Insigh in o he chemical and nu i ional a p o ile o Teneb io moli o
la ae ea ed on di e en Ag i- ood by-p oduc s
M. Lou des Mo ales
a
, M. Pila Segu a-Bo ego
a,*
, Jos´
e Raúl Aguile a-Vel´
azquez
b
,
Raquel M. Callej´
on
a
, Daniel Gu i´
e ez-P aena
c
, C is ina Ubeda
a
a
´
A ea de Nu ici´
on y B oma ología, Dp o. Nu ici´
on y B oma ología, Toxicología y Medicina Legal, Facul ad de Fa macia, Uni e sidad de Se illa, Se illa, Spain
b
Dp o. Bioquímica y Biología Molecula , Facul ad de Fa macia, Uni e sidad de Se illa, Se illa, Spain
c
´
A ea o Toxicology, Dp o. Nu ici´
on y B oma ología, Toxicología y Medicina Legal, Facul ad de Fa macia, Uni e sidad de Se illa, Se illa, Spain
ARTICLE INFO
Keywo ds:
Fa y acid
Alpe ujo
Oli e lea
Mush oom
Heal h-p omo ing index
ABSTRACT
Insec s a e capable o eeding wi h a a ie y o subs a es, being possible ob aining p oduc s ich in p o ein using
by-p oduc s om ag i- ood indus y. Hence, he aim o his wo k was e alua ing he e ec o di e en die s based
on h ee ag i- ood indus y by-p oduc s o Teneb io moli o la ae’s o al a con en s and a y acid p o iles.
S abilized Alpe ujo (ALP), Pleo o us os ea us spen subs a e (POS) and oli e lea lou (OLF) we e used mixed
wi h whea b an, con ol die , a di e en pe cen ages (20–100 %). All subs a es s udied modi ied he pa ame e
analysed espec o he con ol die . OLF sub ac led o he g ea es e ec s on he o al a con en a pe cen age
highe han 40 %. Respec o a y acid p o ile, he mos ele an changes we e he inc eases o o al MUFA
con en in samples om all T. moli o la ae ed wi h ALP and in mos o hose ed wi h POS, due o he speci ic
inc eases o oleic acid con en . On he con a y, o al PUFA con en dec eased in mos cases excep o OLF
subs a e when he addi ion pe cen ages we e lowe han 100 %. Mo eo e , OLF subs a e enhanced he p esence
o
ω
-3.
Finally, acco ding o Eu opean egula ions on nu i ional claims, his a ac ion could be labelled as “sou ce
o
ω
-3”, “high in monounsa u a ed a ” and “high in unsa u a ed a s”. We can he e o e conclude ha eeding
T. moli o wi h hese ag i- ood by-p oduc s, especially om oli e oil indus y, enhanced he a y acids p o ile o
hei a ac ion.
1. In oduc ion
Teneb io moli o la ae ha e been au ho ized as human ood by he
Eu opean Food Sa e y Au ho i y (EFSA) (Eu opean Commission, 2021).
These insec s a e a g ea sou ce o eme gen p o eins conside ing ha
hei con en s may ange be ween 48.8 and 68.9 % o d y weigh and
may p o ide essen ial amino acids, highligh ing he con en o leucine,
aline, o isoleucine (Mu˜
noz-Seijas e al., 2024). Thus, hei inclusion in
he human die can help educing he g eenhouse e ec and he wa e
de ici s, hanks o he dec ease o he mea p o ein p oduc ion and due
o hei low wa e consump ion.
The de elopmen cycle o T. moli o consis s o 4 dis inc li e s ages:
egg, la a, pupa and adul . The cycle begins wi h he laying o eggs by he
emale. The eggs ha ch in 4 days a empe a u es o 26–30 ◦C. The la al
s age has an a e age du a ion o 112 o 203 days, depending on g ow h
condi ions. The pupal s age las s be ween 6 and 20 days, a e which
adul s eme ge as whi e and so bee les, g adually ha dening and
da kening hei exoskele on. Ma ing and egg-laying begin app oxi-
ma ely 3 days a e eme gence, ha ing he adul s an a e age li espan o
32 o 62 days (Ribei o e al., 2018).
T. moli o ha e showed he abili y o inges di e en kind o die .
The e o e, a ious ea ing subs a es ha e been s udied, including
ege ables disca ded due o quali y s anda ds (L´
opez-G´
amez e al.,
2024), as well as co n s o e , soybean meal, and dis ille s’ g ain by-
p oduc s (Zhang e al., 2019), among o he . They a e e en capable o
deg ading di e en ypes o plas ics (B andon e al., 2018; Tsocha zis
e al., 2021). Hence, he p oduc ion in a high scale mode o human
nu i ion ( ood human supply) could enable o p oduce “ ood ich in
p o ein” and he ag i- ood was es exploi a ion simul aneously. On one
hand, oli e oil p oduc ion gene a es la ge quan i ies o was es, including
lea es, alpe ujo, wash wa e , among o he s. Alpe ujo is a mix u e o
ege a ion wa e s o alpechines; solid pa s o he oli e, such as he
* Co esponding au ho .
E-mail add ess: [email p o ec ed] (M.P. Segu a-Bo ego).
Con en s lis s a ailable a ScienceDi ec
Food Resea ch In e na ional
jou nal homepage: www.else ie .com/loca e/ ood es
h ps://doi.o g/10.1016/j. ood es.2025.116223
Recei ed 19 No embe 2024; Recei ed in e ised o m 6 Feb ua y 2025; Accep ed 11 Ma ch 2025
Food Resea ch In e na ional 209 (2025) 116223
A ailable online 19 Ma ch 2025
0963-9969/© 2025 The Au ho s. Published by Else ie L d. This is an open access a icle unde he CC BY-NC-ND license (
h p://c ea i ecommons.o g/licenses/by-
nc-nd/4.0/ ).
s one, mesoca p and skin; and a y emains, p oduced du ing oil
ex ac ion. This was e is highly pollu ing, so a way is being sough o
ake ad an age o hem and educe hei accumula ion. This p oblem
a ec especially o Spain, as i is he wo ld’s leading coun y in e ms o
oli e oil p oduc ion and he amoun o land de o ed o oli e cul i a ion.
Mo eo e , acco ding o FAO (FAO (Food and Ag icul u e O ganiza ion
o he Uni ed Na ions, 2025), global mush oom p oduc ion in 2023 was
50.01 million ons. In cul i a ion o ungi i is p oduced as was e he
exhaus ed subs a e. Thus, in pa allel wi h he inc ease in mush oom
p oduc ion o human consump ion, he quan i y o his was e has also
augmen ed. The e o e, i hese esidues could be used o eed T. moli o i
would be a good al e na i e o educe hem. Thus, wi hin he amewo k
o he ci cula economy, he use o by-p oduc s o was es om he ag i-
ood sec o as a subs a e o he die o T. moli o is a p omising op ion.
On he o he hand, he subs a es used in he ea ing o T. moli o a e
impo an since changes we e obse ed in p oximal composi ion o
la ae. Thus, he use o ege able was e (L´
opez-G´
amez e al., 2024), co n
s o e , soybean meal, and dis ille s’ g ains by p oduc s (Zhang e al.,
2019) o Mo inga olei e a lea es (Ko sou e al., 2023) p oduced, in mos
cases, signi ican inc eases o o al p o ein con en s o la ae, as well as
dec eases in ash con en compa ed o he con ol. Mo eo e , insec a
con en and a y acid p o ile a e a ec ed by he ea ing subs a e
(Pino i e al., 2019). In gene al, T. moli o la ae can ha e o al a
con en s ha can ange be ween 19.12 and 50.15 % o d y weigh
(Huang e al., 2025). The mos abundan a y acids in he la ae o hese
insec s a e oleic acid, ollowed by linoleic acid and palmi ic acid. I he
a y acids a e g ouped acco ding o he numbe o unsa u a ions, in
descending o de , he con en s o monounsa u a ed a y acids (MUFA)
a e he highes , ollowed by polyunsa u a ed (PUFA) and, inally sa u-
a ed (SFA) (dos San os Aguila , 2021).
The amoun and he ype o a s (sa u a ed, monounsa u a ed o
polyunsa u a ed) ha human inges ha e a ele an e ec on hei
heal h. Besides, he in ake o high o al SFA le els, and conc e ely
my is ic, palmi ic, and s ea ic acids, a e linked o an inc eased cance
isk (Mei e al., 2024). Scien i ic e idence has led o he Wo ld Heal h
O ganiza ion (WHO) o upda e i s guidance on o al a and sa u a ed a
consump ion. Hence, WHO ea i ms ha adul s should limi o al a
in ake o 30 % o o al ene gy in ake o less wi h a p ima ily composi ion
o unsa u a ed a y acids, wi h no mo e han 10 % o o al ene gy in ake
coming om sa u a ed a y acids (WHO, 2023). Mo eo e , PUFA inges
ecommenda ions should ange 6–11 % o o al ene gy in ake (FAO
(Food and Ag icul u e O ganiza ion o he Uni ed Na ions), 2010).
Consequen ly, i is no only impo an o explo e new p o ein sou ces
bu also o ensu e hey o e a heal hie a p o ile, as is he case wi h
insec -based p o eins, which a e known o hei high nu i ional alue.
Thus, he aims o his wo k we e o s udy he e ec o di e en ag i- ood
by-p oduc s as ea ing subs a e on he a con en and a y acid p o ile
o T. moli o la ae assessing he po en ial ele ance o each o he a s
ob ained in di e en “heal h s a emen s”. Fo his pu pose, T. moli o ’s
die s based on h ee kinds o ag i- ood by-p oduc s a di e en pe -
cen ages we e s udied.
2. Ma e ials and me hods
2.1. Chemicals
Hexane and me hanol we e supplied by Me ck (Da ms ad , Ge -
many), and hyd ochlo ic acid 37 % / was supplied by PanReac (Ba -
celona, Spain). Two in e nal s anda ds we e used nonadecanoic acid
om Sigma-Ald ich (Me ck, Da ms ad , Ge many) o es he ans-
es e i ica ion eac ion (IST) and 2-phenyle hanol p o ided by Me ck
(Da ms ad , Ge many) as IS o injec ion. To quan i ica ion we e used he
ollowing s anda ds comme cial mix u e: 37 componen FAME mix
(CRM47885) and FAME mix C8-C24 (CRM-18918) om Supelco
(Me ck, Da ms ad , Ge many).
2.2. Collec ion and ea men o eeding subs a es
Th ee di e en ypes o subs a es based on ag i- ood by-p oduc s
has been used in his s udy: s abilized alpe ujo (ALP), mush oom
(Pleo o us os ea us) spen subs a e (POS), and oli e lea s lou (OLF).
Mush oom (Pleo o us os ea us) spen subs a e was p o ided by Se aco
company (Villa anca de C´
o doba, C´
o doba, Spain) and did no unde go
any p e ea men . Alpe ujo and oli e lea s we e p o ided by he Ag i-
cul u al Coope a i e “Nues a Se˜
no a de los ´
Angeles” (Mon ellano,
Se illa, Spain). To ob ain oli e lea es lou (OLF), he lea es we e pul-
e ized. The alpe ujo, a we by-p oduc om he ex ac ion o oil, was
ob ained in Decembe 2022. To s abilize he ma e ial, esh alpe ujo was
subjec ed o a d ying p ocess a 80 ◦C o app oxima ely 43 ±4 h. This
p ocedu e was conduc ed using an ai -d ying o en (Memme UF260).
Following he d ying s ep, he d y alpe ujo was inely g ound using a
Re sch SM 100 mill i ed wi h a 750
μ
m sie e in o de o make i
a ailable o insec s.
2.3. Feeding ials
T. moli o la ae we e ed wi h 3 di e en ypes o subs a es based
on ag i- ood by-p oduc s: s abilized Alpe ujo, mush oom (Pleo o us
os ea us) spen subs a e, and oli e lea es lou . These by-p oduc s
we e mix u e wi h whea b an a di e en pe cen ages (20, 40, 60, 80
and 100 %) and he con ol die was 100 % o whea b an. Th ee assays
o each pe cen age and subs a e we e ca ied ou . Fo logis ical easons,
ini ial eeding assays wi h ALP and POS we e conduc ed, ollowed by
eeding assays wi h he OLF subs a e. Th ee assays we e pe o med o
each pe cen age and subs a e, and each ound o es s included a con-
ol g oup.
2.4. P ocessing o la ae
When 10 % o he indi iduals had pupa ed, he ea ing assays we e
concluded, and he la ae we e sepa a ed om he ea ing subs a es
and as ed o 48 h. A e ha , la ae we e killed by subme ging hem in
ho wa e a 50 ◦C o 2 min ( he mos a ic wa e ba h Raypa, BAD-2,
Te asa, Spain). Then, la ae we e d ied in an o en (Memme model
UF260, Ge many) a 40 ◦C o 3 days, milled in a semi-indus ial mill
Res ch model GM 300 (Haan, Ge many), and s o ed in plas ic bags wi h
Zip lock (he me ic seal) a −20 ◦C un il analysis.
2.5. De e mina ion o o al a con en and a y acid composi ion
The o al con en o a was de e mined by Soxhle me hod based on
AOAC (2012). Ten g ams o samples we e ex ac ed wi h hexane o 2 h.
The a ac ion o samples was s o age a −20 ◦C.
Fa y acids we e de e mined u ning hem in o hei co esponding
me hyl es e s and analysing hem by GC–MS. An acid ca alysed ans-
es e i ica ion was pe o med (Osimani e al., 2017). Fo i , 100
μ
L o
nonadecanoic acid (4.06 mg/mL) (IST) and 2 mL o HCl 3 N in me hanol
we e added o 200 mg o T. moli o la ae a . The mix u e was pu in o
an o en a 70 ◦C, o 2 h. Then, le i cool a oom empe a u e and 1 mL
o dis illed H
2
O and 2 mL o hexane we e added, he mix u e was
cen i uged a 3000 pm du ing 3 min o make easie he sepa a ion o
di e en phase. The o ganic laye was ans e ed o a olume ic lask,
made up o 2 mL wi h hexane and s o age in a ial a −20 ◦C un il
analysis.
An Agilen 6890 GC sys em coupled o an Agilen 5975 ine quad-
upole mass spec ome e (Agilen , San a Cla a, CA, US) was used o
analysis. The olume o sample used was 148
μ
L ha join o 2
μ
L o IS
(2-phenyle hanol, 6.06 mg/mL hexane) we e pu in o a ial mic o-inse
and 1
μ
L we e injec ed in spli mode 10:1 a 250 ◦C. Gas ca ie used was
He a 0.8 mL/min. Fa y acid sepa a ion was ca ied ou using he DB-
Fas FAME column o 20 m ×180
μ
m i.d. x 0.2
μ
m o ilm hickness
(Agilen , San a Cla a, CA, US). The o en empe a u e p og am was:
M.L. Mo ales e al.
Food Resea ch In e na ional 209 (2025) 116223
2
70 ◦C o 0.5 min, hen i was inc eased 20 ◦C/min o 110 ◦C, 10 ◦C/min
o 171 ◦C, 3 ◦C/min 185 ◦C hold du ing 1 min, 0.5 ◦C/min o 190 ◦C,
7 ◦C/min o 204 ◦C du ing 2 min, 25 ◦C/min o 240 ◦C du ing 2 min
(adap ed om Tossa ainen e al., 2017). De ec ion was pe o med on
selec ed ion moni o ing (SIM) mode a 70 eV. The ions moni o ed we e:
55, 67,74, 79, 91, 107, 108, 122, 143, 227, 236, 241, 250, 255, 264, 273,
312, 320, 322, 340, 350, 368, and 382. The MS quad upole, sou ce, and
ans e line empe a u es we e 150 ◦C, 230 ◦C and 250 ◦C, espec i ely.
Fo compounds iden i ica ion, spec a and e en ion imes o eal s an-
da ds we e used, o ha , a mix u e o s anda ds was injec ed in ull scan
mode (29–400 m/z). Me hyl es e s o a y acids (FAMEs) we e quan i-
ied building eg ession lines wi h 6 le els o concen a ion (Table S1,
supplemen a y ma e ial).
2.6. Die a y indica o s
Se e al indexes ela ed o ca dio ascula heal h ha e been calcu-
la ed. Thus, he a ios be ween PUFA:SFA, MUFA:PUFA,
ω
-6:
ω
-3, join o
a he ogenic index (AI), h ombogenic index (TI), hypocholes e olemic/
hype choles e olemic a io (h/H) and heal h-p omo ing index we e
calcula ed (HPI) (Chen & Liu, 2020).
2.7. S a is ical analysis
Analysis o a iance (ANOVA) was pe o med o check whe he he
changes in he alues o a y acids and heal hy indexes we e s a is ical
signi ican (p <0.05). Fo ha , we ha e used In os a so wa e (FCA,
Uni e sidad Nacional de C´
o doba, A gen ina) (pos hoc LSD Fishe ).
Pea son co ela ion analysis (p <0.05) was unde aken in o de o
asce ain he possible ela ionship and he di ec ion o his, be ween
a y acids con en s and he pe cen ages o subs a e added o T. moli o ’s
die . S a so S a is ical, e sion 7.0 (S a so , Tulsa, OK) was used o
his pu pose.
Table 1
To al a con en and p o ile o a y acids o T. moli o c ushed samples ed wi h con ol and wi h di e en pe cen age o ALP subs a es (means alues o iplica e o
biological assays ±SD).
Samples Con ol-1 ALP20 ALP40 ALP60 ALP80 ALP100
To al a con en (%w/w) 21.0 ±1.0
b
22.3 ±0.6
b,B
23.3 ±2.5
b
24.0 ±1.7
b
25.0 ±1.7
b,A
30.3 ±5.9
a,A
Fa y acid con en (% o o al a y acids con en )
SFA
Bu anoic acid C4:0 0.49 ±0.26 0.51 ±0.44 n.q. 0.43 ±0.40 0.48 ±0.70 n.q.
Dodecanoic acid C12:0 0.359 ±0.018
a
0.305 ±0.003
b,B
0.289 ±0.015
b
0.268 ±0.035
b,c,B
0.296 ±0.033
b
0.246 ±0.011
c,B
T idecanoic acid C 13:0 0.113 ±0.002
a
0.105 ±0.001
a,B
0.101 ±0.010
a,b,B
0.089 ±0.010
b,B
0.092 ±0.010
b,B
0.052 ±0.004
c,B
My is ic acid C14:0 3.10 ±0.13
a
2.76 ±0.10
b,c,B
2.82 ±0.21
a,b,c
2.61 ±0.19
c,B
2.97 ±0.24
a,b
2.60 ±0.20
c
Pen adecanoic acid C 15:0 0.303 ±0.009
a
0.273 ±0.020
a,b
0.288 ±0.020
a,b
0.266 ±0.019
b,B
0.265 ±0.013
b
0.228 ±0.030
c,C
Palmi ic acid C16:0 17.1 ±1.6
a
13.8 ±1.9
b,c
12.1 ±1.8
c
12.7 ±1.1
b,c,B
13.9 ±0.7
b,c,B
15.5 ±2.4
a,b,A
Hep adecanoic acid C17:0 0.438 ±0.014
b,c
0.471 ±0.009
a,b,A
0.512 ±0.027
a
0.449 ±0.010
b,c,B
0.409 ±0.029
c,B
0.320 ±0.070
d,B
S ea ic acid C18:0 2.57 ±0.20
c
2.71 ±0.15
b,c
2.82 ±0.10
a,b
2.66 ±0.10
b,c,B
2.71 ±0.0.4
b,c,B
2.97 ±0.11
a,B
A achidic acid C20:0 0.437 ±0.010
b
0.535 ±0.061
a,b
0.590 ±0.077
a
0.553 ±0.032
a
0.524 ±0.026
a,b,C
0.596 ±0.095
a,B
Heneicosanoic acid C21:0 0.009 ±0.001
a
n.q.
b
n.q.
b
n.q.
b
n.q.
b,B
-
b,B
Behenic acid C22:0 0.158 ±0.007
a,b
0.168 ±0.033
a,b
0.195 ±0.033
a
0.150 ±0.032
a,b
0.161 ±0.008
a,b,B
0.128 ±0.039
b,B
T icosanoic acid C23:0 0.013 ±0.003
b,c
0.048 ±0.042
a
n.q.
c,B
0.031 ±0.005
a,b
n.q.
c,B
n.q.
c,B
Lignoce ic acid C24:0 –n.q. n.q. n.q. n.q. –
MUFA
Palmi oleic acid C16:1 (
ω
-7) 1.86 ±0.20
b
1.88 ±0.07
b,B
1.86 ±0.08
b,B
1.93 ±0.12
b,C
1.99 ±0.07
b
2.80 ±0.12
a,B
cis-10-Hep adecenoic acid C17:1 (
ω
-7) 0.352 ±0.029
b
0.419 ±0.011ª
,B
0.415 ±0.011ª
,B
0.431 ±0.009ª
,B
0.439 ±0.007ª
,B
0.375 ±0.044
b,B
Oleic acid C18:1 (
ω
-9) 31.0 ±1.3
c
38.2 ±1.4
b,A
39.8 ±6.4
b
43.4 ±4.2
b,A
44.0 ±2.5
b,A
61.0 ±0.6
a,A
cis-11-Eicosenoic acid C20:1 (
ω
-9) 0.326 ±0.019
b
0.389 ±0.045
a,A
0.373 ±0.035
a,b
0.391 ±0.031
a
0.380 ±0.016
a,b,A
0.393 ±0.031
a
PUFA
Linoleic acid C18:2 (
ω
-6) 39.9 ±1.2
a
36.0 ±0.4
a,b
36.7 ±8.4
a,b
32.3 ±4.3
b,B
30.5 ±2.7
b,B
12.4 ±1.6
c,B
Linolenic acid C18:3 (
ω
-3) 1.3 ±0.04
a
1.26 ±0.01
a,b,A
1.04 ±0.13
c,d,B
1.13 ±0.13
b,c,B
0.90 ±0.05
d,B
0.35 ±0.04
e,B
cis-11,14-Eicosadienoic acid C20:2 (
ω
-6) 0.166 ±0.026ª
,b
0.182 ±0.004ª
,A
0.087 ±0.075
b
0.137 ±0.019ª
,b
0.086 ±0.075
b
n.q.
c,B
To al con en s and Fa y acid indexes
SFA 25.0 ±2.1
a
21.6 ±1.3
b,c,A
19.7 ±1.7
c,B
20.2 ±1.0
b,c,B
21.8 ±0.2
b,c,B
22.7 ±2.3
a,b
MUFA 33.6 ±1.1
c
40.9 ±1.4
b
42.4 ±6.5
b
46.2 ±4.3
b,A
46.8 ±2.5
b,A
64.6 ±0.7
a,A
PUFA 41.4 ±1.2
a
37.5 ±0.4
a,b
37.8 ±8.2
a,b
33.6 ±4.4
b,B
31.5 ±2.5
b,B
12.7 ±1.7
c,B
ω
-6 40.1 ±1.2
a
36.2 ±0.4
a,b
36.8 ±8.3
a,b
32.5 ±4.3
b,B
30.6 ±2.7
b,B
12.4 ±1.6
c,B
ω
-3 1.3 ±0.04
a
1.26 ±0.01
a,b,A
1.04 ±0.13
c,d,B
1.13 ±0.13
b,c,B
0.90 ±0.05
d,B
0.35 ±0.04
e,B
PUFA:SFA 1.66 ±0.18
a
1.74 ±0.10
a
1.94 ±0.59
a
1.66 ±0.25
a,A
1.45 ±0.13
a,B
0.57 ±0.14
b,B
MUFA:PUFA 0.812 ±0.022
c
1.09 ±0.04
b,c
1.18 ±0.38
b,c
1.40 ±0.32
b,c
1.50 ±0.21
b,A
5.12 ±0.61
a,A
ω
-6:
ω
-3 30.8 ±1.1 28.8 ±0.3
B
36.3 ±13.4 28.8 ±1.0
B
34.0 ±3.8 35.2 ±4.1
A,B
AI 0.398 ±0.040
a
0.32 ±0.03
b
0.30 ±0.03
b
0.294 ±0.015
b,B
0.333 ±0.022
b,A
0.34 ±0.05
a,b,A
TI 0.56 ±0.06
a
0.45 ±0.05
b,c
0.42 ±0.05
c,B
0.42 ±0.03
c,B
0.473 ±0.023
a,b,c,B
0.54 ±0.08
a,b,A
h/H 3.6 ±0.4
c
4.5 ±0.6
a,b,c
5.2 ±0.8
a
4.9 ±0.4
a,b,A
4.4 ±0.3
a,b,c,B
4.1 ±0.7
b,c,B
HPI 2.53 ±0.24
b
3.1 ±0.3
a
3.4 ±0.4
a
3.40 ±0.17
a,A
3.01 ±0.20
a,b,B
3.0 ±0.5
a,b,B
-: no peak de ec ed; n.q.: amoun unquan i iable, n.d.: amoun unde ec able.
SFA: sa u a ed a y acid; MUFA: monounsa u a ed a y acid; PUFA: polyunsa u a ed a y acid; AI: a he ogenic index; TI: h ombogenic index;
h/H: hypocholes e olemic/hype choles e olemic a io; HPI: heal h-p omo ing index.
Di e en lowe case le e in he same line means signi ican di e ences (p <0.05) be ween samples ed wi h he same subs a e wi h di e en pe cen age (Con ol-100
%).
Di e en capi al le e means signi ican di e ences (p <0.05) be ween samples o la ae ed wi h he same pe cen age o he h ee subs a es s udied.
M.L. Mo ales e al.
Food Resea ch In e na ional 209 (2025) 116223
3
3. Resul s and discussion
3.1. E ec o he di e en T. moli o die s on o al a con en s and a y
acid p o iles
The o al con en o a in he eeding subs a es used in his wo k
we e di e en , hus, he subs a e ha p o ided mo e a o he la ae
die was Alpe ujo (ALP) (14 % w/w), hen, in he second place he oli e
lea subs a e (OLF) (7.5 %) wi h he hal con en o he p e ious one.
Whea b an subs a e used as con ol con ained a 6 % o a and Pleo o us
os ea us spen subs a e (POS) showed he lowes con en o a (5 %).
In he c ushed o T. moli o la ae, o al a con en anged om 11.1
o 30 % w/w in he case o la ae ed wi h 100 % o OLF and in he case
o 100 % o ALP subs a e, espec i ely (Table 1–3). An obse a ion o
highligh was he inc ease in he o al con en o a in he c ushed o
la ae ed wi h ALP di ec ly p opo ional o he pe cen age o his
subs a e in he die ( =0.91), whe eas an in e se co ela ion was ound
in he case o OLF subs a e pe cen ages ( = − 0.89). Zhang e al. (2019)
and Lawal e al. (2021) also obse ed ha he o al con en o a in
T. moli o la ae was no co ela ed o he a con en o subs a es. I we
compa e he a con en a he same pe cen age o each subs a e, he
con en s we e signi ican ly highe in he case o OLF a 20 %, in ALP a
80 % and in ALP and POS a 100 %. The e o e, he e was in luence o he
eeding subs a e in he o al con en o a , al hough his only clea ly
depended on he amoun o a o he die in he case o alpe ujo sub-
s a e. In consequence, i he objec i e o b eeding T. moli o was o
p oduce oil, a good s a egy would be o include a high pe cen age o
alpe ujo in i s die .
Due o he di e en e ec s o a y acid p o iles o ood on human
heal h, i is impo an o know i changes occu in he a y acid p o ile
and which a e hem.
The a o T. moli o la ae ed wi h he con ol die was ich in PUFA,
con aining p ima ily C18:2
ω
-6 (Linoleic acid) ollowed by C18:1
ω
-9
(oleic acid) and palmi ic acid as sa u a ed a y acid (Tables 1 and 3). A
possible explana ion could be ha C18:2 a y acid was he mos
abundan in he con ol subs a e. This esul ag ees wi h hose ound by
Ko sou e al. (2023), who also used whea b an as die con ol. In con ol
samples, he o al con en o PUFA eached he highes signi ican
alues, ollowed by he MUFA o al con en , being also his signi ican ly
highe han o al con en o SFA. Howe e , i was obse ed ha his
end changed as he die o he la ae was modi ied. In la ae ed wi h
ALP subs a e (Table 1), a clea shi in a y acids composi ion was
obse ed, he o al con en o MUFA inc eased while he o al con en o
PUFA dec eased. I can also be obse ed a end o inc ease o he pe -
cen age o MUFA as he pe cen age o ALP subs a e in he die inc eased
( =0.90). This was p ima ily due o he signi ican inc eases o C18:1
(
ω
-9) con en s, which becomes he mos abundan a y acid in hese
samples. The highes s a is ically signi ican pe cen age o o al MUFA
con en s was eached when he la ae we e ed wi h a 100 % o ALP
(Fig. 1). MUFAs such as C16:1(n-7), C18:1(n-9), and C20:1(n-9), p esen
in ou samples, ha e been associa ed wi h a dec ease in he impac o
obesi y and ela ed me abolic synd omes (Gue endiain e al., 2018).
On he con a y, he C18:2 (
ω
-6) con en s dec eased, being signi i-
can ly lowe when he die con ained pe cen age o alpe ujo highe han
40 espec o he con ol samples. A simila end was obse ed o C18:3
(
ω
-3), o his compound in mos o he ALP pe cen age s udied, excep
o 20 %, he con en s in T. moli o c ushed we e signi ican ly lowe han
in he con ol which in u n mean a dec ease in he o al pe cen age o
ω
-3 (Table 1). Bo h changes lead o a dec ease o he o al PUFA con en s
whish we e in e sely p opo ional o he pe cen age o alpe ujo used (
= − 0.86), being he la ae samples ed wi h he 100 % o ALP die hose
which accoun ed he signi ican ly lowes alues o o al PUFA con en s
(Fig. 1). Palmi ic, ollowed by my is ic and s ea ic acids, hese wo las
wi h simila alues, we e, as in he con ol samples, he sa u a ed a y
acids mos abundan when ALP is used in he die o T. moli o (Table 1).
We did no obse e ele an changes o o al con en o SFA, emaining
Table 2
To al con en and p o ile o a y acids o T. moli o c ushed samples ed wi h
di e en pe cen age o POS subs a e (means alues o iplica e o biological
assays ±SD).
Samples POS20 POS40 POS60 POS80 POS100
To al a con en
(%w/w)
21.0 ±
1.0
a,b,B
22.0 ±
3.5
a,b
22.0 ±
4.4
a,b
18.3 ±
1.5
b,C
25.3 ±
2.9
a,A
Fa y acid con en (% o o al a y acids con en )
SFA
Bu anoic acid C4:0 0.62 ±
0.49
a,b
1.23 ±
1.37
a,b
1.43 ±
1.26
a
n.q.
b
0.16 ±
0.28
a,b
Dodecanoic acid
C12:0
0.319 ±
0.021
B
0.305 ±
0.066
0.305 ±
0.017
A,B
0.323 ±
0.008
0.372 ±
0.056
A
T idecanoic acid C
13:0
0.110 ±
0.008
a,B
0.096 ±
0.022
a,b,
B
0.094 ±
0.009
a,b,
B
0.098 ±
0.016
a,B
0.073 ±
0.013
b,B,
*
My is ic acid C14:0 2.81 ±
0.13
B
2.59 ±
0.58
2.62 ±
0.10
B
2.82 ±
0.13
3.14 ±
0.55
Pen adecanoic acid
C 15:0
0.283 ±
0.026
0.318 ±
0.074
0.396 ±
0.067
A
0.356 ±
0.125
0.333 ±
0.037
B
Palmi ic acid C16:0 12.7 ±
2.3
b,
*
15.1 ±
1.6
a,b
17.2 ±
1.8
a,A
16.2 ±
1.0
a,A
16.2 ±
1.8
a,A
Hep adecanoic
acid C17:0
0.413 ±
0.027
a,B
0.447 ±
0.090
a
0.420 ±
0.053
a,B
0.502 ±
0.108
a
0.284 ±
0.048
b,B,
*
S ea ic acid C18:0 2.73 ±
0.22
2.99 ±
0.55
2.93 ±
0.16
A
2.92 ±
0.18
2.98 ±
0.23
B
A achidic acid
C20:0
0.667 ±
0.131
0.663 ±
0.231
0.664 ±
0.158
0.637 ±
0.038
B
0.637 ±
0.123
B
Heneicosanoic acid
C21:0 n.q.* n.q.* n.q.* n.q.
B,
* n.q.
B,
*
Behenic acid C22:0 0.245 ±
0.08
a
0.233 ±
0.09
a
0.174 ±
0.08
a,b
0.190 ±
0.02
a,b,B
0.110 ±
0.06
b,B
T icosanoic acid
C23:0
0.019 ±
0.011
a
0.017 ±
0.004
a,A
0.012 ±
0.011
a
-
b,B,
* n.q.
b,B,
*
Lignoce ic acid
C24:0 – – n.q. n.q. –
MUFA
Palmi oleic acid
C16:1 (
ω
-7)
1.80 ±
0.01
b,B
1.91 ±
0.15
b,B
2.18 ±
0.06
b,B
2.18 ±
0.33
b
3.51 ±
0.43
a,A,
*
cis-10-
Hep adecenoic
acid C17:1 (
ω
-7)
0.383 ±
0.010
b,C
0.408 ±
0.025ª
,b,
B
0.463 ±
0.045ª
,
B,
*
0.445 ±
0.051ª
,
B,
*
0.386 ±
0.022
b,B
Oleic acid C18:1
(
ω
-9)
37.2 ±
3.8
b,c, A,
B,
*
34.6 ±
1.9
c,d
38.5 ±
3.6
b,c,A,
B,
*
42.8 ±
3.9
b,A,
*
56.6 ±
3.7
a,A,
*
cis-11-Eicosenoic
acid C20:1 (
ω
-9)
0.362 ±
0.039
A,B
0.351 ±
0.068
0.356 ±
0.048
0.365 ±
0.006
A,B
0.366 ±
0.078
PUFA
Linoleic acid C18:2
(
ω
-6)
38.2 ±
4.9
a
37.4 ±
1.7
a,b
31.2 ±
3.5
b,c,B,
*
29.2 ±
4.2
c,B,
*
14.6 ±
5.4
d,B,
*
Linolenic acid
C18:3 (
ω
-3)
1.03 ±
0.08
b,c,
B,
*
1.16 ±
0.9
a,b,B
0.93 ±
0.1
c,B,
*
0.83 ±
0.21
c,B,
*
0.28 ±
0.07
d,B,
*
cis-11,14-
Eicosadienoic
acid C20:2 (
ω
-6)
0.152 ±
0.012ª
,B
0.153 ±
0.007ª
0.166 ±
0.019ª
0.148 ±
0.031
a
-
b,B,
*
To al con en s and Fa y acid indexes
SFA 20.9 ±
1.3
b,A,B,
*
24.0 ±
0.5
a,A
26.3 ±
0.6
a,A
24.1 ±
1.1
a,A
24.3 ±
1.8
a
MUFA 39.7 ±
3.8
c,
*
37.3 ±
1.9
c
41.5 ±
3.6
b,c,A,
B,
*
45.8 ±
4.1
b,A,
*
60.8 ±
3.6
a,A,
*
PUFA 39.3 ±
4.8
a
38.7 ±
1.6
a,b
32.3 ±
3.6
b,c,B,
*
30.1 ±
4.4
c,B,
*
14.9 ±
5.3
d,B,
*
ω
-6 38.3 ±
4.8
a
37.6 ±
1.7
a,b
31.3 ±
3.6
b,c,B,
*
29.3 ±
4.2
c,B,
*
14.6 ±
5.4
d,B,
*
ω
-3
1.03 ±
0.08
b,c,
B,
*
1.16 ±
0.9
a,b,B
0.93 ±
0.1
c,B,
*
0.83 ±
0.21
c,B,
*
0.28 ±
0.07
d,B,
*
PUFA:SFA 1.89 ±
0.33
a
1.61 ±
0.05
a,b
1.23 ±
0.15
b,B,
*
1.26 ±
0.22
b,B,
*
0.63 ±
0.27
c,B,
*
(con inued on nex page)
M.L. Mo ales e al.
Food Resea ch In e na ional 209 (2025) 116223
4
hei alues below he o al alues o MUFA and PUFA, despi e he de-
c eases o he la e , excep o 100 % o ALP die , in his case i was
signi ican ly highe han o al PUFA con en .
Alpe ujo, de i ed om oli es, was ich in C18:1
ω
-9, esul ing his
a y acid o be he mos abundan in he subs a e (Table S2, supple-
men a y), ollowed by palmi ic, linoleic, and s ea ic acids. The high
C18:1 con en in his subs a e esul s in he en ichmen o la ae ed
wi h i , eaching in he ALP100 samples pe cen ages simila o hose in
he subs a e.
Rega ding he POS ea ing subs a e (Table 2), a simila end was
obse ed. The pe cen age o o al MUFA con en inc eased signi ican ly
in mos samples compa ed o he con ol, excep a 40 %, wi h he
highes signi ican alue being achie ed in he 100 % POS die . Mo e-
o e , a di ec co ela ion was obse ed be ween he inc ease in o al
MUFA con en and he pe cen age o POS in he die ( =0.88). The
lowes signi ican pe cen age o o al PUFA con en was also obse ed
wi h he 100 % POS die . As wi h he ALP ea ing subs a e, he mos
ele an changes occu ed in he con en o MUFA and PUFA. Thus, as in
he con ol samples, he mos abundan a y acid was C18:2 when he
die con ained 20 % and 40 % POS. Howe e , when he pe cen age o
POS exceeded 40 %, C18:1 became he mos abundan a y acid. This
esul shows he in luence o he subs a e included in he die since, as
occu ed wi h ALP, POS subs a e p esen s a high con en o C18:1 and
o al MUFA (Table S2, supplemen a y). Rega ding SFA, he amoun was
simila o ha ound in he c ushed T. moli o ed wi h ALP. In mos
cases, o al SFA con en was signi ican ly lowe han MUFA and PUFA
con en s. Howe e , in samples ed wi h 100 % POS, he dec ease in
C18:2 made palmi ic acid he second mos abundan FA a e C18:1, and
he o al SFA con en became signi ican ly highe han PUFA con en ,
simila o wha occu ed wi h he ALP subs a e.
In he case o T. moli o la ae ed wi h he OLF subs a e (Table 3),
he o al PUFA con en s we e signi ican ly highe han SFA and MUFA
con en s in all cases excep wi h he 100 % OLF die , in which a sub-
s an ial inc ease in o al MUFA con en was obse ed due o he ise in
C18:1 con en . Con e sely, he o al SFA con en was signi ican ly lowe
in all cases, showing an in e se co ela ion wi h he pe cen age o OLF
used in he die ( = − 0.90). In hese samples, he palmi ic acid con en
was he lowes , while signi ican inc eases in s ea ic and a achidic acids
we e obse ed when he wo highes pe cen ages o OLF we e consumed
by la ae. Addi ionally, unlike wi h he wo p e ious subs a es, an in-
c ease in C18:3 con en was obse ed in hese samples, di ec ly
co ela ed wi h he pe cen age o OLF subs a e ( =0.89) and s a is i-
cally signi ican when he die con ained mo e han 20 % o OLF. This
may be because his subs a e had he highes con en o C18:3 a y
acids (Table S2, supplemen a y). This sugges s ha OLF may be a
p omising subs a e o enhancing
ω
-3 a y acids le els in T. moli o
la ae a ac ion.
O e all, he mos impo an change obse ed in ou expe imen s was
he dec ease o C18:2 con en s whils C18:1 con en s inc eased as he
pe cen age o con ol subs a e dec eased o ALP and POS subs a es. A
simila end was obse ed by Ko sou e al. (2023) when M. olei e a
lea es we e added a di e en a es o in he T. moli o die .
When compa ing he a y acid p o ile o c ushed T. moli o ed wi h
he same pe cen age o addi ion o he h ee di e en subs a es s udied,
he highes o al PUFA con en s we e obse ed o OLF a 60, 80 and
100 %. In con as , he highe o al MUFA con en was ound in he case
o ALP and POS a 80 and 100 %; whe eas he lowes o al SFA con en
was ob ained wi h he die s o 40, 60 and 80 % o POS. Fu he mo e, he
highes o al con en s o
ω
-3 we e achie ed in he case o OLF wi h
addi ion pe cen ages be ween 40 and 100 %. Hence, OLF subs a e could
no only enhance he p esence o
ω
-3, as men ioned abo e, bu also he
o al con en o PUFA in he a ac ion o hese la ae.
Se e al au ho s ha e ca ied ou assays o compa e he e ec o
di e en die in he nu i ional composi ion o T. moli o la ae. Lawal
e al. (2021) s udied conc e ely he e ec o se e al seed such as lax
seed, chia seed, hemp seed, and apeseed a di e en pe cen age on FA
p o ile, using whole-whea meal as con ol die . These die s inc emen ed
he amoun o a in he la ae. They ound le els o o al SFA and MUFA
con en s highe han ou alues and o al PUFA alues clea ly lowe han
hose ob ained by us. On he con a y, when mush oom spen co n
s o e , highly dena u ed soybean meal, and spi i dis ille s’ g ains we e
used as subs a e o ea T. moli o la ae (Zhang e al., 2019), he alues
o SFA and MUFA we e lowe han hose o ou samples and he o al
PUFA alues we e highe han ou ones.
3.2. Heal h ela ed indexes
The di e en esul s ela i es o he o al con en s o SFA, MUFA and
PUFA can make us hink ha bo h he c ushes o T. moli o and hei a
ac ion isola e may ha e a di e en impac on he consume ’s heal h
depending on he ea ing die used. Fo his eason, di e en die a y
indica o s, ela ed o he consume ’s heal h, we e calcula ed.
PUFA:SFA a io is an index ha has been ela ed wi h a a ou able
e ec o die a y a agains he ca dio ascula diseases (CVD) (Chen &
Liu, 2020), among o he s. In he case o he die s wi h he addi ion o
di e en pe cen age o ALP, he e was no signi ican di e ences espec
o he con ol, excep o he 100 % ALP die , being signi ican ly lowe
han he con ol (Table 1). In he case o POS addi ion, he alues o his
a io we e signi ican lowe when pe cen age equal o highe han 60
we e used (Table 2). Howe e , he samples o T. moli o ed wi h a die
wi h 80 % o OLF, eached he highes signi ican alues. In simila way
ha occu wi h ALP subs a e, he only case wi h alues signi ican ly
lowe han he con ol was 100 % o OLF, al hough he a io was no as
low as wi h he o he wo subs a es (Table 3). Thus, he lowes alues o
PUFA:SFA a io we e p esen ed by ALP and POS a 100 %, bu all
showed alues highe han ecommended 0.4 (Simopoulos, 2003). Chen
and Liu (2020) ca ied ou a e iew in which included he alues o
se e al die a y o nu i ional indexes ela ed o he a y acid con en
om di e en ood p o ein sou ces. Hence, we e i ied ha he alues o
he PUFA:SFA a io o ou samples we e highe han hose o se e al
dai y p oduc s, many mea s and simila o hose o shell ish and ish.
The e o e, he a y acid p o ile o ou T. moli o c ushed p o ides
g ea e ca dio ascula bene i s han some ep esen a i e human die
p o ein oods, especially when he OLF subs a e was used in pe cen age
less han 100.
The a he ogenici y index (AI) ela es he con en o SFA conside ed
p o-a he ogenic and he unsa u a ed a y acids desc ibed as an i-
Table 2 (con inued)
Samples POS20 POS40 POS60 POS80 POS100
MUFA:PUFA 1.03 ±
0.24
b
0.96 ±
0.09
b
1.30 ±
0.25
b
1.56 ±
0.38
b,A
4.45 ±
1.58
a,A,
*
ω
-6:
ω
-3 37.6 ±
6.8
a,b,A
32.8 ±
4.2
a,b
33.7 ±
2.5
a,b,A
36.4 ±
5.6
a,b
55.9 ±
30.5
a,A,
*
AI 0.31 ±
0.03
b,
*
0.34 ±
0.05
a,b
0.38 ±
0.03
a,A
0.367 ±
0.013
a,b,
A
0.39 ±
0.06
a,A
TI 0.43 ±
0.05
b,
*
0.51 ±
0.04
a,b,A
0.58 ±
0.05
a,A
0.55 ±
0.03
a,A
0.58 ±
0.07
a,A
h/H 4.9 ±
0.8
a,
*
4.1 ±
0.5
a,b
3.5 ±
0.4
b,B
3.8 ±
0.3
b,B
3.7 ±
0.5
b,B
HPI 3.3 ±
0.3
a,
*
3.0 ±
0.4
a,b
2.65 ±
0.22
b,B
2.73 ±
0.10
a,b,B
2.6 ±
0.4
b,B
-: no peak de ec ed; n.q.: amoun unquan i iable, n.d.: amoun unde ec able.
SFA: sa u a ed a y acid; MUFA: monounsa u a ed a y acid; PUFA: poly-
unsa u a ed a y acid; AI: a he ogenic index; TI: h ombogenic index; h/H:
hypocholes e olemic/hype choles e olemic a io; HPI: heal h-p omo ing index.
*Mean signi ican di e ence wi h con ol sample ( alues in Table 1) p <0.05.
Di e en lowe case le e in he same line means signi ican di e ences (p <
0.05) be ween samples ed wi h he same subs a e wi h di e en pe cen age
(20–100 %).
Di e en capi al le e means signi ican di e ences (p <0.05) be ween samples
o la ae ed wi h he same pe cen age o he h ee subs a es s udied.
M.L. Mo ales e al.
Food Resea ch In e na ional 209 (2025) 116223
5
a he ogenic (O kusz e al., 2024). Thus, a a wi h low AI is conside ed
heal hy. The alues o AI in ou samples anged om 0.231 (HHO100) o
0.398 (C1) (Tables 1–3). I we compa e hem wi h o he oods which a e
ich in p o eins, ou samples p esen ed alues much lowe han dai y
p oduc s, wi h alues a leas h ee imes highe , lowe han mos o
mea s and simila o lowe han ish (Chen & Liu, 2020). The di e en
die s s udied modi ied his index in di e en ways espec o he con ol
die . Thus, in he die in which Alpe ujo was added, he alues o his
index dec eased posi i ely, excep in 100 % ALP, due o he impo an
dec ease o
ω
-6 a y acid con en s in his sample. On he con a y, in he
die con aining POS, only in he case o he lowes pe cen age o addi ion
p esen ed signi ican ly lowe alues han in he con ol. In he case o
OLF subs a e, he a ou able e ec on heal h occu s wi h pe cen ages
equal o o g ea e han 60.
Rega ding he h ombogenici y index (TI), ha e lec s he p edis-
posi ion o p opensi y o ce ain sa u a ed a y acids o o m clo s in
blood essels (Ulb ich & Sou hga e, 1991) agains he an i h ombo-
genic e ec o ce ain unsa u a ed a y acids. The e o e, his index may
show he posi i e e ec o die a y a agains CVD when i s alue is low.
The inclusion o ALP in he die o T. moli o was bene icial when
pe cen age be ween 20 and 60 we e used, while in he case o OLF
subs a e is necessa y o go up o pe cen ages o he o de o 80 o 100.
The end o his index in T. moli o samples ed wi h POS is equal o AI.
Ou esul s showed ha ou p oduc s we e again heal hie han mea
and dai y p oduc s (Chen & Liu, 2020). Fo bo h indexes, AI and TI,
compa ing he same pe cen age o he h ee di e en subs a es in he
die , lowe signi ican alues we e only obse ed o 80 and 100 % o
OLF.
The hypocholes e olemic/hype choles e olemic (h/H) a io was also
calcula ed as an indica o o he ca dio ascula heal h (CVH) p omo ion
o a die a y. The alues de e mined in ou s udy anged om 3.54
(POS60) o 6.89 in OLF100. Fo his index, he alues o ou samples
we e highe han ish, mea , dai y p oduc s, and mos o shell ish (Chen
& Liu, 2020). Respec o he alues o he con ol samples, he only case
ha showed signi ican highe alues we e ALP40 and 60; POS20;
OLF80 and 100.
Table 3
To al a con en and p o ile o a y acids o T. moli o c ushed samples ed wi h con ol and wi h di e en pe cen age o OLF subs a es (means alues o iplica e o
biological assays ±SD).
Samples Con ol-2 OLF20 OLF40 OLF60 OLF80 OLF100
To al a con en (%w/w) 26.3 ±6.6
a,b
28.5 ±1.8
a,A
23.6 ±1.4
a,b,c
19.7 ±4.3
c
21.1 ±0.4
b,c,B
11.1 ±1.4
d,B
Fa y acid con en (% o o al a y acids con en )
SFA
Bu anoic acid C4:0 0.79 ±0.3 0.98 ±0.76 0.35 ±0.22 0.79 ±0.36 0.96 ±0.53 0.63 ±0.54
Dodecanoic acid C12:0 0.415 ±0.027
a
0.384 ±0.008
a,b,A
0.360 ±0.016
b,c
0.328 ±0.012
c,d,A
0.298 ±0.014
d,e
0.270 ±0.061
e,B
T idecanoic acid C 13:0 0.151 ±0.007
a
0.139 ±0.003
a,b,A
0.148 ±0.009
a,A
0.144 ±0.008
a,b,A
0.146 ±0.001
a,b,A
0.125 ±0.025
b,A
My is ic acid C14:0 3.55 ±0.007
a
3.12 ±0.11
b,A
3.08 ±0.11
b
2.99 ±0.10
b,A
2.76 ±0.03
b
2.39 ±0.48
c
Pen adecanoic acid C 15:0 0.290 ±0.015
d
0.286 ±0.012
d
0.312 ±0.009
c
0.314 ±0.020
c,A,B
0.369 ±0.009
b
0.433 ±0.012
a,A
Palmi ic acid C16:0 13.2 ±0.2
a
12.8 ±0.3
a
12.7 ±1.7
a
12.2 ±0.5
a,B
9.9 ±1.6
b,C
8.6 ±1.6
b,B
Hep adecanoic acid C17:0 0.443 ±0.025
d
0.450 ±0.026
d,A,B
0.539 ±0.027
c
0.570 ±0.014
c,A
0.762 ±0.006
b,A
0.933 ±0.058
a,A
S ea ic acid C18:0 2.52 ±0.07
d
2.68 ±0.07
c,d
2.68 ±0.06
c,d
2.89 ±0.08
c,A,B
3.24 ±0.08
a,B
4.86 ±0.27
a,A
A achidic acid C20:0 0.593 ±0.070
c
0.614 ±0.023
c
0.549 ±0.077
c
0.622 ±0.054
c
0.796 ±0.056
b,A
1.30 ±0.12
a,A
Heneicosanoic acid C21:0 n.q.
c
n.q.
c
n.q.
c
n.q.
c
0.039 ±0.006
b,A
0.052 ±0.004
a,A
Behenic acid C22:0 0.155 ±0.015
c
0.205 ±0.014
b,c
0.168 ±0.051
c
0.203 ±0.048
b,c
0.275 ±0.046
b,A
0.538 ±0.059
a,A
T icosanoic acid C23:0 n.q.
c
n.q.
c
n.q.
c,B
0.010 ±0.017
c
0.045 ±0.005
b,A
0.082 ±0.010
a,A
Lignoce ic acid C24:0 n.q. n.q. n.q. 0.022 ±0.038 n.q. 0.019 ±0.033
MUFA
Palmi oleic acid C16:1 (
ω
-7) 2.50 ±0.03
a
2.38 ±0.05
b,c,A
2.38 ±0.04
b,c,A
2.41 ±0.02
b,A
2.35 ±0.03
b,c
2.32 ±0.04
c,B
cis-10-Hep adecenoic acid C17:1 (
ω
-7) 0.547 ±0.02
d
0.562 ±0.002
d,A
0.615 ±0.023
c,d,A
0.658 ±0.012
c,A
0.870 ±0.076
b,A
1.17 ±0.10
a,A
Oleic acid C18:1 (
ω
-9) 33.1 ±0.8
b,c
32.7 ±1.0
c,B
33.7 ±1.0
b,c
34.9 ±1.3
b,B
33.7 ±1.1
b,c,B
46.6 ±1.3
a,B
cis-11-Eicosenoic acid C20:1 (
ω
-9) 0.328 ±0.027
c,d
0.317 ±0.017
d,B
0.357 ±0.008
b
0.354 ±0.006
b,c
0.356 ±0.005
b,B
0.453 ±0.013
a
PUFA
Linoleic acid C18:2 (
ω
-6) 40.1 ±1.0
a
41.1 ±1.9
a
40.5 ±2.5
a
39.0 ±1.3
a,A
41.4 ±2.3
a,A
27.1 ±0.9
b,A
Linolenic acid C18:3 (
ω
-3) 1.18 ±0.02
c
1.27 ±0.04
c,A
1.45 ±0.09
b,A
1.42 ±0.10
b,A
1.51 ±0.12
b,A
2.00 ±0.06
a,A
cis-11,14-Eicosadienoic acid C20:2 (
ω
-6) 0.126 ±0.006
c
n.q.
d,C
0.150 ±0.017
a,b
0.164 ±0.013
a
0.149 ±0.021
a,b
0.130 ±0.004
b,c,A
To al con en s and Fa y acid indexes
SFA 22.1 ±0.1
a
21.7 ±0.9
a,b,B
20.9 ±1.6
a,b,B
21.1 ±0.3
a,b,B
19.6 ±1.1
b,C
20.2 ±2.2
a,b
MUFA 36.4 ±0.9
b,c
36.0 ±1.1
c
37.0 ±0.9
b,c
38.3 ±1.3
b,B
37.3 ±1.0
c,b,B
50.6 ±1.4
a,B
PUFA 41.4 ±0.9
a
42.4 ±1.9
a
42.1 ±2.4
a
40.6 ±1.2
a,A
43.1 ±2.1
a,A
29.2 ±0.9
b,A
ω
-6 40.2 ±1.0
a
41.1 ±1.9
a
40.6 ±2.4
a
39.2 ±1.3
a,A
41.6 ±2.2
a,A
27.2 ±0.9
b,A
ω
-3 1.18 ±0.02
c
1.27 ±0.04
c,A
1.45 ±0.09
b,A
1.42 ±0.10
b,A
1.51 ±0.12
b,A
2.00 ±0.06
a,A
PUFA:SFA 1.87 ±0.05
b
1.96 ±0.17
a,b
2.03 ±0.28
a,b
1.93 ±0.06
a,b,A
2.21 ±0.23
a,A
1.46 ±0.21
c,A
MUFA:PUFA 0.88 ±0.04
b
0.85 ±0.06
b
0.88 ±0.07
b
0.94 ±0.06
b
0.87 ±0.07
b,B
1.73 ±0.04
a,B
ω
-6:
ω
-3 34.1 ±1.3
a
32.3 ±1.2
a,b,A,B
28.2 ±3.4
b,c
27.7 ±2.9
c,B
27.8 ±3.6
c
13.57 ±0.04
d,B
AI 0.36 ±0.01
a
0.33 ±0.01
a,b
0.32 ±0.01
a,b
0.31 ±0.01
b,B
0.26 ±0.03
c,B
0.23 ±0.04
c,B
TI 0.46 ±0.01
a
0.44 ±0.01
a
0.43 ±0.04
a,A,B
0.42 ±0.01
a,B
0.36 ±0.04
b,C
0.35 ±0.05
b,B
h/H 4.3 ±0.1
c
4.6 ±0.2
c
4.7 ±0.6
c
4.9 ±0.2
b,c,A
6.0 ±0.8
a,b,A
6.9 ±1.4
a,A
HPI 2.8 ±0.0
c
3.1 ±0.1
c
3.1 ±0.2
c
3.2 ±0.1
b,c,A
3.8 ±0.4
a,b,A
4.4 ±0.8
a,A
n.q.: amoun unquan i iable, n.d.: amoun unde ec able.
SFA: sa u a ed a y acid; MUFA: monounsa u a ed a y acid; PUFA: polyunsa u a ed a y acid; AI: a he ogenic index; TI: h ombogenic index;
h/H: hypocholes e olemic/hype choles e olemic a io; HPI: heal h-p omo ing index.
Di e en lowe case le e in he same line means signi ican di e ences (p <0.05) be ween samples ed wi h he same subs a e wi h di e en pe cen age (Con ol-100
%).
Di e en capi al le e means signi ican di e ences (p <0.05) be ween samples o la ae ed wi h he same pe cen age o he h ee subs a es s udied.
M.L. Mo ales e al.
Food Resea ch In e na ional 209 (2025) 116223
6
Finally, we also conside ed he heal hy-p omo ing index (HPI),
al hough his index is he in e se o he AI (Chen & Liu, 2020). Fo ha ,
he e ec o di e en pe cen age o subs a es s udied p o ided simila
esul s o AI. Howe e , o his index, when we compa e all s udied
samples, he sample OLF100 showed he highes signi ican alues.
I we compa e he alues o indices ela ed wi h ca dio ascula
heal h ob ained in his wo k wi h hose ob ained by o he au ho s, we
obse ed simila esul s o e en be e . In he case o O kusz e al.’s
s udy (2024), in which T. moli o om ma ke we e analysed, hese
u ned ou o be less heal hy han ou s, showing PUFA:SFA and h/H
a ios lowe and TI and AI alues highe han ou s. Mo eo e , in he case
o he inclusion o ege al was es (L´
opez-G´
amez e al., 2024) o
M. olei e a lea es (Ko sou e al., 2023) in he T. moli o ’s die , he PUFA:
SFA a ios we e simila o ou s, bu TI and AI alues we e highe and h/
H a ios and HPI lowe han ou s, showing ha he die s assayed by us
seem o be sligh ly mo e heal hie , specially espec o he las kind o
die .
Ano he impo an ac o in a heal hy die is a balanced in ake o
omega-6 e sus omega-3 a s. The desi ed
ω
-6 o
ω
-3 a io seem o be
be ween 1/1 and 5/1 (Lupe e & Benning, 2020). In ou case, he a
ac ion o la ae ed wi h modi ied die s as well as wi h con ol die
showed high
ω
-6:
ω
-3 a ios compa ed o he ecommended alues, he
lowes signi ican alue was ound in he case o he OLF100 die . In he
case o hese insec s, when di e en die we e s udied, only he inclusion
o a pe cen age o 10 % o lax o chia seed was able o adequa ely he
alues o his a io (F anca di e al., 2017; Lawal e al., 2021). The e o e,
his a io would be easily imp o ed wi h he addi ion o a sou ce o
ω
-3.
Fu he mo e, i has ecen ly been sugges ed ha
ω
-9 FA may be as
impo an as
ω
-3 and
ω
-6 FAs o human heal h (Wang e al., 2024). The
a ac ions ob ained in ou s udy we e ich in
ω
-9 MUFAs, and hei
con en s anged be ween 33.04 and 61.42 %. Se e al au ho s ha e
sugges ed ha his ype o FA may con ibu e posi i ely o ca dio as-
cula diseases, neu odegene a i e diseases, and exe an an i-
in lamma o y e ec (Hu e al., 2021; San a-Ma ía e al., 2023; Wahle
e al., 2004). Al hough
ω
-9 MUFAs can be syn hesized endogenously by
he human body he main sou ce is he die (Wang e al., 2024). How-
e e , a speci ic ecommended in ake ange has no ye been de ined.
Since he MU a ac ion is p ima y consis ed o
ω
-9 MUFA, he
ω
-9
MUFA in ake ecommenda ions could be p ac ically equa ed wi h MUFA
in ake.
Among he di e en
ω
-9 ha exis , we ha e ound in ou samples
wo, oleic acid and cis-11-eicosenoic acid (gondoic acid). The i s , as i
has been poin ed abo e, is he mos abundan a y acid in all la ae
samples ed wi h ALP subs a e, in hose ed wi h a pe cen age o POS
equal o g ea e han 60 and in he case o OLF100 die .
Among he heal hy p ope ies a ibu ed o his FA, we can highligh
he possible con ibu ion o imp o ing he a he oscle o ic p ocess and
plaque s abili y (Pe domo e al., 2015). Mo eo e , cis-11-eicosenoic acid
seems o ha e e ec on oxida i e s ess ha p oduce sho - e m memo y
de ici (Cojoca iu e al., 2020). Ano he heal hy e ec is eplacing some
sa u a ed a y acids (SFA) wi h
ω
-9 MUFAs in he die (Maki e al.,
2018). In ou case, all samples p esen ed high alues o
ω
-9 espec o
SFA con en s, especially in he case o ALP subs a e.
Hence, in making an assessmen o he esul s o he mos ele an
heal h ela ed indexes p esen ed by he samples s udied, s ood ou he
a ac ion o T. moli o ed wi h 80 % and 100 % o OLF.
Howe e , he weaknesses o his s udy should be conside ed which
a e ha hese heal h ela ed indexes a e ma hema ical calcula ions om
he esul s de i ed o labo a o y scale expe imen s and, he e o e, o
con i m hei heal hy e ec , u u e assays in animal models could be
necessa y.
3.3. Heal h claims
The heal h- ela ed o ganiza ions ha e gi en a ious ecommenda-
ions on a in ake. On one hand, acco ding o he WHO ecommenda-
ion (2023), in he die o adul s, he 30 % o he o al ene gy in ake
should come om a s, wi h no mo e han 10 % o his o al ene gy
in ake coming om SFA. On he o he hand, Eu opean Food Sa e y
Au ho i y (EFSA) (2017) ecommended ha he o al ene gy in ake om
a should be be ween 20 and 35 % maximum and he pe cen age om
SFA should be as low as possible. Mo eo e , he adequa e in ake ec-
ommended o linoleic acid is 4 % o o al ene gy in ake and 0.5 % o
al a-linolenic acid. Conside ing hese ecommenda ions, we will p oceed
Fig. 1. Pe cen age o o al SFA, MUFA and PUFA con en s in c ushed o T. moli o la ae’s a ac ion ed wi h di e en subs a e: Con ol die (100 % o whea
b an), S abilized Alpe ujo (ALP), Pleo o us os ea us spen subs a e (POS) and oli e lea lou (OLF) mixed wi h di e en pe cen age o whea b an.
M.L. Mo ales e al.
Food Resea ch In e na ional 209 (2025) 116223
7
o analyze he quali ies o he a ac ion o he c ushed p oduc s ob-
ained wi h die s modi ied wi h was e.
Acco ding o he popula ion e e ence in ake (PRI) o ene gy o
adul s by Eu opean Food Sa e y Au ho i y (2013), his should be be-
ween 2000 and 2600 Kcal/day. Conside ing an a e age alue o ene gy
o 2300 Kcal/day, 100 g o c ushed T. moli o ob ained in his wo k
would p o ide be ween 4.3 and 11.8 % o he o al PRI o ene gy p o-
ided by he a ac ion, mos o hem wi h alues a ound 8–9 % o he
PRI o ene gy.
Fu he mo e, be ween 0.9 and 2.7 % o his ene gy come om SFA,
2.2–7.6 % om MUFA and 1.3–4.7 % om PUFA, i.e. mainly MUFA and
PUFA.
In addi ion, he con en linoleic acid was close o he alues equi ed
o mee he ecommended adequa e in ake equi emen s o his a y
acid since, mos o hem, p o ided mo e han 3 % o he o al PRI o
ene gy.
I we ocus on he ex ac ed a ac ion and i would be ma ke ed as
ood p oduc , acco ding o Eu opean egula ions on nu i ional claims
allowed o be included in a label (Eu opean Commission, 2006), hese
could be labelled as “sou ce o
ω
-3” excep ALP100, POS100 and OLF20,
since hei con en s exceed 0.3 g o
α
-linolenic acid in 100 g o p oduc .
Some o hem, such as ALP60, ALP80, ALP100, POS80, POS100 and
OLP100, also mee he equi emen s o be e e ed o as “high in
monounsa u a ed a ” con ibu ing o he o al con en o a y acids
wi h pe cen ages anging be ween 45.8 % and 64.6 %, mo e han 45 %
o he equi ed a y acids and also ep esen ing mo e han 20 % o he
ene gy alue o he p oduc . Finally, all o hem can be labelled as “high
in unsa u a ed a s” because hei con en (73.7–80.4 %) exceeded he
equi ed alue (70 %).
4. Conclusions
The use o di e en pe cen ages o alpe ujo, mush oom spen sub-
s a e and oli e lea lou in he die o T. moli o la ae in luences bo h
he o al a con en s and he a y acid p o iles.
The a y acid p o iles expe ienced changes, especially no able in he
case o die spiked wi h he ALP subs a e, compa ed o he con ol die .
The mos ema kable change was he inc ease o oleic acid, a
ω
-9 a y
acid, which ha e ecen ly been linked o heal h bene i s. This ac is o
g ea ele ance since, among he h ee subs a es s udied, he one wi h
he g ea es ecological in e es is he euse o Alpe ujo, as i is a highly
p oduc i e was e and he mos pollu ing o he h ee. Mo eo e , OLF
subs a e enhanced he p esence o
ω
-3 and he o al con en o PUFA in
he a ac ion o hese la ae.
Rega ding he impac on he consume ’s heal h o he a ac ion o
c ushed T. moli o la ae ob ained in his wo k, o mos o heal h ela ed
indexes we ob ained mo e adequa e alues han hose o o he con-
en ional p o ein sou ces. In addi ion, in all cases he e was an inc ease
o he o al con en o unsa u a ed a y acids wi h espec o ha o
sa u a ed. In addi ion, high pe cen ages o OLF subs a e in he die
p esen ed he mos posi i e esul s espec o heal h indexes. Hence, he
was e om oli e oil p oduc ion appea ed o p o ide he heal hies a
ac ion.
Al hough mos o c ushed T. moli o ob ained p esen ed high o al a
con en s, i was a heal hy a ac ion, ich in unsa u a ed a s, especially
ω
-9, and may be conside ed, in some cases, as “sou ce o
ω
-3”, which
would make his ac ion sui able as an ing edien o imp o e he
heal hiness o ood.
CRediT au ho ship con ibu ion s a emen
Mo ales M. Lou des: W i ing – o iginal d a , Me hodology, In es-
iga ion, Fo mal analysis, Concep ualiza ion. M. Pila Segu a-Bo ego:
Visualiza ion, In es iga ion, Fo mal analysis. Jos´
e Raúl Aguile a-
Vel´
azquez: Resou ces, In es iga ion. Raquel M. Callej´
on: W i ing –
e iew & edi ing. Daniel Gu i´
e ez-P aena: W i ing – e iew & edi ing,
P ojec adminis a ion, Funding acquisi ion. C is ina Ubeda: W i ing –
e iew & edi ing, P ojec adminis a ion, Funding acquisi ion.
Funding sou ces
This wo k is pa o he R +D +i p ojec TED2021-129351 A-I00,
unded by MICIU/AEI/10.13039/501100011033/ and by he “Eu o-
pean Union Nex Gene a ionEU/PRTR”.
Decla a ion o compe ing in e es
The au ho s decla e ha hey ha e no known compe ing inancial
in e es s o pe sonal ela ionships ha could ha e appea ed o in luence
he wo k epo ed in his pape .
Acknowledgmen
The au ho s wish o hank Almaza a Vi gen de los ´
Angeles o sup-
plying he alpe ujo.
Appendix A. Supplemen a y da a
Supplemen a y da a o his a icle can be ound online a h ps://doi.
o g/10.1016/j. ood es.2025.116223.
Da a a ailabili y
Da a will be made a ailable on eques .
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