Biological production of synthetic aviation fuels (SAFs) from biomass

Abstract

Synthetic aviation fuels (SAFs) produced from biological routes are emerging as a practical way to decarbonize the aviation sector. This review summarizes progress in producing jet-range molecules from renewable carbon and benchmarks them against the Jet A and Jet A-1 property window defined by the standard specification for aviation turbine fuels (ASTM D1655). We group candidates into three families, fatty acid-derived streams, alcohol-based precursors, and isoprenoid-derived hydrocarbons, and map their properties and upgrading chemistry to the standard specification for aviation turbine fuel containing synthesized hydrocarbons (ASTM D7566) certification framework. We survey diverse microbial hosts on renewable feedstocks and highlight advances in pathway and chassis engineering and bioprocess intensification that raise titers, productivities, and carbon efficiency. Together, these advances chart a practical route to certified, scalable, and cost-competitive SAF supply.

Keywords

Synthetic aviation fuel / Sustainable aviation fuel / Standard specification of aviation fuel / Fatty acid-derived fuel / Alcohol-based fuel / Isoprenoid-derived fuel / Microbial hosts

References

[1] Abeln F, Chuck CJ. The history, state of the art and future prospects for oleaginous yeast research. Microb Cell Fact, 2021, 20: 221.

[2] Adamczyk PA, Hwang HJ, Chang T-H, Gao Y, Baidoo EEK, Kim J, Webb-Robertson B-JM, Flores JE, Quijano KC, Burnet MC, Munoz N, Sundstrom E, Gladden JM, Liu D. The oleaginous yeast Rhodosporidium toruloides engineered for biomass hydrolysate-derived (E)-α-bisabolene production. Metab Eng, 2025, 90: 92-105.

[3] Akita H, Shibata S, Komoriya T, Kamei S, Asamoto H, Matsumoto M. Simultaneous Saccharification and Fermentation for Isobutanol Production from Banana Peel. Fermentation, 2024, 10: 161.

[4] Alonso-Gutierrez J, Kim E-M, Batth TS, Cho N, Hu Q, Chan LJG, Petzold CJ, Hillson NJ, Adams PD, Keasling JD, Garcia Martin H, Lee TS. Principal component analysis of proteomics (PCAP) as a tool to direct metabolic engineering. Metab Eng, 2015, 28: 123-33.

[5] Argyros DA, Tripathi SA, Barrett TF, Rogers SR, Feinberg LF, Olson DG, Foden JM, Miller BB, Lynd LR, Hogsett DA, Caiazza NC. High ethanol titers from cellulose by using metabolically engineered thermophilic, anaerobic microbes. Appl Environ Microbiol, 2011, 77: 8288-94.

[6] Atnoorkar S, Wiatrowski M, Newes E, Davis R, Peterson S. Algae toHEFA: Economics and potential deployment in the United States. Biofuels Bioprod Bioref, 2024.

[7] Atsumi S, Hanai T, Liao JC. Non-fermentative pathways for synthesis of branched-chain higher alcohols as biofuels. Nature, 2008, 451: 86-9.

[8] Atsumi S, Wu T-Y, Eckl E-M, Hawkins SD, Buelter T, Liao JC. Engineering the isobutanol biosynthetic pathway in Escherichia coli by comparison of three aldehyde reductase/alcohol dehydrogenase genes. Appl Microbiol Biotechnol, 2010, 85: 651-7.

[9] Atsumi S, Wu T-Y, Machado IMP, Huang W-C, Chen P-Y, Pellegrini M, Liao JC. Evolution, genomic analysis, and reconstruction of isobutanol tolerance in Escherichia coli. Mol Syst Biol, 2010, 6: 449.

[10] Baena A, Orjuela A, Rakshit SK, Clark JH. Enzymatic hydrolysis of waste fats, oils and greases (FOGs): Status, prospective, and process intensification alternatives. Chem Eng Process - Process Intensif, 2022, 175: 108930.

[11] Banerjee D, Yunus IS, Wang X, Kim, Jinho, Srinivasan A, Menchavez R, Chen Y, Gin JW, Petzold CJ, Martin HG, Magnuson JK, Adams PD, Simmons BA, Mukhopadhyay A, Kim J, Lee TS. Genome-scale and pathway engineering for the sustainable aviation fuel precursor isoprenol production in Pseudomonas putida. Metab Eng, 2024, 82: 157-70.

[12] Baral NR, Yang M, Harvey BG, Simmons BA, Mukhopadhyay A, Lee TS, Scown CD. Production cost and carbon footprint of biomass-derived dimethylcyclooctane as a high-performance jet fuel blendstock. ACS Sustain Chem Eng, 2021, 9: 11872-82.

[13] Baral NR, Banerjee D, Mukhopadhyay A, Simmons BA, Singer SW, Scown CD. Integration of genome-scale metabolic model with biorefinery process model reveals market-competitive carbon-negative sustainable aviation fuel utilizing microbial cell mass lipids and biogenic CO2. BioResources, 2024, 19: 4056-86.

[14] Batten R, Karanjikar M, Spatari S. A sustainable aviation fuel pathway from biomass: life cycle environmental and cost evaluation for dimethylcyclooctane jet fuel. Sustainable Energy Fuels, 2024, 8: 1924-35.

[15] Bettencourt S, Miranda C, Pozdniakova TA, Sampaio P, Franco-Duarte R, Pais C. Single cell oil production by oleaginous yeasts grown in synthetic and waste-derived volatile fatty acids. Microorganisms, 2020, 8: 1809.

[16] Bi H, Xv C, Su C, Feng P, Zhang C, Wang M, Fang Y, Tan T. β-Farnesene Production from Low-Cost Glucose in Lignocellulosic Hydrolysate by Engineered Yarrowia lipolytica. Fermentation, 2022, 8: 532.

[17] Bierkandt T, Gaiser N, Bachmann J, Oßwald P, Köhler M. Terpene speciation: Analytical insights into the oxidation and pyrolysis of limonene and 1,8-cineole via molecular-beam mass spectrometry. Combust Flame, 2025, 272: 113854.

[18] biofuels international. 2017. BP and DuPont’s bio-isobutanol joint venture, Butamax, acquires ethanol facility in Kansas [WWW Document]. URL https://biofuels-news.com/news/bp-and-duponts-bio-isobutanol-joint-venture-butamax-acquires-ethanol-facility-in-kansas/?utm_source=chatgpt.com (accessed 10.2.25).

[19] Blakey S, Rye L, Wilson CW. Aviation gas turbine alternative fuels: A review. Proc Combust Inst, 2011, 33: 2863-85.

[20] Brennan TCR, Williams TC, Schulz BL, Palfreyman RW, Krömer JO, Nielsen LK. Evolutionary Engineering Improves Tolerance for Replacement Jet Fuels in Saccharomyces cerevisiae. Appl Environ Microbiol, 2015, 81: 3316-25.

[21] Bruder S, Moldenhauer EJ, Lemke RD, Ledesma-Amaro R, Kabisch J. Drop-in biofuel production using fatty acid photodecarboxylase from Chlorella variabilis in the oleaginous yeast Yarrowia lipolytica. Biotechnol Biofuels, 2019, 12: 202.

[22] Cai P, Li Y, Zhai X, Yao L, Ma X, Jia L, Zhou YJ. Microbial synthesis of long-chain α-alkenes from methanol by engineering Pichia pastoris. Bioresour Bioprocess, 2022, 9: 58.

[23] Cai P, Wu X, Deng J, Gao L, Shen Y, Yao L, Zhou YJ. Methanol biotransformation toward high-level production of fatty acid derivatives by engineering the industrial yeast Pichia pastoris. Proc Natl Acad Sci USA, 2022, 119: e2201711119.

[24] Cai W-L, Yu S-Y, Hu Y-H. Synergistic mechanisms of DGAT and PDAT in shaping triacylglycerol diversity: evolutionary insights and metabolic engineering strategies. Front Plant Sci, 2025, 16: 1598815.

[25] Calderon OR, Tao L, Abdullah Z, Moriarty K, Smolinski S, Milbrandt A, Talmadge M, Bhatt A, Zhang Y, Ravi V, Skangos C, Tan E, Payne C. 2024a. Sustainable Aviation Fuel (SAF) State-of-Industry Report:State of SAF Production Process. National Renewable Energy Laboratory. https://doi.org/10.2172/2426562

[26] Calderon OR, Tao L, Abdullah Z, Talmadge M, Milbrandt A, Smolinski S, Moriarty K, Bhatt A, Zhang Y, Ravi V, Skangos C, Davis R, Payne C. 2024b. Sustainable Aviation Fuel State-of-Industry Report: Hydroprocessed Esters and Fatty Acids Pathway. National Renewable Energy Laboratory. https://doi.org/10.2172/2426563

[27] Cao Y-X, Xiao W-H, Zhang J-L, Xie Z-X, Ding M-Z, Yuan Y-J. Heterologous biosynthesis and manipulation of alkanes in Escherichia coli. Metab Eng, 2016, 38: 19-28.

[28] Carruthers DN, Kim J, Mendez-Perez D, Monroe E, Myllenbeck N, Liu Y, Davis RW, Sundstrom E, Lee TS. Microbial production of high octane and high sensitivity olefinic ester biofuels. Biotechnol Biofuels Bioprod, 2023, 16: 60.

[29] Carruthers DN, Donnell I, Sundstrom E, Keasling JD, Lee TS. Prenol production in a microbial host via the Repass. Pathways Metab Eng, 2025, 88: 261-74.

[30] Carvalho LC, Oliveira ALS, Carsanba E, Lopes A, Leal T, Ribeiro M, Fernandes S, Pintado M, Oliveira C. Removal of phenolic compounds from sugarcane syrup and impact on Saccharomyces cerevisiae fermentation for beta-farnesene production. Biotechnol J, 2024, 19: 2300465.

[31] Chatterjee A, DeLorenzo DM, Carr R, Moon TS. Bioconversion of renewable feedstocks by Rhodococcus opacus. Curr Opin Biotechnol, 2020, 64: 10-6.

[32] Chen C-T, Liao JC. Frontiers in microbial 1-butanol and isobutanol production. FEMS Microbiol Lett, 2016, 363: fnw020.

[33] Chen B, Lee D-Y, Chang MW. Combinatorial metabolic engineering of Saccharomyces cerevisiae for terminal alkene production. Metab Eng, 2015, 31: 53-61.

[34] Choi YJ, Lee SY. Microbial production of short-chain alkanes. Nature, 2013, 502: 571-4.

[35] Christopher S. Evaluation of ARA Catalytic Hydrothermolysis (CH) Fuel. FAA Office of Environment and Energy; 2014.

[36] Chuck C, Donnelly J. The compatibility of potential bioderived fuels with Jet A-1 aviation kerosene. Appl Energy, 2014, 118: 83-91.

[37] Chung D, Cha M, Guss AM, Westpheling J. Direct conversion of plant biomass to ethanol by engineered Caldicellulosiruptor bescii. Proc Natl Acad Sci USA, 2014, 111: 8931-6.

[38] Cid Gomes L, Rana A, Berglund M, Wiklund P, Ottosson H. Light-driven (cross-)dimerization of terpenes as a route to renewable C₁₅ –C₃₀ crudes for fuel and lubricant oil applications. Sustainable Energy Fuels, 2023, 7: 868-82.

[39] Cid Gomes L, Vajravel S, Siljebo W, Rana A, Gustafsson T, Bairaktari A, Thomsen M, Ottosson H. Multiliter-Scale Photosensitized Dimerization of Isoprene to Sustainable Aviation Fuel Precursors. ACS Sustain Chem Eng, 2025, 13: 2467-76.

[40] Clomburg JM, Qian S, Tan Z, Cheong S, Gonzalez R. The isoprenoid alcohol pathway, a synthetic route for isoprenoid biosynthesis. Proc Natl Acad Sci USA, 2019, 116: 12810-5.

[41] Crépin L, Lombard E, Guillouet SE. Metabolic engineering of Cupriavidus necator for heterotrophic and autotrophic alka(e)ne production. Metab Eng, 2016, 37: 92-101.

[42] da Silva VZ, Ourique LJ, de David C, Ayub MAZ. Construction of Recombinant Klebsiella pneumoniae to Increase Ethanol Production on Residual Glycerol Fed-Batch Cultivations. Appl Biochem Biotechnol, 2020, 192: 1147-62.

[43] de Pastore AE, Wrobel RL, Paul B, Anthony LC, Sato TK, Zhang Y, Hittinger CT, Maravelias CT. High yield co-production of isobutanol and ethanol from switchgrass: experiments, and process synthesis and analysis. Sustainable Energy Fuels, 2023, 7: 3266-75.

[44] de Pastore AE, Coplien J, Anthony LC, Sato TK, Zhang Y, Karlen SD, Hittinger CT, Maravelias CT. On the synthesis of biorefineries for high-yield isobutanol production: from biomass-to-alcohol experiments to system level analysis. RSC Sustain, 2024, 2: 2532-40.

[45] Degenhardt J, Köllner TG, Gershenzon J. Monoterpene and sesquiterpene synthases and the origin of terpene skeletal diversity in plants. Phytochemistry, 2009, 70: 1621-37.

[46] Dellomonaco C, Clomburg JM, Miller EN, Gonzalez R. Engineered reversal of the β-oxidation cycle for the synthesis of fuels and chemicals. Nature, 2011, 476: 355-9.

[47] Desai SH, Rabinovitch-Deere CA, Fan Z, Atsumi S. Isobutanol production from cellobionic acid in Escherichia coli. Microb Cell Fact, 2015, 14: 52.

[48] Devarapalli M, Lewis R, Atiyeh H. Continuous Ethanol Production from Synthesis Gas by Clostridium ragsdalei in a Trickle-Bed Reactor. Fermentation, 2017, 3: 23.

[49] Díaz MJ, Moya M, Castro E. Bioethanol Production from Steam-Exploded Barley Straw by Co-Fermentation with Escherichia coli SL100. Agronomy. 2022;12:874. https://doi.org/10.3390/agronomy12040874

[50] Ding J, You S, Ba W, Zhang H, Chang H, Qi W, Su R, He Z. Bifunctional utilization of whey powder as a substrate and inducer for β-farnesene production in an engineered Escherichia coli. Bioresour Technol, 2021, 341: 125739.

[51] Dischl R, Sauer D, Voigt C, Harlaß T, Sakellariou F, Märkl R, Schumann U, Scheibe M, Kaufmann S, Roiger A, Dörnbrack A, Renard C, Gauthier M, Swann P, Madden P, Luff D, Johnson M, Ahrens D, Sallinen R, Schripp T, Eckel G, Bauder U, Le Clercq P. Measurements of particle emissions of an A350-941 burning 100% sustainable aviation fuels in cruise. Atmos Chem Phys, 2024, 24: 11255-73.

[52] Dobrowolski A, Drzymała K, Mituła P, Mirończuk AM. Production of tailor-made fatty acids from crude glycerol at low pH by Yarrowia lipolytica. Bioresour Technol, 2020, 314: 123746.

[53] Dong T, Fei Q, Genelot M, Smith H, Laurens LML, Watson MJ, Pienkos PT. A novel integrated biorefinery process for diesel fuel blendstock production using lipids from the methanotroph, Methylomicrobium buryatense. Energy Conv Manag, 2017, 140: 62-70.

[54] Dong C, Wang Y, Zhang H, Leu S-Y. Feasibility of high-concentration cellulosic bioethanol production from undetoxified whole Monterey pine slurry. Bioresour Technol, 2018, 250: 102-9.

[55] Donoso D, García D, Ballesteros R, Lapuerta M, Canoira L. Hydrogenated or oxyfunctionalized turpentine: options for automotive fuel components. RSC Adv, 2021, 11: 18342-50.

[56] Donoso D, Bolonio D, Ballesteros R, Lapuerta M, Canoira L. Hydrogenated orange oil: A waste derived drop-in biojet fuel. Renew Energy, 2022, 188: 1049-58.

[57] Dou X, Li M, Ge Y, Yin G, Wang X, Xue S, Jia B, Zi L, Wan H, Xi Y, Chi Z, Kong F. Photoproduction of Aviation Fuel β-Caryophyllene From the Eukaryotic Green Microalga Chlamydomonas reinhardtii. Biotechnol Bioeng, 2025, 122: 698-709.

[58] Dragomanova S, Tancheva L, Georgieva M. A review: Biological activity of myrtenal and some myrtenal-containing medicinal plant essential oils. Scripta Scientifica Pharm, 2018, 5: 22.

[59] Du R, Su R, Li X, Tantai X, Liu Z, Yang J, Qi W, He Z. Controlled adsorption of cellulase onto pretreated corncob by pH adjustment. Cellulose, 2012, 19: 371-80.

[60] El-Maghraby RM. A Study on Bio-Diesel and Jet Fuel Blending for the Production of Renewable Aviation Fuel. MSF, 2020, 1008: 231-44.

[61] Fackler N, Heijstra BD, Rasor BJ, Brown H, Martin J, Ni Z, Shebek KM, Rosin RR, Simpson SD, Tyo KE, Giannone RJ, Hettich RL, Tschaplinski TJ, Leang C, Brown SD, Jewett MC, Köpke M. Stepping on the Gas to a Circular Economy: Accelerating Development of Carbon-Negative Chemical Production from Gas Fermentation. Annu Rev Chem Biomol Eng, 2021, 12: 439-70.

[62] Fang L, Hao X, Fan J, Liu X, Chen Y, Wang L, Huang X, Song H, Cao Y. Genome-scale CRISPRi screen identifies pcnB repression conferring improved physiology for overproduction of free fatty acids in Escherichia coli. Nat Commun, 2025, 16: 3060.

[63] Feng P, Sun B, Bi H, Bao Y, Wang M, Zhang H, Fang Y. Developing Thermosensitive Metabolic Regulation Strategies in the Fermentation Process of Saccharomyces cerevisiae to Enhance α-Bisabolene Production. ACS Synth Biol, 2025, 14: 1129-41.

[64] Förster AH, Gescher J. Metabolic Engineering of Escherichia coli for Production of Mixed-Acid Fermentation End Products. Front Bioeng Biotechnol, 2014, 2: 16.

[65] Gao M, Ploessl D, Shao Z. Enhancing the Co-utilization of Biomass-Derived Mixed Sugars by Yeasts. Front Microbiol, 2018, 9: 3264.

[66] Gao R, Li Z, Zhou X, Bao W, Cheng S, Zheng L. Enhanced lipid production by Yarrowia lipolytica cultured with synthetic and waste-derived high-content volatile fatty acids under alkaline conditions. Biotechnol Biofuels, 2020, 13: 3.

[67] Gao J, Li Y, Yu W, Zhou YJ. Rescuing yeast from cell death enables overproduction of fatty acids from sole methanol. Nat Metab, 2022, 4: 932-43.

[68] Gao L, Zhang K, Shen Y, Cai P, Zhou YJ. Engineering a versatile yeast platform for sesquiterpene production from glucose or methanol. Biotechnol J, 2024, 19: 2400261.

[69] Gao L, Hou R, Cai P, Yao L, Wu X, Li Y, Zhang L, Zhou YJ. Engineering Yeast Peroxisomes for α-Bisabolene Production from Sole Methanol with the Aid of Proteomic Analysis. JACS Au, 2024, 4: 2474-83.

[70] Geiselman GM, Kirby J, Landera A, Otoupal P, Papa G, Barcelos C, Sundstrom ER, Das L, Magurudeniya HD, Wehrs M, Rodriguez A, Simmons BA, Magnuson JK, Mukhopadhyay A, Lee TS, George A, Gladden JM. Conversion of poplar biomass into high-energy density tricyclic sesquiterpene jet fuel blendstocks. Microb Cell Fact, 2020, 19: 208.

[71] Geleynse S, Brandt K, Garcia-Perez M, Wolcott M, Zhang X. The Alcohol-to-Jet Conversion Pathway for Drop-In Biofuels: Techno-Economic Evaluation. Chemsuschem, 2018, 11: 3728-41.

[72] George KW, Thompson MG, Kang A, Baidoo E, Wang G, Chan LJG, Adams PD, Petzold CJ, Keasling JD, Lee TS. Metabolic engineering for the high-yield production of isoprenoid-based C₅ alcohols in E. coli. Sci Rep, 2015, 5: 11128.

[73] Gevo. An overview of Gevo’s biobased isobutanol production process. Gevo 2023.

[74] Gevo. 2025. Developing Isobutanol and Ethanol Processes in Luverne, Minnesota [WWW Document]. URL https://gevo.com/location/luverne-mn/ (accessed 7.10.26).

[75] Główka M, Wójcik J, Boberski P, Białecki T, Gawron B, Skolniak M, Suchocki T. Sustainable aviation fuel – Comprehensive study on highly selective isomerization route towards HEFA based bioadditives. Renew Energy, 2024, 220: 119696.

[76] Gomes D, Cruz M, de Resende M, Ribeiro E, Teixeira J, Domingues L. Very high gravity bioethanol revisited: main challenges and advances. Fermentation, 2021, 7: 38.

[77] GreenAir News. 2014. Amyris/Total renewable jet fuel gets ASTM green light as SIP fuel for commercial aviation [WWW Document]. URL https://archives.greenairnews.com/www.greenaironline.com/newsddff.html?viewStory=1937 (accessed 7.10.26).

[78] Gu P, Li F, Huang Z. Engineering Escherichia coli for Isobutanol Production from Xylose or Glucose-Xylose Mixture. Microorganisms. 2023;11. https://doi.org/10.3390/microorganisms11102573.

[79] Gu Z, Chai T, Chen X. Evolutionary engineering of Pichia pastoris for α-farnesene biosynthesis from methanol. Syst Microbiol Biomanuf, 2025, 5: 1039-49.

[80] Guo S, Du J, Li D, Xiong J, Chen Y. Versatile xylose and arabinose genetic switches development for yeasts. Metab Eng, 2025, 87: 21-36.

[81] Hamilton L, Luning-Prak D, Cowart J, McDaniel A, Williams S, Leung R. Direct sugar to hydrocarbon (DSH) fuel performance evaluation in multiple diesel engines. SAE Int J Fuels Lubr, 2014, 7: 270-82.

[82] Han GH, Kim SK, Yoon PK-S, Kang Y, Kim BS, Fu Y, Sung BH, Jung HC, Lee D-H, Kim S-W, Lee S-G. Fermentative production and direct extraction of (-)-α-bisabolol in metabolically engineered Escherichia coli. Microb Cell Fact, 2016, 15: 185.

[83] Harrison KW, Harvey BG. Renewable high density fuels containing tricyclic sesquiterpanes and alkyl diamondoids. Sustainable Energy Fuels, 2017, 1: 467-73.

[84] Harvey BG. Fuels and lubricants from bisaboline. US10053643B1; 2018.

[85] Harvey BG, Wright ME, Quintana RL. High-Density Renewable Fuels Based on the Selective Dimerization of Pinenes. Energy Fuels, 2010, 24: 267-73.

[86] Harvey BG, Meylemans HA, Gough RV, Quintana RL, Garrison MD, Bruno TJ. High-density biosynthetic fuels: the intersection of heterogeneous catalysis and metabolic engineering. Phys Chem Chem Phys, 2014, 16: 9448-57.

[87] Harvey BG, Merriman WW, Koontz TA. High-Density Renewable Diesel and Jet Fuels Prepared from Multicyclic Sesquiterpanes and a 1-Hexene-Derived Synthetic Paraffinic Kerosene. Energy Fuels, 2015, 29: 2431-6.

[88] Higashide W, Li Y, Yang Y, Liao JC. Metabolic engineering of Clostridium cellulolyticum for production of isobutanol from cellulose. Appl Environ Microbiol, 2011, 77: 2727-33.

[89] Hill P, Benjamin K, Bhattacharjee B, Garcia F, Leng J, Liu C-L, Murarka A, Pitera D, Rodriguez Porcel EM, da Silva I, Kraft C. Clean manufacturing powered by biology: how Amyris has deployed technology and aims to do it better. J Ind Microbiol Biotechnol, 2020, 47: 965-75.

[90] Holladay J, Abdullah Z, Heyne J. Sustainable Aviation Fuel: Review of Technical Pathways. Department of Energy 2020.

[91] Hu P, Chakraborty S, Kumar A, Woolston B, Liu H, Emerson D, Stephanopoulos G. Integrated bioprocess for conversion of gaseous substrates to liquids. Proc Natl Acad Sci USA, 2016, 113: 3773-8.

[92] Huang Q, Kamal R, Shen H, Lu H, Song J, Chu Y, Xue C, Zhao ZK. Pilot-scale conversion of corn stover into lipids by the red yeast Rhodosporidium toruloides. J Environ Chem Eng, 2022, 10: 108858.

[93] Huang Y, Ye Z, Wan X, Yao G, Duan J, Liu J, Yao M, Sun X, Deng Z, Shen K, Jiang H, Liu T. Systematic mining and evaluation of the sesquiterpene skeletons as high energy aviation fuel molecules. Adv Sci (Weinh), 2023, 10: e2300889.

[94] Huq NA, Hafenstine GR, Huo X, Nguyen H, Tifft SM, Conklin DR, Stück D, Stunkel J, Yang Z, Heyne JS, Wiatrowski MR, Zhang Y, Tao L, Zhu J, McEnally CS, Christensen ED, Hays C, Van Allsburg KM, Unocic KA, Meyer HM, Abdullah Z, Vardon DR. Toward net-zero sustainable aviation fuel with wet waste-derived volatile fatty acids. Proc Natl Acad Sci USA. 2021;118. https://doi.org/10.1073/pnas.2023008118.

[95] IATA. Net zero 2050: sustainable aviation fuels (SAF). IATA; 2025.

[96] Jain S, Kumar S. A comprehensive review of bioethanol production from diverse feedstocks: Current advancements and economic perspectives. Energy, 2024, 296: 131130.

[97] Jansen MLA, Bracher JM, Papapetridis I, Verhoeven MD, de Bruijn H, de Waal PP, van Maris AJA, Klaassen P, Pronk JT. Saccharomyces cerevisiae strains for second-generation ethanol production: from academic exploration to industrial implementation. FEMS Yeast Res 17, 2017.

[98] Jung H-M, Lee JY, Lee J-H, Oh M-K. Improved production of isobutanol in pervaporation-coupled bioreactor using sugarcane bagasse hydrolysate in engineered Enterobacter aerogenes. Bioresour Technol, 2018, 259: 373-80.

[99] Kamal R, Liu Y, Li Q, Huang Q, Wang Q, Yu X, Zhao ZK. Exogenous l-proline improved Rhodosporidium toruloides lipid production on crude glycerol. Biotechnol Biofuels, 2020, 13: 159.

[100] Kang A, George KW, Wang G, Baidoo E, Keasling JD, Lee TS. Isopentenyl diphosphate (IPP)-bypass mevalonate pathways for isopentenol production. Metab Eng, 2016, 34: 25-35.

[101] Kang A, Mendez-Perez D, Goh E-B, Baidoo EEK, Benites VT, Beller HR, Keasling JD, Adams PD, Mukhopadhyay A, Lee TS. Optimization of the IPP-bypass mevalonate pathway and fed-batch fermentation for the production of isoprenol in Escherichia coli. Metab Eng, 2019, 56: 85-96.

[102] Keller CL, Doppalapudi KR, Woodroffe J-D, Harvey BG. Solvent-free dehydration, cyclization, and hydrogenation of linalool with a dual heterogeneous catalyst system to generate a high-performance sustainable aviation fuel. Commun Chem, 2022, 5: 113.

[103] Khamwachirapithak P, Sae-Tang K, Mhuantong W, Tanapongpipat S, Zhao X-Q, Liu C-G, Wei D-Q, Champreda V, Runguphan W. Optimizing Ethanol Production in Saccharomyces cerevisiae at Ambient and Elevated Temperatures through Machine Learning-Guided Combinatorial Promoter Modifications. ACS Synth Biol, 2023, 12: 2897-908.

[104] Khattab SMR, Watanabe T. Efficient Conversion of Glycerol to Ethanol by an Engineered Saccharomyces cerevisiae Strain. Appl Environ Microbiol, 2021, 87: e0026821.

[105] Kim J-H, Wang C, Jang H-J, Cha M-S, Park J-E, Jo S-Y, Choi E-S, Kim S-W. Isoprene production by Escherichia coli through the exogenous mevalonate pathway with reduced formation of fermentation byproducts. Microb Cell Fact, 2016, 15: 214.

[106] Kim J, Baidoo EEK, Amer B, Mukhopadhyay A, Adams PD, Simmons BA, Lee TS. Engineering Saccharomyces cerevisiae for isoprenol production. Metab Eng, 2021, 64: 154-66.

[107] Ko JK, Um Y, Woo HM, Kim KH, Lee S-M. Ethanol production from lignocellulosic hydrolysates using engineered Saccharomyces cerevisiae harboring xylose isomerase-based pathway. Bioresour Technol, 2016, 209: 290-6.

[108] Kothandaraman J, Cosimbescu L, Swita MS. Solvent-Induced Selectivity of Isoprene From Bio-Derived Prenol. Front Chem, 2022, 10: 879129.

[109] Kotogán A, Furka ZT, Kovács T, Volford B, Papp DA, Varga M, Huynh T, Szekeres A, Papp T, Vágvölgyi C, Mondal KC, Kerekes EB, Takó M. Hydrolysis of Edible Oils by Fungal Lipases: An Effective Tool to Produce Bioactive Extracts with Antioxidant and Antimicrobial Potential. Foods. 2022;11. https://doi.org/10.3390/foods11121711.

[110] Kugler S, Ossowicz P, Malarczyk-Matusiak K, Wierzbicka E. Advances in Rosin-Based Chemicals: The Latest Recipes, Applications and Future Trends. Molecules. 2019;24. https://doi.org/10.3390/molecules24091651.

[111] Kumar M, Saha A, deva P P, Karmakar S, Reddy V. , Combustion characteristics of potential alternative aviation fuels and Jet A-1 in a swirl-stabilized combustor. 2nd Natl. Aero Propuls. Conf., Indian Inst. Technol. Kharagpur, Kharagpur, West Bengal, India. 2018.

[112] Kurosawa K, Boccazzi P, de Almeida NM, Sinskey AJ. High-cell-density batch fermentation of Rhodococcus opacus PD630 using a high glucose concentration for triacylglycerol production. J Biotechnol, 2010, 147: 212-8.

[113] Lange J, Müller F, Takors R, Blombach B. Harnessing novel chromosomal integration loci to utilize an organosolv-derived hemicellulose fraction for isobutanol production with engineered Corynebacterium glutamicum. Microb Biotechnol, 2018, 11: 257-63.

[114] LanzaJet. 2024. Freedom Pines Fuels [WWW Document]. URL https://www.lanzajet.com/freedom-pines (accessed 7.10.26).

[115] Ledesma-Amaro R, Nicaud J-M. Yarrowia lipolytica as a biotechnological chassis to produce usual and unusual fatty acids. Prog Lipid Res, 2016, 61: 40-50.

[116] Lee J-E, Vadlani PV, Guragain YN, San K-Y, Min D-H. Production of free fatty acids from switchgrass using recombinant Escherichia coli. Biotechnol Prog, 2018, 34: 91-8.

[117] Lee D, Park H, Kim J-E, Kim Y, Park JH, Lee H, Yoon BH, Han B, Jung JY, Cha S, Lee P, Hahn J-S. Engineering Yarrowia lipolytica for enhanced lipid productivity in nutrient-rich conditions: A scalable approach to microbial lipid production. Metab Eng, 2025, 91: 302-12.

[118] Lee S, Pintor DL, Cho S. Enabling ethanol mixing-controlled compression-ignition combustion using 2-Ethylhexyl nitrate in an off-road engine. Appl Energy, 2025, 377: 124445.

[119] Lehtinen T, Virtanen H, Santala S, Santala V. Production of alkanes from CO2 by engineered bacteria. Biotechnol Biofuels, 2018, 11: 228.

[120] Lei Y, Wang X, Sun S, He B, Sun W, Wang K, Chen Z, Guo Z, Li Z. A review of lipid accumulation by oleaginous yeasts: Culture mode. Sci Total Environ, 2024, 919: 170385.

[121] Lennen RM, Pfleger BF. Engineering Escherichia coli to synthesize free fatty acids. Trends Biotechnol, 2012, 30: 659-67.

[122] Li Y, Zhao ZK, Bai F. High-density cultivation of oleaginous yeast Rhodosporidium toruloides Y4 in fed-batch culture. Enzyme Microb Technol, 2007, 41: 312-7.

[123] Li H, Opgenorth PH, Wernick DG, Rogers S, Wu T-Y, Higashide W, Malati P, Huo Y-X, Cho KM, Liao JC. Integrated electromicrobial conversion of CO2 to higher alcohols. Science, 2012, 335: 1596.

[124] Li J, Ma Y, Liu N, Eser BE, Guo Z, Jensen PR, Stephanopoulos G. Synthesis of high-titer alka(e)nes in Yarrowia lipolytica is enabled by a discovered mechanism. Nat Commun, 2020, 11: 6198.

[125] Li S, Rong L, Wang S, Liu S, Lu Z, Miao L, Zhao B, Zhang C, Xiao D, Pushpanathan K, Wong A, Yu A. Enhanced limonene production by metabolically engineered Yarrowia lipolytica from cheap carbon sources. Chem Eng Sci, 2022, 249: 117342.

[126] Lin PP, Mi L, Morioka AH, Yoshino KM, Konishi S, Xu SC, Papanek BA, Riley LA, Guss AM, Liao JC. Consolidated bioprocessing of cellulose to isobutanol using Clostridium thermocellum. Metab Eng, 2015, 31: 44-52.

[127] Lin H-H, Mendez-Perez D, Park J, Wang X, Cheng Y, Huo J, Mukhopadhyay A, Lee TS, Shanks BH. Precursor prioritization for p-cymene production through synergistic integration of biology and chemistry. Biotechnol Biofuels Bioprod, 2022, 15: 126.

[128] Liu X, Sheng J, Curtiss R. Fatty acid production in genetically modified cyanobacteria. Proc Natl Acad Sci USA, 2011, 108: 6899-904.

[129] Liu Z, Inokuma K, Ho S-H, Haan R, den, Hasunuma T, van Zyl WH, Kondo A. Combined cell-surface display- and secretion-based strategies for production of cellulosic ethanol with Saccharomyces cerevisiae. Biotechnol Biofuels, 2015, 8: 162.

[130] Liu J, Yuan M, Liu J-N, Huang X-F. Bioconversion of mixed volatile fatty acids into microbial lipids by Cryptococcus curvatus ATCC 20509. Bioresour Technol, 2017, 241: 645-51.

[131] Liu C-L, Tian T, Alonso-Gutierrez J, Garabedian B, Wang S, Baidoo EEK, Benites V, Chen Y, Petzold CJ, Adams PD, Keasling JD, Tan T, Lee TS. Renewable production of high density jet fuel precursor sesquiterpenes from Escherichia coli. Biotechnol Biofuels, 2018, 11: 285.

[132] Liu Y, Jiang X, Cui Z, Wang Z, Qi Q, Hou J. Engineering the oleaginous yeast Yarrowia lipolytica for production of α-farnesene. Biotechnol Biofuels, 2019, 12: 296.

[133] Liu S-C, Liu Z, Wei L-J, Hua Q. Pathway engineering and medium optimization for α-farnesene biosynthesis in oleaginous yeast Yarrowia lipolytica. J Biotechnol, 2020, 319: 74-81.

[134] Liu H, Chen S-L, Xu J-Z, Zhang W-G. Dual Regulation of Cytoplasm and Peroxisomes for Improved Α-Farnesene Production in Recombinant Pichia pastoris. ACS Synth Biol, 2021, 10: 1563-73.

[135] Liu Y, Benitez MG, Chen J, Harrison E, Khusnutdinova AN, Mahadevan R. Opportunities and Challenges for Microbial Synthesis of Fatty Acid-Derived Chemicals (FACs). Front Bioeng Biotechnol, 2021, 9: 613322.

[136] Liu Y, Wang Z, Cui Z, Qi Q, Hou J. α-Farnesene production from lipid by engineered Yarrowia lipolytica. Bioresour Bioprocess, 2021, 8: 78.

[137] Liu Y, Zhang J, Li Q, Wang Z, Cui Z, Su T, Lu X, Qi Q, Hou J. Engineering Yarrowia lipolytica for the sustainable production of β-farnesene from waste oil feedstock. Biotechnol Biofuels Bioprod, 2022, 15: 101.

[138] Liu J, Lin M, Han P, Yao G, Jiang H. Biosynthesis Progress of High-Energy-Density Liquid Fuels Derived from Terpenes. Microorganisms. 2024;12. https://doi.org/10.3390/microorganisms12040706.

[139] Lu J, Brigham CJ, Gai CS, Sinskey AJ. Studies on the production of branched-chain alcohols in engineered Ralstonia eutropha. Appl Microbiol Biotechnol, 2012, 96: 283-97.

[140] Luckie BA, Kashyap M, Pearson AN, Chen Y, Liu Y, Valencia LE, Carrillo Romero A, Hudson GA, Tao XB, Wu B, Petzold CJ, Keasling JD. Development of Corynebacterium glutamicum as a monoterpene production platform. Metab Eng, 2024, 81: 110-22.

[141] Luo Y, He Y, Zhang R, Lou D, Zhu R, Zhu C, Li Q. A novel integrated process for selective preparation of jet-range fuels from low carbon alcohols and ABE. Fuel Process Technol, 2023, 241: 107603.

[142] Ma K, He M, You H, Pan L, Hu G, Cui Y, Maeda T. Enhanced fuel ethanol production from rice straw hydrolysate by an inhibitor-tolerant mutant strain of Scheffersomyces stipitis. RSC Adv, 2017, 7: 31180-8.

[143] Ma Z-X, Zhang M, Zhang C-T, Zhang H, Mo X-H, Xing X-H, Yang S. Metabolomic analysis improves bioconversion of methanol to isobutanol in Methylorubrum extorquens AM1. Biotechnol J, 2021, 16: e2000413.

[144] Matsuda F, Ishii J, Kondo T, Ida K, Tezuka H, Kondo A. Increased isobutanol production in Saccharomyces cerevisiae by eliminating competing pathways and resolving cofactor imbalance. Microb Cell Fact, 2013, 12: 119.

[145] Matsudaira A, Hoshino Y, Uesaka K, Takatani N, Omata T, Usuda Y. Production of glutamate and stereospecific flavors, (S)-linalool and (+)-valencene, by Synechocystis sp. PCC6803. J Biosci Bioeng, 2020, 130: 464-70.

[146] Meadows AL, Hawkins KM, Tsegaye Y, Antipov E, Kim Y, Raetz L, Dahl RH, Tai A, Mahatdejkul-Meadows T, Xu L, Zhao L, Dasika MS, Murarka A, Lenihan J, Eng D, Leng JS, Liu C-L, Wenger JW, Jiang H, Chao L, Westfall P, Lai J, Ganesan S, Jackson P, Mans R, Platt D, Reeves CD, Saija PR, Wichmann G, Holmes VF, Benjamin K, Hill PW, Gardner TS, Tsong AE. Rewriting yeast central carbon metabolism for industrial isoprenoid production. Nature, 2016, 537: 694-7.

[147] Melcher F, Vogelgsang F, Haack M, Masri M, Ringel M, Roth A, Garbe D, Brück T. Lipase-mediated plant oil hydrolysis—Toward a quantitative glycerol recovery for the synthesis of pure allyl alcohol and acrylonitrile. Eur J Lipid Sci Technol. 2023;125. https://doi.org/10.1002/ejlt.202200196.

[148] Meylemans HA, Quintana RL, Goldsmith BR, Harvey BG. Solvent-free conversion of linalool to methylcyclopentadiene dimers: a route to renewable high-density fuels. Chemsuschem, 2011, 4: 465-9.

[149] Miao R, Xie H, Lindblad P. Enhancement of photosynthetic isobutanol production in engineered cells of Synechocystis PCC 6803. Biotechnol Biofuels, 2018, 11: 267.

[150] Minty JJ, Singer ME, Scholz SA, Bae C-H, Ahn J-H, Foster CE, Liao JC, Lin XN. Design and characterization of synthetic fungal-bacterial consortia for direct production of isobutanol from cellulosic biomass. Proc Natl Acad Sci USA, 2013, 110: 14592-7.

[151] Misra P, Alvarez-Majmutov A, Chen J. Isomerization catalysts and technologies for biorefining: Opportunities for producing sustainable aviation fuels. Fuel, 2023, 351: 128994.

[152] Monteiro RRC, dos Santos IA, Arcanjo MRA, Cavalcante CL, de Luna FMT, Fernandez-Lafuente R, Vieira RS. Production of jet biofuels by catalytic hydroprocessing of esters and fatty acids: A review. Catalysts, 2022, 12: 237.

[153] Moriarty K, McCormick R. Sustainable Aviation Fuel Blending and Logistics. National Renewable Energy Laboratory; 2024.

[154] Moser S, Pichler H. Identifying and engineering the ideal microbial terpenoid production host. Appl Microbiol Biotechnol, 2019, 103: 5501-16.

[155] Muldoon JA, Harvey BG. Bio-Based Cycloalkanes: The Missing Link to High-Performance Sustainable Jet Fuels. Chemsuschem, 2020, 13: 5777-807.

[156] Muller G, de Godoy VR, Dário MG, Duval EH, Alves-Jr SL, Bücker A, Rosa CA, Dunn B, Sherlock G, Stambuk BU. Improved Sugarcane-Based Fermentation Processes by an Industrial Fuel-Ethanol Yeast Strain. J Fungi (Basel). 2023;9. https://doi.org/10.3390/jof9080803.

[157] Naqvi YA, Mekala Y, Ravi R, Krishnaiah G, Zschocke A, Hull A, Normark M. PureSAF^SM (Alcohol-to-Jet) Fuel Technology: Enabling Sustainable Aviation Fuel. KBR Technical Journal; 2023.

[158] Nazemieh A, Acree WE, Jouyban A. Further computations on physico-chemical properties of binary mixtures of p-cymene with α-pinene, limonene and citral. J Mol Liq, 2022, 350: 118211.

[159] Nazloo EK, Danesh M, Sarrafzadeh M-H, Moheimani NR, Ennaceri H. Biomass and hydrocarbon production from Botryococcus braunii: A review focusing on cultivation methods. Sci Total Environ, 2024, 926: 171734.

[160] Neste. 2025. Sustainable aviation fuel [WWW Document]. URL https://www.neste.com/en-us/products-and-innovation/sustainable-aviation-fuel (accessed 7.10.26).

[161] Ni X, Li J, Yu W, Bai F, Zhao ZK, Gao J, Yang F, Zhou YJ. High-level production of free fatty acids from lignocellulose hydrolysate by co-utilizing glucose and xylose in yeast. Synth Syst Biotechnol, 2025, 10: 401-9.

[162] Ochsenreither K, Glück C, Stressler T, Fischer L, Syldatk C. Production strategies and applications of microbial single cell oils. Front Microbiol, 2016, 7: 1539.

[163] Ojagh H, Creaser D, Salam MA, Grennfelt EL, Olsson L. Hydroconversion of abietic acid into value-added fuel components over sulfided NiMo catalysts with varying support acidity. Fuel Process Technol, 2019, 190: 55-66.

[164] Okonkwo O, Dou C, Oksen E, Narani A, Marcondes W, Chen X, Kim J, Gao Y, Burnet MC, Webb-Robertson B-JM, Poirier BC, Tanjore D, Magnuson JK, Munoz NM, Gardner J. Corn stover variability drives differences in bisabolene production by engineered Rhodotorula toruloides. J Ind Microbiol Biotechnol. 2024;51. https://doi.org/10.1093/jimb/kuae034.

[165] Olson AL, Tunér M, Verhelst S. A review of isobutanol as a fuel for internal combustion engines. Energies, 2023, 16: 7470.

[166] Oßwald P, Whitside R, Schäffer J, Köhler M. An experimental flow reactor study of the combustion kinetics of terpenoid jet fuel compounds: Farnesane, p-menthane and p-cymene. Fuel, 2017, 187: 43-50.

[167] Özaydın B, Burd H, Lee TS, Keasling JD. Carotenoid-based phenotypic screen of the yeast deletion collection reveals new genes with roles in isoprenoid production. Metab Eng, 2013, 15: 174-83.

[168] Park Y-K, Sellés Vidal L, Bell D, Zabret J, Soldat M, Kavšček M, Ledesma-Amaro R. Efficient synthesis of limonene production in Yarrowia lipolytica by combinatorial engineering strategies. Biotechnol Biofuels Bioprod, 2024, 17: 94.

[169] Park Y-K, Studena L, Hapeta P, Haddouche R, Bell DJ, Torres-Montero P, Martinez JL, Nicaud J-M, Botes A, Ledesma-Amaro R. Efficient biosynthesis of β-caryophyllene by engineered Yarrowia lipolytica. Microb Cell Fact, 2025, 24: 38.

[170] Pavan M, Reinmets K, Garg S, Mueller AP, Marcellin E, Köpke M, Valgepea K. Advances in systems metabolic engineering of autotrophic carbon oxide-fixing biocatalysts towards a circular economy. Metab Eng, 2022, 71: 117-41.

[171] Pereira FB, Guimarães PMR, Teixeira JA, Domingues L. Optimization of low-cost medium for very high gravity ethanol fermentations by Saccharomyces cerevisiae using statistical experimental designs. Bioresour Technol, 2010, 101: 7856-63.

[172] Peters MA, Alves CT, Onwudili JA. A Review of Current and Emerging Production Technologies for Biomass-Derived Sustainable Aviation Fuels. Energies, 2023, 16: 6100.

[173] Phulara SC, Chaturvedi P, Chaurasia D, Diwan B, Gupta P. Modulation of culture medium confers high-specificity production of isopentenol in Bacillus subtilis. J Biosci Bioeng, 2019, 127: 458-64.

[174] Qiao K, Wasylenko TM, Zhou K, Xu P, Stephanopoulos G. Lipid production in Yarrowia lipolytica is maximized by engineering cytosolic redox metabolism. Nat Biotechnol, 2017, 35: 173-7.

[175] Ren Y, Liu S, Jin G, Yang X, Zhou YJ. Microbial production of limonene and its derivatives: Achievements and perspectives. Biotechnol. Adv, 2020, 44: 107628.

[176] RFA. Ethanol industry outlook. Renewable fuels association; 2024.

[177] Richter H, Martin M, Angenent L. A Two-Stage Continuous Fermentation System for Conversion of Syngas into Ethanol. Energies, 2013, 6: 3987-4000.

[178] Rinaldi MA, Tait S, Toogood HS, Scrutton NS. Bioproduction of linalool from paper mill waste. Front Bioeng Biotechnol, 2022, 10: 892896.

[179] Rodewald D, Severson SM. Gevo Produces First Cellulosic Renewable Jet Fuel Specified for Use on Commercial Airline Flights. Gevo 2016.

[180] Rolf J, Julsing MK, Rosenthal K, Lütz S. A Gram-Scale Limonene Production Process with Engineered Escherichia coli. Molecules. 2020;25. https://doi.org/10.3390/molecules25081881.

[181] Rosenkoetter KE, Kennedy CR, Chirik PJ, Harvey BG. [4 + 4]-cycloaddition of isoprene for the production of high-performance bio-based jet fuel. Green Chem, 2019, 21: 5616-23.

[182] Rude MA, Baron TS, Brubaker S, Alibhai M, Cardayre D, Schirmer SB. Terminal olefin (1-alkene) biosynthesis by a novel p450 fatty acid decarboxylase from Jeotgalicoccus species. Appl Environ Microbiol, 2011, 77: 1718-27.

[183] Ruffing AM. Improved Free Fatty Acid Production in Cyanobacteria with Synechococcus sp. PCC 7002 as Host. Front Bioeng Biotechnol, 2014, 2: 17.

[184] Rui Z, Harris NC, Zhu X, Huang W, Zhang W. Discovery of a Family of Desaturase-Like Enzymes for 1-Alkene Biosynthesis. ACS Catal, 2015, 5: 7091-4.

[185] Santner P, Chanquia SN, Petrovai N, Benfeldt FV, Kara S, Eser BE. Biocatalytic conversion of fatty acids into drop-in biofuels: Towards sustainable energy sources. EFB Bioeconomy J, 2023, 3: 100049.

[186] Sara M, Brar SK, Blais JF. Lipid production by Yarrowia lipolytica grown on biodiesel-derived crude glycerol: optimization of growth parameters and their effects on the fermentation efficiency. RSC Adv, 2016, 6: 90547-58.

[187] Schirmer A, Rude MA, Li X, Popova E, del Cardayre SB. Microbial biosynthesis of alkanes. Science, 2010, 329: 559-62.

[188] Schleyer P, von Gleicher R, Cupas GJ. Adamantane Rearrangements. The Isomerization of Dihydrocedrene to 1-Ethyl-3,5,7-trimethyladamantane. J Org Chem, 1966, 31: 2014-5.

[189] Schripp T, Grein T, Zinsmeister J, Oßwald P, Köhler M, Müller-Langer F, Hauschild S, Marquardt C, Scheuermann S, Zschocke A, Posselt D. Technical application of a ternary alternative jet fuel blend – Chemical characterization and impact on jet engine particle emission. Fuel, 2021, 288: 119606.

[190] Scown CD, Baral NR, Yang M, Vora N, Huntington T. Technoeconomic analysis for biofuels and bioproducts. Curr Opin Biotechnol, 2021, 67: 58-64.

[191] Shen Y, Brown RC, Wen Z. Syngas fermentation by Clostridium carboxidivorans P7 in a horizontal rotating packed bed biofilm reactor with enhanced ethanol production. Appl Energy, 2017, 187: 585-94.

[192] Shesterkina AA, Strekalova AA, Shuvalova EV, Kapustin GI, Tkachenko OP, Kustov LM. Catalytic synthesis of isoprenol from fatty acid ester over bimetallic Cu–Fe catalysts. Mendeleev Commun, 2022, 32: 672-4.

[193] Songdech P, Butkinaree C, Yingchutrakul Y, Promdonkoy P, Runguphan W, Soontorngun N. Increased production of isobutanol from xylose through metabolic engineering of Saccharomyces cerevisiae overexpressing transcription factor Znf1 and exogenous genes. FEMS Yeast Res. 2024;24. https://doi.org/10.1093/femsyr/foae006.

[194] Sorigué D, Légeret B, Cuiné S, Blangy S, Moulin S, Billon E, Richaud P, Brugière S, Couté Y, Nurizzo D, Müller P, Brettel K, Pignol D, Arnoux P, Li-Beisson Y, Peltier G, Beisson F. An algal photoenzyme converts fatty acids to hydrocarbons. Science, 2017, 357: 903-7.

[195] Su H, Jiang J, Lu Q, Zhao Z, Xie T, Zhao H, Wang M. Engineering Corynebacterium crenatum to produce higher alcohols for biofuel using hydrolysates of duckweed (Landoltia punctata) as feedstock. Microb Cell Fact, 2015, 14: 16.

[196] Sun J, Tian K, Wang J, Dong Z, Liu X, Permaul K, Singh S, Prior BA, Wang Z. Improved ethanol productivity from lignocellulosic hydrolysates by Escherichia coli with regulated glucose utilization. Microb Cell Fact, 2018, 17: 66.

[197] Sundstrom E, Yaegashi J, Yan J, Masson F, Papa G, Rodriguez A, Mirsiaghi M, Liang L, He Q, Tanjore D, Pray TR, Singh S, Simmons B, Sun N, Magnuson J, Gladden J. Demonstrating a separation-free process coupling ionic liquid pretreatment, saccharification, and fermentation with Rhodosporidium toruloides to produce advanced biofuels. Green Chem, 2018, 20: 2870-9.

[198] Tian L, Papanek B, Olson DG, Rydzak T, Holwerda EK, Zheng T, Zhou J, Maloney M, Jiang N, Giannone RJ, Hettich RL, Guss AM, Lynd LR. Simultaneous achievement of high ethanol yield and titer in Clostridium thermocellum. Biotechnol Biofuels, 2016, 9: 116.

[199] TotalEnergies. 2014. Total and Amyris renewable jet fuel ready for use in commercial aviation [WWW Document]. URL https://totalenergies.com/media/news/press-releases/total-and-amyris-renewable-jet-fuel-ready-use-commercial-aviation?utm_source=chatgpt.com (accessed 10.5.25).

[200] Treece TR, Pattanayak S, Matson MM, Cepeda MM, Berben LA, Atsumi S. Electrical-biological hybrid system for carbon efficient isobutanol production. Metab Eng, 2023, 80: 142-50.

[201] Uddin MM, Lee U, Xu H, Li Y, Kwon H, Zhang Y, Smolinski S, Cai H, Tao L. Sustainable aviation fuel from ethanol: Techno-economic analysis and life cycle analysis. Appl Energy, 2025, 398: 126373.

[202] Valencia LE, Incha MR, Schmidt M, Pearson AN, Thompson MG, Roberts JB, Mehling M, Yin K, Sun N, Oka A, Shih PM, Blank LM, Gladden J, Keasling JD. Engineering Pseudomonas putida KT2440 for chain length tailored free fatty acid and oleochemical production. Commun Biol, 2022, 5: 1363.

[203] van Dyk S, Saddler J. Progress in Commercialization of Biojet/Sustainable Aviation Fuels (SAF): Technologies, potential and challenges. IEA Bioenergy; 2021.

[204] van Dyk S, Saddler J. Progress in Commercialization of Biojet/Sustainable Aviation Fuels (SAF): Technologies and policies. IEA Bioenergy; 2024.

[205] van Dyk S, Su J, Mcmillan JD, Saddler JJ. Potential synergies of drop-in biofuel production with further co-processing at oil refineries. Biofuels. Bioprod Bioref, 2019.

[206] van Dyk J, Görgens JF, van Rensburg E. Valorisation of Industrial Waste for Second-Generation Ethanol Production: Towards the Scale-Up of High-Solids Paper Sludge Fermentation2025

[207] Varghese S, Demeke MM, Verhé R, Redant E, Cruyssen CV, Thevelein JM. Process optimization for saccharification and fermentation of the Organic Fraction of Municipal Solid Waste (OFMSW) to maximize ethanol production performance. Bioresource Technol Rep, 2023, 24: 101681.

[208] Wagner E, Sierra-Ibarra E, Rojas NL, Martinez A. One-pot bioethanol production from brewery spent grain using the ethanologenic Escherichia coli MS04. Renew Energy, 2022, 189: 717-25.

[209] Walkling CJ, Zhang DD, Harvey BG. Extended fuel properties of sustainable aviation fuel blends derived from linalool and isoprene. Fuel, 2024, 356: 129554.

[210] Walkling CJ, Doppalapudi KR, Zhang DD, Harvey BG. Renewable diesel fuel prepared by the isomerization and hydrogenation of biosynthetic farnesene. Energy Fuels, 2025, 39: 8495-500.

[211] Walls LE, Rios-Solis L. Sustainable production of microbial isoprenoid derived advanced biojet fuels using different generation feedstocks: A review. Front Bioeng Biotechnol, 2020, 8: 599560.

[212] Walls LE, Otoupal P, Ledesma-Amaro R, Velasquez-Orta SB, Gladden JM, Rios-Solis L. Bioconversion of cellulose into bisabolene using Ruminococcus flavefaciens and Rhodosporidium toruloides. Bioresour Technol, 2023, 368: 128216.

[213] Walsh K, McDonnell K, Weigh P, Evans S, Cornelis J. 2024. ArcelorMittal and LanzaTech Announce Ethanol Production Milestone and Shipment of First Barge from Flagship Steelanol Facility in Belgium [WWW Document]. URL https://lanzatech.com/arcelormittal-and-lanzatech-announce-ethanol-production-milestone-and-shipment-of-first-barge-from-flagship-steelanol-facility-in-belgium/?utm_source=chatgpt.com (accessed 9.26.25).

[214] Wang W, Lu X. Microbial Synthesis of Alka(e)nes. Front. Bioeng Biotechnol, 2013, 1: 10.

[215] Wang Z, Lv Z, Du J, Mo C, Yang X, Tian S. Combined process for ethanol fermentation at high-solids loading and biogas digestion from unwashed steam-exploded corn stover. Bioresour Technol, 2014, 166: 282-7.

[216] Wang D, Thakker C, Liu P, Bennett GN, San K-Y. Efficient production of free fatty acids from soybean meal carbohydrates. Biotechnol Bioeng, 2015, 112: 2324-33.

[217] Wang W-C, Tao L, Markham J, Zhang Y, Tan E, Batan L, Warner E, Biddy M. Review of Biojet Fuel Conversion Technologies. National Renewable Energy Laboratory. 2016.

[218] Wang S, Wang Z, Wang Y, Nie Q, Yi X, Ge W, Yang J, Xian M. Production of isoprene, one of the high-density fuel precursors, from peanut hull using the high-efficient lignin-removal pretreatment method. Biotechnol Biofuels, 2017, 10: 297.

[219] Wang G, Haringa C, Noorman H, Chu J, Zhuang Y. Developing a Computational Framework To Advance Bioprocess Scale-Up. Trends Biotechnol, 2020, 38: 846-56.

[220] Wang W, Liu Y, Shi C, Pan L, Zhang X, Zou J-J. High energy density renewable fuels based on multicyclic sesquiterpene: Synthesis and performance. Fuel, 2022, 318: 123665.

[221] Wang X, Zhang X, Zhang J, Xiao L, Zhou Y, Wang F, Li X. Metabolic engineering of Escherichia coli for efficient production of linalool from biodiesel-derived glycerol by targeting cofactors regeneration and reducing acetate accumulation. Ind Crops Prod, 2023, 203: 117152.

[222] Wang B, Ting ZJ, Zhao M. Sustainable aviation fuels: Key opportunities and challenges in lowering carbon emissions for aviation industry. Carbon Capture Sci Technol, 2024, 13: 100263.

[223] Wargacki AJ, Leonard E, Win MN, Regitsky DD, Santos CNS, Kim PB, Cooper SR, Raisner RM, Herman A, Sivitz AB, Lakshmanaswamy A, Kashiyama Y, Baker D, Yoshikuni Y. An engineered microbial platform for direct biofuel production from brown macroalgae. Science, 2012, 335: 308-13.

[224] Weitz S, Hermann M, Linder S, Bengelsdorf FR, Takors R, Dürre P. Isobutanol production by autotrophic acetogenic bacteria. Front Bioeng Biotechnol, 2021, 9: 657253.

[225] Whited GM, Feher FJ, Benko DA, Cervin MA, Chotani GK, McAuliffe JC, LaDuca RJ, Ben-Shoshan EA, Sanford KJ. TECHNOLOGY UPDATE: Development of a gas-phase bioprocess for isoprene-monomer production using metabolic pathway engineering. Ind Biotechnol, 2010, 6: 152-63.

[226] Willrodt C, David C, Cornelissen S, Bühler B, Julsing MK, Schmid A. Engineering the productivity of recombinant Escherichia coli for limonene formation from glycerol in minimal media. Biotechnol J, 2014, 9: 1000-12.

[227] Wong MS, Li M, Black RW, Le TQ, Puthli S, Campbell P, Monticello DJ. Microaerobic conversion of glycerol to ethanol in Escherichia coli. Appl Environ Microbiol, 2014, 80: 3276-82.

[228] Woodroffe J, Harvey BG. High-Performance, Bio-Based, Jet Fuel Blends Containing Hydrogenated Monoterpenes and Synthetic Paraffinic Kerosenes. Energy Fuels, 2020, 34: 5929-37.

[229] Woodroffe J-D, Harvey BG. Chemoselective hydrogenation of ring-strained monoterpenes: A route to high‐performance sustainable aviation fuels. Energy Technol. 2021;9. https://doi.org/10.1002/ente.202100221.

[230] Woodroffe J-D, Harvey BG. Thermal cyclodimerization of isoprene for the production of high-performance sustainable aviation fuel. Energy Adv, 2022, 1: 338-43.

[231] Woodroffe J-D, Harvey BG. A Simple Process for the Dimerization and Cross-Coupling of Isoprene and Myrcene to High-Performance Jet and Diesel Blendstocks. Energy Fuels, 2022, 36: 2630-8.

[232] Woodroffe J-D, Lupton DV, Garrison MD, Nagel EM, Siirila MJ, Harvey BG. Synthesis and fuel properties of high-energy density cyclopropanated monoterpenes. Fuel Process Technol, 2021, 222: 106952.

[233] Wu H, Karanjikar M, San K-Y. Metabolic engineering of Escherichia coli for efficient free fatty acid production from glycerol. Metab Eng, 2014, 25: 82-91.

[234] Wu H, Lee J, Karanjikar M, San K-Y. Simultaneous utilization of glucose and mannose from woody hydrolysate for free fatty acid production by metabolically engineered Escherichia coli. Bioresour Technol, 2015, 185: 431-5.

[235] Wu J, Cheng S, Cao J, Qiao J, Zhao G-R. Systematic Optimization of Limonene Production in Engineered Escherichia coli. J Agric Food Chem, 2019, 67: 7087-97.

[236] Wu T, Liu J, Li M, Zhang G, Liu L, Li X, Men X, Xian M, Zhang H. Improvement of sabinene tolerance of Escherichia coli using adaptive laboratory evolution and omics technologies. Biotechnol Biofuels, 2020, 13: 79.

[237] Xiao Y, Ruan Z, Liu Z, Wu SG, Varman AM, Liu Y, Tang YJ. Engineering Escherichia coli to convert acetic acid to free fatty acids. Biochem Eng J, 2013, 76: 60-9.

[238] Xie L, Yu W, Gao J, Wang H, Zhou YJ. Ogataea polymorpha as a next-generation chassis for industrial biotechnology. Trends Biotechnol, 2024, 42: 1363-78.

[239] Xu J-Z, Chen S-L, Niu S-Q. Systematic modulating carbon metabolism to improve α-farnesene production in Pichia pastoris. Fuel Process Technol, 2023, 247: 107757.

[240] Xue C, Cheng C. Butanol production by Clostridium. Advances in Bioenergy. Elsevier; 2019. pp. 35–77. https://doi.org/10.1016/bs.aibe.2018.12.001.

[241] Yan Z, Zhang J, Bao J. Increasing cellulosic ethanol production by enhancing phenolic tolerance of Zymomonas mobilis in adaptive evolution. Bioresour Technol, 2021, 329: 124926.

[242] Yang H, Yang Z-J, Huang T-X, Pan L, Wei X-M, Hu Y-F, Yuan Y-Q, Wang L-L, Ding J-J. Accurate density prediction of sesquiterpenoid hedfs and the multiproperty computing server sesquiterpre. ACS Omega, 2024, 9: 26213-21.

[243] Yao F, Liu S-C, Wang D-N, Liu Z-J, Hua Q, Wei L-J. Engineering oleaginous yeast Yarrowia lipolytica for enhanced limonene production from xylose and lignocellulosic hydrolysate. FEMS Yeast Res. 2020a;20. https://doi.org/10.1093/femsyr/foaa046.

[244] Yao P, You S, Qi W, Su R, He Z. Investigation of fermentation conditions of biodiesel by-products for high production of β-farnesene by an engineered Escherichia coli. Environ Sci Pollut Res Int, 2020, 27: 22758-69.

[245] You S, Yin Q, Zhang J, Zhang C, Qi W, Gao L, Tao Z, Su R, He Z. Utilization of biodiesel by-product as substrate for high-production of β-farnesene via relatively balanced mevalonate pathway in Escherichia coli. Bioresour Technol, 2017, 243: 228-36.

[246] Yu X, Liu B, Zhang Y. Effect of Si/Al ratio of high-silica HZSM-5 catalysts on the prins condensation of isobutylene and formaldehyde to isoprene. Heliyon, 2019, 5: e01640.

[247] Yunus IS, Anfelt J, Sporre E, Miao R, Hudson EP, Jones PR. Synthetic metabolic pathways for conversion of CO2 into secreted short-to medium-chain hydrocarbons using cyanobacteria. Metab Eng, 2022, 72: 14-23.

[248] Zappi A, Fortela DL, Holmes WE. An Assessment of Methanotrophs Producing Industrial-Grade Lipids for Biofuels and Other Commercial Chemicals. Energies, 2020, 13: 3887.

[249] Zhang J, Webber MS, Pu Y, Li Z, Meng X, Stone ML, Wei B, Wang X, Yuan S, Klein B, Seemala B, Wyman CE, Ramasamy KK, Thorson M, Langholtz MH, Heyne JS, Koishybay A, Adhikari S, Cao S, Sutton AD, Tuskan GA, Román-Leshkov Y, Ragauskas AJ, Ling T, Davison BH. Sustainable aviation fuels from biomass and biowaste via bio- and chemo-catalytic conversion: Catalysis, process challenges, and opportunities. Green Energy Environ, 2025, 10: 1210-34.

[250] Zhang Y, Ma Z, Li W, Liu C, Gao H, Wang M, Li L, Zhang Q, Lv B, Qin L, Li C. Dynamic regulation and enhancement of synthetic network for efficient biosynthesis of monoterpenoid α-pinene in yeast cell factory. Bioresour Technol, 2025, 419: 132064.

[251] Zhao Y, Zhu K, Li J, Zhao Y, Li S, Zhang C, Xiao D, Yu A. High-efficiency production of bisabolene from waste cooking oil by metabolically engineered Yarrowia lipolytica. Microb Biotechnol, 2021, 14: 2497-513.

[252] Zhao B, Zhang Y, Wang Y, Lu Z, Miao L, Wang S, Li Z, Sun X, Han Y, He S, Zhang Z, Xiao D, Zhang C, Foo JL, Wong A, Yu A. Biosynthesis of α-bisabolene from low-cost renewable feedstocks by peroxisome engineering and systems metabolic engineering of the yeast Yarrowia lipolytica. Green Chem, 2023.

[253] Zheng X, Li Y, Che C, Gao Q, Wang X, Ledesma-Amaro R, Luo ZW, Lü X, Wang Y, Yang X. Metabolic Engineering of Rhodotorula toruloides for the Production of Linalool. J Agric Food Chem, 2025, 73: 5395-404.

[254] Zhou YJ, Kerkhoven EJ, Nielsen J. Barriers and opportunities in bio-based production of hydrocarbons. Nat Energy, 2018, 3: 925-35.

[255] Zhou YJ, Hu Y, Zhu Z, Siewers V, Nielsen J. Engineering 1-Alkene Biosynthesis and Secretion by Dynamic Regulation in Yeast. ACS Synth Biol, 2018, 7: 584-90.

[256] Zhu J, Alegre-Requena JV, Cherry P, Curtis D, Harvey BG, Jabed MA, Kim S, McEnally CS, Pfefferle LD, Woodroffe J-D. Sooting tendencies of terpenes and hydrogenated terpenes as sustainable transportation biofuels. Proc Combust Inst, 2022.

Funding

U.S. Department of Energy(DE-AC02-05CH11231)

National Research Foundation of Korea(RS-2024-00346173)

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