Dynamic Changes in Health‐Promoting Properties and Eating Quality During Off‐Vine Ripening of Tomatoes

Full text

Dynamic Changes in Health-Promoting Properties
and Eating Quality During Off-Vine Ripening
of Tomatoes
Mohammed Wasim Siddiqui , Isabel Lara, Riadh Ilahy, Imen Tlili, Asgar Ali, Fozia Homa, Kamlesh Prasad ,
Vinayak Deshi, Marcello Salvatore Lenucci, and Chafik Hdider
Abstract: Tomato (Solanum lycopersicon L.) fruit is rich in various nutrients, vitamins and health-promoting molecules.
Fresh tomatoes are an important part of the Mediterranean gastronomy, and their consumption is thought to contribute
substantially to the reduced incidence of some chronic diseases in the Mediterranean populations in comparison with
those of other world areas. Unfortunately, tomato fruit is highly perishable, resulting in important economic losses
and posing a challenge to storage, logistic and supply management. This review summarizes the current knowledge on
some important health-promoting and eating quality traits of tomato fruits after harvest and highlights the existence of
substantial cultivar-to-cultivar variation in the postharvest evolution of the considered traits according to maturity stage
at harvest and in response to postharvest manipulations. It also suggests the need for adapting postharvest procedures to
the characteristics of each particular genotype to preserve the optimal quality of the fresh product.
Keywords: eating quality, health-promoting properties, physical attributes, postharvest, tomato
Introduction
Fresh tomato (Solanum lycopersicon L.) fruits pose an important
set of challenges for postharvest storage due to their high-water
content and soft texture. These attributes make tomatoes highly
perishable and difficult to store for long periods without incur-
ring substantial losses and additional costs. After harvest, tomato
fruits are no longer supplied with water and solutes by the parental
plant; thus, storage conditions play a fundamental role in slowing
down fresh product decay quality traits. During tomato fruit ripen-
ing and senescence several biodegradation processes occur, in-
cluding macromolecule depolymerization, substrate consumption,
chloroplasts-to-chromoplasts transition and pigment alterations,
mostly due to the hydrolytic activity of glycosidases, esterases, de-
hydrogenases, oxidases, phosphatases and ribonucleases (Tadesse,
CRF3-2018-0122 Submitted 6/1/2018, Accepted 8/30/2018. Authors Siddiqui
and Deshi are with the Dept. of Food Science and Postharvest Technology, Bihar
Agricultural Univ., Sabour - 813210, Bhagalpur, Bihar, India. Author Lara is with
the Dept. de Quı́mica, Unitat de Postcollita-XaRTA, Univ. de Lleida, Rovira Roure
191, 25198 Lleida, Spain. Authors Ilahy, Tlili, and Hdider are with the Lab. of
Horticulture, Natl Agricultural Research Inst. of Tunisia (INRAT), Univ. of Carthage,
Tunis, Rue H´
edi Karray 2049 Ariana, Tunisia. Author Ali is with the Centre of
Excellence for Postharvest Biotechnology (CEPB), School of Biosciences, The Univ.
of Nottingham Malaysia Campus, Semenyih 43500, Selangor, Malaysia. Author
Homa is with the Dept. of Statistics, Mathematics, and Computer Appplication,
Bihar Agricultural University, Sabour - 813210, Bhagalpur, Bihar, India. Author
Prasad is with Dept. of Food Engineering and Technology, Sant Longowal Inst. of
Engineering and Technology, Longowal - 148106, Punjab, India. Author Lenucci
is with Dipt. di Scienze e Tecnologie Biologiche ed Ambientali, Univ. del Salento
(DiSTeBA), Via Prov.le Lecce-Monteroni, 73100 Lecce, Italy. Direct inquiries to
author Siddiqui (E-mail: wasim_serene@yahoo.com).
Workneh, & Woldetsadik, 2012). Ripening and senescence are
also associated with the de novo biosynthesis of proteins, nucleic
acids, lipids and secondary metabolites including carotenoids (par-
ticularly lycopene) and flavor-related aroma volatiles, as well as to
processes involved in mitochondria maintenance through tran-
scriptional, posttranscriptional, translational and/or posttransla-
tional regulation mechanisms (Workneh & Osthoff, 2010).
In order to preserve satisfactory health-promoting, eating
and processing quality, consider all the major physiological and
biochemical characteristics of tomato fruits is important. Besides
flavor, good quality involves appearance, texture and functional
properties, attributes that generally deteriorate over time until
delivery to the final consumer. The major issue with fresh
tomato storage, transport and marketing is the relatively fast
quality deterioration resulting in short shelf-life potential. Hence,
more intensive research efforts are required to cut quality loss
and extending shelf-life. Tomato fruit ripening regulation has
been a central investigation focus throughout the last years. This
paper reports a brief review of the recent investigations related
to postharvest alterations occurring in the content of bioactive
molecules, health-promoting properties, biochemical attributes
and physical parameters of tomato fruit in response to different
factors, including genotype and postharvest manipulation.
Changes in Bioactive Molecules and
Health-Promoting Properties during Off-Vine
Ripening of Tomato
Health-promoting properties of fruits originate from their
content in bioactive molecules capable of partially preventing or
1540 ComprehensiveReviews in Food Science and Food Safety rVol. 17, 2018
C2018 Institute of Food Technologists®
doi: 10.1111/1541-4337.12395

Dynamic changes in health-promoting properties . . .
delaying oxidative reactions arising from the presence of metabol-
ically or environmentally originated free radicals. These extremely
reactive species display one or more unpaired electrons, and com-
prise mainly superoxide anions, hydroxyl and peroxyl radicals.
Although human cells possess endogenous antioxidant systems, the
dietary intake of exogenous phytochemicals is required to match
the overall antioxidant activity and efficiently counteract radical-
driven damage, especially during aging and/or stressful conditions.
Thus, the antioxidant power and chemical composition of fresh
fruits are becoming determinant attributes in tomato marketing as
they purportedly contribute to the health-promoting properties
of the product. A large part of the health benefits derived from
the consumption of plant-derived foods has been attributed to
hydrophilic and lipophilic bioactives, mainly ascorbic acid (AsA),
glutathione, folates, tocols, carotenoids, and phenolics, although
the health claims remain in many cases to be clearly established in
vivo (Esp´
ın, Garc´
ıa-Conesa, & Tom´
as-Barber´
an, 2007).
In this context, tomato is thought to contribute substantially
to decrease the occurrence of some chronic diseases in the
Mediterranean population compared to other world areas, as it
is a major source of the above-mentioned nutrients (Abushita,
Daood, & Biacs, 2000; Carluccio, Lenucci, Piro, Siems, & Lu˜
no,
2016; Mart´
ınez-Valverde, Periago, Provan, & Chesson, 2002).
Carotenoids are the major phytochemicals in tomato, lycopene
accounting for up to 90% there of (Ilahy et al., 2018; Ilahy, Hdider,
Lenucci, Tlili, & Dalessandro, 2011). However, available informa-
tion on changes in the content of bioactive compounds during
postharvest ripening is generally scarce. In the following subsec-
tions, we provide a brief overview of the published reports on
postharvest modifications in the qualitative and quantitative pro-
files of some of the most important bioactive molecules of fresh
tomato fruits.
Total phenolics
Phenolic acids and two flavonoid families (flavanones and
flavonols) represent the most abundant phenolics in tomato. Phe-
nolic acids account for up to 75% of total phenolics in tomato fruit,
with large genotype variability. They are distributed both in the
pericarp (skin) and inner (mesocarp and endocarp) fruit tissues,
mainly as chlorogenic acid (Mart´
ınez-Valverde et al., 2002; Moco
et al., 2007). Most (98%) of flavonols occur, instead, in the fruit
skin with concentrations largely variable across cultivars (Stewart
et al., 2000). Finally, the stilbenoid resveratrol has also been found
in tomato fruit skin at full ripeness (Ragab, Fleet, Jankowski, Park,
& Bobzin, 2006).
The metabolic pathways involved in the biosynthesis of the
different families of phenolics are complex, closely interrelated,
and profoundly influenced by endogenous and exogenous factors
(Dixon & Steele, 1999; Manach, Scalbert, Morand, R´
em´
esy, &
Jim´
enez, 2004). Accordingly, significant changes in the content
of total phenolics in tomato are expected in response to pre-and
postharvest conditions (Dumas, Dadomo, Di Lucca, & Grolier,
2003; Slimestad & Verheul, 2005). Owing to the quantitative
and qualitative relevance of lycopene and β-carotene, research
on health-promoting properties of tomato fruits has preferentially
focused on these constituents, generally overlooking flavonoids.
Substantial cultivar-to-cultivar variation in the metabolism of
phenolics and flavonoids during tomato ripening and postharvest
has been reported (Table 1). Focusing on phenolics and flavonoids
accumulation in ripening ordinary and high-lycopene tomato cul-
tivars, Ilahy et al. (2011) reported significant differences in the
total phenolic levels even among cultivars of the same typology.
Cultivar “HLY18” attained a peak [310 mg Gallic Acid Equivalent
(GAE)/kg Fresh Weight (FW)] of phenols at the orange-red stage,
while in “HLY13” fruits, two peaks were detected at the green
and orange-red ripening stages (223 and 240 mg GAE/kg FW,
respectively). However, at the same ripening stages, the ordinary
cultivar “Rio Grande” exhibited the lowest levels of total phenols
(113 and 138 mg GAE/kg FW, respectively). The flavonoid levels
varied widely throughout ripening stages. The dynamic of change
in flavonoids was identical in high-lycopene cultivars, but quanti-
tatively different in “Lyco2” fruits. Flavonoid contents remained
essentially stable at later maturity stages. It should be underlined
that flavonoid contents were higher in high-lycopene tomato cul-
tivars studied throughout ripening.
The effect of storage on phenolics is well documented in
tomato fruits. The amount of chlorogenic acid and chalconarin-
genin, quantitatively prominent flavonoids in cherry tomatoes,
has been reported to decrease sharply during 3 weeks postharvest
storage at 20 °C (Slimestad & Verheul, 2005), although this loss
was less pronounced at lower storage temperature. This change
is an important issue if the health-promoting properties of the
product must be preserved, because direct correlation was ob-
served between chalconaringenin levels and antioxidant activity.
However, the total amount of phenolics was unchanged during
the same period, suggesting that other compounds compensate
the decrease. Actually, an earlier report found higher amounts of
total phenolics in “Moneymaker” tomatoes after 16 days at 20 °C
(Giovanelli, Lavelli, Peri, & Nobili, 1999). Cold storage (6 °C) for
up to 4 weeks also led to a decreased content of total phenols and
chlorogenic acid in Micro-Tom’ tomato fruit (G´
omez et al., 2009).
Some treatments have been proposed to alleviate the postharvest
decrease in total phenolics. Brassinolide treatments (immersion in
3or6μM solution for 5 min) were found to significantly increase
the total phenolic content of tomatoes after 3 weeks storage
at 1 °C compared to untreated fruits (Table 1). Interestingly it
was associated with the simultaneous increase of phenylalanine
ammonia-lyase activity, a key enzyme of phenol biosynthesis
(Aghdam, Asghari, Farmani, Mohayeji, & Moradbeygie, 2012).
High-voltage electrostatic field (HVEF) pretreatments also in-
creased the levels of total phenols of green ripe tomato fruits after
24 days storage at 13 °C, compared to the control samples (Zhao,
Hao, Xue, Liu, & Li, 2011). Similarly, direct-electric-current
application in “Pannovy” tomatoes increased total phenols by
up to 120% in the 24 hr following the treatment (Dannehl,
Huyskens-keil, Eichholz, Ulrichs, & Schmidt, 2011). Delactosed
whey permeate (DWP), a novel bio-active product for fresh
products storage, has been shown to improve total phenols in
“Moneymaker” tomatoes after 21 days at 15 °C, concurrently
preserving firmness, appearance and aroma, and reducing decay
incidence (Ahmed, Mart´
ın-Diana, Rico, & Barry-Ryan, 2013).
Furthermore, tissue-specific expression of AtMYB12 (an
Arabidopsis thaliana transcriptional activator of the caffeoyl quinic
acid biosynthesis) in “Micro-Tom” and “Moneymaker” tomato
backgrounds was found to trigger the accumulation, at very high
levels (up to 65-fold higher than controls), of flavonol antioxidants
in the ripe fruits, as a result of the up-regulation of most genes
involved in phenyl propanoid biosynthetic pathway, including
those encoding for phenylalanine ammonia-lyase, chalcone
synthase and flavonol-3-glucosyltransferase, whose expression was
increased over 100-fold (Luo et al., 2008). This expression led
to a significant increase of the hydrophilic antioxidant activity in
the transgenic fruits, and exemplifies the possibility of obtaining
fruits fortified in phenolics. Although transgenic approaches
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Dynamic changes in health-promoting properties . . .
Table 1–Variation in total phenolics and flavonoid content during ripening and postharvest of tomato fruit.
Compounds Tomato cultivar name
Harvest conditions and
applied treatments Observed changes References
Total phenolics Cv. Rio Grande and
high-lycopene tomato
cvs HLY13, HLY18,
and Lyco2
Ripening from green to
red-ripe stage
Cultivar-dependent (Ilahy et al., 2011)
Flavonoid Almost unchanged
Chalconaringenin and
chlorogenic acid
Cherry cv. Jennita Postharvest ripening at 20 °C
for 3 weeks in darkness of
green to red ripe tomato
fruits
Sharp decrease (Slimsestad and
Verheul, 2005)
Postharvest ripening at low
temperature
Low decrease
Total phenolics Cv. Moneymaker 7 different ripening
stages/Postharvest storage
for16daysat20°C
Increased content (Giovanelli et al.,
1999)
Total phenols and
chlorogenic acid
Cv. Micro-Tom Cold storage of breaker and
red-ripe tomato fruits for
4 weeks at 4 °C
Decreased content (G´
omez et al., 2009)
Total phenolics Cv. Newton Storage of mature green fruit
at 1 °Cfor3weeksfollowed
by 3 or 6 μM brassinolide
dip (5min)
Increased content (Aghdam et al.,
2012)
Cv. Chaoyan-219 HVEF treatment applied to
green mature tomato fruits
followed by storage at 13 °C
(24 days)
Increased content (Zhao et al., 2011)
Cv.Pannovy DC application on red-ripe
fruits
Increased content (Dannehl et al.,
2011)
Cv. Moneymaker DWP treatment of ripe
tomato followed by a
storage at 15 °C (21 days)
Increased content (Ahmed et al., 2013)
Flavonol Cvs Micro-Tom and
“Moneymaker
Expression of flavonol-specific
transcriptional activator
65-fold increase (Luo et al., 2008)
Total phenolics and
flavonoids
Manapal (hp-2dg) Integration of hp and ip
mutations in tomato
cultivars compared to
ordinary cvs (green, breaker
and ripe stages of maturity)
High initial content
and lower
postharvest
decrease than the
wild type
(Bino et al., 2005)
Line (n-935) (hp-2dg)(Kolotilin et al.,
2007)
Different hp and ip cvs (Ilahy et al., 2017)
Cvs, cultivars; DC, Direct Electric Current; DWP, Delactosed Whey Permeate; hp, high pigment; hp-2dg, high-pig;ent dqrk green; HVEF, high voltqge electric field; ip, intense pigment.
achieved promising results in increasing several phytochemicals in
tomato fruit (Fraser et al., 2002; Ronen, Carmel-Goren, Zamir,
& Hirschberg, 2000; Rosati et al., 2000), some criticism occurred
because only a single or few compounds are enhanced. However,
the use of hp and ip genotypes ensures a simultaneous increase in
most secondary metabolites without quality compromise (Bino
et al., 2005; Ilahy et al., 2017; Kolotilin et al., 2007).
Ascorbic acid
AsA is a major indicator of the nutritional value of fresh plant
products; thus, the monitoring of the dynamic changes in its level
after harvest and during storage is of interest. A cultivar-dependent
pattern of change in AsA levels has been reported during ripening
(Table 2). AsA was found to increase in the first phases of ripening
and remain either steadily stable or slight decline at the end of the
process (Giovanelli et al., 1999; Tigist, Workneh, & Woldetsadi,
2013). The decline was attributed to the involvement of AsA
in detoxifying the reactive radicals generated by the increase in
respiration rates typical of climacteric fruits (D´
avila-Avi˜
na et al.,
2011). Accordingly, a survey on different cultivars found the
highest AsA contents (184 to 233 mg/kg FW, depending on the
cultivar assessed) in firm ripe fruits, while a slight decrease (165 to
217 mg/kg FW) was observed in the soft ripe ones (Singh, Ray, &
Mishra, 1983). When AsA levels were evaluated in the fruits of the
tomato cultivar “Floriset” at four sequential ripening stages, the
highest concentration was observed when the fruits were turning
yellow, followed by a decrease at more advanced maturity stages
(Abushita, Hebshi, Daood, & Biacs, 1997). In turn, Islam, Matsui,
and Yoshida (1996) and Pila, Gol, and Rao (2010) observed the
highest AsA amounts at the pink stage. In contrast, AsA contents
in “Marmande-Cuarenteno” and “Ailsa Craig” tomatoes were
observed to remain essentially stable along ripening, and to
increase slightly in fully ripe fruit (Cano, Acosta, & Arnao, 2003;
Jim´
enez et al., 2002). Similarly, a progressive increase in AsA
content between the green and the red-ripe stages was reported in
the fruits of “Pant T-3,” “Pant 2466-27,” and “Pusa Hybrid-1”
tomato genotypes, whereas in “SG-12” and “MTH-1” lines AsA
peaked at the yellow stage (Siddiqui, Gupta, & Pandey, 1986).
High-pigment or high-lycopene tomato cultivars were claimed
to have superior functional quality, leading to good postharvest
quality. Therefore, Ilahy et al. (2011), 2018) compared the
levels of AsA, dehydroascorbic acid (DHA) and total vitamin C
(AsA +DHA) in various high-lycopene tomato cultivars during
ripening and the ordinary cultivar “Rio Grande” (Table 2). Again,
the levels of AsA, DHA, and total vitamin C were significantly
different throughout ripening and a genotype-dependent pattern
of change was observed. The fruits of the cultivars “HLY18,”
“HLY13,” and “Rio Grande” exhibited a peak in total vitamin
C content at the orange-red ripening stage (333, 230, and
221 mg/kg FW, respectively). However, “Lyco2” fruits showed
the highest total Vitamin C content at the green-orange and
red-ripe stages. Nevertheless, the fruits of both “Lyco2” and
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Dynamic changes in health-promoting properties . . .
Table 2–Variation in ascorbic acid content during ripening and postharvest of tomato fruit.
Compounds Cultivar name
Harvest conditions and
applied treatments Observed changes References
AsA Cv. Moneymaker Different ripening stages Initial increase followed by
a decrease at final stages
(Giovanelli et al.,
1999)
Roma VF, Melkasalsa,
Melkashola, Metadel,
Eshete, Marglobe Improved,
Fetane, Heinz-1350 and
Bishola)
(Tigist et al., 2013)
Cv Sabour Prabha (Singh et al., 1983)
Cvs DRW 3042, Primato,
Floriset, Katinka, Selma,
Revido, DRW 3126, Gitana,
Ultimo, Relento, Pankor,
Delfino, Tampo, Monica and
Falcato
Peak at turning yellow stage
followed by a decrease
(Abushita et al., 1997)
Cv. Momotaro Peak at the pink stage (Islam et al., 1996)
Cv. Himsona (Pila et al., 2010)
Cv. Marmande-Cuarenteno Constant during maturation
and increase at full
red-ripe stage
(Cano et al., 2003)
Cv. Ailsa Craig (Jim´
enez et al., 2002)
Cvs SG-12, Pusa Hybrid-1,
Marglobe, Roma and Pusa
Ruby,
Cultivar-dependent trend (Siddiqui et al., 1986)
Total
Vitamin C
Cv. Rio Grande and
high-lycopene tomato cvs
HLY13, HLY18, and Lyco2)
Ripening of hp tomato
cultivars
Higher trends and
accumulation levels
compared to “Rio Grande’
(Ilahy et al., 2011)
AsA Cv. Moneymaker Off-vine ripening Lowest levels at green stage
increasing till the red-ripe
stage
(Giovanelli et al.,
1999)
Cvs Marglobe and Roma VF (Getinet et al., 2008)
Cvs Roma VF, Melkasalsa,
Melkashola, Metadel,
Eshete, Marglobe Improved,
Fetane, Heinz-1350 and
Bishola
Postharvest storage of
green-mature fruits for
20 days at room air
temperature 15.4-16.2 °C
and relative humidity of
34.8-52.4%
First increase followed by a
decrease after 20 days
(Tigist et al., 2013)
Cv. Zhenfen Storage of tomato fruits at
different ripening stages in
the dark at 14 °C, 95% RH
for up to 37 days
Increase from green to
red-ripe stage
(Liu et al., 2011)
Rio Grande Different packaging systems
(CaCl2-treated and
nontreated green mature
tomato fruits)
Significant increase along
ripening with a peak
between the pink-red and
the red stages of ripening.
Pretreatememt with CaCl2
prior to packaging
resulted with the highest
ascorbic acid levels
followed by nontreated
packed fruits
(Sammi & Masud,
2007)
Cv. Tradiro Storage the light-red fruits for
7, 15, 25 °C for a period of
10 days.
Slight increase (Toor & Savage, 2006)
Cv. Roma VF Long-term storage of
half-ripen tomato fruits
Sharp decrease (Moneruzzaman et al.,
2008)
Red-fruited cv. Cheers
Yelow-fruited cv. 6205
Processing of red and yellow
tomato cultivars
80% decrease (Georg´
e et al., 2011)
Vitamin C Red intense tomatoes from
commercial varieties
Pasteurization of tomato
pur´
ee prepared from
red-ripe tomato fruits
90% decrease (P´
erez-Conesa et al.,
2009)
AsA Peeled canned tomato
product
Cooking, boiling, frying,
drying Considerable loss (Giovanelli et al.,
2002)
Cvs Excell and Aranca (Sahlin et al., 2004)
NA Milder treatment and lower
temperature
Better retention (Davey et al., 2000)
AsA:Ascrobic acid; hp:high-pigment; NA:not available; RH:relative humidity
“HLY18” high-lycopene cultivars exhibited higher amounts
of total vitamin C than the ordinary cultivar “R´
ıo Grande”
all along ripening. Therefore, besides higher functional quality,
high-lycopene cultivars should exhibit higher postharvest storage
potential without quality compromise (Ilahy et al., 2017).
In addition to cultivar-dependent var iation, substantial differ-
ences in AsA content during postharvest storage have been ob-
served according to the maturity stage at harvest. Mature-green
harvested tomato fruits showed the lowest AsA content, with in-
creasing levels as the r ipening process advanced (Getinet, Sey-
oum, & Woldetsadik, 2008; Giovanelli et al., 1999). Accordingly,
Liu et al. (2011) reported that AsA contents increased from the
green till the red-ripe stage of ripening (from 27.2 to 92.3 mg/kg
FW) during dark storage of “Zhenfen” tomato fruits for up to
37 days at 14 °C, 95% relative humidity (RH). Generally speaking,
higher AsA contents were detected in light-red tomato fruits, but
C2018 Institute of Food Technologists®Vol.17,2018 rComprehensiveReviews in Food Science and Food Safety 1543

Dynamic changes in health-promoting properties . . .
Table 3–Variation in carotenoid content during off-vine ripening of tomato fruit.
Compounds Cultivar name
Harvest conditions and
applied treatments Observed changes References
Lycopene NA Different ripening of ordinary
tomato cultivars
Linear increase throughout
the ripening stages
(Collins et al., 2006)
Cvs Neris, Svara, Vyt˙
enųdidieji,
Jurgiai and Vaisa F1
(Radzeviˇ
cius et al.,
2009)
Cv. Arka Ahuti (Namitha et al., 2011)
Cv. Laura On-vine vs off-vine ripened
greenhouse tomato fruits
On-vine tomato has 32% less
lycopene than off-vine
ripened
(Arias et al., 2000)
NA Tomato fruit harvested before
full redness (at either the
breaker or turning stages)
Similarorhigherlycopene
content accumulated
compared to those
harvested at the soft red
stage
(Collins et al., 2006)
Carotenoids and
lycopene
Cvs HLY02, HLY13,
HLY18, and Kalvert
Ripening of hp tomato
cultivars
Higher trend in hp cultivars
with respect to ordinary cvs
(Lenucci et al., 2006)
Different hp and ip cvs (Ilahy et al., 2018)
Lycopene Cvs Excell and Aranca Long storage periods of
green-mature and red-ripe
Increased (Sahlin et al., 2004)
Cvs Roma VF, Melkasalsa,
Melkashola, Metadel, Eshete,
Marglobe Improved, Fetane,
Heinz-1350 and Bishola
(Tigist et al., 2013)
Cvs Vaisa, Svara and Neris Different ripening stages Lycopene content ranged
from 2.5 to 14.2 mg/kg FW
in Vaisa and Svara at the
green stage respectively.
Cultivar Neris had the highest
lycopene contents attaining
125.1 mg/kg FW at the
red-ripe stage
(Radzeviˇ
cius et al.,
2009)
Cv. Tradiro Storage of light-red tomato
fruits at 7, 15 and 25 °C
Brighter red color at 15 to
25 °Cthanat7°C
(Toor & Savage, 2006)
Cv Pyramid and a fresh market cv Hydroponic and ordinary
red-ripe tomato fruits
divided into halves and
stored at 22 °C for 14 days
Sharp increase (Ajlouni et al., 2001)
Cv. Himsona 10 days storage at ambient
conditions
Progressive increase up to
33.1 mg/kg FW
(Pila et al., 2010)
Cvs FL7692D, Suncoast carrying
old gold mutation, 97E212S
carrying ripenin inhibitor gene
and cvs Agriset, Equinox,
FL7655 and Solar Set
Storage of tomatoes
harvested at the breaker
stage
Peak after 6 days at room
temperature
(Thompson et al.,
2000)
Cvs FL7692D, Suncoast carrying
(og), 97E212S (rin)andcvs
Agriset, Equinox, FL7655 and
Solar Set
Transition from pink to firm or
soft red stages, over 3 to 8
days of storage
Lycopene contents doubled (Thompson et al.,
2000)
Cv Lemance F1 (Brandt et al., 2006)
NA (Collins et al., 2006)
Cv. Zhenfen Storage in the dark at 14 °C,
95% RH for up to 37 days
Increase from 0.16 to 6.80
mg/kg FW
(Liu et al., 2011)
Cv. Clermon Tomatoes harvested at
light-red to red ripe stages
and stored up to 14 days at
room temperature and
under refrigeration
Higher increases at room
temperature
(Javanmardi &
Kubota, 2006)
Cv. UC-82B
Cv. Money Maker
Mature-green tomato fruit
storage in the dark at 23 °C
and 80% relative humidity
for 16 days
Lycopene accumulation (Alba et al., 2000)
Brief postharvest red-light
treatment of mature-green
fruit
2.3-fold accumulation during
ripening
Red-light treatment of tomato
fruit
Lycopene accumulation
Far-red light treatment Decreased content
Cv Red Ruby tomato fruits harvested at the
breaker stage and stored in
the dark at 12–14 °C
Increased 3.5-fold in
comparison with those at
day 4
(Liu et al., 2009)
Carotenoids Cv. Ailsa Craig or Rapsodie 4 days storage of red-ripe
tomato discs
No change observed (Schofield & Paliyath,
2005)
4 days of incubation in
darkness
Increased content
Treatment by red light or red
light followed by far-red
light
Increased content
(Continued)
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Dynamic changes in health-promoting properties . . .
Table 3–Continued.
Compounds Cultivar name
Harvest conditions and
applied treatments Observed changes References
β-carotene Cvs DRW 3042, Primato, Floriset,
Katinka, Selma, Revido, DRW
3126, Gitana, Ultimo, Relento,
Pankor, Delfino, Tampo, Monica
and Falcato
Different ripening stages Increased content in
proportion to the advanced
ripeness in immature fruit
(Abushita et al., 1997)
Cv. Moneymaker (Giovanelli et al.,
1999)
Cv. Arka Ahuti (Namitha et al.,2011)
Cvs Neris, Svara, Vyt˙
enųDidieji,
Jurgiai and Vaisa F1
(Radzeviˇ
cius et al.,
2009)
Cv Ailsa Craig Increase from the green stage
to the fully ripe stage
(Fraser et al., 1994)
Cvs Pusa Ruby and Vashali hybrid Increase up to the light-pink
stage and declined
afterwards during full and
over ripe stages
(Thiagu et al., 1993)
Cv. Ventura Maximum level in yellow
colored fruit and then
decline
(Biacs et al., 1987)
Cv. Ventura Increase from green to
breaker stage and then
decreased
(Biacs et al., 1987)
Cv. Marmande-Cuarenteno (Cano et al., 2003)
High-pigment cvs Lyco1, Lyco2,
HLY02, HLY13, HLY18 and
Kalvert
Ripening of hp compared to
ordinary tomato cultivars
High accumulation pattern
and levels in hp compared
to ordinary tomato cv
(Hdider et al., 2013)
Cv Red Ruby 21 days of storage and
treatment (untreated, red
light and UV-C treated)
No significant change (Liu et al., 2009)
21 days of storage and
sun-light treatment
Content decreased
significantly
Cvs, cultivars; FW, fresh weight; hp, high-pigment; NA, not available; og, old gold mutation; RH, relative humidity; rin, ripening inhibitor; UV-C, ultraviolet-C.
decreased fast following storage under ambient conditions (Getinet
et al., 2008).
Cultivar-to-cultivar variation was also observed in AsA content
after harvest: although AsA levels of six fresh market tomato vari-
eties exhibited a similar increasing trend during 20 days postharvest
storage under ambient conditions (15.4 to 16.2 °C) and (34.8 to
52.4% RH) the content declined thereafter, the processing culti-
vars maintaining roughly 60% higher contents with respect to the
fresh market var ieties at day 32 after harvest (Tigist et al., 2013)
(Table 2).
Sammi and Masud (2007) studied the effect of ripening and
packaging systems on the postharvest storage and quality of “Rio
Grande” fruits. The authors found that AsA level was significantly
increased along ripening, the highest amounts being attained
between the pink-red and red-ripe stages. A prepackaging
fruit treatment with calcium chloride increased AsA content
compared the untreated packed fruits. Slight AsA accumulation
was observed during storage of hydroponically grown tomatoes
at 7, 15, and 25 °C (Toor & Savage, 2006). Moneruzzaman,
Hossain, Sani, and Saifuddin (2008) detected the highest AsA
content in half-ripe tomato (200.5 mg/kg FW) and the lowest
content in mature-green fruit (85.8 mg/kg FW). It has been
noticed a sharp decrease in AsA content following longer storage
periods. The maximal AsA content (122.3 mg/kg FW) was
recorded in half-ripe tomato fruits following 12 days of storage.
It is widely recognized that high temperature treatments (cook-
ing, boiling, frying, pasteurization, and drying) of fresh and pro-
cessed tomato products lead to extensive AsA loss (Giovanelli,
Zanoni, Lavelli, & Nani, 2002; Sahlin, Savage, & Lister, 2004).
Georg´
e et al. (2011) reported about 80% AsA loss after thermal
processing of red and yellow tomato cultivars. Similarly P´
erez-
Conesa et al. (2009) found that pasteurization of tomato pur´
ee
caused 90% loss of vitamin C. Satisfying AsA retention was, in-
stead, achieved by milder ther mal treatments (Davey et al., 2000).
Carotenoids (lycopene and β-carotene)
One of the main evident changes during tomato ripening is the
sharp increase in the levels of carotenoids resulting in a progres-
sive shift from the green to the orange/red pigmentation of the
fruit. This change of color is the outcome of the de novo synthesis
of lycopene and β-carotene occurring during the chloroplasts-
to-chromoplasts transition and of the concurrent fast degrada-
tion of chlorophylls and thilacoidal pigments (D´
avila-Avi˜
na et al.,
2011; Lenucci et al., 2012). Radzeviˇ
cius et al. (2009) reported
that lycopene concentration significantly increase throughout fruit
ripening of different tomato cultivars (“Neris,” “Svara,” “Vyt˙
enų
didieji,” “Jurgiai,” and “Vaisa F1”). Accordingly, Collins, Perkins-
Veazie, and Roberts (2006) reported a lycopene content 50%
higher in soft red-ripe tomato fruits than in those at the pink
stage of ripening, which was, in turn, 70% higher than that of
light red fruits. Similarly, Namitha, Archana, and Negi (2011) ob-
served a gradual lycopene increase between the green and the 5th
day postbreaker stages, up to 153.3 mg/kg FW in “Arka Ahuti”
tomatoes (Table 3). Arias, Lee, Logendra, and Janes (2000) found
that hydroponically grown on-vine ripened greenhouse tomato
fruits had 32% lower lycopene content than off-ripened fruits.
Fruit harvested before full redness (at either the breaker or tur ning
stages) developed similar or higher lycopene content than those
harvested at the soft red-ripe stage (Collins et al., 2006).
Various researchers focused on high-lycopene tomato cultivars
as these offer higher functional quality and, possibly, longer
shelf-life than ordinary cultivars (Ilahy et al., 2017; Lenucci,
Cadinu, Taurino, Piro, & Dalessandro, 2006). Ilahy et al. (2011),
2018) monitored carotenoid accumulation in the fruits of
C2018 Institute of Food Technologists®Vol.17,2018 rComprehensiveReviews in Food Science and Food Safety 1545

Dynamic changes in health-promoting properties . . .
Table 4–Variation in antioxidant activity during ripening and postharvest of tomato fruit.
In vitro assays Cultivar name
Harvest conditions and
applied treatments Observed changes References
Hydrophilic antioxidant
activity (TEAC +FRAP)
Cv. Rio Grande and
high-lycopene tomato
cvs HLY13, HLY18, and
Lyco2
Different ripening stages Decrease along ripening (Ilahy et al., 2011)
Lipophilic antioxidant activity
(TEAC +FRAP)
Increase along ripening (Ilahy et al., 2011)
Antioxidant activity using in
vitro radical scavenging
assay
Cv. Belissimo Storage of fresh-cut tomato
from 3 different maturity
stages at 5 °C.
Decrease depending on the
initial levels at harvest
(Lana & Tijskens,
2006)
Antioxidant activity using rat
liver microsomes
Antioxidant content and
enzymes
Cv. Rhapsody Storage of mature-green
tomato fruits at 4 °Cforup
to 4 weeks
Decrease along storage (Yahia et al., 2007)
Antioxidant content and
activity
Cv. Micro-Tom Storage at 6 °C of tomato fruit
harvested at breaker and
red-ripe stages for 27 days
Significant decrease in
antioxidant activity,
phenolics, ascorbic acid and
lycopene
(G´
omez et al., 2009)
Antioxidant system Cv. Rhapsody Hot air treatment (38 °C) of
green-mature tomato fruit
and storage for up to
4 weeks
Detrimental effects (Yahia et al., 2007)
Antioxidant activity and
antioxidant system
Exposure of green-mature
fruits to 34 °C for 24 hr and
storage at 4 or 20 °C
Promoted antioxidant system
Cv. Chaoyan-219 Postharvest HVEF
pretreatments prior to
storage of green mature
tomato fruits
Enhanced activity of
antioxidant enzymes and
content of nonenzyme
antioxidant compounds
(phenols, glutathione and
ascorbic acid)
(Zhao et al., 2011)
Total antioxidant activity
using the TEAC assay
Cv.Pannovy Postharvest DC applications
in red-ripe tomato fruits
Substantial increase in total
antioxidant activity with
augmented levels of
phenolics, lycopene and
β-carotene
(Dannehl et al., 2011)
Antioxidant activity Cv. Moneymaker DWP treatments and storage
at 15 °C of red-ripe tomato
fruits
Increased 26% the
antioxidant activity with
higher ascorbic acid and
total phenol levels
(Ahmed et al., 2013)
Cvs, cultivars; DC, direct electric current; DWP, delactosed whey permeate; FRAP, ferric reducing antioxidant power; HVEF, high-voltage electric field; TEAC, trolox equivalent antioxidant capacity.
high-lycopene tomato cultivars and revealed that the amount of
total carotenoids and lycopene notably increased during ripening.
Regardless of ripening stage, the carotenoid concentration was
considerably higher in high-lycopene tomato cultivars (“HLY18,”
“HLY13,” and “Lyco2”) than in the ordinary “Rio Grande”
cultivar. Red-ripe “HLY18” fruits displayed the highest levels
of total carotenoids (278 mg β-Carotene Equivalent/kg FW)
and lycopene (254 mg/kg FW). In “HLY18,” “HLY13,” and
“Lyco2” cultivars, lycopene amount was respectively 2.6-, 2.2-,
and 1.9-fold higher than in the ordinary cultivar “Rio Grande.”
Total carotenoids followed a similar trend as lycopene. This
important discrepancy between ordinary and high-lycopene
tomato cultivars was primarily attributed to their genome
carrying spontaneous high-pigment mutations leading to more
deeply pigmented fruits compared to the ordinary currently
grown tomato cultivars (Armend´
ariz, Macua, Lahoz, Gamica, &
Bozal, 2006; Mustilli, Fenzi, Ciliento, Alfano, & Bowler, 1999).
Lycopene content showed considerable cultivar-dependent
variability (Sahlin et al., 2004; Tigist et al., 2013) and increased
following prolonged storage periods. Inherent genetic variation
across genotypes underlies this variation in carotenoid contents
(Tigist et al., 2013). In a survey on different tomato genotypes,
lycopene concentration at the green stage ranged from 2.5 mg/kg
FW in the fruits of the “Vaisa” hybrid to 14.2 mg/kg FW in those
of the cultivar “Svara,” while the highest content (125.1 mg/kg
FW) was observed in the fully ripe fruits of the cultivar “Neris”
(Radzeviˇ
cius et al., 2009; Table 3).
Carotenoid levels are also affected by storage conditions
(Table 3). Toor and Savage (2006) pointed out that tomato fruits
stored at 15 and 25 °C exhibited visually deep red color compared
to those kept at 7 °C, due to the accumulation of up to 1.8-fold
higher lycopene. In another study conducted on two different
medium-sized tomato cultivars from hydroponic (“Pyramid”)
and nonhydroponic production bought from a local supermarket,
Ajlouni, Kremer, and Masih (2001) noted an increase in lycopene
levels during storage at 22 °C for 14 days, from an initial level
of 36 mg/kg FW in both cultivars to 90 and 115 mg/kg FW for
hydroponic- and nonhydroponic fruits, respectively. Similarly, Pila
et al. (2010) studied lycopene accumulation patterns in partially
ripened, orange-yellow and uniformly sized “Himsona” tomato
fruit freshly grown under open field conditions in Gujarat, India,
throughout 10 days storage at ambient conditions, and revealed
progressive increases during the experimental period, r ipe fruit
reaching values of up to 33.1 mg/kg FW.
Besides storage conditions, maturity stage at harvest is an
influential factor on postharvest lycopene levels. Tomato fruits
harvested at the breaker ripening stage attained a peak in lycopene
after 6 days storage at room temperature (Thompson et al., 2000).
Lycopene content doubled between the pink and the firm or soft
red stages of ripening following a 3 to 8 days storage, depending
1546 ComprehensiveReviews in Food Science and Food Safety rVol. 17, 2018 C2018 Institute of Food Technologists®

Dynamic changes in health-promoting properties . . .
Table 5–Variation in shelf-life and physical attributes in harvested tomato fruit.
Traits Cultivar name
Harvest conditions and applied
treatments Observed changes References
Shelf-life
Cv. DRK 453 Combination of different pressure and
temperature during the shelf-life of
early-breaker stage tomato fruits
Hyperbaric treatment at 20 °C
extended tomato shelf-life
during short treatment duration
(Liplap et al.,
2013)
Beefsteak cv. Grando F1 AVG (1 g/L) application on postharvest
storage and shelf-life in tomato fruits
harvested at the breaker stage
Vacuum pressure of -30KPa
reduced ethylene production
rate, lycopene content, a∗and c∗
color indices and fruit firmness.
Extended shelf-life and storage
up to 20 days at 12 °C
(Candir et al.,
2017)
Cv. Dotaerang Blue light (440 to 450 nm) treatment of
mature green fruit for 7 days
Extended shelf-life (Dhakal & Baeck,
2014)
Cv. Zhenzhu Ultraviolet irradiation at a dose of 4.2
Kj/m²of tomato fruits harvested at the
green mature stage
Extended (up to 35 days)
shelf-life at 18 °C
(Bu, Yu,
Aisikaern, &
Ying, 2013)
Cv. Climberley Pulsed light application at fluence of
2.68 and 5.36 j/cm²on whole red-ripe
tomato fruits
Reduced microbial load during
storage
(Aguil´
o-Aguayo
et al., 2013)
Physiological
loss in
weight
(PWL)
Cv. Grandela Mineral oil coating and carnauba wax
treatment of fruit picked at the breaker
stage and storage at 10 °C for 28 days
Decreased PWL in comparison
with control fruit
(D´
avila-Avi˜
na
et al., 2011)
Mineral oil coating and carnauba wax
treatment of fruit picked at the pink
stage and storage at 10 °C for 28 days
Decreased PWL in comparison
with control fruit
Exposure of fruit to 20 °C for 2 days after
cold storage
Decreased PWL in comparison
with control fruit
Cvs Akoma, Pectomech
and Power
Storage of red-ripe fruits for 7, 14 and 21
days at 10 °C then transfer to ambient
condition (20.49 °C and 54.05% RH)
Increased PWL.
Similar levels among varieties
(Kumah et al.,
2011)
Cv Money Maker Storage of green mature tomato fruits PLW ranging from 9- 11% (Ali et al., 2010)
Storage of green mature tomato fruits
coated with gum Arabic
Decreased PWL in comparison
with control fruit
Cv 508 Storage (20 days at 12 and 22 °C) of
tomato fruit harvested at pink to light
red stage
Increased PLW along storage.
Higher PLW at 22 °Cthanat12
°C
(Assi et al.,
2009)
Cvs Marglobe and Roma
VF
Storage of fruit from different cultivars
picked at different maturity stages
PLW levels highly dependent on
all 3 factors considered
(Getinet et al.,
2008)
Cv. Clermon Storage at ambient conditions of light-red
and red-ripe tomato fruits from both
open pollinated varieties and hybrids
Linear decrease along storage (Javanmardi &
Kubota, 2006)
9402x Azad Type-3,
8731 x Azad Type-3
andAzadType-1
(Kumar et al.,
2007)
NA Storage at ambient condition of tomato
fruits at different maturity stages
Higher PWL when harvested at
breaker or turning stages
(Collins et al.,
2006)
Early pear-type and Cv.
“S-12”
Tomato fruits harvested at red ripe stage
and stored 7 days at room temperature
55 and 33% PWL respectively (Kaur et al.,
1977)
Tomato fruits harvested at breaker stage
and stored 7 days at room temperature
Minimum PWL of 23 and 46%
respectively
Cv. Kuber Storage during 12 days of tomatoes
harvested at turning stage and at red
ripe stage
Minimum PWL when compared to
those harvested at the red-ripe
stage
(Gaur & Bajpai,
1982)
Cv. Roma VF Storage of mature green tomato fruits Increase during storage (up to
13.31% at 12th day)
(Moneruzzaman
et al., 2008)
Storage of full-ripen tomato fruits Total PWL was lowest during
storage, being 5.72% at 3rd day
and 11.96% at 12th day of
storage
Cv. Rio Grande Storage of tomato fruits from different
ripening stages in different packaging
systems (systems (CaCl2-treated and
nontreated)
Increase with advancing ripening.
Reduction (50% less) in packed
fruit
(Sammi &
Masud, 2007)
Cv. Roma VF Storage of tomato fruits during 6 days
under ambient condition.
7.7 to 9.7% (Mallik et al.,
1996)
Cv. Clermon Light-red and red-ripe tomato fruits
stored at room and cold temperatures
(5 and 12 °C)
Increased PWL along storage
irrespective of temperature.
Reduced PWL in cold-stored fruit
(Javanmardi &
Kubota, 2006)
Cv. Himsona Storage of partially ripe orange-yellow
tomato fruits at 34 ±1°for 10 days
Progressive increase till full
ripeness
(Pila et al.,
2010)
(Continued)
C2018 Institute of Food Technologists®Vol.17,2018 rComprehensiveReviews in Food Science and Food Safety 1547

Dynamic changes in health-promoting properties . . .
Table 5–Continued.
Traits Cultivar name
Harvest conditions and applied
treatments Observed changes References
Treatment of tomato fruits with
chemicals (GA3, CaCl2, and salicylic
acid) then storage at 34±1°for 10 days
Lesser weight loss in relation to
controls
Cv. Josefina modified atmosphere packaging (5% O2
and 5% CO2)
Extend the shelf-life of the cherry
tomato cv Josefina up to 25
days
(Fagundes et al.,
2015)
Fruit firmness Cv. Belissimo Storage for up to 32 days of tomato fruits
at different ripening stages
Decreased firmness (Lana et al.,
2005)
Cv. “870” Storage of light-red tomato fruits Decreased firmness (Mizrach, 2007)
Cvs Roma VF,
Melkasalsa,
Melkashola, Metadel,
Eshete, Marglobe
Improved, Fetane,
Heinz-1350 and
Bishola
Storage of mature-green tomato fruits
under ambient conditions
Decreased firmness (Tigist et al.,
2013)
Cvs Akoma, Pectomech
and Power
Full ripe fruits stored for7, 14 and 21 days
at 10 °C and then transferred to
ambient condition (20.49 °C and 54.05
%RH)
Decreased firmness (Kumah et al.,
2011)
Cv. Money Maker Gum Arabic coating of green-mature
tomato fruits stored at 20 °Cand
80–90% RH for 20 days.
Decreased firmness down to 10N (Ali et al., 2010)
Cv. Grandela Storage (28 days at 10 °C) of tomato
fruits cv. “Grandela,” harvested at
breaker and pink color stage
Initially, similar firmness (15-16
N) in breaker and pink samples
decreasing with storage
(D´
avila-Avi˜
na
et al., 2011)
Coating with mineral oil or carnauba, and
then storage
(28 days at 10 °C) of “Grandela” fruit
harvested at breaker and pink color
stage
Firmness of breaker fruit was
7.73, 5.43 and 7.03 N for
control, mineral oil-coated and
carnauba-coated samples,
respectively
Firmness of pink fruit was 6.5,
8.08 and 8.13 N for the control,
mineral oil-coated and
carnauba-coated samples,
respectively
Cv. 508 Storage of pink to light-red tomato fruits
at 12 and 22 °C for 10 days
Rapid decline in firmness. Firmer
fruit at 12 °Cthanat22°C
(Assi et al.,
2009)
Cv. Zhenfen UV-B irradiation and storage during of
mature-green fruits 37 days of
Decreased firmness (26.68 to
8.59 N)
(Liu et al., 2011)
Cv Red Ruby 21 days of storage of mature-green
treated daily with short bursts of UV-C,
redlight or sun light.
Gradual decrease in firmness
No effects of red light treatment.
Significant decrease in UV-C-
and sun light- treated tomatoes
(Liu et al., 2009)
a∗,b
∗,c
∗, color indexes; AVG, aminoethoxyvinylglycine; GA3, gibberellic acid; PLW, physiological loss in weight; RH, relative humidity.
on the considered genotype (Brandt, P´
ek, Barna, Lugasi, &
Helyes, 2006; Collins et al., 2006; Thompson et al., 2000). In
accordance with these reports, lycopene content in mature green
tomatoes increased from 1.6 to 68.0 mg/kg FW during a 37-day
storage period (Liu et al., 2011). The lycopene contents of
hydroponically grown tomatoes harvested at light-red to red-ripe
stages increased significantly during storage for up to 14 days, and
were higher in fruits kept at room temperature with respect to
those refrigerated (Javanmardi & Kubota, 2006). Light conditions
have also been found to strongly impact lycopene biosynthesis
and accumulation during off-vine ripening. Dark-stored tomato
fruits showed a notable increase of lycopene content throughout
16 days. Alba, Cordonnier-Pratt, and Pratt (2000) showed that
lycopene biosynthesis was stimulated in mature-green harvested
tomato fruits following a brief red-light treatment (2.3-fold
higher) during fruit r ipening. A far-red light treatment of reversed
the observed light-induced accumulation of lycopene, suggesting
the regulation by fruit-localized phytochromes. When lycopene
was assayed in “Red Ruby” tomato fruits harvested at the breaker
stage and stored at 12 to 14 °C in the dark, a 3.5-fold increase
(85 mg/g Dry Weight) was observed after 15 days storage (Liu,
Zabaras, Bennett, Aguas, & Woonton, 2009). Carotenoid content
in tomato discs remained unchanged until the fourth day of
storage, and started to increase afterwards following 4 days of
darkness incubation or the exposure to either red-light or red-light
followed by far-red light treatment (Schofield & Paliyath, 2005).
Abushita et al. (1997) and Giovanelli et al. (1999) found a
simultaneous increase of β-carotene and lycopene concentra-
tion during tomato fruit ripening (Table 3). Namitha et al.
(2011) reported a gradual increase in β-carotene content (4.6 to
103.7 mg/kg FW) between the mature-green and 10 days post-
breaker stages of ripening. It has been reported that β-carotene
increases linearly from the green (3.3 mg/kg FW) stage to the
full-ripe stage (36.8 mg/kg at 3 weeks postbreaker; Fraser, Trues-
dale, Bird, Schuch, & Bramley, 1994). Thiagu, Chand, and Ra-
mana (1993) showed that the level of β-carotene continuously
increased till the pink stage of ripening and sharply declined after-
wards. Radzeviˇ
cius et al. (2009) showed that β-carotene contents
in tomato fruits increased during ripening. A nonsignificant de-
crease in β-carotene level after full ripeness stage was noted only
in “Svara” tomato fruit. A limited increase, between not fully
ripened and fully ripened stages, was noted in “Vaisa” F1 fruits.
Biacs, Daood, Czinkotai, Hajd´
u, and Kiss-Kutz (1987) reported a
peak of β-carotene in yellow-colored fruits of the processing cul-
tivar “Ventura,” followed by a reduction in the following ripening
stages. Biacs et al. (1987) and Cano et al. (2003) highlighted that
1548 ComprehensiveReviews in Food Science and Food Safety rVol. 17, 2018 C2018 Institute of Food Technologists®

Dynamic changes in health-promoting properties . . .
Table 6–Variation in eating quality-related attributes during ripening and postharvest of tomato fruit.
Traits Cultivar name
Harvest conditions and applied
treatments Observed changes References
Total soluble
solids (TSS)
Cv. Roma VF Different ripening stages Highest (6.82%) and lowest
(5.85%) TSS values in fully
ripe and mature green fruit,
respectively
(Moneruzzaman et al.,
2008)
NA No significant changes (Collins et al., 2006)
Cvs Roma VF, Melkasalsa,
Melkashola, Metadel, Eshete,
Marglobe Improved, Fetane,
Heinz-1350 and Bishola
Fresh market mature-green
tomato varieties stored for
20 days at ambient conditions
Highest TSS contents at day 16 (Tigist et al., 2013)
Processing tomato varieties
stored for 20 days at ambient
conditions
Highest TSS contents at day 20
Cv. Money Maker Storage of gum Arabic-coated
mature-green tomato fruits
Increasing trend of TSS during
storage with treated fruit
characterized by lower TSS at
final day of storage
(Ali et al., 2010)
Cv. Kuber 8 days storage of tomatoes
harvested at the turning and
pink stages
5.2% and 5.9% increase in fruit
harvested at turning and pink
stages, respectively
(Gaur & Bajpai, 1982)
8 days storage of red-ripe tomato
fruit
Substantial decline (6.6 to 4.3)
Cvs Akoma, Pectomech and
Power
Storage of tomato fruits
harvested at mature green
stage for 7, 14 and 21 days at
10 °C then transferred to
ambient condition (20.49 °C
and 54.05 % RH)
Increase up to 8 days of storage
depending on the cultivars
and storage temperature
(Kumah et al., 2011)
Cv. Malike Storage of tomato fruits at the
middle-red ripe stage for
21 days at 10 °C
Slight increase (Znidarcic and Pozrl,
2006)
9402x Azad Type-3, 8731 x Azad
Type-3 and Azad Type-1
Storage of different red-ripe
tomato fruits from open
pollinated and hybrids cvs for 9
days at ambient temperatures
Poststorage increase in all
genotypes
(Kumar et al., 2007)
Cvs Marglobe and Roma VF Storage for 32 days of Marglobe
and Roma tomato fruits
harvested at mature-green,
turning, and light red maturity
stages
Increase with significant
interaction between cultivars
and maturity stages
(Getinet et al., 2008)
Storage of tomato fruit harvested
at different ripening stages at
different temperatures
Increase with color and
maturity, depending on the
stage of ripeness at harvest
and storage temperature
(Trejo & Cantwell,
1996)
Cv. Malike (Znidarcic and Pozrl,
2006
Cv. Sunny (Atta-Aly et al., 2000)
Cv. Grandela Storage at 10 °C of breaker
tomato fruits
Minor changes (D´
avila-Avi˜
na et al.,
2011)
Storage at 10 °C of pink tomato
fruits
15% to 20% decrease with
respect to the initial TSS
Cv. Himsona Storage at 34 ±1°C for 10 days Increasing trend (Pila et al., 2010)
GA, CaCl2, and salicylic acid
treatments prior to storage for
10 days at 34 ±1°C
Lower TSS in treated samples
Cv 508 Pink or light red tomatoes held at
12 and 22 °C
Significant increase over time (Assi et al., 2009)
Cv. Rio Grande Packaging of ripening fruit at
ambient conditions
Increase along ripening.Highest
TSS in packed fruit up to pink
red stage
(Sammi & Masud,
2007)
Cv. Clermon Tomatoes harvested at light red
to red ripe stage and stored at
different temperatures during
14 days
No variation (Javanmardi &
Kubota, 2006)
Cv Red Ruby Breaker tomato fruits storage at
12 to 14 °Cduring21days
No significant changes (Liu et al., 2009)
Cv. Clarion 14 days room temperature stored
mature green and full ripe
tomato fruits
No significant changes (Wills & Ku, 2002)
Cv. BR124 10 days of storage at 12 °C No significant changes in TSS (Kagan-Zur & Mizrahi,
1993)
Different high-pigment tomato
cvs
Processing of high-pigment
tomato cultivars into pur´
ee
Important TSS loss in all cases (Siddiqui and Singh,
2015)
Ordinary tomato cvs Processing of ordinary tomato
cultivars into pur´
ee
Important TSS loss in all cases
Acidity
Cv Ailsa Craig Fruit ripening of standard
cultivars
Decrease in some organic acids (Chen et al., 2001)
(Continued)
C2018 Institute of Food Technologists®Vol.17,2018 rComprehensiveReviews in Food Science and Food Safety 1549

Dynamic changes in health-promoting properties . . .
Table 6–Continued.
Traits Cultivar name
Harvest conditions and applied
treatments Observed changes References
Cv Santa Clara (Castro et al., 2005)
9402x Azad Type-3, 8731 x Azad
Type-3 and Azad Type-1
(Kumar et al., 2007)
Cvs Marglobe and Roma VF (Getinet et al., 2008)
Cv. Himsona (Pila et al., 2010)
Cv. Micro-Tom Ripening “Micro-Tom” fruit stored
at different temperature
Significant decreases in tartaric,
malic, ascorbic, and citric acids
Slow increase in succinic acid
Low acid content in immature
fruit, with the highest levels
at the breaker stage and a
rapid decrease thereafter
(G´
omez et al., 2009)
Cv. Roma VF Ripening conditions (different
temperature and relative
humidity)
Differences in ascorbic acid
content
(Moneruzzaman et al.,
2008).
Cv. Ohio 7814 Ripening Highest organic acid content at
the pink stage, and then a
decline with ripening.
Highest citric than malic acid
concentration throughout
fruit development
Very small amounts of oxalic
acid, with similar change
patterns as those for citric and
malic acids
(Knee & Finger, 1992)
Cv. Momotaro (Islam et al., 1996)
Cv. Roma VF Ripening TA content peaks at the pink
stage
(Moneruzzaman et al.,
2008)
Cv. Momotaro Storage at 15 °C Linearly decreased
concentration with increasing
temperature
(Islam et al., 1996)
Cv. Grandela Coating with mineral oil and
carnauba wax tomato during
ripening
Decrease with maturity
irrespective of coating
treatments
(D´
avila-Avi˜
na et al.,
2011)
Cvs Marglobe and Roma VF Storage for 32 days of Marglobe
and Roma tomato fruits
harvested at mature-green,
turning, and light red maturity
stages
Declining trend
(cultivar-specific)
(Getinet et al., 2008)
9402x Azad Type-3, 8731 x Azad
Type-3 and Azad Type-1
Storage of different open
pollinated and hybrids varieties
cvs and hybrids for 9 days at
ambient temperatures
Decline during storage (Kumar et al., 2007)
Cv. Money Maker Storage of coated and uncoated
tomato fruits
Declining trend irrespective of
treatments
(Ali et al., 2010)
Cv. Pronto Short term storage (4 days after
harvest: air temperature 20 °C,
relative air humidity 55%, air
velocity B 0.1 ms−1)
Increased by 22% (Auerswald et al.,
1999)
Cv. Rio Grande Freshly harvested mature green
tomatoes packed in
polyethylene packaging with or
without treating with calcium
chloride, boric acid and
potassium permanganate
within the different stages of
ripening.
Decrease during ripening with
faster rate in packed fruits
(Sammi & Masud,
2007)
Cv. Tradiro Storage under refrigeration, at 15
and 25 °C
Significantly higher TA at 15
and 25 °C than that of
refrigerated tomatoes
(Toor & Savage, 2006)
Cv. Red Spring Bottled tomato pulp stored at
room temperature (20.0 ±1.8
°C) for 0 to 180 days
Significant decreases in malic
(51%) and citric acid (71%)
(Ord´
o˜
nez-Santos et al.,
2009)
Tomato paste stored at different
temperatures
Gradual increase in acidity (Gould, 1992)
Total sugars
Sugar
content
Roma VF, Melkasalsa,
Melkashola, Metadel, Eshete,
Marglobe Improved, Fetane,
Heinz-1350 and Bishola)
Ripening
Different ripening stages
Increase from the green mature
to the red ripe stage
(Tadesse et al., 2012)
Total sugars Cvs. Sunny and Solar Set Initial increment followed by no
further changes or a slight
decrease
(Baldwin et al., 1991)
Sugars
content
Cv. Moneymaker Content depending on harvest
maturity
(Sinaga, 1986)
(Continued)
1550 ComprehensiveReviews in Food Science and Food Safety rVol. 17, 2018 C2018 Institute of Food Technologists®

Dynamic changes in health-promoting properties . . .
Table 6–Continued.
Traits Cultivar name
Harvest conditions and applied
treatments Observed changes References
Reducing
sugars and
soluble
sugars
Cvs Moscow and Fireball Reducing sugars increased
during ripening and storage
(Dalal et al., 1965)
Sugars Cv. Rio Grande Peak at green to turning stage
and then decrease as ripening
proceeds
(Sammi & Masud,
2007)
Reducing
sugars
Sucrose
Cv. Momotaro More rapid increase at later
than at earlier stages of
ripening
Highest sucrose concentration
in immature and mature
green fruit and then decline
with maturation
(Islam et al., 1996)
Glucose and
fructose
Cv. Micro-Tom Storage at 20 °C of tomato fruits
at different maturity stages
Increased accumulation was
lowered under chilling storage
(G´
omez et al., 2009)
Reducing
sugar
content
9402 x Azad Type-3, 8731 x
Azad Type-3 and Azad Type-1
Storage of tomato fruits at
different ripening stages from
for 9 days at ambient
temperatures
Increase gradually during
storage
(Kumar et al., 2007)
Total sugars Cv. Marglobe Mature green fruit stored at 14 to
19 °C for 28 days
Increased levels up to 8 days of
storage and decreased
afterwards
(Melkamu et al.,
2008)
Reducing
sugars
Cv. Pronto Short term storage of ripe tomato
fruits (4 days after harvest: air
temperature 20 °C, relative air
humidity 55%, air velocity B 0.1
ms−1)
No important change up to
7 days
(Auerswald et al.,
1999)
Total sugars Cv. Roma VF Tomato fruits harvested at
Mature green, half ripen and
full ripen were kept under 3
different conditions; open
condition (control), covering
with white polythene and finally
treatment by CaC2+polythene.
Highest total sugars in packed
fruit at the end of the storage
Increase with ripening
irrespective of maturity
condition
(Moneruzzaman et al.,
2008)
Aroma
Cv. Vanessa Off-vine ripening of tomato Hexanal content increased and
correlated positively with
sweetness
(Krumbein et al.,
2004)
Ordinary fresh market tomatoes
harvested at mature green
Cherry tomato cvs
Roma type tomato fruits,
Cv. Early Girl
Ripening Phenyl acetaldehyde and
3-methylbutanal formed
during ripening
Contribution of furaneol to the
flavour of fresh tomato
(Buttery et al., 2001)
Volatile monoterpenes present
in minute quantity
(Galliard et al., 1977)
NA Ripening of vine vs off-vine
tomato
Higher content of
benzaldehyde, citronellyl
propionate, citronellyl
butryrate, decanal,
dodecanal, geranyl acetate,
geranyl butyrate, nonanal,
and neral in vine-ripened
tomato as compared to
artificially ripened tomatoes
Higher content of butanol,
2,3-butanedione,
isopentanal, isopentyl
acetate, 2-methyl-3-hexanol,
3-pentanol, and propyl
acetate in artificially ripened
tomatoes, as compared to
field-ripened tomato
(Madhavi & Salunkhe,
1998)
Cvs, cultivars; GA3, gibberellic acid; NA, not available; TA, total acidity; TSS, total soluble solids.
β-carotene level increased from the green to the breaker stages
up to 4.9 mmol/kg FW and decreased afterward to 3.4 mmol/kg
FW.
Hdider, Ilahy, Tlili, Lenucci, and Dalessandro (2013) assessed
six high-pigment tomato cultivars (“Lyco1,” “Lyco2,” “HLY02,”
“HLY13,” “HLY18,” and “Kalvert”) in comparison to the or-
dinary “Donald” variety. These authors reported that β-carotene
and lycopene contents showed similar variation trends. At the red
ripe stage, “HLY13” and “HLY18” tomatoes exhibited the high-
est level of β-carotene (19.8 and 19.3 mg/kg FW, respectively)
indicating that, in these varieties, high lycopene amounts were
associated with high β-carotene levels. Such contrasting differ-
ences between high-lycopene and ordinary tomato cultivars were
ascribed to genotypic differences and growing conditions (Dumas
et al., 2003; Ilahy et al., 2016, 2017, 2018). High-pigment tomato
cultivars carry spontaneous mutations leading to exaggerated light-
responsiveness and deeply red-pigment mature fruits compared to
ordinary cultivars (Atanassova, Stoeva-Popova, & Balacheva, 2007;
C2018 Institute of Food Technologists®Vol.17,2018 rComprehensiveReviews in Food Science and Food Safety 1551

Dynamic changes in health-promoting properties . . .
Figure 1–Major changes during ripening of tomato fruits. hp, high pigment; ip, intense pigment; ogc, old gold crimson; +, positive effect; –, negative
effect.
Mustilli et al., 1999). For all the studied cultivars, β-carotene
levels were lowest at the green stage, increasing afterwards till the
red-ripe stage. This increase was 3.7-fold in “Donald” tomato
fruits, whereas in high-lycopene tomato cultivars (with the excep-
tion of “HLY02”) it was between 3.7- to 7.1-fold higher than the
ordinary tomato cultivar (Hdider et al., 2013). The β-carotene
contents remained almost unchanged (average of 12 μg/g D.W.)
in nontreated, red-light-treated and UV-C-treated tomato fruits
throughout 21 days of treatment and storage, in contrast to the
observations for sun light-treated fruits (Liu et al., 2009; Table 3).
Changes in antioxidant activity
Different analytical assays have been developed to measure an-
tioxidant capacity, none of which reflects accurately all ROS
sources or all antioxidant systems existing in plants (Prior, Wu,
& Schaich, 2005). Radical scavenging activity based methods are
mostly used, even though results may not always be transposable
to the in vivo situation. The lack of a standardized method may
also lead to inconsistent results, and thus hinder the interpretation
of published data. Even so, very few reports focused on posthar-
vest dynamic changes affecting antioxidant activity in fresh tomato
fruits, although some reports exist on changes during on-vine
ripening (Cano et al., 2003; Jim´
enez et al., 2002).
Ilahy et al. (2011, 2018) monitored the hydrophilic (HAA) and
lipophilic (LAA) antioxidant activity using the Trolox equivalent
antioxidant capacity (TEAC) and the ferric reducing antioxidant
power (FRAP) assays in ordinary and high-pigment tomato culti-
vars during the ripening process (Table 4). Regardless of the an-
alytical method and cultivars, the highest and lowest HAA values
were found in the green-mature and red-ripe fruits, respectively.
Although HAA significantly dropped throughout ripening, a si-
multaneous increase of LAA was observed in all assessed tomato
cultivars. LAA increase was between 50% and 91% using TEAC
and the FRAP assays, respectively. Although the HAA values de-
creased and LAA increased during tomato fruits ripening in all
cultivars under analysis, at the red-ripe stage, values were 28%,
61%, 110% and 66%, 59%, 124%, respectively, higher in the high-
pigment tomato cultivars “Lyco2,” “HLY13,” and “HLY18” com-
pared to Rio Grande. All of the above reported data highlight the
higher antioxidant profile of high-pigment cultivars, which suit the
ever-increasing consumer demand for nutritive and healthy foods.
Lana and Tijskens (2006) focused on the changes affecting the
antioxidant activity of fresh-cut tomato dur ing 5 °C postharvest
storage. Fruits were harvest at three different maturity stages, and
two methods were used for antioxidant activity determination,
one of them being an in vitro radical scavenging assay, while the
second used rat liver microsomes to mimic an in vivo system.
Although antioxidant activity generally decreased along storage,
the major factor determining this property was apparently the
initial levels at harvest. This observation highlights the need
to harvest the fruits at an adequate maturity stage in order to
optimize the levels of the health-promoting property. Storage of
“Rhapsody” tomato fruit at 4 °C was also reported to decrease the
content of antioxidant molecules (Yahia, Soto-Zamora, Brecht,
1552 ComprehensiveReviews in Food Science and Food Safety rVol. 17, 2018 C2018 Institute of Food Technologists®

Dynamic changes in health-promoting properties . . .
Figure 2–Major changes during postharvest storage of tomato fruits. ogc, old gold crimson; +, positive effect; –, negative effect; AVG,
aminoethoxyvinylglycine; DC, direct electric current; DWP, delactosed whey permeate; GA, gibberellic acid; GA3, gibberellic acid; GMO, genetically
modified organisms; hp, high pigment; HVEF, high-voltage electric field; ip, intense pigment; UV, ultraviolet; UVB, ultraviolet B; UVC, ultraviolet C.
K., & Gardea, 2007). Similarly, significant decreases in phenolics,
AsA and lycopene occurred in “Micro-Tom” fruits after 27
days of storage at 6 °C, although glutathione content increased
and the antioxidant capacity, determined by the 2,2-diphenyl-
1-picrylhydrazyl analytical method, resulted unchanged (G´
omez
et al., 2009).
Some reports suggest that specific postharvest treatments may
partially reduce the detrimental effects of cold storage on the
antioxidant properties of tomato fruits if conditions are carefully
optimized (Table 4). For instance, when “Rhapsody” fruits are
submitted to hot-air pre-treatments to improve storability and
decrease the incidence of chilling injury, detrimental effects on
antioxidant activity were found at 38 °C, while 34 °C exposure
was found to promote the antioxidant capacity of tomatoes (Yahia,
Soto-Zamora, Brecht, & Gardea, 2007). Postharvest pre-treatment
of “Chaoyan-219” tomato fruits by high-voltage electrostatic
field (HVEF), enhanced the antioxidative enzymatic system as
well as the levels of nonenzymatic antioxidant compounds like
phenols, glutathione and AsA (Zhao et al., 2011). Postharvest
direct-electric-current applications in “Pannovy” fruits also
increased substantially the total antioxidant activity measured by
TEAC, with concomitantly augmented phenolics, lycopene and
β-carotene contents (Dannehl et al., 2011). DWP treatments
increased antioxidant activity of “Moneymaker” tomatoes by 26%
at the end of storage at 15 °C, parallel to higher AsA and total
phenols levels (Ahmed et al., 2013).
Changes in Physical Attributes
Shelf-life potential
Liplap et al. (2013) studied the impact of the combination of
different pressure levels and temperatures on tomato fruit shelf-life
(Table 5) finding that a hyperbaric treatment at 20 °C was able
C2018 Institute of Food Technologists®Vol.17,2018 rComprehensiveReviews in Food Science and Food Safety 1553

Dynamic changes in health-promoting properties . . .
to significantly prolong storage time without adverse effects on
eating quality. Similarly, Candir, Candir, and Sen (2017) reported
that postharvest shelf-life of Beefsteak “Grando F1” tomato fruits
was extended by a treatment with 1 g/L aminoethoxyvinylglycine
(AVG) at a vacuum pressure of −30 KPa. Generally, AVG-treated
fruits exhibited lower ethylene production, decreased lycopene
biosynthesis, altered color changes and increased firmness than
nontreated ones.
Dhakal and Baeck (2014) reported that short time (1 week)
irradiation of mature-green tomato fruits with light emitting
diode-generated blue-light (440 to 450 nm) is a practical approach
to delay fruit ripening and softening thus extending shelf-life.
Similarly, UV irradiation (4.2 Kj/m²) prolonged the shelf-life
of green-mature harvested “Zhenzhu” tomato fruits throughout
5 weeks at 18 °C (Bu, Yu, Aisikaer, and Ying, 2013). In the same
context, pulsed light (2.68 and 5.36 j/cm²) was proposed as an
efficient nonthermal food grade technology to reduce microbial
charge of fresh tomatoes during postharvest storage, with no
adverse effects on the the nutritional value of the product (Aguil´
o-
Aguayo, Florence-Charles, Renard, Page, & Carlin, 2013).
Physiological loss in weight
The physiological loss in weight (PLW) is among the main
changes affecting postharvest storage of fresh product. The shelf
life threshold of fresh fruits and vegetables is attained at about
10% PLW (Acedo, 1997; Pal, Roy, and Srivastava, 1997). Storage
duration, temperature and genotype significantly affect PLW
(Javanmardi & Kubota, 2006). PLW may also be attributed to
changes in the levels of soluble sugars since monosaccharides are
used as substrates for respiratory purposes throughout storage
(Singh & Reddy, 2006). Several reports have been published on
PLW in tomato during postharvest storage (Table 5).
D´
avila-Avi˜
na et al. (2011) revealed that PLW of untreated, min-
eral oil-coated and carnauba wax-coated tomatoes treated at the
breaker stage reached values of 3.19%, 1.60%, and 2.20% after
1 month storage at 10 °C, respectively, whereas PLW of fruit sub-
mitted to the same treatments at the pink stage was 3.76%, 1.67%,
and 2.53%, in the same order. After exposure of tomato fruits
to 20 °C for 2 days, untreated, carnauba wax-coated and min-
eral oil-coated fruit, lost 5.82%, 3.15%, and 3.30% of their initial
weight, respectively. Kumah, Olympio, and Tayviah (2011) re-
ported increasing PLW of tomato fruits dur ing storage under vari-
able temperatures. Nevertheless, no significant changes in PLW
were noted among different varieties. Ali, Maqbool, Ramachan-
dran, and Alderson (2010) reported 9% to 11% PLW in gum
arabic-coated tomatoes during storage, lower than those of un-
treated samples. When “508” tomato cultivar harvested at pink to
light-red ripening stages were stored at 12 and 22 °Cduring20
days, PLW increased with subsequent storage, with higher values
at 22 °Cthanat12°C (Assi, Jabarin, & Al-Debei, 2009).
Getinet et al. (2008) investigated cultivar-, maturity- and stor-
age condition-related effects on PLW of tomatoes. The light-red
fruits of cultivar “Marglobe” exhibited the most important PLW
when stored under room temperature. Green-mature “Roma
VF” tomato fruits showed the lowest PLW when stored in an
evaporative cooler. Javanmardi and Kubota (2006) and Kumar,
Singh, Singh, Singh, and Prasad (2007) reported that the av-
erage fruit weight of different varieties exhibited a significantly
linear decrease with increasing storage duration at ambient con-
ditions. Collins et al. (2006) focused on the effect of ripening
stage on PLW throughout storage of tomato fruits at room tem-
perature. Generally, breaker or turning tomato fruits displayed
higher PLW. Red-ripe early pear-type and “S-12” tomato fruits
showed 55% and 33% PLW, respectively, after a week of stor-
age at ambient conditions. In contrast, 23% and 46%, PLW, in
the same order, were observed when fruit were harvested at the
breaker stage (Kaur, Kanwar, & Nandpuri, 1977) (Table 5). Min-
imal PLW were reported after 12 days of storage for turning with
respect to red-ripe tomato fruits (Gaur & Bajpai, 1982). Toma-
toes stored at room temperature showed higher PLW as compared
to those packed in polyethylene bags due to higher transpiration
and water loss rates (Lingaiah, 1982). Total PLW in mature-green
tomato fruits throughout storage was reported to be increased from
6.28% to 13.31% between the 3rd and the 12th days of storage
(Moneruzzaman, Hossain, Sani, Saifuddin, & Alenazi, 2009). In
fully ripe tomato fruits, PLW was the lowest with 5.72% after 3 days
and 11.96% after 12 days of storage. Sammi and Masud (2007) ob-
served that PLW in tomato fruits stored in different packaging sys-
tems increased significantly as the ripening proceeded. Packaging
reduced PLW of fruits by 50% compared to controls at all ripen-
ing stages. Mallik, Bhattacharja, and Bhattacharja (1996) reported
7.7% to 9.7% PLW in fruits of “Roma VF” tomato after 6 days of
storage under ambient conditions. Javanmardi and Kubota (2006)
noted an increase in PLW of hydroponically grown tomato fruits
stored at ambient conditions and under refrigeration (5 and 12 °C)
irrespective of temperature. However, tomatoes held at room tem-
perature showed higher PLW (0.68% per day) as compared to those
kept at 5 °C (0.15% per day) or 12 °C (0.49% per day). Similarly,
Pila et al. (2010) observed that PLW of tomato fruits increased
progressively during their storage, and this progression continued
till the fruit attained full ripeness. Treatment with chemicals such
as gibberellic acid, CaCl2and salicylic acid led to comparatively
lesser PLW in relation to untreated fruit (19.89%) during storage
(Table 5). Active or smart packaging is being increasingly used
in food industries to prolong the shelf-life of different perishable
products. Fagundes et al. (2015) highlighted the efficiency of mod-
ified atmosphere packaging (5% O2and 5% CO2)inextending
the shelf-life of the cherry tomato cultivar “Josefina” until 25 days.
Fruit firmness
Firmness is an important trait governing acceptability and com-
mercial quality evaluation of tomato fruits, and it is altered by mor-
phological and physiological fruit characteristics such as pericarp
firmness, the importance of locule tissue as well as the ripening
stage (Chiesa et al., 1998). Kumah et al. (2011), Lana, Tijskens, and
van Kooten (2005), Mizrach (2007), and Tigist et al. (2013) noted
a loss in tomato firmness throughout storage (Table 5). Firmness
levels and softening rates are cultivar-dependent (Xin et al., 2010),
which could be attributed to differences in metabolic activity
during the ripening process. Firmness loss-related events include
deterioration of the cell structure and intracellular materials, com-
positional changes and disassembly of cell walls (Seymour, Taylor,
& Tucker, 1993), by cell wall-modifying enzyme activities (very
prominently pectinesterase and polygalacturonase; Page, Marty,
Bouchet, Gouble, & Causse, 2008). Loosening of cell wall struc-
ture of fruit epidermis, together with changes in fruit cuticle, result
in softening, higher skin permeability and higher moisture loss, de-
pending upon the genotype. Moisture loss, in turn, contributes to
wilting, shrinkage and firmness loss. Reports on firmness changes
in tomato during postharvest storage are discussed in the next
section.
Kumah et al. (2011) observed that fruit firmness generally
dropped during storage from day 1 till day 7 ir respective of
storage temperature (cold or ambient). Fruit firmness decreased
1554 ComprehensiveReviews in Food Science and Food Safety rVol. 17, 2018 C2018 Institute of Food Technologists®

Dynamic changes in health-promoting properties . . .
significantly during storage as measured in gum-arabic coated and
noncoated tomato fruits (Ali et al., 2010). Tomato fruits kept under
ambient temperature exhibited the lowest firmness values (10 N)
at the end of storage. D´
avila-Avi˜
na et al. (2011) outlined that
firmness of mineral oil and carnauba-coated “Grandela” tomatoes
harvested at breaker and pink color stages showed a decreasing
trend throughout a storage period of 28 days at 10 °C, regardless
of treatment. Tomato fruits at the breaker and pink ripening stages
had initially the same firmness (15 to 16 N) which decreased after-
wards attaining values in the range of 5.4 to 8.1 N. Assi et al. (2009)
studied the storage performance of tomatoes against traditional and
modern handling methods followed in Jordan. Tomato fruits were
stored during 10 days at 12 and 22 °C, displayed a rapid decline
in firmness, although those held at 12 °C remained firmer than
those held at 22 °C after 10 days storage. Sammi and Masud (2007)
evaluated the effect of three different packaging systems and their
efficiency to prolong the storability and upgrade mature-green
fruit quality of the cultivar Rio Grande. The authors observed
that sensory texture scores increased with ripening, but remained
lower in untreated fruits. The firmness of UV-B-irradiated toma-
toes decreased from 26.7 to 8.6 N during storage for 37 days (Liu
et al., 2011). Similarly, firmness of UV-C- and sun-light-treated
tomatoes was significantly lower in comparison with controls after
a storage period of 3 weeks (Liu et al., 2009; Table 5).
Changes in Eating Quality-Related Attributes
Total soluble solids
Total soluble solids (TSS) values are considered one of the most
important ripening-associated qualitative parameters in various
products, including fresh tomatoes (Tehrani, Chandran, Sharif
Hossain, & Nasrulhaq-Boyce, 2011). Changes in TSS content
are mainly related to the hydrolysis of starch into soluble sugars
(sucrose, glucose and fructose) and to the accumulation of organic
acids, thus high TSS values, usually in the range 4.80% to 8.80%,
are a good index of tomato fruit maturity and eating quality
during postharvest storage (Sammi & Masud, 2007). Different
studies reported a gradual increase of TSS throughout storage
(Table 6). Moneruzzaman et al. (2008) found significant variation
in TSS values of tomato juice according to the maturity stage
of fruits, with the highest level (6.82%) measured at the fully
ripe stage. Collins et al. (2006), instead, reported no significant
changes during ripening.
Tigist et al. (2013) observed that fresh market and processing
tomato cultivars reached their TSS peak after 16 and 20 days of
storage at ambient conditions, respectively, to diminish thereafter
in all cases (Table 6). Ali et al. (2010) examined the effect of fruit
coating with gum arabic coating on tomato quality, and found
increasing trend of TSS during subsequent storage, even though
final levels were lower in comparison with untreated samples. An
increase in TSS from 5.2% to 5.9% and 5.8% to 5.9% was noticed
after 8 days of storage for tomato fruits harvested respectively at the
turning and pink stage, but a decline in TSS from 6.6% to 4.3% was
found for red-ripe tomato fruits (Gaur & Bajpai, 1982). Fruits har-
vested at the mature-green ripening stage also displayed an increase
in TSS levels after 8 days of storage, though to a different extent ac-
cording to cultivar and storage temperature (Kumah et al., 2011).
ˇ
Znidarcic and Pozrl (2006) found that °Brix values of tomato fruit
following storage for 3 weeks at 10 °C increased slightly from 5.06
to 6.92. Kumar et al. (2007) studied different open-pollinated and
hybrid varieties and established a range of TSS content between
3.88 and 6.35 °Brix. The TSS contents increased in all genotypes
during 9 days of storage at ambient temperature. Getinet et al.
(2008) observed significant interactions between genotype and
maturity stage influencing tomato fruits TSS content variability,
which increased during storage. TSS increase throughout matura-
tion and ripening in parallel with color, and postharvest changes
were reported to be related to ripening stage at harvest and the
temperature of the storage conditions (Atta-Aly, Brecht, & Huber,
2000; Trejo & Cantwell, 1996; ˇ
Znidarcic & Pozrl, 2006). D´
avila-
Avi ˜
na et al. (2011) pointed out that TSS in tomato fruits harvested
at the breaker stage remained rather unchanged throughout storage
at 10 °C, except for control fruit which displayed a 15% increase
when the storage period expired. Pink tomatoes showed a decrease
in TSS of approximately 15% to 20% with respect to the initial
value. Pila et al. (2010) observed an increasing trend for TSS in
tomatoes throughout 10 days storage period at 34 ±1°Cafter
different treatments including gibberellic acid, CaCl2, and salicylic
acid. Untreated fruit showed higher TSS values as compared with
treated fruits. Assi et al. (2009) compared traditional and modern
tomato handling methods used in Jordan, in which tomato fruits
harvested at the pink or light-red stages were held at 12 or 22 °C,
and found an increase in TSS with storage period, with signifi-
cant differences between both storage temperatures. Sammi and
Masud (2007) studied the impact of different packaging systems
on TSS of tomatoes held at ambient conditions, and found that
TSS increased with ripening stage in both unpacked and packed
samples. Javanmardi and Kubota (2006) analyzed red-ripe cluster
tomato fruits of cv. “Clermon” grown under a hydroponic sys-
tem in greenhouses for TSS changes during consecutive 14 days of
storage at 12 °Cand5°C with respect to 7 days room temperature
storage for the control. The authors reported that TSS values in
tomatoes harvested at the light-red to the red-ripe ripening stages
and stored at different temperatures did not show any variation
during up to 14 days. TSS values remained unchanged for 21 days
of storage of untreated and treated tomato fruits during 3 weeks
of storage at 12 to 14 °C, and were not significantly affected by all
the applied light treatments applied (Liu et al., 2009). Similarly, no
significant variations in TSS contents were detected in tomatoes
kept for 14 days at room temperature (Wills & Ku, 2002) or at
12 °C during 10 days (Kagan-Zur & Mizrahi, 1993).
Siddiqui and Singh (2015) demonstrated that puree prepared
from tomato fruit of high-pigment cultivars Berika and BCT-
119, lost about 45% to 58% of their original TSS values respec-
tively, whereas the loss in ordinary cvs Patharkutchi and Punjab
Chhuhara was only 43% to 56%, respectively. Similarly, Safdar,
Mumtaz, Amjad, Siddiqui, and Hameed (2010) reported increased
TSS contents in tomato paste during storage for 240 days at dif-
ferent temperatures. Since TSS is referred to as the sum of acids
sugars and other secondary components (Beckles, 2012), the con-
sumption of a part of them by micro-organisms as a food source
is likely to lead to decreased TSS levels in puree during storage.
Total acidity
In many fruits, including tomatoes, titratable acidity (TA) is a
good index to assess fruit maturity as its values are known to de-
crease in the later stages of ripening. Usually, in tomato TA is
low in immature-green fruits, reaches a maximum at the turning
stage and decreases rapidly afterwards (Table 6; Castro, Vigneault,
Charles, & Cortez, 2005; Chen, Wilson, Kim, & Grierson, 2001;
Getinet et al., 2008; Kumar et al., 2007; Pila et al., 2010). Accord-
ingly, Islam et al. (1996) and Knee and Finger (1992) reported a
peak in the amount of organic acids at the pink ripening stage.
C2018 Institute of Food Technologists®Vol.17,2018 rComprehensiveReviews in Food Science and Food Safety 1555

Dynamic changes in health-promoting properties . . .
Citric acid content was much higher than that of malic acid and
oxalic acid, whose level was very limited.
A decline of TA levels also occurs during storage and has been
associated with fruit quality deterioration, being strongly related
to a loss of the typical sour taste of tomatoes. The progressive
reduction of tomato fruit TA during ripening and storage is the
result of organic acid catabolism and is partially related to the
increase of respiration rates occurring in climacteric fruits, when
citric and malic acids are rapidly consumed as key respiration
substrates (El-Anany, Hassan, Rehab, & Ali, 2009).
It has been demonstrated that fruit pretreatments and/or
postharvest storage conditions significantly affect TA decline
(Hern´
andez-Su´
arez, Rodr´
ıguez-.Rodr´
ıguez, & D´
ıaz-Romero,
2008; Majidi, Minaei, Almasi, & Mostofi, 2011; Table 6).
“Micro-Tom” tomatoes stored at 20 or 6 °C displayed patterns
of change diverse for each organic acid. At both temperatures
citric, malic, ascorbic and tartaric acids were slight but signifi-
cantly reduced throughout maturity, while succinic acid slowly
accumulated. Refrigeration slowed the specific kinetics of most
acids, without stopping it (G´
omez et al., 2009). Significant
variations in TA values were also reported by Moneruzzaman
et al. (2008, 2009) in freshly harvested tomato fruits grown
in an open field at Mymensing (Bangladesh) and Pealing
Jaya (Malaysia) and subjected to 3 different storage condition.
Ripening stage, storage time and conditions (mainly temperature
and RH) were all found to affect TA levels. After 9 days
of storage, half-ripe harvested tomatoes had the highest TA
(0.48%), followed by fully ripe (0.47%) and mature-green tomato
fruits (0.44%).
Islam et al. (1996) found that organic acid levels significantly
dropped with the increase of storage temperature, with fruits kept
at 15 °C showing values higher than those kept at 25 or 30 °C.
D´
avila-Avi˜
na et al. (2011) noticed that TA of tomatoes decreased
with maturity irrespective of coating treatments. However, TA of
breaker fruits treated with mineral oil and carnauba wax dropped
40% and 25%, respectively, compared to nontreated fruits. Getinet
et al. (2008) found a declining trend for TA during storage of 2 dif-
ferent tomato (“Marglobe” and “Roma VF”) cultivars; however,
the extent of this decline was cultivar-specific. Kumar et al. (2007)
opined that TA of different tomato genotypes (open-pollinated
varieties and hybrids) ranged from 0.34% to 0.47% and decreased
during subsequent storage for 9 days. Ali et al. (2010) found a
declining trend for TA during storage of coated and uncoated
tomatoes irrespective of treatment. Auerswald, Peters, Br ¨
uckner,
Krumbein, and Kuchenbuch (1999) reported a 22% increase in the
TA level of hydroponically grown tomato fruits after a postharvest
storage period of 4 days. Sammi and Masud (2007) reported a
decrease in TA during postharvest tomato ripening, with a faster
rate in packed tomato fruits. Regardless of the ripening stage, the
highest value of TA was monitored in control tomato fruits during
the storage period. Whereas, Toor and Savage (2006) reported that
hydroponically grown tomato fruits stored at 15 and 25 °Chad
TA values of 0.97% and 1.06% citric acid, respectively, significantly
higher with respect to fruits subjected to refrigeration (0.77% cit-
ric acid), and increased during subsequent storage, particularly at
ambient temperature.
Ord´
o˜
nez-Santos, V´
azquez-Od´
eriz, Arbon´
es-Maci˜
neira, and
Romero-Rodr´
ıguez (2009) reported that the level of malic and
citric acids of tomato pulp decreased significantly by 51% and
71%, respectively, during storage for 180 days. However, Gould
(1992) reported that following storage of tomato paste at different
temperatures, TA values increased linearly throughout the period,
with higher levels (18.39%) at ambient temperature than at –10 °C
(7.47%; Table 6).
Total sugar
Total sugar (TS) is also considered an important trait for tomato
quality assessment. Tomatoes accumulate fructose and glucose
more than sucrose (Siddiqui, Ayala-Zavala, & Dhua, 2015). Sugar
content was found to increase throughout r ipening from the green
to the red-ripe stages (Tadesse et al., 2012) (Table 6). In fruit tis-
sues, sucrose synthesis is driven by an increase in sucrose synthase
(SS) activity (Islam et al., 1996), indicating that the enzyme plays a
central role in sucrose accumulation. The hydrolysis of starch also
contributes to the increase of soluble-sugar content during tomato
ripening (Pila et al., 2010).
An initial increment in tomato fruit TS values over-ripening
has been noted, subsequently, TS remained unchanged or exhib-
ited a minor decrease (Baldwin, Nisperos-Carriedo, & Moshonas,
1991). Sugar content varies with maturity stage at harvest (Sinaga,
1986). Dalal, Salunkhe, Boe, and Olson (1965) found that at the
mature-green, breaker, pink, red, and red-ripe tomato ripening
stages, reducing sugars accounted for about 2.4%, 2.90%, 3.10%,
3.45%, and 3.65% on a FW basis, respectively. Regardless of tem-
perature, the level of soluble sugar showed an increasing trend
during storage. Sammi and Masud (2007) observed that during
the transition from the green to the turning maturity stages, con-
trol fruits displayed a peak of sugars. Islam et al. (1996) showed that
reducing sugar accumulated more rapidly at the final than early
ripening stages. A peak of sucrose was detected in immature-green
and mature-green tomato fruits and dropped in later stages of ma-
turity. In tomatoes about 95% of total soluble sugars are reducing
sugars, with fructose levels higher than glucose.
G´
omez et al. (2009) reported increasing levels of glucose and
fructose throughout 20 °C storage period, in breaker tomato fruits,
but this accumulation was decreased under refrigerated storage,
with final values attaining approximately 80% of those measured in
control fruits. Kumar et al. (2007) observed different tomato geno-
types in which the reducing sugar content ranged between 1.98%
and 3.54% and found a gradual increase during storage. Mature-
green tomato fruit stored at low temperature (14 to 19 °C) for
28 days exhibited increasing TS levels up to 8 days and decreased
afterwards (Melkamu, Seyoum, & Woldetsadik, 2008). Auerswald
et al. (1999) reported that the level of reducing sugars in hydropon-
ically grown tomato fruits was unaffected after a week postharvest
period (Table 6). Packed fruits showed the highest TS content
by the end of storage (pink-red to red-ripe maturity stages). Sig-
nificant variation among different maturity stages was reported
by Moneruzzaman et al. (2008) for TS content in fruit pulp. TS
content increased with advancing ripening of fruit irrespective of
maturity stage. A peak of TS (4.03%) was detected in fully ripe
tomato fruits, while the lowest value (3.30%) was measured in
mature-green harvested tomatoes after 12 days of storage.
Aroma
Being a climacteric fruit, tomatoes exhibit the characteristic in-
crease in respiration and rate of ethylene production, as well as
various typical ripening-associated quality characteristics such as
chemical composition, color, texture, taste, and aroma. Aroma is
a major quality attribute determining consumer choice of tomato
for its use, as fresh or processing pur poses. These specific process-
ing purposes help determine the most suitable maturity stage at
which to harvest the tomatoes. Since ethylene is closely associ-
ated with the initiation and subsequent integration of biochemical
1556 ComprehensiveReviews in Food Science and Food Safety rVol. 17, 2018 C2018 Institute of Food Technologists®

Dynamic changes in health-promoting properties . . .
changes during tomato fruit r ipening, most postharvest processes
emphasize the control of ripening aiming to either expanding the
shelf-life potential or accelerating maturity. The exogenous ap-
plication of ethephon, an ethylene-releasing chemical, has been
frequently used to fasten the process of off-vine ripening of com-
mercial tomatoes. Tomato fruit aroma profiles are complex. Of
the over 400 identified volatile compounds, about 30 molecules
seem implicated in providing the characteristic pleasant flavors of
fresh fruits and processed tomato products (Petro-Turza, 1987).
The biosynthesis of aroma molecules in tomatoes, as well as in
most other fruits and vegetables, depends on several pathways
(El Hadi, Zhang, Wu, Zhou, & Tao, 2013; Salles, Nicklaus, &
Septier, 2003). Many aroma-contributing volatiles (alcohols, car-
bonyls, acids and esters) originate from the metabolism of amino
acids, including aspartic and glutamic acids, leucine and glutamine.
Conversion of amino acids to keto acids by aminotransferases and
further oxidation to aldehydes by enzymatically catalyzed decar-
boxylation leading to the formation of various volatile esters has
been reported (Petro-Turza, 1987). The alcohol 3-methylbutanol,
synthesized from leucine, is an important volatile contributing to
sweet and fresh r ipe tomato aroma notes (Buttery & Ling, 1993).
The increase in nonprotein nitrogen associated with the decrease
in protein level has been correlated to an increment in the synthesis
of aroma volatiles. Hexanal, cis-3-hexenal, trans-2-hexenal, cis-3-
hexenol, and hexenol are other important C6 volatile chemical
compounds prominent in tomato fruit flavor (Ruiz et al., 2005;
Yilmaz, Tandon, Scout, Baldwin, & Shewfelt, 2001). The increase
in hexanal production throughout off-vine tomato fruit ripening
has been found to correlate negatively with perceived sourness
and positively with sweetness (Krumbein, Peters, & Br ¨
uckner,
2004) (Table 6). Phenylacetaldehyde and 3-methylbutanal arise
from glycosides hydrolysis throughout maturity. Furaneol con-
tributes to the “fresh” notes in tomato fruits (Buttery, Takeoka,
Naim, Rabinowitch, & Nam, 2001). Volatile monoter penes are
also present in tomato, but in small quantity. Some of the aroma-
contributing compounds are synthesized enzymatically through
the oxidation of membrane lipids, mainly after damage of fruit
tissues at later ripening stages (Galliard, Matthew, Wright, &
Fishwick, 1977). Carbonyls, short-chain alcohols and hydrocar-
bons, long-chain alcohols and esters typically form the aroma
of field-ripened tomato when present in the ratio of 32:10:58,
respectively (Shah, Salunkhe, & Olson, 1969). Benzaldehyde, cit-
ronellyl propionoate, citronellyl butyrate, decanal, dodecanal, ger-
anyl acetate, geranyl butanoate, nonanal, and neral in plant-ripened
tomato fruits were reported to be released in higher concentrations
than in those artificially ripened, which in turn displayed higher
emissions of butanol, 2,3-butanedione, isopentanal, isopentyl ac-
etate, 2-methyl-3-hexanol, 3-pentanol, and propyl acetate (Mad-
havi & Salunkhe, 1998) (Table 6). Off-flavors are associated with
increased productions of 2-methyl-1-butanal, particularly by off-
vine ripened tomato.
Conclusions
Increasing evidence is available regarding the positive role of
a regular dietary intake of fresh tomato fruits and tomato pro-
cessed products on human health and wellbeing. In this review,
the health-promoting, physical and eating-related properties of
tomato fruit are presented and discussed as affected by ripening
stages, storage conditions and their combinations. During off-vine
ripening of tomato, some attributes increase such as lycopene, total
carotenoids, LAA, PLW, TSS, and aroma. However, HAA, fruit
firmness, and TA decrease. AsA show cultivar-dependent accu-
mulation trends during off-vine ripening.
Storage influence health-promoting, physical as well as eating-
related properties of tomato, negatively or positively according to
storage duration and conditions. This review highlights that to-
tal carotenoids, particularly lycopene, total antioxidant activity, and
TS increase during storage, whereas phenol content, AsA content,
and fruit firmness decrease. It is relevant to emphasize that the ef-
fects of off-vine ripening on tomato quality depend mostly on the
initial content of each bioactive compounds since high-pigment
and ordinary cultivars will not reach the same content of lycopene
after the same storage period. Storage effects on tomato quality
will also depend mostly on the applied treatment and temperature.
Generally high quality is obtained under low storage temperature
and mild treatment. Tomato fruit is subjected to complex changes
during ripening and postharvest affecting bioactive molecules and
health-promoting properties, physical and eating quality-related
attributes (Figures 1 and 2). All of the above-reported changes
are aiming to accumulate health-promoting compounds during
ripening and to preserve as long as possible the shelf-life of the
fruit during postharvest storage of tomato fruits under various
conditions.
Author Contributions
Siddiqui MW conceptualized the idea of this review. Siddiqui
MW, Lara I, Ilahy R, and Tlili I scanned the literature, retrieved
and processed papers referenced in the review and wrote the
manuscript. Prasad K, Asghar A, Lenucci MS, and Hdider C
critically reviewed the manuscript and enriched key parts in the
manuscript. Homa F and Deshi V helped in revising manuscript.
All authors contributed to the preparation of the tables and the
revision of the paper before submission.
Conflict of Interests
Authors declare no conflict of interests.
Abbreviations
GAE Gallic acid equivalent
FW Fresh weight
HVEF high-voltage electrostatic field
DW P delactosed whey permeate
AsA ascorbic acid
UV-C ultraviolet-C
HAA hydrophilic antioxidant activity
LAA lipophilic antioxidant activity
AVG Aminoethoxyvinylglycine
PLW physiological loss in weight
RH relative humidity
UV-B ultraviolet-B
TSS total soluble solids
Brix degree Brix
TA titratable acidity
TS total sugars
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