molecules

Review
Role of Lipids in Food Flavor Generation
Fereidoon Shahidi * and Abul Hossain

Department of Biochemistry, Memorial University of Newfoundland, St. John’s, NL A1C 5S7, Canada
* Correspondence: [email protected]

Abstract: Lipids in food are a source of essential fatty acids and also play a crucial role in flavor
and off-flavor development. Lipids contribute to food flavor generation due to their degradation
to volatile compounds during food processing, heating/cooking, and storage and/or interactions
with other constituents developed from the Maillard reaction and Strecker degradation, among
others. The degradation of lipids mainly occurs via autoxidation, photooxidation, and enzymatic
oxidation, which produce a myriad of volatile compounds. The oxidation of unsaturated fatty
acids generates hydroperoxides that then further break down to odor-active volatile secondary lipid
oxidation products including aldehydes, alcohols, and ketones. In this contribution, a summary of the
most relevant and recent findings on the production of volatile compounds from lipid degradation
and Maillard reactions and their interaction has been compiled and discussed. In particular, the
effects of processing such as cooking, drying, and fermentation as well as the storage of lipid-based
foods on flavor generation are briefly discussed.

Keywords: flavor chemistry; lipid oxidation; Maillard reaction; volatile formation

1. Introduction
Lipids, in addition to providing energy, texture, and mouthfeel, play an important role
in the odor and flavor development of food. This could be due to the ability of lipids to
Citation: Shahidi, F.; Hossain, A.
generate odors and flavors, act as precursors of odor and flavor compounds, or modify the
Role of Lipids in Food Flavor
odor and flavor of other components [1]. Lipids are responsible for both the undesirable
Generation. Molecules 2022, 27, 5014.
and desirable flavors of food; the oxidation of lipids mainly results in the development
https://doi.org/10.3390/
of off-flavor and lipoxygenase-derived lipid-based volatiles that are responsible for flavor
molecules27155014
generation. Food flavor is the manifestation of interactions between aroma, taste, and
Academic Editor: Michael C. Qian oral sensations, where the aroma is linked with mostly volatile compounds, and taste is
Received: 18 July 2022
associated with non-volatile high-molecular-weight components [2]. Generally, raw foods,
Accepted: 4 August 2022
mainly meat, provide little aroma and have a mild taste, but this can change to a strong
Published: 6 August 2022
aroma and taste upon processing. For example, odorants of raw and cooked sheep meat
have been reported to be due to cis-1,5-octadien-3-one (geranium-like), 4-ethyloctanoic acid
Publisher’s Note: MDPI stays neutral
(mutton-like), trans-4,5-epoxy-2-decenal (metallic), and trans-2,4-decadienal (deep-fried).
with regard to jurisdictional claims in
Moreover, flavor dilution (FD) factors of aroma compounds such as 2-acetyl-1-pyrroline,
published maps and institutional affil-
4-hydroxy-2,5-dimethyl-3(2H)-furanone, and 2-aminoacetophenone were clearly increased
iations.
upon cooking [3]. However, aquatic foods are slightly different as they have a comparatively
strong flavor in their raw state. This is because of the presence of amines, lipoxygenase-
derived lipid-based volatile components, and the development of secondary oxidation
Copyright: © 2022 by the authors.
products in the raw stage [4,5]. On the other hand, lipids with low and high volatility are
Licensee MDPI, Basel, Switzerland. almost odorless and tasteless, respectively, partly because of their non-polar nature.
This article is an open access article Due to the lipid oxidation in food, an off-flavor is developed, and some bioactive
distributed under the terms and compounds and fat-soluble vitamins are lost. The oxidation of unsaturated fatty acids is a
conditions of the Creative Commons complex process that can occur in the presence of oxygen or via non-enzymatic (autoxida-
Attribution (CC BY) license (https:// tion and photooxidation) and enzymatic (lipoxygenase) pathways. Lipid oxidation and the
creativecommons.org/licenses/by/ development of off-flavor compounds are catalyzed by light, heat, oxygen, photosensitiz-
4.0/). ers, and transition metal ions (e.g., Fe2+ and Cu2+ ). Autoxidation occurs in the presence

Molecules 2022, 27, 5014. https://doi.org/10.3390/molecules27155014 https://www.mdpi.com/journal/molecules

Molecules 2022, 27, 5014 2 of 16

of triplet oxygen (3 O2 ), while photooxidation happens in the presence of singlet oxygen

(1 O2 ) [6]. In autoxidation, primary oxidation products (e.g., hydroperoxides) are formed,
and their subsequent break down into volatile secondary lipid oxidation products such
as ketones, alcohols, and aldehydes, among others, are believed to play the main role in
the development of an off-flavor. Due to the multiple methylene-interrupted cis-double
bonds, polyunsaturated fatty acids (PUFAs) are highly prone to oxidation. For example,
edible oils and fish lipids are rich in omega-3 fatty acids, where propanal and acrolein act
as good indicators for assessing the degree of oxidation. Similarly, meat and meat products
are rich in omega-6 fatty acids, and hexanal serves as a reliable indicator for flavor deterio-
ration [7,8]. Moreover, heterocyclic compounds (e.g., pyrazines, oxazoles, pyridines, and
thiazole) are formed mainly via non-enzymatic browning reactions, which generally play a
key and desirable role in flavor generation [9]. On the other hand, upon thermal processing
and heating experienced during frying, cooking, and grilling, hundreds of compounds can
possibly be generated from lipid degradation, Maillard reactions, or Strecker degradation,
which contribute to the overall desirable aroma upon interacting with each other. This
could occur via forming a wide range of new volatile products upon interaction or entirely
or partially blocking the presence of volatiles from others [10]. For instance, at both the
initial and later stages of thermal processing, aldehydes that are produced upon the thermal
degradation of lipids could join in the Maillard reactions [11]. In addition, the thermal
generation of the oxidation products of lipids may be accompanied by the production of
polar lipids and polymers that have no real effect on flavor [12]. Hence, the aim of this
review is to summarize the most recent literature and knowledge of the role of lipids on
flavor formation and their physiology and chemistry. An overview of the flavor generation
of foods rich in PUFAs is also provided.

2. Physiology of Flavor
Flavor comprises taste, odor, aroma, and other mouth/oral cavity responses to texture,
which is linked to the overall olfactory response to the food taken. Hence, flavor develops
as a complex response of the mouth, tongue, nasal, and sinus cavities. Basic aroma, taste,
and somatosensory perception are sensed by the olfactory bulb, gustatory sensory cells,
and by trigeminal nerves, respectively [10]. Aroma, odor, and smell are synonymous terms
used to describe olfactory responses, though aroma mostly refers to good flavor, while
odor and smell are used to describe the off-flavor. Moreover, the term taste is related
to the tongue response to sour, sweet, bitter, salty, and umami (monosodium glutamate
(MSG)-like taste stimulated by MSG) sensations. Furthermore, astringent and metallic
senses in the nasal and oral cavities can play an important role in flavor. Taste is mainly
related to the high molecular weight of non-volatile lactic acid, quinine, sucrose, sodium
chloride, and tannins, while the aroma is a sensual response when volatiles hit the back
of the nose [2,13]. The interaction with the food matrix and static and dynamic factors is
linked to the release of volatiles from food during chewing. For example, mass transfer,
which is an example of a dynamic factor, mainly depends on the product viscosity and its
interfaces; thus, volatiles can release more slowly in a viscous medium. Therefore, lipids
act as modulators and transporters of volatiles in food [2].
The minimum concentration required to detect volatiles is known as the aroma thresh-
old, which is mainly measured in parts per billion (ppb) or parts per million (ppm). For
example, the thresholds for saturated, monounsaturated, and diunsaturated aldehydes
are 0.014–1, 0.04–2.5, and 0.002–0.6 ppm, respectively, while the thresholds for alcohol,
furans, and ketones are 0.001–3, 1–27, and 0.0002–5.5, respectively [6]. The polarity of the
flavor compounds plays an important role in the flavor development. Usually, the odor
and flavor potential is less in an oily medium than in an aqueous solution. Hence, low
polarity fatty acids, mainly long-chain fatty acids, have a higher flavor threshold in an
oily medium and a lower threshold in aqueous solutions. In contrast, polar compounds
such as short-chain fatty acids have lower and higher thresholds in oils and water, re-
spectively [1]. However, Maillard reaction-derived heterocyclic compounds containing
Molecules 2022, 27, 5014 3 of 16

sulfur and nitrogen have lower thresholds than those for lipid-derived volatiles. Therefore,
greater/higher concentrations of lipid-derived components are required to exhibit the
presence of aroma [10].

3. Role of Lipids Oxidation in Flavor or Off-Flavor Development

Lipids are susceptible to oxidation, which is believed to play the main non-microbial
role in the quality degradation of food and food products including meat, fish, and oil-
based products. Usually, off-flavors are linked very closely to lipids compared to proteins
and carbohydrates. Lipid oxidation not only lessens the nutritional benefits of products
due to the changes in essential fatty acids and vitamins but also negatively affects the
sensory qualities such as flavor, texture, and color, which impacts on the overall consumer
acceptance [6,14]. However, in some cases, lipid oxidation promotes the development
of pleasant aromas, mainly during the ripening or dry-cured stages of meat products,
which is one of the quality parameters of these products [15]. Lipid oxidation is a very
complex phenomenon that undergoes a variety of reactions and produces a wide range
of compounds. In a nutshell, fatty acids, mainly PUFAs, react with molecular oxygen and
produce primary oxidation products (hydroperoxides) via free radical mechanisms. These
compounds are highly unstable due to a weak oxygen–oxygen bond and are believed to
make no contribution to odor and aroma development, but they break down rapidly and
produce a wide variety of secondary constituents, which are responsible for the formation
of off-flavors [16]. However, not all of these constituents similarly contribute to the flavor
profiles, since the overall aroma perception is mainly dependent on the olfactory threshold,
concentration, and type of products. Among these compounds, aldehydes seem to be the
major compounds contributing to flavor development due to their low odor threshold and
higher concentration. The most common aldehydes generated from lipid oxidation are
n-alkanals, 4-hydroxy-2-alkenals, 2-alkenals, and malondialdehyde (MDA) [15]. Hence,
lipid oxidation can occur in various pathways including autoxidation, photooxidation,
thermal oxidation, and enzyme-catalyzed oxidation. Autoxidation is a very common lipid
oxidation pathway, occurring through a continuous free-radical chain reaction. However,
hydroperoxides are formed during the initiation stage of autoxidation, which is the main
difference between the mechanisms of photo and enzymatic oxidation [12].

3.1. Formation of Oxidation Products via Autoxidation

Autoxidation is a combination of three different phases, namely, initiation, propaga-
tion, and termination. These steps are mainly responsible for producing radicals, followed
by the multiplication of reactive compounds, finally degrading or reacting with each other
in order to produce non-reactive products [17]. Autoxidation is the spontaneous oxidation
of lipids in the presence of oxygen, mainly 3 O2 , which can interact with radical species
due to its diradical nature. However, the interaction between unsaturated fatty acids
(singlet electronic state) and oxygens (triplet electronic state) is not possible because of
their chemical natures. Moreover, the triplet oxygen (3 O2 ) cannot change its state to singlet
states (1 O2 ) without external support. Therefore, an initiator is necessary to convert 3 O2
into 1 O2 , or reactive oxygen species (ROS) such as hydroxyl radicals, hydrogen peroxide
(H2 O2 ), superoxide anions, or to generate a radical by removing an electron from the
unsaturated lipids [6,15]. The activation of oxygen mainly occurs in the presence of light,
temperature, or metal ions such as Fe2+ and Cu2+ . At the end of the propagation stage,
primary oxidation products such as lipid hydroperoxides and conjugated dienes/trienes
are produced and these further break down into a series of secondary oxidation products
such as aldehydes, ketones, alcohols, hydrocarbons, epoxy compounds, and volatile or-
ganic acids. Among them, some compounds are responsible for the off-flavor at very low
threshold values [12]. For example, linoleic acid produces 9- and 13-hydroperoxides in
the presence of oxygen, and finally generates various volatiles including 2,4-decadienal,
2-octenal, and hexanal, among others, due to the cleavage of the C–O bond by alkoxy
radicals derived from these hydroperoxides (Figure 1). Similarly, oleic acid forms 8-, 9-,
Molecules 2022, 27, x FOR PEER REVIEW 4 of 17

[12]. For example, linoleic acid produces 9- and 13-hydroperoxides in the presence of ox-
Molecules 2022, 27, 5014
ygen, and finally generates various volatiles including 2,4-decadienal, 2-octenal, and4 of 16
hex-
anal, among others, due to the cleavage of the C–O bond by alkoxy radicals derived from
these hydroperoxides (Figure 1). Similarly, oleic acid forms 8-, 9-, 10-, and 11-hydroper-
10-, and 11-hydroperoxides,
oxides, which leads to many which leads tocompounds
secondary many secondary compounds
such as pentanal, such as pentanal,
octanal, hexanal,
octanal, hexanal,
and heptanal, and heptanal,
among among others [9,13].
others [9,13].

Figure1.1.The
Figure Theformation
formationof
ofsome
somevolatiles
volatilesfrom
fromthe
theautoxidation
autoxidationof
oflinoleic
linoleicacid.
acid.

3.2.
3.2. Formation
Formation of of Oxidation
OxidationProducts
ProductsviaviaPhotooxidation
Photooxidation
A possible route for lipid-derived
A possible route for lipid-derived flavor flavor generation
generation is lipolysis,
is lipolysis, which
which raises
raises the
the level
level of free fatty acids (FFAs). The FFAs may then be oxidized via various
of free fatty acids (FFAs). The FFAs may then be oxidized via various oxidation mecha- oxidation
mechanisms and produces hydroperoxides, and further decompose to a series of volatile
nisms and produces hydroperoxides, and further decompose to a series of volatile flavor
flavor compounds [18]. Photooxidation is much faster than autoxidation, occurring in
compounds [18]. Photooxidation1 is much faster than autoxidation, occurring in the pres-
the presence of singlet oxygen ( O2 ), which can be generated by ultraviolet (UV) light,
ence of singlet oxygen (1O2), which can be generated by ultraviolet (UV) light, and/or pho-
and/or photosensitizers including riboflavin, chlorophyll, and myoglobin. Singlet oxygen
tosensitizers including riboflavin, chlorophyll, and myoglobin. Singlet oxygen is more re-
is more reactive than triplet oxygen due to its higher electrophilicity. For instance, the
active than triplet oxygen due to its higher electrophilicity. For instance, the interaction
interaction1 between 1 O2 and linoleic acid is about 1500 times faster than that with 3 O [19].
between O2 and linoleic acid is about 1500 times faster than that with 3O2 [19]. The2 pho-
The photooxidation also produces hydroperoxides in many ways, where excited triplet
tooxidation also produces hydroperoxides in many ways, where excited triplet sensitizers
sensitizers (e.g., myoglobin and hemoglobin) react with molecular oxygen (3 O2 ) and pro-
(e.g., 1myoglobin and hemoglobin) react with molecular oxygen (3O2) and produce 1O2.
1 O and
duce O2 . Subsequently, hydroperoxides are formed upon interactions between 2
Subsequently, hydroperoxides are formed upon interactions between 1O2 and double
double bonds of unsaturated fatty acids without forming alkyl radical [20]. Moreover, ROS
bonds of unsaturated fatty acids without forming alkyl radical [20]. Moreover, ROS such
such as superoxide radical anion can be formed when the sensitizer interacts with 3 O2 ,
as superoxide
resulting radical
in lipid anion due
oxidation can be
to formed when theofsensitizer
the abstraction hydrogen interacts with 3unsaturated
atoms from O2, resulting
in lipid oxidation due to the abstraction of hydrogen
1 atoms from unsaturated
fatty acids. In addition, hydroxyl radicals and O2 produced from the interaction fatty acids.
between
superoxide radical anions and H2 O2 can initiate lipid oxidation in the presence of metal
ions. Furthermore, alkyl radicals produced from the reaction of fatty acids and sensitizers
can react with molecular oxygen and produce peroxyl radicals, initiating oxidation via
Molecules 2022, 27, 5014 5 of 16

a free radical chain mechanism [15]. Finally, primary oxidation products such as vari-
ous hydroperoxides including 9- and 10-hydroperoxides for oleic acid, 9-, 10-, 12-, 13-,
15-, and 16-hydroperoxides for linolenic acid, and 9-, 10-, 12-, and 13-hydroperoxides for
linoleic acid are formed, which lead to a series of volatile compounds [6]. The formation
of these hydroperoxides mainly depends on the nature of the fatty acids involved in the
oxidation reaction.

3.3. Formation of Oxidation Products via Enzymatic Oxidation

Lipoxygenase, the main enzyme involved in enzymatic oxidation, is found abundantly
in various species of plants, animals, and fish. It is a globular protein comprising a single
polypeptide chain with a molecular mass of 75–80 kDa in animals and 94–104 kDa in
plants. Lipoxygenase plays an important role in the degradation of unsaturated fatty
acids, resulting in flavor generation. Lipoxygenase-mediated reactions produce the aromas
of freshly harvested fish through the production of both volatile alcohols and carbonyl
compounds [21]. For instance, lipoxygenase in fish can react with fatty acids and produces
cis-4-heptenal and 2,4,7-decatrienal isomers, which are responsible for a fishy aroma [16].
The rate of enzyme-catalyzed lipid oxidation mainly depends on the concentration of
enzymes, which is proportional to the oxidation [15]. Lipoxygenase produces conjugated
hydroperoxides through oxidation when the active site of the enzyme abstracts a hydrogen
atom from the methylene group of fatty acids. In order to exhibit activity, the active site of
the enzyme containing iron needs to be in the ferrous form [22]. In short, lipase catalyzes the
first step of the lipid by hydrolysis and produces free fatty acids, followed by the production
of conjugated hydroperoxy fatty acids through lipoxygenase and finally, the formation of
volatiles such as carbonyl compounds. In plants, 9- and 13-hydroperoxy-octadecadienoates
are transformed into reactive intermediates by hydroperoxide dehydrases, hydroperoxide
lyases, epoxide hydrolases, and hydroperoxide epoxygenase [21,23]. For example, the
oxidation of linolenic acid by a lipoxygenase and a subsequent lyase cleavage reaction
produces trans-2, cis-6-nonadienal in cucumbers, and trans-2-hexenal in fresh tomatoes.
Moreover, these carbonyl compounds may further degrade into alcohol (e.g., trans-2, cis-6-
nonadienol and trans-2-hexenol), which provides a stronger aroma than the precursor of
carbonyls [16] (Figure 2). Apart from lipoxygenase, alcohol dehydrogenase, hydroperoxide
lyase, and 3Z, 2E (3cis, 2trans) enal isomerase are also involved in the biosynthetic pathway.
For instance, hydroperoxy fatty acids further break down into C6 or C9 volatiles such as
3-hexenal and 3,6-nonadienal, respectively, in the presence of hydroperoxide lyase [6].
In addition to the enzymatic and non-enzymatic pathways, there is a heat-mediated
approach that also initiates lipid oxidation. The thermal oxidation of frying oil is the effect
of the development of a complex form of the decomposition of fatty acids. The forma-
tion of cyclic and dimeric compounds leads to the thermolytic reaction in unsaturated
fatty acids. Even saturated fatty acids become oxidized when heated at a high tempera-
ture (~150 ◦ C) and produce n-alkanols, n-alkanes, lactones, 2-alkanones, 1-alkenes, and
carboxylic acid [24].
Several analytical techniques have been used to identify and quantify lipid oxidation
products. Generally, techniques that are used to measure the changes in primary oxida-
tion products are changes in the fatty acid contents, peroxide value (PV) (e.g., iodometric
titration and ferric-xylenol orange), and conjugated dienes/trienes, while the thiobarbi-
turic acid reactive substances (TBARS) assay, p-anisidine value, TOTOX value (2 PV +
p-anisidine), and volatile measurements using GC-MS are used to determine secondary
oxidation products. Both primary and secondary oxidation products are recommended to
be measured for better reliability regarding the state of oxidation.
Molecules2022,
Molecules 2022,27,
27,5014
x FOR PEER REVIEW 66 of 16
17

Figure 2. The
Figure2. The formation
formation of
of carbonyls
carbonyls and
and alcohols
alcoholsfrom
fromlinolenic
linolenicacid.
acid.
4. Flavor Chemistry: Lipid Participation in Maillard Interaction
In addition to the enzymatic and non-enzymatic pathways, there is a heat-mediated
The Maillard
approach that alsoreaction
initiates involves a seriesThe
lipid oxidation. of complex chemicalof
thermal oxidation reactions between
frying oil the
is the effect
carbonyls of reducing sugars and proteins, mainly primary or secondary amines,
of the development of a complex form of the decomposition of fatty acids. The formation which is
responsible for the browning of food and its distinctive flavor [25]. The Maillard
of cyclic and dimeric compounds leads to the thermolytic reaction in unsaturated fatty reaction
develops a wide range of chemical constituents belonging to oxazole, thiophene, furan,
acids. Even saturated fatty acids become oxidized when heated at a high temperature
thiazole, pyrrole, pyrazine, and pyridine, among others. Generally, compounds such as
(~150 °C) and produce n-alkanols, n-alkanes, lactones, 2-alkanones, 1-alkenes, and carbox-
peptides, amino acids, thiamine, carbohydrates, nucleotides, and lipids in food promote the
ylic acid [24].
development of flavor. Hence, the generation of aroma volatiles is basically related to the
Several analytical techniques have been used to identify and quantify lipid oxidation
break down and oxidation of lipids, the Maillard reaction, and the degradation of vitamins,
products. Generally, techniques that are used to measure the changes in primary oxida-
mainly thiamine [11]. For example, the Maillard reaction between ribose and cysteine pro-
tion products are changes in the fatty acid contents, peroxide value (PV) (e.g., iodometric
duces sulfur-containing compounds including thiophan-3-thiol and furan-3-thiol, which
titration and ferric-xylenol orange), and conjugated dienes/trienes, while the thiobarbitu-
are responsible for the desirable meaty aroma of cooked meat [9]. In particular, the products
ric acid reactive substances (TBARS) assay, p-anisidine value, TOTOX value (2 PV + p-
of these reactions can act as precursors and interact with other degradation constituents of
anisidine), and volatile measurements using GC-MS are used to determine secondary ox-
food and produce a number of long-chain heterocyclic compounds during cooking. For
idation products. Both primary and secondary oxidation products are recommended to
instance, Henderson and Nawar [26] reported the formation of 2-pentylpyridine through
be measured
the interactionfor better(linoleic
of lipid reliability regarding
acid) the state
and Maillard of oxidation.
reaction by-products (Figure 3). Amadori
products are produced during the first stages of the reaction via glycosylamine because of
4. Flavor Chemistry: Lipid Participation in Maillard Interaction
the interaction between the carbonyls of reducing sugars and primary amines of amino
acids,The Maillard
peptides, reaction
or other involves a The
components. series of complex
formation chemicalaldehydes
of Strecker reactions through
between the
the
carbonyls of of
degradation reducing sugars
amino acids andand proteins,
lipid mainly
oxidation primary
is another or secondary
example amines,aroma
of developing which
is responsible
compounds for These
[27]. the browning
compounds of food and
react its Maillard
with distinctive flavor [25]. Thecarbonyls
reaction-derived Maillard reac-
and
tion develops
form a wide
intermediates, rangefurther
which of chemical
break constituents belonging
down into flavor to oxazole,
compounds. thiophene,
Apart from aminofu-
ran, thiazole, pyrrole, pyrazine, and pyridine, among others. Generally, compounds
acids, phenolic compounds may promote the generation of Strecker aldehydes. For exam- such
as peptides,
ple, amino acids amino acids, thiamine,
and quinones carbohydrates,
originating from phenolics nucleotides,
could produceandflavor
lipidsprecursor
in food
 by-products (Figure 3). Amadori products are produced during the first stages of the re-
action via glycosylamine because of the interaction between the carbonyls of reducing
sugars and primary amines of amino acids, peptides, or other components. The formation
of Strecker aldehydes through the degradation of amino acids and lipid oxidation is an-
other example of developing aroma compounds [27]. These compounds react with Mail-
Molecules 2022, 27, 5014 7 of 16
lard reaction-derived carbonyls and form intermediates, which further break down into
flavor compounds. Apart from amino acids, phenolic compounds may promote the gen-
eration of Strecker aldehydes. For example, amino acids and quinones originating from
volatile aldehydes
phenolics through
could produce Strecker
flavor degradation
precursor volatile[28]. The same
aldehydes studyStrecker
through also reported that
degrada-
amino acids
tion [28]. (e.g.,
The samephenylalanine and methionine)
study also reported that amino were capable
acids of interacting with
(e.g., phenylalanine phenolic
and methio-
compounds (e.g., chlorogenic
nine) were capable acid,
of interacting caffeic
with acid,compounds
phenolic epicatechin, andchlorogenic
(e.g., catechin) and
acid,develop-
caffeic
ing Strecker aldehydes (e.g., phenylacetaldehyde and methional) in a ferricyanide-based
acid, epicatechin, and catechin) and developing Strecker aldehydes (e.g., phenylacetalde-
model system.
hyde and methional) in a ferricyanide-based model system.

Figure3.3. The
Figure The formation
formation of
of 2-pentylpyridine
2-pentylpyridine by
by lipid–Maillard
lipid–Maillard interaction.
interaction.

Numerous sugar dehydration and degradation compounds such as dicarbonyl com-

pounds, furanone and furfural derivatives, and hydroxy ketones are developed by the
dehydration and rearrangement of the subsequent compounds [6,29]. However, the forma-
tion of Maillard reaction products depends on the moisture content, cooking temperature
and time, pH, and the nature of the reactants involved. The reaction rate increases markedly
with the temperature at low moisture levels, and the production of flavor compounds is
linked with areas of the food dehydrated by heat. Moreover, the formation of Maillard
reaction products is affected by the presence of a discrete non-polar environment and the
reactants’ location. For example, the interaction between lipid oxidation and Maillard
reaction products was examined in a model system of oil-in-water emulsion containing
canola oil, water, glucose, phenylalanine, and a surfactant (Tween 20) [30]. It was found
that the development of Maillard reaction products was 16 times greater in the emulsion
system than in the aqueous solution.
Maillard reaction produces a myriad of components that are involved in flavor genera-
tion alone or with the lipid oxidation products. The interaction of these products produces
new volatile compounds in some cases, or partially/wholly blocks other compounds [10].
For instance, aldehydes generated from lipid oxidation could participate in the initial and
later stages of the Maillard reaction during cooking to form volatiles including pyrazines,
Molecules 2022, 27, 5014 8 of 16

thiophenes, pyridines, oxazoles, and thiazoles with alkyl side chains [11,31]. Several thia-
zoles and thiophenes such as 2-alkyl-3-thiazolines, 2-alkylthiazoles, and 2-alkylthiophenes
have been reported in roasted meat. Particularly, carbonyls and sulfur compounds derived
from ribose and cysteine are the major precursors to the flavor of meat. It has been reported
that phospholipid oxidation produced 2,4-decadienal, which participated in the Maillard
reaction and developed 2-alkyl heterocyclic products [9]. Usually, volatiles obtained from
the combination lipid–Maillard reaction exhibit a higher odor threshold than those devel-
oped from its parent reaction, resulting in a weak odor intensity from the lipid–Maillard
by-products. However, it may modify the aroma profiles obtained from this complex
system and provide an indirect impact on the aroma compounds [10,11].

5. Effect of Processing on the Flavor Compounds of Meat and Eggs

Effect of various processing methods on major flavor components of different types of
food is summarized in Table 1.

Table 1. Important flavor components of selected food.

Processing Method Type of Food Volatile Compounds References

Butanal, pentanal, hexanal, octanal, nonanal, hexadecanal,
Roasting Chicken octadecanal, 2-metbylpyrazine, 2,3-dimetbylpyrazine, pyridine, [32]
2-methylpyridine, N-metbylpyrrole, and 2-methylthiazole
Hexanal, heptanal, octanal, nonanal, hexadecanal, trans-2-pentenal,
Cooking Chicken trans-2-heptenal, trans-2-octenal, trans-2-decenal, trans-2-undecenaI, [32]
trans,cis-2,4-decadienal, and trans,trans-2,4-decadienal
1-Octene-3-ol, hexanal, 2-ethyl hexyl acetate, linalool, eugenol,
Frozen storage Chicken [33]
diallyl disulfide, anisole, and α-pinene
Pentanal, hexanal, heptanal, nonanal, 12-methyltridecanal,
Cooking Beef nona-2-trans-enal, decan-2-one, 1-cctene-3-ol, pyrazines, [1,6]
2-methyl-3-furan, 2-pentyl furan
Octanal, phenylacetaldehyde, 2-ethyl-1-hexanol, hexanal,
Frozen storage Beef [34]
1-heptanol, and isoeugenol
1-Octene-3-ol, hexanal, 2-ethyl hexyl acetate, linalool, eugenol,
Frozen storage Meatballs (beef) [35]
diallyl disulfide, anisole, and α-pinene
Hexanal, heptanal, decanal, nonanal, trans-cinnamaldehyde,
Fermentation Sausages (pork) 2-heptanone, 3-hydroxy-2-butanone, linalool, terpinen-4-ol, and [36]
ethyl acetate
Modified-atmosphere 1,8-Cineole, linalool, L–carvone, cinnamaldehyde menthol, and
Cooked ham [37]
packaging cinnamaldehyde
Benzaldehyde, hexanal, 2-heptanone, limonene, hexanol, octanol,
Curing Ham [38,39]
pentanol, 3-methylbutanal, 2-nonanone, butanol, and propanone
3-Hydroxy-cyclohexanone, hexanal, D-limonene, 2-pentyl-furan,
Boiling Egg yolk [40]
phenylacetaldehyde, and 2-ethyl-1-hexanol
Tuna, conger, sardine, 2-Methyl-3-furanthiol, l-penten-3-ol, 2-phenylethanol, and
Cooking/canning [41]
and pale chub dimethyl sulfide
3-Methylbutanal, octanal, 2-nonenal, n-hexanal, nonanal,
Cold plasma treatment Cured black carp [18]
2,4-decadienal, 2,4-nonadienal,1-octene-3-ol, ketone 1-octene-3-one
Soybean and 2-Heptenal, ethyl butyrate, 2,4-pentanedione, acetyl pyrazine,
Frying [42]
canola oils 1-octanoland, 3-methylbutanal, pyridine, and linalool
Butanal, pentanal, hexanal, heptanal, octanal, nonanal, decanal,
undecanal, dodecanal, trans-2-butenal, trans-2-pentenal,
Heating Soybean oil trans-2-hexenal, cis-4-heptenal, cis-2-heptenal, trans-2-octenal, [43]
cis-2-nonenal, cis-2-decenal, cis-2-decenal, 2-undecenal,
trans-2-hepten-1-o, 2-butanone, and 2-pentylfuran
trans-2, cis-6-Nonadienal, trans-2-nonenal, nonanal, n-hexanal,
Bagging Cucumber [44]
trans-2-hexenal, propanal, and cis-2-heptenal
1-Octen-3-one, 3-octanone, 1-octen-3-ol, 3-octanol, 2-octen-1-ol,
Drying Mushroom [45]
1-octanol, benzaldehyde, benzeneacetaldehyde, and decanal
2-Methylbutanal, 3-methylbutanal, n-hexanal,
Drying Tomato 6-methyl-5-hepten-2-ol, 3-methyl-1-butanol, and [46]
6-methyl-5-hepten-2-one

Generally, salty, metallic, and bloody, with a bit of sweet aroma, are the common flavor
of raw meats. Upon processing, mainly heating, meat and meat products produce a series
of reactions including lipid oxidation, which are responsible for the development of specific
meat flavors. During the thermal processing of meat, volatile compounds such as alcohols,
Molecules 2022, 27, 5014 9 of 16

ketones, aldehydes, furans, esters, hydrocarbons, carboxylic acids, pyrans, pyrazines,

lactones, phenols, pyrroles, pyridines, thiazoles, thiazolines, oxazoles, thiophenes, and
other nitrogen- or sulfur-containing compounds are formed [47]. Peptides, amino acids,
nucleotides, organic acids, and other flavor enhancers are the most taste-active constituents
of meats. Hundreds of volatile compounds have been characterized from various muscle
foods, but no single component has so far been documented as being solely responsible for
the aroma development, though hexanal has been found to be the predominant volatile in
cooked meat. For example, Mottram [48] reported around 900 volatiles from cooked beef
in which only a small number of this wide range of components contributed to the flavor
profile. It has been reported that around 3% out of 10,000 identified volatiles contributed
to the flavor development in foods [49]. During heat processing, lipids in the Maillard
reaction produce species-specific flavors in meats. Generally, carbohydrates and free amino
acids of different meats are similar, and thus, a similar flavor profile is expected upon
cooking. Nevertheless, lipids in meat from different species are not always the same,
especially intramuscular lipids, which modify the flavor profile upon heating. Particularly,
the development of meaty aromas is mainly responsible for phospholipids present in the
intramuscular lipids. Moreover, the impact of phospholipids is much higher than that
of triacylglycerols (TAG) in terms of flavor generation due to the higher proportion of
unsaturated fatty acids (e.g., arachidonic acid) in phospholipids [50,51]. Due to the presence
of medium-chain branched fatty acids, the distinct strong flavor of lamb or mutton meat is
observed. The adipose tissue of lamb meat contains 4-ethyl- and 4-methyloctanoic acids,
which are responsible for the distinct flavor [6]. In contrast, chicken has a higher level of
unsaturated fatty acids than red meat, resulting in the generation of more volatile aldehydes
that are related to the distinct aroma of chicken. In particular, lipid-derived carbonyls,
mainly aliphatic aldehydes, are responsible for the fatty flavors of roasted chicken meat.
For example, 193 flavor compounds were identified from the roasted chicken; among
them, 41 were lipid-derived aldehyde compounds [32]. They also reported that primary
oxidation products of linoleic acid such as hexanal and 2,4-decadienal were the major flavor
compounds in chicken meat. Apart from this, heat treatment also helps in lipid migration,
which has both positive and negative effects on food quality. Xiang et al. [40] reported that
lipid migration caused by heating promotes more lipid-specific flavor volatiles including
3-hydroxy-cyclohexanone, hexanal, D-limonene, 2-pentyl-furan, phenylacetaldehyde, and
2-ethyl-1-hexanol in the upper part of egg yolk.
Frozen storage also affects the flavor and sensory characteristics of meat due to the
lipid oxidation catalyzed by iron and myoglobin. For instance, the effect of frozen storage
on the flavor profile of marinated raw beef was investigated, and it was found that the con-
centration of flavor compounds including octanal, phenylacetaldehyde, 2-ethyl-1-hexanol,
hexanal, 1-heptanol, and isoeugenol fluctuated along with the frozen storage [34]. Most
of these compounds belong to alcohols and aldehydes, indicating that they were derived
from lipid oxidation. This is because unsaturated fatty acids (e.g., linolenic, linoleic, and
oleic acids) can undergo autoxidation or enzymatic oxidation and produce various hy-
droperoxides (e.g., 8-, 9-, 10-, 11- or 13-ROOH) and then volatiles (e.g., aldehydes) after
homolysis. Similarly, the flavor of raw chicken meat was evaluated during frozen storage
(0–8 weeks) [33]. It was found that the short-term frozen storage of raw meat improved its
flavor profile. This could be due to the lipolysis, proteolysis, and oxidation degradation
of flavor compounds in the long-term. Moreover, flavor components in cooked beef meat-
balls during frozen storage (0 to 90 days) were measured using various techniques, and
80 volatiles were identified in which 32, mainly 1-octene-3-ol, hexanal, 2-ethyl hexyl acetate,
linalool, eugenol, diallyl disulfide, anisole, and α-pinene, were aroma-active [35]. Their
sensory evaluation results demonstrated that the overall aroma profile started to decrease
with the frozen storage.
Apart from this, the fermentation of meat improves its flavor and inhibits lipid oxi-
dation. Chen et al. [36] suggested that fermentation of Harbin dry sausages lowered the
number of volatiles (e.g., alcohols, aldehydes, hydrocarbons, and acids) generated from
Molecules 2022, 27, 5014 10 of 16

lipid autoxidation. Moreover, dry-aging is a common method for the development of

distinct flavors due to microbial activity, dehydration, and lipid oxidation during storage.
Particularly, dry-aging involves lipid oxidation and produces intermediates, which may
interact with other flavor precursors such as peptides, amino acids, and sugars during heat-
ing and generate a dry-aged flavor [52]. On the other hand, the volatile profile of wet-cured
cooked ham was compared with non-cured meat [37]. Carbonyls found in the cured meat
were lower than the non-cured meat, and the major volatiles were terpenes (1,8-cineole,
linalool, L–carvone, cinnamaldehyde menthol, and cinnamaldehyde), which mainly devel-
oped from seasonings, and sulfur components and 3-methylbutanoic acid originating from
Strecker degradation. In contrast, aldehydes, ketones, and alcohols play a significant role
in the flavor characteristic of dry-cured ham. Among the various volatiles, benzaldehyde,
hexanal, 2-heptanone, limonene, hexanol, octanol, pentanol, 3-methylbutanal, 2-nonanone,
butanol, and propanone were abundant in the dry-cured products [38,39].

Factors Influencing the Flavor Formation of Meat

Several factors including the type of meat, the parameters of the production process,
and components of meat are responsible for its flavor development. For example, the
common volatile compounds of cooked beef are octanal, 2,4-decadienal, nonanal, me-
thional, 2-furfurylthiol, 2-metyl-3-furanthiol, methanethiol, 3-mercapto-2-pentanone, and
4-hydroxy-2,5-dimethyl-3-(2H)-furanone. Interestingly, an almost similar flavor profile
also occurs in cooked chicken and pork, but their concentrations differ among species. A
high concentration of 2-methyl-3-furanthiol, 4-hydroxy-2,5-dimethyl-3-(2H)-furanone, and
2-furfurylthiol is associated with the meaty-caramel odor in cooked beef. In contrast, a
lower concentration of 4-hydroxy-2,5-dimethyl-3-(2H)-furanone is enough to provide the
odor of pork meat, and similarly, a lower level of carbonyls (e.g., hexanal and nonanal) is
responsible for the greasy odor in pork than in beef meat [6,53].
Breed, sex, feed, nutrition, and age notably influence the quality and quantity of fats,
which affect the overall flavor profile of meats. Fats and fatty acid composition are the
principal contributors to the flavor generation in meat. The composition of fatty acids is
mainly dependent on diet, and the flavor is produced upon melting fats. For instance, grass-
fed beef is less susceptible to lipid oxidation than grain-fed beef, and this is because of the
reduced content of flavonoids, vitamins A, C, and E, and carotenoids present in feeds [54].
Due to the differences in the digestive system, fatty acids, mainly polyunsaturated fatty
acids, deposition is higher in beef or lamb. Thus, the meaty flavor is generated by the
breakdown of these fatty acids upon degradation to aldehydes, ketones, and alcohols,
among others. Apart from the type and content of fatty acids, protein content, temperature,
time, water activity, reaction media, pH, aging, marbling, and cooking technique affect
the flavor of meat [27]. Cooking methods modify the chemical composition of meat and
promote lipid oxidation. Compared to other cooking methods, roasting results in increased
oxidation and volatiles due to the longer time and high temperature experienced during
cooking. Hence, the generation of flavor is increased through the Maillard reaction and lipid
oxidation [55]. Similarly, grilling increases the content of pyrazines, a series of nitrogen-
containing compounds responsible for the nutty and roasty flavor, which contribute up
to 80% to the volatile compounds in grilled meat. Moreover, microwave cooking at a
lower temperature and shorter time also causes increased lipid oxidation, which could be
due to the interaction between microwave and meat fat. Frying can change the fatty acid
composition due to added oil for frying, thus increasing the lipid oxidation [55–57].

6. Effect of Processing on the Flavor Compounds of Fish

Both enzymatic and non-enzymatic oxidative reactions are involved in the develop-
ment of fish aromas and flavors. The distinctive plant-green aroma of fresh fish is mainly
produced from lipoxygenase-derived carbonyls and alcohols, which depends on the type
and content of compounds present in different species. Lipoxygenase available in fish can
generate alcohols and then further break down into carbonyls such as 1,5-octadien-3-one
Molecules 2022, 27, 5014 11 of 16

(Figure 4). Moreover, secondary oxidation products play the main role in the flavor de-
velopment in certain species of fish [58]. The fresh fish aromas are linked mainly with
alcohols (l-penten-3-ol, cis-3-hexen-l-ol, l-octen-3-ol, trans-2- octen-1-ol, l,5-octadien-3-ol,
2,5-octadien-l-ol, trans-2-nonen-l-ol, cis-6-nonen-l-ol, cis-3-nonen-l-ol, and 3,6-nonadien-l-ol)
and carbonyls (trans-2-penten-l-al, hexan-l-al, trans-2-hexen-l-al, trans-octen-l-al, trans-2-
nonen-l-al, l-octen-3-one, 2,3-octadien-l-one, and 1,5-octadien-3-one) [41]. Both desirable
and non-desirable flavors are the result of lipid oxidation in marine-based food. In par-
ticular, secondary oxidation products including 2,4,7-decatrienals and other carbonyls of
omega-3
Molecules 2022, 27, x FORfatty acids are responsible for the fishy off-flavor [9]. Apart from volatiles in 12 of 17
PEER REVIEW
seafood, non-volatiles including nucleotides, free amino acids, peptides, minerals, and
sugars are also responsible for the development of taste and flavor [59]. Upon processing,
mainly cooking, the flavornon-volatiles
seafood, of fish changes dramatically,
including nucleotides, where lipid oxidation,
free amino Maillard
acids, peptides, minerals, and
reaction, and Strecker sugars degradation play afor
are also responsible leading role in generating
the development of taste andmeaty
flavor aromas.
[59]. UponFor processing,
example, during tuna canning, 2-methyl-3-furanthiol is produced upon the thermally me-Maillard
mainly cooking, the flavor of fish changes dramatically, where lipid oxidation,
diated reaction of reaction,
cysteineand andStrecker
ribose,degradation
which provides play a the
leading
meatyrole flavor.
in generating
Some meaty aromas. For
fish (e.g.,
example, during tuna canning, 2-methyl-3-furanthiol
conger, sardine, and pale chub) produce a distinct odor upon cooking and these are relatedis produced upon the thermally me-
diated reaction of cysteine and ribose, which provides the meaty flavor. Some fish (e.g.,
to l-penten-3-ol, 2-phenylethanol, and dimethyl sulfide [41]. Moreover, carbonyls are the
conger, sardine, and pale chub) produce a distinct odor upon cooking and these are re-
most important compounds in salted and dried fish. However, [60] reported that drying
lated to l-penten-3-ol, 2-phenylethanol, and dimethyl sulfide [41]. Moreover, carbonyls
such as hot air, microwave,
are the mostand microwave-vacuum
important compounds in salted drying andremoved
dried fish.aHowever,
part of the[60]fishy
reported that
off-odor (e.g., 2-methylisoborneol and cyclic alcohols) and increased
drying such as hot air, microwave, and microwave-vacuum drying removed the grilled flavor
a part of the
of silver carp slices. fishyHowever, the2-methylisoborneol
off-odor (e.g., level of volatilesand generated through
cyclic alcohols) andlipid oxidation
increased the grilled fla-
is lower in hot air-dried samples
vor of silver than However,
carp slices. those prepared
the levelby sun-drying
of volatiles [61]. through
generated On the lipid
otheroxidation
hand, Ke et al. [18]issuggested
lower in hotthat air-dried samples than those
cold plasma-treated airprepared
enhanced bythe
sun-drying [61]. Onofthe other
development
hand, Ke in
volatile flavor compounds et al. [18] suggested
dry-cured that cold plasma-treated
fish originating from the oxidationair enhanced the development of
of unsaturated
volatile flavor compounds in dry-cured fish originating
fatty acids. The major volatiles were aldehydes (3-methylbutanal, octanal, 2-nonenal, from the oxidation of unsaturated
n-
fatty acids. The major volatiles were aldehydes (3-methylbutanal, octanal, 2-nonenal, n-
hexanal, nonanal, 2,4-decadienal, and 2,4-nonadienal), alcohols (1-octene-3-ol), and ketones
hexanal, nonanal, 2,4-decadienal, and 2,4-nonadienal), alcohols (1-octene-3-ol), and ke-
(1-octene-3- one). Among them, n-hexanal derived from the degradation of linoleic acid is
tones (1-octene-3- one). Among them, n-hexanal derived from the degradation of linoleic
responsible for theacid tallowy and green
is responsible for leafy notesand
the tallowy to the fish
green flavor.
leafy notes to the fish flavor.

Figure
Figure 4. The enzymatic 4. The enzymatic
breakdown breakdown of eicosapentaenoic
of eicosapentaenoic acid (EPA)
acid (EPA) (adapted (adapted
from from
Shahidi Shahidi [16]).
[16]).

7. Effect of Processing on the Flavor Compounds of Edible Oils

Edible oils are mainly composed of TAG (~98%) and other minor components such
as glycolipids, phospholipids, waxes, tocopherols, sterols, chlorophylls, and other phe-
nolic compounds. These minor components are collectively referred to as unsaponifiable
matters. During processing, many of these compounds are removed. In particular, chlo-
Molecules 2022, 27, 5014 12 of 16

7. Effect of Processing on the Flavor Compounds of Edible Oils

Edible oils are mainly composed of TAG (~98%) and other minor components such
as glycolipids, phospholipids, waxes, tocopherols, sterols, chlorophylls, and other phe-
nolic compounds. These minor components are collectively referred to as unsaponifiable
matters. During processing, many of these compounds are removed. In particular, chloro-
phylls in oils are responsible for the generation of odor-active aldehydes under fluorescent
light [62,63]. However, the most common flavor components of fats and oils are gener-
ated from unsaturated fatty acids in TAG or polar lipids upon reacting with oxygen. For
example, the major aldehydes produced upon oxidation of linolenic acid are propanal,
2-pentenal, 2-butenal, 2-hexenal, 3-hexenal, 2,4-heptadienal, 2-heptenal, 2,5 octadienal,
2,6-nonadienal, and 2,4,7-decatrienal, whereas pentanal, heptanal, hexanal, 2-octenal, oc-
tanal, 2-heptenal, 2-nonenal, 3-nonenal, 2,4-decadienal, 2-decenal, and 2,4-nonadienal are
formed from the oxidation of linoleic acid and heptanal, octanal, decanal, 2-undecenal,
decanal, and 2-decenal are developed from oleic acid [53]. Generally, the aromas of alde-
hydes are painty, metallic, green, beany, and rancid, and they are frequently related to the
off-flavor. For instance, the main flavor compound responsible for deep-fat fried foods
is 2,4-decadienal. This compound can further break down to trans-epoxy-trans-decenal,
which is considered as one of the most effective odorants of soybean oil [64]. However,
2-pentylfuran and 3-cis-hexenal are mainly responsible for the beany or grassy flavor of
soybean oil [65]. The cooking, mainly frying, of oils plays an important role in flavor
development. For instance, Lee et al. [42] found that the frying of soybean and canola
oils increased volatile concentration, and 2-heptenal, ethyl butyrate, and 2,4-pentanedione
were the major volatiles in soybean oil, whereas ethyl butyrate and linalool were abundant
in canola oil. Moreover, Xiao et al. [43] stated that the content of volatile aldehydes and
alcohols was developed at 120 ◦ C, whereas ketones and furans were produced at 150 ◦ C,
and acids were formed at 180 ◦ C during the heating of soybean oil. The principal volatiles
in most virgin or extra virgin olive oils (EVOO) are C5 and C6 aliphatic components such as
1-hexanol, hexanal, trans-2-hexenal, trans-2-hexen-1-ol, cis-3-hexen-1-ol, cis-2-penten-1-ol,
and 3-methylbutanol [6]. However, the concentration of volatiles is species-specific and
depends mainly on the fatty acid composition. Tian et al. [66] reported that the major
volatile components of rapeseed, soybean, peanut, and sunflower oils were nearly the
same including hexanal, trans-2-heptenal, nonanal, 2,4-decadienal, trans-2,4-nonadienal,
cis-2-heptenal, 1-octene-3-ol, and 1-pentanol, but their concentration varied significantly.

8. Effect of Processing on the Flavor Compounds of Fruits and Vegetables

The major flavor precursors of fruits and vegetables are related to their fatty acids,
which are mainly dependent on the degree of maturity, cultivar, geographic location,
and processing methods. β-oxidation is the primary metabolic pathway for generating
aroma compounds, whereas lipoxygenase also plays an important role in developing
flavor compounds from fatty acids. For example, the biosynthesis of lactones is related to
the flavor development of pineapple (δ-octalactone), coconut (γ-octalactone), peach and
nectarine (γ-decalactone and γ-dodecalactone, respectively) via the β-oxidation pathway.
In contrast, the degradation of linoleic and linolenic acids to aldehydes, alcohols, acids, and
esters via the lipoxygenase pathway is responsible for the flavor development of fruits and
vegetables [67,68]. For instance, lipoxygenase-derived volatiles including cis-3-hexenol, cis-
3-hexenal, trans-2-hexenol, and trans-2-hexenal, which are related to the fresh green flavor
of tomatoes [69]. However, various processing methods have a significant effect on the
flavor profile of fruits and vegetables. Shan et al. [44] reported that the bagging treatment
enhanced the flavor quality of cucumbers, which were related to trans-2, cis-6-nonadienal,
and trans-2-nonenal. Similarly, Lomelí-Martín [70] suggested that high hydrostatic pressure
increased the content of aldehydes and ketones and decreased that of alcohols in the fruits
and vegetables. This could be due to the enzymatic activities and chemical reactions upon
high-pressure treatment. For example, hexanal is associated with the smell of foliage and
grass, which could increase upon high-pressure treatment due to the oxidation of free fatty
Molecules 2022, 27, 5014 13 of 16

acids [71]. Furthermore, autoxidation and the Maillard reaction are the major chemical
reactions responsible for the flavor development of fruits and vegetables during drying.
Zhang et al. [45] found that drying methods (e.g., hot-air drying, freeze-drying, and natural
drying) changed the contents of volatiles (e.g., 1-octen-3-one, 1-octen-3-ol, and 3-octanone)
in mushroom and resulted in a weaker mushroom flavor. Likewise, the volatiles (e.g.,
2-methylbutanal, 3-methylbutanal, n-hexanal, 6-methyl-5-hepten-2-ol, 3-methyl-1-butanol,
and 6-methyl-5-hepten-2-one) of tomatoes were higher in the oven-dried than the sun-dried
samples, indicating that volatiles were mainly formed by oxidation during the thermal
treatment upon oven drying [46].

9. Volatile Measurements
Various techniques can be used to assess volatiles such as aldehydes, ketones, furans,
and alcohols. Headspace analysis is a very common method of determining volatiles.
Generally, gas chromatography coupled with a flame ionizing detector (FID) or mass
spectrometry (GC-MS) and/or an olfactory port is used to analyze volatiles [10]. Steam
distillation-extraction (SDE), solid-phase microextraction (SPME), and dynamic headspace
are very common methods to extract volatiles. Moreover, it has been reported that the
combination of two or more methods (e.g., gas chromatography-olfactometry, GC-O)
results in the better separation of volatiles. In particular, GC-O techniques including
combined hedonic response measurement, aroma extraction dilution analysis (AEDA),
aroma extract concentration analysis (AECA), surface of nasal impact frequency (SNIF),
and finger span cross modality (FSCM) are useful to determine the intensity of aroma
(odor) and the characteristics of volatiles [9,55]. Among them, AEDA is the most common
technique used to analyze volatiles due to its simplicity. Furthermore, multidimensional
gas chromatography (MDGC) coupled with MS and olfactometry detection has recently
been used to investigate the flavor profile [72]. Apart from these, the electronic nose
(E-nose) has gained in popularity in determining the volatile profiles. For example, Xu
et al. [73] suggested that the E-nose has a huge potential for the rapid inspection of volatiles
(e.g., aldehydes, alcohols, 2-acetyl-1-pyrroline, and heterocycles) in rice aging.

10. Conclusions
The development of volatile components in various foods mainly depends on their
constituent compounds and how they process. The generation of flavor volatiles is primarily
linked to lipid degradation, the Maillard reaction, and interactions between the Maillard
reaction and lipid oxidation compounds, among others. Processing such as slow cooking is
mostly responsible for lipid oxidation, while fast cooking is associated with the generation
of Maillard reaction products. Hence, the development of off-flavor can be controlled, and a
desirable flavor can be produced by understanding the detailed mechanisms and inclusion
of biologically active compounds.

Author Contributions: Conceptualization, F.S.; Resources, F.S.; Data curation, A.H.; Writing—
original draft preparation, A.H.; Writing—review and editing, A.H. and F.S.; Supervision, F.S. All
authors have read and agreed to the published version of the manuscript.
Funding: This research was funded by the Natural Sciences and Engineering Research Council
(NSERC) of Canada, RGPIN-2016-04468.
Informed Consent Statement: Not applicable.
Data Availability Statement: Not applicable.
Conflicts of Interest: The authors declare no conflict of interest.
Molecules 2022, 27, 5014 14 of 16

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