Integrated Strategies to Enhance Lipid Productivity in Microalgae for Sustainable Biodiesel Production

Fozy Binhweel; Abdalah Makaranga; Mardiana Idayu Ahmad; Pannaga Pavan Jutur; Nor Hawani Salikin; Mohammad Aliff Shakir

doi:10.65582/gti.2026.011

Authors

Fozy Binhweel School of Industrial Technology, Universiti Sains Malaysia, 11800, Penang, Malaysia; Faculty of Environmental Science and Marine Biology, Hadhramout University, Hadhramout, Yemen. https://orcid.org/0000-0002-7188-0725
Abdalah Makaranga Omics of Algae Group, Industrial Biotechnology, International Centre for Genetic Engineering and Biotechnology, Aruna Asaf Ali Marg, New Delhi 110067, India.
Mardiana Idayu Ahmad School of Industrial Technology, Universiti Sains Malaysia, 11800, Penang, Malaysia. https://orcid.org/0000-0003-1412-3119
Pannaga Pavan Jutur Omics of Algae Group, Industrial Biotechnology, International Centre for Genetic Engineering and Biotechnology, Aruna Asaf Ali Marg, New Delhi 110067, India. https://orcid.org/0000-0001-7988-2883
Nor Hawani Salikin School of Industrial Technology, Universiti Sains Malaysia, 11800, Penang, Malaysia. https://orcid.org/0000-0002-6531-9747
Mohammad Aliff Shakir School of Industrial Technology, Universiti Sains Malaysia, 11800, Penang, Malaysia. https://orcid.org/0000-0003-3137-4446

DOI:

https://doi.org/10.65582/gti.2026.011

Keywords:

Microalgae, Lipid Enhancement, Biodiesel Production, Genetic Engineering, Transesterification

Abstract

Microalgae have emerged as a promising third-generation feedstock for sustainable biodiesel production due to their rapid growth, high photosynthetic efficiency, and ability to grow on non-arable land without competing with food resources. However, their relatively low natural lipid content remains a critical challenge for commercial-scale biodiesel viability. This review presents a comprehensive overview of integrated strategies to enhance lipid productivity in microalgal biomass, including optimization of cultivation media under nutrient stress, nanoparticle-mediated stimulation, and genetic engineering interventions. Recent advancements in cell wall disruption and lipid extraction techniques are also discussed, alongside the development of in situ transesterification to improve biodiesel conversion efficiency. Furthermore, a techno-economic analysis highlights the potential and limitations of these approaches in industrial applications. By integrating biological, chemical, and engineering innovations, this paper underscores the viability of microalgae as a renewable and scalable source for biodiesel, contributing toward global energy sustainability.

Author Biography

Mardiana Idayu Ahmad, School of Industrial Technology, Universiti Sains Malaysia, 11800, Penang, Malaysia.

*Corresponding author

References

Abbriano, R. et al., 2018. Manipulation of a glycolytic regulator alters growth and carbon partitioning in the marine diatom Thalassiosira pseudonana, Algal Research, 32, pp. 250–258. DOI: https://doi.org/10.1016/j.algal.2018.03.018. DOI: https://doi.org/10.1016/j.algal.2018.03.018

Ahmad, M.I. et al., 2023. Optimization of base catalytic transesterification toward maximum biodiesel yield from Azolla filiculoides macroalgae feedstock, Industrial Crops and Products, 197, p. 116590. DOI: https://doi.org/10.1016/j.indcrop.2023.116590. DOI: https://doi.org/10.1016/j.indcrop.2023.116590

Alsaadi, S., Shakir, M.A. and Ahmad, M.I., 2025. A Comparative Review of Vegetative and Animal-Based Biodiesel: Feedstock Selection, Synthesis Method, Characterization Technique, and Economic Perspective, Journal of Climate Change, 11(2), p. 22. DOI: https://doi.org/10.70917/jcc-2025-013. DOI: https://doi.org/10.70917/jcc-2025-013

An, M. et al., 2020. Effects of Nitrogen Forms and Supply Mode on Lipid Production of Microalga Scenedesmus obliquus, Energies, 13(3), p. 697. DOI: https://doi.org/10.3390/en13030697. DOI: https://doi.org/10.3390/en13030697

Araujo, G.S. et al., 2013. Extraction of lipids from microalgae by ultrasound application: Prospection of the optimal extraction method, Ultrasonics Sonochemistry, 20(1), pp. 95–98. DOI: https://doi.org/10.1016/j.ultsonch.2012.07.027. DOI: https://doi.org/10.1016/j.ultsonch.2012.07.027

Arora, N. et al., 2018. Leveraging algal omics to reveal potential targets for augmenting TAG accumulation, Biotechnology Advances, 36(4), pp. 1274–1292. DOI: https://doi.org/10.1016/j.biotechadv.2018.04.005. DOI: https://doi.org/10.1016/j.biotechadv.2018.04.005

Atiku, H. et al., 2016. Harvesting of microalgae biomass from the phycoremediation process of greywater, Environmental Science and Pollution Research, 23(24), pp. 24624–24641. DOI: https://doi.org/10.1007/s11356-016-7456-9. DOI: https://doi.org/10.1007/s11356-016-7456-9

Binhweel, F. et al., 2023. Utilization of marine ulva lactuca seaweed and freshwater azolla filiculoides macroalgae feedstocks toward biodiesel production: Kinetics, thermodynamics, and optimization studies, Renewable Energy, 205, pp. 717–730. DOI: https://doi.org/10.1016/j.renene.2023.01.114. DOI: https://doi.org/10.1016/j.renene.2023.01.114

Binhweel, F. et al., 2024. Kinetics, thermodynamics, and optimization analyses of lipid extraction from discarded beef tallow for bioenergy production, Separation Science and Technology, pp. 1–20. DOI: https://doi.org/10.1080/01496395.2024.2387277. DOI: https://doi.org/10.1080/01496395.2024.2387277

Binhweel, F. et al., 2025. Biodiesel production from Ulva lactuca: Insights into kinetic and thermodynamic evaluation of lipid transesterification, Biocatalysis and Agricultural Biotechnology, 70, p. 103856. DOI: https://doi.org/https://doi.org/10.1016/j.bcab.2025.103856. DOI: https://doi.org/10.1016/j.bcab.2025.103856

Binhweel, F. et al., 2026. A review on the chemical and physical characteristics of biodiesels produced from animal fat waste, Research and Reviews in Sustainability, 2(1 SE-Reviews), pp. 31–48. DOI: https://doi.org/10.65582/rrs.2026.004. DOI: https://doi.org/10.65582/rrs.2026.004

Binhweel, F., Ahmad, M.I. and Zaki, S.A., 2022. Utilization of Polymeric Materials toward Sustainable Biodiesel Industry: A Recent Review, Polymers, 14(19), p. 3950. DOI: https://doi.org/10.3390/polym14193950. DOI: https://doi.org/10.3390/polym14193950

Binhweel, F., Mardiana and Shakir, M.A., 2025. Harnessing waste lipids for biodiesel: Optimization and thermodynamic investigation of beef tallow transesterification, International Journal of Green Energy, pp. 1–13. DOI: https://doi.org/10.1080/15435075.2025.2508474. DOI: https://doi.org/10.1080/15435075.2025.2508474

Boni, J., Aida, S. and Leila, K., 2018. Lipid Extraction Method From Microalgae Botryococcus Braunii As Raw Material To Make Biodiesel With Soxhlet Extraction, Journal of Physics: Conference Series, 1095, p. 012004. DOI: https://doi.org/10.1088/1742-6596/1095/1/012004. DOI: https://doi.org/10.1088/1742-6596/1095/1/012004

Brennan, L. and Owende, P., 2010. Biofuels from microalgae—A review of technologies for production, processing, and extractions of biofuels and co-products, Renewable and Sustainable Energy Reviews, 14(2), pp. 557–577. DOI: https://doi.org/10.1016/j.rser.2009.10.009. DOI: https://doi.org/10.1016/j.rser.2009.10.009

Cao, H. et al., 2013. Direct Biodiesel Production from Wet Microalgae Biomass of Chlorella pyrenoidosa through In Situ Transesterification, BioMed Research International, 2013, pp. 1–6. DOI: https://doi.org/10.1155/2013/930686. DOI: https://doi.org/10.1155/2013/930686

Carvalho Júnior, R.M. et al., 2011. Microalgae biodiesel via in situ methanolysis, Journal of Chemical Technology & Biotechnology, 86(11), pp. 1418–1427. DOI: https://doi.org/10.1002/jctb.2652. DOI: https://doi.org/10.1002/jctb.2652

Cerff, M. et al., 2012. Harvesting fresh water and marine algae by magnetic separation: Screening of separation parameters and high gradient magnetic filtration, Bioresource Technology, 118, pp. 289–295. DOI: https://doi.org/10.1016/j.biortech.2012.05.020. DOI: https://doi.org/10.1016/j.biortech.2012.05.020

Chen, X. et al., 2015. Ionic liquid‐assisted subcritical water promotes the extraction of lipids from wet microalgae Scenedesmus sp, European Journal of Lipid Science and Technology, 117(8), pp. 1192–1198. DOI: https://doi.org/10.1002/ejlt.201400189. DOI: https://doi.org/10.1002/ejlt.201400189

Cheng, J. et al., 2016. Enhancing the growth rate and astaxanthin yield of Haematococcus pluvialis by nuclear irradiation and high concentration of carbon dioxide stress, Bioresource Technology, 204, pp. 49–54. DOI: https://doi.org/10.1016/j.biortech.2015.12.076. DOI: https://doi.org/10.1016/j.biortech.2015.12.076

Chew, K.W. et al., 2017. Microalgae biorefinery: High value products perspectives, Bioresource Technology, 229, pp. 53–62. DOI: https://doi.org/10.1016/j.biortech.2017.01.006. DOI: https://doi.org/10.1016/j.biortech.2017.01.006

Chu, F.-F. et al. 2013. Phosphorus plays an important role in enhancing biodiesel productivity of Chlorella vulgaris under nitrogen deficiency, Bioresource Technology, 134, pp. 341–346. DOI: https://doi.org/10.1016/j.biortech.2013.01.131. DOI: https://doi.org/10.1016/j.biortech.2013.01.131

Chungjatupornchai, W. and Fa-aroonsawat, S., 2021. Enhanced triacylglycerol production in oleaginous microalga Neochloris oleoabundans by co-overexpression of lipogenic genes: Plastidial LPAAT1 and ER-located DGAT2, Journal of Bioscience and Bioengineering, 131(2), pp. 124–130. DOI: https://doi.org/10.1016/j.jbiosc.2020.09.012. DOI: https://doi.org/10.1016/j.jbiosc.2020.09.012

Cornish-Bowden, A., 2014. Current IUBMB recommendations on enzyme nomenclature and kinetics, Perspectives in Science, 1(1–6), pp. 74–87. DOI: https://doi.org/10.1016/j.pisc.2014.02.006. DOI: https://doi.org/10.1016/j.pisc.2014.02.006

Dejoye Tanzi, C., Abert Vian, M. and Chemat, F., 2013. New procedure for extraction of algal lipids from wet biomass: A green clean and scalable process, Bioresource Technology, 134, pp. 271–275. DOI: https://doi.org/10.1016/j.biortech.2013.01.168. DOI: https://doi.org/10.1016/j.biortech.2013.01.168

Delgado, R. et al., 2021. Effect of nitrogen limitation on growth, biochemical composition, and cell ultrastructure of the microalga Picocystis salinarum, Journal of Applied Phycology, 33(4), pp. 2083–2092. DOI: https://doi.org/10.1007/s10811-021-02462-8. DOI: https://doi.org/10.1007/s10811-021-02462-8

Dong, T. et al., 2013. Two-step in situ biodiesel production from microalgae with high free fatty acid content, Bioresource Technology, 136, pp. 8–15. DOI: https://doi.org/10.1016/j.biortech.2013.02.105. DOI: https://doi.org/10.1016/j.biortech.2013.02.105

Feng, P. et al., 2012. Lipid accumulation and growth characteristics of Chlorella zofingiensis under different nitrate and phosphate concentrations, Journal of Bioscience and Bioengineering, 114(4), pp. 405–410. DOI: https://doi.org/10.1016/j.jbiosc.2012.05.007. DOI: https://doi.org/10.1016/j.jbiosc.2012.05.007

Golmakani, M.-T. et al., 2014. Pressurized limonene as an alternative bio-solvent for the extraction of lipids from marine microorganisms, The Journal of Supercritical Fluids, 92, pp. 1–7. DOI: https://doi.org/10.1016/j.supflu.2014.05.001. DOI: https://doi.org/10.1016/j.supflu.2014.05.001

Gómez, B. et al., 2019. Application of pulsed electric fields in meat and fish processing industries: An overview, Food Research International, 123, pp. 95–105. DOI: https://doi.org/10.1016/j.foodres.2019.04.047. DOI: https://doi.org/10.1016/j.foodres.2019.04.047

Gomma, A.E. et al., 2015. Improvement in Oil Production by Increasing Malonyl-CoA and Glycerol-3-Phosphate Pools in Scenedesmus quadricauda, Indian Journal of Microbiology, 55(4), pp. 447–455. DOI: https://doi.org/10.1007/s12088-015-0546-4. DOI: https://doi.org/10.1007/s12088-015-0546-4

Han, H.-S. et al., 2016. Regulation of glucose metabolism from a liver-centric perspective, Experimental & Molecular Medicine, 48(3), pp. e218–e218. DOI: https://doi.org/10.1038/emm.2015.122. DOI: https://doi.org/10.1038/emm.2015.122

Han, S.-F. et al., 2019. Application of pulse electric field pretreatment for enhancing lipid extraction from Chlorella pyrenoidosa grown in wastewater, Renewable Energy, 133, pp. 233–239. DOI: https://doi.org/10.1016/j.renene.2018.10.034. DOI: https://doi.org/10.1016/j.renene.2018.10.034

Hannon, M. et al., 2010. Biofuels from algae: challenges and potential, Biofuels, 1(5), pp. 763–784. DOI: https://doi.org/10.4155/bfs.10.44. DOI: https://doi.org/10.4155/bfs.10.44

He, M. et al., 2017. Improvement on lipid production by Scenedesmus obliquus triggered by low dose exposure to nanoparticles, Scientific Reports, 7(1), p. 15526. DOI: https://doi.org/10.1038/s41598-017-15667-0. DOI: https://doi.org/10.1038/s41598-017-15667-0

He, Y. et al., 2020. Sustainable biodiesel production from the green microalgae Nannochloropsis: Novel integrated processes from cultivation to enzyme-assisted extraction and ethanolysis of lipids, Energy Conversion and Management, 209, p. 112618. DOI: https://doi.org/10.1016/j.enconman.2020.112618. DOI: https://doi.org/10.1016/j.enconman.2020.112618

Hossain, N., Mahlia, T.M.I. and Saidur, R., 2019. Latest development in microalgae-biofuel production with nano-additives, Biotechnology for Biofuels, 12(1), p. 125. DOI: https://doi.org/10.1186/s13068-019-1465-0. DOI: https://doi.org/10.1186/s13068-019-1465-0

Huang, L.-F. et al., 2015. Overexpressing Ferredoxins in Chlamydomonas reinhardtii Increase Starch and Oil Yields and Enhance Electric Power Production in a Photo Microbial Fuel Cell, International Journal of Molecular Sciences, 16(8), pp. 19308–19325. DOI: https://doi.org/10.3390/ijms160819308. DOI: https://doi.org/10.3390/ijms160819308

Hwangbo, K. et al., 2014. Overexpression of stearoyl-ACP desaturase enhances accumulations of oleic acid in the green alga Chlamydomonas reinhardtii, Plant Biotechnology Reports, 8(2), pp. 135–142. DOI: https://doi.org/10.1007/s11816-013-0302-3. DOI: https://doi.org/10.1007/s11816-013-0302-3

Jia, J. et al., 2015. Molecular mechanisms for photosynthetic carbon partitioning into storage neutral lipids in Nannochloropsis oceanica under nitrogen-depletion conditions, Algal Research, 7, pp. 66–77. DOI: https://doi.org/10.1016/j.algal.2014.11.005. DOI: https://doi.org/10.1016/j.algal.2014.11.005

Kim, B. et al., 2019. Simplifying biodiesel production from microalgae via wet in situ transesterification: A review in current research and future prospects, Algal Research, 41, p. 101557. DOI: https://doi.org/10.1016/j.algal.2019.101557. DOI: https://doi.org/10.1016/j.algal.2019.101557

Koh, H.G. et al., 2019. Heterologous synthesis of chlorophyll b in Nannochloropsis salina enhances growth and lipid production by increasing photosynthetic efficiency, Biotechnology for Biofuels, 12(1), p. 122. DOI: https://doi.org/10.1186/s13068-019-1462-3. DOI: https://doi.org/10.1186/s13068-019-1462-3

Kwon, S. et al., 2018. Enhancement of biomass and lipid productivity by overexpression of a bZIP transcription factor in Nannochloropsis salina, Biotechnology and Bioengineering, 115(2), pp. 331–340. DOI: https://doi.org/10.1002/bit.26465. DOI: https://doi.org/10.1002/bit.26465

Lai, Y.S. et al., 2014. Effects of pulsed electric field treatment on enhancing lipid recovery from the microalga, Scenedesmus, Bioresource Technology, 173, pp. 457–461. DOI: https://doi.org/10.1016/j.biortech.2014.09.124. DOI: https://doi.org/10.1016/j.biortech.2014.09.124

Latsos, C., van Houcke, J. and Timmermans, K.R., 2020. The Effect of Nitrogen Starvation on Biomass Yield and Biochemical Constituents of Rhodomonas sp., Frontiers in Marine Science, 7. DOI: https://doi.org/10.3389/fmars.2020.563333. DOI: https://doi.org/10.3389/fmars.2020.563333

Lee, S.Y. et al., 2021. Techniques of lipid extraction from microalgae for biofuel production: a review, Environmental Chemistry Letters, 19(1), pp. 231–251. DOI: https://doi.org/10.1007/s10311-020-01088-5. DOI: https://doi.org/10.1007/s10311-020-01088-5

Li-Beisson, Y. et al., 2019. The lipid biochemistry of eukaryotic algae, Progress in Lipid Research, 74, pp. 31–68. DOI: https://doi.org/10.1016/j.plipres.2019.01.003. DOI: https://doi.org/10.1016/j.plipres.2019.01.003

Li, K.-Y. et al., 2025. Advances of microalgae-based enhancement strategies in industrial flue gas treatment: From carbon sequestration to lipid production, Bioresource Technology, 423, p. 132250. DOI: https://doi.org/10.1016/j.biortech.2025.132250. DOI: https://doi.org/10.1016/j.biortech.2025.132250

Li, P. et al., 2011. In Situ Biodiesel Production from Fast-Growing and High Oil Content Chlorella pyrenoidosa in Rice Straw Hydrolysate, Journal of Biomedicine and Biotechnology, 2011, pp. 1–8. DOI: https://doi.org/10.1155/2011/141207. DOI: https://doi.org/10.1155/2011/141207

Liang, K. et al., 2013. Effect of phosphorus on lipid accumulation in freshwater microalga Chlorella sp., Journal of Applied Phycology, 25(1), pp. 311–318. DOI: https://doi.org/10.1007/s10811-012-9865-6. DOI: https://doi.org/10.1007/s10811-012-9865-6

Lim, J.K. et al., 2012. Rapid Magnetophoretic Separation of Microalgae, Small, 8(11), pp. 1683–1692. DOI: https://doi.org/10.1002/smll.201102400. DOI: https://doi.org/10.1002/smll.201102400

Liu, H. et al., 2024. A review of the strategy to promote microalgae value in CO2 conversion-lipid enrichment-biodiesel production, Journal of Cleaner Production, 436, p. 140538. DOI: https://doi.org/10.1016/j.jclepro.2023.140538. DOI: https://doi.org/10.1016/j.jclepro.2023.140538

Liu, Z. et al., 2023. Technologies for harvesting the microalgae for industrial applications: Current trends and perspectives, Bioresource Technology, 387, p. 129631. DOI: https://doi.org/10.1016/j.biortech.2023.129631. DOI: https://doi.org/10.1016/j.biortech.2023.129631

Morozova, S., Karabuga, A. and Utlu, Z., 2025. Geopolitical Conflicts: Blockchain Use in International Energy Trading, Energy Catalyst, 1, pp. 79–88. DOI: https://doi.org/10.61552/EC.2025.006. DOI: https://doi.org/10.61552/EC.2025.006

Mulchandani, K., Kar, J.R. and Singhal, R.S., 2015. Extraction of Lipids from Chlorella saccharophila Using High-Pressure Homogenization Followed by Three Phase Partitioning, Applied Biochemistry and Biotechnology, 176(6), pp. 1613–1626. DOI: https://doi.org/10.1007/s12010-015-1665-4. DOI: https://doi.org/10.1007/s12010-015-1665-4

Muñoz, C.F. et al., 2021. Genetic engineering of microalgae for enhanced lipid production, Biotechnology Advances, 52, p. 107836. DOI: https://doi.org/10.1016/j.biotechadv.2021.107836. DOI: https://doi.org/10.1016/j.biotechadv.2021.107836

Muto, M. et al., 2015. Enhancement of glycerol metabolism in the oleaginous marine diatom Fistulifera solaris JPCC DA0580 to improve triacylglycerol productivity, Biotechnology for Biofuels, 8(1), p. 4. DOI: https://doi.org/10.1186/s13068-014-0184-9. DOI: https://doi.org/10.1186/s13068-014-0184-9

Nobre, B.P. et al., 2013. A biorefinery from Nannochloropsis sp. microalga – Extraction of oils and pigments. Production of biohydrogen from the leftover biomass, Bioresource Technology, 135, pp. 128–136. DOI: https://doi.org/10.1016/j.biortech.2012.11.084. DOI: https://doi.org/10.1016/j.biortech.2012.11.084

Obeid, S. et al., 2018. Supercritical carbon dioxide extraction and fractionation of lipids from freeze-dried microalgae Nannochloropsis oculata and Chlorella vulgaris, Algal Research, 34, pp. 49–56. DOI: https://doi.org/10.1016/j.algal.2018.07.003. DOI: https://doi.org/10.1016/j.algal.2018.07.003

Pan, J. et al., 2016. Microwave-assisted extraction of lipids from microalgae using an ionic liquid solvent [BMIM][HSO4], Fuel, 178, pp. 49–55. DOI: https://doi.org/10.1016/j.fuel.2016.03.037. DOI: https://doi.org/10.1016/j.fuel.2016.03.037

Pandey, S. et al., 2024. Biodiesel production from microalgae: A comprehensive review on influential factors, transesterification processes, and challenges, Fuel, 367, p. 131547. DOI: https://doi.org/10.1016/j.fuel.2024.131547. DOI: https://doi.org/10.1016/j.fuel.2024.131547

Pandit, P.R. and Fulekar, M.H., 2019. Biodiesel production from microalgal biomass using CaO catalyst synthesized from natural waste material, Renewable Energy, 136, pp. 837–845. DOI: https://doi.org/10.1016/j.renene.2019.01.047. DOI: https://doi.org/10.1016/j.renene.2019.01.047

Patel, A., Pruthi, V. and Pruthi, P.A., 2017.Synchronized nutrient stress conditions trigger the diversion of CDP-DG pathway of phospholipids synthesis towards de novo TAG synthesis in oleaginous yeast escalating biodiesel production, Energy, 139, pp. 962–974. DOI: https://doi.org/10.1016/j.energy.2017.08.052. DOI: https://doi.org/10.1016/j.energy.2017.08.052

Peng, K.-T. et al., 2014. Delta 5 Fatty Acid Desaturase Upregulates the Synthesis of Polyunsaturated Fatty Acids in the Marine Diatom Phaeodactylum tricornutum, Journal of Agricultural and Food Chemistry, 62(35), pp. 8773–8776. DOI: https://doi.org/10.1021/jf5031086. DOI: https://doi.org/10.1021/jf5031086

Pieber, S., Schober, S. and Mittelbach, M., 2012. Pressurized fluid extraction of polyunsaturated fatty acids from the microalga Nannochloropsis oculata, Biomass and Bioenergy, 47, pp. 474–482. DOI: https://doi.org/10.1016/j.biombioe.2012.10.019. DOI: https://doi.org/10.1016/j.biombioe.2012.10.019

Poliner, E. et al., 2018. A toolkit for Nannochloropsis oceanica CCMP 1779 enables gene stacking and genetic engineering of the eicosapentaenoic acid pathway for enhanced long‐chain polyunsaturated fatty acid production, Plant Biotechnology Journal, 16(1), pp. 298–309. DOI: https://doi.org/10.1111/pbi.12772. DOI: https://doi.org/10.1111/pbi.12772

Procházková, G. et al., 2014. Effect of nutrient supply status on biomass composition of eukaryotic green microalgae, Journal of Applied Phycology, 26(3), pp. 1359–1377. DOI: https://doi.org/10.1007/s10811-013-0154-9. DOI: https://doi.org/10.1007/s10811-013-0154-9

Qv, X., Zhou, Q. and Jiang, J., 2014. Ultrasound‐enhanced and microwave‐assisted extraction of lipid from Dunaliella tertiolecta and fatty acid profile analysis, Journal of Separation Science, 37(20), pp. 2991–2999. DOI: https://doi.org/10.1002/jssc.201400458. DOI: https://doi.org/10.1002/jssc.201400458

Rawat, J. et al., 2022. Latest Expansions in Lipid Enhancement of Microalgae for Biodiesel Production: An Update, Energies, 15(4), p. 1550. DOI: https://doi.org/10.3390/en15041550. DOI: https://doi.org/10.3390/en15041550

Razak, T.R. et al., 2026. An AI-Based Hybrid Prophet–LSTM Model for Forecasting and Financial Optimisation in Sustainable Energy Grids, Artificial Intelligence for Sustainable Cities, 1(1), pp. 42–60. DOI: https://doi.org/10.65582/aifsc.2026.004. DOI: https://doi.org/10.65582/aifsc.2026.004

Rodolfi, L. et al., 2009. Microalgae for oil: Strain selection, induction of lipid synthesis and outdoor mass cultivation in a low‐cost photobioreactor, Biotechnology and Bioengineering, 102(1), pp. 100–112. DOI: https://doi.org/10.1002/bit.22033. DOI: https://doi.org/10.1002/bit.22033

Roopnarain, A., Gray, V.M. and Sym, S.D., 2014. Phosphorus limitation and starvation effects on cell growth and lipid accumulation in Isochrysis galbana U4 for biodiesel production’, Bioresource Technology, 156, pp. 408–411. DOI: https://doi.org/10.1016/j.biortech.2014.01.092. DOI: https://doi.org/10.1016/j.biortech.2014.01.092

Salem, O.M.A., El-Ardy, O. and Abd El-Rahman, M.A., 2013. Effect of Nitrogen and Phosphorus Concentrations in Growth Medium and Salt Stress on Growth, Lipid Content, and Biodiesel Producing Ability of Microalgae’, in J. Bot. 3rd international conf, pp. 17–18.

Satpati, G.G., Gorain, P.C. and Pal, R., 2016. Efficacy of EDTA and Phosphorous on Biomass Yield and Total Lipid Accumulation in Two Green Microalgae with Special Emphasis on Neutral Lipid Detection by Flow Cytometry, Advances in Biology, 2016, pp. 1–12. DOI: https://doi.org/10.1155/2016/8712470. DOI: https://doi.org/10.1155/2016/8712470

Senusi, W. et al., 2024. Biodiesel production and characteristics from waste frying oils: sources, challenges, and circular economic perspective, Environmental Science and Pollution Research, 31(23), pp. 33239–33258. DOI: https://doi.org/10.1007/s11356-024-33533-1. DOI: https://doi.org/10.1007/s11356-024-33533-1

Shakir, M.A. et al., 2024. Biomass to biofuel: Palm kernel shells as catalyst supports for enhanced biodiesel production, Biofuels, Bioproducts and Biorefining, 18(6), pp. 2038–2052. DOI: https://doi.org/10.1002/bbb.2683. DOI: https://doi.org/10.1002/bbb.2683

Shakir, M.A. and Ahmad, M.I., 2025. Biodiesel production from Ulva lactuca oil extracted by supercritical carbon dioxide using a biogenic activated carbon catalyst, Bioresource Technology Reports, 32, p. 102393. DOI: https://doi.org/https://doi.org/10.1016/j.biteb.2025.102393. DOI: https://doi.org/10.1016/j.biteb.2025.102393

Shalfoh, E. et al., 2024. Fish waste oil extraction using supercritical CO2 extraction for biodiesel production: Mathematical, and kinetic modeling, Renewable Energy, 220, p. 119659. DOI: https://doi.org/10.1016/j.renene.2023.119659. DOI: https://doi.org/10.1016/j.renene.2023.119659

Shi, H. et al., 2018. Production of eicosapentaenoic acid by application of a delta-6 desaturase with the highest ALA catalytic activity in algae, Microbial Cell Factories, 17(1), p. 7. DOI: https://doi.org/10.1186/s12934-018-0857-3. DOI: https://doi.org/10.1186/s12934-018-0857-3

Sierra, L.S., Dixon, C.K. and Wilken, L.R., 2017. Enzymatic cell disruption of the microalgae Chlamydomonas reinhardtii for lipid and protein extraction, Algal Research, 25, pp. 149–159. DOI: https://doi.org/10.1016/j.algal.2017.04.004. DOI: https://doi.org/10.1016/j.algal.2017.04.004

Silve, A. et al., 2018. Extraction of lipids from wet microalga Auxenochlorella protothecoides using pulsed electric field treatment and ethanol-hexane blends, Algal Research, 29, pp. 212–222. DOI: https://doi.org/10.1016/j.algal.2017.11.016. DOI: https://doi.org/10.1016/j.algal.2017.11.016

Solovchenko, A. et al., 2020. Phosphorus Feast and Famine in Cyanobacteria: Is Luxury Uptake of the Nutrient Just a Consequence of Acclimation to Its Shortage?, Cells, 9(9), p. 1933. DOI: https://doi.org/10.3390/cells9091933. DOI: https://doi.org/10.3390/cells9091933

Subramanian, S. et al., 2013. Comparative energetics and kinetics of autotrophic lipid and starch metabolism in chlorophytic microalgae: implications for biomass and biofuel production, Biotechnology for Biofuels, 6(1), p. 150. DOI: https://doi.org/10.1186/1754-6834-6-150. DOI: https://doi.org/10.1186/1754-6834-6-150

Südfeld, C. et al., 2021. High-throughput insertional mutagenesis reveals novel targets for enhancing lipid accumulation in Nannochloropsis oceanica, Metabolic Engineering, 66, pp. 239–258. DOI: https://doi.org/10.1016/j.ymben.2021.04.012. DOI: https://doi.org/10.1016/j.ymben.2021.04.012

Sun, J. et al., 2019. Microalgae biodiesel production in China: A preliminary economic analysis, Renewable and Sustainable Energy Reviews, 104, pp. 296–306. DOI: https://doi.org/10.1016/j.rser.2019.01.021. DOI: https://doi.org/10.1016/j.rser.2019.01.021

Tan, J. Sen et al., 2020. A review on microalgae cultivation and harvesting, and their biomass extraction processing using ionic liquids, Bioengineered, 11(1), pp. 116–129. DOI: https://doi.org/10.1080/21655979.2020.1711626. DOI: https://doi.org/10.1080/21655979.2020.1711626

Tarigan, E., 2023. Comparison of Energy Production Between Fixed-Mount and Tracking Systems of Solar PV Systems in Jakarta, Indonesia, Future Cities and Environment, 9, p. 8. DOI: https://doi.org/10.5334/fce.167. DOI: https://doi.org/10.5334/fce.167

Tran, D.-T., Chen, C.-L. and Chang, J.-S., 2013. Effect of solvents and oil content on direct transesterification of wet oil-bearing microalgal biomass of Chlorella vulgaris ESP-31 for biodiesel synthesis using immobilized lipase as the biocatalyst, Bioresource Technology, 135, pp. 213–221. DOI: https://doi.org/10.1016/j.biortech.2012.09.101. DOI: https://doi.org/10.1016/j.biortech.2012.09.101

Tsigie, Y.A. et al., 2012. In situ biodiesel production from wet Chlorella vulgaris under subcritical condition, Chemical Engineering Journal, 213, pp. 104–108. DOI: https://doi.org/10.1016/j.cej.2012.09.112. DOI: https://doi.org/10.1016/j.cej.2012.09.112

Uyar, M. and Hayber, S.E., 2025. Impact of Clean Energy Integration on CO2 Intensity in European Power Systems, Global Decarbonisation, 1(1 SE-Technical Articles), pp. 1–18. Available here: https://globaldecarbonisation.com/index.php/gd/article/view/1056. DOI: https://doi.org/10.17184/eac.9904

Vasistha, S. et al., 2021. Current advances in microalgae harvesting and lipid extraction processes for improved biodiesel production: A review, Renewable and Sustainable Energy Reviews, 137, p. 110498. DOI: https://doi.org/10.1016/j.rser.2020.110498. DOI: https://doi.org/10.1016/j.rser.2020.110498

Velasquez-Orta, S.B., Lee, J.G.M. and Harvey, A., 2012. Alkaline in situ transesterification of Chlorella vulgaris, Fuel, 94, pp. 544–550. DOI: https://doi.org/10.1016/j.fuel.2011.11.045. DOI: https://doi.org/10.1016/j.fuel.2011.11.045

Viguera, M. et al., 2016. The process parameters and solid conditions that affect the supercritical CO2 extraction of the lipids produced by microalgae, The Journal of Supercritical Fluids, 113, pp. 16–22. DOI: https://doi.org/10.1016/j.supflu.2016.03.001. DOI: https://doi.org/10.1016/j.supflu.2016.03.001

Wang, J. et al., 2015. From microalgae oil to produce novel structured triacylglycerols enriched with unsaturated fatty acids, Bioresource Technology, 184, pp. 405–414. DOI: https://doi.org/10.1016/j.biortech.2014.09.133. DOI: https://doi.org/10.1016/j.biortech.2014.09.133

Wang, X. et al., 2018. Heterogeneous expression of human PNPLA3 triggers algal lipid accumulation and lipid droplet enlargement, Algal Research, 31, pp. 276–281. DOI: https://doi.org/10.1016/j.algal.2018.02.019. DOI: https://doi.org/10.1016/j.algal.2018.02.019

Wang, X. et al., 2020. TAG pathway engineering via GPAT2 concurrently potentiates abiotic stress tolerance and oleaginicity in Phaeodactylum tricornutum, Biotechnology for Biofuels, 13(1), p. 160. DOI: https://doi.org/10.1186/s13068-020-01799-5. DOI: https://doi.org/10.1186/s13068-020-01799-5

Wood, D.A., 2021. Microalgae to biodiesel - Review of recent progress, Bioresource Technology Reports, 14, p. 100665. DOI: https://doi.org/10.1016/j.biteb.2021.100665. DOI: https://doi.org/10.1016/j.biteb.2021.100665

Xu, L. et al., 2011. A simple and rapid harvesting method for microalgae by in situ magnetic separation, Bioresource Technology, 102(21), pp. 10047–10051. DOI: https://doi.org/10.1016/j.biortech.2011.08.021. DOI: https://doi.org/10.1016/j.biortech.2011.08.021

Yaakob, M.A. et al., 2021. Influence of Nitrogen and Phosphorus on Microalgal Growth, Biomass, Lipid, and Fatty Acid Production: An Overview, Cells, 10(2), p. 393. DOI: https://doi.org/10.3390/cells10020393. DOI: https://doi.org/10.3390/cells10020393

Yan, J. et al., 2013. Overexpression of acetyl-CoA synthetase increased the biomass and fatty acid proportion in microalga Schizochytrium, Applied Microbiology and Biotechnology, 97(5), pp. 1933–1939. DOI: https://doi.org/10.1007/s00253-012-4481-6. DOI: https://doi.org/10.1007/s00253-012-4481-6

Yang, F. et al., 2018. Transcriptome analysis for phosphorus starvation-induced lipid accumulation in Scenedesmus sp, Scientific Reports, 8(1), p. 16420. DOI: https://doi.org/10.1038/s41598-018-34650-x. DOI: https://doi.org/10.1038/s41598-018-34650-x

Yao, Y. et al., 2014. Glycerol and neutral lipid production in the oleaginous marine diatom Phaeodactylum tricornutum promoted by overexpression of glycerol-3-phosphate dehydrogenase, Biotechnology for Biofuels, 7(1), p. 110. DOI: https://doi.org/10.1186/1754-6834-7-110. DOI: https://doi.org/10.1186/1754-6834-7-110

Yodsuwan, N., Sawayama, S. and Sirisansaneeyakul, S., 2017. Effect of nitrogen concentration on growth, lipid production and fatty acid profiles of the marine diatom Phaeodactylum tricornutum, Agriculture and Natural Resources, 51(3), pp. 190–197. DOI: https://doi.org/10.1016/j.anres.2017.02.004. DOI: https://doi.org/10.1016/j.anres.2017.02.004

Yoneda, K. et al., 2018. Homologous expression of lipid droplet protein-enhanced neutral lipid accumulation in the marine diatom Phaeodactylum tricornutum, Journal of Applied Phycology, 30(5), pp. 2793–2802. DOI: https://doi.org/10.1007/s10811-018-1402-9. DOI: https://doi.org/10.1007/s10811-018-1402-9

Yu, W.-L. et al., 2011. Modifications of the metabolic pathways of lipid and triacylglycerol production in microalgae, Microbial Cell Factories, 10(1), p. 91. DOI: https://doi.org/10.1186/1475-2859-10-91. DOI: https://doi.org/10.1186/1475-2859-10-91

Zarrinmehr, M.J. et al., 2020. Effect of nitrogen concentration on the growth rate and biochemical composition of the microalga, Isochrysis galbana, The Egyptian Journal of Aquatic Research, 46(2), pp. 153–158. DOI: https://doi.org/10.1016/j.ejar.2019.11.003. DOI: https://doi.org/10.1016/j.ejar.2019.11.003

Zhang, J. et al., 2014. Overexpression of the soybean transcription factor GmDof4 significantly enhances the lipid content of Chlorella ellipsoidea, Biotechnology for Biofuels, 7(1), p. 128. DOI: https://doi.org/10.1186/s13068-014-0128-4. DOI: https://doi.org/10.1186/s13068-014-0128-4

Zhang, S. et al., 2018. Alleviation of reactive oxygen species enhances PUFA accumulation in Schizochytrium sp. through regulating genes involved in lipid metabolism, Metabolic Engineering Communications, 6, pp. 39–48. DOI: https://doi.org/10.1016/j.meteno.2018.03.002. DOI: https://doi.org/10.1016/j.meteno.2018.03.002

Zhang, Y. et al., 2022. Assessment of enzyme addition strategies on the enhancement of lipid yield from microalgae, Biochemical Engineering Journal, 177, p. 108198. DOI: https://doi.org/10.1016/j.bej.2021.108198. DOI: https://doi.org/10.1016/j.bej.2021.108198

Zhou, J. et al., 2022. Extraction of lipids from microalgae using classical and innovative approaches, Food Chemistry, 384, p. 132236. DOI: https://doi.org/10.1016/j.foodchem.2022.132236. DOI: https://doi.org/10.1016/j.foodchem.2022.132236

Zhu, H. et al., 2010. Effect of functionalized MCM41 nanoparticles on syngas fermentation, Biomass and Bioenergy, 34(11), pp. 1624–1627. DOI: https://doi.org/10.1016/j.biombioe.2010.06.008. DOI: https://doi.org/10.1016/j.biombioe.2010.06.008

Zhu, S. et al., 2014. Metabolic changes of starch and lipid triggered by nitrogen starvation in the microalga Chlorella zofingiensis, Bioresource Technology, 152, pp. 292–298. DOI: https://doi.org/10.1016/j.biortech.2013.10.092. DOI: https://doi.org/10.1016/j.biortech.2013.10.092

Zhu, Z. et al., 2018. The synchronous TAG production with the growth by the expression of chloroplast transit peptide-fused ScPDAT in Chlamydomonas reinhardtii, Biotechnology for Biofuels, 11(1), p. 156. DOI: https://doi.org/10.1186/s13068-018-1160-6. DOI: https://doi.org/10.1186/s13068-018-1160-6