Boosting plant oil yields: the role of genetic engineering in industrial applications

* Corresponding author at: E-mail address: [email protected] ‡: These authors contributed equally. Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Biofuel Research Journal 42 (2024) 2105-2145 Review Paper Boosting plant oil yields: the role of genetic engineering in industrial applications Nima Hajinajaf 1,‡, Ahmad Fayyazbakhsh2,‡, Sara Kamal Shahsavar3, Forough Sanjarian4, Hassan Rahnama5,* 1Chemical Engineering Program, School for Engineering of Matter, Transport, and Energy, Arizona State University, Tempe, AZ, USA. 2Department of Environmental Protection Engineering, Faculty of Technology, Tomas Bata University in Zlín, T. G. Masaryka Square 5555, 760 01 Zlín, Czech Republic. 3Department of Microbiology and Virology, School of Medicine, Mashhad University of Medical Sciences, Mashhad, Iran. 4National Institute of Genetic Engineering and Biotechnology, Tehran, Iran. 5Agricultural Biotechnology Research Institute of Iran (ABRII), Agricultural Research Education and Extension Organization (AREEO), Karaj, Iran. HIGHLIGHTS GRAPHICAL ABSTRACT ➢ Plant oil-based products have emerged as ecofriendly alternatives to petroleum counterparts. ➢ Plant oils are versatile, with applications in cooking, lubrication, cosmetics, polymers, and medicine. ➢ Non-edible plant oils present new opportunities for biodiesel and bioproduct production. ➢ Genetic engineering can enhance both the yield and quality of plant oils for bio-based industries. ARTICLE INFO ABSTRACT Article history: Received 25 March 2024 Received in revised form 20 May 2024 Accepted 26 May 2024 Published 1 June 2024 Keywords: Biofuel Genetic engineering Plant oils Triacylglycerol Bio-based economy Sustainability As climate change intensifies and the need to reduce human-caused emissions becomes more urgent, transitioning to a bio-based economy is essential. This paper explores the diverse industrial applications of plant oils as sustainable alternatives to petroleumbased products, including their use in food, polymers, lubricants, surfactants, pesticides, emollients, and biofuels. This review delves into biosynthetic pathways, detailing the key enzymes and processes involved in the synthesis of triacylglycerol. It thoroughly discusses how genetic and metabolic engineering can not only increase oil yields but also modify fatty acid compositions to better meet industrial requirements. By understanding genetics and utilizing advanced biotechnologies, the oil content and quality of plant sources can be significantly enhanced, aligning with both sustainability goals and industrial demands. This paper provides a comprehensive overview of the current uses and genetic engineering of plant oil production, proposing innovative strategies such as utilizing oils from biomass or cultivating non-edible oil crops. These approaches aim to establish a sustainable industrial system, reduce reliance on fossil fuels, and promote the growth of an environmentally responsible biobased economy. Additionally, the review highlights future directions, examining the economic implications and environmental benefits of adopting plant oils across various sectors and positioning them as pivotal to achieving an eco-friendly, bio-based economy. ©2024 Alpha Creation Enterprise CC BY 4.0 Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2106 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Contents 1. Introduction .......................................................................................................................................................................................................................... 2106 2. Industrial applications of plant oil and the challenges faced ................................................................................................................................................. 2107 2.1. Polymers ...................................................................................................................................................................................................................... 2109 2.2. Surfactants ................................................................................................................................................................................................................... 2110 2.3. Lubricants .................................................................................................................................................................................................................... 2115 2.4. Metal working fluid ..................................................................................................................................................................................................... 2115 2.5. Stabilizers and plasticizers ........................................................................................................................................................................................... 2115 2.6. Composites................................................................................................................................................................................................................... 2116 2.7. Pesticides ..................................................................................................................................................................................................................... 2118 2.8. Paint, coating and adhesives ......................................................................................................................................................................................... 2118 2.9. Inks .............................................................................................................................................................................................................................. 2118 2.10. Emollients .................................................................................................................................................................................................................. 2118 2.11. Wax esters .................................................................................................................................................................................................................. 2118 2.12. Textile finishing ......................................................................................................................................................................................................... 2122 3. Plant oil production pathways .............................................................................................................................................................................................. 2122 3.1. Fatty acids synthesis ..................................................................................................................................................................................................... 2123 3.2. Triacylglycerol biosynthesis......................................................................................................................................................................................... 2124 3.3. Triacylglycerol storage ................................................................................................................................................................................................. 2124 4. Genetic engineering for improved oil production ................................................................................................................................................................. 2124 4.1. Metabolic engineering to enhance oil production ......................................................................................................................................................... 2124 4.1.1. Increasing the content of oil per seed ................................................................................................................................................................... 2126 4.1.1.1. FA synthesis (Carbon flux redirection) ........................................................................................................................................................ 2127 4.1.1.2. Glycerol backbone ....................................................................................................................................................................................... 2127 4.1.1.3. TAG biosynthesis ........................................................................................................................................................................................ 2127 4.1.1.4. Lipid transfer proteins ................................................................................................................................................................................. 2127 4.1.1.5. Transcription factors (TFs) .......................................................................................................................................................................... 2127 4.2. Increasing seed oil content by enhancing seed size ...................................................................................................................................................... 2128 4.3. Genetic Engineering for new oil resources: biomass-derived oil .................................................................................................................................. 2128 4.4. Modifying the composition of vegetable oils for industrial applications ...................................................................................................................... 2130 4.4.1. Monounsaturated fatty acids ................................................................................................................................................................................ 2131 4.4.2. Medium-chain saturates ....................................................................................................................................................................................... 2132 4.4.3. Engineering wax ester synthesis .......................................................................................................................................................................... 2133 5. Policy and practical implications .......................................................................................................................................................................................... 2133 6. Challenges and prospects ...................................................................................................................................................................................................... 2133 7. Conclusions .......................................................................................................................................................................................................................... 2133 References ................................................................................................................................................................................................................................ 2134 1. Introduction Climate change and its adverse impacts on various aspects of human health were recently projected in the latest global report released by the Lancet Countdown: Tracking Progress on Health and Climate Change (Watts et al., 2021). Among the reported findings was an increased rate of exposure to heatwaves globally between 2000 and 2016, affecting an additional 125 million medically vulnerable adults. The widespread use of petroleum-derived products has been linked to rising atmospheric CO2 levels, which are associated with the frequency of heatwaves (Verma et al., 2019; Rej et al., 2022). These findings highlight the need to decrease anthropogenic greenhouse gas (GHG) emissions through strategies such as transitioning towards a bio-based economy (Bergfreund et al., 2021; Hajinajaf et al., 2022c; Hajinajaf et al., 2024). Replacing petroleum-derived products with eco-friendly alternatives is a key feature of this transition (De Vrieze et al., 2020). There are growing concerns about the future availability of petroleumderived products (Siracusa and Blanco, 2020). Utilizing renewable raw materials for daily life products seems crucial for sustainable development (Bergfreund et al., 2021; Hajinajaf et al., 2022b). Figure 1a shows the share of different sources in the global primary energy supply. Using renewable materials can reduce CO2 emissions and offer additional advantages Abbreviations ABA Abscisic acid G3PDH Glycerol-3-phosphate dehydrogenase ACCase Acetyl-CoA carboxylase GAPDH Glyceraldehyde 3-phosphate dehydrogenase AP2 Apetala2 GPAT Glycerol-3-phosphate acyltransferase ARF2 Auxin Response Factor 2 LPAAT Lysophosphatidic acid acyltransferase CAGR Compound annual growth rate MWFs Metalworking fluids CPT Choline phosphotransferase PC Phosphatidylcholine CRISPR Clustered regularly interspaced short palindromic repeats PDAT Phospholipid: diacylglycerol acyltransferase DGAT Diacylglycerol acyltransferase PVC Polyvinyl chloride ER Endoplasmic reticulum PXA1 Peroxisomal ABC transporter 1 ESBO Epoxidized soybean oil SDP1 SUGAR-DEPENDENT1 FA Fatty Acids TAG Triacylglycerol FAD2 Fatty acid desaturase 2 TFs Multiple transcription factors FAR Fatty Acid Reductases TP Triose-phosphates G3P Glycerol-3-phosphate TTG2 Transparent Testa Glabra 2 Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2107 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Fig. 1. a) Share (%) of different sources in global primary energy supply (Kan et al., 2019); b) Shares of world oil consumption in various sectors and industries in the year 2018 (IEA., 2018). The “Other” category encompasses agriculture, commercial and public services, non-specified other, pipeline, and non-specified transport; c) Area equipped for irrigation (OECD, 2012); d) Total arable land in use (FAO, 2011). associated with green chemistry, such as biodegradability and lower toxicity (Fallahi et al., 2021; Hajinajaf et al., 2021). One of the most important needs of various industries, including the food industry, is industrial oils, ranging from lubricating to hydraulic and cutting oils. Currently, a significant portion of industrial oil resources used even in the food industry are derived from petroleum, leading to substantial oleochemical pollution (Metzger and Hüttermann, 2009; Hayes, 2021; Fayyazbakhsh et al., 2022). Figure 1b shows the distribution of global oil consumption across different sectors and industries in 2018 (IEA., 2018). Plant oils are considered ideal environmentally friendly, renewable, and sustainable feedstocks that could potentially replace petroleum-derived oil in the mentioned industries (El-Dalatony et al., 2022; Yaashikaa et al., 2022). With a market cap comparable to that of fossil-based fuels (Hoang et al., 2021), industrial oil is a key sector for transitioning from petroleumbased to plant-based oils. However, since over 85% of the plant oils produced globally are used for human nutrition, it is unrealistic to fully replace fossil oils with plant oils in industry and transportation (Hajjari et al., 2017). It has been highlighted that replacing just 40% of the fossil oils in these industries would require global plant oil production to triple by 2030 (Carlsson et al., 2011). The increasing scarcity of renewable water resources (as highlighted in Figure 1c and detailed in Table 1), along with the diminishing availability of arable land (as shown in Fig. 1d), adds significant complexity to this endeavor. This challenge is particularly pronounced when we depend on existing oil crops with stabilized yields and oil content. Table 2 shows the yield and oil content of various plant oil feedstocks. In addition to yield and oil content, challenges associated with existing plant oils extend to unfavorable fatty acid profiles in certain feedstocks, leading to bio-oil properties unsuitable for specific industrial applications. Moreover, the presence of toxic or allergenic compounds further complicates the utilization of these oils (Baskar et al., 2019; Nomanbhay et al., 2018). Therefore, it is critical to develop innovative and promising industrial plant oil platforms to tackle these challenges. However, it is equally crucial to ensure that these platforms do not intensify competition with food crops, thereby safeguarding global food security. This importance is highlighted by global plant oil production statistics, which forecast an increase from 149 million tonnes in 2005 to 282 million tonnes in 2050, aimed at meeting the demands of a growing world population (Alexandratos and Bruinsma, 2012). Table 3 provides an overview of global plant oil production, current demands, and projected demands beyond 2050. Besides increasing plant oil production yield per hectare, one of the main approaches to meeting industrial demand for plant oils is developing new oil crops that can utilize marginal or non-agricultural lands and waters. In alignment with these goals, this review aims to comprehensively explore the diverse applications of plant oils across various industries while addressing the associated challenges. It also delves into plant metabolic pathways for oil production and the use of genetic engineering to enhance oil quantity and quality. Additionally, the review critically examines innovative alternative strategies, such as biomass-derived oils. Table 4 summarizes the various aspects of plant oils, their applications, and related research covered in this review, comparing them to discussions in review articles published from 2017 to 2023. 2. Industrial applications of plant oil and the challenges faced Major oil crops on which global oil production depends include palm, soybean, rapeseed (also known as canola), corn, sunflower, cottonseed, olive, and peanut. Minor oil feedstocks include safflower, coconut, sesame, and linseed (Table 2) (Alexandratos and Bruinsma, 2012; Wan et al., 2017b). While the majority of global oil production over the last decade has been directed toward food/feed applications, approximately one-fifth has been used for industrial and bioenergy applications, and this proportion is expected to increase (Scarlat et al., 2015). In other words, the ratio between food, feed, and industrial/bioenergy applications of global oil production generally stands at 80:6:14 (Quispe et al., 2013). However, with growing biodiesel production, this ratio has shifted to 74:6:20 (Biermann et al., 2011). These proportions are still anticipated to change further in favor of non-food applications, including bioenergy and industrial plant-based oil production (Rathour et al., 2023). Therefore, the main challenge is increasing the quantity of global plant oil production to meet these growing Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2108 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 1. Annual renewable water resources and irrigation water withdrawal. Precipitation (mm/annum) Renewable Water Resources (km3) Water Use Efficiency Ratio (%) Irrigation Water Withdrawal (km3) Pressure on Water Resources due to Irrigation (%) * 2005/2007 2050 2005/2007 2050 2005/2007 2050 World 800 4200 44 46 2620 2906 6 7 Developed Countries 540 14000 42 43 505 493 4 4 Developing Countries 990 28000 44 47 2115 2413 8 9 *Water withdrawal for irrigation as a percentage of total annual renewable water resources. Sources: Bruinsma (2009); Nachtergaele et al. (2023); http://www.fao.org/nr/solaw/thematic-reports/en/. Table 2. Different plant oil feedstocks, their production parameters, and commercial applications. Source: Wan et al. (2017b). Commercial Uses (non-food uses in Bold) Global Oil Production (million tonnes) in 2016 Global Seed Production (million tonnes) in 2016 Global Cultivation Area (million ha) in 2016 L oil/ha kg oil/ha Oil Content (wt%) Oil Crop Food, biodiesel, inks, plasticizers, crayons, paints, and soy candles 54.47 345.97 120.41 446 375 15-20 Soybean (Glycine max) Food, biodiesel 27.31 68.52 34.05 1190 1000 38-46 Rapeseed (Brassica napus L.) Food, coatings 17.11 45.36 23.36 952 800 25-35 Sunflower (Helianthus annuus) Food, lubricants, inks 2.854 2.713 2.700 9.7 1212 1019 10-30 Olive (Olea europaea) Medicine, biodiesel, cosmetics, massage oils and soaps, confectionery and bakery industries 5.78 42.28 24.77 1059 890 45-55 Peanut (Arachis hypogaea) Food and feed, margarine, salad dressings, medicine 5.09 38.88 29.58 325 273 18-25 Cotton seed (Gossypium hirsutum) Food, margarine, shortening, cooking oil, confectionary, soaps, sauces, fat substitutes, biodiesel 7.58 kernel 63.86 palm Palm kernel 17.09 - 5950 5000 30-60 Palm (Arecaceae) Food, margarine, painting, skin moisturizer, and creams for softening and smoothing the skin - 0.8 <1 779 655 20-45 Safflower (Carthamus tinctorius L.) Food, biodiesel, carrier for drug molecules in pharmaceutical preparations, soap, salve, inks and paint, and textile industries 0.523 1608.62 35 172 145 3.3-15.9 Corn (Zea mays) Food, wholesome tonic, medicine, hair treatment, body massage, worship 1.926 4.4 8 696 585 52-63 Sesame (Sesamum indicum) Food, medicine, healtcare, fuel 3.44 61.44 - 2689 2260 63-65 Coconut (Cocos nucifera) Food, medicine, soap production, beverages, pharmaceuticals - 8.6 - 572 481 25-30 Mustard (Brassica alba) Food, margarine, medicinal 1.8 63 - 828 696 12.1-25 16-32 Rice bran (Oryza sativa L.) Enamels, varnishes, resins, coatings - - - 940 790 14-22 Tung fruit (Aleurites fordii) - - 8.9 - 2638 2217 11.2318.8 Avocado (Persea americana) - 0.015 - - 1892 1590 35-40 Jatropha (Jatropha curcas L.) - 0.055 - - - - 27-39 Karanja (Pongamia pinnata) Plasticizers, lubricants, medicine, adhesives, cosmetics, hair oils, food containers, fuel additives, insulation, nylon, synthetic resins, fibers, paints, varnishes, plastics, inks, textiles, drying oils, fungus-growth-inhibiting compounds, embalming fluid, soaps, dyeing aids, cleaning products, detergents, personal care products, styling gel, and adhesive remover 0.73 1.8 - 1413 1188 53 37-60 Castor (Ricinus communis) - - - - 583 490 30-40 Camelina (Camelina sativa) - - - - 1818 1528 44-59 Jojoba (Simmondsia chinensis) Medicine, feeds, sealants, caulking compounds, linoleum, earthen floors, adobe, textiles, fixative, rust inhibitor, lubricant, leather treatment, polishes, varnishes, oil paints, composition ornament for molded decoration, animal care products, wood preservation, industrial lubricant 0.8 3.9 - 478 402 38 -44 Linseed (Linum usitatissimum) Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2109 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 2. continued. Commercial Uses (non-food uses in Bold) Global Oil Production (million tonnes) in 2016 Global Seed Production (million tonnes) in 2016 Global Cultivation Area (million ha) in 2016 L oil/ha kg oil/ha Oil Content (wt%) Oil Crop - - 5.8 - 1026 863 40-45 Coca (Cacao) Food, chocolates, bakery, confectionery, mixed nuts, medicine - - - 482 405 50-55 Hazelnut Table 3. Global plant oil production; current demands and envisioned demands beyond 2050 (Alexandratos and Bruinsma, 2012). Key Variables 2005/2007 2050 2080 2100 Population (million) UN 2010 Revision 6584 9306 9969 10125 Oil crops (oil equivalent), food (kg/capita) 12.1 16.2 16.9 - Oil crops (oil equivalent), all uses (kg/capita) 21.9 30.5 33.8 - Arable land area 1592 1661 1630 - Oil crop production (million tonnes) 149 282 367 - Oil crops (for biofuel) (million tonne) 7 29 - - Oil crops (for biofuel) (% total uses) 4.8 10.3 - - industrial and bioenergy demands (Haregu et al., 2023). The various industrial uses of plant oils and their derivatives, such as oleochemicals, fatty acids (FA), fatty alcohols, and glycerin across different industries, are presented in Figure 2. Moreover, industrial applications and the market size of plant oil are discussed in the subsequent sections (Fig. 3). 2.1. Polymers Over the last few decades, commercially available polymers have been derived from non-renewable fossil resources. The global annual consumption of polymers is approximately 300 million tonnes, with an annual growth rate of 5% (Halden, 2010). This significant quantity, along with the resultant waste streams often disposed of into various ecosystems Table 4. Comparative analysis of plant oil applications and research topics in recently published review articles. Reference Applications Genetic Engineering Global Market Oil-seed Production Policy and Practical Implications Industrial Applications Energy Biosynthetic Pathway Oil Content Oil Composition Oil in Biomass Afonso et al. (2023) √ ˟ ˟ ˟ ˟ ˟ ˟ ˟ ˟ Xu et al. (2018) ˟ √ √ ˟ ˟ √ √ ˟ ˟ Sagun et al. (2023) ˟ ˟ √ √ √ √ ˟ √ ˟ Zhou et al. (2023) ˟ ˟ √ √ √ ˟ ˟ ˟ ˟ Wan et al. (2017b) ˟ √ √ √ √ √ ˟ √ ˟ Rauf et al. (2023) ˟ ˟ √ √ √ √ ˟ √ ˟ Qi et al. (2020) √ ˟ √ √ √ √ ˟ ˟ ˟ Salehi Jouzani et al. (2018) ˟ √ √ √ √ ˟ ˟ ˟ ˟ Msanne et al. (2020) ˟ ˟ √ √ √ ˟ ˟ √ ˟ Present Review √ √ √ √ √ √ √ √ √ without treatment in many parts of the world, has led to growing environmental and health concerns, particularly regarding the use of petrochemical-based polymers. Conversely, this situation has sparked a surge in interest in biobased polymers (Desroches et al., 2012). These polymers are not only renewable but also biodegradable and eco-friendly (Adekunle and Okolie, 2015; Hajinajaf et al., 2022a). Biobased polymers can be synthesized from polysaccharides, fibers, polylactic acid, and other materials, with triacylglycerol (TAG) oils and FA also serving as reliable starting materials during the production process (Acquavia et al., 2021; Zubair et al., 2021). In 2019, the total production volume of biobased polymers reached 3.8 million tonnes (https://www.bioplasticsmagazine.com/en/). Plant oil-based polymers can undergo fabrication using various copolymerization techniques, such as cationic, free radical, and thermal methods, which involve combining plant oils with a variety of petroleumbased co-monomers (Gogoi et al., 2022; Zhu et al., 2023). However, it is important to note that most plant oils require modifications at their naturally occurring reactive sites, such as ester groups and carbon-carbon double bonds, before they can be utilized in biopolymer production (Ike et al., 2021; Ruiz-Rico and Barat, 2021). Therefore, these reactive sites, including the carbon-carbon double bonds found in fatty acid chains, play a crucial role during polymerization (Biermann et al., 2021; Rajput et al., 2023). In simpler terms, these double bonds act as excellent starting points for biopolymer production. For instance, converting double bonds in FA into hydroxyl groups, followed by their reaction with isocyanate, could result in the formation of polyurethanes (Dyer et al., 2008). Additionally, other polymerization techniques, such as acyclic metathesis polymerization (Piccini et al., 2021; Quirino et al., 2021) and ring-opening metathesis polymerization (ROMP) (Ganewatta et al., 2021; Yarolimek et al., 2021), have been employed to synthesize plant oil-based polymers (Garrison et al., 2016). Neves et al. (2018) also demonstrated that modified vegetable oils containing acrylic double bonds exhibit high reactivity and form thermosetting biopolymers through free radical polymerization. Table 5 provides examples of commercially available plant oil-based polymers and their real-world applications. Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2110 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Fig. 2. Global market break-up of (a) oleochemical, (b) fatty acids, (c) fatty alcohols, and (d) glycerin by different industries. Fig. 3. Industrial applications of plant oils and their market size (Zhou et al., 2020; https://www.marketsandmarkets.com; and https://www.statista.com). 2.2. Surfactants Surfactants are amphipathic compounds that influence the surface or interfacial energy of materials (Bergfreund et al., 2021). They constitute a widely used class of chemicals predominantly sourced from petroleum (Nagtode et al., 2023). Typically, surfactants comprise a hydrophilic group linked to a hydrophobic moiety (Polarz et al., 2018; Lamch et al., 2020). The hydrophilic groups vary depending on whether the surfactants are anionic or cationic. Anionic surfactants encompass carboxylate, sulfate, sulfonate, or phosphate groups, while cationic surfactants comprise amine or ammonium groups (Rocky et al., 2023). Surfactants find application in both edible and non-edible products, ranging from soaps and detergents to food emulsifiers and cosmetics (Bergfreund et al., 2021; De Luca et al., 2021; Mohammed and Ikiensikimama, 2023). As illustrated in Table 6, surfactants derived from petrochemicals have raised significant health and environmental concerns due to their toxicological properties. Consequently, safer alternatives, known as biosurfactants, have garnered considerable attention, particularly in the food and cosmetics industries. Indeed, eco-friendly alternatives, like biosurfactants, offer comparable physicochemical properties (such as emulsification, de-emulsification, foaming, and wetting) to their petroleum-derived counterparts (AhmadiAshtiani et al., 2020; Sarubbo et al., 2022). They also possess several advantages, including lower toxicity, biodegradability, and increased resistance across a broader range of pH, salinity, and temperature conditions (Abbot et al., 2022; Sarubbo et al., 2022). Oilseed crops are considered promising feedstocks for biosurfactant production. Specifically, the FA present in plant oils or their corresponding methyl esters can be reduced to produce fatty alcohols, which are then utilized in biosurfactant formulations (Van Renterghem et al., 2018). Despite their favorable environmental attributes, biosurfactants still present less favorable economic characteristics compared to surfactants derived from petroleum (Gaur et al., 2022; Joshi et al., 2022). This issue is Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2111 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 5. Examples of commercially available plant oil-based polymers (Desroches et al., 2012; McKeon et al., 2016; Zhang et al., 2017). Plant oil Sources Products Trade name Company Applications Soybean oil Polymerized Soybean Oil (polySOY) - - - Acrylated Epoxidized Soybean Oil (AESO) Ebecryl 860 UCB Chemicals Company, Advent International (Boston, MA, USA) Surface coatings Maleic acid reacted AESO (MAESO) also “maleated acrylated epoxidized soy oil (MAESO)” - - Sheet molding compound Soybean oil monoglyceride (SOMG), also “maleinated soybean oil monoglyceride” - - - Thermosets prepared by the cationic copolymerization of soybean Oil - - - Polymerized epoxidized soybean oil (ESO) PlastiSoy™ CHS (USA), Makwell (India), The Chemical Company (USA), MultiPlus (Thailand), PolyMar Enterprises (USA), FMC (USA) Rubbers, Resins, Coatings, Paints, Plasticizers, Adhesives, Polyols (polyurethanes), Thermosets Epoxidized Soybean Oil Vikoflex® Arkema (USA) - Norbornenyl-functionalized fatty alcohols derived from soybean oil (NMSA) - - - Polyurethanes BiOH® Cargill (USA) - Agrol® BioBased Technologies (USA) Lubricants, Building Products, Printing Inks, Diesel Additives, Coatings, Furniture, Adhesives, Automotive, Agricultural Products Renuva® Dow Chemical (USA) Adhesive, Conventional Flexible Polyurethane Foam (FPF), Viscoelastic Foam, High Resilience Foam (HR), Molded Foam Sovermol® polyols BASF (Cogins) Oleochemicals (Malesia) Adhesives, Binders, Floor coatings, Castings, Electroplating Linseed oil Linseed Oil Monoglyceride (LOMG) - - - Polymerized Linseed Oil (Linoleum) Linoville, Marmoleum®, Forbo's Topshield™, NATURCote™, Forbo (Switzerland), Armstrong (USA), Torlys (Newzeland) Floor covering Polyurethane (Boiled Linseed Oil (BLO)) Crown® Boiled Linseed Oil W. M. Barr, USA; Crown (USA) Wood finisher Epoxidized linseed oil HiBond® Polar Industries (Canada) Paints, plasticizers, adhesives, coatings, or any application for an epoxidized oil Vikoflex Arkema (USA) - Cyclopentadiene polymers DilulineTM Cargill (USA) Dring oil ML189 Archer Daniel Midland (USA) Varnish, Enamel, Aluminum paint, Reinforced oil Castor oil Maleated alcoholized castor oils (MACOs) - - - 2-(Acryloyloxy) Ethyl Oleate (AEO) HeloxyTM Flexibilizer Momentive Specialty Chemicals (USA) Epoxy Resin, Coating, Construction, Composites, Adhesives, Electrical castings, Electrical laminates and Fibers Estolides Heloxy™ Modifiers48 Hexion (USA) - ERISYSTMGE-35H Emerald Performance Materials (USA) Concrete Patching Compounds, Floor Coatings, Adhesives, Bridge Decking Compounds, Joint Sealants Vorite®Polymerized Castor Oil Vertellus (USA) Elastomers, Adhesives, Coatings, Inks, Polyols, Sealants Zenigloss ® Zenitech (Canada) Emollient, Lip gloss agent, Personal care applications Polyglycerol polyricinoleate (PGPR) Fraken Biochemical (China), Spell Organics (India) Food emulsifier/texture and viscosity control (Chocolate) Polyamides Rilsan®PA11, Rilsan®Fine Powder, Arkema (USA, France) Electrical cable, fuel line, fluid transfer, quick connectors, fasteners and clips, friction parts, pneumatic and hydraulic hose Polyurethane Ultramid® BALANCE BASF (USA) Automotive Agrol Star™ BioBased Technologies (USA) Inks and Coatings Lupranol® Balance 50 BASF (Germany) Foam - Jayant Agro Organics Limited (India) - Polymer Pebax Rnew® Arkema (USA, France) Sports, Medical, Packaging and Industrial applications EcoPaXX™ DSM Automotive and Electrical markets Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2112 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 6. Toxicity of different surfactants against various organisms (Ivancovic and Hrenovic, 2010; Cowan-Ellsberry et al., 2014; Yuan et al., 2014; Theis et al., 2016; Badmus et al., 2021). Surfactant Group Petrochemical surfactants Toxicological features of petrochemical surfactants Plant oil-based alternatives Application Anionic Linear alkylbenzene sulfonates (LAS) Bacteria: Vibrio fischeri (EC50-Luminescence 30 min: 2.6 mg/l) Pseudomonase putida (EC50-Growth inhibition 16 h: 33.4 mg/l) Algae: Dunaliella sp (EC50-24 h: 3.5 mg/l) Crustaceans: Ceriodaphnia dubia (EC50Immobilization 48 h: 5.96 mg/l) Daphnia magna LC50– 48 h, 13.9 mg/ LC50– 48 h, 8.1 mg/l LC50– 48 h, 1.22 mg/l Fish: Carassius auratus (EC50Immobilization 48 h: 5.1 mg/l) Salmo gairdneri (Immobilization EC50– 48 h, 33.61 mg/l) Gammbusia affinis (mosquito fish) Immobilization EC50-48 h, 40.15 mg/l Carassius auratus (goldfish) Immobilization EC50-48 h, 38.04 mg/l Plant: Bush beans, radish and grasses: Yield and growth NOEC–76 days, 27 mg/kg Potato: Yield and growth NOEC–106 days, 16 mg/kg Brand Name: Eurasol (Ammonium oleate, Potassium tallate, Mixed fatty acid salts, Potassium cocoate, Potassium palmate) EOC Co. Belgium Brand name: SERVO® BRILLANT (castor oil sulphonate) Elementis Co. The Netherlands As detergents, foaming agents, emulsifiers, antistatic agents, dispersants, stabilizers in the family and chemical aspects of life, Cosmetic, Pharmaceutical and Petrochemical products Linear ether sulfate - Octylphenol polyoxyethylene sodium - Soaps Alga: EC50: 10-50 mg/L Sodium dodecyl sulphate (SDS) Bacteria: Vibrio fischeri (EC50Luminescence 15 min: 2.6 mg/L) Algae: Raphidocelis subcapitata (IC50 - Cell density 72 h: 36.58 mg/L) Crustaceans: Artemia salina (LC50 - Larvae mortality 24 h: 41.04 mg/L) Gastropod: Physa acuta (LC50 - Mortality 24 h: 27.2 mg/L) Sea urchin: Paracentrotus lividus (EC50Fertilization rate: 3.2 mg/L) Fish: Gammbusia affinis (EC50Immobilization 48 h: 13.64 mg/L) Salmo gairdneri (rainbow trout) Immobilization (EC50– 48 h, 10.84 mg/L) Carassius auratus (goldfish) Immobilization (EC50– 48 h, 12.35 mg/L) Alkyl sulphate (AS) - Sodium lauryl sulphate (SLS) - Alkyl ethoxysulphate (AES) Algae: Pseudokirchneriella subcapitata (EC50 - Cell density 72 h 3.5 mg/L) Raphidocelis subcapitata (IC50 - Cell density 72 h 2.18 mg/L) Crustaceans: Artemia franciscana (LC50Nauplii mortality 72 h 23.92 mg/L) Fish: Salmo gairdneri (EC50Immobilization 48 h 10.84 mg/L) Secondary alkane sulphonates (SAS) - Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2113 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 6. continued. Surfactant Group Petrochemical surfactants Toxicological features of petrochemical surfactants Plant oil-based alternatives Application Non-ionic Polyoxyethylenestearyl ether(20EO) Water flea: EC50: 48 mg/L Brand Name: Eur Amid (cocamide diethanolamine, monoethanolamine); Euroxide (Cocamine oxide, Cocamidopropylamine Oxide, EOC Co. Belgium Brand Name: SERDOLAMIDE (coconut oil diethanol amide (Superamide), oleic acid diethanol amide (Kritchevsky), soya oil diethanol amide in TEA/water (Super-amide)), SERDOX® (oleic acid monoethanol amide), ECOSURF™ SA Surfactants (Dow co, USA), EcoSense™ Surfactants (DOW company, USA) Textile, paper, food, plastic, glass, fiber, medicines, pesticides, dyes, other industries, emulsifiers, wetting agents, and foam stabilisation agents, in various biotechnological processes, and to facilitate solubilisation and increase drug carrier stability Polyoxyethylenestearyl ether(10EO) Australian native frogs: Full narcosis EC50– 48 h, 2.8– 3.8 mg/L Fish: Fathead minnow: LC50– 96 h, 4.6 mg/L Lauryl alcohol ethoxylates(7EO) - Nonylphenolethoxylates (9EO~11EO) (NPE) Crustaceans: Daphnia magna LC50– 48 h, 14 mg/L Fish: Pimphales promelas (fathead minnow): LC50– 10 d, 2.7 mg/L Fathead minnow LC50– 96 h, 4.6 mg/L Effects on reproductive health of fish: NP induce the production of vitellogenin in male fish, a protein usually only found in sexually mature females under the influence of estrogens Alkylphenol ethoxylate (APE) - Octyl phenol ethoxyales (OPE) Effects on reproductive health of fish:OP induce the production of vitellogenin in male fish, a protein usually only found in sexually mature females under the influence of estrogens Alcohol ethoxylate (AE) Bacteria: Microcystis aeruginosa (Estimated EC10 - Cell density 0.154 mg/L) Algae: Lemna minor (Estimated EC10 - Frond count 0.101 mg/L) Navicula pelliculosa (Estimated EC10 - Cell density 0.140 mg/L) Crustaceans: Ceriodaphnia dubia (EC50 - Immobilization 48 h 0.39 mg/L) Fish: Pimephales promelas (NOEC - Survival 4.35 mg/L) Fatty acid ethoxylate (FAE) - Cationic Cetyltrimethyl ammonium chloride - - Sterilization, rust, corrosion, breaking, corrosion and mineral flotation, detergents, fabric softeners, and hair conditioners Quaternary ammonium compound (QAC) QACs are toxic to mammalian cells and are not recommended for systemic application, damaging effects of cationic surfactants on human lymphocytes Bacteria: Vibrio fischeri (EC50Luminescence 30 min 0.5 mg/L) Pseudomonas putida (EC50 - Growth inhibition 16 h 6.9 mg/L) Algae: Dunaliella sp. (EC50 - 24 h 0.79 mg/L) Crustaceans: Daphnia magna (EC50Immobilization 24 h 0.38 mg/L) Fish: Salmo gairdneri (EC50Immobilization 48 h 1.21 mg/L) Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2120 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 10. Research studies on using plant-based pesticides on various organisms. Reference Plant Oil Oil concentration Application Targeted Species Time (h) Mortality /Inhibitory (%) Results Kavetsou et al. (2019) Encapsulated Mentha pulegium EO 0.10% Used in yeast cell microcarriers Insect pest Myzus persicae 24 5 • Encapsulated-oil showed higher insecticidal activity than non-encapsulated one. 48 5 Non-encapsulated M. pulegium EO 0.10% 24 0 48 5 VasanthaSrinivasan et al. (2018) Betel leaf oil 500 mg/kg In soil against earthworm and redworm Eudrilus eugeniae (Kinberg) 168 ≈1.5 • Innovative and safe insecticides for soildwelling creatures. • The enzyme level was not affected by the essential oil of P. betle. Thus, the plantderived volatile oil did not affect biochemical reactions 336 ≈1.2 1000 mg/kg 168 ≈3.5 336 ≈2.5 500 mg/kg Eisenia fetida (Savigny) 168 ≈1.9 336 ≈1.5 1000 mg/ kg 168 ≈3.2 336 ≈2.5 Tabari et al. (2017) α-thujone-rich Artemisia sieberi essential oil 2 μg/cm3 Poultry industry Dermanyssus gallinae (Dermanyssida) 24 3.22 • Prolonged toxicity of the oil. • On adults of the poultry red mite, D. gallinae, -thujone-rich A. sieberi essential oil showed promising toxicity and repellant action. 5 μg/cm3 10.88 10 μg/cm3 25.77 Adak et al. (2020) Eucalyptus 1 μg/cm3 Rice Sitophilus oryzae 24 55 • Downsized essential oils have higher insecticide potential than normal size. • Nanoemulsion eucalyptol can improve efficiency and reduce the cost of essential oils. 2 μg/cm3 100 3 μg/cm3 Tribolium castaneum 17 7 μg/cm3 58 Klein et al. (2020) Thyme 0.148% (v/v) Laboratory and greenhouse bioassays Deroceras reticulatum 24 50 • By 1% (v/v), after 24 hours, the mortality by all types of essential oils reached 100%, showing that these plant extracts might be usable as new rational molluscicides. • Thyme showed the highest performance as a pesticide than other oils. • Although thyme proved beneficial against the targeted species, the open environment could influence the results. 0.26% (v/v) 99 Garlic 0.204% (v/v) 50 0.329% (v/v) 99 Rosemary 0.307% (v/v) 50 0.554% (v/v) 99 Lemongrass 0.32% (v/v) 50 0.72% (v/v) 99 Cinnamon Cassia 0.42% (v/v) 50 0.799% (v/v) 99 Idoko and Ileke (2020) Aframomum melegueta 1 mL/L Cowpea seeds Callosobruchus maculatus 12 13.33 • Protect stored cowpeas against C. maculatus. • The influences of essential oils on C. maculatus mortality increased with the exposure time and treatment rates. • The significant mortality induced by essential oils may be owing to the essential oils’ suffocating odor, which must have disturbed the insects’ regular respiratory mechanism. 5 mL/L 26.67 Annona muricata 1 mL/L 13.33 5 mL/L 16.67 Eucalyptus globules 1 mL/L 13.33 5 mL/L 20 Ficus exasperate 1 mL/L 16.67 5 mL/L 20 Tetrapleura tetraptera 1 mL/L 6.67 5 mL/L 20 Papadimitriou et al. (2019) Pulegone extracted from M. pulegium 500 µL/L Cucumber Tomato Tetranychus urticae females 72 47 • Incorporating these two EOs did not influence the mortality rate of helpful insects (Nesidiocoris tenuis). • No phytotoxic influences were recorded after using EOs on the plants. 1000 µL/L 51 Piperitone extrcted from M. pulegium 500 µL/L 65 1000 µL/L 71 Janaki et al. (2018) Cyperus rotundus 0.4 μl/cm2 • Cowpea • Chickpea • Dates C. maculatus 24 64 • The used EO exhibited a repellent effect on all three kinds of insects examined in the study. • The repellent effect in O. surinamensis and T. granarium was more than in C. maculatus. 1 μl/cm2 88 0.4 μl/cm2 Trogoderma granarium 66 1 μl/cm2 77 0.4 μl/cm2 Oryzaephilus surinamensis 44 1 μl/cm2 72 Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2121 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 11. Use of vegetable oils in coating materials and their important properties. Reference Plant oil Oil content (%) or concentration Purpose Time (d) (factor) Important property Influence on the property Important information Dong and Wang (2017) Garlic essential oil 1 The influence of essential oils coating for improving the quality of strawberries that contain carboxymethyl cellulose 3 (storage time) Decay percentage (%) ≈-13.5% • Using carboxymethyl cellulose in conjunction with garlic essential oil improved decay percentage, weight loss, total soluble solids, titratable acidity, and ascorbic acid content, as well as retaining greater total phenol and anthocyanin concentrations in strawberries. Total phenol conc.(mg/g) ≈0.3 6 (storage time) Decay percentage (%) ≈-40% Total phenol conc.(mg/g) ≈0.12 3 3 (storage time) Decay percentage (%) ≈-12.5% Total phenol conc.(mg/g) ≈0.37 6 (storage time) Decay percentage (%) ≈-42% Total phenol conc.(mg/g) ≈0.21 Alotaibi and Tahergorabi (2018) Thyme essential oil 2 Sweet potato starch based-coating of refrigerated storage with essential oil and studying the influence on the shrimp quality 4 Hardness (N) 5.8 • Significant reduction in the bacterial population by thyme. • A sweet potato starch-based coating incorporated with thyme essential oil might be a feasible option for preserving shrimp meat quality and reducing losses. Resilience -0.03 4 Hardness (N) 7.06 Resilience -0.01 Fernández et al. (2020) Citronellol 2 (mg/mL) Develop an antifungal hybrid filler for coatings that is both environmentally friendly and affordable 10 C. globosum inhibition (%) 77 • Plant extracts containing EOs can be utilized as antifungal agents. • The main active component was found to be citronellol. • Producing functional bioactive hybrids was the benefit of blending citronellol. A. alternata inhibition (%) 78 5 (mg/mL) C. globosum inhibition (%) 96 A. alternata inhibition (%) 98 Citral 2 (mg/mL) C. globosum inhibition (%) 75 A. alternata inhibition (%) 75 5 (mg/mL C. globosum inhibition (%) 98 A. alternata inhibition (%) 98 Vital et al. (2018) Ginger 0.10% Acceptability of fish fillet with an alginatebased coating containing essential oils 7 Shear force (N) -1.85 • Reduction in color losses and` lipid oxidation of fish fillet. • The antioxidant activity increased. • Using these additives is an effective way to eliminate undesirable properties of the food. Weight loss (%) -0.03 pH -0.04 Oregano Shear force (N) 1.58 Weight loss (%) 0.18 pH -0.06 Buendía et al. (2020) Carvacrol:or egano: cinnamon (70:10:20) ≈1% Coating of cardboard, including EO entrapped within cyclodextrins nanotube 6 TA (% citric acid) 0.001 • The lowest decay incidences and the highest firmness were for cyclodextrins nanotube blended with EOs. • That blend showed the highest antimicrobial activity compared to samples without plant oil. • The changes in pH were negligible. • The additives did not influence the physicochemical quality of the product (tomatoes). Firmness (N) 3.2 6 with a supplementary commercialization period TA (% citric acid) 0.015 Firmness (N) 1.5 Coating of cardboard including EO entrapped within cyclodextrins halloysite nanotube 6 TA (% citric acid) -0.01 Firmness (N) -2.7 6 with a supplementary commercialization period TA (% citric acid) -0.029 Firmness (N) 2.7 Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2122 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 11. continued. Reference Plant oil Oil content (%) or concentration Purpose Time (d) (factor) Important property Influence on the property Important information Klangmuang and Sothornvit (2018) Plai 15g/L Incorporating essential oils into hydroxypropyl methylcellulose-based nanocomposite on mango (cv. Namdokmai Sithong) against an anthracnose disease 15 Weight loss (%) From ≈9 to ≈7.4 • Incorporating essential oils into the matrix inhibited C. gloeosporioides fungus. • Ginger showed the highest performance. • Essential oils did not influence the taste, flavor, and quality of mango and extend the fruit’s shelf life. Disease severity (score) From ≈3 to ≈2.2 Firmness (N) From ≈1.95 to ≈7 Color From ≈4.35 to ≈4 Ginger Weight loss (%) From ≈9 to ≈7.8 Disease severity (score) From ≈3 to ≈1.9 Firmness (N) From ≈1.95 to ≈6.8 Color From ≈4.35 to ≈3.7 Majdinasab et al. (2020) Shirazi thyme 2% The influence of used EOs on antimicrobial and antioxidant coating and shelf-life extension of chicken fillet 10 Cooking loss (%) -15.5 • The change in color by thyme was less than summer savory. • The shelf-life of chicken fillets is extended. • The overall acceptability was more than the control (Basil-seed gum) when they used EOs, and thyme influence was higher than summer savory. Overall acceptability From 2.4 to 3.8 Texture From 2.4 to 4.4 Summer savory Cooking loss (%) -13.5 Overall acceptability From 2.4 to 3.5 Texture From 2.4 to 4.3 printing inks, candles, and polishes due to their distinctive properties (Zhukov and Popov, 2022). The global wax and wax ester markets, particularly in the UK, the USA, Brazil, Japan, and other countries, are highly appealing and anticipated to grow at a rate of 3.83, reaching USD 15.91 billion by 2030 (Fig. 3) (Verified Market Research, 2023). While some plants naturally produce wax esters in their seed oil, like jojoba, they are unsuitable for large-scale cultivation as they yield an undesirable mixture of very long-chained wax esters unfit for technological applications (Langsdorf et al., 2021; Simonsen et al., 2023). However, numerous other plants show promise in producing wax esters, such as sunflower seed, olive, palm, Camelina sativa, and Arabidopsis thaliana, exhibiting high performance (Qi et al., 2020; Clews et al., 2023). Olive and palm oil-based wax esters are particularly noteworthy, with those derived from olive oil consisting of long, straight-chain fatty alcohols esterified with FA (Abdelmoez and Mustafa, 2014; Mariani et al., 2018). Several factors influence the rate of esterification in olive oil, including refining process variables, storage conditions, and reagent concentration (Mariani et al., 2018; Diarte et al., 2021). 2.12. Textile finishing Washing, bleaching, dyeing, and coating constitute the primary stages of the textile finishing process, utilized to enhance the bulk of textiles or garments post-weaving and synthetic material manufacturing (Achaw and Danso-Boateng, 2021; Al-Sayed and Abdelrahman, 2021). Textile finishing plays a crucial role in determining the final appearance and aesthetic qualities of textiles while also imparting desirable properties such as flame retardancy, wrinkle resistance, water-and-oil repellency, and more (Haule and Nambela, 2022; Javaid et al., 2024). Over the projected period (2017-2024), the global market for textile finishing chemicals is forecasted to grow at a CAGR of 3.8%, rising from USD 8.9 billion in 2022 to USD 12 billion in 2030 (Prescient and Strategic Intelligence, 2023). However, a significant portion of textile finishes currently available in the market pose potential harm to the environment and human health, with compounds like triclosan known for accumulating in aquatic environments and posing toxicity risks to aquatic organisms (Bhat et al., 2022; Periyasamy, 2023). In light of these concerns, plant oils present a sustainable alternative for use as additives in textile finishing processes (Natarajan et al., 2022). One notable application is their use as insect repellents, leveraging their natural properties to repel insects, along with their utilization in aromatherapy textiles for antibacterial, olfactory, and medicinal purposes (Mittal et al., 2019). It is important to note that antimicrobial textiles, while effective, maintain eco-friendly credentials, making them favorable in contemporary contexts, particularly amidst the COVID-19 crisis, which has underscored the importance of controlling infectious diseases (Bouaziz et al., 2021). Beyond textile applications, plant oils serve various industries. In medicine, they demonstrate potent antifungal, antimicrobial, anticancer, and wound-healing properties. In the food and food packaging sectors, they function as natural antifungal and antibacterial agents. Additionally, in the cosmetics and perfume industries, their aromatic qualities make them invaluable ingredients. Plant oils also contribute to mining processes through ore froth flotation and serve as surfactants in oil-well drilling muds. Moreover, in rubber production, plant oils serve as vulcanizing agents, softeners, and mold release agents. They also find use as solvents and antidust agents, and they aid in paper recycling by efficiently removing printing inks. Subsequent sections will delve into plant oil production pathways and fatty acid synthesis. 3. Plant oil production pathways Plant oils primarily consist of TAGs and are predominantly stored in the seeds or mesocarp of fruits (Ge et al., 2021; Hernández et al., 2021). However, notable exceptions exist, such as Simmondsia chinensis (jojoba), where oils accumulate in the form of esters of long-chain alcohols and FA (Guzha et al., 2023). From a chemical standpoint, TAGs result from the esterification of FA (C8–C24) with glycerol (Wei et al., 2024). It is intriguing to note the close similarity between the generic chemical formulas of these FA and those of fossil-oriented hydrocarbons, i.e., CH3(CH2)nCOOH vs. CH3(CH2)nCH3, respectively (Rajaeifar et al., 2019). This similarity has sparked widespread investigations into using plant oils and their derivatives as alternatives to fossil fuels. Overall, the properties of plant oils and their applications are largely attributed to their fatty acid composition (Mannu et al., 2020). Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2123 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. It is important to acknowledge that while palmitic acid (16:0), stearic acid (18:0), oleic acid (18:1), linoleic acid (18:2), and linolenic acid (18:3) are the major FA found in plant oils (Wei et al., 2019; Piovesana et al., 2021), there is remarkable diversity among different plant species in terms of their oil compositions. For instance, there are over 300 variations in fatty acid profiles in higher plants (Liu et al., 2022b). Moreover, some plants produce uncommon FA, further enriching plant oil compositions. Examples of such uncommon FA include short carbon chain FA (C8 to C14) in Cuphea, long carbon chain FA (C20 to C24) in rape, Crambe, B. napus, and A. thaliana, and hydroxy fatty acid (C18:0-OH) in castor bean (R. communis), among others (Roscoe et al., 2015). From an application standpoint, these FA impart specific characteristics that make these oils suitable for use across various industries. Consequently, such plants offer unique gene pools that could be leveraged to modify or manipulate conventional oil crops. To achieve this objective, it is crucial to gain in-depth insights into the metabolic pathways governing plant oil production, including the involved genes and their regulatory mechanisms. Therefore, fatty acid synthesis and modification (elongation and desaturation), as well as TAG synthesis and accumulation, are presented and discussed in the subsequent sections. 3.1. Fatty acids synthesis The synthesis of FAs, primarily occurring in plastids, serves as the cornerstone of plant oil production pathways. As illustrated in Figure 4, the process commences with the carboxylation of acetyl-CoA, leading to the formation of malonyl-CoA. It is noteworthy that acetyl-CoA is synthesized differently in photosynthetic (e.g., leaves) and non-photosynthetic (e.g., fruits and seeds) plant tissues (Li-Beisson et al., 2016). In photosynthetic tissues, acetyl-CoA is produced in chloroplast stroma through the fixation of CO2 into triose-phosphates (TP) via the Calvin cycle (Fig. 4a). These TPs are subsequently converted into pyruvate and eventually acetyl-CoA through the glycolysis pathway (Tang et al., 2022). Alternatively, TPs may be directed into the starch synthesis pathway (Xu et al., 2024) (Fig. 4a). In contrast, sucrose acts as the precursor for acetyl-CoA synthesis in nonphotosynthetic tissues. However, as sucrose cannot enter plastids where acetyl-CoA is synthesized, it undergoes cleavage into its constituent components by invertases or sucrose synthases. The resulting hexoses are then converted into hexose phosphates (i.e., glucose 6-phosphate and fructose 6-phosphate) (Xu et al., 2024). These hexose phosphates undergo Fig. 4. Overview of major reactions involved in fatty acid and triacylglycerol synthesis. (a) Glycolysis is used to make proteins, storage starches, or lipids from absorbed carbohydrates from photosynthesis. Sucrose is transported from photosynthetic tissues into developing seeds, where it is metabolized into precursors such as glucose 6-phosphate and phosphoenolpyruvate in the cytosol of embryo and/or endosperm cells before being transferred to plastids for fatty acid synthesis.; (b) De novo fatty acid production and modification: the precursors for the synthesis of C8–C18 saturated fatty acyl-ACPs on a plastidial multienzyme fatty acid synthetase complex include acetyl-CoA and malonyl-CoA. Unsaturated and monounsaturated fatty acids are transported from plastids to the endoplasmic reticulum for further processing by an acyl-CoA transporter (ACT). (c) TAG synthesis. A complex process combining successive acylation of a glycerol moiety and substantial acyl editing via phosphatidylcholine-dependent desaturases or desaturase-like enzymes results in triacylglycerols. Abbreviations: TP: Triose phosphate; G6P: Glucose 6phosphate; OPP: Oxidative pentose phosphate; Pyr, pyruvate; ACCase, acetyl-CoA carboxylase; FAS, FA synthase; FA, fatty acid; FFA, Free Faty Acid; CoA, coenzyme A; LACS, Long Chain acyl-CoA Synthetase; G3P, Glycerol 3-phosphate; GPAT: G3P acyltransferase; LPA: lysophosphatidic acid; LPAAT: lysophosphatidic acid acyltransferase; PA: phosphatidic acid; PP: phosphatase; DAG: diacylglycerol; DAGAT: DAG acyltransferase; TAG: triacylglycerol; DGAT: diacylglycerol acyl trans acylase; PDAT: phosphatidylcholine-dependent acyltransferase; PDCT: phosphatidylcholine: diacylglycerol choline phosphotransferase; PC: phosphatidylcholine; PLA2: phospholipase A2; LPC: Lys phosphatidylcholine; LPCAT: Lys phosphatidylcholine acyltransferase; DES: Desaturase; CPT: CDPcholine: DAG choline phosphotransferase. Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2124 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. further conversion into various intermediates, such as phosphoenolpyruvate and malate, through the glycolysis and/or the oxidative pentose phosphate (OPP) pathway before being transported into plastids for acetyl-CoA production (Seliniski and Scheibe, 2019; Bu et al., 2023). In the subsequent step, the pivotal reaction catalyzed by acetyl-CoA carboxylase (ACCase) results in the formation of malonyl-CoA, the building block of the FA synthesis pathway, via acetyl-CoA carboxylation (Fig. 4b). The fatty acid synthase enzyme complex (including 3-ketoacylACP synthase of type III, type I, and type II, or KASIII, KASI, and KASII, respectively, as well as acyl-ACP thioesterases, FAT A or FAT B) then utilizes malonyl-CoA to synthesize FAs ranging from 4 to 18 carbons in length (Bates et al., 2013). Initially, KASIII catalyzes the condensation, yielding C4:0 FAs. Subsequent condensation reactions are carried out by KASI, producing FAs up to C16:0. Finally, KASII facilitates the final steps of FA elongation from C16:0 to C18:0 (Manan et al., 2017). FAT A or FAT B plays a crucial role in determining the chain length of the synthesized FAs within the enzyme complex (Liu et al., 2022a and b). Notably, the accumulation of medium-chain FAs, such as C10:0 and C12:0, involves the presence of evolved FAT A or FAT B, capable of premature hydrolysis of the growing acyl thioesters. Considering the significant role of acetyl-CoA in the overall FA synthesis pathway, the reaction it is involved in is regarded as a major rate-limiting step (He et al., 2020). Furthermore, FA ranging from C8:0–C18:0 may undergo unsaturation catalyzed by fatty acid desaturases (FADs), specifically FAD6, 7, and 8 (Manan et al., 2017), while still within plastids (Elahi et al., 2016). The resulting saturated and monounsaturated FAs in the C8–C18 range are transported as acyl-CoAs to the endoplasmic reticulum (ER) for further modification and elongation (Bu et al., 2023; Fell et al., 2023). Continued FA elongation in the ER involves a series of reactions utilizing cytosolic malonyl-CoA and imported acyl-CoA (Bu et al., 2023; Fell et al., 2023). These reactions are catalyzed by an ER-specific fatty acid elongation complex comprising four enzymes: 3-ketoacyl-CoA synthase, 3-ketoacylCoA reductase, 3-hydroxyacyl-CoA dehydrase, and enoyl-CoA reductase (Bu et al., 2023; Fell et al., 2023). This enzymatic complex facilitates the production of FAs up to C24. Additionally, Fatty acid desaturase 2 (FAD2) and Fatty acid desaturase 3 (FAD3) are responsible for sequential desaturation reactions occurring in the ER, leading to the formation of polyunsaturated FA (Manan et al., 2017). 3.2. Triacylglycerol biosynthesis TAG synthesis generally occurs through a series of reactions known as the Kennedy pathway, located in the endoplasmic reticulum (ER) (Kim, 2020) (Fig. 4c). This pathway initiates with the introduction of glycerol-3phosphate (G3P), which is sequentially acylated by acyl-CoAs of chloroplast origin (Schmid, 2021). G3P is generated from dihydroxyacetone phosphate (DHAP) through a reaction catalyzed by glycerol-3-phosphate dehydrogenase (G3PDH) (Kim, 2020). The enzymes involved in this sequential acylation process include glycerol-3-phosphate acyltransferase (GPAT), lysophosphatidic acid acyltransferase (LPAAT), and diacylglycerol acyltransferase (DGAT) (Li et al., 2015a). Prior to the third acylation by DGAT, a dephosphorylation step is necessary, which is catalyzed by phosphatidic acid phosphatase (PAP) (Kim, 2020; Lutkewitte and Finck, 2020). It is noteworthy that TAG biosynthesis may involve additional complexities beyond the traditional Kennedy pathway. For instance, acylCoAs could also be supplied from membrane lipid phosphatidylcholine (PC) through alternative pathways. Acyl-CoAs may be incorporated or released from PC through a series of reactions known as acyl editing reactions, catalyzed by lyso-PC acyltransferase (LPCAT). Alternatively, a reaction catalyzed by phospholipase A2 (PLA2) may result in the release of free FA, which can then be acylated to form acyl-CoA (Lutkewitte and Finck, 2020; Schmid, 2021) (Fig. 4c). Another alternative pathway complicating TAG biosynthesis is the incorporation of diacylglycerol (DAG) into phosphatidylcholine (PC), catalyzed by choline phosphotransferase (CPT) (Kim, 2020). The existing acyl groups can be desaturated by fatty acid desaturase (FAD). These PCincorporated DAG molecules can then be acylated into TAG through reactions catalyzed by phospholipid: diacylglycerol acyltransferase (PDAT) or PC:DAG phosphocholine transferase (PDCT) (Kim, 2020). It is important to note that the reaction catalyzed by CPT is reversible, allowing DAG to be released at later stages and converted into TAG through the catalytic action of DGAT (Eichmann and Lass, 2015). 3.3. Triacylglycerol storage Upon completion of TAG biosynthesis, TAGs are stored within oil bodies (OBs), also known as oleosomes, ranging from 0.2 to 2.5 µm (Lutkewitte and Finck, 2020). The structure of OBs comprises a stabilizing monolayer of phospholipids containing proteins such as oleosins, caleosins, steroleosins, and aquaporins (Lutkewitte and Finck, 2020). Among these proteins, oleosins, being the most abundant, play pivotal roles in the TAG storage capacity of various plant tissues and across different plant species. Indeed, the absence of these proteins in vegetative tissues is the reason behind their inability to store lipids (Xu et al., 2024). Mechanistically, these proteins prevent the coalescence of oil bodies during storage through steric hindrance and electronegative repulsion, thus determining the final size of the OBs (De Chiriko et al., 2018; Kanai et al., 2019). For example, Hu et al. (2009) suggested that a low concentration of oleosins is associated with the formation of larger OBs and, consequently, lower oil contents. 4. Genetic engineering for improved oil production In recent decades, researchers have devoted significant attention to enhancing the quality and quantity of seed oils for both food and non-food purposes, as well as exploring new oil sources (Subedi et al., 2020a and b; Bhati et al., 2021; Chen et al., 2021a; Sheri et al., 2021). The fatty acid composition of plant oils plays a crucial role in determining their quality and suitability for various applications, primarily by distinguishing between saturated and unsaturated FA. Saturated FA lack double bonds in their carbon structure and exhibit a linear configuration, whereas unsaturated FA contain at least one double bond in their carbon chains. Vegetable oils with a higher monounsaturated-to-saturated fatty acid ratio demonstrate improved stability under high-temperature conditions, such as during frying, and exhibit prolonged shelf life when stored. In oils intended for human consumption, cooking oils typically contain higher proportions of monounsaturated FA, such as oleic acid, while salad oils predominantly comprise polyunsaturated FA like linoleic and α-linolenic acids (Kapoor et al., 2021; Saini et al., 2021). To meet the increasing demand for oils, whether for human consumption or other applications like biofuels, metabolic and genetic engineering techniques, including the utilization of recombinant DNA technology, have been employed to manipulate oil content in plants and broaden the range of oil varieties (Fig. 5). Table 12 provides an overview of the fatty acid composition in various plants and vegetable oils. 4.1. Metabolic engineering to enhance oil production Augmenting oil production in seeds has long been a primary objective for both plant breeders and genetic engineers (Subedi et al., 2020a and b; Bhat et al., 2022; Rauf et al., 2023). Oilseed crops inherently possess higher seed oil content compared to other agriculturally significant crops. However, there exists considerable variation among oilseed crops themselves, with oil content ranging from 20% in soybeans to 60% in sesame, despite similarities in their lipid biosynthesis pathways (Zafar et al., 2019). Remarkably, a mere 1% improvement in soybean oil production per hectare could contribute over USD 1 billion annually to the crop’s global value (Bates et al., 2014), emphasizing the profound impact of metabolic engineering in oilseed crops, with the potential to significantly enhance seed oil contents. Metabolic engineering broadly aims to regulate flux into metabolic pathways by either increasing the availability of upstream substrates or strengthening the sink in the final stages of the pathway. Both strategies have been harnessed to increase seed oil accumulation. In the case of oilseeds, yield improvements can be achieved by increasing the amount of oil per seed, enlarging the size of the seed, or boosting the number of seeds per plant. Strategies to enhance oil production in plants include enhancing the availability of fatty acid (FA) precursors, elevating the rate of FA synthesis, improving TAG assembly processes, and restricting TAG degradation pathways (He et al., 2020; Subedi et al., 2020a and b; Kapoor et al., 2021). In the following section, we review the application of metabolic engineering for the improvement of oil content and its quality. Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2125 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Fig. 5. Genetic engineering of plant oils for different applications. Table 12. Fatty acid composition of plants and vegetables. Plant Fatty acid composition (%) Reference Saturated fatty acids Unsaturated fatty acids Others Caprylic 8:0 capric 10:00 Lauric 12:00 Myristic 14:0 Palmitic 16:0 Stearic 18:0 Arachidic 20:0 Behenic 22:0 Lignoceric 24:0 Cerotic 26:0 myristoleic 14:1 Palmitoleic 16:1 Sapienic 16:1 Oleic 18:1 Linoleic 18:2 α-linolenic 18:3 Eicosenoic 20:1 Erucic 22:1 Avocado - - - 0.33 23.6 1 - - - - - 3.58 - 47.2 13.4 0 - - 11.7 MorenoCamacho et al. (2019) Castor bean - - - - 1 1 - - - - - - - 3 4 trace - - 91 Aid (2020) Cocoa butter - - 0-1 0-4 24.533.7 33.740.2 1 - - - - 0-4 - 26.335 1.7-3 trace - - - Naik and Kumar (2014) Coconut 8 7 49 8 8 2 - - - - - - - 6 2 0 - - - Boateng et al. (2016) Corn - - - - 6.7-16.5 0.7-6.6 0-1 - - - - - - 16.243.8 39.569.5 0-3.1 - - - White (2007) Linseed - - - - 5--6 4--5 - - - - - - - 15-20 14 50-55 - - - Aid (2020); Bayrak et al. (2010) Olive - - - - 7.5-20 0.5-5 0-0.8 0-0.2 0-1 - - 0.3-3.5 - 55-83 3.5-21 0-1.5 - - - Tsimidou et al. (2003) Palm - - 0.2 1.1 44 4.5 0.1 - - - - - - 39.2 10.1 0.4 - - - Mancini et al. (2015) Palm kernel 3.3 3.5 47.8 16.3 8.5 2.4 0.1 - - - - - - 15.4 2.4 - - - - Mancini et al. (2015) Rapeseed - - - - 3.63 2.32 - - - - - - - 85.31 3.41 4.4 0.94 - - Guan et al. (2016) Soybean - - - - 10 4 - - - - - - - 18 55 13 - - - Clemente and Cahoon (2009) Sunflower - - - - 6 5 - - - - - - - 19 68 trace - - - Aid (2020) Cottonseed - - - - 25.39 2.33 - - - - 1 0.6 - 16.35 52.89 - - - - Yang et al. (2019) Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2126 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 12. continued. Plant Fatty acid composition (%) Reference Saturated fatty acids Unsaturated fatty acids Others Caprylic 8:0 capric 10:00 Lauric 12:00 Myristic 14:0 Palmitic 16:0 Stearic 18:0 Arachidic 20:0 Behenic 22:0 Lignoceric 24:0 Cerotic 26:0 myristoleic 14:1 Palmitoleic 16:1 Sapienic 16:1 Oleic 18:1 Linoleic 18:2 α-linolenic 18:3 Eicosenoic 20:1 Erucic 22:1 Almond - - - - 5.076.78 - - - - - - - - 57.5473.94 19.3235.18 0.040.1 - - - Sathe et al. (2008) Flaxseed - - - - 4.9-8 2.444.59 - - - - - - - 13.4419.39 12.2517.44 39.960.42 - - - Goyal et al. (2014) Hempseed - - - - 6 3 1 0.44 0.197 - - 0.098 - 16 55 15 - - 3.265 Sova et al. (2018) Walnut - - - - 5.615.82 - - - - - - - - 22.6227.27 49.9354.41 14.3217.82 - - - Dogan and Akgul (2005) Rice bran - - - 0.3 22.7 1.8 0.9 - - - - - - 43.9 29.2 1.25 - - - Latha and Nasirullah (2014) Pumpkin seed - - - 0.233 14.82 6.67 0.433 0.058 - - - - - 25.81 - 50.88 - 0.055 1.041 Bardaa et al. (2016) Sesame - - - - 7.9-12 4.8-6.1 - - - - - - - 35.942.3 41.547.9 - - - - Wacal et al. (2019) Peanut - - - - 8.2-15.1 1.1-7.2 0.8-3.2 1.85.4 0.5-2.5 - - - - 31.560.2 19.945.4 - 0.6-2.6 - - Hammond et al. (1997) Grapeseed - - - - 8.4-6.51 3.863.07 - - - - - - - 16.111.62 77.5972.5 - - - - Al Juhaimi et al. (2017) Beech nut - - - 0.18 10.2 5.81 0.47 0.61 - - - 0.25 - 37.2 34.1 4.1 5.63 0.83 0.27 Ayaz et al. (2011) Macadamia - - - - 9 2 - - - - - 22 - 60 2 - - - - Tan et al. (2020) Hazelnut - - - - 4.398.85 1.673.18 - - - - - - - 73.4881.57 10.4614.95 0.020.34 - - - Balta et al. (2006) Cashew - - - 0.01 10.3 8.8 0.14 0.13 0.14 0.01 - 0.37 - 61.8 17.3 0.14 - 0.01 0.43 Griffin and Dean (2017) Pecan - - - 0.4 6.012 - - - - - - - - 23.91 66.81 1.83 - - - Rivera-Rangel et al. (2018) Orange seed - - 2.96 0.89 12.6 8.9 0.54 - - - - - - 43.03 25.11 4.3 0.67 - - Iwuagwu et al. (2018) Citrus seed - - - - 25.832.2 2.8-4.4 - - - - - - - 21.924.1 36.139.8 3.4-4.4 - - - Reazai et al. (2014) Watermelon - - - - 9.88 6.96 0.26 - - - - - - 14.25 68.07 - - - - Rezig et al. (2019) Argan - - - - 11--14 4--7 - - - - - - - 46-48 31-35 - - - - Charrouf and Guillaume (2008) Mustard - - - - 1.87 1.52 1.13 1.4 0.41 - - - 0.1 15.7 12.99 6.18 - 48.8 9.83 El-Shenawy et al. (2014) Ben 0.03 - - 0.12 6.25 4.97 3.23 6.02 0.36 0.92 - - - 73.57 0.46 - - 0.11 1.88 Leone et al. (2016) Eucalyptol - - - - 36 3 - - - - - 7.3 - 27.2 19.3 - - - 3.6 Rekkab et al. (2012) Thyme 0.52 0.15 0.18 1.07 12.07 6.15 - - - - - 2.3 - 33.04 41.73 0.61 - - 2.18 Attia et al. (2017) 4.1.1. Increasing the content of oil per seed As mentioned previously, TAG biosynthesis involves three primary metabolic steps: glycolysis, FA synthesis, and TAG production (Yang et al., 2022b). Glycolysis, as the initial step in TAG synthesis, provides the carbon resources necessary for TAG production. Metabolic engineering strategies aimed at increasing oil accumulation in plants may involve promoting the production of precursors for lipid synthesis (e.g., G3P and acetyl-CoA), enhancing the rate of FA synthesis, increasing TAG assembly, and inhibiting TAG degradation pathways (Xu et al., 2024). Augmenting FA accumulation is achievable through various approaches, including increasing the carbon flux towards FA synthesis, overexpressing transcriptional regulators in FA synthesis, promoting the expression of ACCase, the pivotal enzyme in plastid FA synthesis, suppressing competing pathways for FA utilization (e.g., membrane lipid synthesis), and enhancing the allocation of carbon for FA synthesis by limiting starch synthesis (Mulgund, 2022; Ranjbar and Malcata, 2022). The lipid metabolic pathways in plants are highly intricate, involving at least 120 enzymatic reactions and over 600 genes (Lee et al., 2017; Vanhercke et al., 2019). However, physiological and developmental factors can influence oil biosynthesis in plants (Li et al., 2020a; Yang et al., 2022b). Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2127 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Additionally, regulatory pathways governing carbon flux to storage and membrane lipid biosynthesis remain incompletely understood (Li et al., 2020b; Yang et al., 2022b). Therefore, a comprehensive understanding of the complex networks of oil biosynthesis and its regulatory framework in plants can aid in identifying and engineering key genes involved in increasing oil content (Savadi et al., 2017). 4.1.1.1. FA synthesis (Carbon flux redirection) Increasing seed oil content necessitates redirecting carbon flux towards fatty acid biosynthesis, as it can significantly impact TAG production (Muñoz et al., 2021; Song et al., 2023). Genes encoding enzymes associated with the FA biosynthetic pathway are key targets for genetic modification (Savadi et al., 2017). However, it is important to note that attempts to enhance the availability of precursors for FA synthesis by altering central carbon metabolism have not always been successful, as flux control in a pathway typically involves multiple components rather than individual enzymes (Song et al., 2023). ACCase, the rate-determining enzyme governing FA biosynthesis, has been extensively studied in various model plants (Wan et al., 2017a). ACCase catalyzes the production of malonyl-CoA, providing two carbon units to the FA synthase complex. Overexpression of the cytosolic multifunctional form of ACCase in the plastids of rapeseed led to a modest increase in seed lipid content (Zhukov and Popov, 2022). Conversely, overexpression of the gene encoding the biotin carboxyl carrier protein 2 (BCCP2), a component of plastidial ACCase, resulted in decreased seed oil content and increased levels of sugar and protein (Zhukov and Popov, 2022), indicating that ACCase activity is not the sole limiting factor in FA synthesis (Savadi et al., 2017). Mitochondrial pyruvate dehydrogenase (PDH), responsible for converting pyruvate and coenzyme A (CoA) into acetyl-CoA, is regulated negatively by pyruvate dehydrogenase kinase (PDHK). Repression of PDHK activity through antisense RNA technology during seed maturation has been shown to enhance seed oil content and seed weight in A. thaliana (Subedi et al., 2020a and b). Silencing the Pyruvate dehydrogenase kinase cDNA (AtPDHK) gene using RNAi technology increased seed oil in transgenic plants by up to 50%, altering fatty acid composition (Schewender and Hey, 2012). Additionally, pyruvate, a key product of glycolysis, serves as a major carbon source for fatty acid production in plastids (Schewender and Hey, 2012). Seed-specific overexpression of a pyruvate transporter, BASS2, increased oil production by 24-43% in transgenic Arabidopsis compared to wild-type plants (Lee et al., 2017), highlighting the potential of promoting oil biosynthesis by enhancing pyruvate flux into plastids. 4.1.1.2. Glycerol backbone G3P plays a critical role in regulating the production and retention of TAGs in plant tissues. However, G3P supply has been shown to be insufficient to maintain high G3P levels during the peak oil accumulation period in oilseed crops (Pollard and Hill, 2021). G3P is generated in plants through two distinct enzymes: cytosolic glyceraldehyde 3-phosphate dehydrogenase (GAPDH) and glycerol kinase (GK) (Jeelani et al., 2023). Glycerol kinase converts glycerol into G3P, primarily in germinating seeds, while cytosolic GAPDH catalyzes the conversion of dihydroxyacetone phosphate (DHAP) into G3P in various plant tissues, including seeds. Genetic modification to enhance G3P levels has been successful in boosting seed oil content. For instance, the seedspecific expression of yeast cytosolic GAPDH in B. napus resulted in a 34-fold increase in G3P levels and a 40% increase in seed oil content. Similarly, engineering C. sativa by co-expressing A. thaliana diacylglycerol acyltransferase1 (DGAT1) and yeast cytosolic GAPDH led to a 13% increase in seed oil content and up to a 52% increase in seed mass compared to wild-type plants (Chhikara et al., 2018). Moreover, overexpression of DGAT1 in transgenic tobacco plants resulted in a 2.19-fold increase in oil content compared to non-transgenic control lines (Chenarani et al., 2022). 4.1.1.3. TAG biosynthesis TAGs in plants are synthesized through the Kennedy pathway in the ER via a sequential acylation of G3P by membrane-bound glycolipid acyltransferases. These enzymes include GPAT, LPAAT, and diacylglycerol acyltransferase (DGAT) (Chenarani et al., 2022). The first acylation of G3P is carried out by GPAT, which produces lysophosphatidic acid in the ER, chloroplast (CHP), or mitochondria (MIT). LPAAT, located in the ER, CHP, MIT, plasma membrane (PM), and cytoplasm (CP), performs the second acylation of lysophosphatidic acid to generate phosphatidic acid. DGAT completes the final step in the Kennedy pathway by converting diacylglycerol (DAG) into TAG (Wan et al., 2017a). Since these enzymes are rate-limiting in TAG synthesis, researchers have explored genetic modifications to enhance LPAAT, GPAT, and DGAT activity in plants. Among these, DGAT is considered a critical rate-limiting step due to its comparatively lower enzyme activity compared to the other enzymes in the pathway (Wan et al., 2017a). Enhancing GPAT and LPAAT activity has proven effective in increasing seed oil content. Arabidopsis genetically engineered to express plastidial safflower GPAT and E. coli GPAT exhibited increased seed oil content during seed maturation (Wan et al., 2017b). Similarly, the expression of genes from B. napus encoding microsomal LPAAT isoforms in Arabidopsis seeds unexpectedly increased seed total FA content and seed weight (Zhang et al., 2022). Furthermore, the expression of a mutant yeast LPAAT gene (SLC1-1) under the control of the CaMV35S promoter in Arabidopsis and high-erucic acid B. napus resulted in a 48% increase in seed oil content (Wan et al., 2017b). 4.1.1.4. Lipid transfer proteins Lipid transfer proteins (LTPs), also known as plant lipid transfer proteins (PLTPs), are small, compact proteins with a hydrophobic cavity that facilitates the transfer of phospholipids and FA between cell membranes (Getz, 2018). Known for nearly half a century, LTPs are divided into two structurally related subfamilies based on their molecular weights: LTP1s (9 kDa) and LTP2s (7 kDa) (Fan et al., 2013). Puroindoline A and B (PINA and PINB) proteins exhibit a structure similar to that of non-specific lipid-transfer proteins found in plants (Zhang et al., 2019b). Heterologous expression of wheat (Triticum aestivum L.) puroindoline genes (PINA and PINB) in corn plants (Zea mays L.) increased the germ size without affecting the seed size. As a result, the overall seed oil content increased by 25%, and the germ yield rose by approximately 34% (Zhang et al., 2019b). 4.1.1.5. Transcription factors (TFs) In general, the accumulation of seed oil requires coordinated transcriptional control of numerous biosynthetic pathways (Kong et al., 2019; Niu et al., 2020; Yang et al., 2022b). Multiple transcription factors (TFs) play a pivotal role in synchronizing the regulatory networks of genes involved in fatty acid and TAG biosynthesis. These TFs control gene expression by recognizing and binding to specific sequences in gene promoters (Kong et al., 2019; Papadimitriou et al., 2019; Tian et al., 2019; Kumar et al., 2020). Manipulating TFs represents an alternative strategy for enhancing seed oil yield alongside single-enzyme approaches (Santin et al., 2021; Yang et al., 2022a). Positive TFs, such as LEAFY COTYLEDON genes (LEC1 and LEC2), FUSCA3 (FUS3), ABSCISIC ACID INSENSITIVE3 (ABI3), and WRINKLED1 (WRI1), are known for their pivotal roles in regulating the activities of other TFs involved in seed maturation and oil accumulation (Fatihi et al., 2013; Kumar et al., 2020). However, to regulate the expression of these positive TFs, plants have evolved mechanisms to suppress their expression during the vegetative phase of development. Negative regulators, including TFs, microRNAs (miRNAs), and proteins, act by modulating the chromatin conformation of their target genes (Kumar et al., 2020). A common strategy for manipulating seed oil content in plants is overexpression of positive regulators such as WRI1, LEC1/2, and MYB96 or suppression of negative regulators like MYB89 and WRKY6. Ideally, genetic engineering should focus on TFs that activate FA and TAG biosynthesis genes. Overexpressing TFs involved in FA synthesis can lead to increased TAG content. For example, overexpressing MYB96, a TF that activates TAG biosynthesis genes DGAT1 and PDAT1, can significantly boost TAG storage in seeds. However, overexpressing certain TFs, chromatin remodeling agents, and miRNAs can have unintended consequences on plant growth, development, and yield. Therefore, tissue- Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2128 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. specific overexpression or silencing of desired genes in transgenic seed oil plants is recommended. WRINKLED1 (WRI1), belonging to the APETALA 2/ethyleneresponsive element binding protein (AP2/EREBP) subfamily of TFs, plays a crucial role in regulating plant oil accumulation by promoting carbon partitioning into FA synthesis within the glycolysis pathway. WRI1 regulates the expression of at least 15 enzymes involved in FA synthesis and the glycolytic pathway, making it an ideal target for genetic manipulation to enhance oil accumulation in plants. Overexpressing transcription factors that regulate WRI1 expression have been shown to increase seed oil content in transgenic plants. For example, overexpressing soybean ZF351 and GmDREBL increased oil content in transgenic Arabidopsis by binding to the WRI1 promoter. Similarly, overexpressing maize ZmLEC1, an activator of WRI1, led to elevated oil content in Arabidopsis, Camelina, and maize, though it had unintended effects on seed germination and leaf growth. Conversely, overexpressing maize ZmWRI1 increased oil content without these side effects. Numerous studies have shown that overexpressing WRI1 or its orthologs significantly boosts seed oil content in transgenic plants. For instance, constitutive expression of WRI1 in Arabidopsis leads to a 10% to 40% increase in seed oil content and an increase in seed size. Selecting an appropriate promoter to drive WRI1 expression is critical for genetically engineering oil accumulation in the seeds of transgenic plants. For instance, using the FUS3 or Oleosin promoter to drive WRI1 can increase seed oil content, while using the ZEIN promoter does not result in distinguishable changes in oil content. Table 13 shows the transcription factors and their overexpression on oil content. 4.2. Increasing seed oil content by enhancing seed size Both genetic and environmental factors shape the final weight and dimensions of plant organs during growth and development. In oilseed crops, seed size significantly impacts crop productivity and yield. Thus, increasing seed size and biomass emerges as a strategy to enhance oilseed crop yield. However, it is crucial to consider that augmenting seed weight might necessitate a reduction in seed numbers due to limited assimilation, potentially affecting seed filling. Furthermore, altering seed oil levels can have repercussions on total seed protein and vice versa when working with fixed assimilates. Hence, the key aspect in boosting oilseed plant yield is ensuring genetic manipulation does not adversely affect seed numbers. Redirecting carbon flux toward oil production rather than protein becomes imperative in this regard. Research on seed size regulation has identified around 88 key regulators responsible for organ size, associated with pathways like KLUH, ubiquitinproteasome, G-protein signaling, Mitogen-Activated Protein Kinase (MAPK), and plant hormone pathways. Enhancing seed oil content involves expressing genes related to seed development, embryo size, and oil storage tissues in crops such as Arabidopsis, maize, and rice. One approach is to increase oil-carrying tissues while minimally altering non-oil-carrying seed tissues. For instance, enhancing seed oil content in the endosperm, a novel storage tissue in plants, is a viable strategy. These insights into seed size regulation and oil content enhancement pave the way for improving oilseed crop productivity and yield. Seed size regulation involves a complex interplay of factors, such as plant hormones, ubiquitin, microRNAs, and cytochrome P450s (CYPs). The CYP78A subfamily, a plant-specific gene family, plays a crucial role in controlling plant growth and organ size. Various subfamilies of CYP78A identified in Arabidopsis, rice, wheat, tomato, and soybean regulate organ size and development. Overexpressing CYP78A genes in different plant species has demonstrated significant effects on seed size and organ development. Additionally, genes and regulatory elements like SWEET, BS1, LEC1, ARF2, DA1, DA2, Abscisic acid biosynthesis-related genes (ABA2 and ABI5), TTG2, AP2, RING-type E3 ubiquitin ligases, Enhancer of DA1 (EOD1), and miRNA172 impact seed size through diverse mechanisms. Enhancing the sink strength of seed oil can increase the number of oilstoring bodies in seeds. Elevating the oleosin protein content in developing seeds enhances oil storage capacity and promotes oil biosynthesis and storage in oil bodies, consequently increasing seed oil content. Overexpressing oleosin genes has proven effective in increasing seed oil content in plants like Arabidopsis and rice. Achieving embryo-specific overexpression of the appropriate oleosin gene is crucial to augmenting seed oil content without unintended effects. Moreover, transcription factor ARF2, responsive to auxin, acts as a repressor of cell division, elongation, and organ growth. Mutations in ARF2 can lead to enlarged seed size and weight. Tissue-specific expression of wild-type ARF2 restores normal flower morphology and fertility, underscoring its role in seed setting. Similarly, overexpression of WRI1, a member of the AP2/EREBP family, has been shown to increase seed size in transgenic plants. These findings shed light on the intricate regulatory mechanisms governing seed size and oil content in plants, offering avenues for enhancing crop productivity and yield. 4.3. Genetic Engineering for new oil resources: biomass-derived oil In addition to traditional plant seeds and fruits, various plant tissues can synthesize TAG or plant oils. This capability has spurred research into genetic engineering strategies to increase oil content in leaves and other vegetative tissues of high biomass crops in an environmentally sustainable manner. Such modified plant biomass can serve as an energy-dense, nutritionally valuable resource suitable for electricity production, biodiesel, renewable fuel production, and nutritionally enhanced animal feed. Vegetative plant organs typically contain around 1.5% TAG and 5-10% FAs on a dry weight basis, making TAG an essential buffer and intermediate storage pool for toxic and excess FAs released during membrane fatty acid turnover. Despite this, accumulating high levels of TAG in vegetative tissues similar to oilseeds is challenging due to the complex nature of its biosynthetic pathways. Metabolic engineering studies in model plants have suggested that crops with a 15% improvement in biomass oil content could produce up to ten times more oil per unit area compared to canola oilseed crops. Early efforts focused on the overexpression or downregulation of individual genes involved in FA or TAG biosynthesis, resulting in only modest increases in TAG content in vegetative tissues. Recent approaches have combined multiple metabolic engineering strategies to achieve higher TAG levels in plant biomass. 1. Triacylglycerol assembly (Pull) approach: This strategy involves targeting the TAG assembly process to increase the demand for precursors that influence the flux toward TAG accumulation in plant tissues (Vanhercke et al., 2017; Wan et al., 2017b; Park et al., 2021). Overexpressing TAG assembly enzymes has been suggested as an effective means to boost TAG levels in vegetative tissues (Xu and Shanklin, 2016). For instance, overexpressing Arabidopsis DGAT1 in N. benthamiana led to a 20-fold increase in leaf TAG content, while targeted expression of the same gene in xylem tissues resulted in a 63% increase in FAs production (Nookaraju et al., 2014). Additionally, overexpression of Chlamydomonas DGAT2 in Arabidopsis resulted in a 25-fold increase in TAG content and changes in TAG composition (Park et al., 2021; Sagun et al., 2023). Notably, overexpressing PDAT1 in transgenic Arabidopsis plants resulted in a remarkable 28-fold increase in TAG content in the leaves (Fan et al., 2013). Additionally, overexpressing mouse monoacylglycerol acyltransferase 1 (MGAT1) and MGAT2 led to diacylglycerol (DAG) accumulation, which subsequently resulted in approximately 7-9-fold increases in TAG content in transgenic tobacco leaves (Petrie et al., 2012; Sagun et al., 2023). 2. de novo FAs biosynthesis (Push) approach: This approach involves manipulating key factors like ACCase, WRI, and transcription factors such as LEC1, LEC2, FUS3, and ABI3 to boost the de novo synthesis of FAs (Weselake, 2016; Xu and Shanklin, 2016; Vanhercke et al., 2017). For example, the heterologous expression of Arabidopsis acetylCoA carboxylase (ACC1) in potato plants resulted in a five-fold increase in TAG content in transgenic tubers (Vanhercke et al., 2019; Luo et al., 2022). Overexpression of transcription factors (TFs) such as LEC2 and WRI1 has also been shown to enhance lipid content in leaves (Park et al., 2021; Luo et al., 2022). Constitutive expression of Arabidopsis WRI1 resulted in a 22-fold increase in TAG content in the transgenic vegetative tissues of Nicotiana benthamiana (Vanhercke et al., 2019). Interestingly, unlike LEC1 and LEC2, overexpression of WRI1 does not appear to have unintended effects on plant growth and development. Therefore, combining WRI1 overexpression with the regulation of other genes may be an effective approach to achieve a significant increase in oil content in the vegetative tissues of plants (Weselake, 2016; Wan et al., 2017b). Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2129 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 13. Transcription factors and their overexpression on oil content. Gene Function Host Plant Reference WRI1 Increased seed oil and TAG content Arabidopsis thaliana Sun et al. (2017) Lim et al. (2022), Chen et al. (2020) Camelina sativa An and Suh, (2015) Glycine max Chen et al. (2018), Wang et al. (2022c) Zea mays Pouvreau et al. (2011) Oryza sativa Sun et al. (2017) Jatropha curcas Ye et al. (2018) LEC1 Increased seed oil content Brassica napus Elahi et al. (2016) A. thaliana Zhu et al. (2018) C. sativa Zhu et al. (2018) Z. mays Shen et al. (2010) LEC2 Increased seed oil content A. A. thaliana Manan et al. (2017) TAG accumulation in leaves Kim et al. (2015) FUS3 TAG accumulation in vegetative tissues A. thaliana Zhang et al. (2016) ABI3 Increased oil accumulation in leaves A. thaliana Yang et al. (2021) DOF-type factors Gossypium hirsutum Su et al. (2017) GLABRA2 Increased seed oil content A. thaliana Chai et al. (2010) Basic leucine zipper (bZIPs) Elevated seed oil content A. thaliana Song et al. (2013) MYB TFs Increased seed oil content Arabidopsis and Lotus Li et al. (2017), Khan et al. (2019) Improved seed oil C. sativa Kim et al. (2019) SPT Correlation with seed oil content A. thaliana Liu et al. (2017) G-protein γ subunit 3 (AGG3) Increased seed size C. sativa Roy Choudhury et al. (2014) Purple acid phosphatase 2 (PAP2) Increased seed size C. sativa Zhang et al. (2012) RNAi suppression of AGPase Increased seed size C. sativa Na et al. (2018) CYP78A Increased seed size A. thaliana Fang et al. (2012) Increased seed size O. sativa Xu et al. (2015) Increased seed size C. sativa Holz and Dormann (2021) Increased seed size Nicotiana tabaccum Tian et al. (2016) Correlation with seed size B. napus, G. hirsutum Shi et al. (2019) Sun et al. (2017) GmSWEET10a Simultaneous increases in seed size and oil content G. max Wang et al. (2020b) BIG SEEDS1 (BS1) Increased seed size and weight G. max Ge et al. (2016) BIG SEEDS1 (SHB1) Increased seed size Brassica juncea A. thaliana, G. max Savadi et al. (2015) Ge et al (2016) KLUH Increased seed size A. thaliana, G. max Adamski et al. (2009) Zhao et al. (2016) IKU2 Increased seed size A. thaliana Fatihi et al. (2013) AUXIN RESPONSE FACTOR 2 (ARF2) Increased seed size A. thaliana Schruff et al. (2006) DA1 Increased seed size A . thaliana Li et al. (2008) Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2136 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. preliminary evaluation of the functionality as pharmaceutical ingredients. Environ. Technol. Innovat. 21, 101379. [81] De Luca, M., Pappalardo, I., Limongi, A.R., Viviano, E., Radice, R.P., Todisco, S., Martelli, G., Infantino, V., Vassallo, A., 2021. Lipids from microalgae for cosmetic applications. Cosmetics. 8. [82] De Vrieze, J., Verbeeck, K., Pikaar, I., Boere, J., Van , A., Rabaey, K., Verstraete, W., 2020. The hydrogen gas bio-based economy and the production of renewable building block chemicals, food and energy. New Biotechnol. 55, 12-18. [83] Demorest, Z.L., Coffman, A., Baltes, N.J., Stoddard, T.J., Clasen, B.M., Luo, S., Retterath, A., Yabandith, A., Gamo, M.E., Bissen, J., Mathis, L., Voytas, D.F., Zhang, F., 2016. Direct stacking of sequence-specific nuclease-induced mutations to produce high oleic and low linolenic soybean oil. BMC Plant Biol. 16(1), 225. [84] Desroches, M., Caillol, S., Auvergne, R., Boutevin, B., David, G., 2012. Biobased cross-linked polyurethanes obtained from ester/amide pseudo-diols of fatty acid derivatives synthesized by thiol-ene coupling. Polym. Chem. 3, 450-457. [85] Diarte, C., Romero, A., Romero, M.P., Graell, J., Lara, I., 2021. Chemical and sensory characterization of nine spanish monovarietal olive oils: an emphasis on wax esters. Agriculture. 11, 170. [86] Díez-Pascual, A.M., Rahdar, R., 2021. Composites of vegetable oilbased polymers and carbon nanomaterials. Macromol. 1, 276-292. [87] Ding, L.N., Gu, S.L., Zhu, F.G., Ma, Z.Y., Li, J., Li, M., Wang, Z., Tan, X.L., 2020. Long-chain acyl-CoA synthetase 2 is involved in seed oil production in Brassica napus. BMC Plant Biol. 20, 1-14. [88] Ding, L.N., Guo, X.J., Li, M., Fu, Z.L., Yan, S.Z., Zhu, K.M., Wang, Z. et al. (2019) Improving seed germination and oil contents by regulating the GDSL transcriptional level in Brassica napus. Plant Cell Rep. 38, 243-253. [89] Do, P.T., Nguyen, C.X., Bui, H.T., Tran, L.T.N., Stacey, G., Gillman, J.D., Zhang, Z.J., Stacey, M.G., 2019. Demonstration of highly efficient dual gRNA CRISPR/Cas9 editing of the homeologous GmFAD2-1A and GmFAD2-1B genes to yield a high oleic, low linoleic and α-linolenic acid phenotype in soybean. BMC Plant Boil. 19(1), 311. [90] Dogan, M., Akgul, A., 2005. Fatty acid composition of some walnut (Juglans regia L.) cultivars from east Anatolia. Grasas y Aceites 56(4), 328-331. [91] Dong, F., Wang, X., 2017. Effects of carboxymethyl cellulose incorporated with garlic essential oil composite coatings for improving quality of strawberries. Int. J .Biol. Macromol. 104, 821826. [92] Dong, Z., Zhao, H., He, J., Huai, J., Lin, H., Zheng, J., Liu, Y., Wang, G., 2002. Overexpression of a foxtail millet acetyl-CoA carboxylase gene in maize increases sethoxydim resistance and oil content. Afr. J. Biotechnol. 10, 3986-3995. [93] Dyer, J.M., Stymne, S., Green, A.G., Carlsson, A.S., 2008. High-value oils from plants. Plant J. 54, 640-655. [94] Ebrahimnezhad-Khaljiri, H., Ghadi, A., 2023. Recent advancement in synthesizing bio-epoxy nanocomposites using lignin, plant oils, saccharides, polyphenols, and natural rubbers: a review. Int. J. Biol. Macromol. 128041. [95] Eichmann, T.O., Lass, A., 2015. DAG tales: the multiple faces of diacylglycerol - stereochemistry, metabolism, and signaling. Cell Mol. Life Sci. 72, 3931-3952. [96] Elahi, N., Duncan, R.W., Stasolla, C., 20 16. Modification of oil and glucosinolate content in canola seeds with altered expression of Brassica napus LEAFY COTYLEDON1. Plant Physiol. Biochem. 100, 52-63. [97] El-Dalatony, M.M., Sharma, P., Hussein, E.E., Elnaggar, A.Y., Salama, E.S., 2022. Pigand vegetable-cooked waste oils as feedstock for biodiesel, biogas, and biopolymer production. Biomass Convers. Biorefinery. [98] Ellafi, A., Jabbari, H., Tomomewo, O.S., Mann, M.D., Geri, M.B., Tang, C., 2020. Future of hydraulic fracturing application in terms of water management and environmental issues: a critical review. In SPE Canada Unconventional Resources Conference?. D053S11R01. SPE. [99] Elmowafy, M., Musa, A., Alnusaire, T.S., Shalaby, K., Fouda, M.M., Salama, A., Al-Sanea, M.M., Abdelgawad, M.A., Gamal, M., Fouad, S.A., 2021. Olive oil/pluronic oleogels for skin delivery of quercetin: In vitro characterization and ex vivo skin permeability. Polymers. 13, 1808. [100] El-Shenawy, N.S., Abu Zaid, A., AlHarbi, M.S., Al-Thomali, A.W., 2014. Effects of mustard oil on oxidative stress parameters of male mice Nahla. Adv. Food Sci. 36, 78-85. [101] Ettaki, H., Troncoso-Ponce, M.A., To, A., Barthole, G., Lepiniec, L., Baud, S., 2018. Overexpression of MYB115, AAD2, or AAD3 in Arabidopsis thaliana seeds yields contrasting omega-7 contents. PloS One. 13(1), e0192156. [102] Fallahi, A., Hajinajaf, N., Tavakoli, O., Mehrabadi, A., 2021. Effects of simultaneous CO2 addition and biomass recycling on growth characteristics of microalgal mixed culture. J. Chem. Technol. Biotechnol. 96, 3398-407. [103] Fan, J., Yan, C., Zhang, X., Xu, C., 2013. Dual role for phospholipid: Diacylglycerol acyltransferase: Enhancing fatty acid synthesis and diverting fatty acids from membrane lipids to triacylglycerol in Arabidopsis leaves. Plant Cell. 25(9), 3506-3518. [104] Fang, W., Wang, Z., Cui, R., Li, J., Li, Y., 2012. Maternal control of seed size by EOD3/CYP78A6 in Arabidopsis thaliana. Plant J. 70, 929-939. [105] FAO, 2011. The state of the world’s land and water resources: managing systems at risk. [106] FAO, 2023. Agricultural production statistics 2000–2022. FAOSTAT Analytical Briefs, No. 79. Rome. [107] Fatihi, A., Zbierzak, A.M., Dörmann, P., 2013. Alterations in seed development gene expression affect size and oil content of Arabidopsis seeds. Plant Physiol. 163(2), 973-985. [108] Fayyazbakhsh, A., Bell, M.L., Zhu, X., Mei, X., Koutný, M., Hajinajaf, N., Zhang, Y., 2022. Engine emissions with air pollutants and greenhouse gases and their control technologies. J. Clean. Prod. 376, 134260. [109] Fell, D. A., Taylor, D. C., Weselake, R. J., Harwood, J. L., 2023. Metabolic Control Analysis of triacylglycerol accumulation in oilseed rape. Bio Systems. 227-228, 104905. [110] Fernández, M.A., Roque, L.B., Espinosa, E.G., Deyá, C., Bellotti, N., 2020. Organo-montmorillonite with biogenic compounds to be applied in antifungal coatings. Appl. Clay Sci. 184, 105369. [111] Fernando, W.L.R., Sarmilan, N., Wickramasinghe, K.C., Herath, H., Perera, G.I.P., 2020. Experimental investigation of minimum quantity lubrication (MQL) of coconut oil based Metal Working Fluid. Mater. Today Proceed. 23, 23-26. [112] Freitas-Silva, J., de Oliveira, B.F.R., Dias, G.R., de Carvalho, M.M., Laport, M.S., 2023. Unravelling the sponge microbiome as a promising source of biosurfactants. Crit. Rev. Microbiol.49(1), 101106. [113] Gan, L., Park, K., Chai, J., Updike, E. M., Kim, H., Voshall, A., Behera, S., Yu, X. H., Cai, Y., Zhang, C., Wilson, M. A., Mower, J. P., Moriyama, E. N., Zhang, C., Kaewsuwan, S., Liu, Q., Shanklin, J., Cahoon, E. B., 2022. Divergent evolution of extreme production of variant plant monounsaturated fatty acids. PNAS. 119(30), e2201160119. [114] Ganewatta, M.S., Wang, Z., Tang, C., 2021. Chemical syntheses of bioinspired and biomimetic polymers toward biobased materials. Nat. Rev. Chem. 5, 753-772. [115] Garrison, T.F., Murawski, A., Quirino, R.L., 2016. Bio-based polymers with potential for biodegradability. Polymers. 8, 1-22. [116] Gaur, V.K., Sharma, P., Sirohi, R., Varjani, S., Taherzadeh, M.J,, Chang, J.S., Yong, Ng. H., Wong, J.W.C., Kim, S.H., 2022. Production of biosurfactants from agro-industrial waste and waste cooking oil in a circular bioeconomy: an overview. Bioresour. Technol. 343, 126059. [117] Ge, L., Yu, J., Wang, H., Luth, D., Bai, G., Wang, K., Chen, R., 2016. Increasing seed size and quality by manipulating BIG SEEDS1 in legume species. Proc. Natl. Acad. Sci. USA. 113, 12414-12419. [118] Ge, Y., Dong, X., Liu, Y., Yang, Y., Zhan, R., 2021. Molecular and biochemical analyses of avocado (Persea americana) reveal differences in the oil accumulation pattern between the mesocarp and seed during the fruit developmental period. Sci. Hortic. 276, 109717. Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2137 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. [119] Gemsprim, M.S., Babu, N., and Udhayakumar, S., 2021. Tribological evaluation of vegetable oil-based lubricant blends. Mat. Today: Proc. 37, 2660-65. [120] Getz G.S., 2018. Lipid transfer proteins: introduction to the thematic review series. J. Lipid Res. 59(5), 745-748. [121] Ghatge, D., Ramanujam, R.,. 2023. Sustainable machining: a review. Materialtoday Proc. [122] Giannakas, A., Patsaoura, A., Barkoula, N.M,, Ladavos, A., 2017. A novel solution blending method for using olive oil and corn oil as plasticizers in chitosan based organoclay nanocomposites. Carbohydr Polym. 157, 550-557. [123] Gogoi, G., Thakur, A.J., Maji, T.K., 2022. Effect of Natural Crosslinker on the Properties of Chicken Feather and Modified Vegetable Oil Based Green Composites. J. Nat. Fibers. 19, 7896-908. [124] Goyal, A., Sharma, V., Upadhyay, N., Gill, S., Sihag, M., 2014. Flax and flaxseed oil: an ancient medicine & modern functional food. J. Food Sci. Technol. 51(9), 1633-1653. [125] Griffin, L., Dean, L., 2017. Nutrient composition of raw, dry-roasted, and skin-on cashew. [126] Guan, M., Chen, H., Xiong, X., Lu, X., Li, X., Huang, F., Guan, C., 2016. A study on triacylglycerol composition and the structure of high-oleic rapeseed oil. Engineering. 2, 258-262. [127] Guo, Z.H., Haslam, R.P., Michaelson, L.V., Yeung, E.C., Lung, S.C., Napier, J.A., Chye, M.L., 2019. The overexpression of rice ACYL‐ COA‐BINDING PROTEIN 2 increases grain size and bran oil content in transgenic rice. TPJ. 100, 1132-1147. [128] Guzha, A., Whitehead, P., Ischebeck, T., Chapman, K.D., 2023. Lipid droplets: packing hydrophobic molecules within the aqueous cytoplasm. Annu. Rev. Plant Biol. 74, 195-223. [129] Hajinajaf, N., Mehrabadi, A., Tavakoli, O., 2021. Practical strategies to improve harvestable biomass energy yield in microalgal culture: a review. Biomass Bioenerg. 145, 105941. [130] Hajinajaf, N., Das, M., Patra, P., Ghosh, A., Varman, A.M., 2022a. Recycling of Multiple Organic Solid Wastes into Biogas via Anaerobic Digestion. In: Fang, Z., Smith Jr., R.L., Xu, L. (eds) Production of Biofuels and Chemicals from Sustainable Recycling of Organic Solid Waste. Biofuels and Biorefineries, vol 11. Springer, Singapore. [131] Hajinajaf, N., Fallahi, A., Rabbani, Y., Tavakoli, O., Sarrafzadeh, M.H., 2022b. Integrated CO2 capture and nutrient removal by microalgae Chlorella vulgaris and optimization using neural network and support vector regression. Waste Biomass Valori. 13, 4749-70. [132] Hajinajaf, N., Rabbani, Y., Mehrabadi, A., Tavakoli, O., 2022c. Experimental and modeling assessment of large-scale cultivation of microalgae Nannochloropsis sp. PTCC 6016 to reach high efficiency lipid extraction. Int. J. Environ. Sci. Technol. 19, 5511-5528. [133] Hajinajaf, N., Fallahi, A., Eustance, E., Sarnaik, A., Askari, A., Najafi, M., Davis, R.W., Rittmann, B.E., Varman A.M. 2024. Managing carbon dioxide mass transfer in photobioreactors for enhancing microalgal biomass productivity. Algal Res. 103506. [134] Hajjari, M., Tabatabaei M., Aghbashlo, M., Ghanavati, G., 2017. A review on the prospects of sustainable biodiesel production: a global scenario with an emphasis on waste-oil biodiesel utilization. Renew. Sus. Eenrg. Rev. 72, 445-64. [135] Halden, R.U., 2010. Plastics and health risks. Annu. Rev. Public Health. 31, 179-194. [136] Hammond, E.G., Duvick, D., Wang, T., Dodo, H., Pittman, R., 1997. Survey of the fatty acid composition of peanut (Arachis hypogaea) germplasm and characterization of their epoxy and eicosenoic acids. J. Amer. Oil Chem. Soc. 74, 1235-1239. [137] Hamnas, A., Unnikrishnan, G., 2023.Bio-lubricants from vegetable oils: Characterization, modifications, applications and challenges– Review. Renew. Sust. Energy. Rev. 182, 113413. [138] Haregu, S., Likna, Y., Tadesse, D., Masi, C., 2023. Recent Development of Biomass Energy as a Sustainable Energy Source to Mitigate Environmental Change. In: Ramanujam, P.K., Parameswaran, B., Bharathiraja, B., Magesh, A. (eds) Bioenergy. Energy, Environment, and Sustainability. 119-138, Springer, Singapore. [139] Harikrishnan, S., George, S.D., Chidangil, S., Unnikrishnnan VK. 2024. Archaeophotonics: applications of laser spectroscopic techniques for the analysis of archaeological samples. Appl. Spectrosc. Rev. 59, 187-223. [140] Hasan, K.M.F., Horváth, P.G., Alpár, T., 2020. Potential natural fiber polymeric nanobiocomposites: a review. Polymers. 12, 1072. [141] Hassan, A.A., Abbas, A., Rasheed, T., Bilal, M., Iqbal, H.M.N., Wang, S., 2019. Development, influencing parameters and interactions of bioplasticizers: An environmentally friendlier alternative to petro industry-based sources. Sci. Total Environ. 682, 394-404. [142] Hatanaka, T., Tomita, Y., Matsuoka, D., Sasayama, D., Fukayama, H., Azuma, T., Soltani Gishini, M. F., Hildebrand, D. 2022. Different acyl-CoA:diacylglycerol acyltransferases vary widely in function, and a targeted amino acid substitution enhances oil accumulation. J. Exp. Bot. 73(9), 3030-3043. [143] Haule, L.V., Nambela, L., 2022. Sustainable application of nanomaterial for finishing of textile material. Green Nanomat. Ind. Appl. 177-206. [144] Haun, W., Coffman, A., Clasen, B..M, Demorest, Z.L., Lowy, A., Ray, E., Retterath, A., Stoddard, T., Juillerat, A., Cedrone, F., Mathis, L., Voytas, D.F., Zhang, F., 2014. Improved soybean oil quality by targeted mutagenesis of the fatty acid desaturase 2 gene family. Plant Biotechnol. J. 12, 934-940. [145] Hayes, D.G., 2021. Oils and their use beyond the food industry. In: Oil Oilseed Process, John Wiley & Sons Ltd.119-48. [146] Hayta, P., Oktav, M., Ateş Duru, Ö., 2022. Evaluation of plant‐based oils for production of offset printing ink. J. Am. Oil Chem. Soc. 99(8), 711-719. [147] He, M., Qin, C. X., Wang, X., Ding, N. Z., 2020. Plant unsaturated fatty acids: biosynthesis and regulation. Front. Plant Sci. 11, 390. [148] Hernández, M.L., Sicardo, M.D., Belaj, A., Martínez-Rivas, J.M., 2021. The oleic/linoleic acid ratio in olive (Olea europaea L.) fruit mesocarp is mainly controlled by OeFAD2-2 and OeFAD2-5 genes together with the different specificity of extraplastidial acyltransferase enzymes. Front. Plant Sci. 12, 653997. [149] Hoang, A.T., Tabatabaei, M., Aghbashlo, M., Carlucci, A.P., Ölçer, A.I., Le, A.T., Ghassemi, A., 2021. Rice bran oil-based biodiesel as a promising renewable fuel alternative to petrodiesel: a review. Renew Sustain. Energy Rev. 135, 110204. [150] Hölzl, G., Dörmann, P., 2021. Alterations of flower fertility, plant size, seed weight, and seed oil content in transgenic Camelina sativa plants overexpressing CYP78A. Ind. Crops Prod. 170, 113794. [151] Hu, Z., Wang, X., Zhan, G., Liu, G., Hua, W., Wang, H., 2009. Unusually large oilbodies are highly correlated with lower oil content in Brassica napus. Plant Cell Rep. 28, 541-549. [152] Ibrahim, A.M.M., Wei, L.I., Mourad, A.H.I., Mohamed, A.E., Abd ElNaby, A.M., Al Soufi, M.S., Ezzat, M.F. Elsheikh, A., 2023. Cooling and lubrication techniques in grinding: a state-of-the-art review, applications, and sustainability assessment. Chinese J. Aeronaut. 36(7), 76-113. [153] Idoko, J.E., Ileke, K.D., 2020. Comparative evaluation of insecticidal properties of essential oils of some selected botanicals as biopesticides against Cowpea bruchid, Callosobruchus maculatus (Fabricius) [Coleoptera: Chrysomelidae]. Bull. Natl. Res. Cent. 44, 17. [154] IEA., 2018. World oil final consumption by sector, 2018. [155] Ike, D.C., Ibezim-Ezeani, M.U., Akaranta, O., 2021. Cashew nutshell liquid and its derivatives in oil field applications: an update. Green Chem. Lett. Rev. 14, 618-631. [156] Insight, G.M., 2023. Emollients Market - By Source (Botanical, Animal, Mineral, Synthetic), By Form (Creams & Lotions, Oils & Serums, Balms & Butters, Sprays), By Function, By End-use & Forecast. 20232032. [157] Isman, M.B., 2020. Commercial development of plant essential oils and their constituents as active ingredients in bioinsecticides. Phytochem Rev. 19, 235-241. [158] Ivankovic, T., Hrenovic, J., 2010. Surfactant in the environment. Arh. Hig. Rada. Toksikol. 61, 95-110. [159] Ivarson, E., Iven, T., Sturtevant, D., Ahlman, A., Cai, Y., Chapman, K., Feussner, I., Zhu, L.H., 2017. Production of wax esters in the wild oil species Lepidium campestre. Ind. Crop. Prod. 108, 535-42. Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2138 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. [160] Iven, T., Hornung, E., Heilmann, M., Feussner, I., 2016. Synthesis of oleyl oleate wax esters in Arabidopsis thaliana and Camelina sativa seed oil. Plant Biotechnol. J. 14(1), 252-259. [161] Iwuagwu, M.O., Solomon, C.U., Amanze, J.E., 2018. Physicochemical analysis and characterization of edible oil from seeds of orange (Citrus sinensis L.) and pumpkin (Cucurbita pepo L.). Eur. J. Biotech. Biosci. 6, 35-40. [162] Janaki, S., Zandi-Sohani, N., Ramezani, L., Szumny, A., 2018. Chemical composition and insecticidal efficacy of Cyperus rotundus essential oil against three stored product pests. Int. Biodeterior. Biodegrad. 133, 93-98. [163] Javaid, S., Saleem, N., Rehman, S.U., 2024. Polymeric nano-emulsion in functional textile finishing. nanoemulsions-design and applications. IntechOpen. [164] Jia, P., Hu, L., Zhang, M., Feng, G., Zhou, Y., 2017. Phosphorus containing castor oil based derivatives: Potential non-migratory flame retardant plasticizer. Eur. Polym. J. 87, 209-220. [165] Jia, P., Xia, H., Tang, K., Zhou, Y., 2018. Plasticizers derived from biomass resources: a short review. Polymers. 10, 1303. [166] Jiang, X., Guan, D., 2017. The global CO2 emissions growth after international crisis and the role of international trade. Energy Policy. 109, 734-746. [167] Joshi, S.J., Al-Farsi, H., Al-Bahry, S., Al-Wahaibi, Y., 2022. “Glycolipid biosurfactant-silica nanoparticles” based green application for enhancement of oil recovery. Pet. Sci. Technol. 40(17), 2064-2081. [168] Jung, S.H., Kim, R.J., Kim, K.J., Lee, D.H., Suh, M.C., 2019. Plastidial and mitochondrial malonyl CoA-ACP malonyltransferase is essential for cell division and its overexpression increases storage oil content. Plant Cell Physiol. 60, 1239-1249. [169] Kalita, D., Tarnavchyk, I., Webster, D.C., Chisholm, B.J., 2022. Synthesis and evaluation of novel plant oil-based polymers as binders for artist paints: controllable drying behavior and low yellowness. Prog. Org. Coatings 163, 106607. [170] Kalscheuer, R., Stolting, T., Steinbuchel, A., 2006. Microdiesel: Escherichia coli engineered for fuel production. Microbiology. 152(9), 2529-2536. [171] Kan, S., Chen, B., Chen, G., 2019. Worldwide energy use across global supply chains: decoupled from economic growth? Appl. Energ. 250, 1235-1245. [172] Kanai, M., Mano, S., Kondo, M., Hayashi, M., Nishimura, M., 2016. Extension of oil biosynthesis during the mid-phase of seed development enhances oil content in Arabidopsis seeds. Plant Biotechnol. J. 14, 1241-1250. [173] Kanai, M., Yamada, T., Hayashi, M., Mano, S., Nishimura, M., 2019. Soybean (Glycine max L.) triacylglycerol lipase GmSDP1 regulates the quality and quantity of seed oil. Sci. Rep.9, 8924. [174] Kapoor, B., Kapoor, D., Gautam, S., Singh, R. and Bhardwaj, S., 2021. Dietary polyunsaturated fatty acids (PUFAs): Uses and potential health benefits. Curr. Nutr. Rep. 10, 232-42. [175] Karunarathna, N.L., Wang, H., Harloff, H.J., Jiang, L., Jung, C., 2020. Elevating seed oil content in a polyploid crop by induced mutations in SEED FATTY ACID REDUCER genes. Plant Biotechnol. J. 18, 2251-2266. [176] Kavetsou, E., Koutsoukos, S., Daferera, D., Polissiou, M.G., Karagiannis, D., Perdikis, D.C, Detsi, A., 2019. Encapsulation of Mentha pulegium essential oil in yeast cell microcarriers: an approach to environmentally friendly pesticides. J. Agric. Food Chem. 67, 47464753. [177] Kawall, K., 2021. Genome-edited Camelina sativa with a unique fatty acid content and its potential impact on ecosystems. Environmen. Sci. Eur. 33, 38. [178] Kazeem, R.A., Fadare, D.A., Ikumapayi, O.M., Adediran, A.A., Aliyu, S.J., Akinlabi, S.A., Jen, T.C., Akinlabi, E.T., 2022. Advances in the application of vegetable-oil-based cutting fluids to sustainable machining operations-a review. Lubricants 10, 69. [179] Khan, K., Kumar, V., Niranjan, A., Shanware, A., Sane, V.A., 2019. JcMYB1, a Jatropha R2R3MYB transcription factor gene, modulates lipid biosynthesis in transgenic plants. Plant Cell Physiol. 60, 462-475. [180] Khan, M.A.A., Hussain, M., Lodhi, S.K., Zazoum, B., Asad, M., Afzal, A., 2022. Green metalworking fluids for sustainable machining operations and other sustainable systems: a review. Metals. 12, 1466. [181] Kim H. U., 2020. Lipid metabolism in plants. Plants (Basel, Switzerland), 9(7), 871. [182] Kim, H.U., Lee, K.R., Go, Y.S., Jung, J.H., Suh, M.C., Kim, J.B., 2011. Endoplasmic reticulum-located PDAT1-2 from castor bean enhances hydroxy fatty acid accumulation in transgenic plants. Plant Cell Physiol. 52(6), 983-993. [183] Kim, H.U., Lee, K.R., Jung, S.J., Shin, H.A., Go, Y.S., Suh, M.C., Kim, J.B., 2015. Senescence inducible LEC2 enhances triacylglycerol accumulation in leaves without negatively affecting plant growth. Plant Biotechnol. J. 13, 1346-1359. [184] Kim, R.J., Kim, H.U., Suh, M.C., 2019. Development of Camelina enhanced with drought stress resistance and seed oil production by cooverexpression of MYB96A and DGAT1C. Ind.Crops Prod. 138, 111475. [185] Klangmuang, P., Sothornvit, R., 2018. Active coating from hydroxypropyl methylcellulose-based nanocomposite incorporated with Thai essential oils on mango (cv. Namdokmai Sithong). Food Biosci. 23, 9-15. [186] Klein, M.L., Chastain, T.G., Garbacik, C.J., Qian, Y.P.L., Mc Donnell, R.J., 2020. Acute toxicity of essential oils to the pest slug Deroceras reticulatum in laboratory and greenhouse bioassays. J. Pest Sci. 93,415-425. [187] Kong, Q., Yang, Y., Guo, L., Yuan, L., Ma, W., 2020. Molecular basis of plant oil biosynthesis: insights gained from studying the WRINKLED1 transcription factor. Front. Plant Sci.11, 24. [188] Krishna, P.V., Srikant, R.R., Rao, D.N., 2010. Experimental investigation on the performance of nanoboric acid suspensions in SAE-40 and coconut oil during turning of AISI 1040 steel. Int. J. Mach. Tools Manuf. 50, 911-916. [189] Kumar, A., Sharma, A., Upadhyaya, K.C., 2016. Vegetable oil: nutritional and industrial perspective. Cur. Genom. 17(3), 230-240. [190] Kumar, N., Chaudhary, A., Singh, D., Teotia, S., 2020. Transcriptional regulation of seed oil accumulation in Arabidopsis thaliana: role of transcription factors and chromatin remodelers. J. Plant Biochem. Biotechnol. 29, 754-68. [191] Kumar, S., 2019. Recent developments of biobased plasticizers and their effect on mechanical and thermal properties of poly (vinyl chloride): a review. Ind. Eng. Chem. Res. 58, 11659-72. [192] Lamch, Ł., Witek, K., Jarek, E., Obłąk, E., Warszyński, P., Wilk, K.A., 2020. New mild amphoteric sulfohydroxybetaine-type surfactants containing different labile spacers: Synthesis, surface properties and performance. J. Colloid Interface Sci. 558, 220-229. [193] Langsdorf, A., Volkmar, M., Holtmann, D., Ulber, R., 2021. Material utilization of green waste: a review on potential valorization methods. Bioresour. Bioprocessing 8, 1-26. [194] Latha, R.B., Nasirullah, D.R., 2014. Physico-chemical changes in rice bran oil during heating at frying temperature. J. Food Sci. Technol. 51, 335-340. [195] Lee, E.J., Oh, M., Hwang, J.U., Li-Beisson, Y., Nishida, I., Lee, Y., 2017. Seed-specific overexpression of the pyruvate transporter BASS2 increases oil content in Arabidopsis seeds. Front. Plant Sci. 8, 241057. [196] Lee, H.G., Park, M.E., Park, B.Y., Kim, H.U., Seo, P. J., 2019a. The Arabidopsis MYB96 transcription factor mediates ABA-dependent triacylglycerol accumulation in vegetative tissues under drought stress conditions. Plants (Basel, Switzerland), 8(9), 296. [197] Lee, J.Y., Garcia, C.V., Shin, G.H., Kim, J.T., 2019b. Antibacterial and antioxidant properties of hydroxypropyl methylcellulose-based active composite films incorporating oregano essential oil nanoemulsions. Lwt 106,164-171. [198] Lee, K.R., Jeon, I., Yu, H., Kim, S.G., Kim, H.S., Ahn, S.J., Lee,, J., Lee, S.K., Kim, H.U., 2021. Increasing monounsaturated fatty acid contents in hexaploid Camelina sativa seed oil by FAD2 gene knockout using CRISPR-Cas9. Front. Plant Sci. 29, 12:702930. [199] Lee, S.B., Kim, H., Kim, R.J., Suh, M.C., 2014. Overexpression of Arabidopsis MYB96 confers drought resistance in Camelina sativa via cuticular wax accumulation. Plant Cell Rep. 33, 1535-1546. [200] Leone, A., Spada, A., Battezzati, A., Schiraldi, A., Aristil, J., Bertoli, S., 2016. Moringa oleifera seeds and oil: Characteristics and uses for human health. Int. J. Mol. Sci.17, 1-14. Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2139 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. [201] Li, Y., Zheng, L., Corke, F., Smith, C. and Bevan, M.W., 2008. Control of final seed and organ size by the DA1 gene family in Arabidopsis thaliana. Genes Dev. 22(10), 1331-1336. [202] Li, C., Li, L., Lian, J., Watts, R., Nelson, R., Goodwin, B., Lehner, R., 2015a. Roles of Acyl-CoA: Diacylglycerol acyltransferases 1 and 2 in triacylglycerol synthesis and secretion in primary hepatocytes. Arterioscler Thromb Vasc. Biol. 35, 1080-1091. [203] Li, M., Wei, F., Tawfall, A., Tang, M., Saettele, A., Wang, X., 2015b. Overexpression of patatin-related phospholipase AIIId altered plant growth and increased seed oil content in Camelina. Plant Biotechnol. J. 13, 766-778. [204] Li, D., Jin, C., Duan, S., Zhu, Y., Qi, S., Liu, K., Gao, C., Ma, H., Zhang, M., Liao, Y., Chaen, M., 2017. MYB89 transcription factor represses seed oil accumulation. Plant Physiol. 173, 1211-1225. [205] Li, H., Zhou, R., Liu, P., Yang, M., Xin, D., Liu, C., Zhang, Z., Wu, X., Chen, Q., Zhao, Y., 2023. Design of high-monounsaturated fatty acid soybean seed oil using GmPDCTs knockout via a CRISPR-Cas9 system. Plant Biotechnol. J. 21(7), 1317-1319. [206] Li, N., Meng, H., Li, S., Zhang, Z., Zhao, X., Wang, S., Liu, A., Li, Q., Song, Q., Li, X., 2020a. Two plastid fatty acid exporters contribute to seed oil accumulation in Arabidopsis. Plant Physiol. 182(4), 19101919. [207] Li, Y., Kong, D., Fu, Y., Sussman, M.R., Wu, H., 2020b. The effect of developmental and environmental factors on secondary metabolites in medicinal plants. Plant Physiol. Biochem.148, 80-89. [208] Li, R., Yu, K., Wu, Y., Tateno, M., Hatanaka, T., Hildebrand, D.F., 2012a. Vernonia DGATs can complement the disrupted oil and protein metabolism in epoxygenase-expressing soybean seeds. Metabol. Engin. 14(1), 29-38. [209] Li, X., van Loo, E.N., Gruber, J., Fan, J., Guan, R., Frentzen, M., Stymne, S., Zhu, L.H., 2012b. Development of ultra-high erucic acid oil in the industrial oil crop Crambe abyssinica. Plant Biotechnol. J. 10(7), 862-870. [210] Li, X., Guan, R., Fan, J., Zhu, L.H., 2019. Development of industrial oil crop Crambe abyssinica for wax ester production through metabolic engineering and cross breeding. Plant Cell Physiol. 60(6), 1274-1283. [211] Li-Beisson, Y., Nakamura, Y. and Harwood, J., 2016. Lipids: from chemical structures, biosynthesis, and analyses to industrial applications. Lipids in plant and algae development pringer, Cham., 118. [212] Lim, A.R.Q., Kong, Q., Singh, S.K., Guo, L., Yuan, L., Ma, W., 2022. Sunflower WRINKLED1 plays a key role in transcriptional regulation of oil biosynthesis. Int. J. Mol. Sci. 23, 3054. [213] Liu, B., Sun, Y., Wang, X., Xue, J., Wang, J., Jia, X., Li, R., 2022a. Identification and functional characterization of Acyl-ACP thioesterases B (GhFatBs) responsible for palmitic acid accumulation in cotton seeds. Int. J. Mol. Sci, 23(21), 12805. [214] Liu, Y., Han, J., Li, Z., Jiang, Z., Luo, L., Zhang, Y., Chen, M., Yang, Y., Liu, Z., 2022b. Heterologous expression of Jatropha curcas fatty acyl-ACP thioesterase A (JcFATA) and B (JcFATB) affects fatty acid accumulation and promotes plant growth and development in Arabidopsis. Int. J. Mol. Sci. 23: 4209. [215] Liu, W., Qiu, J., Chen, T., Fei, M., Qiu, R., Sakai, E., 2019. Regulating tannic acid-crosslinked epoxidized soybean oil oligomers for strengthening and toughening bamboo fibers-reinforced poly (lactic acid) biocomposites. Compos. Sci. Technol. 181, 107709. [216] Liu, W.X., Liu, H.L., Qu, L. Q., 2013. Embryo-specific expression of soybean oleosin altered oil body morphogenesis and increased lipid content in transgenic rice seeds. Theor. Appl. Genet. 126, 2289-2297. [217] Lu, C., Fulda, M., Wallis, J.G., Browse, J., 2006. A high-throughput screen for genes from castor that boost hydroxy fatty acid accumulation in seed oils of transgenic Arabidopsis. The Plant J. Cell and Mol. Biol, 45(5), 847-856. [218] Lu, Q., Zhang, Z.S., Zhan, R.T., He, R., 2018. Proteomic analysis of Zanthoxylum nitidum seeds dormancy release: Influence of stratification and gibberellin. Ind. Crops Prod. 122, 7-15. [219] Lunn, D., Wallis, J. G., Browse, J., 2018. Overexpression of Seipin1 increases oil in hydroxy fatty acid-accumulating seeds. Plant Cell Physiol. 59, 205-214. [220] Luo, G., Cao, V.D., Kannan, B., Liu, H., Shanklin J., Altpeter, F. 2022. Metabolic engineering of energycane to hyperaccumulate lipids in vegetative biomass. BMC Biotechnol. 22, 24 (2022). [221] Lutkewitte, A.J. and Finck, B.N., 2020. Regulation of signaling and metabolism by lipin-mediated phosphatidic acid phosphohydrolase activity. Biomol.10, 1386. [222] Machado, M. Rodriguez-Alcalá, L.M., Gomes, A.M., Pintado, M., 2023. Vegetable oils oxidation: mechanisms, consequences and protective strategies. Food Rev. Int. 39, 4180-97. [223] Mahesh, K., Philip, J.T., Joshi, S., Kuriachen, B., 2021. Machinability of Inconel 718: A critical review on the impact of cutting temperatures. Mater. Manuf. Proces. 36(7), 753-791. [224] Majdinasab, M., Niakousari, M., Shaghaghian, S., Dehghani, H., 2020. Antimicrobial and antioxidant coating based on basil seed gum incorporated with Shirazi thyme and summer savory essential oils emulsions for shelf-life extension of refrigerated chicken fillets. Food Hydrocoll. 108,106011. [225] Manan, S., Ahmad, M.Z., Zhang, G., Chen, B., Haq, B.U., Yang, J., Zhao, J., 2017. Soybean LEC2 regulates subsets of genes involved in controlling the biosynthesis and catabolism of seed storage substances and seed development. Front. Plant Sci. 8, 1-16. [226] Mancini, A., Imperlini, E., Nigro, E., Montagnese, C., Daniele, A., Orrù, S., Buono, P., 2015. Biological and nutritional properties of palm oil and palmitic acid: Effects on health. Molecules. 20, 1733917361. [227] Mangeon, C., Michely, L., Rios de Anda, A., Thevenieau, F., Renard, E., Langlois, V., 2018. Natural terpenes used as plasticizers for poly (3-hydroxybutyrate). ACS Sustain Chem. Eng. 6, 16160-16168. . [228] Mannu, A., Garroni, S., Ibanez Porras, J., Mele, A., 2020. Available technologies and materials for waste cooking oil recycling. Processes. 8(3), 366. [229] Maraschin, F. dos S., Kulcheski, F.R., Segatto, A.L.A., Trenz, T.S., Barrientos-Diaz, O., Margis-Pinheiro, M., Margis, R., TurchettoZolet, A.C., 2019. Enzymes of glycerol-3-phosphate pathway in triacylglycerol synthesis in plants: Function, biotechnological application and evolution. Prog. Lipid Res. 73, 46-64. [230] Mariani, C., Lucci, P., Conte, L., 2018. Identification of phytyl vaccinate as a major component of wax ester fraction of extra virgin olive oil. Eur. J. Lipid Sci. Technol. 120, 1800154. [231] McKeon, T.A., Hayes D.G., Hildebrand, D.F., Weselake, R.J., 2016. Chapter 1 - Introduction to Industrial Oil Crops. In: McKeon TA, Hayes DG, Hildebrand DF, Weselake RJ (eds) Industrial Oil Crops. AOCS Press, pp 1-13. [232] Metzger, J.O., Hüttermann, A., 2009. Sustainable global energy supply based on lignocellulosic biomass from afforestation of degraded areas. Naturwissenschaften. 96, 279-288. [233] Miklaszewska, M., Zienkiewicz, K., Inchana, P., Zienkiewicz, A., 2021. Lipid metabolism and accumulation in oilseed crops. OCL, 28, p.50. [234] Mittal, R.P., Rana, A., Jaitak, V., 2019. Essential oils: an impending substitute of synthetic antimicrobial agents to overcome antimicrobial resistance. Curr. Drug Targets 20, 605-624. [235] Mohammed, S., and Ikiensikimama, S.S., 2023. Vegetable oils as surfactant feedstocks for enhanced oil recovery: a review. Chem. Eng. Res. Des. 200, 693-705. [236] Mohanty, S.S., Koul, Y., Varjani, S., Pandey, A., Ngo, H.H., Chang, J.S., Wong, J.W.C., Bui, X.T., 2021. A critical review on various feedstocks as sustainable substrates for biosurfactants production: a way towards cleaner production. Microb. Cell Fac. 20, 120. [237] Montava-Jordà, S., Quiles-Carrillo, L., Richart, N., Torres-Giner, S., Montanes, N., 2019. Enhanced interfacial adhesion of polylactide/poly (ε-caprolactone)/walnut shell flour composites by reactive extrusion with maleinized linseed oil. Polymers. 11, 758. [238] Moreno-Camacho, C.A., Montoya-Torres, J.R., Jaegler, A., Gondran, N., 2019. Sustainability metrics for real case applications of the supply chain network design problem: A systematic literature review. J. Clean. Prod. 231, 600-618. [239] Msanne, J., Kim, H., Cahoon, E.B., 2020. Biotechnology tools and applications for development of oilseed crops with healthy vegetable oils. Biochimie. 178, 4-14. Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2140 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. [240] Mujtaba, M.A., Cho, H.M., Masjuki, H.H., Kalam, M.A., Ong, H.C., Gul, M., Harith, M.H., Yusoff, H.N.M.A., 2020. Critical review on sesame seed oil and its methyl ester on cold flow and oxidation stability. Energy. Rep. 6, 40-54. [241] Mulgund, A., 2022. Increasing lipid accumulation in microalgae through environmental manipulation, metabolic and genetic engineering: a review in the energy NEXUS framework. Energy Nexus. 5, 100054. [242] Muñoz, C.F., Südfeld, C., Naduthodi, M.I., Weusthuis, R.A., Barbosa, M.J., Wijffels, R.H., D’Adamo, S., 2021. Genetic engineering of microalgae for enhanced lipid production. Biotechnol. Adv. 52, 107836. [243] Muralidhar, V., Chaganti, P.K., 2020. A review on testing methods of metalworking fluids for environmental health. Mater. Today Proc. 26, 2405-2411. [244] Na, G.N., Aryal, N., Fatihi, A., Kang, J., Lu, C., 2018. Seed-specific suppression of ADP-glucose pyrophosphorylase in Camelina sativa increases seed size and weight. Biotechnol. Biofuels. 11, 330. . [245] Nachtergaele, F., van Velthuizen, H., Verelst, L.), Wiberg, D., Henry, M., Chiozza, F., Yigini, Y., Aksoy, E., Batjes, N., Boateng, E., Fisher, G., Jones, A., Montanarella, L., Shi, X., Tramberend, S., 2023. Harmonized World Soil Database version 2.0. FAO. [246] Nagtode V.S., Cardoza, C., Yasin, H.K.A., Mali, S.N., Tambe, S.M., Roy, P., Singh, K., Goel, A., Amin, P.D., Thorat, B.R., 2023. Green surfactants (biosurfactants): a petroleum-free substitute for Sustainability-Comparison, applications, market, and future prospects. ACS omega. 8, 11674-99. [247] Naik, B., Kumar, V., 2014. Cocoa butter and its alternatives: a Review. J. Bioresour. Engin. Technol. 1, 7-17. [248] Napier, J.A., 2007. The production of unusual fatty acids in transgenic plants. Annu. Rev/ Plant Biol. 58, 295-319. [249] Napier, J.A., Haslam, R.P., Beaudoin, F., Cahoon, E.B., 2014. Understanding and manipulating plant lipid composition: Metabolic engineering leads the way. Curr. Opin. Plant Biol. 19, 68-75. [250] Natarajan, G., Rajan, T.P., Das, S., 2022. Application of sustainable textile finishing using natural biomolecules. J. Nat. Fibers 19, 43504367. [251] Neves, J.S., Valadares L.F., Machado, F., 2018. Tailoring acrylated soybean oil-containing terpolymers through emulsion polymerization. Colloids Interfaces. 2, 46. [252] Nguyen, H.T., Park, H., Koster, K.L., Cahoon, R.E., Nguyen, H.T., Shanklin, J., Clemente, T.E., Cahoon, E.B., 2015. Redirection of metabolic flux for high levels of omega-7 monounsaturated fatty acid accumulation in Camelina seeds. Plant Biotechnol. J. 13(1), 38-50. [253] Niu, Y., Wu, L., Li, Y., Huang, H., Qian, M., Sun, W., Zhu, H., Xu, Y., Fan, Y., Mahmood, U., 2020. Deciphering the transcriptional regulatory networks that control size, color, and oil content in Brassica rapa seeds. Biotechnol.Biofuels 13, 1-20. [254] Njuguna, J. , Siddique, S., Kwroffie, L.B., Piromrat, S., AddaeAfoakwa, K., Ekeh-Adegbotolu, U., Oluyemi, G., Yates, K., Mishra, A.K., Moller, L., 2022. The fate of waste drilling fluids from oil & gas industry activities in the exploration and production operations. Waste Manage. 139, 362-80. [255] Nomanbhay, S., Hussein, R., Ong, M.Y., 2018. Sustainability of biodiesel production in Malaysia by production of bio-oil from crude glycerol using microwave pyrolysis: A review. Green Chem. Lett. Rev. 11, 135-157. [256] Nookaraju, A., Pandey, S.K., Fujino, T., Kim, J.Y., Suh, M.C., Joshi, C.P., 2014. Enhanced accumulation of fatty acids and triacylglycerols in transgenic tobacco stems for enhanced bioenergy production. Plant Cell Rep. 33, 1041-1052 . [257] OECD, 2012. Economic, environmental and social statistics. [258] Okuzaki, A., Ogawa, T., Koizuka, C., Kaneko, K., Inaba, M., Imamura, J., Koizuka, N., 2018. CRISPR/Cas9-mediated genome editing of the fatty acid desaturase 2 gene in Brassica napus. Plant Physiol. Biochem. PPB. 131, 63-69. [259] Orue, A., Eceiza, A., Arbelaiz, A., 2018. Preparation and characterization of poly (lactic acid) plasticized with vegetable oils and reinforced with sisal fibers. Ind. Crops Prod. 112, 170-180. [260] Owuna, F.J., 2020. Stability of vegetable based oils used in the formulation of ecofriendly lubricants–a review. Egypt. J. Pet. 29, 25156. [261] Papadimitriou, D.M., Petrakis, E.A., Arvaniti, K.A., Kimbaris, A.C., Polissiou, M.G., Perdikis, D.C., 2019. Comparative bioactivity of essential oils from two Mentha pulegium (Lamiaceae) chemotypes against Aphis gossypii, Aphis spiraecola, Tetranychus urticae and the generalist predator Nesidiocoris tenuis. Phytoparasitica. 47, 683-692. [262] Park, K., Sanjaya, S., Quach, T., Cahoon, E., 2021. Toward sustainable production of value-added bioenergy and industrial oils in oilseed and biomass feedstocks. Glob. Change Biol. Bioenrg. 13, 1610-1623. [263] Park, M.E., Lee, K.R., Chen, G.Q., Kim, H.U., 2022. Enhanced production of hydroxy fatty acids in Arabidopsis seed through modification of multiple gene expression. Biotechnol. Biofuel Bioprod. 15(1), 66. [264] Patole, P., Kulkarni, V., 2018. Parametric optimization of minimum quantity lubrication in turning of AISI 4340 using nano fluids. Materials Today: Proceedings 5(5), 12419-12425. [265] Pattnaik, M., Mishra, H.N. 2022. Amelioration of the stability of polyunsaturated fatty acids and bioactive enriched vegetable oil: blending, encapsulation, and its application. Crit. Rev. Food Sci. Nut. 62, 6253-76. [266] Pavlačková, J., Egner, P., Slavík, R., Mokrejš, P., Gál, R., 2020. Hydration and barrier potential of cosmetic matrices with bee products. Molecules. 25, 2510. . [267] Peivasteh-Roudsari, L., Barzegar-Bafrouei, R., Aghbolagh Sharifi, K., Azimisalim, S., Karami, M., Abedinzadeh, S., Asadinezhad, S., Tajdar-Oranj, B., Mahdavi, V., Mirza Alizadeh, A., 2023. Origin, dietary exposure, and toxicity of endocrine-disrupting food chemical contaminants: a comprehensive review. Heliyon. 9(7), e18140. [268] Periyasamy, A.P., 2023. Microfiber emissions from functionalized textiles: potential threat for human health and environmental risks. Toxics. 11, 406. [269] Petrie, J.R., Shrestha, P., Belide, S., Mansour, M.P., Liu, Q., Horne, Nichos, P.D., Singh, S.P., 2012. Transgenic production of arachidonic acid in oilseeds. Transgenic Res. 21, 139-147. [270] Phulpoto, A.H., Maitlo, M.A., Kanhar, N.A., 2021. Culture-dependent to culture-independent approaches for the bioremediation of paints: a review. Int. J. Environ. Sci. Technol. 18, 241-62. [271] Piccini, M., Lightfoot, J., Dominguez B.C., Buchard, A., 2021. Xylose-based polyethers and polyesters via ADMET polymerization toward polyethylene-like materials. ACS Appl. Polym. Mater. 3, 5870-5881. [272] Pinheiro, C.T., Quina, M.J.,, Gando-Ferreira L.M., 2021. Management of waste lubricant oil in Europe: a circular economy approach. Crit. Rev. Environ. Sci. Technol. 51, 2015-2050. [273] Piovesana, S., Aita, S.E., Cannazza, G., Capriotti, A.L., Cavaliere, C., Cerrato, A., Guarnaccia, P., Montone, C.M., Laganà, A., 2021. Indepth cannabis fatty acid profiling by ultra-high performance liquid chromatography coupled to high resolution mass spectrometry. Talanta. 228, 122249. [274] Pizzimenti, S., Bernazzani, L., Tine, M.R., Treil, V., Duce, C., Bonaduce, I., 2021. Oxidation and cross-linking in the curing of airdrying artists’ oil paints. ACS Appl. Polym. Mat. 3, 1912-22. [275] Polaris Market Research, 2024.Composites Market Share, Size, Trends, Industry Analysis Report, By Fiber Type (Glass Fiber, Carbon Fiber, Others); By Resin Type; By Manufacturing Process; By End Use; By Region; Segment Forecast, 20242032. Report ID: PM4827. [276] Polarz, S., Kunkel, M., Donner, A., Schlötter, M., 2018. Added‐value surfactants. Chem.–A Europ. J. 24(71), 18842-18856. [277] Poljšak, N., Kreft, S., Kočevar Glavač, N., 2020. Vegetable butters and oils in skin wound healing: Scientific evidence for new opportunities in dermatology. Phytother. Res. 34, 254-269. [278] Pollard, M., Shachar-Hill, Y., 2022. Kinetic complexities of triacylglycerol accumulation in developing embryos from Camelina sativa provide evidence for multiple biosynthetic systems. J. Biol. Chem. 298(1), 101396. [279] Porokhovinova, E.A., Matveeva, T.V., Khafizova, G.V., Bemova, V.D., Doubovskaya, A.G., Kishlyan, N.V., Podolnaya, L.P., Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2141 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Gavrilova, V.A., 2022. Fatty acid composition of oil crops: genetics and genetic engineering. Genet. Resour. Crop Evol. 69, 2029-45. [280] Pouvreau, B., Baud, S., Vernoud, V., Morin, V., Py, C., Gendrot, G., Pichon, J.P., Rouster, J., Paul, W., Rogowsky, P.M., 2011. Duplicate maize Wrinkled1 transcription factors activate target genes involved in seed oil biosynthesis. Plant Physiol. 156(2), 674-686. [281] Prescient and Strategic Intelligence (2023) Textile Finishing Chemical Market. https://www.psmarketresearch.com/market-analysis/textilefinishing-chemicals-market. [282] Price, A.M., Doner, N.M., Gidda, S.K., Jambunathan, S., James, C.N., Schami, A., Yurchenko, O., Mullen, R.T., Dyer, J.M., Puri, V., Chapman, K.D., 2020. Mouse fat-specific protein 27 (FSP27) expressed in plant cells localizes to lipid droplets and promotes lipid droplet accumulation and fusion. Biochimie. 169, 41-53. [283] Qi, W., Lu, H., Zhang, Y., Cheng, J., Huang, B., Lu, X., Sheteiwy, M.S.A., Kuang, S., Shao, H., 2020. Oil crop genetic modification for producing added value lipids. Crit. Rev. Biotechnol. 40, 777-786. [284] Qin, Y., Li, W., Liu, D., Yuan, M., Li, L., 2017 Development of active packaging film made from poly (lactic acid) incorporated essential oil. Prog. Org. Coatings. 103, 76-82. [285] Quirino, R.L., Monroe, K., Fleischer, C.H., Biswas, E., Kessler, M.R., 2021. Thermosetting polymers from renewable sources. Polym. Int. 70, 167-180. [286] Quispe, C.A.G., Coronado, C.J.R., Carvalho, J.A., 2013. Glycerol: Production, consumption, prices, characterization and new trends in combustion. Renew. Sustain. Energy Rev. 27, 475-493. [287] Rajaeifar, M.A., Hemayati, S.S., Tabatabaei, M., Aghbashlo, M., Mahmoudi, S.B., 2019. A review on beet sugar industry with a focus on implementation of waste-to-energy strategy for power supply. Renew. Sustain. Energy Rev. 103, 423-442. [288] Rajput, C.V., Sastry, N.V., Chikhaliya, N.P., 2023. Vegetable oils based precursors: modifications and scope for futuristic bio-based polymeric materials. J. Polym. Res. 30, 159. [289] Ranjbar, S., Malcata, F.X., 2022. Challenges and prospects for sustainable microalga-based oil: A comprehensive review, with a focus on metabolic and genetic engineering. Fuel 324, 124567. [290] Rathnakumar, A.L., Sujatha, M., 2022. Breeding major oilseed crops: Prospects and future research needs, in: Accelerated Plant Breeding, Volume 4: Oil Crops. 1-40. [291] Rathour, R.K., Behl, B., Dhashmana, K., Sakhuja, D., Ghai, H., Sharma, N., Meena, K.R., Bhatt, A.K. Bhatia, R.K., 2023. Non-food crops derived lignocellulose biorefinery fo r sustainable production of biomaterials, biochemicals and bioenergy: a review on trends and techniques. Ind. Crop Prod. 204, 117220. [292] Rauf, S., Fatima, S., Ortiz, R., 2023. Modification of Fatty Acid Profile and Oil Contents Using Gene Editing in Oilseed Crops for a Changing Climate. GM crops Food. 14(1), 1-12. [293] Reazai, M., Mohammadpourfard, I., Nazmara, S., Jahanbakhsh, M., Shiri, L., 2014. Physicochemical characteristics of citrus seed oils from Kerman, Iran. J Lipids. 2014, 1-3. [294] Rehan, Z.A., Usman, A., 2023. Polymeric Paints and Coatings.' in, Advanced Functional Polymers: Synthesis to Applications (Springer). [295] Rej, S., Bandyopadhyay, A., Mahmood, H., Murshed, M., Mahmud, S., 2022. The role of liquefied petroleum gas in decarbonizing India: fresh evidence from wavelet–partial wavelet coherence approach. Environ. Sci. Pollut. Res. 29, 35862-35883. [296] Rekkab, S., Zarrok, H., Salghi, R., Zarrouk, A., Bazzi, L., Hammouti, B., Kabouche, Z., Touzani, R., Zougagh, M., 2012. Green corrosion inhibitor from essential oil of Eucalyptus globulus (Myrtaceae) for C38 steel in sulfuric acid solution. J. Mater. Environ. Sci. 3, 613-627. Corpus ID: 201083326. [297] Rezig, L., Chouaibi, M., Meddeb, W., Msaada, K., Hamdi, S., 2019. Chemical composition and bioactive compounds of Cucurbitaceae seeds: Potential sources for new trends of plant oils. Process Saf. Environ. Prot. 127, 73-81. [298] Rivera-Rangel, L., Aguilera-Campos, K., García-Triana, A., AyalaSoto, J., Chavez-Flores, D., Hernández-Ochoa, L., 2018. Comparison of oil content and fatty acids profile of Western Schley, Wichita, and native pecan nuts cultured in Chihuahua, Mexico. J. Lipids. 2018. 4781345. [299] Rocky, M.M.H., Rahman, I.I.M., Biswas, F.B., Rahman, S., Endo, M., Wong, K.H., Mashio, A.S., Hasegawa. H., 2023. Cellulose-based materials for scavenging toxic and precious metals from water and wastewater: a review. Chem. Eng. J. 472, 144677. [300] Roscoe, T.T., Guilleminot, J., Bessoule, J.J., Berger, F., Devic, M., 2015. Complementation of seed maturation phenotypes by ectopic expression of ABSCISIC ACID INSENSITIVE3, FUSCA3 and LEAFY COTYLEDON2 in Arabidopsis. Plant Cell Physiol. 56(6), 1215-1228. [301] Roy Choudhury, S., Riesselman, A.J., Pandey, S., 2014. Constitutive or seed-specific overexpression of Arabidopsis G-protein γ subunit 3 (AGG3) results in increased seed and oil production and improved stress tolerance in Camelina sativa. Plant Biotechnol. J. 12, 49-59. [302] Ruiz-Lopez, N., Broughton, R., Usher, S., Salas, J.J., Haslam, R.P., Napier, J.A., Beaudoin, F., 2017. Tailoring the composition of novel wax esters in the seeds of transgenic Camelina sativa through systematic metabolic engineering. Plant Biotechnol. J. 15(7), 837849. [303] Ruiz-Rico, M., Barat, J.M., 2021. Natural antimicrobial-coated supports as filter aids for the microbiological stabilisation of drinks. Lwt. 147, 111634. [304] Sabbahi, R., Azzaoui, K., Rhazi, L., Ayerdi-Gotor, A., Aussenac, T., Depeint, F., Taleb, M., Hammouti, B., 2023. Factors affecting the quality of canola grains and their implications for grain-based foods. Foods. 12(11), 2219. [305] Sagun, J.V., Yadav, U.P., Alonso, A.P., 2023. Progress in understanding and improving oil content and quality in seeds. Fron. Plant Sci. 14, 1116894. [306] Saini, R.K., Prasad, P., Sreedhar, R.V., Akhilender Naidu, K., Shang, X. and Keum, Y.S., 2021. Omega− 3 polyunsaturated fatty acids (PUFAs): Emerging plant and microbial sources, oxidative stability, bioavailability, and health benefits—A review. Antioxidants 10,0 1627. [307] Salaheldeen, M., Mariod, A.A., Aroua, M.K., Rahman, S.M.A., Soudagar, M.E.M., Fattah, I.M.R., 2021. Current state and perspectives on transesterification of triglycerides for biodiesel production. Catalysts 11, 1121. [308] Salehi Jouzani, G., Sharafi, R., Soheilivand, S., 2018. Fueling the future; plant genetic engineering for sustainable biodiesel production. Biofuel Res. J. 5(3), 829-845. [309] Salih, N., Salimon, J., 2021. A review on eco-friendly green biolubricants from renewable and sustainable plant oil sources. Biointerface Res. Appl. Chem, 11, 13303-27. [310] Salimon, J., Salih, N., Yousif, E., 2012. Industrial development and applications of plant oils and their biobased oleochemicals. Arab. J. Chem. 5, 135-145. [311] Sancheti, S.V., Yadav, G.D., 2022. Synthesis of environment‐friendly, sustainable, and nontoxic bio-lubricants: A critical review of advances and a path forward. Biofuels, Bioprod. Biorefin. 16, 1172-95. [312] Sankaranarayanan, R., Krolczyk, G.M., 2021. A comprehensive review on research developments of vegetable-oil based cutting fluids for sustainable machining challenges. J. Manuf. Process. 67, 286-313. [313] Santika, T., Budiharta, S., Law, E.A., Struebig, M., Ancrenaz, M., Poh, T.M., Ancrenaz, M., Struebig, M.J. and Meijaard, E., 2019. Does oil palm agriculture help alleviate poverty? A multidimensional counterfactual assessment of oil palm development in Indonesia. World Dev. 120, 10517. [314] Santin, A., Russo, M.T., Ferrante, M.I., Balzano, S., Orefice, I., Sardo, A., 2021. Highly valuable polyunsaturated fatty acids from microalgae: strategies to improve their yields and their potential exploitation in aquaculture. Molecules 26, 7697. [315] Sarubbo, L.A., da Gloria, C.S.M., Durval, I.J.B., Bezerra, K.G.O., Ribeiro, B.G., Silva, I.A., Twigg, M.S., Banat, I.M., 2022. Biosurfactants: Production, properties, applications, trends, and general perspectives. Biochem. Eng. J. 181, 108377. [316] Sarwer, A., Hussain, M., Al-Muhtaseb, A.H., Inayat, A., Rafiq, S., Khurram, M.S., Ul-Haq, N., Shah, N.S., Din, A.A., Ahmad, I., 2022. Suitability of biofuels production on commercial scale from various feedstocks: a critical review. Chem. BioEng. Rev. 9, 423-41. Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2142 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. [317] Sathe, S.K., Seeram, N.P., Kshirsagar, H.H., Heber, D., Lapsley, K.A., 2008. Fatty acid composition of California grown almonds. J. Food Sci. 73. [318] Savadi S., Vasupalli N., Kumar V., Dargan S., Gupta N. C., Chamola R., Bhat, S.R., 2015. Effect of overexpression of Arabidopsis thaliana SHB1 and KLUH genes on seed weight and yield contributing traits in Indian mustard (Brassica juncea L. (Czern.) Indian J. Genet. 75, 349-356. [319] Savadi, S., Lambani, N., Kashyap, P.L., Bisht, D.S., 2017. Genetic engineering approaches to enhance oil content in oilseed crops. Plant Growth Regul. 83, 207-222. [320] Scarlat, N., Dallemand, J.F., Monforti-Ferrario, F., Nita, V., 2015. The role of biomass and bioenergy in a future bioeconomy: Policies and facts. Environ. Dev. 15, 3-34. [321] Schmid, K.M., 2021. Lipid metabolism in plants. In Biochemistry of lipids, lipoproteins and membranes (pp. 121-159). Elsevier. [322] Schruff, M.C., Spielman, M., Tiwari, S., Adams, S., Fenby, N., Scott, R.J, 2006. The AUXIN RESPONSE FACTOR 2 gene of Arabidopsis links auxin signalling, cell division, and the size of seeds and other organs. Development. 133, 251-261. [323] Schwender, J., Hay, J.O., 2012. Predictive modeling of biomass component tradeoffs in Brassica napus developing oilseeds based on in silico manipulation of storage metabolism. Plant Physiol. 160(3), 1218-1236. [324] Selinski, J., Scheibe, R., 2019. Malate valves: old shuttles with new perspectives. Plant Biol. (Stuttgart, Germany), 21 Suppl 1(Suppl Suppl 1), 21-30. [325] Shah, R., Woydt, M., Zhang, S., 2021. The economic and environmental significance of sustainable lubricants. Lubricants 9, 111. [326] Shao, Q., Liu, X., Su, T., Ma, C., Wang, P., 2019. New insights into the role of seed oil body proteins in metabolism and plant development. Front. Plant Sci. 10, 1568. [327] Shen, B., Allen, W.B., Zheng, P., Li, C., Glassman, K., Ranch, J., Nubel, D., Tarczynski, M.C., 2010. Expression of ZmLEC1 and ZmWRI1 increases seed oil production in maize. Plant Physiol. 153(3), 980-987. [328] Sheri, V., Muddanuru, T., Mulpuri, S., 2021. Genetic Engineering of Sunflower (Helianthus annuus L.) for Important Agronomic Traits. In: Genetically Modified Crops. Springer. 175-200. [329] Shi, L., Song, J., Guo, C., Wang, B., Guan, Z., Yang, P., Chen, X., Zhang, Q., King, G.J., Wang, J., Liu, K., 2019. A CACTA-like transposable element in the upstream region of BnaA9.CYP78A9 acts as an enhancer to increase silique length and seed weight in rapeseed. Plant J. 98, 524-539. [330] Sia, C.B., Kansedo, J., Tan, Y.H., Lee, K.T., 2020. Evaluation on biodiesel cold flow properties, oxidative stability and enhancement strategies: a review. Biocatal. Agric. Biotechnol. 24, 101514. [331] Silva, R. de C.F.S., Almeida, D.G., Rufino, R.D., Luna, J.M., Santos, V.A., Sarubbo, L.A., 2014. Applications of biosurfactants in the petroleum industry and the remediation of oil spills. Int. J. Mol. Sci. 15, 12523-12542. [332] Simonsen, G., Ravotti, R., O'Neill, P., Stamatiou, A., 2023. Biobased phase change materials in energy storage and thermal management technologies. Renew. Sust. Energy Rev. 184, 113546. [333] Singh, H., Sharma, V.S., Dogra, M., 2020a. Exploration of graphene assisted vegetables oil based minimum quantity lubrication for surface grinding of TI-6AL-4V-ELI. Tribol. Int. 144, 106113. [334] Singh, Y., Sharma, A., Singh, N., Singla, M., Rastogi, P.M., 2020b. Prospects of inedible plant oil-driven bio-lubricants for tribological characteristics-a review. Int. J. Ambient Energy. 41, 1534-47. [335] Sinha, S., Jha, J.K., Maiti, M.K., Basu, A., Mukhopadhyay, U.K., Sen, S.K., 2007. Metabolic engineering of fatty acid biosynthesis in Indian mustard (Brassica juncea) improves nutritional quality of seed oil. Plant Biotechnol. Rep. 1, 185-197. [336] Siracusa, V., Blanco, I., 2020. Bio-polyethylene (Bio-PE), Biopolypropylene (Bio-PP) and Bio-poly(ethylene terephthalate) (BioPET): Recent developments in bio-based polymers analogous to petroleum-derived ones for packaging and engineering applications. Polymers. 12, 1641. [337] Song, H., Taylor, D.C., Zhang, M., 2023. Bioengineering of soybean oil and its impact on agronomic traits. Int. J. Mol. Sci. 24(3), 2256. [338] Song, Q.X., Li, Q.T., Liu, Y.F., Zhang, F.X., Ma, B., Zhang, W.K., Man, W.Q., Du, W.G., Wang, G.D., Chen, S.Y., Zhang, J.S., 2013. Soybean GmbZIP123 gene enhances lipid content in the seeds of transgenic Arabidopsis plants. J. Exp. Bot. 64, 4329-4341. [339] Souza, A.C., Goto, G.E.O., Mainardi, J.A., Coelho, A.C.V Tadini, C.C., 2013. Cassava starch composite films incorporated with cinnamon essential oil: Antimicrobial activity, microstructure, mechanical and barrier properties. LWT-Food Sci. Technol. 54, 346352. [340] Sova, N., Lutsenko, M., Korchmaryova, A., Andrusevych, K., 2018. Research of physical and chemical parameters of the oil obtained from organic and conversion hemp seeds varieties “Hliana”. Ukrainian Food Journal 7(2), 244-252. . [341] Statista, 2023a. Polyvinyl chloride (PVC) production volume worldwide in 2018 and 2025. [342] Statista, 2023b. Bio-pesticides market size worldwide in 2016 and 2021, with a forecast for 2027. [343] Stolp, L.J., Gronlund, P.J., Kodali, D.R., 2019. Soybean oil fatty acid ester estolides as potential plasticizers. J. Am. Oil Chem. Soc. 96, 727738. [344] Stoutjesdijk, P.A., Hurleston,e C., Singh, S.P., Green, A.G., 2000. High-oleic acid Australian Brassica napus and B. juncea varieties produced by co-suppression of endogenous Δ12-desaturases. Biochem. Soc. Trans. 28, 938-940. [345] Su, Y., Liang, W., Liu, Z., Wang, Y., Zhao, Y., Ijaz, B., Hua, J., 2017. Overexpression of GhDof1 improved salt and cold tolerance and seed oil content in Gossypium hirsutum. J. Plant Physiol. 218, 222-234. [346] Subedi, U., Jayawardhane, K.N., Pan, X., Ozga, J., Chen, G., Foroud, N.A., Singer, S.D., 2020a. The potential of genome editing for improving seed oil content and fatty acid composition in oilseed crops. Lipids. 55(5), 495-512. [347] Subedi, U., Ozga, J.A., Chen, G., Foroud, N.A., Singer, S.D., 2020b. CRISPR/Cas-mediated genome editing for the improvement of oilseed crop productivity. Crit. Rev. Plant Sci. 39, 195-221. [348] Sun, R., Ye, R., Gao, L., Zhang, L., Wang, R., Mao, T., Zheng, Y., Li, D., Lin, Y., 2017. Characterization and ectopic expression of CoWRI1, an AP2/EREBP domain-containing transcription factor from coconut (Cocos nucifera L.) endosperm, changes the seeds oil content in transgenic Arabidopsis thaliana and rice (Oryza sativa L.). Front. Plant Sci. 8, 63. [349] Tabari, M.A., Youssefi, M.R., Benelli, G., 2017. Eco-friendly control of the poultry red mite, Dermanyssus gallinae (Dermanyssidae), using the α-thujone-rich essential oil of Artemisia sieberi (Asteraceae): toxic and repellent potential. Parasitol. Res. 116, 1545-1551. [350] Talib, N., Rahim, E.A., 2015. Performance evaluation of chemically modified crude jatropha oil as a bio-based metalworking fluids for machining process. Procedia. Cirp. 26, 346-350. [351] Tan, C.X., Tan, S.S., Tan, S.T., 2020. Chapter 52 - Cold pressed macadamia oil, in: Ramadan, M.F. (Ed.) Cold Pressed Oils. Academic Press, pp. 587-595. [352] Tanasă, F., Teacă C.A., Zănoagă, M., 2021. Protective coatings for wood. in, Handbook of Modern Coating Technologies. Elsevier. [353] Tang, S., Guo, N., Tang, Q., Peng, F., Liu, Y., Xia, H., Lu, S., Guo, L., 2022. Pyruvate transporter BnaBASS2 impacts seed oil accumulation in Brassica napus. Plant Biotechnol. J. 20(12), 24062417. [354] Telaumbanua, M., Savitri, E.A., Shofi, A.B., Suharyatun, S,. Wisnu, F.K., Haryanto, A., 2021. Plant-based pesticide using citronella (Cymbopogon nardus L.) extract to control insect pests on rice plants. In: IOP Conference Series: Earth and Environmental Science. IOP Publishing. p 12071. [355] Thangadurai, D., Dabire, S.S., Sangeetha, J., Al-Tawaha, A.R.M.S., Adetunji, C.O., Islam, S., Shettar, A.K., David, M., Hospet, R., Adetunji, J.B., 2020. Greener composites from plant fibers: Preparation, structure, and properties. Handbook of Nanomaterials and Nanocomposites for Energy and Environmental Applications. 119. Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2143 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. [356] Theis, K.R., Venkataraman, A., Wagner, A.P., Holekamp, K.E., Schmidt, T.M., 2016. Age-Related Variation in the Scent Pouch (Hyaena hyaena). In: Chemical Signals in Vertebrates 13, 87-103. [357] Tian, Y., Lv, X., Xie, G., Wang, L., Dai, T., Qin, X., Chen, F., Xu, Y., 2019. FAX2 mediates fatty acid export from plastids in developing Arabidopsis seeds. Plant Cell Physiol. 60, 2231-2242. [358] Tian, Y., Lv, X., Xie, G., Zhang, J., Xu, Y., Chen, F., 2018. Seedspecific overexpression of AtFAX1 increases seed oil content in Arabidopsis. Biochem. Biophys. Res. Commun. 500, 370-375. [359] Tian, Y., Zhang, M., Hu, X., Wang, L., Dai, J., Xu, Y., Chen, F., 2016. Overexpression of CYP78A98, a cytochrome P450 gene from Jatropha curcas L., increases seed size of transgenic tobacco. Electron. J. Biotechnol. 19, 15-22. [360] Tsimidou, M., Blekas, G., Boskou, D., 2003. OLIVE OIL, in: Caballero, B. (Ed.) Encyclopedia of Food Sciences and Nutrition (Second Edition). Academic Press, Oxford, pp. 4252-4260. [361] Ullah, S, Dhar, N.R., 2018. Effects of vegetable oil based cutting fluid in machining Kevlar composite material. Am. J. Mech. Eng. 6, 54-60. [362] Uppar, R., Dinesha, P., Kumar, S., 2023. A critical review on vegetable oil-based bio-lubricants: Preparation, characterization, and challenges. Environ. Dev. Sustain. 25(9), pp.9011-9046. [363] Vaikuntapu, P.R., Kumar, V.D., 2023. Applications and challenges of harnessing genome editing in oilseed crops. J. Plant Biochem. Biotechnol. 32, 751-72. [364] Van Acker, J., den Bulcke, J.V., Forsthuber, B, Grüll, G., 2023. Wood preservation and wood finishing. in, Springer Handbook of Wood Science and Technology. Springer. [365] van Erp, H., Kelly, A.A., Menard, G., Eastmond, P.J., 2014. Multigene engineering of triacylglycerol metabolism boosts seed oil content in Arabidopsis. Plant Physiol. 165, 30-36. [366] Van Renterghem, L., Roelants, S.L.K.W., Baccile, N., Uyttersprot, K., Taelman, M.C., Everaert, B., Mincke, S., Ledegen, S., Debrouwer, S., Scholtens, K., 2018. From lab to market: an integrated bioprocess design approach for new‐to‐nature biosurfactants produced by Starmerella bombicol. Biotechnol. Bioeng. 115, 1195-206. [367] Vanhercke, T., Divi, U.K., El Tahchy, A., Liu, Q., Mitchell, M., Taylor, M.C., Eastmond, P.J., Bryant, F., Mechanicos, A., Blundell, C., Zhi, Y., Belide, S., Shrestha, P., Zhou, X.R., Ral, J.P., White, R.G., Green, A., Singh, S.P., Petrie, J.R., 2017. Step changes in leaf oil accumulation via iterative metabolic engineering. Metab. Eng. 39, 237-246. [368] Vanhercke, T., Dyer, J.M., Mullen, R.T., Kilaru, A., Rahman, M.M., Petrie, J.R., Green, A.G., Yurchenko, O., Singh, S.P., 2019. Metabolic engineering for enhanced oil in biomass. Prog. Lipid Res. 74, 103-129. [369] Vanhercke, T., El Tahchy, A., Shrestha, P., Zhou, X.R., Singh, S.P., Petrie, J.R., 2013. Synergistic effect of WRI1 and DGAT1 coexpression on triacylglycerol biosynthesis in plants. FEBS Lett. 587, 364-369. [370] Vantage Market Research, 2022. Biopesticide market research – Global industry assessment & forcast. Report Code: VMR-1854. [371] Vasantha-Srinivasan, P., Chellappandian, M., Senthil-Nathan, S., Ponsankar, A., Thanigaivel, A., Karthi, S., Edwin, E.S., Selin-Rani, S., Kalaivani, K., Maggi, F., Benelli, G., 2018. A novel herbal product based on Piper betle and Sphaeranthus indicus essential oils: Toxicity, repellent activity and impact on detoxifying enzymes GST and CYP450 of Aedes aegypti Liston (Diptera: Culicidae). J. Asia. Pac. Entomol. 21(4), 1466-1472. [372] Vaughn, A.R., Clark, A.K., Sivamani, R.K., Shi, V.Y., 2018. Natural oils for skin-barrier repair: Ancient compounds now backed by modern science. Am. J. Clin. Dermatol. 19, 103-117. [373] Verified Market Research, 2023. Global wax market size by end-user industry, by form, by application, by geographic scope and forecast. Report ID: 40682. [374] Verma, S., Lu, S., Kenis, P.J.A. 2019. Co-electrolysis of CO and glycerol as a pathway to carbon chemicals with improved technoeconomics due to low electricity consumption. Nat. Energy 4, 466-474. [375] Vital, A.C.P., Guerrero, A., Ornaghi, M.G., Kempinski, E.M.B.C., Sary, C., de Oliveira Monteschio, J., Matumoto-Pintro, P.T., Ribeiro, R.P., do Prado, I.N., 2018. Quality and sensory acceptability of fish fillet (Oreochromis niloticus) with alginate-based coating containing essential oils. J. Food Sci. Technol. 55, 4945-4955. [376] Wacal, C., Ogata, N., Basalirwa, D., Sasagawa, D., Kato, M., Handa, T., Masunaga, T., Yamamoto, S., Nishihara, E., 2019. Fatty Acid Composition of Sesame (Sesamum indicum L.) Seeds in Relation to Yield and Soil Chemical Properties on Continuously Monocropped Upland Fields Converted from Paddy Fields. Agronomy. 9, 1-19. [377] Wallis, J.G., Bengtsson, J.D., Browse, J., 2022. Molecular approaches reduce saturates and eliminate trans fats in food oils. Front. Plant Sci. 13, 908608. [378] Wan, H., Cui, Y., Ding, Y., Mei, J., Dong, H., Zhang, W., Wu, S., Liang, Y., Zhang, C., Li, J., 2017a. Time-series analyses of transcriptomes and proteomes reveal molecular networks underlying oil accumulation in canola. Front. Plant Sci.7, 2007. [379] Wan, S., Truong-Trieu, V.M., Ward, T., Whalen, J.K., Altosaar, I., 2017b. Advances in the use of genetically modified plant biomass for biodiesel generation. Biofuels, Bioprod Biorefin. [380] Wang, K., Froehlich, J. E., Zienkiewicz, A., Hersh, H. L., Benning, C., 2017. A plastid phosphatidylglycerol lipase contributes to the export of acyl groups from plastids for seed oil biosynthesis. Plant Cell 29, 1678-1696. [381] Wang, L., Wei, X., Wang, G., Zhao, S., Cui, J., Gao, A., Zhang, G., Yan, Y., 2020a. A facile and industrially feasible one-pot approach to prepare graphene-decorated PVC particles and their application in multifunctional PVC/graphene composites with segregated structure. Compos. B: Eng. 185, 107775. [382] Wang, S., Liu, S., Wang, J., Yokosho, K., Zhou, B., Yu, Y.C., Liu, Z., Frommer, W.B,. Ma, J.F., Chen, L.Q., Guan, Y., Shou, H., Tian, Z., 2020b. Simultaneous changes in seed size, oil content and protein content driven by selection of SWEET homologues during soybean domestication. Natl. Sci. Rev. 7, 1776-1786. [383] Wang, X., Xiao, B., Yang, G., Chen, J., Liu, W., 2021. Enzymatic preparation of phytosterol esters with fatty acids from high-oleic sunflower seed oil using response surface methodology. R.S.C. Adv. 11, 15204-15212. [384] Wang, M., Garneau, M.G., Poudel, A.N., Lamm, D., Koo, A.J., Bates, P.D., Thelen, J.J., 2022a. Overexpression of pea a-carboxyltransferase in Arabidopsis and Camelina increases fatty acid synthesis leading to improved seed oil content. Plant J. 110, 1035-104. [385] Wang, P., Xiong, X., Zhang, X., Wu, G., Liu, F., 2022b. A Review of erucic acid production in Brassicaceae oilseeds: Progress and prospects for the genetic engineering of high and low-erucic acid rapeseeds (Brassica napus). Fron. Plant Sci. 13, 899076. [386] Wang, Z., Wang, Y., Shang, P., Yang, C., Yang, M., Huang, J., Ren, B., Zuo, Z., Zhang, Q., Li, W., 2022c. Overexpression of soybean GmWRI1a stably increases the seed oil content in soybean. Int. J. Mol. Sci. 23, 5084. [387] Watts, N., Amann, M., Arnell, N., Ayeb-Karlsson, S., Beagley, J., Belesova, K., Boykoff, M., et al., 2021. The 2020 report of The Lancet Countdown on health and climate change: responding to converging crises. The Lancet. 397(10269), 129-170. [388] Wei, T., Mueed, A., Luo, T., Sun, Y., Zhang, B., Zheng, L., Deng, Z., Li, J., 2024. 1, 3-dioleoyl-2-palmitoyl-glycerol and 1-oleoyl-2palmitoyl-3-linoleoyl-glycerol: Structure-function relationship, triacylglycerols preparation, nutrition value. Food Chem. 138560. [389] Wei, W., Sun, C., Jiang, W., Zhang, X., Hong, Y., Jin, Q., Tao, G., Wang, X., Yang, Z., 2019. Triacylglycerols fingerprint of edible vegetable oils by ultra-performance liquid chromatography-Q-ToFMS. Lwt. 112, 108261. [390] Wenning, L., Yu, T., David, F., Nielsen, J., Siewers, V., 2017. Establishing very long‐chain fatty alcohol and wax ester biosynthesis in Saccharomyces cerevisiae. Biotechnol. Bioengin, 114(5), 10251035. [391] Weselake, R.J., 2016. Chapter 15 - Engineering Oil Accumulation in Vegetative Tissue. In: McKeon TA, Hayes DG, Hildebrand DF, Weselake RJ (eds) Industrial Oil Crops. AOCS Press. 413-434. [392] White, W.B., 2007. Clinical assessment of early morning blood pressure in patients with hypertension. Prev. Cardiol. 10(4), 210-214. [393] Wiberg, E., Edwards, P., Byrne, J., Stymne, S., Dehesh, K., 2000. The distribution of caprylate, caprate and laurate in lipids from developing Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2144 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. and mature seeds of transgenic Brassica napus L. Planta. 212(1), 3340. [394] Wickramasinghe, K.C., Sasahara, H., Rahim, E.A., Perera, G.I.P., 2021. Recent advances on high performance machining of aerospace materials and composites using vegetable oil-based metal working fluids. J. Clean. Prod. 310, 127459. [395] Wood, C.C., Okada, S., Taylor, M.C., Menon, A., Mathew, A., Cullerne, D., Stephen, S.J., Allen, R.S., Zhou, X.R., Liu, Q., Oakeshott, J.G., Singh, S.P., Green, A.G., 2018. Seed-specific RNAi in safflower generates a superhigh oleic oil with extended oxidative stability. Plant Biotechnol. J. 16(10), 1788-1796. [396] Xiao, Z., Tang, F., Zhang, L., Li, S., Wang, S., Huo, Q., Yang, B., Zhang, C., Wang, D., Li, Q., Wei, L., Guo, T., Qu, C., Lu, K., Zhang, Y., Guo, L., Li, J., Li, N., 2021. The Brassica napus fatty acid exporter FAX1-1 contributes to biological yield, seed oil content, and oil quality. Biotechnol. Biofuels 14, 190. [397] Xu, C., Shanklin, J., 2016. Triacylglycerol Metabolism, Function, and Accumulation in Plant Vegetative Tissues. Annu. Rev. Plant Biol .67, 179-206. [398] Xu, F., Fang, J., Ou, S., Gao, S., Zhang, F., Du, L., Xiao, Y., Wang, H., Sun, X., Chu, J., 2015. Variations in CYP 78 A 13 coding region influence grain size and yield in rice. Plant Cell Environ. 38(4), 800811. [399] Xu, X.Y., Yang, H.K., Singh, S.P., Sharp, P.J., Liu, Q., 2018. Genetic manipulation of non-classic oilseed plants for enhancement of their potential as a biofactory for triacylglycerol production. Engineering, 4(4), 523-533. [400] Xu, Y., Schmiege, S.C. and Sharkey, T.D., 2024. The oxidative pentose phosphate pathway in photosynthesis: a tale of two shunts. New Phytol. 242: 2453-2463. [401] Xuan, W., Odelius. K., Hakkarainen, M., 2020. Dual-Functioning Antibacterial Eugenol-Derived Plasticizers for Polylactide. Biomolecules 10, 1077. [402] Yaashikaa, P.R., Kumar, P.S., Karishma, S., 2022. Bio-derived catalysts for production of biodiesel: A review on feedstock, oil extraction methodologies, reactors and lifecycle assessment of biodiesel. Fuel. 316, 123379. [403] Yang, Y., Huang, J., Zhang, R., Zhu, J., 2017. Designing bio-based plasticizers: Effect of alkyl chain length on plasticization properties of isosorbide diesters in PVC blends. Mater .Des. 126, 29-36. [404] Yang, A., Qi, M., Wang, X., Wang, S., Sun, L., Qi, D., Zhu, L., Duan, Y., Gao, X., Ali Rajput, S., 2019. Refined cottonseed oil as a replacement for soybean oil in broiler diet. Food Sci. Nutr. 7(3), 10271034. [405] Yang, Z., Liu, X., Wang, K., Li, Z., Jia, Q., Zhao, C., Zhang, M., 2021. ABA-INSENSITIVE 3 with or without FUSCA3 highly up-regulates lipid droplet proteins and activates oil accumulation. J. Exp. Bot. 73, 2077-2092. [406] Yang, J., Chen, B., Manan, S., Li, P., Liu, C., She, G., Zhao, S., Zhao, J., 2022a. Critical metabolic pathways and SAD/FADs, WRI1s, and DGATs cooperate for high-oleic acid oil production in developing oil tea (Camellia oleifera) seeds. Hortic. Res. 9, uhac087. [407] Yang, Y., Kong, Q., Lim, A.R.Q., Lu, S., Zhao, H., Guo, L., Yuan, L., Ma, W., 2022b. Transcriptional regulation of oil biosynthesis in seed plants: Current understanding, applications, and perspectives. Plant Commun. 3, 100328. [408] Yarolimek, M.R., Bookbinder, H.R., Coia, B.M., Kennemur, J.G., 2021. Ring-Opening Metathesis Polymerization of δ-Pinene: WellDefined Polyolefins from Pine Sap. A.C.S. Macro. Lett. 10760-766. [409] Ye, J., Wang, C., Sun, Y., Qu, J., Mao, H., Chua, N.H. 2018. Overexpression of a transcription factor increases lipid content in a Woody perennial Jatropha curcas. Front. Plant Sci. 9, 1479. [410] Yin, Q,. Li, C., Dong, L., Bai, X., Zhang, Y., Yang, M., Jia, D., Li, R., Liu, Z., 2021. Effects of physicochemical properties of different base oils on friction coefficient and surface roughness in MQL milling AISI 1045. Int. J. Precis Eng. Manuf. - Green Technol. 8, 1629-1647. [411] Yu, D., Hornung, E., Iven, T., Feussner, I., 2018. High-level accumulation of oleyl oleate in plant seed oil by abundant supply of oleic acid substrates to efficient wax ester synthesis enzymes. Biotechnol. Biofuel. 11, 53. [412] Yuan, H., Yao, J., Masakorala, K., Wang, F., Cai, M., Yu, C., 2014. Isolation and characterization of a newly isolated pyrene-degrading Acinetobacter strain USTB-X. Environ. Sci. Pollut. Res. 21, 27242732. [413] Yurchenko, O., Shockey, J.M., Gidda, S.K., Silver, M.I., Chapman, K.D., Mullen, R.T., Dyer, J. M., 2017. Engineering the production of conjugated fatty acids in Arabidopsis thaliana leaves. Plant Biotechnol. J. 15(8), 1010-1023. [414] Zafar, S., Li, Y.L., Li, N.N., Zhu, K.M., Tan, X.L., 2019. Recent advances in enhancement of oil content in oilseed crops. J. Biotechnol. 301, 35-44. [415] Zale, J, Jung, H., Kim, J.Y., Pathak, B., Karan, R., Liu, H., Chen, X., Wu, H., Candreva, J., Zhai, Z., 2016. Metabolic engineering of sugarcane to accumulate energy‐dense triacylglycerols in vegetative biomass. Plant Biotechnol. J. 14, 661-69. [416] Zhai, Z., Liu, H., Shanklin, J., 2021. Ectopic expression of OLEOSIN 1 and inactivation of GBSS1 have a synergistic effect on oil accumulation in plant leaves. Plants. 10(3), 513. [417] Zhang, C., Garrison,T.F., Madbouly, S.A., Kessler, M.R., 2017. Recent advances in vegetable oil-based polymers and their composites. Prog. Polym. Sci. 71, 91-143. [418] Zhang, C., Iskandarov, U., Klotz, E.T., Stevens, R.L., Cahoon, R.E., Nazarenus, T.J., Pereira, S.L., Cahoon, E. B., 2013. A thraustochytrid diacylglycerol acyltransferase 2 with broad substrate specificity strongly increases oleic acid content in engineered Arabidopsis thaliana seeds. J. Exp. Bot. 64(11), 3189-3200. [419] Zhang, L., Yang, X.D., Zhang, Y.Y., Yang, J., Qi, G.X., Guo, D.Q., Xing, G.J., Yao, Y., Xu, W.J., Li, H.Y., Li, Q.Y., Dong, Y.S., 2014. Changes in oleic Acid content of transgenic soybeans by antisense RNA mediated posttranscriptional gene silencing. Int. J. Genom. 921950. [420] Zhang, M., Cao, X., Jia, Q., Ohlrogge, J., 2016. FUSCA 3 activates triacylglycerol accumulation in Arabidopsis seedlings and tobacco BY 2 cells. Plant J. 88, 95-107. [421] Zhang, D., Zhang, H., Hu, Z., Chu, S., Yu, K., Lv, L., Yang, Y., Zhang, X., Chen, X., Kan, G., Tang, Y., An, Y.Q.C., Yu, D., 2019a. Artificial selection on GmOLEO1 contributes to the increase in seed oil during soybean domestication. PloS Genet. 15, 1008267. [422] Zhang, X., Hong, M., Wan, H., Luo, L., Yu, Z., Guo, R., 2019b. Identification of key genes involved in embryo development and differential oil accumulation in two contrasting maize genotypes. Genes, 10(12), 993. [423] Zhang, Z., Jiang, P., Liu, D., Feng, S., Zhang, P., Wang, Y., Fu, J., Agus, H., 2021. Research progress of novel bio-based plasticizers and their applications in poly (vinyl chloride). J. Mater. Sci. 56, 1015510182. [424] Zhang, K., He, J., Yin, Y., Chen, K., Deng, X., Yu, P., Li, H., Zhao, W., Yan, S. and Li, M., 2022. Lysophosphatidic acid acyltransferase 2 and 5 commonly, but differently, promote seed oil accumulation in Brassica napus. Biotechnol. biofuels bioprod. 15, 83. [425] Zhao, B., Dai, A., Wei, H., Yang, S., Wang, B., Jiang, N., Feng, X., 2016. Arabidopsis KLU homologue GmCYP78A72 regulates seed size in soybean. Plant. Mol. Biol. 90, 33-47. [426] Zhao, Y., Cao, P., Cui, Y., Liu, D., Li, J., Zhao, Y., Yang, S., Zhang, B., Zhou, R., Sun, M., Guo, X., Yang, M., Xin, D., Zhang, Z., Li, X., Lv, C., Liu, C., Qi, Z., Xu, J., Wu, X., Chen, Q., 2021. Enhanced production of seed oil with improved fatty acid composition by overexpressing NAD+-dependent glycerol-3-phosphate dehydrogenase in soybean. J. Integr. Plant Biol. 63, 1036-1053. [427] Zhang, Y., Yu, L., Yung, K.F., Leung, D.Y., Sun, F., Lim, B.L., 2012. Over-expression of AtPAP2 in Camelina sativa leads to faster plant growth and higher seed yield. Biotechnol. Biofuels, 5, 1-10. [428] Zheng, T., Wu, Z., Xie, Q., Fang, J,. Hu, Y., Lu, M., Xia, F., Nie, Y., Ji, J. 2018. Structural modification of waste cooking oil methyl esters as cleaner plasticizer to substitute toxic dioctyl phthalate. J. Clean Prod. 186, 1021-1030. [429] Zhou, Y., Zhao, W., Lai, Y., Zhang, B., Zhang, D., 2020. Edible plant oil: global status, health issues, and perspectives. Front. Plant Sci. 11,1315. Hajinajaf et al. / Biofuel Research Journal 42 (2024) 2105-2145 2145 Please cite this article as: Hajinajaf N., Fayyazbakhsh A., Kamal Shahsavar S., Sanjarian F., Rahnama H. Boosting plant oil yields: the role of genetic engineering in industrial applications. Biofuel Research Journal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. [430] Zhu, F., Farnung, L., Kaasinen, E., Sahu, B., Yin, Y., Wei, B., Dodonova, S.O., Nitta, K.R., Morgunova, E., Taipale, M., Cramer, P., 2018. The interaction landscape between transcription factors and the nucleosome. Nature, 562(7725), 76-81. [431] Zhu, G., Liu, C., Zhang, C., 2023. Plant oil-based polymers. Phys. Sci. Rev. 8, 895-936. [432] Zhukov, A., Popov, V., 2022. Synthesis of C20-38 fatty acids in plant tissues. Int. J. Mol. Sci. 23, 4731. [433] Zubair, M., Pradhan, R.A., Arshad, M., Ullah, A., 2021. Recent advances in lipid derived bio‐based materials for food packaging applications. Macromol. Mater. Eng. 306, 2000799. Nima Hajinajaf is a Ph.D. candidate in the Chemical Engineering department at Arizona State University, Tempe, USA. He holds a Master's degree in Chemical Engineering – Process Design from the University of Tehran, Iran, and a Bachelor's Degree in Chemical Engineering from the Petroleum University of Technology, Ahvaz, Iran. Nima has authored over 17 peer-reviewed journal papers with an h-index of 10, addressing various aspects of microbial biotechnology and bioprocess engineering. His research focuses on (1) CRISPR-Cas genome editing for metabolic enhancement in cyanobacteria and E. coli; (2) flux balance analysis for optimizing biochemical production; (3) advanced bioreactor design and scale-up; and (4) techno-economic evaluations of biobased processes. His comprehensive research profile is available at: https://scholar.google.com/citations?user=pcuiynEAAAAJ&hl=en&oi=ao. Ahmad Fayyzbakhsh holds a PhD in Chemistry from Tomas Bata University in Czechia, where their research under the Horizon 2020 project focused on controlling the biodegradation of three polyesters. Currently, he works as a researcher at Montan University in Leoben, Austria, which continues to pioneer sustainable solutions in energy science. His research profile can be found at: https://scholar.google.com/citations?user=eQ-zZ_IAAAAJ&hl=en. Sara Kamal Shahsavar is a researcher with a Master of Science in Medical Microbiology from the Mashhad University of Medical Sciences, Iran. Her thesis explored the inhibitory potential of IgY antibodies in the treatment and prophylaxis of Helicobacter pylori infection. Sara's academic journey began when she earned her Bachelor of Science in Cellular and Molecular Biology. Currently, Sara's research interests span the development of novel biomedical applications of natural compounds, with a focus on the therapeutic potentials of antibodies. Her research interests include (1) Gene expression, (2) Recombinant Protein Expression and Purification, and (3) Plasmid Cloning. Her research profile is available at: http://orcid.org/0009-0009-2480-5106. Dr. Hassan Rahnama got his PhD in Plant Physiology from the Faculty of Science, University of Tehran, Iran, in 2004. Currently, he works as an associate Prof. in the Plant Genetic Engineering and Biosafety Department of the Agricultural Biotechnology Research Institute of Iran (ABRII). His research focuses on the genetic engineering of oilseed crops to improve their quality and quantity characteristics. Dr. Rahnama published several papers and books on plant biotechnology, plant genetic engineering, bioethics and biosafety, etc. His comprehensive research profile is available at: https://scholar.google.com/citations?user=dWJDZZQAAAAJ&hl=en&oi= ao. Dr. Forough Sanjarian earned her PhD in Cell and Molecular Biology from the Faculty of Science at Razi University in Iran in 2006. She currently serves as a faculty member in the Plant Bio-product Department at the National Institute for Genetic Engineering and Biotechnology (NIGEB) in Tehran, Iran. Her research primarily focuses on plant genetic engineering, physiology, and biotechnology. Dr. Sanjarian specializes in studying plant responses to biotic and abiotic stresses, with a specific emphasis on genetic transformation and a deep understanding of molecular mechanisms. Her research profile can be found at: https://scholar.google.com/citations?user=qJbNtukAAAAJ&hl=en.