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

* Co esponding au ho a : E-mail add ess: h ahnama@ab ii.ac.i ‡: These au ho s con ibu ed equally. Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Bio uel Resea ch Jou nal 42 (2024) 2105-2145 Re iew Pape Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions Nima Hajinaja 1,‡, Ahmad Fayyazbakhsh2,‡, Sa a Kamal Shahsa a 3, Fo ough Sanja ian4, Hassan Rahnama5,* 1Chemical Enginee ing P og am, School o Enginee ing o Ma e , T anspo , and Ene gy, A izona S a e Uni e si y, Tempe, AZ, USA. 2Depa men o En i onmen al P o ec ion Enginee ing, Facul y o Technology, Tomas Ba a Uni e si y in Zlín, T. G. Masa yka Squa e 5555, 760 01 Zlín, Czech Republic. 3Depa men o Mic obiology and Vi ology, School o Medicine, Mashhad Uni e si y o Medical Sciences, Mashhad, I an. 4Na ional Ins i u e o Gene ic Enginee ing and Bio echnology, Teh an, I an. 5Ag icul u al Bio echnology Resea ch Ins i u e o I an (ABRII), Ag icul u al Resea ch Educa ion and Ex ension O ganiza ion (AREEO), Ka aj, I an. HIGHLIGHTS GRAPHICAL ABSTRACT ➢ Plan oil-based p oduc s ha e eme ged as eco- iendly al e na i es o pe oleum coun e pa s. ➢ Plan oils a e e sa ile, wi h applica ions in cooking, lub ica ion, cosme ics, polyme s, and medicine. ➢ Non-edible plan oils p esen new oppo uni ies o biodiesel and biop oduc p oduc ion. ➢ Gene ic enginee ing can enhance bo h he yield and quali y o plan oils o bio-based indus ies. ARTICLE INFO ABSTRACT A icle his o y: Recei ed 25 Ma ch 2024 Recei ed in e ised o m 20 May 2024 Accep ed 26 May 2024 Published 1 June 2024 Keywo ds: Bio uel Gene ic enginee ing Plan oils T iacylglyce ol Bio-based economy Sus ainabili y As clima e change in ensi ies and he need o educe human-caused emissions becomes mo e u gen , ansi ioning o a bio-based economy is essen ial. This pape explo es he di e se indus ial applica ions o plan oils as sus ainable al e na i es o pe oleum- based p oduc s, including hei use in ood, polyme s, lub ican s, su ac an s, pes icides, emollien s, and bio uels. This e iew del es in o biosyn he ic pa hways, de ailing he key enzymes and p ocesses in ol ed in he syn hesis o iacylglyce ol. I ho oughly discusses how gene ic and me abolic enginee ing can no only inc ease oil yields bu also modi y a y acid composi ions o be e mee indus ial equi emen s. By unde s anding gene ics and u ilizing ad anced bio echnologies, he oil con en and quali y o plan sou ces can be signi ican ly enhanced, aligning wi h bo h sus ainabili y goals and indus ial demands. This pape p o ides a comp ehensi e o e iew o he cu en uses and gene ic enginee ing o plan oil p oduc ion, p oposing inno a i e s a egies such as u ilizing oils om biomass o cul i a ing non-edible oil c ops. These app oaches aim o es ablish a sus ainable indus ial sys em, educe eliance on ossil uels, and p omo e he g ow h o an en i onmen ally esponsible bio- based economy. Addi ionally, he e iew highligh s u u e di ec ions, examining he economic implica ions and en i onmen al bene i s o adop ing plan oils ac oss a ious sec o s and posi ioning hem as pi o al o achie ing an eco- iendly, bio-based economy. ©2024 Alpha C ea ion En e p ise CC BY 4.0 Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2106 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Con en s 1. In oduc ion .......................................................................................................................................................................................................................... 2106 2. Indus ial applica ions o plan oil and he challenges aced ................................................................................................................................................. 2107 2.1. Polyme s ...................................................................................................................................................................................................................... 2109 2.2. Su ac an s ................................................................................................................................................................................................................... 2110 2.3. Lub ican s .................................................................................................................................................................................................................... 2115 2.4. Me al wo king luid ..................................................................................................................................................................................................... 2115 2.5. S abilize s and plas icize s ........................................................................................................................................................................................... 2115 2.6. Composi es................................................................................................................................................................................................................... 2116 2.7. Pes icides ..................................................................................................................................................................................................................... 2118 2.8. Pain , coa ing and adhesi es ......................................................................................................................................................................................... 2118 2.9. Inks .............................................................................................................................................................................................................................. 2118 2.10. Emollien s .................................................................................................................................................................................................................. 2118 2.11. Wax es e s .................................................................................................................................................................................................................. 2118 2.12. Tex ile inishing ......................................................................................................................................................................................................... 2122 3. Plan oil p oduc ion pa hways .............................................................................................................................................................................................. 2122 3.1. Fa y acids syn hesis ..................................................................................................................................................................................................... 2123 3.2. T iacylglyce ol biosyn hesis......................................................................................................................................................................................... 2124 3.3. T iacylglyce ol s o age ................................................................................................................................................................................................. 2124 4. Gene ic enginee ing o imp o ed oil p oduc ion ................................................................................................................................................................. 2124 4.1. Me abolic enginee ing o enhance oil p oduc ion ......................................................................................................................................................... 2124 4.1.1. Inc easing he con en o oil pe seed ................................................................................................................................................................... 2126 4.1.1.1. FA syn hesis (Ca bon lux edi ec ion) ........................................................................................................................................................ 2127 4.1.1.2. Glyce ol backbone ....................................................................................................................................................................................... 2127 4.1.1.3. TAG biosyn hesis ........................................................................................................................................................................................ 2127 4.1.1.4. Lipid ans e p o eins ................................................................................................................................................................................. 2127 4.1.1.5. T ansc ip ion ac o s (TFs) .......................................................................................................................................................................... 2127 4.2. Inc easing seed oil con en by enhancing seed size ...................................................................................................................................................... 2128 4.3. Gene ic Enginee ing o new oil esou ces: biomass-de i ed oil .................................................................................................................................. 2128 4.4. Modi ying he composi ion o ege able oils o indus ial applica ions ...................................................................................................................... 2130 4.4.1. Monounsa u a ed a y acids ................................................................................................................................................................................ 2131 4.4.2. Medium-chain sa u a es ....................................................................................................................................................................................... 2132 4.4.3. Enginee ing wax es e syn hesis .......................................................................................................................................................................... 2133 5. Policy and p ac ical implica ions .......................................................................................................................................................................................... 2133 6. Challenges and p ospec s ...................................................................................................................................................................................................... 2133 7. Conclusions .......................................................................................................................................................................................................................... 2133 Re e ences ................................................................................................................................................................................................................................ 2134 1. In oduc ion Clima e change and i s ad e se impac s on a ious aspec s o human heal h we e ecen ly p ojec ed in he la es global epo eleased by he Lance Coun down: T acking P og ess on Heal h and Clima e Change (Wa s e al., 2021). Among he epo ed indings was an inc eased a e o exposu e o hea wa es globally be ween 2000 and 2016, a ec ing an addi ional 125 million medically ulne able adul s. The widesp ead use o pe oleum-de i ed p oduc s has been linked o ising a mosphe ic CO2 le els, which a e associa ed wi h he equency o hea wa es (Ve ma e al., 2019; Rej e al., 2022). These indings highligh he need o dec ease an h opogenic g eenhouse gas (GHG) emissions h ough s a egies such as ansi ioning owa ds a bio-based economy (Be g eund e al., 2021; Hajinaja e al., 2022c; Hajinaja e al., 2024). Replacing pe oleum-de i ed p oduc s wi h eco- iendly al e na i es is a key ea u e o his ansi ion (De V ieze e al., 2020). The e a e g owing conce ns abou he u u e a ailabili y o pe oleum- de i ed p oduc s (Si acusa and Blanco, 2020). U ilizing enewable aw ma e ials o daily li e p oduc s seems c ucial o sus ainable de elopmen (Be g eund e al., 2021; Hajinaja e al., 2022b). Figu e 1a shows he sha e o di e en sou ces in he global p ima y ene gy supply. Using enewable ma e ials can educe CO2 emissions and o e addi ional ad an ages Abb e ia ions ABA Abscisic acid G3PDH Glyce ol-3-phospha e dehyd ogenase ACCase Ace yl-CoA ca boxylase GAPDH Glyce aldehyde 3-phospha e dehyd ogenase AP2 Ape ala2 GPAT Glyce ol-3-phospha e acyl ans e ase ARF2 Auxin Response Fac o 2 LPAAT Lysophospha idic acid acyl ans e ase CAGR Compound annual g ow h a e MWFs Me alwo king luids CPT Choline phospho ans e ase PC Phospha idylcholine CRISPR Clus e ed egula ly in e spaced sho palind omic epea s PDAT Phospholipid: diacylglyce ol acyl ans e ase DGAT Diacylglyce ol acyl ans e ase PVC Poly inyl chlo ide ER Endoplasmic e iculum PXA1 Pe oxisomal ABC anspo e 1 ESBO Epoxidized soybean oil SDP1 SUGAR-DEPENDENT1 FA Fa y Acids TAG T iacylglyce ol FAD2 Fa y acid desa u ase 2 TFs Mul iple ansc ip ion ac o s FAR Fa y Acid Reduc ases TP T iose-phospha es G3P Glyce ol-3-phospha e TTG2 T anspa en Tes a Glab a 2 Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2107 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Fig. 1. a) Sha e (%) o di e en sou ces in global p ima y ene gy supply (Kan e al., 2019); b) Sha es o wo ld oil consump ion in a ious sec o s and indus ies in he yea 2018 (IEA., 2018). The “O he ” ca ego y encompasses ag icul u e, comme cial and public se ices, non-speci ied o he , pipeline, and non-speci ied anspo ; c) A ea equipped o i iga ion (OECD, 2012); d) To al a able land in use (FAO, 2011). associa ed wi h g een chemis y, such as biodeg adabili y and lowe oxici y (Fallahi e al., 2021; Hajinaja e al., 2021). One o he mos impo an needs o a ious indus ies, including he ood indus y, is indus ial oils, anging om lub ica ing o hyd aulic and cu ing oils. Cu en ly, a signi ican po ion o indus ial oil esou ces used e en in he ood indus y a e de i ed om pe oleum, leading o subs an ial oleochemical pollu ion (Me zge and Hü e mann, 2009; Hayes, 2021; Fayyazbakhsh e al., 2022). Figu e 1b shows he dis ibu ion o global oil consump ion ac oss di e en sec o s and indus ies in 2018 (IEA., 2018). Plan oils a e conside ed ideal en i onmen ally iendly, enewable, and sus ainable eeds ocks ha could po en ially eplace pe oleum-de i ed oil in he men ioned indus ies (El-Dala ony e al., 2022; Yaashikaa e al., 2022). Wi h a ma ke cap compa able o ha o ossil-based uels (Hoang e al., 2021), indus ial oil is a key sec o o ansi ioning om pe oleum- based o plan -based oils. Howe e , since o e 85% o he plan oils p oduced globally a e used o human nu i ion, i is un ealis ic o ully eplace ossil oils wi h plan oils in indus y and anspo a ion (Hajja i e al., 2017). I has been highligh ed ha eplacing jus 40% o he ossil oils in hese indus ies would equi e global plan oil p oduc ion o iple by 2030 (Ca lsson e al., 2011). The inc easing sca ci y o enewable wa e esou ces (as highligh ed in Figu e 1c and de ailed in Table 1), along wi h he diminishing a ailabili y o a able land (as shown in Fig. 1d), adds signi ican complexi y o his endea o . This challenge is pa icula ly p onounced when we depend on exis ing oil c ops wi h s abilized yields and oil con en . Table 2 shows he yield and oil con en o a ious plan oil eeds ocks. In addi ion o yield and oil con en , challenges associa ed wi h exis ing plan oils ex end o un a o able a y acid p o iles in ce ain eeds ocks, leading o bio-oil p ope ies unsui able o speci ic indus ial applica ions. Mo eo e , he p esence o oxic o alle genic compounds u he complica es he u iliza ion o hese oils (Baska e al., 2019; Nomanbhay e al., 2018). The e o e, i is c i ical o de elop inno a i e and p omising indus ial plan oil pla o ms o ackle hese challenges. Howe e , i is equally c ucial o ensu e ha hese pla o ms do no in ensi y compe i ion wi h ood c ops, he eby sa egua ding global ood secu i y. This impo ance is highligh ed by global plan oil p oduc ion s a is ics, which o ecas an inc ease om 149 million onnes in 2005 o 282 million onnes in 2050, aimed a mee ing he demands o a g owing wo ld popula ion (Alexand a os and B uinsma, 2012). Table 3 p o ides an o e iew o global plan oil p oduc ion, cu en demands, and p ojec ed demands beyond 2050. Besides inc easing plan oil p oduc ion yield pe hec a e, one o he main app oaches o mee ing indus ial demand o plan oils is de eloping new oil c ops ha can u ilize ma ginal o non-ag icul u al lands and wa e s. In alignmen wi h hese goals, his e iew aims o comp ehensi ely explo e he di e se applica ions o plan oils ac oss a ious indus ies while add essing he associa ed challenges. I also del es in o plan me abolic pa hways o oil p oduc ion and he use o gene ic enginee ing o enhance oil quan i y and quali y. Addi ionally, he e iew c i ically examines inno a i e al e na i e s a egies, such as biomass-de i ed oils. Table 4 summa izes he a ious aspec s o plan oils, hei applica ions, and ela ed esea ch co e ed in his e iew, compa ing hem o discussions in e iew a icles published om 2017 o 2023. 2. Indus ial applica ions o plan oil and he challenges aced Majo oil c ops on which global oil p oduc ion depends include palm, soybean, apeseed (also known as canola), co n, sun lowe , co onseed, oli e, and peanu . Mino oil eeds ocks include sa lowe , coconu , sesame, and linseed (Table 2) (Alexand a os and B uinsma, 2012; Wan e al., 2017b). While he majo i y o global oil p oduc ion o e he las decade has been di ec ed owa d ood/ eed applica ions, app oxima ely one- i h has been used o indus ial and bioene gy applica ions, and his p opo ion is expec ed o inc ease (Sca la e al., 2015). In o he wo ds, he a io be ween ood, eed, and indus ial/bioene gy applica ions o global oil p oduc ion gene ally s ands a 80:6:14 (Quispe e al., 2013). Howe e , wi h g owing biodiesel p oduc ion, his a io has shi ed o 74:6:20 (Bie mann e al., 2011). These p opo ions a e s ill an icipa ed o change u he in a o o non- ood applica ions, including bioene gy and indus ial plan -based oil p oduc ion (Ra hou e al., 2023). The e o e, he main challenge is inc easing he quan i y o global plan oil p oduc ion o mee hese g owing Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2108 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 1. Annual enewable wa e esou ces and i iga ion wa e wi hd awal. P ecipi a ion (mm/annum) Renewable Wa e Resou ces (km3) Wa e Use E iciency Ra io (%) I iga ion Wa e Wi hd awal (km3) P essu e on Wa e Resou ces due o I iga ion (%) * 2005/2007 2050 2005/2007 2050 2005/2007 2050 Wo ld 800 4200 44 46 2620 2906 6 7 De eloped Coun ies 540 14000 42 43 505 493 4 4 De eloping Coun ies 990 28000 44 47 2115 2413 8 9 *Wa e wi hd awal o i iga ion as a pe cen age o o al annual enewable wa e esou ces. Sou ces: B uinsma (2009); Nach e gaele e al. (2023); h p://www. ao.o g/n /solaw/ hema ic- epo s/en/. Table 2. Di e en plan oil eeds ocks, hei p oduc ion pa ame e s, and comme cial applica ions. Sou ce: Wan e al. (2017b). Comme cial Uses (non- ood uses in Bold) Global Oil P oduc ion (million onnes) in 2016 Global Seed P oduc ion (million onnes) in 2016 Global Cul i a ion A ea (million ha) in 2016 L oil/ha kg oil/ha Oil Con en (w %) Oil C op Food, biodiesel, inks, plas icize s, c ayons, pain s, 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 (B assica napus L.) Food, coa ings 17.11 45.36 23.36 952 800 25-35 Sun lowe (Helian hus annuus) Food, lub ican s, inks 2.854 2.713 2.700 9.7 1212 1019 10-30 Oli e (Olea eu opaea) Medicine, biodiesel, cosme ics, massage oils and soaps, con ec ione y and bake y indus ies 5.78 42.28 24.77 1059 890 45-55 Peanu (A achis hypogaea) Food and eed, ma ga ine, salad d essings, medicine 5.09 38.88 29.58 325 273 18-25 Co on seed (Gossypium hi su um) Food, ma ga ine, sho ening, cooking oil, con ec iona y, soaps, sauces, a subs i u es, biodiesel 7.58 ke nel 63.86 palm Palm ke nel 17.09 - 5950 5000 30-60 Palm (A ecaceae) Food, ma ga ine, pain ing, skin mois u ize , and c eams o so ening and smoo hing he skin - 0.8 <1 779 655 20-45 Sa lowe (Ca hamus inc o ius L.) Food, biodiesel, ca ie o d ug molecules in pha maceu ical p epa a ions, soap, sal e, inks and pain , and ex ile indus ies 0.523 1608.62 35 172 145 3.3-15.9 Co n (Zea mays) Food, wholesome onic, medicine, hai ea men , body massage, wo ship 1.926 4.4 8 696 585 52-63 Sesame (Sesamum indicum) Food, medicine, heal ca e, uel 3.44 61.44 - 2689 2260 63-65 Coconu (Cocos nuci e a) Food, medicine, soap p oduc ion, be e ages, pha maceu icals - 8.6 - 572 481 25-30 Mus a d (B assica alba) Food, ma ga ine, medicinal 1.8 63 - 828 696 12.1-25 16-32 Rice b an (O yza sa i a L.) Enamels, a nishes, esins, coa ings - - - 940 790 14-22 Tung ui (Aleu i es o dii) - - 8.9 - 2638 2217 11.23- 18.8 A ocado (Pe sea ame icana) - 0.015 - - 1892 1590 35-40 Ja opha (Ja opha cu cas L.) - 0.055 - - - - 27-39 Ka anja (Pongamia pinna a) Plas icize s, lub ican s, medicine, adhesi es, cosme ics, hai oils, ood con aine s, uel addi i es, insula ion, nylon, syn he ic esins, ibe s, pain s, a nishes, plas ics, inks, ex iles, d ying oils, ungus-g ow h-inhibi ing compounds, embalming luid, soaps, dyeing aids, cleaning p oduc s, de e gen s, pe sonal ca e p oduc s, s yling gel, and adhesi e emo e 0.73 1.8 - 1413 1188 53 37-60 Cas o (Ricinus communis) - - - - 583 490 30-40 Camelina (Camelina sa i a) - - - - 1818 1528 44-59 Jojoba (Simmondsia chinensis) Medicine, eeds, sealan s, caulking compounds, linoleum, ea hen loo s, adobe, ex iles, ixa i e, us inhibi o , lub ican , lea he ea men , polishes, a nishes, oil pain s, composi ion o namen o molded deco a ion, animal ca e p oduc s, wood p ese a ion, indus ial lub ican 0.8 3.9 - 478 402 38 -44 Linseed (Linum usi a issimum) Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2109 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 2. con inued. Comme cial Uses (non- ood uses in Bold) Global Oil P oduc ion (million onnes) in 2016 Global Seed P oduc ion (million onnes) in 2016 Global Cul i a ion A ea (million ha) in 2016 L oil/ha kg oil/ha Oil Con en (w %) Oil C op - - 5.8 - 1026 863 40-45 Coca (Cacao) Food, chocola es, bake y, con ec ione y, mixed nu s, medicine - - - 482 405 50-55 Hazelnu Table 3. Global plan oil p oduc ion; cu en demands and en isioned demands beyond 2050 (Alexand a os and B uinsma, 2012). Key Va iables 2005/2007 2050 2080 2100 Popula ion (million) UN 2010 Re ision 6584 9306 9969 10125 Oil c ops (oil equi alen ), ood (kg/capi a) 12.1 16.2 16.9 - Oil c ops (oil equi alen ), all uses (kg/capi a) 21.9 30.5 33.8 - A able land a ea 1592 1661 1630 - Oil c op p oduc ion (million onnes) 149 282 367 - Oil c ops ( o bio uel) (million onne) 7 29 - - Oil c ops ( o bio uel) (% o al uses) 4.8 10.3 - - indus ial and bioene gy demands (Ha egu e al., 2023). The a ious indus ial uses o plan oils and hei de i a i es, such as oleochemicals, a y acids (FA), a y alcohols, and glyce in ac oss di e en indus ies, a e p esen ed in Figu e 2. Mo eo e , indus ial applica ions and he ma ke size o plan oil a e discussed in he subsequen sec ions (Fig. 3). 2.1. Polyme s O e he las ew decades, comme cially a ailable polyme s ha e been de i ed om non- enewable ossil esou ces. The global annual consump ion o polyme s is app oxima ely 300 million onnes, wi h an annual g ow h a e o 5% (Halden, 2010). This signi ican quan i y, along wi h he esul an was e s eams o en disposed o in o a ious ecosys ems Table 4. Compa a i e analysis o plan oil applica ions and esea ch opics in ecen ly published e iew a icles. Re e ence Applica ions Gene ic Enginee ing Global Ma ke Oil-seed P oduc ion Policy and P ac ical Implica ions Indus ial Applica ions Ene gy Biosyn he ic Pa hway Oil Con en Oil Composi ion Oil in Biomass A onso e al. (2023) √ ˟ ˟ ˟ ˟ ˟ ˟ ˟ ˟ Xu e al. (2018) ˟ √ √ ˟ ˟ √ √ ˟ ˟ Sagun e al. (2023) ˟ ˟ √ √ √ √ ˟ √ ˟ Zhou e al. (2023) ˟ ˟ √ √ √ ˟ ˟ ˟ ˟ Wan e al. (2017b) ˟ √ √ √ √ √ ˟ √ ˟ Rau e al. (2023) ˟ ˟ √ √ √ √ ˟ √ ˟ Qi e al. (2020) √ ˟ √ √ √ √ ˟ ˟ ˟ Salehi Jouzani e al. (2018) ˟ √ √ √ √ ˟ ˟ ˟ ˟ Msanne e al. (2020) ˟ ˟ √ √ √ ˟ ˟ √ ˟ P esen Re iew √ √ √ √ √ √ √ √ √ wi hou ea men in many pa s o he wo ld, has led o g owing en i onmen al and heal h conce ns, pa icula ly ega ding he use o pe ochemical-based polyme s. Con e sely, his si ua ion has spa ked a su ge in in e es in biobased polyme s (Des oches e al., 2012). These polyme s a e no only enewable bu also biodeg adable and eco- iendly (Adekunle and Okolie, 2015; Hajinaja e al., 2022a). Biobased polyme s can be syn hesized om polysaccha ides, ibe s, polylac ic acid, and o he ma e ials, wi h iacylglyce ol (TAG) oils and FA also se ing as eliable s a ing ma e ials du ing he p oduc ion p ocess (Acqua ia e al., 2021; Zubai e al., 2021). In 2019, he o al p oduc ion olume o biobased polyme s eached 3.8 million onnes (h ps://www.bioplas icsmagazine.com/en/). Plan oil-based polyme s can unde go ab ica ion using a ious copolyme iza ion echniques, such as ca ionic, ee adical, and he mal me hods, which in ol e combining plan oils wi h a a ie y o pe oleum- based co-monome s (Gogoi e al., 2022; Zhu e al., 2023). Howe e , i is impo an o no e ha mos plan oils equi e modi ica ions a hei na u ally occu ing eac i e si es, such as es e g oups and ca bon-ca bon double bonds, be o e hey can be u ilized in biopolyme p oduc ion (Ike e al., 2021; Ruiz-Rico and Ba a , 2021). The e o e, hese eac i e si es, including he ca bon-ca bon double bonds ound in a y acid chains, play a c ucial ole du ing polyme iza ion (Bie mann e al., 2021; Rajpu e al., 2023). In simple e ms, hese double bonds ac as excellen s a ing poin s o biopolyme p oduc ion. Fo ins ance, con e ing double bonds in FA in o hyd oxyl g oups, ollowed by hei eac ion wi h isocyana e, could esul in he o ma ion o polyu e hanes (Dye e al., 2008). Addi ionally, o he polyme iza ion echniques, such as acyclic me a hesis polyme iza ion (Piccini e al., 2021; Qui ino e al., 2021) and ing-opening me a hesis polyme iza ion (ROMP) (Ganewa a e al., 2021; Ya olimek e al., 2021), ha e been employed o syn hesize plan oil-based polyme s (Ga ison e al., 2016). Ne es e al. (2018) also demons a ed ha modi ied ege able oils con aining ac ylic double bonds exhibi high eac i i y and o m he mose ing biopolyme s h ough ee adical polyme iza ion. Table 5 p o ides examples o comme cially a ailable plan oil-based polyme s and hei eal-wo ld applica ions. Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2110 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Fig. 2. Global ma ke b eak-up o (a) oleochemical, (b) a y acids, (c) a y alcohols, and (d) glyce in by di e en indus ies. Fig. 3. Indus ial applica ions o plan oils and hei ma ke size (Zhou e al., 2020; h ps://www.ma ke sandma ke s.com; and h ps://www.s a is a.com). 2.2. Su ac an s Su ac an s a e amphipa hic compounds ha in luence he su ace o in e acial ene gy o ma e ials (Be g eund e al., 2021). They cons i u e a widely used class o chemicals p edominan ly sou ced om pe oleum (Nag ode e al., 2023). Typically, su ac an s comp ise a hyd ophilic g oup linked o a hyd ophobic moie y (Pola z e al., 2018; Lamch e al., 2020). The hyd ophilic g oups a y depending on whe he he su ac an s a e anionic o ca ionic. Anionic su ac an s encompass ca boxyla e, sul a e, sul ona e, o phospha e g oups, while ca ionic su ac an s comp ise amine o ammonium g oups (Rocky e al., 2023). Su ac an s ind applica ion in bo h edible and non-edible p oduc s, anging om soaps and de e gen s o ood emulsi ie s and cosme ics (Be g eund e al., 2021; De Luca e al., 2021; Mohammed and Ikiensikimama, 2023). As illus a ed in Table 6, su ac an s de i ed om pe ochemicals ha e aised signi ican heal h and en i onmen al conce ns due o hei oxicological p ope ies. Consequen ly, sa e al e na i es, known as biosu ac an s, ha e ga ne ed conside able a en ion, pa icula ly in he ood and cosme ics indus ies. Indeed, eco- iendly al e na i es, like biosu ac an s, o e compa able physicochemical p ope ies (such as emulsi ica ion, de-emulsi ica ion, oaming, and we ing) o hei pe oleum-de i ed coun e pa s (Ahmadi- Ash iani e al., 2020; Sa ubbo e al., 2022). They also possess se e al ad an ages, including lowe oxici y, biodeg adabili y, and inc eased esis ance ac oss a b oade ange o pH, salini y, and empe a u e condi ions (Abbo e al., 2022; Sa ubbo e al., 2022). Oilseed c ops a e conside ed p omising eeds ocks o biosu ac an p oduc ion. Speci ically, he FA p esen in plan oils o hei co esponding me hyl es e s can be educed o p oduce a y alcohols, which a e hen u ilized in biosu ac an o mula ions (Van Ren e ghem e al., 2018). Despi e hei a o able en i onmen al a ibu es, biosu ac an s s ill p esen less a o able economic cha ac e is ics compa ed o su ac an s de i ed om pe oleum (Gau e al., 2022; Joshi e al., 2022). This issue is Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2111 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 5. Examples o comme cially a ailable plan oil-based polyme s (Des oches e al., 2012; McKeon e al., 2016; Zhang e al., 2017). Plan oil Sou ces P oduc s T ade name Company Applica ions Soybean oil Polyme ized Soybean Oil (polySOY) - - - Ac yla ed Epoxidized Soybean Oil (AESO) Ebec yl 860 UCB Chemicals Company, Ad en In e na ional (Bos on, MA, USA) Su ace coa ings Maleic acid eac ed AESO (MAESO) also “malea ed ac yla ed epoxidized soy oil (MAESO)” - - Shee molding compound Soybean oil monoglyce ide (SOMG), also “maleina ed soybean oil monoglyce ide” - - - The mose s p epa ed by he ca ionic copolyme iza ion o soybean Oil - - - Polyme ized epoxidized soybean oil (ESO) Plas iSoy™ CHS (USA), Makwell (India), The Chemical Company (USA), Mul iPlus (Thailand), PolyMa En e p ises (USA), FMC (USA) Rubbe s, Resins, Coa ings, Pain s, Plas icize s, Adhesi es, Polyols (polyu e hanes), The mose s Epoxidized Soybean Oil Viko lex® A kema (USA) - No bo nenyl- unc ionalized a y alcohols de i ed om soybean oil (NMSA) - - - Polyu e hanes BiOH® Ca gill (USA) - Ag ol® BioBased Technologies (USA) Lub ican s, Building P oduc s, P in ing Inks, Diesel Addi i es, Coa ings, Fu ni u e, Adhesi es, Au omo i e, Ag icul u al P oduc s Renu a® Dow Chemical (USA) Adhesi e, Con en ional Flexible Polyu e hane Foam (FPF), Viscoelas ic Foam, High Resilience Foam (HR), Molded Foam So e mol® polyols BASF (Cogins) Oleochemicals (Malesia) Adhesi es, Binde s, Floo coa ings, Cas ings, Elec opla ing Linseed oil Linseed Oil Monoglyce ide (LOMG) - - - Polyme ized Linseed Oil (Linoleum) Lino ille, Ma moleum®, Fo bo's Topshield™, NATURCo e™, Fo bo (Swi ze land), A ms ong (USA), To lys (Newzeland) Floo co e ing Polyu e hane (Boiled Linseed Oil (BLO)) C own® Boiled Linseed Oil W. M. Ba , USA; C own (USA) Wood inishe Epoxidized linseed oil HiBond® Pola Indus ies (Canada) Pain s, plas icize s, adhesi es, coa ings, o any applica ion o an epoxidized oil Viko lex A kema (USA) - Cyclopen adiene polyme s DilulineTM Ca gill (USA) D ing oil ML189 A che Daniel Midland (USA) Va nish, Enamel, Aluminum pain , Rein o ced oil Cas o oil Malea ed alcoholized cas o oils (MACOs) - - - 2-(Ac yloyloxy) E hyl Olea e (AEO) HeloxyTM Flexibilize Momen i e Special y Chemicals (USA) Epoxy Resin, Coa ing, Cons uc ion, Composi es, Adhesi es, Elec ical cas ings, Elec ical lamina es and Fibe s Es olides Heloxy™ Modi ie s48 Hexion (USA) - ERISYSTMGE-35H Eme ald Pe o mance Ma e ials (USA) Conc e e Pa ching Compounds, Floo Coa ings, Adhesi es, B idge Decking Compounds, Join Sealan s Vo i e®Polyme ized Cas o Oil Ve ellus (USA) Elas ome s, Adhesi es, Coa ings, Inks, Polyols, Sealan s Zenigloss ® Zeni ech (Canada) Emollien , Lip gloss agen , Pe sonal ca e applica ions Polyglyce ol poly icinolea e (PGPR) F aken Biochemical (China), Spell O ganics (India) Food emulsi ie / ex u e and iscosi y con ol (Chocola e) Polyamides Rilsan®PA11, Rilsan®Fine Powde , A kema (USA, F ance) Elec ical cable, uel line, luid ans e , quick connec o s, as ene s and clips, ic ion pa s, pneuma ic and hyd aulic hose Polyu e hane Ul amid® BALANCE BASF (USA) Au omo i e Ag ol S a ™ BioBased Technologies (USA) Inks and Coa ings Lup anol® Balance 50 BASF (Ge many) Foam - Jayan Ag o O ganics Limi ed (India) - Polyme Pebax Rnew® A kema (USA, F ance) Spo s, Medical, Packaging and Indus ial applica ions EcoPaXX™ DSM Au omo i e and Elec ical ma ke s Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2112 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 6. Toxici y o di e en su ac an s agains a ious o ganisms (I anco ic and H eno ic, 2010; Cowan-Ellsbe y e al., 2014; Yuan e al., 2014; Theis e al., 2016; Badmus e al., 2021). Su ac an G oup Pe ochemical su ac an s Toxicological ea u es o pe ochemical su ac an s Plan oil-based al e na i es Applica ion Anionic Linea alkylbenzene sul ona es (LAS) Bac e ia: Vib io ische i (EC50-Luminescence 30 min: 2.6 mg/l) Pseudomonase pu ida (EC50-G ow h inhibi ion 16 h: 33.4 mg/l) Algae: Dunaliella sp (EC50-24 h: 3.5 mg/l) C us aceans: Ce iodaphnia dubia (EC50- Immobiliza ion 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: Ca assius au a us (EC50- Immobiliza ion 48 h: 5.1 mg/l) Salmo gai dne i (Immobiliza ion EC50– 48 h, 33.61 mg/l) Gammbusia a inis (mosqui o ish) Immobiliza ion EC50-48 h, 40.15 mg/l Ca assius au a us (gold ish) Immobiliza ion EC50-48 h, 38.04 mg/l Plan : Bush beans, adish and g asses: Yield and g ow h NOEC–76 days, 27 mg/kg Po a o: Yield and g ow h NOEC–106 days, 16 mg/kg B and Name: Eu asol (Ammonium olea e, Po assium alla e, Mixed a y acid sal s, Po assium cocoa e, Po assium palma e) EOC Co. Belgium B and name: SERVO® BRILLANT (cas o oil sulphona e) Elemen is Co. The Ne he lands As de e gen s, oaming agen s, emulsi ie s, an is a ic agen s, dispe san s, s abilize s in he amily and chemical aspec s o li e, Cosme ic, Pha maceu ical and Pe ochemical p oduc s Linea e he sul a e - Oc ylphenol polyoxye hylene sodium - Soaps Alga: EC50: 10-50 mg/L Sodium dodecyl sulpha e (SDS) Bac e ia: Vib io ische i (EC50- Luminescence 15 min: 2.6 mg/L) Algae: Raphidocelis subcapi a a (IC50 - Cell densi y 72 h: 36.58 mg/L) C us aceans: A emia salina (LC50 - La ae mo ali y 24 h: 41.04 mg/L) Gas opod: Physa acu a (LC50 - Mo ali y 24 h: 27.2 mg/L) Sea u chin: Pa acen o us li idus (EC50- Fe iliza ion a e: 3.2 mg/L) Fish: Gammbusia a inis (EC50- Immobiliza ion 48 h: 13.64 mg/L) Salmo gai dne i ( ainbow ou ) Immobiliza ion (EC50– 48 h, 10.84 mg/L) Ca assius au a us (gold ish) Immobiliza ion (EC50– 48 h, 12.35 mg/L) Alkyl sulpha e (AS) - Sodium lau yl sulpha e (SLS) - Alkyl e hoxysulpha e (AES) Algae: Pseudoki chne iella subcapi a a (EC50 - Cell densi y 72 h 3.5 mg/L) Raphidocelis subcapi a a (IC50 - Cell densi y 72 h 2.18 mg/L) C us aceans: A emia anciscana (LC50- Nauplii mo ali y 72 h 23.92 mg/L) Fish: Salmo gai dne i (EC50- Immobiliza ion 48 h 10.84 mg/L) Seconda y alkane sulphona es (SAS) - Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2113 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 6. con inued. Su ac an G oup Pe ochemical su ac an s Toxicological ea u es o pe ochemical su ac an s Plan oil-based al e na i es Applica ion Non-ionic Polyoxye hylenes ea yl e he (20EO) Wa e lea: EC50: 48 mg/L B and Name: Eu Amid (cocamide die hanolamine, monoe hanolamine); Eu oxide (Cocamine oxide, Cocamidop opylamine Oxide, EOC Co. Belgium B and Name: SERDOLAMIDE (coconu oil die hanol amide (Supe - amide), oleic acid die hanol amide (K i che sky), soya oil die hanol amide in TEA/wa e (Supe -amide)), SERDOX® (oleic acid monoe hanol amide), ECOSURF™ SA Su ac an s (Dow co, USA), EcoSense™ Su ac an s (DOW company, USA) Tex ile, pape , ood, plas ic, glass, ibe , medicines, pes icides, dyes, o he indus ies, emulsi ie s, we ing agen s, and oam s abilisa ion agen s, in a ious bio echnological p ocesses, and o acili a e solubilisa ion and inc ease d ug ca ie s abili y Polyoxye hylenes ea yl e he (10EO) Aus alian na i e ogs: Full na cosis EC50– 48 h, 2.8– 3.8 mg/L Fish: Fa head minnow: LC50– 96 h, 4.6 mg/L Lau yl alcohol e hoxyla es(7EO) - Nonylphenole hoxyla es (9EO~11EO) (NPE) C us aceans: Daphnia magna LC50– 48 h, 14 mg/L Fish: Pimphales p omelas ( a head minnow): LC50– 10 d, 2.7 mg/L Fa head minnow LC50– 96 h, 4.6 mg/L E ec s on ep oduc i e heal h o ish: NP induce he p oduc ion o i ellogenin in male ish, a p o ein usually only ound in sexually ma u e emales unde he in luence o es ogens Alkylphenol e hoxyla e (APE) - Oc yl phenol e hoxyales (OPE) E ec s on ep oduc i e heal h o ish:OP induce he p oduc ion o i ellogenin in male ish, a p o ein usually only ound in sexually ma u e emales unde he in luence o es ogens Alcohol e hoxyla e (AE) Bac e ia: Mic ocys is ae uginosa (Es ima ed EC10 - Cell densi y 0.154 mg/L) Algae: Lemna mino (Es ima ed EC10 - F ond coun 0.101 mg/L) Na icula pelliculosa (Es ima ed EC10 - Cell densi y 0.140 mg/L) C us aceans: Ce iodaphnia dubia (EC50 - Immobiliza ion 48 h 0.39 mg/L) Fish: Pimephales p omelas (NOEC - Su i al 4.35 mg/L) Fa y acid e hoxyla e (FAE) - Ca ionic Ce yl ime hyl ammonium chlo ide - - S e iliza ion, us , co osion, b eaking, co osion and mine al lo a ion, de e gen s, ab ic so ene s, and hai condi ione s Qua e na y ammonium compound (QAC) QACs a e oxic o mammalian cells and a e no ecommended o sys emic applica ion, damaging e ec s o ca ionic su ac an s on human lymphocy es Bac e ia: Vib io ische i (EC50- Luminescence 30 min 0.5 mg/L) Pseudomonas pu ida (EC50 - G ow h inhibi ion 16 h 6.9 mg/L) Algae: Dunaliella sp. (EC50 - 24 h 0.79 mg/L) C us aceans: Daphnia magna (EC50- Immobiliza ion 24 h 0.38 mg/L) Fish: Salmo gai dne i (EC50- Immobiliza ion 48 h 1.21 mg/L) Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2120 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 10. Resea ch s udies on using plan -based pes icides on a ious o ganisms. Re e ence Plan Oil Oil concen a ion Applica ion Ta ge ed Species Time (h) Mo ali y /Inhibi o y (%) Resul s Ka e sou e al. (2019) Encapsula ed Men ha pulegium EO 0.10% Used in yeas cell mic oca ie s Insec pes Myzus pe sicae 24 5 • Encapsula ed-oil showed highe insec icidal ac i i y han non-encapsula ed one. 48 5 Non-encapsula ed M. pulegium EO 0.10% 24 0 48 5 Vasan ha- S ini asan e al. (2018) Be el lea oil 500 mg/kg In soil agains ea hwo m and edwo m Eud ilus eugeniae (Kinbe g) 168 ≈1.5 • Inno a i e and sa e insec icides o soil- dwelling c ea u es. • The enzyme le el was no a ec ed by he essen ial oil o P. be le. Thus, he plan - de i ed ola ile oil did no a ec biochemical eac ions 336 ≈1.2 1000 mg/kg 168 ≈3.5 336 ≈2.5 500 mg/kg Eisenia e ida (Sa igny) 168 ≈1.9 336 ≈1.5 1000 mg/ kg 168 ≈3.2 336 ≈2.5 Taba i e al. (2017) α- hujone- ich A emisia siebe i essen ial oil 2 μg/cm3 Poul y indus y De manyssus gallinae (De manyssida) 24 3.22 • P olonged oxici y o he oil. • On adul s o he poul y ed mi e, D. gallinae, - hujone- ich A. siebe i essen ial oil showed p omising oxici y and epellan ac ion. 5 μg/cm3 10.88 10 μg/cm3 25.77 Adak e al. (2020) Eucalyp us 1 μg/cm3 Rice Si ophilus o yzae 24 55 • Downsized essen ial oils ha e highe insec icide po en ial han no mal size. • Nanoemulsion eucalyp ol can imp o e e iciency and educe he cos o essen ial oils. 2 μg/cm3 100 3 μg/cm3 T ibolium cas aneum 17 7 μg/cm3 58 Klein e al. (2020) Thyme 0.148% ( / ) Labo a o y and g eenhouse bioassays De oce as e icula um 24 50 • By 1% ( / ), a e 24 hou s, he mo ali y by all ypes o essen ial oils eached 100%, showing ha hese plan ex ac s migh be usable as new a ional molluscicides. • Thyme showed he highes pe o mance as a pes icide han o he oils. • Al hough hyme p o ed bene icial agains he a ge ed species, he open en i onmen could in luence he esul s. 0.26% ( / ) 99 Ga lic 0.204% ( / ) 50 0.329% ( / ) 99 Rosema y 0.307% ( / ) 50 0.554% ( / ) 99 Lemong ass 0.32% ( / ) 50 0.72% ( / ) 99 Cinnamon Cassia 0.42% ( / ) 50 0.799% ( / ) 99 Idoko and Ileke (2020) A amomum melegue a 1 mL/L Cowpea seeds Callosob uchus macula us 12 13.33 • P o ec s o ed cowpeas agains C. macula us. • The in luences o essen ial oils on C. macula us mo ali y inc eased wi h he exposu e ime and ea men a es. • The signi ican mo ali y induced by essen ial oils may be owing o he essen ial oils’ su oca ing odo , which mus ha e dis u bed he insec s’ egula espi a o y mechanism. 5 mL/L 26.67 Annona mu ica a 1 mL/L 13.33 5 mL/L 16.67 Eucalyp us globules 1 mL/L 13.33 5 mL/L 20 Ficus exaspe a e 1 mL/L 16.67 5 mL/L 20 Te apleu a e ap e a 1 mL/L 6.67 5 mL/L 20 Papadimi iou e al. (2019) Pulegone ex ac ed om M. pulegium 500 µL/L Cucumbe Toma o Te anychus u icae emales 72 47 • Inco po a ing hese wo EOs did no in luence he mo ali y a e o help ul insec s (Nesidioco is enuis). • No phy o oxic in luences we e eco ded a e using EOs on he plan s. 1000 µL/L 51 Pipe i one ex c ed om M. pulegium 500 µL/L 65 1000 µL/L 71 Janaki e al. (2018) Cype us o undus 0.4 μl/cm2 • Cowpea • Chickpea • Da es C. macula us 24 64 • The used EO exhibi ed a epellen e ec on all h ee kinds o insec s examined in he s udy. • The epellen e ec in O. su inamensis and T. g ana ium was mo e han in C. macula us. 1 μl/cm2 88 0.4 μl/cm2 T ogode ma g ana ium 66 1 μl/cm2 77 0.4 μl/cm2 O yzaephilus su inamensis 44 1 μl/cm2 72 Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2121 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 11. Use o ege able oils in coa ing ma e ials and hei impo an p ope ies. Re e ence Plan oil Oil con en (%) o concen a ion Pu pose Time (d) ( ac o ) Impo an p ope y In luence on he p ope y Impo an in o ma ion Dong and Wang (2017) Ga lic essen ial oil 1 The in luence o essen ial oils coa ing o imp o ing he quali y o s awbe ies ha con ain ca boxyme hyl cellulose 3 (s o age ime) Decay pe cen age (%) ≈-13.5% • Using ca boxyme hyl cellulose in conjunc ion wi h ga lic essen ial oil imp o ed decay pe cen age, weigh loss, o al soluble solids, i a able acidi y, and asco bic acid con en , as well as e aining g ea e o al phenol and an hocyanin concen a ions in s awbe ies. To al phenol conc.(mg/g) ≈0.3 6 (s o age ime) Decay pe cen age (%) ≈-40% To al phenol conc.(mg/g) ≈0.12 3 3 (s o age ime) Decay pe cen age (%) ≈-12.5% To al phenol conc.(mg/g) ≈0.37 6 (s o age ime) Decay pe cen age (%) ≈-42% To al phenol conc.(mg/g) ≈0.21 Alo aibi and Tahe go abi (2018) Thyme essen ial oil 2 Swee po a o s a ch based-coa ing o e ige a ed s o age wi h essen ial oil and s udying he in luence on he sh imp quali y 4 Ha dness (N) 5.8 • Signi ican educ ion in he bac e ial popula ion by hyme. • A swee po a o s a ch-based coa ing inco po a ed wi h hyme essen ial oil migh be a easible op ion o p ese ing sh imp mea quali y and educing losses. Resilience -0.03 4 Ha dness (N) 7.06 Resilience -0.01 Fe nández e al. (2020) Ci onellol 2 (mg/mL) De elop an an i ungal hyb id ille o coa ings ha is bo h en i onmen ally iendly and a o dable 10 C. globosum inhibi ion (%) 77 • Plan ex ac s con aining EOs can be u ilized as an i ungal agen s. • The main ac i e componen was ound o be ci onellol. • P oducing unc ional bioac i e hyb ids was he bene i o blending ci onellol. A. al e na a inhibi ion (%) 78 5 (mg/mL) C. globosum inhibi ion (%) 96 A. al e na a inhibi ion (%) 98 Ci al 2 (mg/mL) C. globosum inhibi ion (%) 75 A. al e na a inhibi ion (%) 75 5 (mg/mL C. globosum inhibi ion (%) 98 A. al e na a inhibi ion (%) 98 Vi al e al. (2018) Ginge 0.10% Accep abili y o ish ille wi h an algina e- based coa ing con aining essen ial oils 7 Shea o ce (N) -1.85 • Reduc ion in colo losses and` lipid oxida ion o ish ille . • The an ioxidan ac i i y inc eased. • Using hese addi i es is an e ec i e way o elimina e undesi able p ope ies o he ood. Weigh loss (%) -0.03 pH -0.04 O egano Shea o ce (N) 1.58 Weigh loss (%) 0.18 pH -0.06 Buendía e al. (2020) Ca ac ol:o egano: cinnamon (70:10:20) ≈1% Coa ing o ca dboa d, including EO en apped wi hin cyclodex ins nano ube 6 TA (% ci ic acid) 0.001 • The lowes decay incidences and he highes i mness we e o cyclodex ins nano ube blended wi h EOs. • Tha blend showed he highes an imic obial ac i i y compa ed o samples wi hou plan oil. • The changes in pH we e negligible. • The addi i es did no in luence he physicochemical quali y o he p oduc ( oma oes). Fi mness (N) 3.2 6 wi h a supplemen a y comme cializa ion pe iod TA (% ci ic acid) 0.015 Fi mness (N) 1.5 Coa ing o ca dboa d including EO en apped wi hin cyclodex ins halloysi e nano ube 6 TA (% ci ic acid) -0.01 Fi mness (N) -2.7 6 wi h a supplemen a y comme cializa ion pe iod TA (% ci ic acid) -0.029 Fi mness (N) 2.7 Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2122 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 11. con inued. Re e ence Plan oil Oil con en (%) o concen a ion Pu pose Time (d) ( ac o ) Impo an p ope y In luence on he p ope y Impo an in o ma ion Klangmuang and So ho n i (2018) Plai 15g/L Inco po a ing essen ial oils in o hyd oxyp opyl me hylcellulose-based nanocomposi e on mango (c . Namdokmai Si hong) agains an an h acnose disease 15 Weigh loss (%) F om ≈9 o ≈7.4 • Inco po a ing essen ial oils in o he ma ix inhibi ed C. gloeospo ioides ungus. • Ginge showed he highes pe o mance. • Essen ial oils did no in luence he as e, la o , and quali y o mango and ex end he ui ’s shel li e. Disease se e i y (sco e) F om ≈3 o ≈2.2 Fi mness (N) F om ≈1.95 o ≈7 Colo F om ≈4.35 o ≈4 Ginge Weigh loss (%) F om ≈9 o ≈7.8 Disease se e i y (sco e) F om ≈3 o ≈1.9 Fi mness (N) F om ≈1.95 o ≈6.8 Colo F om ≈4.35 o ≈3.7 Majdinasab e al. (2020) Shi azi hyme 2% The in luence o used EOs on an imic obial and an ioxidan coa ing and shel -li e ex ension o chicken ille 10 Cooking loss (%) -15.5 • The change in colo by hyme was less han summe sa o y. • The shel -li e o chicken ille s is ex ended. • The o e all accep abili y was mo e han he con ol (Basil-seed gum) when hey used EOs, and hyme in luence was highe han summe sa o y. O e all accep abili y F om 2.4 o 3.8 Tex u e F om 2.4 o 4.4 Summe sa o y Cooking loss (%) -13.5 O e all accep abili y F om 2.4 o 3.5 Tex u e F om 2.4 o 4.3 p in ing inks, candles, and polishes due o hei dis inc i e p ope ies (Zhuko and Popo , 2022). The global wax and wax es e ma ke s, pa icula ly in he UK, he USA, B azil, Japan, and o he coun ies, a e highly appealing and an icipa ed o g ow a a a e o 3.83, eaching USD 15.91 billion by 2030 (Fig. 3) (Ve i ied Ma ke Resea ch, 2023). While some plan s na u ally p oduce wax es e s in hei seed oil, like jojoba, hey a e unsui able o la ge-scale cul i a ion as hey yield an undesi able mix u e o e y long-chained wax es e s un i o echnological applica ions (Langsdo e al., 2021; Simonsen e al., 2023). Howe e , nume ous o he plan s show p omise in p oducing wax es e s, such as sun lowe seed, oli e, palm, Camelina sa i a, and A abidopsis haliana, exhibi ing high pe o mance (Qi e al., 2020; Clews e al., 2023). Oli e and palm oil-based wax es e s a e pa icula ly no ewo hy, wi h hose de i ed om oli e oil consis ing o long, s aigh -chain a y alcohols es e i ied wi h FA (Abdelmoez and Mus a a, 2014; Ma iani e al., 2018). Se e al ac o s in luence he a e o es e i ica ion in oli e oil, including e ining p ocess a iables, s o age condi ions, and eagen concen a ion (Ma iani e al., 2018; Dia e e al., 2021). 2.12. Tex ile inishing Washing, bleaching, dyeing, and coa ing cons i u e he p ima y s ages o he ex ile inishing p ocess, u ilized o enhance he bulk o ex iles o ga men s pos -wea ing and syn he ic ma e ial manu ac u ing (Achaw and Danso-Boa eng, 2021; Al-Sayed and Abdel ahman, 2021). Tex ile inishing plays a c ucial ole in de e mining he inal appea ance and aes he ic quali ies o ex iles while also impa ing desi able p ope ies such as lame e a dancy, w inkle esis ance, wa e -and-oil epellency, and mo e (Haule and Nambela, 2022; Ja aid e al., 2024). O e he p ojec ed pe iod (2017-2024), he global ma ke o ex ile inishing chemicals is o ecas ed o g ow a a CAGR o 3.8%, ising om USD 8.9 billion in 2022 o USD 12 billion in 2030 (P escien and S a egic In elligence, 2023). Howe e , a signi ican po ion o ex ile inishes cu en ly a ailable in he ma ke pose po en ial ha m o he en i onmen and human heal h, wi h compounds like iclosan known o accumula ing in aqua ic en i onmen s and posing oxici y isks o aqua ic o ganisms (Bha e al., 2022; Pe iyasamy, 2023). In ligh o hese conce ns, plan oils p esen a sus ainable al e na i e o use as addi i es in ex ile inishing p ocesses (Na a ajan e al., 2022). One no able applica ion is hei use as insec epellen s, le e aging hei na u al p ope ies o epel insec s, along wi h hei u iliza ion in a oma he apy ex iles o an ibac e ial, ol ac o y, and medicinal pu poses (Mi al e al., 2019). I is impo an o no e ha an imic obial ex iles, while e ec i e, main ain eco- iendly c eden ials, making hem a o able in con empo a y con ex s, pa icula ly amids he COVID-19 c isis, which has unde sco ed he impo ance o con olling in ec ious diseases (Bouaziz e al., 2021). Beyond ex ile applica ions, plan oils se e a ious indus ies. In medicine, hey demons a e po en an i ungal, an imic obial, an icance , and wound-healing p ope ies. In he ood and ood packaging sec o s, hey unc ion as na u al an i ungal and an ibac e ial agen s. Addi ionally, in he cosme ics and pe ume indus ies, hei a oma ic quali ies make hem in aluable ing edien s. Plan oils also con ibu e o mining p ocesses h ough o e o h lo a ion and se e as su ac an s in oil-well d illing muds. Mo eo e , in ubbe p oduc ion, plan oils se e as ulcanizing agen s, so ene s, and mold elease agen s. They also ind use as sol en s and an i- dus agen s, and hey aid in pape ecycling by e icien ly emo ing p in ing inks. Subsequen sec ions will del e in o plan oil p oduc ion pa hways and a y acid syn hesis. 3. Plan oil p oduc ion pa hways Plan oils p ima ily consis o TAGs and a e p edominan ly s o ed in he seeds o mesoca p o ui s (Ge e al., 2021; He nández e al., 2021). Howe e , no able excep ions exis , such as Simmondsia chinensis (jojoba), whe e oils accumula e in he o m o es e s o long-chain alcohols and FA (Guzha e al., 2023). F om a chemical s andpoin , TAGs esul om he es e i ica ion o FA (C8–C24) wi h glyce ol (Wei e al., 2024). I is in iguing o no e he close simila i y be ween he gene ic chemical o mulas o hese FA and hose o ossil-o ien ed hyd oca bons, i.e., CH3(CH2)nCOOH s. CH3(CH2)nCH3, espec i ely (Rajaei a e al., 2019). This simila i y has spa ked widesp ead in es iga ions in o using plan oils and hei de i a i es as al e na i es o ossil uels. O e all, he p ope ies o plan oils and hei applica ions a e la gely a ibu ed o hei a y acid composi ion (Mannu e al., 2020). Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2123 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. I is impo an o acknowledge ha while palmi ic acid (16:0), s ea ic acid (18:0), oleic acid (18:1), linoleic acid (18:2), and linolenic acid (18:3) a e he majo FA ound in plan oils (Wei e al., 2019; Pio esana e al., 2021), he e is ema kable di e si y among di e en plan species in e ms o hei oil composi ions. Fo ins ance, he e a e o e 300 a ia ions in a y acid p o iles in highe plan s (Liu e al., 2022b). Mo eo e , some plan s p oduce uncommon FA, u he en iching plan oil composi ions. Examples o such uncommon FA include sho ca bon chain FA (C8 o C14) in Cuphea, long ca bon chain FA (C20 o C24) in ape, C ambe, B. napus, and A. haliana, and hyd oxy a y acid (C18:0-OH) in cas o bean (R. communis), among o he s (Roscoe e al., 2015). F om an applica ion s andpoin , hese FA impa speci ic cha ac e is ics ha make hese oils sui able o use ac oss a ious indus ies. Consequen ly, such plan s o e unique gene pools ha could be le e aged o modi y o manipula e con en ional oil c ops. To achie e his objec i e, i is c ucial o gain in-dep h insigh s in o he me abolic pa hways go e ning plan oil p oduc ion, including he in ol ed genes and hei egula o y mechanisms. The e o e, a y acid syn hesis and modi ica ion (elonga ion and desa u a ion), as well as TAG syn hesis and accumula ion, a e p esen ed and discussed in he subsequen sec ions. 3.1. Fa y acids syn hesis The syn hesis o FAs, p ima ily occu ing in plas ids, se es as he co ne s one o plan oil p oduc ion pa hways. As illus a ed in Figu e 4, he p ocess commences wi h he ca boxyla ion o ace yl-CoA, leading o he o ma ion o malonyl-CoA. I is no ewo hy ha ace yl-CoA is syn hesized di e en ly in pho osyn he ic (e.g., lea es) and non-pho osyn he ic (e.g., ui s and seeds) plan issues (Li-Beisson e al., 2016). In pho osyn he ic issues, ace yl-CoA is p oduced in chlo oplas s oma h ough he ixa ion o CO2 in o iose-phospha es (TP) ia he Cal in cycle (Fig. 4a). These TPs a e subsequen ly con e ed in o py u a e and e en ually ace yl-CoA h ough he glycolysis pa hway (Tang e al., 2022). Al e na i ely, TPs may be di ec ed in o he s a ch syn hesis pa hway (Xu e al., 2024) (Fig. 4a). In con as , suc ose ac s as he p ecu so o ace yl-CoA syn hesis in non- pho osyn he ic issues. Howe e , as suc ose canno en e plas ids whe e ace yl-CoA is syn hesized, i unde goes clea age in o i s cons i uen componen s by in e ases o suc ose syn hases. The esul ing hexoses a e hen con e ed in o hexose phospha es (i.e., glucose 6-phospha e and uc ose 6-phospha e) (Xu e al., 2024). These hexose phospha es unde go Fig. 4. O e iew o majo eac ions in ol ed in a y acid and iacylglyce ol syn hesis. (a) Glycolysis is used o make p o eins, s o age s a ches, o lipids om abso bed ca bohyd a es om pho osyn hesis. Suc ose is anspo ed om pho osyn he ic issues in o de eloping seeds, whe e i is me abolized in o p ecu so s such as glucose 6-phospha e and phosphoenolpy u a e in he cy osol o emb yo and/o endospe m cells be o e being ans e ed o plas ids o a y acid syn hesis.; (b) De no o a y acid p oduc ion and modi ica ion: he p ecu so s o he syn hesis o C8–C18 sa u a ed a y acyl-ACPs on a plas idial mul ienzyme a y acid syn he ase complex include ace yl-CoA and malonyl-CoA. Unsa u a ed and monounsa u a ed a y acids a e anspo ed om plas ids o he endoplasmic e iculum o u he p ocessing by an acyl-CoA anspo e (ACT). (c) TAG syn hesis. A complex p ocess combining successi e acyla ion o a glyce ol moie y and subs an ial acyl edi ing ia phospha idylcholine-dependen desa u ases o desa u ase-like enzymes esul s in iacylglyce ols. Abb e ia ions: TP: T iose phospha e; G6P: Glucose 6- phospha e; OPP: Oxida i e pen ose phospha e; Py , py u a e; ACCase, ace yl-CoA ca boxylase; FAS, FA syn hase; FA, a y acid; FFA, F ee Fa y Acid; CoA, coenzyme A; LACS, Long Chain acyl-CoA Syn he ase; G3P, Glyce ol 3-phospha e; GPAT: G3P acyl ans e ase; LPA: lysophospha idic acid; LPAAT: lysophospha idic acid acyl ans e ase; PA: phospha idic acid; PP: phospha ase; DAG: diacylglyce ol; DAGAT: DAG acyl ans e ase; TAG: iacylglyce ol; DGAT: diacylglyce ol acyl ans acylase; PDAT: phospha idylcholine-dependen acyl ans e ase; PDCT: phospha idylcholine: diacylglyce ol choline phospho ans e ase; PC: phospha idylcholine; PLA2: phospholipase A2; LPC: Lys phospha idylcholine; LPCAT: Lys phospha idylcholine acyl ans e ase; DES: Desa u ase; CPT: CDP- choline: DAG choline phospho ans e ase. Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2124 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. u he con e sion in o a ious in e media es, such as phosphoenolpy u a e and mala e, h ough he glycolysis and/o he oxida i e pen ose phospha e (OPP) pa hway be o e being anspo ed in o plas ids o ace yl-CoA p oduc ion (Seliniski and Scheibe, 2019; Bu e al., 2023). In he subsequen s ep, he pi o al eac ion ca alyzed by ace yl-CoA ca boxylase (ACCase) esul s in he o ma ion o malonyl-CoA, he building block o he FA syn hesis pa hway, ia ace yl-CoA ca boxyla ion (Fig. 4b). The a y acid syn hase enzyme complex (including 3-ke oacyl- ACP syn hase o ype III, ype I, and ype II, o KASIII, KASI, and KASII, espec i ely, as well as acyl-ACP hioes e ases, FAT A o FAT B) hen u ilizes malonyl-CoA o syn hesize FAs anging om 4 o 18 ca bons in leng h (Ba es e al., 2013). Ini ially, KASIII ca alyzes he condensa ion, yielding C4:0 FAs. Subsequen condensa ion eac ions a e ca ied ou by KASI, p oducing FAs up o C16:0. Finally, KASII acili a es he inal s eps o FA elonga ion om C16:0 o C18:0 (Manan e al., 2017). FAT A o FAT B plays a c ucial ole in de e mining he chain leng h o he syn hesized FAs wi hin he enzyme complex (Liu e al., 2022a and b). No ably, he accumula ion o medium-chain FAs, such as C10:0 and C12:0, in ol es he p esence o e ol ed FAT A o FAT B, capable o p ema u e hyd olysis o he g owing acyl hioes e s. Conside ing he signi ican ole o ace yl-CoA in he o e all FA syn hesis pa hway, he eac ion i is in ol ed in is ega ded as a majo a e-limi ing s ep (He e al., 2020). Fu he mo e, FA anging om C8:0–C18:0 may unde go unsa u a ion ca alyzed by a y acid desa u ases (FADs), speci ically FAD6, 7, and 8 (Manan e al., 2017), while s ill wi hin plas ids (Elahi e al., 2016). The esul ing sa u a ed and monounsa u a ed FAs in he C8–C18 ange a e anspo ed as acyl-CoAs o he endoplasmic e iculum (ER) o u he modi ica ion and elonga ion (Bu e al., 2023; Fell e al., 2023). Con inued FA elonga ion in he ER in ol es a se ies o eac ions u ilizing cy osolic malonyl-CoA and impo ed acyl-CoA (Bu e al., 2023; Fell e al., 2023). These eac ions a e ca alyzed by an ER-speci ic a y acid elonga ion complex comp ising ou enzymes: 3-ke oacyl-CoA syn hase, 3-ke oacyl- CoA educ ase, 3-hyd oxyacyl-CoA dehyd ase, and enoyl-CoA educ ase (Bu e al., 2023; Fell e al., 2023). This enzyma ic complex acili a es he p oduc ion o FAs up o C24. Addi ionally, Fa y acid desa u ase 2 (FAD2) and Fa y acid desa u ase 3 (FAD3) a e esponsible o sequen ial desa u a ion eac ions occu ing in he ER, leading o he o ma ion o polyunsa u a ed FA (Manan e al., 2017). 3.2. T iacylglyce ol biosyn hesis TAG syn hesis gene ally occu s h ough a se ies o eac ions known as he Kennedy pa hway, loca ed in he endoplasmic e iculum (ER) (Kim, 2020) (Fig. 4c). This pa hway ini ia es wi h he in oduc ion o glyce ol-3- phospha e (G3P), which is sequen ially acyla ed by acyl-CoAs o chlo oplas o igin (Schmid, 2021). G3P is gene a ed om dihyd oxyace one phospha e (DHAP) h ough a eac ion ca alyzed by glyce ol-3-phospha e dehyd ogenase (G3PDH) (Kim, 2020). The enzymes in ol ed in his sequen ial acyla ion p ocess include glyce ol-3-phospha e acyl ans e ase (GPAT), lysophospha idic acid acyl ans e ase (LPAAT), and diacylglyce ol acyl ans e ase (DGAT) (Li e al., 2015a). P io o he hi d acyla ion by DGAT, a dephospho yla ion s ep is necessa y, which is ca alyzed by phospha idic acid phospha ase (PAP) (Kim, 2020; Lu kewi e and Finck, 2020). I is no ewo hy ha TAG biosyn hesis may in ol e addi ional complexi ies beyond he adi ional Kennedy pa hway. Fo ins ance, acyl- CoAs could also be supplied om memb ane lipid phospha idylcholine (PC) h ough al e na i e pa hways. Acyl-CoAs may be inco po a ed o eleased om PC h ough a se ies o eac ions known as acyl edi ing eac ions, ca alyzed by lyso-PC acyl ans e ase (LPCAT). Al e na i ely, a eac ion ca alyzed by phospholipase A2 (PLA2) may esul in he elease o ee FA, which can hen be acyla ed o o m acyl-CoA (Lu kewi e and Finck, 2020; Schmid, 2021) (Fig. 4c). Ano he al e na i e pa hway complica ing TAG biosyn hesis is he inco po a ion o diacylglyce ol (DAG) in o phospha idylcholine (PC), ca alyzed by choline phospho ans e ase (CPT) (Kim, 2020). The exis ing acyl g oups can be desa u a ed by a y acid desa u ase (FAD). These PC- inco po a ed DAG molecules can hen be acyla ed in o TAG h ough eac ions ca alyzed by phospholipid: diacylglyce ol acyl ans e ase (PDAT) o PC:DAG phosphocholine ans e ase (PDCT) (Kim, 2020). I is impo an o no e ha he eac ion ca alyzed by CPT is e e sible, allowing DAG o be eleased a la e s ages and con e ed in o TAG h ough he ca aly ic ac ion o DGAT (Eichmann and Lass, 2015). 3.3. T iacylglyce ol s o age Upon comple ion o TAG biosyn hesis, TAGs a e s o ed wi hin oil bodies (OBs), also known as oleosomes, anging om 0.2 o 2.5 µm (Lu kewi e and Finck, 2020). The s uc u e o OBs comp ises a s abilizing monolaye o phospholipids con aining p o eins such as oleosins, caleosins, s e oleosins, and aquapo ins (Lu kewi e and Finck, 2020). Among hese p o eins, oleosins, being he mos abundan , play pi o al oles in he TAG s o age capaci y o a ious plan issues and ac oss di e en plan species. Indeed, he absence o hese p o eins in ege a i e issues is he eason behind hei inabili y o s o e lipids (Xu e al., 2024). Mechanis ically, hese p o eins p e en he coalescence o oil bodies du ing s o age h ough s e ic hind ance and elec onega i e epulsion, hus de e mining he inal size o he OBs (De Chi iko e al., 2018; Kanai e al., 2019). Fo example, Hu e al. (2009) sugges ed ha a low concen a ion o oleosins is associa ed wi h he o ma ion o la ge OBs and, consequen ly, lowe oil con en s. 4. Gene ic enginee ing o imp o ed oil p oduc ion In ecen decades, esea che s ha e de o ed signi ican a en ion o enhancing he quali y and quan i y o seed oils o bo h ood and non- ood pu poses, as well as explo ing new oil sou ces (Subedi e al., 2020a and b; Bha i e al., 2021; Chen e al., 2021a; She i e al., 2021). The a y acid composi ion o plan oils plays a c ucial ole in de e mining hei quali y and sui abili y o a ious applica ions, p ima ily by dis inguishing be ween sa u a ed and unsa u a ed FA. Sa u a ed FA lack double bonds in hei ca bon s uc u e and exhibi a linea con igu a ion, whe eas unsa u a ed FA con ain a leas one double bond in hei ca bon chains. Vege able oils wi h a highe monounsa u a ed- o-sa u a ed a y acid a io demons a e imp o ed s abili y unde high- empe a u e condi ions, such as du ing ying, and exhibi p olonged shel li e when s o ed. In oils in ended o human consump ion, cooking oils ypically con ain highe p opo ions o monounsa u a ed FA, such as oleic acid, while salad oils p edominan ly comp ise polyunsa u a ed FA like linoleic and α-linolenic acids (Kapoo e al., 2021; Saini e al., 2021). To mee he inc easing demand o oils, whe he o human consump ion o o he applica ions like bio uels, me abolic and gene ic enginee ing echniques, including he u iliza ion o ecombinan DNA echnology, ha e been employed o manipula e oil con en in plan s and b oaden he ange o oil a ie ies (Fig. 5). Table 12 p o ides an o e iew o he a y acid composi ion in a ious plan s and ege able oils. 4.1. Me abolic enginee ing o enhance oil p oduc ion Augmen ing oil p oduc ion in seeds has long been a p ima y objec i e o bo h plan b eede s and gene ic enginee s (Subedi e al., 2020a and b; Bha e al., 2022; Rau e al., 2023). Oilseed c ops inhe en ly possess highe seed oil con en compa ed o o he ag icul u ally signi ican c ops. Howe e , he e exis s conside able a ia ion among oilseed c ops hemsel es, wi h oil con en anging om 20% in soybeans o 60% in sesame, despi e simila i ies in hei lipid biosyn hesis pa hways (Za a e al., 2019). Rema kably, a me e 1% imp o emen in soybean oil p oduc ion pe hec a e could con ibu e o e USD 1 billion annually o he c op’s global alue (Ba es e al., 2014), emphasizing he p o ound impac o me abolic enginee ing in oilseed c ops, wi h he po en ial o signi ican ly enhance seed oil con en s. Me abolic enginee ing b oadly aims o egula e lux in o me abolic pa hways by ei he inc easing he a ailabili y o ups eam subs a es o s eng hening he sink in he inal s ages o he pa hway. Bo h s a egies ha e been ha nessed o inc ease seed oil accumula ion. In he case o oilseeds, yield imp o emen s can be achie ed by inc easing he amoun o oil pe seed, enla ging he size o he seed, o boos ing he numbe o seeds pe plan . S a egies o enhance oil p oduc ion in plan s include enhancing he a ailabili y o a y acid (FA) p ecu so s, ele a ing he a e o FA syn hesis, imp o ing TAG assembly p ocesses, and es ic ing TAG deg ada ion pa hways (He e al., 2020; Subedi e al., 2020a and b; Kapoo e al., 2021). In he ollowing sec ion, we e iew he applica ion o me abolic enginee ing o he imp o emen o oil con en and i s quali y. Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2125 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Fig. 5. Gene ic enginee ing o plan oils o di e en applica ions. Table 12. Fa y acid composi ion o plan s and ege ables. Plan Fa y acid composi ion (%) Re e ence Sa u a ed a y acids Unsa u a ed a y acids O he s Cap ylic 8:0 cap ic 10:00 Lau ic 12:00 My is ic 14:0 Palmi ic 16:0 S ea ic 18:0 A achidic 20:0 Behenic 22:0 Lignoce ic 24:0 Ce o ic 26:0 my is oleic 14:1 Palmi oleic 16:1 Sapienic 16:1 Oleic 18:1 Linoleic 18:2 α-linolenic 18:3 Eicosenoic 20:1 E ucic 22:1 A ocado - - - 0.33 23.6 1 - - - - - 3.58 - 47.2 13.4 0 - - 11.7 Mo eno- Camacho e al. (2019) Cas o bean - - - - 1 1 - - - - - - - 3 4 ace - - 91 Aid (2020) Cocoa bu e - - 0-1 0-4 24.5- 33.7 33.7- 40.2 1 - - - - 0-4 - 26.3- 35 1.7-3 ace - - - Naik and Kuma (2014) Coconu 8 7 49 8 8 2 - - - - - - - 6 2 0 - - - Boa eng e al. (2016) Co n - - - - 6.7-16.5 0.7-6.6 0-1 - - - - - - 16.2- 43.8 39.5- 69.5 0-3.1 - - - Whi e (2007) Linseed - - - - 5--6 4--5 - - - - - - - 15-20 14 50-55 - - - Aid (2020); Bay ak e al. (2010) Oli e - - - - 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 e al. (2003) Palm - - 0.2 1.1 44 4.5 0.1 - - - - - - 39.2 10.1 0.4 - - - Mancini e al. (2015) Palm ke nel 3.3 3.5 47.8 16.3 8.5 2.4 0.1 - - - - - - 15.4 2.4 - - - - Mancini e al. (2015) Rapeseed - - - - 3.63 2.32 - - - - - - - 85.31 3.41 4.4 0.94 - - Guan e al. (2016) Soybean - - - - 10 4 - - - - - - - 18 55 13 - - - Clemen e and Cahoon (2009) Sun lowe - - - - 6 5 - - - - - - - 19 68 ace - - - Aid (2020) Co onseed - - - - 25.39 2.33 - - - - 1 0.6 - 16.35 52.89 - - - - Yang e al. (2019) Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2126 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 12. con inued. Plan Fa y acid composi ion (%) Re e ence Sa u a ed a y acids Unsa u a ed a y acids O he s Cap ylic 8:0 cap ic 10:00 Lau ic 12:00 My is ic 14:0 Palmi ic 16:0 S ea ic 18:0 A achidic 20:0 Behenic 22:0 Lignoce ic 24:0 Ce o ic 26:0 my is oleic 14:1 Palmi oleic 16:1 Sapienic 16:1 Oleic 18:1 Linoleic 18:2 α-linolenic 18:3 Eicosenoic 20:1 E ucic 22:1 Almond - - - - 5.07- 6.78 - - - - - - - - 57.54- 73.94 19.32- 35.18 0.04- 0.1 - - - Sa he e al. (2008) Flaxseed - - - - 4.9-8 2.44- 4.59 - - - - - - - 13.44- 19.39 12.25- 17.44 39.9- 60.42 - - - Goyal e al. (2014) Hempseed - - - - 6 3 1 0.44 0.197 - - 0.098 - 16 55 15 - - 3.265 So a e al. (2018) Walnu - - - - 5.61- 5.82 - - - - - - - - 22.62- 27.27 49.93- 54.41 14.32- 17.82 - - - Dogan and Akgul (2005) Rice b an - - - 0.3 22.7 1.8 0.9 - - - - - - 43.9 29.2 1.25 - - - La ha and Nasi ullah (2014) Pumpkin seed - - - 0.233 14.82 6.67 0.433 0.058 - - - - - 25.81 - 50.88 - 0.055 1.041 Ba daa e al. (2016) Sesame - - - - 7.9-12 4.8-6.1 - - - - - - - 35.9- 42.3 41.5- 47.9 - - - - Wacal e al. (2019) Peanu - - - - 8.2-15.1 1.1-7.2 0.8-3.2 1.8- 5.4 0.5-2.5 - - - - 31.5- 60.2 19.9- 45.4 - 0.6-2.6 - - Hammond e al. (1997) G apeseed - - - - 8.4-6.51 3.86- 3.07 - - - - - - - 16.1- 11.62 77.59- 72.5 - - - - Al Juhaimi e al. (2017) Beech nu - - - 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 e al. (2011) Macadamia - - - - 9 2 - - - - - 22 - 60 2 - - - - Tan e al. (2020) Hazelnu - - - - 4.39- 8.85 1.67- 3.18 - - - - - - - 73.48- 81.57 10.46- 14.95 0.02- 0.34 - - - Bal a e 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 G i in and Dean (2017) Pecan - - - 0.4 6.012 - - - - - - - - 23.91 66.81 1.83 - - - Ri e a-Rangel e al. (2018) O ange seed - - 2.96 0.89 12.6 8.9 0.54 - - - - - - 43.03 25.11 4.3 0.67 - - Iwuagwu e al. (2018) Ci us seed - - - - 25.8- 32.2 2.8-4.4 - - - - - - - 21.9- 24.1 36.1- 39.8 3.4-4.4 - - - Reazai e al. (2014) Wa e melon - - - - 9.88 6.96 0.26 - - - - - - 14.25 68.07 - - - - Rezig e al. (2019) A gan - - - - 11--14 4--7 - - - - - - - 46-48 31-35 - - - - Cha ou and Guillaume (2008) Mus a d - - - - 1.87 1.52 1.13 1.4 0.41 - - - 0.1 15.7 12.99 6.18 - 48.8 9.83 El-Shenawy e 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 e al. (2016) Eucalyp ol - - - - 36 3 - - - - - 7.3 - 27.2 19.3 - - - 3.6 Rekkab e 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 A ia e al. (2017) 4.1.1. Inc easing he con en o oil pe seed As men ioned p e iously, TAG biosyn hesis in ol es h ee p ima y me abolic s eps: glycolysis, FA syn hesis, and TAG p oduc ion (Yang e al., 2022b). Glycolysis, as he ini ial s ep in TAG syn hesis, p o ides he ca bon esou ces necessa y o TAG p oduc ion. Me abolic enginee ing s a egies aimed a inc easing oil accumula ion in plan s may in ol e p omo ing he p oduc ion o p ecu so s o lipid syn hesis (e.g., G3P and ace yl-CoA), enhancing he a e o FA syn hesis, inc easing TAG assembly, and inhibi ing TAG deg ada ion pa hways (Xu e al., 2024). Augmen ing FA accumula ion is achie able h ough a ious app oaches, including inc easing he ca bon lux owa ds FA syn hesis, o e exp essing ansc ip ional egula o s in FA syn hesis, p omo ing he exp ession o ACCase, he pi o al enzyme in plas id FA syn hesis, supp essing compe ing pa hways o FA u iliza ion (e.g., memb ane lipid syn hesis), and enhancing he alloca ion o ca bon o FA syn hesis by limi ing s a ch syn hesis (Mulgund, 2022; Ranjba and Malca a, 2022). The lipid me abolic pa hways in plan s a e highly in ica e, in ol ing a leas 120 enzyma ic eac ions and o e 600 genes (Lee e al., 2017; Vanhe cke e al., 2019). Howe e , physiological and de elopmen al ac o s can in luence oil biosyn hesis in plan s (Li e al., 2020a; Yang e al., 2022b). Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2127 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Addi ionally, egula o y pa hways go e ning ca bon lux o s o age and memb ane lipid biosyn hesis emain incomple ely unde s ood (Li e al., 2020b; Yang e al., 2022b). The e o e, a comp ehensi e unde s anding o he complex ne wo ks o oil biosyn hesis and i s egula o y amewo k in plan s can aid in iden i ying and enginee ing key genes in ol ed in inc easing oil con en (Sa adi e al., 2017). 4.1.1.1. FA syn hesis (Ca bon lux edi ec ion) Inc easing seed oil con en necessi a es edi ec ing ca bon lux owa ds a y acid biosyn hesis, as i can signi ican ly impac TAG p oduc ion (Muñoz e al., 2021; Song e al., 2023). Genes encoding enzymes associa ed wi h he FA biosyn he ic pa hway a e key a ge s o gene ic modi ica ion (Sa adi e al., 2017). Howe e , i is impo an o no e ha a emp s o enhance he a ailabili y o p ecu so s o FA syn hesis by al e ing cen al ca bon me abolism ha e no always been success ul, as lux con ol in a pa hway ypically in ol es mul iple componen s a he han indi idual enzymes (Song e al., 2023). ACCase, he a e-de e mining enzyme go e ning FA biosyn hesis, has been ex ensi ely s udied in a ious model plan s (Wan e al., 2017a). ACCase ca alyzes he p oduc ion o malonyl-CoA, p o iding wo ca bon uni s o he FA syn hase complex. O e exp ession o he cy osolic mul i unc ional o m o ACCase in he plas ids o apeseed led o a modes inc ease in seed lipid con en (Zhuko and Popo , 2022). Con e sely, o e exp ession o he gene encoding he bio in ca boxyl ca ie p o ein 2 (BCCP2), a componen o plas idial ACCase, esul ed in dec eased seed oil con en and inc eased le els o suga and p o ein (Zhuko and Popo , 2022), indica ing ha ACCase ac i i y is no he sole limi ing ac o in FA syn hesis (Sa adi e al., 2017). Mi ochond ial py u a e dehyd ogenase (PDH), esponsible o con e ing py u a e and coenzyme A (CoA) in o ace yl-CoA, is egula ed nega i ely by py u a e dehyd ogenase kinase (PDHK). Rep ession o PDHK ac i i y h ough an isense RNA echnology du ing seed ma u a ion has been shown o enhance seed oil con en and seed weigh in A. haliana (Subedi e al., 2020a and b). Silencing he Py u a e dehyd ogenase kinase cDNA (A PDHK) gene using RNAi echnology inc eased seed oil in ansgenic plan s by up o 50%, al e ing a y acid composi ion (Schewende and Hey, 2012). Addi ionally, py u a e, a key p oduc o glycolysis, se es as a majo ca bon sou ce o a y acid p oduc ion in plas ids (Schewende and Hey, 2012). Seed-speci ic o e exp ession o a py u a e anspo e , BASS2, inc eased oil p oduc ion by 24-43% in ansgenic A abidopsis compa ed o wild- ype plan s (Lee e al., 2017), highligh ing he po en ial o p omo ing oil biosyn hesis by enhancing py u a e lux in o plas ids. 4.1.1.2. Glyce ol backbone G3P plays a c i ical ole in egula ing he p oduc ion and e en ion o TAGs in plan issues. Howe e , G3P supply has been shown o be insu icien o main ain high G3P le els du ing he peak oil accumula ion pe iod in oilseed c ops (Polla d and Hill, 2021). G3P is gene a ed in plan s h ough wo dis inc enzymes: cy osolic glyce aldehyde 3-phospha e dehyd ogenase (GAPDH) and glyce ol kinase (GK) (Jeelani e al., 2023). Glyce ol kinase con e s glyce ol in o G3P, p ima ily in ge mina ing seeds, while cy osolic GAPDH ca alyzes he con e sion o dihyd oxyace one phospha e (DHAP) in o G3P in a ious plan issues, including seeds. Gene ic modi ica ion o enhance G3P le els has been success ul in boos ing seed oil con en . Fo ins ance, he seed- speci ic exp ession o yeas cy osolic GAPDH in B. napus esul ed in a 3- 4- old inc ease in G3P le els and a 40% inc ease in seed oil con en . Simila ly, enginee ing C. sa i a by co-exp essing A. haliana diacylglyce ol acyl ans e ase1 (DGAT1) and yeas cy osolic GAPDH led o a 13% inc ease in seed oil con en and up o a 52% inc ease in seed mass compa ed o wild- ype plan s (Chhika a e al., 2018). Mo eo e , o e exp ession o DGAT1 in ansgenic obacco plan s esul ed in a 2.19- old inc ease in oil con en compa ed o non- ansgenic con ol lines (Chena ani e al., 2022). 4.1.1.3. TAG biosyn hesis TAGs in plan s a e syn hesized h ough he Kennedy pa hway in he ER ia a sequen ial acyla ion o G3P by memb ane-bound glycolipid acyl ans e ases. These enzymes include GPAT, LPAAT, and diacylglyce ol acyl ans e ase (DGAT) (Chena ani e al., 2022). The i s acyla ion o G3P is ca ied ou by GPAT, which p oduces lysophospha idic acid in he ER, chlo oplas (CHP), o mi ochond ia (MIT). LPAAT, loca ed in he ER, CHP, MIT, plasma memb ane (PM), and cy oplasm (CP), pe o ms he second acyla ion o lysophospha idic acid o gene a e phospha idic acid. DGAT comple es he inal s ep in he Kennedy pa hway by con e ing diacylglyce ol (DAG) in o TAG (Wan e al., 2017a). Since hese enzymes a e a e-limi ing in TAG syn hesis, esea che s ha e explo ed gene ic modi ica ions o enhance LPAAT, GPAT, and DGAT ac i i y in plan s. Among hese, DGAT is conside ed a c i ical a e-limi ing s ep due o i s compa a i ely lowe enzyme ac i i y compa ed o he o he enzymes in he pa hway (Wan e al., 2017a). Enhancing GPAT and LPAAT ac i i y has p o en e ec i e in inc easing seed oil con en . A abidopsis gene ically enginee ed o exp ess plas idial sa lowe GPAT and E. coli GPAT exhibi ed inc eased seed oil con en du ing seed ma u a ion (Wan e al., 2017b). Simila ly, he exp ession o genes om B. napus encoding mic osomal LPAAT iso o ms in A abidopsis seeds unexpec edly inc eased seed o al FA con en and seed weigh (Zhang e al., 2022). Fu he mo e, he exp ession o a mu an yeas LPAAT gene (SLC1-1) unde he con ol o he CaMV35S p omo e in A abidopsis and high-e ucic acid B. napus esul ed in a 48% inc ease in seed oil con en (Wan e al., 2017b). 4.1.1.4. Lipid ans e p o eins Lipid ans e p o eins (LTPs), also known as plan lipid ans e p o eins (PLTPs), a e small, compac p o eins wi h a hyd ophobic ca i y ha acili a es he ans e o phospholipids and FA be ween cell memb anes (Ge z, 2018). Known o nea ly hal a cen u y, LTPs a e di ided in o wo s uc u ally ela ed sub amilies based on hei molecula weigh s: LTP1s (9 kDa) and LTP2s (7 kDa) (Fan e al., 2013). Pu oindoline A and B (PINA and PINB) p o eins exhibi a s uc u e simila o ha o non-speci ic lipid- ans e p o eins ound in plan s (Zhang e al., 2019b). He e ologous exp ession o whea (T i icum aes i um L.) pu oindoline genes (PINA and PINB) in co n plan s (Zea mays L.) inc eased he ge m size wi hou a ec ing he seed size. As a esul , he o e all seed oil con en inc eased by 25%, and he ge m yield ose by app oxima ely 34% (Zhang e al., 2019b). 4.1.1.5. T ansc ip ion ac o s (TFs) In gene al, he accumula ion o seed oil equi es coo dina ed ansc ip ional con ol o nume ous biosyn he ic pa hways (Kong e al., 2019; Niu e al., 2020; Yang e al., 2022b). Mul iple ansc ip ion ac o s (TFs) play a pi o al ole in synch onizing he egula o y ne wo ks o genes in ol ed in a y acid and TAG biosyn hesis. These TFs con ol gene exp ession by ecognizing and binding o speci ic sequences in gene p omo e s (Kong e al., 2019; Papadimi iou e al., 2019; Tian e al., 2019; Kuma e al., 2020). Manipula ing TFs ep esen s an al e na i e s a egy o enhancing seed oil yield alongside single-enzyme app oaches (San in e al., 2021; Yang e al., 2022a). Posi i e TFs, such as LEAFY COTYLEDON genes (LEC1 and LEC2), FUSCA3 (FUS3), ABSCISIC ACID INSENSITIVE3 (ABI3), and WRINKLED1 (WRI1), a e known o hei pi o al oles in egula ing he ac i i ies o o he TFs in ol ed in seed ma u a ion and oil accumula ion (Fa ihi e al., 2013; Kuma e al., 2020). Howe e , o egula e he exp ession o hese posi i e TFs, plan s ha e e ol ed mechanisms o supp ess hei exp ession du ing he ege a i e phase o de elopmen . Nega i e egula o s, including TFs, mic oRNAs (miRNAs), and p o eins, ac by modula ing he ch oma in con o ma ion o hei a ge genes (Kuma e al., 2020). A common s a egy o manipula ing seed oil con en in plan s is o e exp ession o posi i e egula o s such as WRI1, LEC1/2, and MYB96 o supp ession o nega i e egula o s like MYB89 and WRKY6. Ideally, gene ic enginee ing should ocus on TFs ha ac i a e FA and TAG biosyn hesis genes. O e exp essing TFs in ol ed in FA syn hesis can lead o inc eased TAG con en . Fo example, o e exp essing MYB96, a TF ha ac i a es TAG biosyn hesis genes DGAT1 and PDAT1, can signi ican ly boos TAG s o age in seeds. Howe e , o e exp essing ce ain TFs, ch oma in emodeling agen s, and miRNAs can ha e unin ended consequences on plan g ow h, de elopmen , and yield. The e o e, issue- Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2128 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. speci ic o e exp ession o silencing o desi ed genes in ansgenic seed oil plan s is ecommended. WRINKLED1 (WRI1), belonging o he APETALA 2/e hylene- esponsi e elemen binding p o ein (AP2/EREBP) sub amily o TFs, plays a c ucial ole in egula ing plan oil accumula ion by p omo ing ca bon pa i ioning in o FA syn hesis wi hin he glycolysis pa hway. WRI1 egula es he exp ession o a leas 15 enzymes in ol ed in FA syn hesis and he glycoly ic pa hway, making i an ideal a ge o gene ic manipula ion o enhance oil accumula ion in plan s. O e exp essing ansc ip ion ac o s ha egula e WRI1 exp ession ha e been shown o inc ease seed oil con en in ansgenic plan s. Fo example, o e exp essing soybean ZF351 and GmDREBL inc eased oil con en in ansgenic A abidopsis by binding o he WRI1 p omo e . Simila ly, o e exp essing maize ZmLEC1, an ac i a o o WRI1, led o ele a ed oil con en in A abidopsis, Camelina, and maize, hough i had unin ended e ec s on seed ge mina ion and lea g ow h. Con e sely, o e exp essing maize ZmWRI1 inc eased oil con en wi hou hese side e ec s. Nume ous s udies ha e shown ha o e exp essing WRI1 o i s o hologs signi ican ly boos s seed oil con en in ansgenic plan s. Fo ins ance, cons i u i e exp ession o WRI1 in A abidopsis leads o a 10% o 40% inc ease in seed oil con en and an inc ease in seed size. Selec ing an app op ia e p omo e o d i e WRI1 exp ession is c i ical o gene ically enginee ing oil accumula ion in he seeds o ansgenic plan s. Fo ins ance, using he FUS3 o Oleosin p omo e o d i e WRI1 can inc ease seed oil con en , while using he ZEIN p omo e does no esul in dis inguishable changes in oil con en . Table 13 shows he ansc ip ion ac o s and hei o e exp ession on oil con en . 4.2. Inc easing seed oil con en by enhancing seed size Bo h gene ic and en i onmen al ac o s shape he inal weigh and dimensions o plan o gans du ing g ow h and de elopmen . In oilseed c ops, seed size signi ican ly impac s c op p oduc i i y and yield. Thus, inc easing seed size and biomass eme ges as a s a egy o enhance oilseed c op yield. Howe e , i is c ucial o conside ha augmen ing seed weigh migh necessi a e a educ ion in seed numbe s due o limi ed assimila ion, po en ially a ec ing seed illing. Fu he mo e, al e ing seed oil le els can ha e epe cussions on o al seed p o ein and ice e sa when wo king wi h ixed assimila es. Hence, he key aspec in boos ing oilseed plan yield is ensu ing gene ic manipula ion does no ad e sely a ec seed numbe s. Redi ec ing ca bon lux owa d oil p oduc ion a he han p o ein becomes impe a i e in his ega d. Resea ch on seed size egula ion has iden i ied a ound 88 key egula o s esponsible o o gan size, associa ed wi h pa hways like KLUH, ubiqui in- p o easome, G-p o ein signaling, Mi ogen-Ac i a ed P o ein Kinase (MAPK), and plan ho mone pa hways. Enhancing seed oil con en in ol es exp essing genes ela ed o seed de elopmen , emb yo size, and oil s o age issues in c ops such as A abidopsis, maize, and ice. One app oach is o inc ease oil-ca ying issues while minimally al e ing non-oil-ca ying seed issues. Fo ins ance, enhancing seed oil con en in he endospe m, a no el s o age issue in plan s, is a iable s a egy. These insigh s in o seed size egula ion and oil con en enhancemen pa e he way o imp o ing oilseed c op p oduc i i y and yield. Seed size egula ion in ol es a complex in e play o ac o s, such as plan ho mones, ubiqui in, mic oRNAs, and cy och ome P450s (CYPs). The CYP78A sub amily, a plan -speci ic gene amily, plays a c ucial ole in con olling plan g ow h and o gan size. Va ious sub amilies o CYP78A iden i ied in A abidopsis, ice, whea , oma o, and soybean egula e o gan size and de elopmen . O e exp essing CYP78A genes in di e en plan species has demons a ed signi ican e ec s on seed size and o gan de elopmen . Addi ionally, genes and egula o y elemen s like SWEET, BS1, LEC1, ARF2, DA1, DA2, Abscisic acid biosyn hesis- ela ed genes (ABA2 and ABI5), TTG2, AP2, RING- ype E3 ubiqui in ligases, Enhance o DA1 (EOD1), and miRNA172 impac seed size h ough di e se mechanisms. Enhancing he sink s eng h o seed oil can inc ease he numbe o oil- s o ing bodies in seeds. Ele a ing he oleosin p o ein con en in de eloping seeds enhances oil s o age capaci y and p omo es oil biosyn hesis and s o age in oil bodies, consequen ly inc easing seed oil con en . O e exp essing oleosin genes has p o en e ec i e in inc easing seed oil con en in plan s like A abidopsis and ice. Achie ing emb yo-speci ic o e exp ession o he app op ia e oleosin gene is c ucial o augmen ing seed oil con en wi hou unin ended e ec s. Mo eo e , ansc ip ion ac o ARF2, esponsi e o auxin, ac s as a ep esso o cell di ision, elonga ion, and o gan g ow h. Mu a ions in ARF2 can lead o enla ged seed size and weigh . Tissue-speci ic exp ession o wild- ype ARF2 es o es no mal lowe mo phology and e ili y, unde sco ing i s ole in seed se ing. Simila ly, o e exp ession o WRI1, a membe o he AP2/EREBP amily, has been shown o inc ease seed size in ansgenic plan s. These indings shed ligh on he in ica e egula o y mechanisms go e ning seed size and oil con en in plan s, o e ing a enues o enhancing c op p oduc i i y and yield. 4.3. Gene ic Enginee ing o new oil esou ces: biomass-de i ed oil In addi ion o adi ional plan seeds and ui s, a ious plan issues can syn hesize TAG o plan oils. This capabili y has spu ed esea ch in o gene ic enginee ing s a egies o inc ease oil con en in lea es and o he ege a i e issues o high biomass c ops in an en i onmen ally sus ainable manne . Such modi ied plan biomass can se e as an ene gy-dense, nu i ionally aluable esou ce sui able o elec ici y p oduc ion, biodiesel, enewable uel p oduc ion, and nu i ionally enhanced animal eed. Vege a i e plan o gans ypically con ain a ound 1.5% TAG and 5-10% FAs on a d y weigh basis, making TAG an essen ial bu e and in e media e s o age pool o oxic and excess FAs eleased du ing memb ane a y acid u no e . Despi e his, accumula ing high le els o TAG in ege a i e issues simila o oilseeds is challenging due o he complex na u e o i s biosyn he ic pa hways. Me abolic enginee ing s udies in model plan s ha e sugges ed ha c ops wi h a 15% imp o emen in biomass oil con en could p oduce up o en imes mo e oil pe uni a ea compa ed o canola oilseed c ops. Ea ly e o s ocused on he o e exp ession o down egula ion o indi idual genes in ol ed in FA o TAG biosyn hesis, esul ing in only modes inc eases in TAG con en in ege a i e issues. Recen app oaches ha e combined mul iple me abolic enginee ing s a egies o achie e highe TAG le els in plan biomass. 1. T iacylglyce ol assembly (Pull) app oach: This s a egy in ol es a ge ing he TAG assembly p ocess o inc ease he demand o p ecu so s ha in luence he lux owa d TAG accumula ion in plan issues (Vanhe cke e al., 2017; Wan e al., 2017b; Pa k e al., 2021). O e exp essing TAG assembly enzymes has been sugges ed as an e ec i e means o boos TAG le els in ege a i e issues (Xu and Shanklin, 2016). Fo ins ance, o e exp essing A abidopsis DGAT1 in N. ben hamiana led o a 20- old inc ease in lea TAG con en , while a ge ed exp ession o he same gene in xylem issues esul ed in a 63% inc ease in FAs p oduc ion (Nooka aju e al., 2014). Addi ionally, o e exp ession o Chlamydomonas DGAT2 in A abidopsis esul ed in a 25- old inc ease in TAG con en and changes in TAG composi ion (Pa k e al., 2021; Sagun e al., 2023). No ably, o e exp essing PDAT1 in ansgenic A abidopsis plan s esul ed in a ema kable 28- old inc ease in TAG con en in he lea es (Fan e al., 2013). Addi ionally, o e exp essing mouse monoacylglyce ol acyl ans e ase 1 (MGAT1) and MGAT2 led o diacylglyce ol (DAG) accumula ion, which subsequen ly esul ed in app oxima ely 7-9- old inc eases in TAG con en in ansgenic obacco lea es (Pe ie e al., 2012; Sagun e al., 2023). 2. de no o FAs biosyn hesis (Push) app oach: This app oach in ol es manipula ing key ac o s like ACCase, WRI, and ansc ip ion ac o s such as LEC1, LEC2, FUS3, and ABI3 o boos he de no o syn hesis o FAs (Weselake, 2016; Xu and Shanklin, 2016; Vanhe cke e al., 2017). Fo example, he he e ologous exp ession o A abidopsis ace yl- CoA ca boxylase (ACC1) in po a o plan s esul ed in a i e- old inc ease in TAG con en in ansgenic ube s (Vanhe cke e al., 2019; Luo e al., 2022). O e exp ession o ansc ip ion ac o s (TFs) such as LEC2 and WRI1 has also been shown o enhance lipid con en in lea es (Pa k e al., 2021; Luo e al., 2022). Cons i u i e exp ession o A abidopsis WRI1 esul ed in a 22- old inc ease in TAG con en in he ansgenic ege a i e issues o Nico iana ben hamiana (Vanhe cke e al., 2019). In e es ingly, unlike LEC1 and LEC2, o e exp ession o WRI1 does no appea o ha e unin ended e ec s on plan g ow h and de elopmen . The e o e, combining WRI1 o e exp ession wi h he egula ion o o he genes may be an e ec i e app oach o achie e a signi ican inc ease in oil con en in he ege a i e issues o plan s (Weselake, 2016; Wan e al., 2017b). Hajinaja e al. / Bio uel Resea ch Jou nal 42 (2024) 2105-2145 2129 Please ci e his a icle as: Hajinaja N., Fayyazbakhsh A., Kamal Shahsa a S., Sanja ian F., Rahnama H. Boos ing plan oil yields: he ole o gene ic enginee ing in indus ial applica ions. Bio uel Resea ch Jou nal 42 (2024) 2105-2145. DOI: 10.18331/BRJ2024.11.2.5. Table 13. T ansc ip ion ac o s and hei o e exp ession on oil con en . Gene Func ion Hos Plan Re e ence WRI1 Inc eased seed oil and TAG con en A abidopsis haliana Sun e al. (2017) Lim e al. (2022), Chen e al. (2020) Camelina sa i a An and Suh, (2015) Glycine max Chen e al. (2018), Wang e al. (2022c) Zea mays Pou eau e al. (2011) O yza sa i a Sun e al. (2017) Ja opha cu cas Ye e al. (2018) LEC1 Inc eased seed oil con en B assica napus Elahi e al. (2016) A. haliana Zhu e al. (2018) C. sa i a Zhu e al. (2018) Z. mays Shen e al. (2010) LEC2 Inc eased seed oil con en A. A. haliana Manan e al. (2017) TAG accumula ion in lea es Kim e al. (2015) FUS3 TAG accumula ion in ege a i e issues A. haliana Zhang e al. (2016) ABI3 Inc eased oil accumula ion in lea es A. haliana Yang e al. (2021) DOF- ype ac o s Gossypium hi su um Su e al. (2017) GLABRA2 Inc eased seed oil con en A. haliana Chai e al. (2010) Basic leucine zippe (bZIPs) Ele a ed seed oil con en A. haliana Song e al. (2013) MYB TFs Inc eased seed oil con en A abidopsis and Lo us Li e al. (2017), Khan e al. (2019) Imp o ed seed oil C. sa i a Kim e al. (2019) SPT Co ela ion wi h seed oil con en A. haliana Liu e al. (2017) G-p o ein γ subuni 3 (AGG3) Inc eased seed size C. sa i a Roy Choudhu y e al. (2014) Pu ple acid phospha ase 2 (PAP2) Inc eased seed size C. sa i a Zhang e al. (2012) RNAi supp ession o AGPase Inc eased seed size C. sa i a Na e al. (2018) CYP78A Inc eased seed size A. haliana Fang e al. (2012) Inc eased seed size O. sa i a Xu e al. (2015) Inc eased seed size C. sa i a Holz and Do mann (2021) Inc eased seed size Nico iana abaccum Tian e al. (2016) Co ela ion wi h seed size B. napus, G. hi su um Shi e al. (2019) Sun e al. (2017) GmSWEET10a Simul aneous inc eases in seed size and oil con en G. max Wang e al. (2020b) BIG SEEDS1 (BS1) Inc eased seed size and weigh G. max Ge e al. (2016) BIG SEEDS1 (SHB1) Inc eased seed size B assica juncea A. haliana, G. max Sa adi e al. (2015) Ge e al (2016) KLUH Inc eased seed size A. haliana, G. max Adamski e al. (2009) Zhao e al. 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Nima Hajinaja is a Ph.D. candida e in he Chemical Enginee ing depa men a A izona S a e Uni e si y, Tempe, USA. He holds a Mas e 's deg ee in Chemical Enginee ing – P ocess Design om he Uni e si y o Teh an, I an, and a Bachelo 's Deg ee in Chemical Enginee ing om he Pe oleum Uni e si y o Technology, Ah az, I an. Nima has au ho ed o e 17 pee - e iewed jou nal pape s wi h an h-index o 10, add essing a ious aspec s o mic obial bio echnology and biop ocess enginee ing. His esea ch ocuses on (1) CRISPR-Cas genome edi ing o me abolic enhancemen in cyanobac e ia and E. coli; (2) lux balance analysis o op imizing biochemical p oduc ion; (3) ad anced bio eac o design and scale-up; and (4) echno-economic e alua ions o bio- based p ocesses. His comp ehensi e esea ch p o ile is a ailable a : h ps://schola .google.com/ci a ions?use =pcuiynEAAAAJ&hl=en&oi=ao. Ahmad Fayyzbakhsh holds a PhD in Chemis y om Tomas Ba a Uni e si y in Czechia, whe e hei esea ch unde he Ho izon 2020 p ojec ocused on con olling he biodeg ada ion o h ee polyes e s. Cu en ly, he wo ks as a esea che a Mon an Uni e si y in Leoben, Aus ia, which con inues o pionee sus ainable solu ions in ene gy science. His esea ch p o ile can be ound a : h ps://schola .google.com/ci a ions?use =eQ-zZ_IAAAAJ&hl=en. Sa a Kamal Shahsa a is a esea che wi h a Mas e o Science in Medical Mic obiology om he Mashhad Uni e si y o Medical Sciences, I an. He hesis explo ed he inhibi o y po en ial o IgY an ibodies in he ea men and p ophylaxis o Helicobac e pylo i in ec ion. Sa a's academic jou ney began when she ea ned he Bachelo o Science in Cellula and Molecula Biology. Cu en ly, Sa a's esea ch in e es s span he de elopmen o no el biomedical applica ions o na u al compounds, wi h a ocus on he he apeu ic po en ials o an ibodies. He esea ch in e es s include (1) Gene exp ession, (2) Recombinan P o ein Exp ession and Pu i ica ion, and (3) Plasmid Cloning. He esea ch p o ile is a ailable a : h p://o cid.o g/0009-0009-2480-5106. D . Hassan Rahnama go his PhD in Plan Physiology om he Facul y o Science, Uni e si y o Teh an, I an, in 2004. Cu en ly, he wo ks as an associa e P o . in he Plan Gene ic Enginee ing and Biosa e y Depa men o he Ag icul u al Bio echnology Resea ch Ins i u e o I an (ABRII). His esea ch ocuses on he gene ic enginee ing o oilseed c ops o imp o e hei quali y and quan i y cha ac e is ics. D . Rahnama published se e al pape s and books on plan bio echnology, plan gene ic enginee ing, bioe hics and biosa e y, e c. His comp ehensi e esea ch p o ile is a ailable a : h ps://schola .google.com/ci a ions?use =dWJDZZQAAAAJ&hl=en&oi= ao. D . Fo ough Sanja ian ea ned he PhD in Cell and Molecula Biology om he Facul y o Science a Razi Uni e si y in I an in 2006. She cu en ly se es as a acul y membe in he Plan Bio-p oduc Depa men a he Na ional Ins i u e o Gene ic Enginee ing and Bio echnology (NIGEB) in Teh an, I an. He esea ch p ima ily ocuses on plan gene ic enginee ing, physiology, and bio echnology. D . Sanja ian specializes in s udying plan esponses o bio ic and abio ic s esses, wi h a speci ic emphasis on gene ic ans o ma ion and a deep unde s anding o molecula mechanisms. He esea ch p o ile can be ound a : h ps://schola .google.com/ci a ions?use =qJbN ukAAAAJ&hl=en.