Review

Received: 5 July 2017 Revised: 12 September 2017 Accepted article published: 30 March 2018 Published online in Wiley Online Library:

(wileyonlinelibrary.com) DOI 10.1002/jsfa.9039

The aroma volatile repertoire in strawberry

fruit: a review
Jia-wei Yan,a† Zhao-jun Ban,b† Hong-yan Lu,a Dong Li,a Elena Poverenov,c
Zi-sheng Luoa* and Li Lia*
Abstract
Aroma significantly contributes to flavor, which directly affects the commercial quality of strawberries. The strawberry aroma is
complex as many kinds of volatile compounds are found in strawberries. In this review, we describe the current knowledge of
the constituents and of the biosynthesis of strawberry volatile compounds, and the effect of postharvest treatments on aroma
profiles. The characteristic strawberry volatile compounds consist of furanones, such as 2,5-dimethyl-4-hydroxy-3(2H)-furanone
and 4-methoxy-2,5-dimethyl-3(2H)-furanone; esters, including ethyl butanoate, ethyl hexanoate, methyl butanoate, and methyl
hexanoate; sulfur compounds such as methanethiol, and terpenoids including linalool and nerolidol. As for postharvest
treatment, the present review discusses the overview of aroma volatiles in response to temperature, atmosphere, and
exogenous hormones, as well as other treatments including ozone, edible coating, and ultraviolet radiation. The future
prospects for strawberry volatile biosynthesis and metabolism are also presented.
© 2018 Society of Chemical Industry

Keywords: aroma; volatile; flavor; postharvest; strawberry

INTRODUCTION of strawberries. These studies mainly focused on the research

Strawberry fruit (Fragaria × ananassa), a genus of the Rosaceae
family, is the most commonly consumed berry fruit crop world-
wide and is valued for its unique flavor and nutritional quality.1
The pursuit of high yields, large size, and long shelf life has affected
the flavor of strawberries, causing a decline in sales; this indicates
that customers commonly purchase the fruit because of its special
flavor.2 The contribution of taste and mouthfeel to the fruit’s flavor
has attracted the attention of consumers. Nevertheless, the volatile
compounds are the main constituents responsible for the volatile
quality that is a direct factor attracting customers, and they have
high correlation with overall liking of strawberry, although it looks
as if sweetness is more important (Fig. 1).3
Volatile compounds are significant components of strawberry
flavor, and slight changes may significantly modify the taste,
although such compounds only account for 0.001–0.01% of the
fruit’s weight.4 Strawberry aroma represents a good example of
a complex fruit aroma. To date, more than 360 volatile chemicals
have been observed in fresh strawberries.5 These compounds
include esters, alcohols, ketones, furans, terpenes, aldehydes,
and sulfur compounds.6 Types and concentrations of volatiles
contributing to strawberries vary according to their cultivar and
maturity.7 However, esters still represent the most abundant
volatile compounds in strawberries, with as many as 131 different
types accounting for 25–90% of all strawberry volatiles.8 Esters
are also the major source of fruity and floral odors in strawberries,
and the content of esters may be the basis for classifying different
volatile patterns.9,10 Although the furanones, sulfur compounds,
terpenoids and some other compounds exhibit much lower
quantities compared with esters, these compounds demonstrated
significant effects on strawberry volatile.
Considering the important influence of volatile compounds on
flavor, numerous studies have been conducted on aroma volatiles
advances regarding the profiles and biosynthesis of specific straw-
berry volatiles. The present review will depict the dynamic changes
of volatile compounds in strawberries in response to exogenous
abiotic stimuli.
CHARACTERISTIC VOLATILE COMPOUNDS IN
STRAWBERRY
Not all volatile compounds observed in strawberries actually
affect the strawberry volatiles. The odor activity value (OAV) (ratio
of concentration to its sensory threshold)11 has usually been uti-
lized to distinguish these compounds. Only when the OAV value
is higher than 1 is the compound regarded as possessing the
characteristics of a volatile compound to a significant extent. The
∗ Correspondence to: Z-s Luo or L Li, College of Biosystems Engineering and Food
Science, Zhejiang University, Yuhangtang Road 866, Xihu District Hangzhou,
310058, China. E-mail: luozisheng@zju.edu.cn (Luo); lili1984@zju.edu.cn (Li)
† These authors contributed equally to this work.
a Key Laboratory of Agro-Products Postharvest Handling of Ministry of Agri-
culture and Rural Affairs, Zhejiang Key Laboratory for Agro-Food Process-
ing, College of Biosystems Engineering and Food Science, Zhejiang University,
Hangzhou, China
b Zhejiang Provincial Key Laboratory of Chemical and Biological Processing
Technology of Farm Products, Zhejiang Provincial Collaborative Innovation
Center of Agricultural Biological Resources Biochemical Manufacturing, School
of Biological and chemical Engineering/School of Light Industry, Zhejiang
University of Science and Technology, Hangzhou, China
c Department of Postharvest Science, ARO, the Volcani Center, RishonLeZion,
Israel

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 www.soci.org J-w Yan et al.

4-Methoxy-2,5-dimethyl-3(2H)-furanone represents the charac-

teristic volatile furanone in strawberries; its threshold value is
4 × 10−5 mg kg−1 in water.5 It was first discovered in 1965 and
was described as caramel-like, sweet, floral, and fruity, offering
a strawberry-like odor.14,15 It exists in natural fruits in four forms,
namely, DMHF-glucoside, DMMF (mesifuran), DMHF-malonyl-
glucoside, and free aglycone.16,17 Of these, DMHF and DMMF are
the characteristic volatile compounds in fresh strawberry.
Furanones correspond to a few natural volatile compounds
derived from hexoses and pentoses without the breakdown of
the carbon skeleton.18 In the 1990s, as a result of investigation
into the four DMHF forms, glucoside was considered as the
probable precursor of free aglycone, whereas DMHF-glucoside
was deemed to be the stable form of DMHF.19 At that time, new
in vitro methods were applied to study biosynthetic pathways,
aiding speculation about the precursors and derivatives of DMHF.
It was previously discovered that D-fructose derivatives, such
as D-fructose-6-phosphate, D-fructose-1,6-bisphosphate, and
6-deoxy-D-fructose, were the possible precursors of furanones
Figure 1. The correlation between sensorial qualities and overall liking of
as they increased the DMHF concentrations and its resulting
strawberry (regressed against coefficient of determination).
derivatives.19,20 Strawberry callus culture was used in many stud-
ies as precursor feeding as no endogenous furanone existed
Table 1. The characteristic volatile and corresponding precursor in fresh tissue.6,21 Subsequently, analysis by high-performance
compounds in strawberry fruits liquid chromatography (HPLC) and radio labeling proved that
D-fructose-1,6-bisphosphate was possibly the closest precursor
Volatile group Characteristic compounds Precursors to DMHF, and D-fructose could be transformed into DMHF.19 As
Furanones DMHF and its derivative Carbohydrate expected, DMHF was converted into DMMF, DMHF-glucoside,
Esters Methyl butanoate, ethyl Fatty acids or and DMHF-malonyl-glucoside during late stages of strawberry
butanoate, ethyl amino acids ripening and development.22 Based on investigation of furanone
hexanoate, methyl precursors, the putative pathway featured D-glucose or D-fructose
butanoate metabolized to D-fructose-6-phosphate before conversion from
Terpenoids Linalool and nerolidol Carbohydrate the resulting compound into D-fructose-1,6-diphosphate.23,24
Sulfur compounds Methanethiol Unknown It was reported that D-fructose-1,6-diphosphate was then con-
Benzenoids Benzyl alcohol and Amino acids verted into 4-hydroxy-5-methyl-2-methylene-3(2H)-furanone
benzyl acetate
(HMMF), which then converted into DMHF; DMHF was
converted into DMMF or DMHF-glucoside and then trans-
formed into DMHF-malonyl-glucoside in later stages of fruit
higher OAV values of volatile compounds in strawberries indicate development.17,23,25,26 Nevertheless, the biosynthetic pathway of
their more pivotal role in aroma.12 Some compounds (such as DMHF remained unclear, providing limited information regarding
ketones and long-chain acids) feature lower sensory thresholds, transformation of D-fructose-1,6-diphosphate into HMMF.
even at higher concentrations. In contrast, certain low-content As more studies on biomolecular aspects in strawberries were
volatiles contribute significantly to the characteristic volatiles of performed, some of the enzymes involved in the pathways were
strawberry.4 Esters, furanones, terpenes, and sulfur compounds reported. It was stated that quinine oxidoreductase catalyzed the
are among the substances that account for the characteristics transformation of HMMF to DMHF,27 O-methyltransferase partic-
of strawberry volatiles. The esters include ethyl butanoate, ethyl ipating in the DMHF methylation, glucosyl transferase, and mal-
hexanoate, methyl butanoate, and methyl hexanoate; the main onyl transferase converting DMHF into DMHF-glucoside and then
furanones were 2,5-dimethyl-4-hydroxy-3(2H)-furanone (DMHF) into DMHF-malonyl-glucoside,28 respectively. Some genes encod-
and 4-methoxy-2,5-dimethyl-3(2H)-furanone (DMMF), and the ing the enzymes involved were identified, i.e. Fragaria × ananassa
major sulfur compounds and terpenes are methanethiol, linalool, quinone oxidoreductase gene (FaQR)27 and Fragaria × ananassa
and nerolidol.4,13 O-methyltransferase gene (FaOMT).28 A study of FaQR showed that
After exploring aroma compounds in strawberries, considerable FaQR cDNA consisted of 969 bp, and FaQR protein featured 322
attention has recently been given to the metabolism of volatiles, amino acids with a calculated molecular mass of 34.3 kDa. The
especially those involved in biomolecule metabolism and gene monomer expressed in Escherichia coli can catalyze the formation
expression. Many attempts have been made to elucidate the of DMHF.27 Specific transcripts of FaOMT accumulated in ripening
molecular mechanism of volatile profiles. The metabolism of fura- strawberries were only observed in fruits instead of in other tissues,
nones, esters, terpenoids, and other volatiles will now be discussed strawberry flowers, leaves, and roots.29 Substrates of FaOMT pro-
(see Table 1). tein exhibited a common structural feature: a o-diphenolic struc-
ture, as shown in DMHF and its diphenolic tautomer. It was also
Furanones supposed that FaOMT significantly influenced the biosynthesis of
Strawberries contain very low amounts of volatile furanones. vanillin and participated in the lignifications of achenes and vas-
However, compared to their threshold values, the furanones cular bundles in fruits.29 According to recent research on FaOMT, a
have a considerable influence on the profiles of the volatiles. homolog of FaOMT was partly responsible for the natural variation

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The aroma volatile repertoire in strawberry fruit
in DMMF content, and the 30 bp homolog could be a significant
tool in regulating gene expression during strawberry ripening.30
Esters
Esters are the characteristic volatile compounds that define the
strawberry volatiles.8,10,31 Different cultivars contained different
kinds and different quantities of ester compounds. The most
important odorants of F. × ananassa are methyl butanoate, ethyl
butanoate, ethyl hexanoate, and 2-methylbutanoate; whereas the
butyl formate, octyl acetate, decyl acetate, benzyl acetate, carveyl
acetate, decyl butanoate, methyl nicotinate, methyl anthranilate,
and methyl N-formylanthranilate were only found in F. × versa.10
In the biosynthesis of strawberry volatiles, the last important
step is the esterification of alcohols with an acyl moiety of acyl-CoA
by catalysis of different alcohol acyl-transferases (AATs).32 The fatty
acids and amino acids are the precursors of alcohols and acyl-CoAs
in fruits.33,34 On the one hand, the fatty acids, and linoleic and
linolenic acids in general, turn into volatile aldehydes, such as
hexanal and (3Z)-hexenal through the oxidative degradation of
lipoxygenase (LOX) or hydroperoxide lyase (HPL),35–37 which then
convert to the alcohols by alcohol dehydrogenases (ADHs).38 The
𝛽-oxidation of fatty acids results in the formation of C2 units
(acetyl-CoA).39 On the other hand, the catabolism of amino acids
by amino transferases forms 𝛼-keto acids, and then 𝛼-keto acids
become the substrates for 𝛼-keto acid decarboxylase or 𝛼-keto
dehydrogenase, converting into the volatile aldehydes or acetyl
CoA. Afterwards, the aldehydes and acetyl-CoA are converted into
the esters by the actions of ADH and AAT.34,40
Recently, considerable attention has been focused on the
enzymes and genes involved in the fruit ester biosynthesis. Exist-
ing studies indicate that both ADH and AAT play a vital role in ester
biosynthetic pathways, which result from their activity and expres-
sion in ripe fruits. Alcohol dehydrogenases contributed to survival
from hypoxia, the production of cinnamyl alcohols for lignification
of cell walls, and protection from chilling, as well as the biogenesis
of flavor and fragrance volatiles in advanced plant tissues.38 Both
nicotinamide dinucleotide phosphate (NADP)-dependent and
nicotinamide adenine dinucleotide (NAD)-dependent ADHs in
strawberry were reported to possess wide substrate specifici-
ties, including alcohols and aldehydes. Nicotinamide adenine
dinucleotide-dependent ADHs activities are mainly against
short-chained alcohols, whereas NADP-dependent ADHs activ-
ities are mainly against aromatic and terpene alcohols. The
ADH activity increased progressively and differentially between
receptacle and achene tissues during strawberry ripening.38 A
previous study revealed that the molecular weight of ADH was
24.6 kDa and the optimum pH was 6 for ADH activity in straw-
berry callus cultures.41 The appropriate substrates of ADH included
1-propanol (Michaelis constant (Km) = 3.54 mmol L−1 ) and ethanol
(Km = 6.66 mmol L−1 ).42 The characteristics of ADH in strawberry
achenes differed depending on callus culture. The optimum sub-
strates for achene ADH were ethanol (Km = 5.950 mmol L−1 ) and
methanol (Km = 12.610 mmol L−1 ).42 The ADH gene was the first
protein-encoding gene completely sequenced in strawberries.43
This gene and its intron-containing sequence were commonly
applied in ongoing phylogenetic studies on Fragaria species.44,45
Until now, limited information about the relationship between
ADH transcript expression and volatile performance has been
reported in strawberries. The rate-limiting and the last step in
biosynthesis of volatile esters is the reaction catalyzed by AAT;46
which depended on substrate availability and AAT specificity.30,47
Several AAT enzyme isoforms and their corresponding genes
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were widely studied in strawberries with high commercial interest
recently because of the critical role of AAT in ester biosynthesis. It
was reported that the structure of AAT proteins featured several
motifs analogous to BAHD, derived from the abbreviations of
the names (BEAT, AHCT, HCBT, and DAT) of the first four enzymes
identified and characterized in this family of acyltransferases.48
Among these compounds, the HXXXD motif and the DFGWG
sequence were characteristic motifs that were located separately
in the middle of the protein sequence and close to the carboxylic
end. Both of the sequences exhibited a decisive effect on the AAT
and its isoforms. The DFGWG sequence maintained the conforma-
tional integration of AAT enzyme structure whereas the HXXXD
motif was out of action without the histidine residue.48–50
As expected, the AAT expression level was positively correlated
with the total ester content as strawberries ripened and devel-
oped, which further confirmed the involvement of AATs in ester
production. Several AAT genes were extracted from F. ananassa
(SAAT, and AAT2),51,52 wild strawberry F. vesca (VAAT),53 and
F. chiloensis (FcAAT1)54 in studies to prove this theory. The max-
imum transcript levels of AATs were observed in red-ripe fruits,
and their expression was parallel with ester production. Never-
theless, different AATs acted on different substrates. Strawberry
alcohol acyl-transferase preferred the medium-chain aliphatic
alcohols in combination with different acyl-CoAs, whereas FaAAT2
exhibited higher activities with short straight-chain alcohol and
aromatic alcohols, such as cinnamyl alcohol.51,52 Wild strawberry
alcohol acyl-transferases and SAAT were shown in phylogenetic
analysis as closely related, and VAAT acted on short-chain alcohol
substrates.53 The discrepancy in AATs and the corresponding sub-
strates resulted in different ester categories and contents among
F. vesca and Tudnew, Carisma, Camarosa, Sweet Charlie, and Eris
in F. ananassa.55 Different cultivars and changes in volatile esters
in strawberries can thus be identified through the analysis of the
transcript expression of AATs.
Terpenoids
Among terpenoid compounds, volatile monoterpenes (C10 )
and sesquiterpenes (C15 ) were identified in most soft fruits.5
In cultivated ripe strawberry, linalool, and nerolidol were
detected as volatile terpenoids, whereas 𝛼-pinene, 𝛽-myrcene,
𝛼-terpineol, 𝛽-phellandrene, and myrtenyl acetate, were iden-
tified in wild strawberries.56 It is interesting that three triter-
penoids – euscaphic, tormentic, and myrianthic acids – were
found as antifungal phytoalexins in unripe Houkouwase fruit
and were not characteristic volatiles.57 Recently, it was reported
that, in Falandi strawberries, nine sesquiterpenoids and three
triterpenoids were isolated as nonphenolic constituents for their
antidiabetic, antitumor, and antioxidant effects.58
According to previous studies performed on terpenes, ter-
penoids were biosynthesized through the mevalonate pathway
(MVA pathway) or the 2-C-methyl-D-erythritol 4-phosphate path-
way (MEP pathway); the former occurred in cytosol, and the
latter transpired in plastids.59–61 The MVA pathway was not abso-
lutely separated from the MEP pathway because of the exchange
across plastidial membrane by unknown transporters, and the
establishment of so-called ‘metabolic cross-talk’ (Fig. 2).62
The gene expression and action substrates of terpene synthase F.
ananassa nerolidol synthase 1 (FaNES1), was revealed in different
stages and fruit tissues of cultivated strawberries. In wild strawber-
ries, F. vesca pinene synthase (FvPINS) was reported to be involved
in a typical monoterpenes biosynthesis, and FvPINS protein was
proven effective in the biosynthesis of 𝛼-pinene, 𝛽-phellandrene,
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Figure 2. The metabolic pathways of terpenoids in strawberries: IPP, isopentenyl diphosphate; DMAPP, dimethylallyl diphosphate; GPP, geranyl
diphosphate; FPP, farnesyl diphosphate; TPS, terpene synthases.

and 𝛽-myrcene from GPP.55 As reported, FvPINS converted GPP
into 𝛼-pinene, and the enzyme encoding cytochrome P450 gene
(pinene hydroxylase) catalyzed the transformation from 𝛼-pinene
to myrtenol.55 In the research, FaNES1 and FvPINS gene expres-
sions accounted for different dominant terpenes in cultivated and
wild strawberries.56
manner; for instance, ‘Strawberry Festival’ and ‘Florida Radiance’
possessed high thioester concentrations, whereas ‘Dover’, ‘Rosa
Linda’, and ‘Florida Belle’ contained higher sulfide and lower
thioester concentrations.66 Through the measurement of these
sulfur compound OAVs, methanethiol was determined to be the
predominantly active volatile sulfur compound.66

Other volatile compounds

Volatile benzenoids
Sulfur compounds
In ripe strawberries, volatile benzenoid concentrations were mea-
Sulfur compounds influence volatiles, because of the low level
sured at trace levels compared with those characteristic volatile
of their odor thresholds.63 Sulfur compounds in strawberries
compounds like esters, terpenes and furnanones. The volatile con-
were first reported in 1963.64 However, there are still difficul-
centrations of benzyl alcohol and benzyl acetate decreased as the
ties in exploring trace contents and the biosynthesis of sulfur
strawberries ripened and developed.68
compounds in strawberries. To date, 19 sulfur compounds
These volatile benzenoids were known to be derived from
have been found in strawberries. Such compounds include 10
the shikimate pathway via L-phenylalanine, and the first step
thioesters, eight alkyl sulfides, and an unknown component. The
of this pathway involves the deamination of L-phenylalanine
thioesters consist of methyl thioacetate, methyl thiobutyrate,
to trans-cinnamic acid by phenylalanine ammonia-lyase (PAL).69
methyl thiopropionate, ethyl thiobutanoate, methyl thiohex-
The reaction catalyzed by phospho-2-dehydro-3-deoxyheptonate
anoate, methyl (methylthio) acetate, ethyl (methylthio) acetate,
aldolase (DAHPS) represented the rate-determining step in this
methyl 2-(methylthio) butyrate, methyl 3-(methylthio) propi-
pathway.70 Benzenoids were also formed via a CoA-dependent,
onate, ethyl 3-(methylthio) propionate, and methyl thiooctanoate,
𝛽-oxidative pathway from L-phenylalanine71 and cinnamate. Coen-
and alkyl sulfides comprise hydrogen sulfide, methanethiol,
zyme A ligase (CNL) was reported as a significant peroxisome that
dimethyl sulfide, dimethyl disulfide, dimethyl trisulfide, and sul-
catalyzed the cinnamoyl-CoA formation in this pathway.72 Later,
fur dioxide.65,66 Except for hydrogen sulfide and methanethiol
through the isolation and analysis of strawberry-volatiles-related
contents remaining relatively consistent during ripening, most
genes, the F. ananassa cinnamate CNL (FaCNL) gene, F. ananassa
sulfur compounds increased as fruit ripened. It was also revealed
DAHPS (FaDAHPS1 and FaDAHPS2) gene, and F. ananassa PAL
that most sulfur compounds increased dramatically during the
(FaPAL1 and FaPAL2) gene were reported to be related to the
commercial ripening, full ripening, and over-ripening stages.65
biosynthesis of volatile benzenoids in strawberries.68
Between the full ripening and the over-ripening stages, in par-
As previous research provided limited information on volatile
ticular, the concentration of total sulfur compounds doubled.65
benzenoids in strawberries, further studies should identify the
Later studies on strawberry sulfur compounds showed that a
influence of volatile benzenoids on strawberries and biosynthetic
puree process in the experimental pretreatment of strawberries
mechanisms.
affected the analysis of volatile sulfur compounds; except for
methanethiol, the dimethyl disulfide and dimethyl trisulfide in
the puree exhibited lower amounts while most sulfur compounds
were similar to, or higher than, those in intact strawberries.67 EFFECTS OF POSTHARVEST FACTORS ON
Principal component analysis (PCA) of sulfur compounds among VOLATILES PERFORMANCE
12 strawberry cultivars showed differences in constituents and Considerable attention has been focused on the variations in the
in the amounts of sulfur compounds in a cultivar dependent characteristic volatile compounds during postharvest storage. We

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The aroma volatile repertoire in strawberry fruit
will thus discuss the influence of postharvest treatment on straw-
berry volatiles. The temperature, atmospheric gas, and exogenous
postharvest treatment (spraying, coating or dipping) contribute to
the maintenance of the fruit’s postharvest quality.
Temperature
Low temperature is widely known to decrease the respiratory
rate and metabolic activity in fruits. Studies have been inconsis-
tent regarding the influence of temperature on different volatile
compounds. Except for 3-hexenyl acetate and ethyl butanoate,
furanones and most volatile esters increased gradually with
increasing temperature.19,73,74 As for the effect of temperature on
volatile terpenes, a report revealed that the terpene content in
‘Akihime’ after 9 days of storage at low temperature declined less
than that at room temperature,75 whereas another study found
that in white ‘Sweet Charlie’ fruit, after storage at 15 and 25 ∘ C
separately, terpenes increased with elevation in temperature.68
The acid and alcohol content in strawberries after storage at low
and room temperatures showed minimal changes with variation
in temperature.75 Volatile benzenoids, including benzyl alco-
hol and benzyl acetate, were down-regulated by an increase in
temperature.68
Recently, further studies concentrated on the effects of molecu-
lar factors on strawberry volatiles. The quantity of FaQR declined
remarkably at low temperature versus room temperature. The
quantity of pyruvate decarboxylase isozyme 2 (PDC2) was also
lower at low temperature.75 In a dark environment, the increase
in temperature was accompanied by up-regulated expressions of
FaQR and FaOMT but decreased transcriptions of FaAAT, FaNES,
and FaPAL1.68 The same research showed that total esters and ter-
penes increased with increased temperature, and the phenomena
were probably caused by the interaction of light and temperature
or some unknown pathway involved in the volatile biosynthetic
compounds.68
Atmosphere
The production of volatiles was affected by certain modified
atmospheric conditions, which were usually applied to main-
tain fruit quality, such as controlled-atmosphere (CA) and
modified-atmosphere packaging (MAP). Nevertheless, inade-
quate or extreme atmospheres could result in the fruit possessing
an ‘off’ flavor.
Modified atmosphere packaging was carried out to fill the pack-
age with a particular ‘atmosphere’ to prolong the shelf life of fresh
fruits. It was reported that both total volatile compounds and
the butanoate amount in strawberry under MAP conditions (11%
CO2 + 11% O2 + 78% N2 or 100% CO2 ) were lower than control.76
However, in another study, the profile of the volatiles in Honeoye
strawberries remained constant during storage in the 11–14% O2
and 9–12% CO2 packages, whereas a considerable increase was
observed in ethyl acetate levels in ‘Korona’ fruits.77 The cultivar-
and temperature-dependent ‘off’ odors were reported to result
from CO2 treatment.78
The atmosphere is adjusted to a particular range in CA packag-
ing to maintain fruit quality and delay senescence. The ethanol and
ethyl acetate content in wild strawberries stored at 15% CO2 /6%
O2 and 10% CO2 /11% O2 increased 42 times and 12 times com-
pared to their initial value, respectively; and the treatment con-
tributed to the elucidation of the relationship between ‘off’ flavor
development and high CO2 concentration.79 The acetaldehyde
concentration increased as CO2 partial pressure decreased, and
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the acetaldehyde content in the untreated control was three times
higher than that in CA storage.79 Furthermore, other research
reported that the formation of volatile esters and volatile fura-
nones was promoted and the decline in volatile acids, alcohol, and
terpenes was delayed in 2% O2 and 12% CO2 CA storage.75
The influence of atmosphere on the biosynthesis of volatiles
varied. It was reported that, in strawberries, AAT activity was in
response to exogenous high CO2 , as methyl and ethyl acetate
were the characteristic ester compounds.80 The abundance of
quinone oxidoreductase dramatically decreased in CA, whereas
PDC2 showed higher abundance in strawberries under CA.75 These
results proved that the expressions of both quinone oxidoreduc-
tase and pyruvate decarboxylase which coincided with volatile
performance were in response to CA condition.75
Exogenous postharvest treatments
Exogenous spraying, coating, or dipping were widely used to pro-
long the shelf life of strawberries.81 Previous studies examined
the effects of exogenous treatments on strawberry quality, but
little information exists on the effects of exogenous treatments
on strawberry volatiles. As expected, in the non-climacteric fruit,
exogenous ethylene and 1-methylcyclopropene (1-MCP) showed
minimal effect on strawberry volatiles.82 However, the applica-
tion of exogenous cytokinin significantly reduced total volatile
amounts, including esters and alcohols, and demonstrated that
exogenous cytokinin may inhibit glycolysis, fatty acid metabolism,
and the conversion from sugar to esters and alcohols in ‘Akihime’
strawberries.83 Strawberries treated with 0.1% hexanal showed
a remarkable decline in LOX activity but a significant increase
in phospholipase D (PLD) activity; these changes are related to
the volatile biosynthesis during ripening.84 Strawberries treated
with 0.5–1 mmol L−1 phenylethyl alcohol after 15-day storage still
showed a similar volatile profile to fresh strawberries; however, no
report was available on the quantities of specific compounds that
might cause an ‘off’ flavor in strawberries.85
Methyl jasmonate (MJ) is a natural hormonal compound that
can reduce decay and maintain the quality of fruits and veg-
etables. Strawberries treated with MJ plus ethanol (MJ-ETOH)
showed higher concentration of the major volatile compounds
including methyl acetate, isoamyl acetate, ethyl hexanoate, butyl
acetate, and hexyl acetate, than independent ethanol treatment
and control.86 Another research also reported that MJ treatment
increased the content of 1-octanol, 1-hexanol, linalool, furaneol,
2-heptanone, and ethyl hexanoate after five or seven-day stor-
age, whereas MJ exhibited less effect on ethyl butanoate.87 There
was evidence that 1-hexanol content declined after 7 days due
to an increase in AAT in response to MJ treatment.87 Different
enantiomers of MJ also affected the volatile profiles differently.
Interestingly, the (+)-MJ inhibited the formation of ethyl 2-methyl
butanoate and isoamyl acetate but promoted the formation of
ethyl hexanoate and hexyl acetate, whereas (−)-MJ inhibited the
formation of these four esters; however, the mixture of (+)/(−)-MJ
exhibited lower activity than separate (+)-MJ or (−)-MJ.88
Polysaccharides, proteins, and lipids, or their combinations, were
used as edible coatings to maintain fruit quality and extend
shelf life. Chitosan, the polysaccharide obtained from seafood,
appeared to be an ideal preservative for fresh berries. Some studies
were thus conducted on the responses of volatiles in strawberries
to chitosan coating. A large delay was noted in build-up of ‘off’
flavors, attributed to acetaldehyde and ethanol, in strawberries
dipped in chitosan acetate solution at 1% or 1.5% (w/w), and
coated fruits exhibited enhanced levels of ethyl butanoate, ethyl
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hexanoate, and acetate esters.89 Another study also reported that
pure chitosan promoted production of esters and dimethyl furfural
after coating, and coatings containing lemon essential oil add
terpenes (limonene, 𝛾-terpinene, p-cymene, and 𝛼-citral) to fruit
volatiles, and thus also accelerated fermentation and modified
the typical composition of the fruit volatiles.90 Strawberry volatiles
may be enriched under the influence of chitosan with lemon
essential oils or other volatile compounds.90
Ozone treatment was reported to reduce the incidence of
decay and affect the development of volatiles. However, no
‘off’ flavor was observed in fruits treated by ozone at 1.5 𝜇L L−1
concentration,91 which was contrary to previous reports that an
‘off’ flavor occurred under an ozone atmosphere.20 Combined
treatment (UV-C light, gaseous O3 , superatmospheric O2 , and
high CO2 ) of strawberries caused sensory property loss because of
off-flavor development.92
Ultra-violet radiation is mainly used to control decay, and stud-
ies have concentrated on its antimicrobial effect, particularly on
cell-wall metabolism.93,94 Recently, volatile ester contents in 3/4
ripe ‘Aromas’ strawberry were increased remarkably increased by
4.35 kJ m−2 UV-C treatment compared to control. Transcript lev-
els of ADH and AAT also exhibited a notable increase on the sec-
ond and fourth days after treatment.95 This research also revealed
that UV-C treatment seemed to induce accumulation of the aller-
genic protein Fra a1.95 However, the linalool content remained
consistent, while both furaneol and mesifurane concentrations
decreased in response to the UV-C treatment.96
FUTURE PROSPECTS AND CONCLUSIONS
Although the characteristic volatile compounds in strawberry have
been identified and quantified, research on the biosynthetic path-
ways and precursor validation of strawberry volatiles is still limited.
Recently, it was reported that the nonvolatile compounds turned
to volatiles after hydrolysis of the bonds between the sugar and the
aglycone.97 Nevertheless, there are still some gaps between the
volatile biosynthesis and performance; for example, the process
by which d-fructose-1,6-diphosphate converts into HMMF is still
unclear and the biosynthesis of sulfur compounds is still unknown.
Recently our laboratory focused on volatile profiles in response
to exogenous hormone and abiotic stress. At the proteomic
level, it was found that the expression of pyruvate decarboxylase
(PDC) 2 and acetyl-CoA carboxylase (ACC) was down-regulated
by application of exogenous forchlorfenuron (CPPU). This result
provided evidence that CPPU suppressed volatile biosynthesis in
strawberries.83 In CA and low-temperature conditions, the expres-
sion of FaQR and PDC2 coincided with the accumulation of
volatiles during storage.75 These results revealed a new trend
of studying strawberry volatile metabolism at the proteomic,
metabolomic and transcriptomic levels.
In conclusion, aroma is a significant factor contributing to straw-
berry flavor, which is an important quality that influences con-
sumer acceptability. Strawberry aroma is one of the most com-
plex aromas, with more than 360 volatile compounds. Fura-
nones, esters, terpenoids, sulfur compounds, and benzenoids are
regarded as the main characteristic volatile compounds. This
review is a comprehensive and updated overview of the features
and metabolisms of the major volatile compounds in strawberry,
and the effects of postharvest treatment factors, such as tempera-
ture, atmosphere, exogenous hormone, edible coating, ozone, UV
radiation, on volatiles in strawberries. However, the research gap
between volatile biosynthesis and performance needs to be filled.
wileyonlinelibrary.com/jsfa © 2018 Society of Chemical Industry
J-w Yan et al.
ACKNOWLEDGEMENTS
This work was supported by the National Key Research and Devel-
opment Program of China (No. 2017YFD0401304), the Provincial
Natural Science Foundation of Zhejiang (No. LY17C200008 and
LY17C200014), and the Research Foundation for Talented Scholars
in ZUST (No. F701103G01).
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