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Review

Arthrospira platensis as Protein-Rich Source for Human Nutrition

by
Steffen Braune
1,2,*,
Conrad G. H. Jung
3,
Jan-Heiner Küpper
1,2,3 and
Friedrich Jung
1
1
Institute of Biotechnology, Molecular Cell Biology, Brandenburg University of Technology Cottbus-Senftenberg, 01968 Senftenberg, Germany
2
Faculty of Health Sciences Brandenburg, Brandenburg University of Technology Cottbus-Senftenberg, 01968 Senftenberg, Germany
3
Carbon Biotech Social Enterprise AG, 01968 Senftenberg, Germany
*
Author to whom correspondence should be addressed.
Life 2025, 15(12), 1789; https://doi.org/10.3390/life15121789
Submission received: 6 November 2024 / Revised: 28 October 2025 / Accepted: 29 October 2025 / Published: 21 November 2025
(This article belongs to the Special Issue Algae—a Step Forward in the Sustainability of Resources: 2nd Edition)
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Abstract

The continuing growth of the world’s population, combined with climate change, poses a growing challenge to ensuring food security in the 21st century. Animal protein, e.g., from beef, is a particularly rich source of protein, but there is by no means enough arable land on earth to transfer the animal meat-rich nutritional style of the early industrialized countries to the global South. A hitherto largely neglected option for the production of proteins is the cultivation of microalgae and cyanobacteria, which already have a long history of use as a human or animal food for their nutritional and environmental merits. In particular, Limnospira platensis (Spirulina and formerly Arthrospira platensis)—a filamentous cyanobacterium—is considered the “food of the future” since it is a viable source of vegan protein. In this manuscript, we review the scientific literature as well as national and intergovernmental agency statements regarding the quality and quantity of AP-based proteins. The content of AP protein is discussed in relation to other species and different cultivation conditions, as well as to traditional crops and animal husbandry. The amino acid profile and quality assessment of AP as a dietary protein are discussed. In addition, the environmental aspects of AP production are considered, and the role of AP in efforts to bridge the ‘protein gap’ to improve nutrition and food security is discussed.

1. Introduction

The world’s population continues to grow and will reach 9.7 billion people by 2050, according to United Nations projections [1]. Along with the increasing use of natural resources, climate change, and biodiversity loss, this poses a growing challenge to ensuring food security in the 21st century. To maintain energy metabolism, humans need sufficient calories in the form of carbohydrates and fats. In addition, a continuous supply of essential amino acids, vitamins, and minerals to maintain basic cell functions is needed. Malnutrition is usually a combination of insufficient intake of calorie-rich foods and, in particular, essential amino acids for protein anabolism [2].
Protein is the major structural component of all cells in the body. Proteins also function as enzymes, as transporters or receptors in membranes, as transport carriers of, e.g., lipids, and as hormones. Amino acids, the building blocks of proteins, serve as precursors for the diversity of cellular proteins to function in immune reactions, muscle contraction, and action potentials, and they are an important nitrogen source for other cellular molecules. The “Recommended Dietary Allowance” for both men and women is 0.8 g of good-quality protein/kg body weight per day [3]. The most important aspect of a protein from a nutritional point of view is its amino acid composition, but the protein’s structure may also influence its digestibility.
Animal protein from beef, pig, or chicken, for example, is a particularly rich source of the nine essential amino acids. But there is by no means enough arable land on earth to transfer the animal meat-rich nutritional style of the early industrialized countries to the global South or developing countries in principle [4,5]. Furthermore, a diet based on animal products leads to a disproportionate amount of CO2 emissions. The Intergovernmental Panel on Climate Change (IPCC) estimates that food production and distribution together are responsible for 21 to 37% of all man-made greenhouse gases [6]. New nutritional concepts are needed to ensure the long-term food security of the world’s population and, at the same time, to preserve our planet.
A hitherto largely neglected option for the large-scale production of proteins for human and animal food [7,8,9,10,11,12,13] as a renewable raw material is the cultivation of microalgae and cyanobacteria [14,15]. However, as early as 1974, the cyanobacterium Limnospira platensis (LP, often referred to as Spirulina [16] and earlier as Arthrospira platensis [17,18]) was identified as the “food of the future” at the United Nations World Food Conference.
This filamentous, helical-shaped, photoautotrophic, and alkaliphilic Limnospira platensis contains over 50 nutrients and is rich in proteins containing all essential amino acids. Figure 1 representatively shows the typical morphology of AP cultured in Zarrouk medium, as well as the major functional categories of proteins in AP [19].
It is further rich in B vitamins, iron, magnesium, potassium, and many other vitamins and minerals, as well as antioxidants [17,20]. AP has received considerable attention due to its production process in open pond systems or closed bioreactors, which requires less water and land than crops or animal-derived foods [21]. Due to these characteristics, AP is currently the most exploited industrial species, aside from Chlorella (Chlorophyta), Dunaliella, Crypthecodinium (Dinoflagellata), Haematococcus (Chlorophyta), and others [22,23,24].
In this brief review, we discuss the characteristics of AP protein with regard to its intracellular quantity and nutritional quality.

2. Protein Quantity

Many microalgae and cyanobacteria species contain high levels of protein, with AP typically comprising 40–60% protein of dry matter (see Table 1 [25,26]). This is one of the highest protein contents of microalgae species and of food in general [19,27,28], and it is one of the reasons why they have historically been used as a human food source [29]. As a bacterium, AP has the advantage over other photoautotrophic microorganisms in that it does not have a cell wall made of cellulose; therefore, it does not require chemical or physical processing to be metabolized [25].
Depending on the strain and the environmental and production conditions, the protein content can range between 17% and 80% of the dry weight (Table 1). Cultivation conditions that affect the protein content include the composition of the culture medium, the habitat, the light source and intensity, the temperature, and the method for the protein extraction and analysis, as well as the growth phase in which the biomass was harvested [37,38]. One of the main influencing factors is the illumination [39,40,41]. Light color (spectra), light intensity, and duration of illumination (light/dark cycles) can influence the quantity and quality of the biomass gained in AP cultures significantly [42].
The protein content in AP is higher than in plant or animal sources such as wheat, maize, rice, soybeans, or dried beef (55.6% [43,44,45] (Table 2). Under well-standardized conditions, protein concentrations of around 60% can be achieved in different cultivation setups (Table 3). Lower values are associated with more experimental approaches, which typically are aimed at optimizing culture conditions or understanding the influence of individual cultivation parameters (e.g., pH, temperature, CO2, etc.) on growth and AP biomass composition [36,46,47]. The same applies to studies focusing on different nutrient sources/compositions for economic or environmental reasons, e.g., carbon or nitrogen from residues of other production processes [48]. Also, approaches for the recycling of used culture media were evaluated in this matter [49,50]. The results shown in Table 2 summarize that about 40 times more protein can be produced per hectare than with plant crops and also significantly more than with meat production.

3. Protein Quality

Not only the quantity but also the quality of the protein plays a major role. From a qualitative point of view, AP proteins are complete, since all nine essential amino acids present meet the Food and Agriculture Organization of the United Nations (FAO) requirements. It has been demonstrated that AP protein is a source of essential amino acids that is comparable to other proteins, including those derived from soybeans and chicken eggs [59,60]. The free amino acids dissolved in the body fluids are only a very small proportion of the body’s total mass of amino acids. Nevertheless, they are very important for the nutritional and metabolic control of the body’s proteins.

3.1. Amino Acid Profile

Since the amino acid profile of dietary proteins is not identical to the amino acid profile of body proteins, proteins are considered to be of lower quality the further their amino acid profile deviates from the body’s own amino acid profile. Therefore, it is not the number and quantity of amino acids in dietary protein that limit protein synthesis, but rather the concentration of the amino acid that is most deficient (called the limiting amino acid). If a limiting amino acid is depleted, the body cannot produce endogenous proteins, since humans can only convert amino acids into endogenous protein to the extent that the smallest amount of an amino acid is available. As with the biological value, the chicken egg white serves as a reference value of 100 (or 1.0). As shown in Table 4, the spectrum of amino acids of AP shows that the biological value of proteins in spirulina is comparable to egg white [60,61,62].
Histidine and lysine were determined to be limiting or slightly under-represented in the case of AP, while in another study, lysine concentration was reported to be adequate [61]. This study revealed that the amino acids methionine and cysteine were the limiting amino acids. Even so, they are present at more than 80% of the ideal level defined by the FAO, calculated on the basis of egg albumin and casein.
It must be taken into account that the drying methods—especially drying on hot drums—can reduce the methionine content by some 30% compared to spray drying by evaporation [66,67,68].

3.2. Quality Scoring of Dietary Protein

In addition to the amino acid composition, different scorings were developed to assess the quality of dietary protein. The Protein Digestibility Corrected Amino Acid Score (PDCAAS, recommended by the FAO, became the industry standard in 1993 [69]. FAO has recently recommended the newer Digestible Indispensable Amino Acid Score (DIAAS) to supersede PDCAAS. The DIAAS system (ileal digestibility of the essential amino acids) was emphasized as a more accurate measure of protein absorption [70,71,72]. The difference between PDCAAS and DIAAS lies in the fact that fecal digestibility, used in PDCAAS, may be affected by microbial degradation, while true ileal digestibility, used in DIAAS, more accurately represents the amounts of amino acids absorbed in the gastrointestinal tract [68,72]. For this reason, DIAAS is promoted as the superior method and preferred over the PDCAAS [73,74].
Devi et al. reported that the DIAAS of AP is 85.2% higher than DIAAS values of certain plants (chickpeas: 56.6% or mung beans: 57.7%) [73]. Burd et al. showed that raw beef meat had clearly higher values (97%) than all plant or microalgae DIAAS [74]. However, grilled or baked (80%) beef meat showed lower values than raw beef meat or even AP.
In other countries (e.g., Canada), the Protein Efficiency Rate (PER) is additionally the official method for assessing the quality of protein [75,76]. This is the weight gain of an individual divided by the weight of proteins ingested. Measurements are usually made on growing rats. The reference proteins are lactalbumin or casein [77].
The PER value for AP determined in growing rats was estimated between 1.8 and 2.6 [78,79,80], as against a PER value for casein of 2.5. The PER values of plants are mostly lower, as for maize (1.23) or wheat (1.15) [78].

3.3. In Vivo Studies

The growth rate of rats fed AP as the sole protein source is higher than or equal to that of control animals. In addition, rats fed AP fixed greater or equal amounts of protein compared to rats supplemented with essential amino acids of equal metabolic energy. These results indicate excellent metabolic use of the amino acids in AP, which is confirmed by the levels of free amino acids found in the blood and muscle of test animals [73].
There are a few studies performed on humans. These studies tend to show results similar to those obtained in animal trials, though digestibility seems slightly lower [61,81,82].

4. Technological Challenges in Production and Processing

4.1. Cultivation Strategies

Cultivation and processing of microalgae require complex processes. Although AP and other microalgae species are characterized by relatively high growth rates and photosynthetic efficacies, upscaling to an industrial level and the associated cost structure are still challenging [83,84].
Autotrophic cultivation—e.g., in open raceway ponds—is relatively inexpensive and still represents the standard for the large-scale production of AP. However, it is reported to yield lower biomass densities in comparison to, e.g., mixotrophic approaches [85,86]. The harvesting and the energy expenditure are the dominant cost factors for autotrophic cultivation systems [87].
In principle, heterotrophic cultivation can result in high biomass and protein yields, but it is more cost-intensive [86,88]. Associated with this are higher production costs, e.g., for sterility of cultivation and processing systems, as well as a strong dependency and sensitivity of the organisms to the cultivation medium and chemical substrates (e.g., glucose) [85,86,89]. Until now, attempts to identify AP strains capable of continuously growing under dark heterotrophic culture conditions have not been successful. Thus, exposure to light plays a crucial role in reaching an industrial scale cultivation and respective growth rates [89,90].
In mixotrophic systems, high growth rates can be achieved by combining the two aforementioned cultivation techniques. This approach requires strong monitoring and regulation of factors such as the carbon source and lighting regime, which increases the complexity of the overall process [85,91,92]. Pereira et al., [91] for instance, presented a successful mixotrophic cultivation approach using dairy wastewater. In this study, an increase in the biomass concentration during 15 cultivation days was achieved by supplementing the reference Zarrouk medium with 5% buffalo mozzarella cheese whey. However, under these cultivation conditions, the protein content of the AP biomass decreased while the carbohydrate content increased [91].

4.2. Biotic and Abiotic Contamination

A major challenge associated with the cultivation of AP for the production of food/feed-related products—and thus the biomass quality and safety—is the management of biotic and abiotic contaminants.
In this context, contamination with other organisms such as toxic and non-toxic heterotrophic bacteria, insects (e.g., Ephydridae), and harmful species of micro-zooplankton (flagellates, amoebae, ciliates, and rotifers) is reported, which can occur in high densities and graze AP [93,94,95].
Particularly in open cultivation systems such as ponds, contaminants can be introduced by animals or airflow, which can survive in the culture despite the relatively high pH value. Contaminations through human handling during the processing of the culture, but particularly of the biomass, are very likely less relevant, but are discussed as well [96].
Vardaka et al. tested 31 commercial AP food supplements and identified 469 different taxonomic units [97]. Pathogenic bacteria—such as Pseudomonas, Clostridium, and Enterococcus—represent major challenges, posing health risks to humans and animals [98]. Likewise, cyanobacterial species that produce toxins such as microcystins (e.g., Microcystis) can be a serious source of contamination in the production of food/feed-related products from AP biomass [99,100].
Beyond biological contaminations, AP cultures can be affected by abiotic factors such as heavy metals and pesticides [101] since the organism can absorb these substances from its environment (biosorption). On the one hand, this is studied as a potential remediation technology (phytoremediation), and on the other, it acts as a source of contamination [93]. Particularly in areas with high industrial pollution and when low-grade agricultural fertilizers are applied as a nutrient source, even relatively low amounts can accumulate in the AP biomass with cultivation time. The same applies for cultures that come into contact with contaminated soil, e.g., in earthened ponds or when heavy metal-contaminated lining materials are used for ponds [102].
Successful production requires a comprehensive understanding of the various sources of contamination (biotic and abiotic) and the implementation of effective prevention and control measures [15]. While open pond systems are more cost-effective and accessible, closed cultivation systems combined with modern sterilization and monitoring technologies provide a promising way to produce high-quality AP biomass free from contamination [103,104,105].

4.3. Functional and Sensory Characteristics

The functional properties of AP protein biomass, such as foaming, gelling, or emulsifying ability, are crucial for the further processing of AP but depend significantly on environmental factors such as pH value, ion concentration, and temperature [106,107,108]. A high foaming capability was observed for AP protein isolates, but with low stability in many pH ranges. Gel formation is also possible, but more difficult at acidic pH and less efficient than with established protein sources such as soy.
In terms of sensory properties, taste, color, and smell represent challenges for consumer acceptance and require technological solutions for masking or processing [109]. For instance, phycocyanin colors protein isolates substantially, and bioactive peptides, as well as free amino acids in the biomass, are associated with bitter taste [110,111].
Lipid oxidation during storage and drying procedures can result in the production of volatile compounds (so-called off-flavors) that can lead to ‘fishy’ or ‘seaweed-like’ odors [112,113]. Sensory evaluation studies show that increased proportions of AP in a food product can limit acceptance due to these sensory properties. Accepted levels of added AP biomass vary among the studied products and range, e.g., for pasta and nectar, between 10 and 20% [114,115] and 1 and 2% for fruit cereal bars, yogurt/cheese products (labneh), vegan pesto, and a sourdough bakery product [116,117,118,119]. Thus, further technological solutions for masking undesired sensory properties and processing are required for improving consumer acceptance.

5. Environmental Considerations

The production of AP proteins offers environmental advantages over conventional protein sources. This approach to protein production is more resource-efficient, requiring less water and land compared to traditional sources like soy or cattle farming. AP can be grown on barren land, making use of areas not suitable for conventional food production.
From an environmental standpoint, AP biomass production stands out for its sustainability. The process does not involve genetic manipulation, pesticides, or herbicides. The alkaline pH environment in the cultivation medium can reduce undesired organisms (co-flora), reducing the need for chemical intervention [120,121]. Essential nutrients can be efficiently incorporated and monitored without the risk of over-fertilization. This nutrient management prevents groundwater contamination, a significant advantage over conventional farming methods. In arid regions, the ability to use sea or brackish water for production further contributes to the conservation of precious groundwater resources [122].
AP production can contribute to climate protection because AP, like other microalgae and cyanobacteria, outperforms many photosynthetic plants in carbon dioxide fixation per unit of biomass. This is due to the significant investments plants make in non-photosynthetic structures, such as roots, stems, and supportive tissues, as well as their lower overall conversion efficiencies [123,124,125,126,127]. Thus, AP-based protein production can help mitigate the emission of greenhouse gases like carbon dioxide and methane, which are typically associated with animal husbandry.
A further advantage of AP protein production lies in its potential impact on land use and reforestation. By partially or completely replacing animal proteins with AP-based products, vast areas currently dedicated to animal feed or pasture could be freed up. These liberated lands could be repurposed for planting new forests, which would serve as additional carbon sinks that actively sequester atmospheric carbon. Furthermore, the trees grown on these reclaimed areas could eventually be harvested for use as construction timber, providing a long-term carbon storage solution.
Life cycle assessment studies indicate that optimizing production and harvesting processes is important for achieving an advantage in terms of the economic and environmental footprint/costs per kg of microalgae-based protein compared to conventional protein sources [86]. This is particularly discussed for regions with relatively low levels of sunshine and average temperatures, such as Central Europe, due to the higher energy consumption/costs for the cultivation and processing of biomass.
The reduction of, e.g., freshwater requirements, utilization of waste heat, and agricultural waste streams has the potential to substantially improve the economic and environmental footprint of microalgae production [128,129].

6. Regulatory Aspects

From a regulatory perspective, AP is generally regarded as safe (GRAS) by the European Food Safety Authority (EFSA) [130] and by the U.S. Food and Drug Administration (FDA). The latter accepted a GRAS notification (GRN No. 417) for the use of dried AP biomass as an ingredient in various foods and drinks without objection in 2012 [131]. Thus, these microalgae have the potential to serve as a substitute for vegetable protein in food products, and their efficacy may even surpass that of soy.
One of the major challenges in the production of AP-derived protein for food and feed applications is the dependency of protein and nutrient composition on environmental conditions and cultivation location. This complicates standardization for the food industry. As described in a previous paragraph, AP can accumulate heavy metals, iodine, or toxic metabolites, which necessitates additional purification and control steps (see Section 4.2).
Despite its GRAS status, the allergenic potential has not been conclusively clarified, as discussed by Gromek et al. [132]. Strong allergic reactions are very rare and mainly reported for predisposed, atopic individuals [133,134]. The French Agency for Food, Environmental and Occupational Health & Safety reported on the “risks associated with the consumption of food supplements containing spirulina” in 2017 [135]. An expert committee concluded that “Apart from the risk of contamination, spirulina does not seem to present a health risk at low doses (up to several grams per day)”. As described above, contamination of cultivation systems with other bacteria (e.g., cyanotoxin-producing species) and heavy metals is possible and can exceed the limits laid by the regulatory authorities. These parameters must be taken into account in the discussion about and the testing of the allergenic potential of AP. The current research suggests that AP biomass has an allergenic potential with minimal clinical relevance in healthy individuals [132,136,137]. It should be noted that beyond fundamental animal trials from the 1990s (see: [132]), recent in vitro and in vivo studies report on the anti-allergic potential of AP, e.g., for the treatment of allergic rhinitis [138,139,140]. These were, however, critically reviewed, and more randomized clinical trials in accordance with the CONSORT standardization have been suggested [141].
According to the regulatory situation, the application of AP protein as a food product is uncritical due to its legal acceptance in the US, the EU, and Asia. However, this applies to AP biomass that has not undergone any significant changes in processing or composition. New technologies—such as protein isolates—or use in novel foods may require a complex approval procedure, e.g., according to the “Novel food” regulation in the EU [142].

7. Conclusions

Arthrospira is, with up to 80% dry mass, very rich in protein. The protein content is higher in comparison to unicellular algae and most other cyanobacteria or to beef, pig, or chicken meat. Without requiring large areas of arable land and less water compared to conventional crops/livestock farming, AP thus offers the possibility to close the “protein gap”. In addition, comprehensive analyses and nutritional studies have demonstrated that AP protein is of high quality and as good as animal proteins and even better than conventional Leguminosae-based protein.
However, the quality of proteins can vary dramatically, depending on digestibility and the availability of essential amino acids [143]. In contrast to vegetable proteins—which often contain only a small amount of one or more EAAs or lack them completely—AP is a considerable source of protein whose EAA composition meets FAO requirements in comparison to egg white. AP is well digestible. The thin cell wall of AP consists of peptidoglycan and polysaccharide layers within the typical organization of Gram-negative bacteria. This is in contrast to eukaryotic microalgae such as Chlorella, in which the cell wall contains an indigestible cellulose scaffold.
Despite the remaining technological challenges, a shift towards AP-based protein production represents a holistic approach to addressing multiple environmental challenges simultaneously. It can offer a sustainable, efficient, and environmentally friendly alternative to traditional protein sources, with far-reaching benefits for resource management, climate protection, and land use optimization.

Author Contributions

C.G.H.J., S.B. and F.J., conceptualization, and writing—original draft preparation; J.-H.K., and F.J., resources and writing—review and editing, S.B. and J.-H.K., funding aquisition. All authors have read and agreed to the published version of the manuscript.

Funding

This work was financially supported by the state of Brandenburg, Germany (Program: Strengthening technological and application-oriented research in research networks of scientific institutions in the state of Brandenburg, StaF-Verbund), and the European Regional Development Fund (EFRE).

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Data sharing is not applicable.

Conflicts of Interest

The authors Jan-Heiner Küpper and Conrad G. H. Jung were employed by the company Carbon Biotech Social Enterprise AG. All 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.

References

  1. United Nations (Ed.) Global Population Growth and Sustainable Development; United Nations publication; United Nations: New York, NY, USA, 2021; ISBN 978-92-1-148350-5. [Google Scholar]
  2. Semba, R.D. The Rise and Fall of Protein Malnutrition in Global Health. Ann. Nutr. Metab. 2016, 69, 79–88. [Google Scholar] [CrossRef]
  3. Wu, G. Dietary Protein Intake and Human Health. Food Funct. 2016, 7, 1251–1265. [Google Scholar] [CrossRef]
  4. Rizvi, S.; Pagnutti, C.; Fraser, E.; Bauch, C.T.; Anand, M. Global Land Use Implications of Dietary Trends. PLoS ONE 2018, 13, e0200781. [Google Scholar] [CrossRef]
  5. Lan, Y.; Xu, B.; Huan, Y.; Guo, J.; Liu, X.; Han, J.; Li, K. Food Security and Land Use under Sustainable Development Goals: Insights from Food Supply to Demand Side and Limited Arable Land in China. Foods 2023, 12, 4168. [Google Scholar] [CrossRef]
  6. Intergovernmental Panel On Climate Change. Climate Change and Land: IPCC Special Report on Climate Change, Desertification, Land Degradation, Sustainable Land Management, Food Security, and Greenhouse Gas Fluxes in Terrestrial Ecosystems, 1st ed.; Cambridge University Press: Cambridge, UK, 2022; ISBN 978-1-009-15798-8. [Google Scholar]
  7. Çelekli, A.; Özbal, B.; Bozkurt, H. Challenges in Functional Food Products with the Incorporation of Some Microalgae. Foods 2024, 13, 725. [Google Scholar] [CrossRef] [PubMed]
  8. Spínola, M.P.; Costa, M.M.; Prates, J.A.M. Effect of Cumulative Spirulina Intake on Broiler Meat Quality, Nutritional and Health-Related Attributes. Foods 2024, 13, 799. [Google Scholar] [CrossRef] [PubMed]
  9. Hernández-López, I.; Alamprese, C.; Cappa, C.; Prieto-Santiago, V.; Abadias, M.; Aguiló-Aguayo, I. Effect of Spirulina in Bread Formulated with Wheat Flours of Different Alveograph Strength. Foods 2023, 12, 3724. [Google Scholar] [CrossRef]
  10. Ruszkowska, M.; Tańska, M.; Miedzianka, J.; Kowalczewski, P.Ł. Field Cricket (Gryllus Bimaculatus) and Spirulina (Arthrospira Platensis) Powders as Environmentally Friendly Protein Enrichment Ingredients in Corn Snacks. Foods 2024, 13, 2390. [Google Scholar] [CrossRef]
  11. Lambiase, C.; Braghieri, A.; Barone, C.M.A.; Di Francia, A.; Pacelli, C.; Serrapica, F.; Lorenzo, J.M.; De Rosa, G. Use of Cyanobacterium Spirulina (Arthrospira Platensis) in Buffalo Feeding: Effect on Mozzarella Cheese Quality. Foods 2023, 12, 4095. [Google Scholar] [CrossRef]
  12. Prandi, B.; Boukid, F.; Van De Walle, S.; Cutroneo, S.; Comaposada, J.; Van Royen, G.; Sforza, S.; Tedeschi, T.; Castellari, M. Protein Quality and Protein Digestibility of Vegetable Creams Reformulated with Microalgae Inclusion. Foods 2023, 12, 2395. [Google Scholar] [CrossRef] [PubMed]
  13. Verni, M.; Demarinis, C.; Rizzello, C.G.; Pontonio, E. Bioprocessing to Preserve and Improve Microalgae Nutritional and Functional Potential: Novel Insight and Perspectives. Foods 2023, 12, 983. [Google Scholar] [CrossRef]
  14. Rotter, A.; Giannakourou, A.; Argente García, J.E.; Quero, G.M.; Auregan, C.; Triantaphyllidis, G.; Venetsanopoulou, A.; De Carolis, R.; Efstratiou, C.; Aboal, M.; et al. Identification of Marine Biotechnology Value Chains with High Potential in the Northern Mediterranean Region. Mar. Drugs 2023, 21, 416. [Google Scholar] [CrossRef]
  15. Novoveská, L.; Nielsen, S.L.; Eroldoğan, O.T.; Haznedaroglu, B.Z.; Rinkevich, B.; Fazi, S.; Robbens, J.; Vasquez, M.; Einarsson, H. Overview and Challenges of Large-Scale Cultivation of Photosynthetic Microalgae and Cyanobacteria. Mar. Drugs 2023, 21, 445. [Google Scholar] [CrossRef] [PubMed]
  16. Fais, G.; Manca, A.; Bolognesi, F.; Borselli, M.; Concas, A.; Busutti, M.; Broggi, G.; Sanna, P.; Castillo-Aleman, Y.M.; Rivero-Jiménez, R.A.; et al. Wide Range Applications of Spirulina: From Earth to Space Missions. Mar. Drugs 2022, 20, 299. [Google Scholar] [CrossRef] [PubMed]
  17. Sinetova, M.A.; Kupriyanova, E.V.; Los, D.A. Spirulina/Arthrospira/Limnospira—Three Names of the Single Organism. Foods 2024, 13, 2762. [Google Scholar] [CrossRef]
  18. Roussel, T.; Halary, S.; Duval, C.; Piquet, B.; Cadoret, J.-P.; Vernès, L.; Bernard, C.; Marie, B. Monospecific Renaming within the Cyanobacterial Genus Limnospira (Spirulina) and Consequences for Food Authorization. J. Appl. Microbiol. 2023, 134, lxad159. [Google Scholar] [CrossRef]
  19. Pan-utai, W.; Iamtham, S.; Roytrakul, S.; Settachaimongkon, S.; Wattanasiritham, L.S.; Boonbumrung, S.; Mookdasanit, J.; Sithtisarn, S. Arthrospira Platensis Mutagenesis for Protein and C-Phycocyanin Improvement and Proteomics Approaches. Life 2022, 12, 911. [Google Scholar] [CrossRef]
  20. Jung, F.; Krüger-Genge, A.; Waldeck, P.; Küpper, J.-H. Spirulina platensis, a Super Food? J. Cell. Biotechnol. 2019, 5, 43–54. [Google Scholar] [CrossRef]
  21. Bleakley, S.; Hayes, M. Algal Proteins: Extraction, Application, and Challenges Concerning Production. Foods 2017, 6, 33. [Google Scholar] [CrossRef]
  22. Ismail, I.; Hwang, Y.-H.; Joo, S.-T. Meat Analog as Future Food: A Review. J. Anim. Sci. Technol. 2020, 62, 111–120. [Google Scholar] [CrossRef]
  23. Barbosa, M.; Inácio, L.G.; Afonso, C.; Maranhão, P. The Microalga Dunaliella and Its Applications: A Review. Appl. Phycol. 2023, 4, 99–120. [Google Scholar] [CrossRef]
  24. Lu, X.; Zhao, W.; Wang, J.; He, Y.; Yang, S.; Sun, H. A Comprehensive Review on the Heterotrophic Production of Bioactive Compounds by Microalgae. World J. Microbiol. Biotechnol. 2024, 40, 210. [Google Scholar] [CrossRef]
  25. Manirafasha, E.; Murwanashyaka, T.; Ndikubwimana, T.; Rashid Ahmed, N.; Liu, J.; Lu, Y.; Zeng, X.; Ling, X.; Jing, K. Enhancement of Cell Growth and Phycocyanin Production in Arthrospira (Spirulina) Platensis by Metabolic Stress and Nitrate Fed-Batch. Bioresour. Technol. 2018, 255, 293–301. [Google Scholar] [CrossRef]
  26. Wang, Y.; Tibbetts, S.; McGinn, P. Microalgae as Sources of High-Quality Protein for Human Food and Protein Supplements. Foods 2021, 10, 3002. [Google Scholar] [CrossRef]
  27. Nosratimovafagh, A.; Fereidouni, A.E.; Krujatz, F. Modeling and Optimizing the Effect of Light Color, Sodium Chloride and Glucose Concentration on Biomass Production and the Quality of Arthrospira Platensis Using Response Surface Methodology (RSM). Life 2022, 12, 371. [Google Scholar] [CrossRef] [PubMed]
  28. Gentscheva, G.; Nikolova, K.; Panayotova, V.; Peycheva, K.; Makedonski, L.; Slavov, P.; Radusheva, P.; Petrova, P.; Yotkovska, I. Application of Arthrospira Platensis for Medicinal Purposes and the Food Industry: A Review of the Literature. Life 2023, 13, 845. [Google Scholar] [CrossRef]
  29. Ciferri, O. Spirulina, the Edible Microorganism. Microbiol. Rev. 1983, 47, 551–578. [Google Scholar] [CrossRef] [PubMed]
  30. Minkova, K.M.; Tchernov, A.A.; Tchorbadjieva, M.I.; Fournadjieva, S.T.; Antova, R.E.; Busheva, M.C. Purification of C-Phycocyanin from Spirulina (Arthrospira) Fusiformis. J. Biotechnol. 2003, 102, 55–59. [Google Scholar] [CrossRef] [PubMed]
  31. Belal, E.B.; Khalafalla, M.M.E.; El-Hais, A.M.A. Use of Spirulina (Arthrospira Fusiformis) for Promoting Growth of Nile Tilapia Fingerlings. Afr. J. Microbiol. Res. 2012, 6, 6423–6431. [Google Scholar] [CrossRef]
  32. Milledge, J.J. Commercial Application of Microalgae Other than as Biofuels: A Brief Review. Rev. Environ. Sci. Biotechnol. 2011, 10, 31–41. [Google Scholar] [CrossRef]
  33. Becker, E.W. Micro-Algae as a Source of Protein. Biotechnol. Adv. 2007, 25, 207–210. [Google Scholar] [CrossRef]
  34. Aaronson, S.; Berner, T.; Dubinsky, Z. Microalgae as a Source of Chemicals and Natural Products. In Algae Biomass: Production and Use; Elsevier/North-Holland Biomedical Press: Amsterdam, The Netherland, 1980; pp. 595–601. [Google Scholar]
  35. Ferreira, L.S.; Rodrigues, M.S.; Converti, A.; Sato, S.; Carvalho, J.C.M. Arthrospira (Spirulina) Platensis Cultivation in Tubular Photobioreactor: Use of No-Cost CO2 from Ethanol Fermentation. Appl. Energy 2012, 92, 379–385. [Google Scholar] [CrossRef]
  36. Jung, C.G.H.; Nghinaunye, T.; Waldeck, P.; Braune, S.; Petrick, I.; Küpper, J.-H.; Jung, F. Decarbonization of Arthrospira Platensis Production by Using Atmospheric CO2 as an Exclusive Carbon Source: Proof of Principle. Int. J. Environ. Sci. Technol. 2024, 21, 4635–4644. [Google Scholar] [CrossRef]
  37. Jung, F.; Jung, C.G.H.; Krüger-Genge, A.; Waldeck, P.; Küpper, J.-H. Factors Influencing the Growth of Spirulina Platensis in Closed Photobioreactors under CO2—O2 Conversion. J. Cell. Biotechnol. 2019, 5, 125–134. [Google Scholar] [CrossRef]
  38. Nghinaunye, T.; Waldeck, P.; Jung, C.G.H.; Küpper, J.-H.; Jung, F.; Braune, S. Response of Arthrospira Platensis to Different Temperatures Regarding Growth and Biochemical Composition. Clin. Hemorheol. Microcirc. 2024, 86, 205–211. [Google Scholar] [CrossRef]
  39. Tredici, M.R.; Zittelli, G.C. Efficiency of Sunlight Utilization: Tubular versus Flat Photobioreactors. Biotechnol. Bioeng. 1998, 57, 187–197. [Google Scholar] [CrossRef]
  40. Niangoran, N.U.F.; Buso, D.; Zissis, G.; Prudhomme, T. Influence of Light Intensity and Photoperiod on Energy Efficiency of Biomass and Pigment Production of Spirulina (Arthrospira Platensis). OCL 2021, 28, 37. [Google Scholar] [CrossRef]
  41. Bezerra, R.P.; Matsudo, M.C.; Converti, A.; Sato, S.; De Carvalho, J.C.M. Influence of Ammonium Chloride Feeding Time and Light Intensity on the Cultivation of Spirulina (Arthrospira) Platensis. Biotechnol. Bioeng. 2008, 100, 297–305. [Google Scholar] [CrossRef]
  42. Jung, C.H.G.; Waldeck, P.; Sykora, S.; Braune, S.; Petrick, I.; Küpper, J.-H.; Jung, F. Influence of Different Light-Emitting Diode Colors on Growth and Phycobiliprotein Generation of Arthrospira Platensis. Life 2022, 12, 895. [Google Scholar] [CrossRef]
  43. Becker, E.W. Microalgae: Biotechnology and Microbiology; Cambridge Studies in Biotechnology; Cambridge University Press: Cambridge, UK; New York, NY, USA; Melbourne, Australia, 1994; ISBN 978-0-521-35020-4. [Google Scholar]
  44. Saito, Y.; Itakura, K.; Kuramoto, M.; Kaho, T.; Ohtake, N.; Hasegawa, H.; Suzuki, T.; Kondo, N. Prediction of Protein and Oil Contents in Soybeans Using Fluorescence Excitation Emission Matrix. Food Chem. 2021, 365, 130403. [Google Scholar] [CrossRef]
  45. Adeyi, A.M.; Osunde, Z.D.; Ogunjirin, O.A.; Awu, J.I.; Ibrahim, A. Effect of Drying on Nutritional Quality of Dried Meat (Kilishi). J. Agric. Eng. Technol. (JAET) 2015, 23, 14–34. Available online: https://jaet.niae.net/index.php/jaet/issue/view/8 (accessed on 28 October 2025).
  46. Arahou, F.; Hassikou, R.; Arahou, M.; Rhazi, L.; Wahby, I. Influence of Culture Conditions on Arthrospira Platensis Growth and Valorization of Biomass as Input for Sustainable Agriculture. Aquac. Int. 2021, 29, 2009–2020. [Google Scholar] [CrossRef]
  47. Fagiri, Y.M.A.; Salleh, A.; El-Nagerabi, S.A.F. Impact of Physico-Chemical Parameters on the Physiological Growth of Arthrospira (Spirulina Platensis) Exogenous Strain UTEXLB2340. Afr. J. Biotechnol. 2013, 12, 5458–5465. [Google Scholar] [CrossRef]
  48. Gómez, C.; Guzmán-Carrasco, A.; Lafarga, T.; Acién-Fernández, F.G. Optimization of a New Culture Medium for the Large-Scale Production of Protein-Rich Arthrospira Platensis (Oscillatoriales, Cyanophyceae). J. Phycol. 2021, 57, 636–644. [Google Scholar] [CrossRef] [PubMed]
  49. Mejia-da-Silva, L.D.C.; Matsudo, M.C.; Morocho-Jacome, A.L.; de Carvalho, J.C.M. Application of Physicochemical Treatment Allows Reutilization of Arthrospira Platensis Exhausted Medium. Appl. Biochem. Biotechnol. 2018, 186, 40–53. [Google Scholar] [CrossRef]
  50. Morocho-Jácome, A.L.; Sato, S.; de Carvalho, J.C.M. Ferric Sulfate Coagulation and Powdered Activated Carbon Adsorption as Simultaneous Treatment to Reuse the Medium in Arthrospira Platensis Cultivation. J. Chem. Technol. Biotechnol. 2016, 91, 901–910. [Google Scholar] [CrossRef]
  51. Coca, M.; Barrocal, V.M.; Lucas, S.; González-Benito, G.; García-Cubero, M.T. Protein Production in Spirulina Platensis Biomass Using Beet Vinasse-Supplemented Culture Media. Food Bioprod. Process. 2015, 94, 306–312. [Google Scholar] [CrossRef]
  52. Avila-Leon, I.; Chuei Matsudo, M.; Sato, S.; de Carvalho, J.C.M. Arthrospira Platensis Biomass with High Protein Content Cultivated in Continuous Process Using Urea as Nitrogen Source. J. Appl. Microbiol. 2012, 112, 1086–1094. [Google Scholar] [CrossRef]
  53. Colla, L.M.; Oliveira Reinehr, C.; Reichert, C.; Costa, J.A.V. Production of Biomass and Nutraceutical Compounds by Spirulina Platensis under Different Temperature and Nitrogen Regimes. Bioresour. Technol. 2007, 98, 1489–1493. [Google Scholar] [CrossRef]
  54. Tokuşoglu, Ö.; üUnal, M.K. Biomass Nutrient Profiles of Three Microalgae: Spirulina Platensis, Chlorella Vulgaris, and Isochrisis Galbana. J. Food Sci. 2003, 68, 1144–1148. [Google Scholar] [CrossRef]
  55. Matsudo, M.C.; Bezerra, R.P.; Sato, S.; Perego, P.; Converti, A.; Carvalho, J.C.M. Repeated Fed-Batch Cultivation of Arthrospira (Spirulina) Platensis Using Urea as Nitrogen Source. Biochem. Eng. J. 2009, 43, 52–57. [Google Scholar] [CrossRef]
  56. Phang, S.M.; Miah, M.S.; Yeoh, B.G.; Hashim, M.A. Spirulina Cultivation in Digested Sago Starch Factory Wastewater. J. Appl. Phycol. 2000, 12, 395–400. [Google Scholar] [CrossRef]
  57. Ranganathan, P.; Amal, J.C.; Savithri, S.; Haridas, A. Experimental and Modelling of Arthrospira Platensis Cultivation in Open Raceway Ponds. Bioresour. Technol. 2017, 242, 197–205. [Google Scholar] [CrossRef]
  58. Dewi, E.N.; Amalia, U.; Mel, M. The Effect of Different Treatments to the Amino Acid Contents of Micro Algae Spirulina sp. Aquat. Procedia 2016, 7, 59–65. [Google Scholar] [CrossRef]
  59. Fleurence, J. Seaweed Proteins: Biochemical, Nutritional Aspects and Potential Uses. Trends Food Sci. Technol. 1999, 10, 25–28. [Google Scholar] [CrossRef]
  60. World Health Organization (Ed.) Protein Quality Evaluation: Report of the Joint FAO/WHO Expert Consultation, Bethesda, MD, USA, 4–8 December 1989; FAO food and nutrition paper; Food and Agriculture Organization of the United Nations: Rome, Italy, 1991; ISBN 978-92-5-103097-4. [Google Scholar]
  61. U.S. Department of Agriculture. Agricultural Research Service Seaweed, Spirulina, Dried. Available online: https://fdc.nal.usda.gov/food-details/170495/nutrients (accessed on 10 March 2025).
  62. Panel on Macronutrients; Panel on the Definition of Dietary Fiber; Subcommittee on Upper Reference Levels of Nutrients; Subcommittee on Interpretation and Uses of Dietary Reference Intakes; Standing Committee on the Scientific Evaluation of Dietary Reference Intakes; Food and Nutrition Board; Institute of Medicine. Chapter 10: Protein and Amino Acids. In Dietary Reference Intakes for Energy, Carbohydrate, Fiber, Fat, Fatty Acids, Cholesterol, Protein, and Amino Acids; National Academies Press: Washington, DC, USA, 2005; p. 10490. ISBN 978-0-309-08525-0. [Google Scholar]
  63. Dietary Protein Quality Evaluation in Human Nutrition: Report of an FAO Expert Consultation [on Protein Quality Evaluation in Human Nutrition], 31 March–2 April 2011, Auckland, New Zealand; FAO food and nutrition paper; Food and Agriculture Organization of the United Nations: Rome, Italy, 2013; ISBN 978-92-5-107417-6.
  64. Brestenský, M.; Nitrayová, S.; Patráš, P.; Nitray, J. Dietary Requirements for Proteins and Amino Acids in Human Nutrition. Curr. Nutr. Food Sci. 2019, 15, 638–645. [Google Scholar] [CrossRef]
  65. U.S. Department of Agriculture. Agricultural Research Service Eggs, Grade A, Large, Egg White. Available online: https://fdc.nal.usda.gov/food-details/747997/nutrients (accessed on 10 March 2025).
  66. Mróz, M.; Parchem, K.; Jóźwik, J.; Domingues, M.R.; Kusznierewicz, B. The Impact of Different Drying Methods on the Metabolomic and Lipidomic Profiles of Arthrospira Platensis. Molecules 2024, 29, 1747. [Google Scholar] [CrossRef] [PubMed]
  67. Kim, G.M.; Kim, Y.-K. Drying Techniques of Microalgal Biomass: A Review. Appl. Chem. Eng. 2022, 33, 145–150. [Google Scholar] [CrossRef]
  68. Vermorel, M.; Toullec, G.; Dumond, D.; Pion, R. Valeur protĕique et energĕtique des algues bleues spirulines supplĕmentĕes en acides aminĕs: Utizisation digestive et mĕtabolique par le rat en croissance. Ann. Nutr. Aliment. 1975, 29, 535–552. [Google Scholar]
  69. Schaafsma, G. The Protein Digestibility-Corrected Amino Acid Score. J. Nutr. 2000, 130, 1865S–1867S. [Google Scholar] [CrossRef]
  70. Hoffman, J.R.; Falvo, M.J. Protein—Which Is Best? J. Sports Sci. Med. 2004, 3, 118–130. [Google Scholar]
  71. Jiménez-Munoz, L.M.; Tavares, G.M.; Corredig, M. Design Future Foods Using Plant Protein Blends for Best Nutritional and Technological Functionality. Trends Food Sci. Technol. 2021, 113, 139–150. [Google Scholar] [CrossRef]
  72. Moughan, P.J.; Lim, W.X.J. Digestible Indispensable Amino Acid Score (DIAAS): 10 Years On. Front. Nutr. 2024, 11, 1389719. [Google Scholar] [CrossRef]
  73. Rutherfurd, S.M.; Fanning, A.C.; Miller, B.J.; Moughan, P.J. Protein Digestibility-Corrected Amino Acid Scores and Digestible Indispensable Amino Acid Scores Differentially Describe Protein Quality in Growing Male Rats. J. Nutr. 2015, 145, 372–379. [Google Scholar] [CrossRef] [PubMed]
  74. Burd, N.A.; McKenna, C.F.; Salvador, A.F.; Paulussen, K.J.M.; Moore, D.R. Dietary Protein Quantity, Quality, and Exercise Are Key to Healthy Living: A Muscle-Centric Perspective Across the Lifespan. Front. Nutr. 2019, 6, 83. [Google Scholar] [CrossRef]
  75. Lamb, M.W.; Harden, M.L. Protein as a Source of Amino Acids. In The Meaning of Human Nutrition; Elsevier: Amsterdam, The Netherlands, 1973; pp. 153–191. ISBN 978-0-08-017079-4. [Google Scholar]
  76. Mansilla, W.D.; Marinangeli, C.P.F.; Cargo-Froom, C.; Franczyk, A.; House, J.D.; Elango, R.; Columbus, D.A.; Kiarie, E.; Rogers, M.; Shoveller, A.K. Comparison of Methodologies Used to Define the Protein Quality of Human Foods and Support Regulatory Claims. Appl. Physiol. Nutr. Metab. 2020, 45, 917–926. [Google Scholar] [CrossRef] [PubMed]
  77. Joint FAO/WHO Ad Hoc Expert Committee on Energy and Protein Requirements; World Health Organization; Food and Agriculture Organization of the United Nations. Energy and Protein Requirements: Report of a Joint FAO/WHO ad hoc Expert Committee, Rome, Italy, 22 March–2 April 1971. 1973. Available online: https://iris.who.int/items/7243fc9d-a5df-44d8-80fb-b732c34e1e54 (accessed on 28 October 2025).
  78. Devi, M.A.; Venkataraman, L.V. Supplementary Value of the Proteins of Blue Green Algae Spirulina Platensis to Rice and Wheat Proteins. Nutr. Rep. Int. 1983, 28, 1029–1035. [Google Scholar]
  79. Soni, R.A.; Sudhakar, K.; Rana, R.S.; Baredar, P. Food Supplements Formulated with Spirulina. In Algae; Mandotra, S.K., Upadhyay, A.K., Ahluwalia, A.S., Eds.; Springer: Singapore, 2021; pp. 201–226. ISBN 978-981-15-7517-4. [Google Scholar]
  80. Furst, P.T. Spirulina a Nutricious Alga, Once a Staple of Aztec Diet, Could Feed Many of the World Hungry People. Hum. Nat. 1978, 3, 60–65. [Google Scholar]
  81. Devi, S.; Varkey, A.; Sheshshayee, M.S.; Preston, T.; Kurpad, A.V. Measurement of Protein Digestibility in Humans by a Dual-Tracer Method. Am. J. Clin. Nutr. 2018, 107, 984–991. [Google Scholar] [CrossRef]
  82. De La Jara, A.; Ruano-Rodriguez, C.; Polifrone, M.; Assunçao, P.; Brito-Casillas, Y.; Wägner, A.M.; Serra-Majem, L. Impact of Dietary Arthrospira (Spirulina) Biomass Consumption on Human Health: Main Health Targets and Systematic Review. J. Appl. Phycol. 2018, 30, 2403–2423. [Google Scholar] [CrossRef]
  83. Yang, S.; Xu, H.; Chen, J.-H.; Liu, B.; Cheng, K.-W. Microalgae as a Sustainable Protein Source: Key Issues Related to Their Production, Application, and the Way Forward. Food Bioprocess Technol. 2024, 17, 1–33. [Google Scholar] [CrossRef]
  84. Safdar, W.; Qazi, A.S.; Ahmed, S.; Tariq, M.R.; Ahmed, H. Nonconventional and Novel Strategies to Produce Spirulina Biomass. In Pharmaceutical and Nutraceutical Potential of Cyanobacteria; Mehmood, M.A., Verma, P., Shah, M.P., Betenbaugh, M.J., Eds.; Springer International Publishing: Cham, Switzerland, 2024; pp. 161–194. ISBN 978-3-031-45522-3. [Google Scholar]
  85. Chojnacka, K.; Zielińska, A. Evaluation of Growth Yield of Spirulina (Arthrospira) Sp. in Photoautotrophic, Heterotrophic and Mixotrophic Cultures. World J. Microbiol. Biotechnol. 2012, 28, 437–445. [Google Scholar] [CrossRef]
  86. Smetana, S.; Sandmann, M.; Rohn, S.; Pleissner, D.; Heinz, V. Autotrophic and Heterotrophic Microalgae and Cyanobacteria Cultivation for Food and Feed: Life Cycle Assessment. Bioresour. Technol. 2017, 245, 162–170. [Google Scholar] [CrossRef]
  87. Tan, J.S.; Lee, S.Y.; Chew, K.W.; Lam, M.K.; Lim, J.W.; Ho, S.-H.; Show, P.L. A Review on Microalgae Cultivation and Harvesting, and Their Biomass Extraction Processing Using Ionic Liquids. Bioengineered 2020, 11, 116–129. [Google Scholar] [CrossRef]
  88. Nagarajan, D.; Lee, D.-J.; Chang, J. Heterotrophic Microalgal Cultivation. In Bioreactors for Microbial Biomass and Energy Conversion; Liao, Q., Chang, J., Herrmann, C., Xia, A., Eds.; Green Energy and Technology; Springer: Singapore, 2018; pp. 117–160. ISBN 978-981-10-7676-3. [Google Scholar]
  89. Mühling, M.; Belay, A.; Whitton, B.A. Screening Arthrospira (Spirulina) Strains for Heterotrophy. J. Appl. Phycol. 2005, 17, 129–135. [Google Scholar] [CrossRef]
  90. Marquez, F.J.; Sasaki, K.; Kakizono, T.; Nishio, N.; Nagai, S. Growth Characteristics of Spirulina Platensis in Mixotrophic and Heterotrophic Conditions. J. Ferment. Bioeng. 1993, 76, 408–410. [Google Scholar] [CrossRef]
  91. Pereira, M.I.B.; Chagas, B.M.E.; Sassi, R.; Medeiros, G.F.; Aguiar, E.M.; Borba, L.H.F.; Silva, E.P.E.; Neto, J.C.A.; Rangel, A.H.N. Mixotrophic Cultivation of Spirulina Platensis in Dairy Wastewater: Effects on the Production of Biomass, Biochemical Composition and Antioxidant Capacity. PLoS ONE 2019, 14, e0224294. [Google Scholar] [CrossRef] [PubMed]
  92. Russo, N.P.; Ballotta, M.; Usai, L.; Torre, S.; Giordano, M.; Fais, G.; Casula, M.; Dessì, D.; Nieri, P.; Damergi, E.; et al. Mixotrophic Cultivation of Arthrospira Platensis (Spirulina) under Salt Stress: Effect on Biomass Composition, FAME Profile and Phycocyanin Content. Mar. Drugs 2024, 22, 381. [Google Scholar] [CrossRef]
  93. Amha Belay Mass Culture of Spirulina Outdoors—The Earthrise Farms Experience. In Spirulina platensis (Arthrospira): Physiology, Cell-Biology and Biotechnology; CRC Press: Boca Raton, FL, USA; Taylor and Francis: Abingdon, UK, 1997; pp. 131–158. ISBN 978-0-429-07994-8.
  94. Wang, L.; Yuan, D.; Li, Y.; Ma, M.; Hu, Q.; Gong, Y. Contaminating Microzooplankton in Outdoor Microalgal Mass Culture Systems: An Ecological Viewpoint. Algal Res. 2016, 20, 258–266. [Google Scholar] [CrossRef]
  95. Yuan, D.; Zhan, X.; Wang, M.; Wang, X.; Feng, W.; Gong, Y.; Hu, Q. Biodiversity and Distribution of Microzooplankton in Spirulina (Arthrospira) Platensis Mass Cultures throughout China. Algal Res. 2018, 30, 38–49. [Google Scholar] [CrossRef]
  96. AlFadhly, N.K.Z.; Alhelfi, N.; Altemimi, A.B.; Verma, D.K.; Cacciola, F. Tendencies Affecting the Growth and Cultivation of Genus Spirulina: An Investigative Review on Current Trends. Plants 2022, 11, 3063. [Google Scholar] [CrossRef]
  97. Vardaka, E.; Kormas, K.A.; Katsiapi, M.; Genitsaris, S.; Moustaka-Gouni, M. Molecular Diversity of Bacteria in Commercially Available “Spirulina” Food Supplements. PeerJ 2016, 4, e1610. [Google Scholar] [CrossRef]
  98. Furmaniak, M.A.; Misztak, A.E.; Franczuk, M.D.; Wilmotte, A.; Waleron, M.; Waleron, K.F. Edible Cyanobacterial Genus Arthrospira: Actual State of the Art in Cultivation Methods, Genetics, and Application in Medicine. Front. Microbiol. 2017, 8, 2541. [Google Scholar] [CrossRef]
  99. Rhoades, J.; Fotiadou, S.; Paschalidou, G.; Papadimitriou, T.; Ordóñez, A.Á.; Kormas, K.; Vardaka, E.; Likotrafiti, E. Microbiota and Cyanotoxin Content of Retail Spirulina Supplements and Spirulina Supplemented Foods. Microorganisms 2023, 11, 1175. [Google Scholar] [CrossRef]
  100. Pinchart, P.-E.; Leruste, A.; Pasqualini, V.; Mastroleo, F. Microcystins and Cyanobacterial Contaminants in the French Small-Scale Productions of Spirulina (Limnospira Sp.). Toxins 2023, 15, 354. [Google Scholar] [CrossRef]
  101. Belay, A. Spirulina (Arthrospira): Production and Quality Assurance. In Spirulina in Human Nutrition and Health; CRC Press: Boca Raton, FL, USA, 2007; pp. 1–25. [Google Scholar]
  102. Joseph, C. Dott Elements of Pond Design and Construction. In Handbook of Microalgal Mass Culture; CRC Press: Boca Raton, FL, USA, 1986; pp. 265–283. [Google Scholar]
  103. Jung, C.H.G.; Braune, S.; Waldeck, P.; Küpper, J.-H.; Petrick, I.; Jung, F. Morphology and Growth of Arthrospira Platensis during Cultivation in a Flat-Type Bioreactor. Life 2021, 11, 536. [Google Scholar] [CrossRef]
  104. Lim, H.R.; Khoo, K.S.; Chew, K.W.; Teo, M.Y.M.; Ling, T.C.; Alharthi, S.; Alsanie, W.F.; Show, P.L. Evaluation of Real-Time Monitoring on the Growth of Spirulina Microalgae: Internet of Things and Microalgae Technologies. IEEE Internet Things J. 2024, 11, 3274–3281. [Google Scholar] [CrossRef]
  105. Havlik, I.; Lindner, P.; Scheper, T.; Reardon, K.F. On-Line Monitoring of Large Cultivations of Microalgae and Cyanobacteria. Trends Biotechnol. 2013, 31, 406–414. [Google Scholar] [CrossRef]
  106. Benelhadj, S.; Gharsallaoui, A.; Degraeve, P.; Attia, H.; Ghorbel, D. Effect of pH on the Functional Properties of Arthrospira (Spirulina) Platensis Protein Isolate. Food Chem. 2016, 194, 1056–1063. [Google Scholar] [CrossRef] [PubMed]
  107. Chronakis, I.S. Gelation of Edible Blue-Green Algae Protein Isolate (Spirulina platensis Strain Pacifica): Thermal Transitions, Rheological Properties, and Molecular Forces Involved. J. Agric. Food Chem. 2001, 49, 888–898. [Google Scholar] [CrossRef] [PubMed]
  108. Guo, X.; Wang, Q.; Yang, Q.; Gong, Z.; Wu, Y.; Liu, X. Effects of Molecular Structure and Charge State on the Foaming and Emulsifying Properties of Spirulina Protein Isolates. Food Res. Int. Ott. Ont 2024, 187, 114407. [Google Scholar] [CrossRef]
  109. Ang, W.S.; Kee, P.E.; Lan, J.C.-W.; Chen, W.-H.; Chang, J.-S.; Khoo, K.S. Unveiling the Rise of Microalgae-Based Foods in the Global Market: Perspective Views and Way Forward. Food Biosci. 2025, 66, 105390. [Google Scholar] [CrossRef]
  110. Tagliamonte, S.; Oliviero, V.; Vitaglione, P. Food Bioactive Peptides: Functionality beyond Bitterness. Nutr. Rev. 2025, 83, 369–381. [Google Scholar] [CrossRef]
  111. Begum, N.; Qi, F.; Yang, F.; Khan, Q.U.; Khan, Q.U.; Faizan, F.Q.; Li, J.; Wang, X.; Wang, X.; Wang, J.; et al. Nutritional Composition and Functional Properties of A. Platensis-Derived Peptides: A Green and Sustainable Protein-Rich Supplement. Processes 2024, 12, 2608. [Google Scholar] [CrossRef]
  112. Ughetti, A.; D’Eusanio, V.; Strani, L.; Russo, A.L.; Roncaglia, F. Influence of Drying and Storage Conditions on the Volatile Organic Compounds Profile of Spirulina Platensis. Separations 2024, 11, 180. [Google Scholar] [CrossRef]
  113. Shahid, A.; Fan, Z.; Su, K.; Zhao, A.; Mehmood, M.A.; Chang, J.-S.; Solovchenko, A.E.; Alam, M.A.; Xu, J. Sensory Chemistry of Spirulina: Unveiling Trends and Innovations in Aromatic Volatile Organic Compound Biosynthesis in off-Flavors and Odor Mitigation Strategies. Trends Food Sci. Technol. 2025, 156, 104886. [Google Scholar] [CrossRef]
  114. Koli, D.; Rudra, S.; Bhowmik, A.; Pabbi, S. Nutritional, Functional, Textural and Sensory Evaluation of Spirulina Enriched Green Pasta: A Potential Dietary and Health Supplement. Foods 2022, 11, 979. [Google Scholar] [CrossRef]
  115. Aljobair, M.O.; Albaridi, N.A.; Alkuraieef, A.N.; AlKehayez, N.M. Physicochemical Properties, Nutritional Value, and Sensory Attributes of a Nectar Developed Using Date Palm Puree and Spirulina. Int. J. Food Prop. 2021, 24, 845–858. [Google Scholar] [CrossRef]
  116. Ebid, W.M.A.; Ali, G.S.; Elewa, N.A.H. Impact of Spirulina Platensis on Physicochemical, Antioxidant, Microbiological and Sensory Properties of Functional Labneh. Discov. Food 2022, 2, 29. [Google Scholar] [CrossRef]
  117. Ottombrino, A.; Cianciabella, M.; Medoro, C.; Picariello, A.; Oliviero, A.; De Sena, V.; Predieri, S.; Rossi, M. Sensory Analyses Driven Formulation of Fruit Cereal Bars Enriched With Arthrospira platensis Dried Powder. Food Sci. Nutr. 2025, 13, e70154. [Google Scholar] [CrossRef]
  118. Niccolai, A.; Venturi, M.; Galli, V.; Pini, N.; Rodolfi, L.; Biondi, N.; D’Ottavio, M.; Batista, A.P.; Raymundo, A.; Granchi, L.; et al. Development of New Microalgae-Based Sourdough “Crostini”: Functional Effects of Arthrospira Platensis (Spirulina) Addition. Sci. Rep. 2019, 9, 19433. [Google Scholar] [CrossRef]
  119. Podgórska-Kryszczuk, I. Effect of Arthrospira Platensis (Spirulina) Fortification on Physicochemical, Nutritional, Bioactive, Textural, and Sensory Properties of Vegan Basil Pesto. Nutrients 2024, 16, 2825. [Google Scholar] [CrossRef] [PubMed]
  120. Mufidatun, A.; Koerniawan, M.D.; Siregar, U.J.; Suwanti, L.T.; Budiman, A.; Suyono, E.A. The Effect of pH on Contamination Reduction and Metabolite Contents in Mass Cultures of Spirulina (Arthrospira Platensis Gomont). Int. J. Adv. Sci. Eng. Inf. Technol. 2023, 13, 84–90. [Google Scholar] [CrossRef]
  121. McBride, R.C.; Lopez, S.; Meenach, C.; Burnett, M.; Lee, P.A.; Nohilly, F.; Behnke, C. Contamination Management in Low Cost Open Algae Ponds for Biofuels Production. Ind. Biotechnol. 2014, 10, 221–227. [Google Scholar] [CrossRef]
  122. Mary Leema, J.T.; Kirubagaran, R.; Vinithkumar, N.V.; Dheenan, P.S.; Karthikayulu, S. High Value Pigment Production from Arthrospira (Spirulina) Platensis Cultured in Seawater. Bioresour. Technol. 2010, 101, 9221–9227. [Google Scholar] [CrossRef]
  123. Franke, S.; Steingröwer, J.; Walther, T.; Krujatz, F. The Oxygen Paradigm—Quantitative Impact of High Concentrations of Dissolved Oxygen on Kinetics and Large-Scale Production of Arthrospira Platensis. ChemEngineering 2022, 6, 14. [Google Scholar] [CrossRef]
  124. Hu, X.; Zhou, J.; Liu, G.; Gui, B. Selection of Microalgae for High CO2 Fixation Efficiency and Lipid Accumulation from Ten Chlorella Strains Using Municipal Wastewater. J. Environ. Sci. 2016, 46, 83–91. [Google Scholar] [CrossRef] [PubMed]
  125. Vonshak, A.; Laorawat, S.; Bunnag, B.; Tanticharoen, M. The Effect of Light Availability on the Photosynthetic Activity and Productivity of Outdoor Cultures of Arthrospira Platensis (Spirulina). J. Appl. Phycol. 2014, 26, 1309–1315. [Google Scholar] [CrossRef]
  126. Sili, C.; Torzillo, G.; Vonshak, A. Arthrospira (Spirulina). In Ecology of Cyanobacteria II; Whitton, B.A., Ed.; Springer: Dordrecht, The Netherlands, 2012; pp. 677–705. ISBN 978-94-007-3854-6. [Google Scholar]
  127. Siva Kiran, R.R.; Madhu, G.; Satyanarayana, S.V. Spirulina in Combating Protein Energy Malnutrition (PEM) and Protein Energy Wasting (PEW)—A Review. J. Nutr. Res. 2015, 3, 62–79. [Google Scholar] [CrossRef]
  128. Melikoglu, M. Waste Valorization Strategies with Inputs for Microalgae Biorefineries: A Global Review. Clean. Water 2025, 4, 100125. [Google Scholar] [CrossRef]
  129. Chen, Y.; Wan, W.-W.; Cui, K.-H.; Lau, B.P.-Y.; Lee, F.W.-F.; Xu, S.J.-L. Feasibility and Efficiency of Microalgae Cultivation for Nutrient Recycling and Energy Recovery from Food Waste Filtrate. PLoS ONE 2025, 20, e0315801. [Google Scholar] [CrossRef] [PubMed]
  130. Chacón-Lee, T.L.; González-Mariño, G.E. Microalgae for “Healthy” Foods—Possibilities and Challenges. Compr. Rev. Food Sci. Food Saf. 2010, 9, 655–675. [Google Scholar] [CrossRef]
  131. U.S. Food and Drug Administration. GRAS Notices, GRN No. 417, Dried Biomass of Arthrospira Platensis Also Known as Spirulina Platensis (Spirulina). 2012. Available online: https://hfpappexternal.fda.gov/scripts/fdcc/index.cfm?set=grasnotices&id=417 (accessed on 28 October 2025).
  132. Gromek, W.; Kołdej, N.; Kurowski, M.; Majsiak, E. Spirulina (Arthrospira Platensis): Antiallergic Agent or Hidden Allergen? A Literature Review. Foods Basel Switz. 2024, 13, 1052. [Google Scholar] [CrossRef]
  133. Le, T.-M.; Knulst, A.C.; Röckmann, H. Anaphylaxis to Spirulina Confirmed by Skin Prick Test with Ingredients of Spirulina Tablets. Food Chem. Toxicol. 2014, 74, 309–310. [Google Scholar] [CrossRef] [PubMed]
  134. Pescosolido, E.; Yerly, D.; Caubet, J.-C.; Bergmann, M.M. Delayed IgE–Mediated Hypersensitivity to Arthrospira Platensis (Spirulina). Ann. Allergy. Asthma. Immunol. 2022, 129, 522–524. [Google Scholar] [CrossRef] [PubMed]
  135. French Agency for Food, Environmental and Occupational Health & Safety. ANSES Opinion Request No 2014-SA-0096: Opinion of the French Agency for Food, Environmental and Occupational Health & Safety on the “Risks Associated with the Consumption of Food Supplements Containing Spirulina”. ANSES Opin. 2017, 38. Available online: https://www.anses.fr/en/system/files/NUT2014SA0096EN.pdf (accessed on 28 October 2025).
  136. Zhou, C.; Jepsen, C.S.; Rigby, N.; Lübeck, M.; Mackie, A.; Sancho, A.I.; Bøgh, K.L. Evaluation of Allergenicity of the Alternative Food Protein Sources Microalga Spirulina and Rapeseed Cake—A Study in Brown Norway Rats. Food Res. Int. 2025, 221, 117357. [Google Scholar] [CrossRef]
  137. Bianco, M.; Ventura, G.; Calvano, C.D.; Losito, I.; Cataldi, T.R.I. Discovery of Marker Peptides of Spirulina Microalga Proteins for Allergen Detection in Processed Foodstuffs. Food Chem. 2022, 393, 133319. [Google Scholar] [CrossRef]
  138. Mao, T.K.; Van de Water, J.; Gershwin, M.E. Effects of a Spirulina-Based Dietary Supplement on Cytokine Production from Allergic Rhinitis Patients. J. Med. Food 2005, 8, 27–30. [Google Scholar] [CrossRef]
  139. Cingi, C.; Conk-Dalay, M.; Cakli, H.; Bal, C. The Effects of Spirulina on Allergic Rhinitis. Eur. Arch. Oto-Rhino-Laryngol. Off. J. Eur. Fed. Oto-Rhino-Laryngol. Soc. EUFOS Affil. Ger. Soc. Oto-Rhino-Laryngol.—Head Neck Surg. 2008, 265, 1219–1223. [Google Scholar] [CrossRef]
  140. Nourollahian, M.; Rasoulian, B.; Gafari, A.; Anoushiravani, M.; Jabari, F.; Bakhshaee, M. Clinical Comparison of the Efficacy of Spirulina Platensis and Cetirizine for Treatment of Allergic Rhinitis. Acta Otorhinolaryngol. Ital. 2020, 40, 224–229. [Google Scholar] [CrossRef]
  141. Batista, I.G.; Person, O.C.; Bogar, P.; Angélico Júnior, F.V. Efficacy of Spirulina for Allergic Rhinitis. medRxiv 2021. [Google Scholar] [CrossRef]
  142. Cruz, J.D.; Vasconcelos, V. Legal Aspects of Microalgae in the European Food Sector. Foods Basel Switz. 2023, 13, 124. [Google Scholar] [CrossRef] [PubMed]
  143. Boisen, S.; Eggum, B.O. Critical Evaluation of in Vitro Methods for Estimating Digestibility in Simple-Stomach Animals. Nutr. Res. Rev. 1991, 4, 141–162. [Google Scholar] [CrossRef] [PubMed]
Life 15 01789 g001
Figure 1. (Left) Typical morphology of spirals and (Right) overview of functional categories of Arthrospira platensis proteins (Left: phase contrast microscopy, 40-fold primary magnification, EVOS™ XL Core Imaging System, Thermo Fisher Scientific, Darmstadt, Germany).
Figure 1. (Left) Typical morphology of spirals and (Right) overview of functional categories of Arthrospira platensis proteins (Left: phase contrast microscopy, 40-fold primary magnification, EVOS™ XL Core Imaging System, Thermo Fisher Scientific, Darmstadt, Germany).
Life 15 01789 g001
Table 1. Protein content of Arthrospira platensis of different strains.
Table 1. Protein content of Arthrospira platensis of different strains.
SpeciesProtein Content
[% Dry Matter]		Reference
Arthrospira fusiformis35–64[30,31]
Arthrospira maxima60–71 [32,33,34]
Arthrospira platensis17–80 [35,36]
Table 2. Total yield, protein content, and protein yield of traditional crops and cultures of Arthrospira platensis.
Table 2. Total yield, protein content, and protein yield of traditional crops and cultures of Arthrospira platensis.
CropTotal Yield
[Tons/(ha per Year)]		Protein Content
[%]		Protein Yield
[Tons/(ha per Year)]
Wheat6.79.50.64
Maize147.41.04
Rice (hulled)87.10.57
Soybeans4351.4
Arthrospira platensis60–706539–45
Table 3. Representative protein contents of Arthrospira platensis under different production conditions.
Table 3. Representative protein contents of Arthrospira platensis under different production conditions.
Culture SystemCulture MediumProtein Content
[%]		Reference
Erlenmeyer flaskSchlösser
(+Beet vinasse)		39–55
35–72		[51]
PhotobioreactorSchlösser
(urea as nitrogen source)		18–71[52]
PhotobioreactorZarrouk
(sodium nitrate modification)		56–70[53]
PhotobioreactorZarrouk69[36]
PhotobioreactorFM-II58–63[48]
Semicontinuous culture systemConwey61–64[54]
Elongated minitank
(outdoor pond simulation)		Schlösser
(urea as nitrogen source)		45–62[55]
Open pondKosaric and digested sago starch factory wastewater53–68[56]
Open pondModified Zarrouk56[57]
Open pondn.d.59[58]
Erlenmeyer flask“Standard” with variations17–32[35]
n.d.n.d.46–63[32]
Table 4. Dietary amino acid requirements as stated by the Food and Agriculture Organization of the United Nations in comparison to profiles of Arthrospira and egg white as stated by the U.S. Department of Agriculture, Agricultural Research Service.
Table 4. Dietary amino acid requirements as stated by the Food and Agriculture Organization of the United Nations in comparison to profiles of Arthrospira and egg white as stated by the U.S. Department of Agriculture, Agricultural Research Service.
Essential
Amino Acids
(EAAs)		Required
[g/100 g Protein]
[63,64]		Arthrospira
[g/100 g Protein]
[61]		Egg White
[g/100 g Protein]
[65]
Histidine1.61.080.257
Isoleucine3.03.210.609
Leucine6.14.951.02
Lysine4.83.020.822
Methionin + Cystine2.31.15 + 0.6620.486 + 0.407
Phenylalanin + Tyrosine4.12.78 + 2.580.726 + 0.466
Threonine2.52.970.567
Tryptophan0.660.9290.188
Valine4.03.510.779
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Braune, S.; Jung, C.G.H.; Küpper, J.-H.; Jung, F. Arthrospira platensis as Protein-Rich Source for Human Nutrition. Life 2025, 15, 1789. https://doi.org/10.3390/life15121789

AMA Style

Braune S, Jung CGH, Küpper J-H, Jung F. Arthrospira platensis as Protein-Rich Source for Human Nutrition. Life. 2025; 15(12):1789. https://doi.org/10.3390/life15121789

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Braune, Steffen, Conrad G. H. Jung, Jan-Heiner Küpper, and Friedrich Jung. 2025. "Arthrospira platensis as Protein-Rich Source for Human Nutrition" Life 15, no. 12: 1789. https://doi.org/10.3390/life15121789

APA Style

Braune, S., Jung, C. G. H., Küpper, J.-H., & Jung, F. (2025). Arthrospira platensis as Protein-Rich Source for Human Nutrition. Life, 15(12), 1789. https://doi.org/10.3390/life15121789

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