Door: Elias Scheldeman
Gebaseerd op masterproefonderzoek aan de Universiteit Gent
Als we denken aan alternatieven voor vlees, komen we al snel uit bij soja uit Zuid-Amerika of andere plantaardige bronnen geteeld op het land. Maar wat als de duurzame eiwitbron van de toekomst gewoon voor onze eigen kust drijft? Masteronderzoek aan de Universiteit Gent toont aan dat zeesla (Ulva), een veel voorkomend groenwier in de Noordzee en bij uitbreiding ter wereld, boordevol hoogwaardige eiwitten én gezondheidsvoordelen zit. Toch laten we deze groene goudmijn momenteel links liggen.
Wie aan zeewier denkt, denkt waarschijnlijk aan een rolletje sushi of aan glibberige planten op het strand. Toch consumeert u onbewust al dagelijks zeewier. De westerse levensmiddelenindustrie gebruikt al op grote schaal zeewier, maar vrijwel uitsluitend voor technologische doeleinden. Hoewel rood- en bruinwieren aanzienlijke hoeveelheden eiwitten bevatten, worden ze vandaag vooral geoogst voor hun hydrocolloïden (zoals alginaat en carrageen). Deze dienen als verdikkingsmiddelen of stabilisatoren in sauzen, ijs en desserts maar worden evengoed in cosmeticaproducten gebruikt.
Groenwieren, specifiek het geslacht Ulva, worden in industriële toepassingen grotendeels over het hoofd gezien. Dit is een gemiste kans. Ulva is wereldwijd verspreid, veruit het meest abundante groenwier ter wereld en bezit, zo blijkt nu, een hoogwaardig eiwitprofiel met alle essentiële aminozuren.

Diversiteit aan macroalgen in de Noordzee.
Cijfers van de Voedsel- en Landbouworganisatie van de VN (FAO) tonen hoe hard Europa achterloopt:
Door onze eigen Noordzee te benutten voor lokale grootschalige kweek, kan de EU haar afhankelijkheid van plantaardige eiwitimport drastisch verkleinen.
Om de werkelijke nutritionele waarde van Ulva-eiwitten te bepalen, volstaat het meten van het totale eiwitgehalte niet; de eiwitten moeten ook biologisch beschikbaar zijn voor het menselijk lichaam. In dit onderzoek werd de menselijke spijsvertering gesimuleerd met het internationaal erkende INFOGEST 2.0-protocol.
Dit is een gestandaardiseerde laboratoriummethode die de reis van voedsel door onze mond, maag en dunne darm nabootst. Door stapsgewijs de juiste menselijke enzymen, zuren en temperaturen toe te passen in labo-omstandigheden, wordt nauwkeurig gemeten in welke mate eiwitten daadwerkelijk worden afgebroken tot opneembare aminozuren.
Workflow van het onderzoek.
Het onderzoek leverde daarnaast interessante nutritionele inzichten op: tijdens deze enzymatische afbraak in de gesimuleerde maag en darm gebeurt er meer dan alleen vertering. Wanneer Ulva-eiwitten in stukjes worden geknipt, komen er kleine eiwitfragmenten vrij die bioactieve peptiden worden genoemd.
Onderzoek heeft uitgewezen dat deze specifieke peptiden actieve functies in ons lichaam kunnen vervullen zodra ze zijn opgenomen. Eerder soortgelijk onderzoek toonde onder andere een anti-diabetische werking aan en een effectieve werking tegen een hoge bloeddruk (ACE-remming). Ulva levert dus niet alleen bouwstoffen, maar produceert tijdens de vertering ook actieve componenten die onze gezondheid direct kunnen ondersteunen.
Het onderzoek nam vier verschillende Ulva-soorten onder de loep (twee Europese en twee Zuid-Koreaanse soorten). Uit de resultaten blijkt dat de eiwitverteerbaarheid enorm varieert: van matig 38% tot sterk 83%.
Het vernieuwende van dit onderzoek ligt in het feit dat het niet stopt bij de conclusie dat één specifieke soort genetisch "beter" presteert. Het onderzoek toont aan dat de biologische beschikbaarheid van het eiwit wordt bepaald door de interactie met andere stoffen in het wier, zoals suikers en fenolen.
Deze componenten binden zich aan de eiwitten en vormen een structurele barrière voor onze spijsverteringsenzymen.
Uit de resultaten van dit onderzoek blijkt dat focussen op één specifieke "super-soort" een valkuil is voor producenten, omdat de kwaliteit bij een volgende oogst plotseling kan afwijken. Dit heeft twee directe gevolgen voor de ontwikkeling van een Europese industrie:
Dit onderzoek bewijst dat we de complexiteit van zeewier niet mogen onderschatten. Eiwitverteerbaarheid is geen vaststaand biologisch feit, maar het resultaat van een dynamisch samenspel tussen de soort en zijn omgeving. Dankzij deze nieuwe inzichten beschikt de sector nu over de concrete biochemische randvoorwaarden om de stap te zetten van onvoorspelbare wildpluk naar een stabiele, hoogwaardige industrie.
De toekomst van onze eiwittransitie ligt dus niet enkel op het land, maar ook voor onze eigen kust. Het is tijd om in de Noordzee te gaan oogsten: zeesla is klaar om van een over het hoofd gezien groenwier te transformeren tot een lokale, niet meer weg te denken plantaardige eiwitbron op ons bord.
Abdollahi, M., Axelsson, J., Carlsson, N.-G., Nylund, G. M., Albers, E., & Undeland, I. (2019). Effect of stabilization method and freeze/thaw-aided precipitation on structural and functional properties of proteins recovered from brown seaweed (Saccharina latissima). Food Hydrocolloids, 96, 140–150. https://doi.org/10.1016/j.foodhyd.2019.05.007
Ahn, J., Cao, M.-J., Yu, Y. Q., & Engen, J. R. (2013). Accessing the reproducibility and specificity of pepsin and other aspartic proteases. Biochimica et Biophysica Acta (BBA) - Proteins and Proteomics, 1834(6), 1222–1229. https://doi.org/10.1016/j.bbapap.2012.10.003
Albers, E., Malmhäll-Bah, E., Olsson, J., Sterner, M., Mayers, J. J., Nylund, G. M., Rupar-Gadd, K., Abdollahi, M., Cvijetinovic, S., Welander, U., Edlund, U., Pavia, H., & Undeland, I. (2021). Influence of preservation methods on biochemical composition and downstream processing of cultivated Saccharina latissima biomass. Algal Research, 55, 102261. https://doi.org/10.1016/j.algal.2021.102261
Angell, A. R., Angell, S. F., De Nys, R., & Paul, N. A. (2016). Seaweed as a protein source for mono-gastric livestock. Trends in Food Science & Technology, 54, 74–84. https://doi.org/10.1016/j.tifs.2016.05.014
Angell, A. R., Paul, N. A., & De Nys, R. (2017). A comparison of protocols for isolating and concentrating protein from the green seaweed Ulva ohnoi. Journal of Applied Phycology, 29(2), 1011–1026. https://doi.org/10.1007/s10811-016-0972-7
Arbita, A. A., Paul, N. A., Cox, J., & Zhao, J. (2024). Screening and isolation of milk-clotting enzymes from seaweed. Journal of Applied Phycology, 36(2), 887–896. https://doi.org/10.1007/s10811-023-02975-4
Archibald, J. M. (2009). The Puzzle of Plastid Evolution. Current Biology, 19(2), R81–R88. https://doi.org/10.1016/j.cub.2008.11.067
Aziz, A., Roy, S., Hassan, A., Haque, M., Shahjadee, U., Mondal, K., Mohanta, L., Mashuk, O., & Saha, B. (2021). Potential of Ulva linza L. and Caulerpa racemosa var. Uvifera seaweeds from Cox’s Bazar, Bangladesh as sea vegetable. Bangladesh Journal of Scientific and Industrial Research, 56(3), 223–230. https://doi.org/10.3329/bjsir.v56i3.55970
Bender, L., Harding, D., & Lee, G. (2005). The Facts on File Illustrated Guide to the Human Body: Digestive System. Facts on File.
Benjama, O., & Masniyom, P. (2011). Nutritional composition and physicochemical properties of two green seaweeds (Ulva pertusa and U. intestinalis) from the Pattani Bay in southern Thailand.
Bennici, A. (2008). Origin and early evolution of land plants: Problems and considerations. Communicative & Integrative Biology, 1(2), 212–218. https://doi.org/10.4161/cib.1.2.6987
Berk, Z. (with FAO). (1992). Technology of production of edible flours and protein products from soybeans. Food and Agriculture Organization of the United Nations.
Bermano, G. (2020). Seaweed-derived bioactives as potential energy regulators in obesity and type 2 diabetes. In Advances in Pharmacology (Vol. 87, pp. 205–256). Elsevier. https://doi.org/10.1016/bs.apha.2019.10.002
Biancarosa, I., Espe, M., Bruckner, C. G., Heesch, S., Liland, N., Waagbø, R., Torstensen, B., & Lock, E. J. (2017). Amino acid composition, protein content, and nitrogen-to-protein conversion factors of 21 seaweed species from Norwegian waters. Journal of Applied Phycology, 29(2), 1001–1009. https://doi.org/10.1007/s10811-016-0984-3
Blomster, J., Maggs, C. A., & Stanhope, M. J. (1998). Molecular and morphological analysis of Enteromorpha intestinalis and E. compressa (chlorophyta) in the british isles. Journal of Phycology, 34(2), 319–340. https://doi.org/10.1046/j.1529-8817.1998.340319.x
Braissant, O., Astasov-Frauenhoffer, M., Waltimo, T., & Bonkat, G. (2020). A Review of Methods to Determine Viability, Vitality, and Metabolic Rates in Microbiology. Frontiers in Microbiology, 11, 547458. https://doi.org/10.3389/fmicb.2020.547458
Brodkorb, A., Egger, L., Alminger, M., Alvito, P., Assunção, R., Ballance, S., Bohn, T., Bourlieu-Lacanal, C., Boutrou, R., Carrière, F., Clemente, A., Corredig, M., Dupont, D., Dufour, C., Edwards, C., Golding, M., Karakaya, S., Kirkhus, B., Le Feunteun, S., … Recio, I. (2019). INFOGEST static in vitro simulation of gastrointestinal food digestion. Nature Protocols, 14(4), 991–1014. https://doi.org/10.1038/s41596-018-0119-1
Buettner, A. (2001). Observation of the Swallowing Process by Application of Videofluoroscopy and Real-time Magnetic Resonance Imaging—Consequences for Retronasal Aroma Stimulation. Chemical Senses, 26(9), 1211–1219. https://doi.org/10.1093/chemse/26.9.1211
Burson, A., Stomp, M., Akil, L., Brussaard, C. P. D., & Huisman, J. (2016). Unbalanced reduction of nutrient loads has created an offshore gradient from phosphorus to nitrogen limitation in the North Sea. Limnology and Oceanography, 61(3), 869–888. https://doi.org/10.1002/lno.10257
Buschmann, A. H., Prescott, S., Potin, P., Faugeron, S., Vásquez, J. A., Camus, C., Infante, J., Hernández-González, M. C., Gutíerrez, A., & Varela, D. A. (2014). The Status of Kelp Exploitation and Marine Agronomy, with Emphasis on Macrocystis pyrifera, in Chile. In Advances in Botanical Research (Vol. 71, pp. 161–188). Elsevier. https://doi.org/10.1016/B978-0-12-408062-1.00006-8
Butré, C. I., Wierenga, P. A., & Gruppen, H. (2012). Effects of Ionic Strength on the Enzymatic Hydrolysis of Diluted and Concentrated Whey Protein Isolate. Journal of Agricultural and Food Chemistry, 60(22), 5644–5651. https://doi.org/10.1021/jf301409n
Cahill, P. L., Hurd, C. L., & Lokman, M. (2010). Keeping the water clean—Seaweed biofiltration outperforms traditional bacterial biofilms in recirculating aquaculture. Aquaculture, 306(1–4), 153–159. https://doi.org/10.1016/j.aquaculture.2010.05.032
Caicedo-Lopez, L. H., Luzardo-Ocampo, I., Cuellar-Nuñez, M. L., Campos-Vega, R., Mendoza, S., & Loarca-Piña, G. (2019). Effect of the in vitro gastrointestinal digestion on free-phenolic compounds and mono/oligosaccharides from Moringa oleifera leaves: Bioaccessibility, intestinal permeability and antioxidant capacity. Food Research International, 120, 631–642. https://doi.org/10.1016/j.foodres.2018.11.017
Camporesi, G., Canzian, C., & Bordoni, A. (2026). Optimizing INFOGEST Digest Conditioning for Reliable in vitro Assessment of Nutrient Bioavailability Using Caco-2 Cell Models. Nutrients, 18(2), 339. https://doi.org/10.3390/nu18020339
Carl, C., De Nys, R., Lawton, R. J., & Paul, N. A. (2014). Methods for the Induction of Reproduction in a Tropical Species of Filamentous Ulva. PLoS ONE, 9(5), e97396. https://doi.org/10.1371/journal.pone.0097396
Cebrián-Lloret, V., Martínez-Abad, A., Recio, I., López-Rubio, A., & Martínez-Sanz, M. (2024). In vitro digestibility of proteins from red seaweeds: Impact of cell wall structure and processing methods. Food Research International, 178, 113990. https://doi.org/10.1016/j.foodres.2024.113990
Černá, M. (2011). Seaweed Proteins and Amino Acids as Nutraceuticals. In Advances in Food and Nutrition Research (Vol. 64, pp. 297–312). Elsevier. https://doi.org/10.1016/B978-0-12-387669-0.00024-7
Chen, J. (2009). Food oral processing—A review. Food Hydrocolloids, 23(1), 1–25. https://doi.org/10.1016/j.foodhyd.2007.11.013
Chen, J., Gaikwad, V., Holmes, M., Murray, B., Povey, M., Wang, Y., & Zhang, Y. (2011). Development of a simple model device for in vitro gastric digestion investigation. Food & Function, 2(3–4), 174. https://doi.org/10.1039/c0fo00159g
Chew, K. W., Chia, S. R., Lee, S. Y., Zhu, L., & Show, P. L. (2019). Enhanced microalgal protein extraction and purification using sustainable microwave-assisted multiphase partitioning technique. Chemical Engineering Journal, 367, 1–8. https://doi.org/10.1016/j.cej.2019.02.131
Childs, P. E. (2021). The history of the chemicals from seaweed industry in Ireland. Bulletin for the History of Chemistry, 46(2), 149–163. https://doi.org/10.70359/bhc2021v046p149
Church, F. C., Porter, D. H., Catignani, G. L., & Swaisgood, H. E. (1985). An o-phthalaldehyde spectrophotometric assay for proteinases. Analytical Biochemistry, 146(2), 343–348. https://doi.org/10.1016/0003-2697(85)90549-4
Cian, R. E., Hernández-Chirlaque, C., Gámez-Belmonte, R., Drago, S. R., Sánchez De Medina, F., & Martínez-Augustin, O. (2018). Green Alga Ulva spp. Hydrolysates and Their Peptide Fractions Regulate Cytokine Production in Splenic Macrophages and Lymphocytes Involving the TLR4-NFκB/MAPK Pathways. Marine Drugs, 16(7), 235. https://doi.org/10.3390/md16070235
Cian, R. E., Nardo, A. E., Garzón, A. G., Añon, M. C., & Drago, S. R. (2022). Identification and in silico study of a novel dipeptidyl peptidase IV inhibitory peptide derived from green seaweed Ulva spp. hydrolysates. LWT, 154, 112738. https://doi.org/10.1016/j.lwt.2021.112738
Coaten, D. J., Guls, H. D., Þorsteinsdóttir, M., & Halldórsson, H. P. (2023). Effect of Environmental Physico-Chemical Parameters on the Biochemical Composition of Wild Icelandic Laminaria digitata and Saccharina latissima (Laminariaceae, Phaeophyceae). Regional Studies in Marine Science, 60, 102839. https://doi.org/10.1016/j.rsma.2023.102839
Copernicus Marine Service. (n.d.). Expert viewer. Copernicus Marine Service. Retrieved 28 October 2025, from https://data.marine.copernicus.eu/viewer/expert
Corino, C., Modina, S. C., Di Giancamillo, A., Chiapparini, S., & Rossi, R. (2019). Seaweeds in Pig Nutrition. Animals, 9(12), 1126. https://doi.org/10.3390/ani9121126
Costa, S. P., Cotas, J., & Pereira, L. (2024). Laminar Ulva Species: A Multi-Tool for Humankind? Applied Sciences, 14(8), 3448. https://doi.org/10.3390/app14083448
Cox, S., Abu-Ghannam, N., & Gupta, S. (2010). An Assessment of the Antioxidant and Antimicrobial Activity of Six Species of Edible Irish Seaweeds. https://doi.org/10.21427/D7HC92
Dawczynski, C., Schubert, R., & Jahreis, G. (2007). Amino acids, fatty acids, and dietary fibre in edible seaweed products. Food Chemistry, 103(3), 891–899. https://doi.org/10.1016/j.foodchem.2006.09.041
Day, L. (2013). Proteins from land plants – Potential resources for human nutrition and food security. Trends in Food Science & Technology, 32(1), 25–42. https://doi.org/10.1016/j.tifs.2013.05.005
DuBois, Michel., Gilles, K. A., Hamilton, J. K., Rebers, P. A., & Smith, Fred. (1956). Colorimetric Method for Determination of Sugars and Related Substances. Analytical Chemistry, 28(3), 350–356. https://doi.org/10.1021/ac60111a017
Duong-Ly, K. C., & Gabelli, S. B. (2014). Salting out of Proteins Using Ammonium Sulfate Precipitation. In Methods in Enzymology (Vol. 541, pp. 85–94). Elsevier. https://doi.org/10.1016/B978-0-12-420119-4.00007-0
Dupont, D., Alric, M., Blanquet-Diot, S., Bornhorst, G., Cueva, C., Deglaire, A., Denis, S., Ferrua, M., Havenaar, R., Lelieveld, J., Mackie, A. R., Marzorati, M., Menard, O., Minekus, M., Miralles, B., Recio, I., & Van Den Abbeele, P. (2019). Can dynamic in vitro digestion systems mimic the physiological reality? Critical Reviews in Food Science and Nutrition, 59(10), 1546–1562. https://doi.org/10.1080/10408398.2017.1421900
Echave, J., Fraga-Corral, M., Garcia-Perez, P., Popović-Djordjević, J., H. Avdović, E., Radulović, M., Xiao, J., A. Prieto, M., & Simal-Gandara, J. (2021). Seaweed Protein Hydrolysates and Bioactive Peptides: Extraction, Purification, and Applications. Marine Drugs, 19(9), 500. https://doi.org/10.3390/md19090500
Echave, J., Otero, P., Garcia-Oliveira, P., Munekata, P. E. S., Pateiro, M., Lorenzo, J. M., Simal-Gandara, J., & Prieto, M. A. (2022). Seaweed-Derived Proteins and Peptides: Promising Marine Bioactives. Antioxidants, 11(1), 176. https://doi.org/10.3390/antiox11010176
El Mellouki, M., Ait Benichou, S., Elhamdani, N., Cherki, M., & Dakir, M. (2025). Chemical and biological properties of green freshwater alga Ulva intestinalis collected from Casablanca, Morocco. Egyptian Journal of Aquatic Research, 51(1), 39–47. https://doi.org/10.1016/j.ejar.2024.12.006
El-Beltagi, H. S., Mohamed, A. A., Mohamed, H. I., Ramadan, K. M. A., Barqawi, A. A., & Mansour, A. T. (2022). Phytochemical and Potential Properties of Seaweeds and Their Recent Applications: A Review. Marine Drugs, 20(6), 342. https://doi.org/10.3390/md20060342
El-Deeb, N. M., Ahmed, T. A. E., Premarathna, A. D., Rjabovs, V., Tuvikene, R., Hammami, R., Boulianne, M., & Hincke, M. T. (2025). Experimental datasets on the extraction of functional ingredients from seaweeds for controlling bacterial infection. Data in Brief, 60, 111569. https://doi.org/10.1016/j.dib.2025.111569
EnCor Biotechnology Inc. (n.d.). Ammonium sulfate precipitation protocol. EnCor Biotechnology. Retrieved 10 March 2026, from https://files.encorbio.com/protocols/AM-SO4.htm
Endres, J. G., American Oil Chemists’ Society, & Soy Protein Council (Eds). (2001). Soy protein products: Characteristics, nutritional aspects, and utilization (Rev. and expanded ed). AOCS Press.
Espinosa-Ramírez, J., Mondragón-Portocarrero, A. C., Rodríguez, J. A., Lorenzo, J. M., & Santos, E. M. (2023). Algae as a potential source of protein meat alternatives. Frontiers in Nutrition, 10, 1254300. https://doi.org/10.3389/fnut.2023.1254300
European Food Safety Authority (EFSA). (2017). Dietary Reference Values for nutrients Summary report. EFSA Supporting Publications, 14(12). https://doi.org/10.2903/sp.efsa.2017.e15121
European Food Safety Authority (EFSA), Dujardin, B., Ferreira de Sousa, R., & Gómez Ruiz, J. Á. (2023). Dietary exposure to heavy metals and iodine intake via consumption of seaweeds and halophytes in the European population. EFSA Journal, 21(1). https://doi.org/10.2903/j.efsa.2023.7798
FAO. (2014). Fisheries and aquaculture information and statistics services (Food and Agriculture Organization of the United Nations.) [Data set]. https://www.fao.org/home/en
FAO. (2018). The global status of seaweed production, trade and utilization. FAO
Farghali, M., Mohamed, I. M. A., Osman, A. I., & Rooney, D. W. (2023). Seaweed for climate mitigation, wastewater treatment, bioenergy, bioplastic, biochar, food, pharmaceuticals, and cosmetics: A review. Environmental Chemistry Letters, 21(1), 97–152. https://doi.org/10.1007/s10311-022-01520-y
Fleurence, J. (1999). Seaweed proteins. Trends in Food Science & Technology, 10(1), 25–28. https://doi.org/10.1016/S0924-2244(99)00015-1
Fleurence, J., Le Coeur, C., Mabeau, S., Maurice, M., & Landrein, A. (1995). Comparison of different extractive procedures for proteins from the edible seaweeds Ulva rigida and Ulva rotundata. Journal of Applied Phycology, 7(6), 577–582. https://doi.org/10.1007/BF00003945
Fleurence, J., Massiani, L., Guyader, O., & Mabeau, S. (1995). Use of enzymatic cell wall degradation for improvement of protein extraction from Chondrus crispus, Gracilaria verrucosa and Palmaria palmata. Journal of Applied Phycology, 7(4), 393–397. https://doi.org/10.1007/BF00003796
Fontes-Candia, C., Díaz-Piñero, L., Carlos Martínez, J., Gómez-Mascaraque, L. G., López-Rubio, A., & Martínez-Sanz, M. (2023). Nanostructural changes in Polysaccharide-Casein Gel-Like structures upon in vitro gastrointestinal digestion. Food Research International, 169, 112862. https://doi.org/10.1016/j.foodres.2023.112862
Forbord, S., Matsson, S., Brodahl, G. E., Bluhm, B. A., Broch, O. J., Handå, A., Metaxas, A., Skjermo, J., Steinhovden, K. B., & Olsen, Y. (2020). Latitudinal, seasonal and depth-dependent variation in growth, chemical composition and biofouling of cultivated Saccharina latissima (Phaeophyceae) along the Norwegian coast. Journal of Applied Phycology, 32(4), 2215–2232. https://doi.org/10.1007/s10811-020-02038-y
Fu, Z., Akula, S., Thorpe, M., & Hellman, L. (2021). Marked difference in efficiency of the digestive enzymes pepsin, trypsin, chymotrypsin, and pancreatic elastase to cleave tightly folded proteins. Biological Chemistry, 402(7), 861–867. https://doi.org/10.1515/hsz-2020-0386
Fujiwara-Arasaki, T., Mino, N., & Kuroda, M. (1984). The protein value in human nutrition of edible marine algae in Japan. Hydrobiologia, 116–117(1), 513–516. https://doi.org/10.1007/BF00027735
Ganesan, A. R., Tiwari, U., & Rajauria, G. (2019). Seaweed nutraceuticals and their therapeutic role in disease prevention. Food Science and Human Wellness, 8(3), 252–263. https://doi.org/10.1016/j.fshw.2019.08.001
Gao, G., Clare, A. S., Rose, C., & Caldwell, G. S. (2017). Reproductive sterility increases the capacity to exploit the green seaweed Ulva rigida for commercial applications. Algal Research, 24, 64–71. https://doi.org/10.1016/j.algal.2017.03.008
Garcia-Vaquero, M., Mora, L., & Hayes, M. (2019). In Vitro and In Silico Approaches to Generating and Identifying Angiotensin-Converting Enzyme I Inhibitory Peptides from Green Macroalga Ulva lactuca. Marine Drugs, 17(4), 204. https://doi.org/10.3390/md17040204
Gephart, J. A., Henriksson, P. J. G., Parker, R. W. R., Shepon, A., Gorospe, K. D., Bergman, K., Eshel, G., Golden, C. D., Halpern, B. S., Hornborg, S., Jonell, M., Metian, M., Mifflin, K., Newton, R., Tyedmers, P., Zhang, W., Ziegler, F., & Troell, M. (2021). Environmental performance of blue foods. Nature, 597(7876), 360–365. https://doi.org/10.1038/s41586-021-03889-2
Gerhardt Analytical Systems. (n.d.). Kjeldahl method. https://www.gerhardt.de/en/know-how/analytical-methods/kjeldahl-method/
Ghaderiardakani, F. (2019). Ulva growth, development and applications[Doctoral disseration]. The University of Birmingham.
Gorissen, S. H. M., Crombag, J. J. R., Senden, J. M. G., Waterval, W. A. H., Bierau, J., Verdijk, L. B., & Van Loon, L. J. C. (2018). Protein content and amino acid composition of commercially available plant-based protein isolates. Amino Acids, 50(12), 1685–1695. https://doi.org/10.1007/s00726-018-2640-5
Grácio, M., Oliveira, S., Lima, A., & Boavida Ferreira, R. (2023). RuBisCO as a protein source for potential food applications: A review. Food Chemistry, 419, 135993. https://doi.org/10.1016/j.foodchem.2023.135993
Graham, L. E., & Wilcox, L. W. (2000). Algae. Prentice Hall.
Guiry, M. D., & Guiry, G. (2022). National University of Ireland. AlgaeBase. https://www.algaebase.org
Haendiges, J., Timme, R., Ramachandran, P., & Balkey, M. (2020). DNA Quantification using the Qubit Fluorometer v1. https://doi.org/10.17504/protocols.io.bi8dkhs6
Han, T., Kang, S.-H., Park, J.-S., Lee, H.-K., & Brown, M. T. (2008). Physiological responses of Ulva pertusa and U. armoricana to copper exposure. Aquatic Toxicology, 86(2), 176–184. https://doi.org/10.1016/j.aquatox.2007.10.016
Harrysson, H., Hayes, M., Eimer, F., Carlsson, N.-G., Toth, G. B., & Undeland, I. (2018). Production of protein extracts from Swedish red, green, and brown seaweeds, Porphyra umbilicalis Kützing, Ulva lactuca Linnaeus, and Saccharina latissima (Linnaeus) J. V. Lamouroux using three different methods. Journal of Applied Phycology, 30(6), 3565–3580. https://doi.org/10.1007/s10811-018-1481-7
Harrysson, H., Konasani, V. R., Toth, G. B., Pavia, H., Albers, E., & Undeland, I. (2019). Strategies for Improving the Protein Yield in pH-Shift Processing of Ulva lactuca Linnaeus: Effects of Ulvan Lyases, pH-Exposure Time, and Temperature. ACS Sustainable Chemistry & Engineering, 7(15), 12688–12691. https://doi.org/10.1021/acssuschemeng.9b02781
Hayden, H. S., Blomster, J., Maggs, C. A., Silva, P. C., Stanhope, M. J., & Waaland, J. R. (2003). Linnaeus was right all along: Ulva and Enteromorpha are not distinct genera. European Journal of Phycology, 38(3), 277–294. https://doi.org/10.1080/1364253031000136321
Hayes, M. (2021). Bioactive Peptides in Preventative Healthcare: An Overview of Bioactivities and Suggested Methods to Assess Potential Applications. Current Pharmaceutical Design, 27(11), 1332–1341. https://doi.org/10.2174/1381612827666210125155048
Holdt, S. L., & Kraan, S. (2011). Bioactive compounds in seaweed: Functional food applications and legislation. Journal of Applied Phycology, 23(3), 543–597. https://doi.org/10.1007/s10811-010-9632-5
Hou, Y., Wu, Z., Dai, Z., Wang, G., & Wu, G. (2017). Protein hydrolysates in animal nutrition: Industrial production, bioactive peptides, and functional significance. Journal of Animal Science and Biotechnology, 8(1), 24. https://doi.org/10.1186/s40104-017-0153-9
Huang, W., Tan, H., & Nie, S. (2022). Beneficial effects of seaweed-derived dietary fiber: Highlights of the sulfated polysaccharides. Food Chemistry, 373, 131608. https://doi.org/10.1016/j.foodchem.2021.131608
Hwang, E. K., & Park, C. S. (2020). Seaweed cultivation and utilization of Korea. ALGAE, 35(2), 107–121. https://doi.org/10.4490/algae.2020.35.5.15
Iñiguez, C., Galmés, J., & Gordillo, F. J. L. (2019). Rubisco carboxylation kinetics and inorganic carbon utilization in polar versus cold-temperate seaweeds. Journal of Experimental Botany, 70(4), 1283–1297. https://doi.org/10.1093/jxb/ery443
Ismail, M. M., Miranda Lopez, J. M., & El‐Sayed, A. A. M. (2025). Evaluation of the Spatial Variations in the Biochemical Composition of Seaweed Species Along the Coast of Alexandria, With a Focus on Fatty Acids and Total Amino Acids of the Prevalent Edible Species. Food Science & Nutrition, 13(7), e70645. https://doi.org/10.1002/fsn3.70645
Jansen, H. M., Bernard, M. S., Nederlof, M. A. J., Van Der Meer, I. M., & Van Der Werf, A. (2022). Seasonal variation in productivity, chemical composition and nutrient uptake of Ulva spp. (Chlorophyta) strains. Journal of Applied Phycology, 34(3), 1649–1660. https://doi.org/10.1007/s10811-022-02708-z
Jueterbock, A., Hoarau-Heemstra, H., Wigger, K., Duarte, B., Bruckner, C., Chapman, A., Duan, D., Engelen, A., Gauci, C., Hill, G., Hu, Z.-M., Khanal, P., Khatei, A., Mackintosh, A., Meland, H., Melo, R., Nilsen, A. M. L., Olsen, L., Rautenberger, R., … Zhang, J. (2025). Roadmap to sustainably develop the European seaweed industry. Npj Ocean Sustainability, 4(1), 22. https://doi.org/10.1038/s44183-025-00122-9
Juul, L., Danielsen, M., Nebel, C., Steinhagen, S., Bruhn, A., Jensen, S. K., Undeland, I., & Dalsgaard, T. K. (2021). Ulva fenestrata protein – Comparison of three extraction methods with respect to protein yield and protein quality. Algal Research, 60, 102496. https://doi.org/10.1016/j.algal.2021.102496
Juul, L., Haue, S. K., Bruhn, A., Boderskov, T., & Kastrup Dalsgaard, T. (2023). Alkaline pH increases protein extraction yield and solubility of the extracted protein from sugar kelp (Saccharina latissima). Food and Bioproducts Processing, 140, 144–150. https://doi.org/10.1016/j.fbp.2023.05.008
Juul, L., Møller, A. H., Bruhn, A., Jensen, S. K., & Dalsgaard, T. K. (2021). Protein solubility is increased by antioxidant addition during protein extraction from the green macroalga, Ulva sp. Journal of Applied Phycology, 33(1), 545–555. https://doi.org/10.1007/s10811-020-02285-z
Juul, L., Steinhagen, S., Bruhn, A., Jensen, S. K., Undeland, I., & Dalsgaard, T. K. (2022). Combining pressing and alkaline extraction to increase protein yield from Ulva fenestrata biomass. Food and Bioproducts Processing, 134, 80–85. https://doi.org/10.1016/j.fbp.2022.05.006
Juul, L., Nissen, S. H., Bruhn, A., Alexi, N., Jensen, S. K., Hammershøj, M., & Dalsgaard, T. K. (2024). Ulva species: A critical review on the green seaweed as a source of food protein. Trends in Food Science & Technology, 149, 104534. https://doi.org/10.1016/j.tifs.2024.104534
Kadam, S. U., Álvarez, C., Tiwari, B. K., & O’Donnell, C. P. (2017). Extraction and characterization of protein from Irish brown seaweed Ascophyllum nodosum. Food Research International, 99, 1021–1027. https://doi.org/10.1016/j.foodres.2016.07.018
Kasmiati, K., Syahrul, S., Badraeni, B., & Rahmi, M. H. (2022). Proximate and mineral compositions of the green seaweeds Caulerpa lentilifera and Caulerpa racemosa from South Sulawesi Coast, Indonesia. IOP Conference Series: Earth and Environmental Science, 1119(1), 012049. https://doi.org/10.1088/1755-1315/1119/1/012049
Kazir, M., Abuhassira, Y., Robin, A., Nahor, O., Luo, J., Israel, A., Golberg, A., & Livney, Y. D. (2019). Extraction of proteins from two marine macroalgae, Ulva sp. And Gracilaria sp., for food application, and evaluating digestibility, amino acid composition and antioxidant properties of the protein concentrates. Food Hydrocolloids, 87, 194–203. https://doi.org/10.1016/j.foodhyd.2018.07.047
Keeling, P. J. (2010). The endosymbiotic origin, diversification and fate of plastids. Philosophical Transactions of the Royal Society B: Biological Sciences, 365(1541), 729–748. https://doi.org/10.1098/rstb.2009.0103
Kendel, M., Wielgosz-Collin, G., Bertrand, S., Roussakis, C., Bourgougnon, N., & Bedoux, G. (2015). Lipid Composition, Fatty Acids and Sterols in the Seaweeds Ulva armoricana, and Solieria chordalis from Brittany (France): An Analysis from Nutritional, Chemotaxonomic, and Antiproliferative Activity Perspectives. Marine Drugs, 13(9), 5606–5628. https://doi.org/10.3390/md13095606
Khairy, H. M., & El-Shafay, S. M. (2013). Seasonal variations in the biochemical composition of some common seaweed species from the coast of Abu Qir Bay, Alexandria, Egypt. Oceanologia, 55(2), 435–452. https://doi.org/10.5697/oc.55-2.435
Khan, W., Rayirath, U. P., Subramanian, S., Jithesh, M. N., Rayorath, P., Hodges, D. M., Critchley, A. T., Craigie, J. S., Norrie, J., & Prithiviraj, B. (2009). Seaweed Extracts as Biostimulants of Plant Growth and Development. Journal of Plant Growth Regulation, 28(4), 386–399. https://doi.org/10.1007/s00344-009-9103-x
Kjeldahl, J. (1883). Neue Methode zur Bestimmung des Stickstoffs in organischen Körpern. Fresenius’ Zeitschrift für analytische Chemie, 22(1), 366–382. https://doi.org/10.1007/BF01338151
Korea Hydrographic and Oceanographic Agency. (n.d.). Ocean data distribution. KHOA Ocean Grid. Retrieved 18 January 2025, from https://www.khoa.go.kr/oceangrid/gis/category/reference/distribution.do
Kumar, C., Ganesan, P., Suresh, P. V., & Baskar, N. (2008). Seaweeds as a source of nutritionally beneficial compounds—A review.
Kumari, P., Kumar, M., Gupta, V., Reddy, C. R. K., & Jha, B. (2010). Tropical marine macroalgae as potential sources of nutritionally important PUFAs. Food Chemistry, 120(3), 749–757. https://doi.org/10.1016/j.foodchem.2009.11.006
Laing, W., & Christeller, J. (2004). Extraction of Proteins from Plant Tissues. Current Protocols in Protein Science, 38(1). https://doi.org/10.1002/0471140864.ps0407s38
Leliaert, F., Smith, D. R., Moreau, H., Herron, M. D., Verbruggen, H., Delwiche, C. F., & De Clerck, O. (2012). Phylogeny and Molecular Evolution of the Green Algae. Critical Reviews in Plant Sciences, 31(1), 1–46. https://doi.org/10.1080/07352689.2011.615705
Liu, W., Fu, D., Zhang, X., Chai, J., Tian, S., & Han, J. (2019). Development and validation of a new artificial gastric digestive system. Food Research International, 122, 183–190. https://doi.org/10.1016/j.foodres.2019.04.015
Livsmedelsverket. (2023a). The Swedish Food Agency food database (Versions 2023-06-13) [Data set]. https://soknaringsinnehall.livsmedelsverket.se/Home/FoodDetails/1225?sokord=Egg raw&soktyp=1&kategoriId=
Livsmedelsverket. (2023b). The Swedish Food Agency food database (Versions 2023-06-13) [Data set]. https://soknaringsinnehall.livsmedelsverket.se/Home/FoodDetails/887?sokord=soy&soktyp=1&kategoriId=
Lozano Muñoz, I., & Díaz, N. F. (2022). Minerals in edible seaweed: Health benefits and food safety issues. Critical Reviews in Food Science and Nutrition, 62(6), 1592–1607. https://doi.org/10.1080/10408398.2020.1844637
Lund, M. N. (2021). Reactions of plant polyphenols in foods: Impact of molecular structure. Trends in Food Science & Technology, 112, 241–251. https://doi.org/10.1016/j.tifs.2021.03.056
Mabeau, S., & Fleurence, J. (1993). Seaweed in food products: Biochemical and nutritional aspects. Trends in Food Science & Technology, 4(4), 103–107. https://doi.org/10.1016/0924-2244(93)90091-N
Mæhre, H., Jensen, I.-J., & Eilertsen, K.-E. (2016). Enzymatic Pre-Treatment Increases the Protein Bioaccessibility and Extractability in Dulse (Palmaria palmata). Marine Drugs, 14(11), 196. https://doi.org/10.3390/md14110196
Maehre, H. K., Edvinsen, G. K., Eilertsen, K.-E., & Elvevoll, E. O. (2016). Heat treatment increases the protein bioaccessibility in the red seaweed dulse (Palmaria palmata), but not in the brown seaweed winged kelp (Alaria esculenta). Journal of Applied Phycology, 28(1), 581–590. https://doi.org/10.1007/s10811-015-0587-4
Magnusson, M., Glasson, C. R. K., Vucko, M. J., Angell, A., Neoh, T. L., & De Nys, R. (2019). Enrichment processes for the production of high-protein feed from the green seaweed Ulva ohnoi. Algal Research, 41, 101555. https://doi.org/10.1016/j.algal.2019.101555
Makki, K., Deehan, E. C., Walter, J., & Bäckhed, F. (2018). The Impact of Dietary Fiber on Gut Microbiota in Host Health and Disease. Cell Host & Microbe, 23(6), 705–715. https://doi.org/10.1016/j.chom.2018.05.012
Mantri, V. A., Kazi, M. A., Balar, N. B., Gupta, V., & Gajaria, T. (2020). Concise review of green algal genus Ulva Linnaeus. Journal of Applied Phycology, 32(5), 2725–2741. https://doi.org/10.1007/s10811-020-02148-7
Manzanares, P., Gandía, M., Garrigues, S., & Marcos, J. F. (2019). Improving Health-Promoting Effects of Food-Derived Bioactive Peptides through Rational Design and Oral Delivery Strategies. Nutrients, 11(10), 2545. https://doi.org/10.3390/nu11102545
Manzoor, M. F., Afraz, M. T., Yılmaz, B. B., Adil, M., Arshad, N., Goksen, G., Ali, M., & Zeng, X.-A. (2024). Recent progress in natural seaweed pigments: Green extraction, health-promoting activities, techno-functional properties and role in intelligent food packaging. Journal of Agriculture and Food Research, 15, 100991. https://doi.org/10.1016/j.jafr.2024.100991
Marrion, O., Schwertz, A., Fleurence, J., Guéant, J. L., & Villaume, C. (2003). Improvement of the digestibility of the proteins of the red alga Palmaria palmata by physical processes and fermentation. Food / Nahrung, 47(5), 339–344. https://doi.org/10.1002/food.200390078
Marsham, S., Scott, G. W., & Tobin, M. L. (2007). Comparison of nutritive chemistry of a range of temperate seaweeds. Food Chemistry, 100(4), 1331–1336. https://doi.org/10.1016/j.foodchem.2005.11.029
Marze, S. (2017a). Bioavailability of Nutrients and Micronutrients: Advances in Modeling and In Vitro Approaches. Annual Review of Food Science and Technology, 8(1), 35–55. https://doi.org/10.1146/annurev-food-030216-030055
Marze, S. (2017b). Modeling of Food Digestion. In Modeling of Microscale Transport in Biological Processes (pp. 353–374). Elsevier. https://doi.org/10.1016/B978-0-12-804595-4.00014-6
Masuko, T., Minami, A., Iwasaki, N., Majima, T., Nishimura, S.-I., & Lee, Y. C. (2005). Carbohydrate analysis by a phenol–sulfuric acid method in microplate format. Analytical Biochemistry, 339(1), 69–72. https://doi.org/10.1016/j.ab.2004.12.001
Mendez, R. L., & Kwon, J. Y. (2021). Effect of extraction condition on protein recovery and phenolic interference in Pacific dulse (Devaleraea mollis). Journal of Applied Phycology, 33(4), 2497–2509. https://doi.org/10.1007/s10811-021-02467-3
Miftahof, R. N. (2017). Biomechanics of the Human Stomach. Springer International Publishing. https://doi.org/10.1007/978-3-319-59677-8
Milinovic, J., Mata, P., Diniz, M., & Noronha, J. P. (2021). Umami taste in edible seaweeds: The current comprehension and perception. International Journal of Gastronomy and Food Science, 23, 100301. https://doi.org/10.1016/j.ijgfs.2020.100301
Minekus, M., Alminger, M., Alvito, P., Ballance, S., Bohn, T., Bourlieu, C., Carrière, F., Boutrou, R., Corredig, M., Dupont, D., Dufour, C., Egger, L., Golding, M., Karakaya, S., Kirkhus, B., Le Feunteun, S., Lesmes, U., Macierzanka, A., Mackie, A., … Brodkorb, A. (2014). A standardised static in vitro digestion method suitable for food – an international consensus. Food Funct., 5(6), 1113–1124. https://doi.org/10.1039/C3FO60702J
Mishra, A., Vishwakarma, K., Malaviya, P., Kumar, N., Pavón, L. R., Shandilya, C., Sharma, R., Bisht, A., & Takkar, S. (2022). Influence of greenhouse gases on plant epigenomes for food security. In Biomass, Biofuels, Biochemicals (pp. 421–450). Elsevier. https://doi.org/10.1016/B978-0-12-823500-3.00003-0
Miyashita, K., Narayan, B., Tsukui, T., Kamogawa, H., Abe, M., & Hosokawa, M. (2011). Brown Seaweed Lipids as Potential Source of Omega‐3 PUFA in Biological Systems. In S. Kim (Ed.), Handbook of Marine Macroalgae (1st edn, pp. 329–339). Wiley. https://doi.org/10.1002/9781119977087.ch16
Molina-Gilarranz, I., Cebrián-Lloret, V., Recio, I., & Martínez-Sanz, M. (2025). Impact of structure and composition on the digestibility and nutritional quality of alternative protein-rich extracts from the green seaweed Ulva lacinulata. Food Research International, 201, 115646. https://doi.org/10.1016/j.foodres.2024.115646
Montgomery, V. A., Lindsey, C. Y., Smith, L. A., & Webb, R. P. (2021). Development of an o-pthalaldehyde (OPA) assay to measure protein content in Ricin Vaccine E. coli (RVEcTM). Vaccine, 39(3), 564–570. https://doi.org/10.1016/j.vaccine.2020.12.008
Morrissey, J., Kraan, S., & Guiry, M. D. (2001). A guide to commercially important seaweeds on the Irish coast. Irish Bord Iascaigh Mhara/IIrish Sea Fisheries Board.
Mosca, A. C., & Chen, J. (2016). Food oral management: Physiology and objective assessment. Current Opinion in Food Science, 9, 11–20. https://doi.org/10.1016/j.cofs.2016.03.003
Moustafa, Y., & Eladel, H. (2015). Amino acids profile of Ulva sp. Macroalgae collected from Alexandria coast in Egypt: A potential food resource. (1), 15–22.
Narayan, B., Kumar, C. S., Sashima, T., Maeda, H., Hosokawa, M., & Miyashita, K. (2008). Composition, Functionality and Potential Applications of Seaweed Lipids. In C. T. Hou & J. Shaw (Eds), Biocatalysis and Bioenergy (1st edn, pp. 463–490). Wiley. https://doi.org/10.1002/9780470385869.ch26
National Institute of Diabetes and Digestive and Kidney Diseases (NIDDK), National Institutes of Health. (n.d.). https://www.niddk.nih.gov/news/media-library/18260
Neori, A., Msuya, F. E., Shauli, L., Schuenhoff, A., Kopel, F., & Shpigel, M. (2003). A novel three-stage seaweed (Ulva lactuca) biofilter design for integrated mariculture. Journal of Applied Phycology, 15(6), 543–553. https://doi.org/10.1023/B:JAPH.0000004382.89142.2d
Neveux, N., Magnusson, M., Mata, L., Whelan, A., De Nys, R., & Paul, N. A. (2016). The treatment of municipal wastewater by the macroalga Oedogonium sp. And its potential for the production of biocrude. Algal Research, 13, 284–292. https://doi.org/10.1016/j.algal.2015.12.010
North Sea seaweeds: DIP and DIN uptake kinetics and management strategies. (2019). University of Groningen.
O’ Brien, R. O., Hayes, M., Sheldrake, G., Tiwari, B., & Walsh, P. (2022). Macroalgal Proteins: A Review. Foods, 11(4), 571. https://doi.org/10.3390/foods11040571
O’ Connor, J., Meaney, S., Williams, G. A., & Hayes, M. (2020). Extraction of Protein from Four Different Seaweeds Using Three Different Physical Pre-Treatment Strategies. Molecules, 25(8), 2005. https://doi.org/10.3390/molecules25082005
OECD. (2025). OECD Review of Fisheries 2025. OECD Publishing. https://doi.org/10.1787/560cd8fc-en
Oliveira, M. N. D., Freitas, A. L. P., Carvalho, A. F. U., Sampaio, T. M. T., Farias, D. F., Alves Teixeira, D. I., Gouveia, S. T., Pereira, J. G., & Sena, M. M. D. C. C. D. (2009). Nutritive and non-nutritive attributes of washed-up seaweeds from the coast of Ceará, Brazil. Food Chemistry, 115(1), 254–259. https://doi.org/10.1016/j.foodchem.2008.12.004
Olsen, J. V., Ong, S.-E., & Mann, M. (2004). Trypsin Cleaves Exclusively C-terminal to Arginine and Lysine Residues. Molecular & Cellular Proteomics, 3(6), 608–614. https://doi.org/10.1074/mcp.T400003-MCP200
Paiva, L., Lima, E., Neto, A. I., & Baptista, J. (2016). Isolation and characterization of angiotensin I-converting enzyme (ACE) inhibitory peptides from Ulva rigida C. Agardh protein hydrolysate. Journal of Functional Foods, 26, 65–76. https://doi.org/10.1016/j.jff.2016.07.006
Pan, S., Wang, S., Jing, L., & Yao, D. (2016). Purification and characterisation of a novel angiotensin-I converting enzyme (ACE)-inhibitory peptide derived from the enzymatic hydrolysate of Enteromorpha clathrata protein. Food Chemistry, 211, 423–430. https://doi.org/10.1016/j.foodchem.2016.05.087
Peasura, N., Laohakunjit, N., Kerdchoechuen, O., & Wanlapa, S. (2015). Characteristics and antioxidant of Ulva intestinalis sulphated polysaccharides extracted with different solvents. International Journal of Biological Macromolecules, 81, 912–919. https://doi.org/10.1016/j.ijbiomac.2015.09.030
Peñalver, R., Lorenzo, J. M., Ros, G., Amarowicz, R., Pateiro, M., & Nieto, G. (2020). Seaweeds as a Functional Ingredient for a Healthy Diet. Marine Drugs, 18(6), 301. https://doi.org/10.3390/md18060301
Pereira, L., Cotas, J., & Gonçalves, A. (2024). Seaweed Proteins: A Step towards Sustainability? Nutrients, 16(8), 1123. https://doi.org/10.3390/nu16081123
Pimentel, F. B., Cermeño, M., Kleekayai, T., Harnedy, P. A., FitzGerald, R. J., Alves, R. C., & Oliveira, M. B. P. P. (2020). Effect of in vitro simulated gastrointestinal digestion on the antioxidant activity of the red seaweed Porphyra dioica. Food Research International, 136, 109309. https://doi.org/10.1016/j.foodres.2020.109309
Politiek, R. G. A., Bruins, M. E., Keppler, J. K., & Schutyser, M. A. I. (2022). Effect of oil content on pin-milling of soybean. Journal of Food Engineering, 334, 111149. https://doi.org/10.1016/j.jfoodeng.2022.111149
Pootthachaya, P., Puangsap, W., Bunchalee, P., Plangklang, P., Reungsang, A., Yuangsoi, B., Cherdthong, A., Tengjaroenkul, B., & Wongtangtintharn, S. (2023). Investigation of nutritional profile, protein solubility and in vitro digestibility of various algae species as an alternative protein source for poultry feed. Algal Research, 72, 103147. https://doi.org/10.1016/j.algal.2023.103147
Porse, H., & Rudolph, B. (2017). The seaweed hydrocolloid industry: 2016 updates, requirements, and outlook. Journal of Applied Phycology, 29(5), 2187–2200. https://doi.org/10.1007/s10811-017-1144-0
Postma, P. R., Cerezo-Chinarro, O., Akkerman, R. J., Olivieri, G., Wijffels, R. H., Brandenburg, W. A., & Eppink, M. H. M. (2018). Biorefinery of the macroalgae Ulva lactuca: Extraction of proteins and carbohydrates by mild disintegration. Journal of Applied Phycology, 30(2), 1281–1293. https://doi.org/10.1007/s10811-017-1319-8
Prabhu, M., Chemodanov, A., Gottlieb, R., Kazir, M., Nahor, O., Gozin, M., Israel, A., Livney, Y. D., & Golberg, A. (2019). Starch from the sea: The green macroalga Ulva ohnoi as a potential source for sustainable starch production in the marine biorefinery. Algal Research, 37, 215–227. https://doi.org/10.1016/j.algal.2018.11.007
Prakash Nirmal, N., Singh Rajput, M., Bhojraj Rathod, N., Mudgil, P., Pati, S., Bono, G., Nalinanon, S., Li, L., & Maqsood, S. (2023). Structural characteristic and molecular docking simulation of fish protein-derived peptides: Recent updates on antioxidant, anti-hypertensive and anti-diabetic peptides. Food Chemistry, 405, 134737. https://doi.org/10.1016/j.foodchem.2022.134737
Pujiastuti, D. Y., Ghoyatul Amin, M. N., Alamsjah, M. A., & Hsu, J.-L. (2019). Marine Organisms as Potential Sources of Bioactive Peptides that Inhibit the Activity of Angiotensin I-Converting Enzyme: A Review. Molecules, 24(14), 2541. https://doi.org/10.3390/molecules24142541
Qiao, K., Zhou, X., Wu, H., Zhang, L., Liang, L., & Zhang, Y. (2025). Research on the release and absorption regularities of free amino acids and peptides in vitro digestion of yeast protein. Food Chemistry, 482, 144176. https://doi.org/10.1016/j.foodchem.2025.144176
Rajpurohit, B., & Li, Y. (2023). Overview on pulse proteins for future foods: Ingredient development and novel applications. Journal of Future Foods, 3(4), 340–356. https://doi.org/10.1016/j.jfutfo.2023.03.005
Ratana-Arporn, P., & Chirapart, A. (2006). Nutritional evaluation of tropical green seaweeds Caulerpa lentillifera and Ulva reticulata.75–83.
Reith, J. H., Deurwaarder, E. P., Hemmes, K., Curvers, A. P. W. M., Kamermans, P., Brandenburg, W., & Zeeman, G. (2005). Bio-offshore. Grootschalige teelt van zeewieren in combinatie met offshore windparken in de Noordzee. https://www.ecn.nl/docs/library/report/2005/c05008.pdf
Robin, A., Kazir, M., Sack, M., Israel, A., Frey, W., Mueller, G., Livney, Y. D., & Golberg, A. (2018). Functional Protein Concentrates Extracted from the Green Marine Macroalga Ulva sp., by High Voltage Pulsed Electric Fields and Mechanical Press. ACS Sustainable Chemistry & Engineering, 6(11), 13696–13705. https://doi.org/10.1021/acssuschemeng.8b01089
Rogers, K. (2011). The Digestive System. Britannica Educational Publishing.
Ruperez, P. (2002). Mineral content of edible marine seaweeds. Food Chemistry, 79(1), 23–26. https://doi.org/10.1016/S0308-8146(02)00171-1
Saladin, K. S. (2017). Human Anatomy (5th edn). McGraw-Hill Education.
Sánchez, A., & Vázquez, A. (2017). Bioactive peptides: A review. Food Quality and Safety, 1(1), 29–46. https://doi.org/10.1093/fqs/fyx006
Schiener, P., Black, K. D., Stanley, M. S., & Green, D. H. (2015). The seasonal variation in the chemical composition of the kelp species Laminaria digitata, Laminaria hyperborea, Saccharina latissima and Alaria esculenta. Journal of Applied Phycology, 27(1), 363–373. https://doi.org/10.1007/s10811-014-0327-1
Semmouri, I., Lee, H., Alahyaribeik, S., Baldominos Botas, J. M., Claeys, P., Han, T., Janssen, C. R., Mirzaei, M., Asselman, J., & Park, J. (2025). The effect of light, aeration and temperature on the growth and phenolic content of cultivated Ecklonia cava gametophytes (Phaeophyceae) and their antioxidant properties. Journal of Applied Phycology, 37(6), 4395–4411. https://doi.org/10.1007/s10811-025-03649-z
Semmouri, I., Vanhercke, S., Ferreira, A. S., Knoop, J., De Clerck, O., Nunes, C., Coimbra, M. A., De Souza Mesquita, L. M., Ventura, S. P. M., Janssen, C. R., & Asselman, J. (2025). The Effect of High Temperatures on the Growth and Biochemical Composition of Cultivated Palmaria Palmata (Rhodophyta). Marine Biotechnology, 27(6), 167. https://doi.org/10.1007/s10126-025-10548-4
Sensoy, I. (2021). A review on the food digestion in the digestive tract and the used in vitro models. Current Research in Food Science, 4, 308–319. https://doi.org/10.1016/j.crfs.2021.04.004
Seracu, D. I. (1987). The Study of UV and VIS Absorption Spectra of the Complexes of Amino Acids with Ninhydrin. Analytical Letters, 20(9), 1417–1428. https://doi.org/10.1080/00032718708066323
Shannon, E., & Abu-Ghannam, N. (2019). Seaweeds as nutraceuticals for health and nutrition. Phycologia, 58(5), 563–577. https://doi.org/10.1080/00318884.2019.1640533
Shi, L., Zhao, W., Yang, Z., Subbiah, V., & Suleria, H. A. R. (2022). Extraction and characterization of phenolic compounds and their potential antioxidant activities. Environmental Science and Pollution Research, 29(54), 81112–81129. https://doi.org/10.1007/s11356-022-23337-6
Smith, M. E., & Morton, D. G. (2010). Systems of The Body. The Digestive System. Basic Science and Clinical Conditions (2nd edn). Elsevier Limited.
Soufi, J., El Hammoudani, Y., Haboubi, K., Hanafi, I., & Dimane, F. (2024). Ulva spp (Ulva intestinalis, U. fasciata, U. lactuca, and U. rigida) composition and abiotic environmental factors. BIO Web of Conferences, 109, 01012. https://doi.org/10.1051/bioconf/202410901012
Sousa, A. I., Martins, I., Lillebø, A. I., Flindt, M. R., & Pardal, M. A. (2007). Influence of salinity, nutrients and light on the germination and growth of Enteromorpha sp. Spores. Journal of Experimental Marine Biology and Ecology, 341(1), 142–150. https://doi.org/10.1016/j.jembe.2006.09.020
Souto-Prieto, A., Cobos, A., Ferreiro, T., Castillo, Z., Díaz-Piñero, L., Martínez-Sanz, M., & Lopez-Sanchez, P. (2025). Evolution of the structure and rheology of processed seaweed Ulva spp. During in vitro digestion. Npj Science of Food, 9(1), 282. https://doi.org/10.1038/s41538-025-00639-w
Steinbruch, E., Wise, J., Levkov, K., Chemodanov, A., Israel, Á., Livney, Y. D., & Golberg, A. (2023). Enzymatic cell wall degradation combined with pulsed electric fields increases yields of water-soluble-protein extraction from the green marine macroalga Ulva sp. Innovative Food Science & Emerging Technologies, 84, 103231. https://doi.org/10.1016/j.ifset.2022.103231
Steinhagen, S., Enge, S., Cervin, G., Larsson, K., Edlund, U., Schmidt, A. E. M., Wahlström, N., Kollander, B., Pavia, H., Undeland, I., & Toth, G. B. (2022). Harvest Time Can Affect the Optimal Yield and Quality of Sea Lettuce (Ulva fenestrata) in a Sustainable Sea-Based Cultivation. Frontiers in Marine Science, 9, 816890. https://doi.org/10.3389/fmars.2022.816890
Steinhagen, S., Enge, S., Larsson, K., Olsson, J., Nylund, G. M., Albers, E., Pavia, H., Undeland, I., & Toth, G. B. (2021). Sustainable Large-Scale Aquaculture of the Northern Hemisphere Sea Lettuce, Ulva fenestrata, in an Off-Shore Seafarm. Journal of Marine Science and Engineering, 9(6), 615. https://doi.org/10.3390/jmse9060615
Stévant, P., Marfaing, H., Rustad, T., Sandbakken, I., Fleurence, J., & Chapman, A. (2017). Nutritional value of the kelps Alaria esculenta and Saccharina latissima and effects of short-term storage on biomass quality. Journal of Applied Phycology, 29(5), 2417–2426. https://doi.org/10.1007/s10811-017-1126-2
Sullivan, R. (2009). The Human Body, How It Works, Digestion and Nutrition. Chelsea House Publications.
Sun, S., Xu, X., Sun, X., Zhang, X., Chen, X., & Xu, N. (2019). Preparation and Identification of ACE Inhibitory Peptides from the Marine Macroalga Ulva intestinalis. Marine Drugs, 17(3), 179. https://doi.org/10.3390/md17030179
Tagliazucchi, D., Martini, S., & Solieri, L. (2019). Bioprospecting for Bioactive Peptide Production by Lactic Acid Bacteria Isolated from Fermented Dairy Food. Fermentation, 5(4), 96. https://doi.org/10.3390/fermentation5040096
Tamayo Tenorio, A., Kyriakopoulou, K. E., Suarez-Garcia, E., Van Den Berg, C., & Van Der Goot, A. J. (2018). Understanding differences in protein fractionation from conventional crops, and herbaceous and aquatic biomass—Consequences for industrial use. Trends in Food Science & Technology, 71, 235–245. https://doi.org/10.1016/j.tifs.2017.11.010
Tanna, B., & Mishra, A. (2019). Nutraceutical Potential of Seaweed Polysaccharides: Structure, Bioactivity, Safety, and Toxicity. Comprehensive Reviews in Food Science and Food Safety, 18(3), 817–831. https://doi.org/10.1111/1541-4337.12441
Tibbetts, S. M., Milley, J. E., & Lall, S. P. (2016). Nutritional quality of some wild and cultivated seaweeds: Nutrient composition, total phenolic content and in vitro digestibility. Journal of Applied Phycology, 28(6), 3575–3585. https://doi.org/10.1007/s10811-016-0863-y
Toth, G. B., Harrysson, H., Wahlström, N., Olsson, J., Oerbekke, A., Steinhagen, S., Kinnby, A., White, J., Albers, E., Edlund, U., Undeland, I., & Pavia, H. (2020). Effects of irradiance, temperature, nutrients, and pCO2 on the growth and biochemical composition of cultivated Ulva fenestrata. Journal of Applied Phycology, 32(5), 3243–3254. https://doi.org/10.1007/s10811-020-02155-8
Trigo, J. (2023). Seaweed as a sustainable source of food protein: Maximizing seaweed protein content, protein recovery and nutritional quality [Doctoral disseration]. Chalmers University of Technology.
Trigo, J. P., Engström, N., Steinhagen, S., Juul, L., Harrysson, H., Toth, G. B., Pavia, H., Scheers, N., & Undeland, I. (2021). In vitro digestibility and Caco-2 cell bioavailability of sea lettuce (Ulva fenestrata) proteins extracted using pH-shift processing. Food Chemistry, 356, 129683. https://doi.org/10.1016/j.foodchem.2021.129683
Umen, J. G. (2014). Green Algae and the Origins of Multicellularity in the Plant Kingdom. Cold Spring Harbor Perspectives in Biology, 6(11), a016170–a016170. https://doi.org/10.1101/cshperspect.a016170
United Nations. (2017). World Population Prospects: The 2017 Revision. United Nations Department of Economic and Social Affairs, Population Division.
USDA. (2018a). USDA National Nutrient Database for Standard Reference [Data set]. https://fdc.nal.usda.gov/fdc-app.html#/food-details/174270/nutrients
USDA. (2018b). USDA National Nutrient Database for Standard Reference [Data set]. https://fdc.nal.usda.gov/fdc-app.html#/food-details/174030/nutrients
Van Der Molen, J., Ruardij, P., Mooney, K., Kerrison, P., O’Connor, N. E., Gorman, E., Timmermans, K., Wright, S., Kelly, M., Hughes, A. D., & Capuzzo, E. (2018). Modelling potential production of macroalgae farms in UK and Dutch coastal waters. Biogeosciences, 15(4), 1123–1147. https://doi.org/10.5194/bg-15-1123-2018
Vásquez, V., Martínez, R., & Bernal, C. (2019). Enzyme-assisted extraction of proteins from the seaweeds Macrocystis pyrifera and Chondracanthus chamissoi: Characterization of the extracts and their bioactive potential. Journal of Applied Phycology, 31(3), 1999–2010. https://doi.org/10.1007/s10811-018-1712-y
Vega-Gómez, L. M., Molina-Gilarranz, I., Fontes-Candia, C., Cebrián-Lloret, V., Recio, I., & Martínez-Sanz, M. (2024). Production of hybrid protein-polysaccharide extracts from Ulva spp. Seaweed with potential as food ingredients. Food Hydrocolloids, 153, 110046. https://doi.org/10.1016/j.foodhyd.2024.110046
Vega-Rojas, L. J., Luzardo-Ocampo, I., Mosqueda, J., Palmerín-Carreño, D. M., Escobedo-Reyes, A., Blanco-Labra, A., Escobar-García, K., & García-Gasca, T. (2021). Bioaccessibility and In Vitro Intestinal Permeability of a Recombinant Lectin from Tepary Bean (Phaseolus acutifolius) Using the Everted Intestine Assay. International Journal of Molecular Sciences, 22(3), 1049. https://doi.org/10.3390/ijms22031049
Vilg, J. V., & Undeland, I. (2017). pH-driven solubilization and isoelectric precipitation of proteins from the brown seaweed Saccharina latissima—Effects of osmotic shock, water volume and temperature. Journal of Applied Phycology, 29(1), 585–593. https://doi.org/10.1007/s10811-016-0957-6
Vizcaíno, A. J., Galafat, A., Sáez, M. I., Martínez, T. F., & Alarcón, F. J. (2020). Partial Characterization of Protease Inhibitors of Ulva ohnoi and Their Effect on Digestive Proteases of Marine Fish. Marine Drugs, 18(6), 319. https://doi.org/10.3390/md18060319
Walker, J. M. (1994). The Bicinchoninic Acid (BCA) Assay for Protein Quantitation. In J. M. Walker, Basic Protein and Peptide Protocols (Vol. 32, pp. 5–8). Humana Press. https://doi.org/10.1385/0-89603-268-X:5
Wei, N., Quarterman, J., & Jin, Y.-S. (2013). Marine macroalgae: An untapped resource for producing fuels and chemicals. Trends in Biotechnology, 31(2), 70–77. https://doi.org/10.1016/j.tibtech.2012.10.009
Wells, M. L., Potin, P., Craigie, J. S., Raven, J. A., Merchant, S. S., Helliwell, K. E., Smith, A. G., Camire, M. E., & Brawley, S. H. (2017). Algae as nutritional and functional food sources: Revisiting our understanding. Journal of Applied Phycology, 29(2), 949–982. https://doi.org/10.1007/s10811-016-0974-5
WHO/FAO/UNU. (2007). Protein and Amino Acid Requirements in Human Nutrition. WHO Technical Report Series.
Wijers, T., Hylkema, A., Visser, T., & Timmermans, K. (2020). Effects of preservation on protein extraction in four seaweed species. Journal of Applied Phycology, 32(5), 3401–3409. https://doi.org/10.1007/s10811-020-02197-y
Wong, K. H., & Cheung, P. C. K. (2001). Nutritional evaluation of some subtropical red and green seaweeds Part II. In vitro protein digestibility and amino acid profiles of protein concentrates. Food Chemistry, 72(1), 11–17. https://doi.org/10.1016/S0308-8146(00)00176-X
Xu, J., Liao, W., Liu, Y., Guo, Y., Jiang, S., & Zhao, C. (2023). An overview on the nutritional and bioactive components of green seaweeds. Food Production, Processing and Nutrition, 5(1), 18. https://doi.org/10.1186/s43014-023-00132-5
Yazawa, K., & Numata, K. (2014). Recent Advances in Chemoenzymatic Peptide Syntheses. Molecules, 19(9), 13755–13774. https://doi.org/10.3390/molecules190913755
Zhang, X., Cao, D., Sun, X., Sun, S., & Xu, N. (2019). Preparation and identification of antioxidant peptides from protein hydrolysate of marine alga Gracilariopsis lemaneiformis. Journal of Applied Phycology, 31(4), 2585–2596. https://doi.org/10.1007/s10811-019-1746-9
Zhong, H., Gao, X., Cheng, C., Liu, C., Wang, Q., & Han, X. (2020). The Structural Characteristics of Seaweed Polysaccharides and Their Application in Gel Drug Delivery Systems. Marine Drugs, 18(12), 658. https://doi.org/10.3390/md18120658