgae in aquaponics remain underexplored. Future research should aim to systematically evaluate a broader range of microalgal species in aquaponic settings, including their interactions with microbial communities, effects on nutrient dynamics, and their contribution to overall system productivity and resilience. Moreover, multidisciplinary approaches combining aquaculture, plant science, and microbial ecology are needed to optimise microalgae integration. By addressing these knowledge gaps, aquaponics can evolve into a more self-sustaining, circular food production system capable of meeting future global demands.</p>
</sec>
</body>
<back>
<sec id="s8" sec-type="author-contributions">
<title>Author contributions</title>
<p>RM: Conceptualization, Writing &#x2013; original draft, Visualization. CS: Data curation, Writing &#x2013; review &amp; editing. DN: Data curation, Writing &#x2013; review &amp; editing. RS: Writing &#x2013; review &amp; editing. ZA:Writing &#x2013; review &amp; editing. LR: Writing &#x2013; review &amp; editing. X-LX: Writing &#x2013; review &amp; editing. M-ZR: Writing &#x2013; review &amp; editing. AJ: Writing &#x2013; review &amp; editing, Funding acquisition.</p>
</sec>
<sec id="s9" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research and/or publication of this article. This research received funding from UAEU in the form of Strategic Research Fund (Grant Code: 12R292) addressing the SDG 2: Zero Hunger, SDG 13: Climate Action and SDG 14: Life below Water.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>The authors would like to thank all the co-authors and corresponding authors for their valuable support and contributions during the preparation of this review. We are grateful to United Arab Emirates University for providing the financial assistance necessary for this work.</p>
</ack>
<sec id="s10" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s11" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
<p>Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.</p>
</sec>
<sec id="s12" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
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