* 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
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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)
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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
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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
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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
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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) -
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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)
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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

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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
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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
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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.
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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.
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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)

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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).
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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-
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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
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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. (2016)
IKU2 Inc eased seed size
A. haliana
Fa ihi e al. (2013)
AUXIN RESPONSE FACTOR 2 (ARF2) Inc eased seed size
A. haliana
Sch u e al. (2006)
DA1 Inc eased seed size
A
. haliana Li e al. (2008)
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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.