Structural properties and biotechnological applications of cyanobacterial phycobiliproteins

Sapana Jha; Varsha K. Singh; Payel Rana; Ashish P. Singh; Amit Gupta; Palak Rana; Riya Tripathi; Rajeshwar P. Sinha; Sapana Jha; Varsha K. Singh; Payel Rana; Ashish P. Singh; Amit Gupta; Palak Rana; Riya Tripathi; Rajeshwar P. Sinha

doi:10.3934/molsci.2026007

AIMS Molecular Science

2026, Volume 13, Issue 2: 119-142. doi: 10.3934/molsci.2026007

Previous Article Next Article

Review Special Issues

Structural properties and biotechnological applications of cyanobacterial phycobiliproteins

Laboratory of Photobiology and Molecular Microbiology, Department of Botany, Institute of Science, Banaras Hindu University, Varanasi-221005, India

Received: 13 October 2025 Revised: 10 February 2026 Accepted: 24 February 2026 Published: 19 March 2026

Cyanobacteria are a promising and sustainable source of numerous bioactive compounds, including phycobiliproteins (PBPs). PBPs are a group of water-soluble pigments that serve as a critical component in the accessory light-harvesting system in cyanobacteria. PBPs are composed of chromophore-binding proteins such as phycocyanin, phycoerythrin, and allophycocyanin, which exhibit distinctive structural and functional properties. PBPs aggregate into larger structures called phycobilisomes, located on the thylakoid membrane of cyanobacteria, consisting of rod and core subunits. PBPs scavenge free radicals and reduce oxidative stress, which plays a pivotal role in the pathogenesis of numerous chronic diseases such as cancer, diabetes, cardiovascular disorders, and neurodegenerative conditions. Due to their pharmacological properties, outstanding therapeutic value, and ecological significance, they have garnered considerable scientific interest in recent years. In this review, we provide an in-depth examination of the structural attributes of PBPs and explore their promising biotechnological applications, offering insights into current developments, challenges, and future directions for utilizing PBPs in scientific and industrial fields.
- bioactive compounds,
- cyanobacteria,
- microalgae,
- phycocyanin,
- phycoerythrin
Citation: Sapana Jha, Varsha K. Singh, Payel Rana, Ashish P. Singh, Amit Gupta, Palak Rana, Riya Tripathi, Rajeshwar P. Sinha. Structural properties and biotechnological applications of cyanobacterial phycobiliproteins[J]. AIMS Molecular Science, 2026, 13(2): 119-142. doi: 10.3934/molsci.2026007

Related Papers:

Abstract

Cyanobacteria are a promising and sustainable source of numerous bioactive compounds, including phycobiliproteins (PBPs). PBPs are a group of water-soluble pigments that serve as a critical component in the accessory light-harvesting system in cyanobacteria. PBPs are composed of chromophore-binding proteins such as phycocyanin, phycoerythrin, and allophycocyanin, which exhibit distinctive structural and functional properties. PBPs aggregate into larger structures called phycobilisomes, located on the thylakoid membrane of cyanobacteria, consisting of rod and core subunits. PBPs scavenge free radicals and reduce oxidative stress, which plays a pivotal role in the pathogenesis of numerous chronic diseases such as cancer, diabetes, cardiovascular disorders, and neurodegenerative conditions. Due to their pharmacological properties, outstanding therapeutic value, and ecological significance, they have garnered considerable scientific interest in recent years. In this review, we provide an in-depth examination of the structural attributes of PBPs and explore their promising biotechnological applications, offering insights into current developments, challenges, and future directions for utilizing PBPs in scientific and industrial fields.

Acknowledgments

Sapana Jha (No. R/Dev. /Sch./ UGC Non-NET Fello./2022-23/52561) is thankful to BHU for providing institutional fellowship. Varsha K. Singh (09/0013(12862)/2021- EMR-1), Amit Gupta (09/0013(0912)/2019-EMR-I), and Palak Rana (09/0013 (16603)/2023-EMR-I) are thankful to the Council of Scientific and Industrial Research (CSIR), New Delhi, India, for the Senior Research Fellowship (SRF). Payel Rana (09/0013(21806)/2025-EMR-I) is thankful to the CSIR, New Delhi, India, for financial assistance in the form of Junior Research Fellowship (JRF). Ashish P. Singh (NTA Ref. No. 191620014505) and Riya Tripathi (NTA Ref. No. 231620041285) are thankful to the University Grants Commission (UGC), New Delhi, India, for the financial assistance in the form of a Senior Research Fellowship (SRF) and Junior Research Fellowship (JRF), respectively. The incentive grant received from IoE (Scheme no. 6031), Banaras Hindu University, Varanasi, India, to Rajeshwar Sinha is highly acknowledged.

Conflict of interest

The authors declare no conflicts of interest in this paper.

References

[1]	Li W, Su HN, Pu Y, et al. (2019) Phycobiliproteins: Molecular structure, production, applications, and prospects. Biotechnol Adv 37: 340-353. https://doi.org/10.1016/j.biotechadv.2019.01.008
[2]	Costa ML, Rodrigues JA, Azevedo J, et al. (2018) Hepatotoxicity induced by paclitaxel interaction with turmeric in association with a microcystin from a contaminated dietary supplement. Toxicon 150: 207-211. https://doi.org/10.1016/j.toxicon.2018.05.022
[3]	Regueiras A, Pereira S, Costa MS, et al. (2018) Differential toxicity of cyanobacteria isolated from marine sponges towards echinoderms and crustaceans. Toxins 10: 297. https://doi.org/10.3390/toxins10070297
[4]	Guan X, Wang J, Zhu J, et al. (2013) Photosystem II photochemistry and phycobiliprotein of the red algae Kappaphycus alvarezii and their implications for light adaptation. BioMed Res Int 2013: 256549. https://doi.org/10.1155/2013/256549
[5]	Zhang J, Ma J, Liu D, et al. (2017) Structure of phycobilisome from the red alga Griffithsia pacifica. Nature 551: 57-63. https://doi.org/10.1038/nature24278
[6]	Bryant DA, Gisriel CJ (2024) The structural basis for light harvesting in organisms producing phycobiliproteins. Plant Cell 36: 4036-4064. https://doi.org/10.1093/plcell/koae126
[7]	Jha S, Singh VK, Singh AP, et al. (2024) The radiant world of cyanobacterial phycobiliproteins: Examining their structure, functions, and biomedical potentials. Targets 2: 32-51. https://doi.org/10.3390/targets2010002
[8]	Singh VK, Jha S, Rana P, et al. (2023) Resilience and mitigation strategies of cyanobacteria under ultraviolet radiation stress. Int J Mol Sci 24: 12381. https://doi.org/10.3390/ijms241512381
[9]	Manirafasha E, Guo L, Jing K (2020) Nutraceutical and pharmaceutical applications of phycobiliproteins. Pigments from microalgae handbook . Cham: Springer 575-584. https://doi.org/10.1007/978-3-030-50971-2_23
[10]	Bryant DA, Guglielmi G, de Marsac NT, et al. (1979) The structure of cyanobacterial phycobilisomes: A model. Arch Microbiol 123: 113-127. https://doi.org/10.1007/BF00446810
[11]	Pagels F, Guedes AC, Amaro HM, et al. (2019) Phycobiliproteins from cyanobacteria: Chemistry and biotechnological applications. Biotechnol Adv 37: 422-443. https://doi.org/10.1016/j.biotechadv.2019.02.010
[12]	Manirafasha E, Ndikubwimana T, Zeng X, et al. (2016) Phycobiliprotein: Potential microalgae derived pharmaceutical and biological reagent. Biochem Eng J 109: 282-296. https://doi.org/10.1016/j.bej.2016.01.025
[13]	Judson OP (2017) The energy expansions of evolution. Nat Ecol Evol 1: 0138. https://doi.org/10.1038/s41559-017-0138
[14]	McKay CP (2016) Water sources for cyanobacteria below desert rocks in the Negev Desert determined by conductivity. Glob Ecol Conserv 6: 145-151. https://doi.org/10.1016/j.gecco.2016.02.010
[15]	Zheng L, Zheng Z, Li X, et al. (2021) Structural insight into the mechanism of energy transfer in cyanobacterial phycobilisomes. Nat Commun 12: 5497. https://doi.org/10.1038/s41467-021-25813-y
[16]	Kawakami K, Hamaguchi T, Hirose Y, et al. (2022) Core and rod structures of a thermophilic cyanobacterial light-harvesting phycobilisome. Nat Commun 13: 3389. https://doi.org/10.1038/s41467-022-30962-9
[17]	Zhang W, Zhou J, Zhang Y, et al. (2025) Metabolic engineering of a high-light tolerant cyanobacterium Synechocystis sp. PCC 6803 for efficient biosynthesis of β-farnesene from CO₂. Syst Microbiol Biomanufact 5: 1464-1474. https://doi.org/10.1007/s43393-025-00391-y
[18]	Aryee AN, Agyei D, Akanbi TO (2018) Recovery and utilization of seaweed pigments in food processing. Curr Opin Food Sci 19: 113-119. https://doi.org/10.1016/j.cofs.2018.03.013
[19]	Chen H, Qi H, Xiong P (2022) Phycobiliproteins—A family of algae-derived biliproteins: productions, characterization and pharmaceutical potentials. Mar Drugs 20: 450. https://doi.org/10.3390/md20070450
[20]	Gan F, Bryant DA (2015) Adaptive and acclimative responses of cyanobacteria to far-red light. Environ Microbiol 17: 3450-3465. https://doi.org/10.1111/1462-2920.12992
[21]	Li Y, Lin Y, Garvey CJ, et al. (2016) Characterization of red-shifted phycobilisomes isolated from the chlorophyll f-containing cyanobacterium Halomicronema hongdechloris. BBA-Bioenergetics 1857: 107-114. https://doi.org/10.1016/j.bbabio.2015.10.009
[22]	Ashaolu TJ, Samborska K, Lee CC, et al. (2021) Phycocyanin, a super functional ingredient from algae; properties, purification characterization, and applications. Int J Biol Macromol 193: 2320-2331. https://doi.org/10.1016/j.ijbiomac.2021.11.064
[23]	Kehoe DM (2010) Chromatic adaptation and the evolution of light color sensing in cyanobacteria. Proc Natl Acad Sci USA 107: 9029-9030. https://doi.org/10.1073/pnas.1004510107
[24]	Sonani RR, Patel S, Bhastana B, et al. (2017) Purification and antioxidant activity of phycocyanin from Synechococcus sp. R42DM isolated from industrially polluted site. Bioresource Technol 245: 325-331. https://doi.org/10.1016/j.biortech.2017.08.129
[25]	Chaubey MG, Patel SN, Rastogi RP, et al. (2020) Cyanobacterial pigment protein allophycocyanin exhibits longevity and reduces Aβ-mediated paralysis in C. elegans: Complicity of FOXO and NRF2 ortholog DAF-16 and SKN-1. 3 Biotech 10: 332. https://doi.org/10.1007/s13205-020-02314-1
[26]	Braune S, Krüger-Genge A, Kammerer S, et al. (2021) Phycocyanin from Arthrospira platensis as potential anti-cancer drug: Review of in vitro and in vivo studies. Life 11: 91. https://doi.org/10.3390/life11020091
[27]	Dagnino-Leone J, Figueroa CP, Castañeda ML, et al. (2022) Phycobiliproteins: Structural aspects, functional characteristics, and biotechnological perspectives. Comput Struct Biotec J 20: 1506-1527. https://doi.org/10.1016/j.csbj.2022.02.016
[28]	Zheng J, Inoguchi T, Sasaki S, et al. (2012) Phycocyanin and phycocyanobilin from Spirulina platensis protect against diabetic nephropathy by inhibiting oxidative stress. Am J Physiol-Reg I 304: R110-R120. https://doi.org/10.1152/ajpregu.00648.2011
[29]	Minic SL, Stanic-Vucinic D, Mihailovic J, et al. (2016) Digestion by pepsin releases biologically active chromopeptides from C-phycocyanin, a blue-colored biliprotein of microalga Spirulina. J Proteomics 147: 132-139. https://doi.org/10.1016/j.jprot.2016.03.043
[30]	Liu J, Bai X, Fu P (2022) In silico and in vitro assessment of bioactive peptides from Arthrospira platensis phycobiliproteins for DPP-IV inhibitory activity, ACE inhibitory activity, and antioxidant activity. J Appl Phycol 34: 1497-1511. https://doi.org/10.1007/s10811-022-02732-z
[31]	Tounsi L, Ben Hlima H, Hentati F, et al. (2023) Microalgae: A promising source of bioactive phycobiliproteins. Mar Drugs 21: 440. https://doi.org/10.3390/md21080440
[32]	Munawaroh HSH, Gumilar GG, Nurjanah F, et al. (2020) In-vitro molecular docking analysis of microalgae extracted phycocyanin as an anti-diabetic candidate. Biochem Eng J 161: 107666. https://doi.org/10.1016/j.bej.2020.107666
[33]	Hao S, Li F, Li Q, et al. (2022) Phycocyanin protects against high glucose high fat diet induced diabetes in mice and participates in AKT and AMPK signaling. Foods 11: 3183. https://doi.org/10.3390/foods11203183
[34]	Husain A, Alouffi S, Khanam A, et al. (2022) Therapeutic efficacy of natural product ‘C-phycocyanin’ in alleviating streptozotocin-induced diabetes via the inhibition of glycation reaction in rats. Int J Mol Sci 23: 14235. https://doi.org/10.3390/ijms232214235
[35]	Ren Z, Xie Z, Cao D, et al. (2018) C-phycocyanin inhibits hepatic gluconeogenesis and increases glycogen synthesis via activating Akt and AMPK in insulin resistance hepatocytes. Food Funct 9: 2829-2839. https://doi.org/10.1039/C8FO00257F
[36]	Ziyaei K, Abdi F, Mokhtari M, et al. (2023) Phycocyanin as a nature-inspired antidiabetic agent: A systematic review. Phytomedicine 119: 154964. https://doi.org/10.1016/j.phymed.2023.154964
[37]	Ou Y, Ren Z, Wang J, et al. (2016) Phycocyanin ameliorates alloxan-induced diabetes mellitus in mice: Involved in insulin signaling pathway and GK expression. Chem Biol Interact 247: 49-54. https://doi.org/10.1016/j.cbi.2016.01.018
[38]	Gao Y, Liu C, Wan G, et al. (2016) Phycocyanin prevents methylglyoxal-induced mitochondrial-dependent apoptosis in INS-1 cells by Nrf2. Food Funct 7: 1129-1137. https://doi.org/10.1039/C5FO01548K
[39]	Li Y, Aiello G, Bollati C, et al. (2020) Phycobiliproteins from Arthrospira platensis (Spirulina): A new source of peptides with dipeptidyl peptidase-IV inhibitory activity. Nutrients 12: 794. https://doi.org/10.3390/nu12030794
[40]	Villaró S, Jiménez-Márquez S, Musari E, et al. (2023) Production of enzymatic hydrolysates with in vitro antioxidant, antihypertensive, and antidiabetic properties from proteins derived from Arthrospira platensis. Food Res Int 163: 112270. https://doi.org/10.1016/j.foodres.2022.112270
[41]	Rimbau V, Camins A, Pubill D, et al. (2001) C-phycocyanin protects cerebellar granule cells from low potassium/serum deprivation-induced apoptosis. Naunyn-Schmied Arch Pharmacol 364: 96-104. https://doi.org/10.1007/s002100100437
[42]	Pentón-Rol G, Martínez-Sánchez G, Cervantes-Llanos M, et al. (2011) C-phycocyanin ameliorates experimental autoimmune encephalomyelitis and induces regulatory T cells. Int Immunopharmacol 11: 29-38. https://doi.org/10.1016/j.intimp.2010.10.001
[43]	Marín-Prida J, Pentón-Rol G, Rodrigues FP, et al. (2012) C-phycocyanin protects SH-SY5Y cells from oxidative injury, rat retina from transient ischemia and rat brain mitochondria from Ca²⁺/phosphate-induced impairment. Brain Res Bull 89: 159-167. https://doi.org/10.1016/j.brainresbull.2012.08.011
[44]	Ou Y, Zheng S, Lin L, et al. (2010) Protective effect of C-phycocyanin against carbon tetrachloride-induced hepatocyte damage in vitro and in vivo. Chem Biol Int 185: 94-100. https://doi.org/10.1016/j.cbi.2010.03.013
[45]	Cervantes-Llanos M, Lagumersindez-Denis N, Marín-Prida J, et al. (2018) Beneficial effects of oral administration of c-phycocyanin and phycocyanobilin in rodent models of experimental autoimmune encephalomyelitis. Life Sci 194: 130-138. https://doi.org/10.1016/j.lfs.2017.12.032
[46]	Wang C, Zhao Y, Wang L, et al. (2021) C-phycocyanin mitigates cognitive impairment in doxorubicin-induced chemobrain: impact on neuroinflammation, oxidative stress, and brain mitochondrial and synaptic alterations. Neurochem Res 46: 149-158. https://doi.org/10.1007/s11064-020-03164-2
[47]	Min SK, Park JS, Luo L, et al. (2015) Assessment of C-phycocyanin effect on astrocytes-mediated neuroprotection against oxidative brain injury using 2D and 3D astrocyte tissue model. Sci Rep 5: 14418. https://doi.org/10.1038/srep14418
[48]	Vadiraja BB, Gaikwad NW, Madyastha KM (1998) Hepatoprotective effect of C-phycocyanin: Protection for carbon tetrachloride and R-(+)-pulegone-mediated hepatotoxicty in rats. Biochem Bioph Res Co 249: 428-431. https://doi.org/10.1006/bbrc.1998.9149
[49]	Nagaraj S, Arulmurugan P, Rajaram MG, et al. (2012) Hepatoprotective and antioxidative effects of C-phycocyanin from Arthrospira maxima SAG 25780 in CCl4-induced hepatic damage rats. Biomed Prev Nutr 2: 81-85. https://doi.org/10.1016/j.bionut.2011.12.001
[50]	Soni B, Visavadiya NP, Madamwar D (2008) Ameliorative action of cyanobacterial phycoerythrin on CCl₄-induced toxicity in rats. Toxicology 248: 59-65. https://doi.org/10.1016/j.tox.2008.03.008
[51]	Fan C, Jiang J, Yin X, et al. (2012) Purification of selenium-containing allophycocyanin from selenium-enriched Spirulina platensis and its hepatoprotective effect against t-BOOH-induced apoptosis. Food Chem 134: 253-261. https://doi.org/10.1016/j.foodchem.2012.02.130
[52]	Nemoto-Kawamura C, Hirahashi T, Nagai T, et al. (2004) Phycocyanin enhances secretary IgA antibody response and suppresses allergic IgE antibody response in mice immunized with antigen-entrapped biodegradable microparticles. J Nutr Sci Vitaminol 50: 129-136. https://doi.org/10.3177/jnsv.50.129
[53]	Ivanova KG, Stankova KG, Nikolov VN, et al. (2010) The biliprotein C-phycocyanin modulates the early radiation response: A pilot study. Mutat Res-Gen Toxi En 695: 40-45. https://doi.org/10.1016/j.mrgentox.2009.11.002
[54]	Lee D, Nishizawa M, Shimizu Y, et al. (2017) Anti-inflammatory effects of dulse (Palmaria palmata) resulting from the simultaneous water-extraction of phycobiliproteins and chlorophyll a. Food Res Int 100: 514-521. https://doi.org/10.1016/j.foodres.2017.06.040
[55]	Chen HW, Yang TS, Chen MJ, et al. (2014) Purification and immunomodulating activity of C-phycocyanin from Spirulina platensis cultured using power plant flue gas. Process Biochem 49: 1337-1344. https://doi.org/10.1016/j.procbio.2014.05.006
[56]	Grover P, Bhatnagar A, Kumari N, et al. (2021) C-phycocyanin-a novel protein from Spirulina platensis-in vivo toxicity, antioxidant and immunomodulatory studies. Saudi J Biol Sci 28: 1853-1859. https://doi.org/10.1016/j.sjbs.2020.12.037
[57]	McCarty MF (2011) Clinical potential of phycocyanobilin for induction of T regulatory cells in the management of inflammatory disorders. Med Hypotheses 77: 1031-1033. https://doi.org/10.1016/j.mehy.2011.08.041
[58]	Hao S, Li S, Wang J, et al. (2019) Phycocyanin exerts anti-proliferative effects through down-regulating TIRAP/NF-κB activity in human non-small cell lung cancer cells. Cells 8: 588. https://doi.org/10.3390/cells8060588
[59]	Liao G, Gao B, Gao Y, et al. (2016) Phycocyanin inhibits tumorigenic potential of pancreatic cancer cells: Role of apoptosis and autophagy. Sci Rep 6: 34564. https://doi.org/10.1038/srep34564
[60]	Jiang L, Wang Y, Zhu F, et al. (2019) Molecular mechanism of anti-cancer activity of the nano-drug C-PC/CMC-CD59sp NPs in cervical cancer. J Cancer 10: 92-104. https://doi.org/10.7150/jca.27462
[61]	Subhashini J, Mahipal SV, Reddy MC, et al. (2004) Molecular mechanisms in C-phycocyanin induced apoptosis in human chronic myeloid leukemia cell line-K562. Biochem Pharmacol 68: 453-462. https://doi.org/10.1016/j.bcp.2004.02.025
[62]	Li B, Gao MH, Zhang XC, et al. (2006) Molecular immune mechanism of C-phycocyanin from Spirulina platensis induces apoptosis in HeLa cells in vitro. Biotechnol Appl Biochem 43: 155-164. https://doi.org/10.1042/BA20050142
[63]	Jiang L, Wang Y, Liu G, et al. (2018) C-Phycocyanin exerts anti-cancer effects via the MAPK signaling pathway in MDA-MB-231 cells. Cancer Cell Int 18: 12. https://doi.org/10.1186/s12935-018-0511-5
[64]	Kaur P, Dhandayuthapani S, Venkatesan T, et al. (2020) Molecular mechanism of C-phycocyanin induced apoptosis in LNCaP cells. Bioorg Med Chem 28: 115272. https://doi.org/10.1016/j.bmc.2019.115272
[65]	Gantar M, Dhandayuthapani S, Rathinavelu A (2012) Phycocyanin induces apoptosis and enhances the effect of topotecan on prostate cell line LNCaP. J Med Food 15: 1091-1095. https://doi.org/10.1089/jmf.2012.0123
[66]	Singh AP, Gupta A, Jaiswal J, et al. (2025) Phycobiliproteins: Phycocyanin, allophycocyanin, and phycoerythrin. Microalgae and one health . Academic Press 171-186. https://doi.org/10.1016/B978-0-443-22080-7.00033-8
[67]	Akbarizare M, Ofoghi H, Hadizadeh M, et al. (2020) In vitro assessment of the cytotoxic effects of secondary metabolites from Spirulina platensis on hepatocellular carcinoma. Egypt Liver J 10: 11. https://doi.org/10.1186/s43066-020-0018-3
[68]	Safaei M, Maleki H, Soleimanpour H, et al. (2019) Development of a novel method for the purification of C-phycocyanin pigment from a local cyanobacterial strain Limnothrix sp. NS01 and evaluation of its anticancer properties. Sci Rep 9: 9474. https://doi.org/10.1038/s41598-019-45905-6
[69]	Bottone C, Camerlingo R, Miceli R, et al. (2019) Antioxidant and anti-proliferative properties of extracts from heterotrophic cultures of Galdieria sulphuraria. Nat Prod Res 33: 1659-1663. https://doi.org/10.1080/14786419.2018.1425853
[70]	Ying J, Tang Z, Zhao G, et al. (2021) Transcriptionomic study on apoptosis of SKOV-3 cells induced by phycoerythrin from Gracilaria lemaneiformis. Anti-Cancer Agent Me 21: 1240-1249. https://doi.org/10.2174/1871520620666200908102621
[71]	Sanattalab E (2024) Insights into marine bioluminescence: Unravelling the intricacies of natural fluorescence probes. Hacettepe J Biol Chem 52: 285-296. https://doi.org/10.15671/hjbc.1465113
[72]	Rodriguez EA, Tran GN, Gross LA, et al. (2016) A far-red fluorescent protein evolved from a cyanobacterial phycobiliprotein. Nat Methods 13: 763-769. https://doi.org/10.1038/nmeth.3935
[73]	Jespersen L, Strømdahl LD, Olsen K, et al. (2005) Heat and light stability of three natural blue colorants for use in confectionery and beverages. Eur Food Res Technol 220: 261-266. https://doi.org/10.1007/s00217-004-1062-7
[74]	Adli SA, Ali F, Azmi AS, et al. (2020) Development of biodegradable cosmetic patch using a polylactic acid/phycocyanin–alginate composite. Polymer 12: 1669. https://doi.org/10.3390/polym12081669
[75]	Zhang Y, Liu H, Dai X, et al. (2021) Cyanobacteria-based near-infrared light-excited self-supplying oxygen system for enhanced photodynamic therapy of hypoxic tumors. Nano Res 14: 667-673. https://doi.org/10.1007/s12274-020-3094-0
[76]	Sekar S, Chandramohan M (2008) Phycobiliproteins as a commodity: Trends in applied research, patents and commercialization. J Appl Phycol 20: 113-136. https://doi.org/10.1007/s10811-007-9188-1
[77]	Xing FL, Zhang ZH, Yang CL, et al. (2022) Phycoerythrobilin/phycourobilin as efficient sensitizers of dye-sensitized solar cell. Soil Energy 243: 494-499. https://doi.org/10.1016/j.solener.2022.08.028
[78]	Qiang X, Wang L, Niu J, et al. (2021) Phycobiliprotein as fluorescent probe and photosensitizer: A systematic review. Int J Biol Macromol 193: 1910-1917. https://doi.org/10.1016/j.ijbiomac.2021.11.022
[79]	Bazylińska U, Kulbacka J, Schmidt J, et al. (2018) Polymer-free cubosomes for simultaneous bioimaging and photodynamic action of photosensitizers in melanoma skin cancer cells. J Colloid Interface Sci 522: 163-173. https://doi.org/10.1016/j.jcis.2018.03.063
[80]	Mokwena MG, Kruger CA, Ivan MT, et al. (2018) A review of nanoparticle photosensitizer drug delivery uptake systems for photodynamic treatment of lung cancer. Photodiagn Photodyn 22: 147-154. https://doi.org/10.1016/j.pdpdt.2018.03.006
[81]	Loshchenov МV, Levkin VV, Chernousov АF, et al. (2018) Laser video fluorescence diagnosis of stomach diseases. Sovrem Tehnol Med 10: 42. https://doi.org/10.17691/stm2018.10.4.05
[82]	Theresia C, Zheng J, Chen XY (2017) Topical ALA-PDT as alternative therapeutic option in treatment-recalcitrant dermatosis: report of 4 cases. Photodiagn Photodyn 20: 189-192. https://doi.org/10.1016/j.pdpdt.2017.10.010
[83]	Grossman AR, Schaefer MR, Chiang GG, et al. (1993) The phycobilisome, a light-harvesting complex responsive to environ mental conditions. Microbiol Rev 57: 725-749. https://doi.org/10.1128/mr.57.3.725-749.1993
[84]	Loos D, Cotlet M, De Schryver F, et al. (2004) Single-molecule spectroscopy selectively probes donor and acceptor chromophores in the phycobiliprotein allophycocyanin. Biophys J 87: 2598-2608. https://doi.org/10.1529/biophysj.104.046219
[85]	Peng PP, Dong LL, Sun YF, et al. (2014) The structure of allophycocyanin B from Synechocystis PCC 6803 reveals the structural basis for the extreme redshift of the terminal emitter in phycobilisomes. Acta Cryst 70: 2558-2569. https://doi.org/10.1107/S1399004714015776
[86]	Scheer H, Zhao KH (2008) Biliprotein maturation: The chromophore attachment. Mol Microbiol 68: 263-276. https://doi.org/10.1111/j.1365-2958.2008.06160.x
[87]	Sonani RR, Gupta GD, Madamwar D, et al. (2015) Crystal structure of allophycocyanin from marine cyanobacterium Phormidium sp. A09DM. PLoS ONE 10: e0124580. https://doi.org/10.1371/journal.pone.0124580
[88]	Biswas A, Vasquez YM, Dragomani TM, et al. (2010) Biosynthesis of cyanobacterial phycobiliproteins in Escherichia coli: chromophorylation efficiency and specificity of all bilin lyases from Synechococcus sp. strain PCC 7002. Appl Environ Microbiol 76: 2729-2739. https://doi.org/10.1128/AEM.03100-09
[89]	Tang K, Ding WL, Höppner A, et al. (2015) The terminal phycobilisome emitter, LCM: A light-harvesting pigment with a phytochrome chromophore. Proc Natl Acad Sci USA 112: 15880-15885. https://doi.org/10.1073/pnas.1519177113
[90]	Sowndarya DS (2021) Extraction and purification of C-phycocyanin from Arthrospira species and its application in LIP-BALM formulation. Int J Creat Res Thoughts 9: b492-b502.
[91]	Feng Y, Lu H, Hu J, et al. (2022) Anti-aging effects of R-phycocyanin from Porphyra haitanensis on HUVEC cells and Drosophila melanogaster. Mar Drugs 20: 468. https://doi.org/10.3390/md20080468
[92]	Wu LC, Lin YY, Yang SY, et al. (2011) Antimelanogenic effect of c-phycocyanin through modulation of tyrosinase expression by upregulation of ERK and downregulation of p38 MAPK signaling pathways. J Biomed Sci 18: 74. https://doi.org/10.1186/1423-0127-18-74
[93]	Nihal B, Gupta NV, Gowda DV, et al. (2018) Formulation and development of topical anti acne formulation of spirulina extract. Int J Appl Pharm 10: 229-233. https://doi.org/10.22159/IJAP.2018V10I6.26334
[94]	Athiyappan KD, Routray W, Paramasivan B (2024) Phycocyanin from Spirulina: A comprehensive review on cultivation, extraction, purification, and its application in food and allied industries. Food Humanity 2: 100235. https://doi.org/10.1016/j.foohum.2024.100235
[95]	Luzardo-Ocampo I, Ramírez-Jiménez AK, Yañez J, et al. (2021) Technological applications of natural colorants in food systems: A review. Foods 10: 634. https://doi.org/10.3390/foods10030634
[96]	Li Y, Li X, Liang ZP, et al. (2022) Progress of microencapsulated phycocyanin in food and pharma industries: A review. Molecules 27: 5854. https://doi.org/10.3390/molecules27185854
[97]	Alam MA, Xu JL, Wang Z (2020) Microalgae biotechnology for food, health and high value products. Springer Singapore. https://doi.org/10.1007/978-981-15-0169-2
[98]	Dewi EN, Kurniasih RA, Purnamayati L (2018) The application of microencapsulated phycocyanin as a blue natural colorant to the quality of jelly candy. IOP Conference Series: Earth Environ Sci 116: 012047. https://doi.org/10.1088/1755-1315/116/1/012047
[99]	de O Moreira I, Passos TS, Chiapinni C, et al. (2012) Colour evaluation of a phycobiliprotein-rich extract obtained from Nostoc PCC9205 in acidic solutions and yogurt. J Sci Food Agric 92: 598-605. https://doi.org/10.1002/jsfa.4614
[100]	Galetovic A, Seura F, Gallardo V, et al. (2020) Use of phycobiliproteins from atacama cyanobacteria as food colorants in a dairy beverage prototype. Foods 9: 244. https://doi.org/10.3390/foods9020244
[101]	García AB, Longo E, Bermejo R (2021) The application of a phycocyanin extract obtained from Arthrospira platensis as a blue natural colorant in beverages. J Appl Phycol 33: 3059-3070. https://doi.org/10.1007/s10811-021-02522-z
[102]	Madji S, Antih J, Tabib M, et al. (2025) Extraction, purification, bioactivity and pharmacological effects of phycobiliproteins (PBPs): A review. Analytica 6: 44. https://doi.org/10.3390/analytica6040044
[103]	Kovaleski G, Kholany M, Dias LMS, et al. (2022) Extraction and purification of phycobiliproteins from algae and their applications. Front Chem 10: 1065355. https://doi.org/10.3389/fchem.2022.1065355
[104]	Zahra Z, Choo DH, Lee H, et al. (2020) Cyanobacteria: Review of current potentials and applications. Environments 7: 13. https://doi.org/10.3390/environments7020013
[105]	Minić S, Gligorijević N, Veličković L, et al. (2024) Narrative review of the current and future perspectives of phycobiliproteins' applications in the food industry: From natural colors to alternative proteins. Int J Mol Sci 25: 7187. https://doi.org/10.3390/ijms25137187
[106]	Rawiwan P, Peng Y, Paramayuda IGPB, et al. (2022) Red seaweed: A promising alternative protein source for global food sustainability. Trends Food Sci Technol 123: 37-56. https://doi.org/10.1016/j.tifs.2022.03.003
[107]	Van der Spiegel M, Noordam MY, Van der Fels-Klerx HJ (2013) Safety of novel protein sources (insects, microalgae, seaweed, duckweed, and rapeseed) and legislative aspects for their application in food and feed production. Compr Rev Food Sci F 12: 662-678. https://doi.org/10.1111/1541-4337.12032
[108]	Liang Y, Deng L, Feng Z, et al. (2023) A chitosan-based flocculation method for efficient recovery of high-purity B-phycoerythrin from a low concentration of phycobilin in wastewater. Molecules 28: 3600. https://doi.org/10.3390/molecules28083600
[109]	Kovaleski G, Kholany M, Dias LMS, et al. (2022) Extraction and purification of phycobiliproteins from algae and their applications. Front Chem 10: 1065355. https://doi.org/10.3389/fchem.2022.1065355
[110]	Zahra Z, Choo DH, Lee H, et al. (2020) Cyanobacteria: Review of current potentials and applications. Environments 7: 13. https://doi.org/10.3390/environments7020013
[111]	Gisriel CJ, Shen G, Brudvig GW, et al. (2024) Structure of the antenna complex expressed during far-red light photoacclimation in Synechococcus sp. PCC 7335. J Biol Chem 300: 105590. https://doi.org/10.1016/j.jbc.2023.105590
[112]	Ko YJ, Lee ME, Cho BH, et al. (2024) Bioproduction of porphyrins, phycobilins, and their proteins using microbial cell factories: engineering, metabolic regulations, challenges, and perspectives. Crit Rev Biotechnol 44: 373-387. https://doi.org/10.1080/07388551.2023.2168512

Reader Comments

Your name:*

Email:*
© 2026 the Author(s), licensee AIMS Press. This is an open access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/4.0)