Department of Plant Breeding

An overview of recent studies of tomato

(Solanum lycopersicum spp) from a social,
biochemical and genetic perspective on
quality parameters

Evelyn E. Villanueva Gutierrez

Introductory Paper at the Faculty of Landscape Architecture,

Horticulture and Crop Production Science 2018:3
Swedish University of Agricultural Sciences
Alnarp, December 2018
An overview of recent studies of tomato
(Solanum lycopersicum spp) from a social,
biochemical and genetic perspective
on quality parameters
Evelyn Elizabeth Villanueva Gutierrez

Swedish University of Agricultural Sciences,

Department of Plant Breeding, [Link]@[Link]

Place of publication: Alnarp-Sweden

Year of publication: 2018
Title of series: Introductory Paper
Part number: 2018:3
Online publication: [Link]
Bibliographic reference: Villanueva, EE (2018). An overview of recent studies of tomato
(Solanum lycopersicum spp) from a social, biochemical and genetic
perspective on quality parameters. Alnarp-Sweden: Sveriges
lantbruksuniversitet. (Introductory Paper, 2018:3).

Keywords: Tomato, postharvest management, quality traits

Swedish University of Agricultural Sciences

Faculty of Landscape Architecture, Horticulture and Crop Production Science
Department of Plant Breeding
Plant Product Quality Group
Abstract

Tomato is the second most important horticultural product cultivated worldwide.

For research, tomato is considered a model organism of the Solanaceae family, and
has therefore been, and is still a major crop subject of studies both in the laboratory
and under field conditions.

This introductory paper aims to explore origin and distribution, economic im-
portance, social impact, and postharvest losses of tomatoes from field to consump-
tion.

The present literature overview summarizes variation in quality traits such as bio-
active compounds in fruit, as well as in sensory factors, e.g. color, shape, texture and
volatile compounds. From the health perspective, regular intake of tomato fruit con-
sumed fresh, as a juice, or canned, increase levels of carotenoids, lycopene, vitamin
C, and polyphenols compounds in the daily diet, which are reported to be beneficial
to health and lower the risk of some diseases, such as some forms of cancer and dia-
betes. Regarding quality traits, internal traits as well as color, size, and texture, are
determined by the presence or absence of various compounds in the tomato, while
resistance to mechanical damage is influencing external quality traits.

Quality traits are affected by environmental conditions during pre- and post-har-
vest stages. Tomato production management influences the yield, acceptability, price
and length of shelf life but differently in each tomato cultivar.

Finally, this introductory paper presents a brief overview of studies concerning

genetic diversity among domesticated and wild relatives and their impact on quality
traits, including recent studies of tomato biodiversity, genetic identity of cultivars and
their wild relatives. In addition, a few studies are mentioned concerning trans-formed
plants, showing the importance of different enzyme activities in relation to effects on
internal and external quality traits

Keywords: Tomato, postharvest management, quality traits.

1
2
Table of contents

List of tables 5

1 General perspective of tomato crop and its societal importance 7

1.1 Origin and distribution of tomato 7
1.2 Nutritional importance of the crop 8
1.3 Economic and social importance of the crop 8
1.4 Losses from the field to consumers 9
1.5 Globalization and its impact on horticultural farmers. A social point of view 10

2 Factors determining the quality of tomato fruits 12

2.1 Nutritional factors 12
2.1.1 Carotenoids, including lycopene 12
2.1.2 Vitamin C including ascorbic acid 14
2.1.3 Polyphenols contents and antioxidant properties 14
2.2 Sensory factors 16
2.2.1 Textural properties 16
2.2.2 Aroma and flavor 17
2.2.3 Total soluble solids and titratable acidity 18
2.2.4 Enzyme activity 18
2.2.5 Volatile organic compounds 19
2.2.6 Color 20

3 Factors affecting quality parameters 24

3.1 Effects of environmental conditions on fruit quality during development of
fruits 24
3.1.1 Temperature 24
3.1.2 Relative Humidity 25
3.1.3 Light 25
3.1.4 Soil nutrients 26
3.1.5 Size 27
3.2 Current post harvest management of tomato 27
3.2.1 Temperature treatments and their implication on shelf life,
appearance, color, firmness, chilling injuries and bioactive com-
pounds 28
3.2.2 Application of organic substances after harvest to improve or extend
shelf life of tomato 29

3
 3.2.3 Application of inorganic substances after harvest to extend shelf life of
tomato 29
3.2.4 UV light application on tomato and impact on shelf life 31
3.2.5 Application of modified or controlled atmosphere using carbon dioxide,
oxygen, nitrogen and ozone treatment and impact on shelf life 32
3.2.6 Mechanical technologies in postharvest management of tomato 33
3.3 Effects of postharvest techniques on fruit quality traits 33
3.3.1 Texture 33
3.3.2 Flavor and volatile compounds 33
3.3.3 Total soluble solids and titratable acidity 34
3.3.4 Treatments effecting fruit color during the ripening process 35
3.3.5 Change in nutrients and nutrient contents 36
3.4 Reduction of losses 37
3.4.1 Reduction of losses by the use of pre-harvest factors 37
3.4.2 Reduction of losses during postharvest handling 38
3.4.3 Biochemical changes caused by mechanical damage 39

4 Genetic studies in tomato 40

4.1 Genetic studies related to biodiversity using molecular markers 40
4.2 Genetic studies related to quality parameters using molecular markers 41
4.2.1 Nutrient contents and sensory factors 41
4.2.2 Texture and yield 42
4.2.3 Color 42

5 General Conclusions 45

Acknowledgements 47

References 48

4
List of tables

Table 1. Levels of phenolic compounds in fresh tomato and tomato sauce with or
without olive oil (Martínez-Huélamo et al., 2016). 16
Table 2 Volatile compounds quantified in three tomato varieties (Preedy & Watson,
2008). 20
Table 3. Relationships between storage temperature and change in color. 36
Table 4 List of genes associated with color traits in tomato flesh fruit (Rick, 2017). 44

5
6
1 General perspective of tomato crop and
its societal importance

1.1 Origin and distribution of tomato

The center of origin of tomatoes have been debated by many, some are suggest-
ing the center to be the dry coastal desert of Peru (Jenkins, 1948, Preedy and Watson,
2008, Blanca et al., 2012), while others have suggested a dual center with one part
in the coastal region between the Andes (Blanca et al., 2012) and the ocean and the
second part from South Mexico to Guatemala (Bauchet and Mathilde, 2012). Wild
relatives of tomato are distributed in the Andes from Ecuador, through Peru and to
Chile (Peralta et al., 2005), growing between sea level and 3300 meters above sea
level (Blanca et al., 2012) in diverse climatic conditions. The domestication is still
unclear but linguistic evidence has postulated Peru and Mexico as the major regions
of domestication (Peralta et al., 2006). Tomatoes are known to be used in cooking
in Mexico by the Aztecs already 500 BC and were transferred to the rest of the world
by the conquistadors after the capture of the Aztecs territory (Bergougnoux, 2014).

Taxonomically, tomato belongs to the Solanaceae family. The cultivated tomato

belongs to the species Solanum lycopersicum, while Solanum pimpinellifolium is
the closest wild relative with a divergence of only 0.6% nucleotide base pairs (The
Tomato Genome et al., 2012). A range of other wild tomato relatives are available,
e.g. Solanum chmiewelskii, Solanum neorickii, Solanum chilense, Solanum habro-
chaites, Solanum pennilli, Solanum juglandifolium, Solanum ochranthum, Solanum
lycopersicoides, Solanum sitiens, Solanume corneliomuelleri, Solanum arranum
and Solanum galapagense (Peralta et al., 2006). Despite the fact that wild tomato
relatives have contributed limitedly with desirable phenotypic traits to current cul-
tivated tomatoes, interesting alleles for future tomato breeding may still be available
in uninvestigated tomato collections (Rendón-Anaya and Herrera-Estrella, 2017).

7
 A large variation has been ascribed to the tomatoes as related to differences in
shape, color, flavor and other parameters. Wild tomato are generally small as com-
pared with the domesticated ones (Bergougnoux, 2014), and the differences in size
is regarded as a result of changes in a total of six quantitative traits loci (QTL) during
the domestication process (Bai and Lindhout, 2007, Bergougnoux, 2014).

1.2 Nutritional importance of the crop

Tomatoes are currently an important food component globally. The tomatoes are
in fact the second largest vegetable both in terms of production and consumption
(FAO, 2016). Reports from the United States show tomato as the second most con-
sumed fresh vegetable with 6 kg/person in 2017 (USDA, 2016). Tomatoes are
known as a source of vitamins and pro-vitamins (vitamin C, pro-vitamin A, β caro-
tene, folate), minerals such as potassium, and secondary metabolites such as lyco-
pene, flavonoids, phytosterols and polyphenols (Beecher, 1998, Luthria et al.,
2006). Thus, 100 g of fresh tomato provides over 46%, 8% and 3.4% of the daily
requirements of vitamin A (being 900 UE), vitamin C (being 82.5 mg) and potas-
sium (being 3500 mg), respectively (Gebhardt and Thomas, 2002, Canene-Adams
et al., 2005). Furthermore, processed tomato such as soup, paste, concentrate, juice
and ketchup (Bergougnoux, 2014) also contribute positively to human health by the
content of the mentioned compounds in these products.

1.3 Economic and social importance of the crop

Tomato is produced in temperate, subtropical and tropical areas around the world
(Blanca et al., 2012) and it is the second horticultural crop produced in terms of
yield in the world (FAO, 2016). Numbers from 2016 showed United States, China,
India and Turkey as the countries with the largest production area (FAO, 2016,
USDA, 2016). Fresh tomato production reached 163.719.357 tons in the world in
2013, and around 4.5% of the produced tomatoes are traded. The same year, the
relatively highest tomatoes export was reported from Mexico followed by The Neth-
erlands and Jordan, exporting 20%, 13% and 8% of their produced tomatoes, re-
spectively. The revenue for the tomato export from these countries during the same
year was 1195, 1675 and 517 US dollar/ton. Countries with the highest relative to-
mato import were during the same time Russia, followed by Germany and France

8
with 12%, 10.3% and 7.8% respectively (FAOSTAT, 2013). Tomatoes are produced
on field, but also under controlled conditions during autumn and winter. They are
also produced in colder climates such as in the Scandinavian countries. Controlled
conditions certify tomatoes production by the use of artificial lighting, increased
temperature using heaters, as well as fertigation to produce high quality tomatoes
(Oda and Saito, 2006).

Tomato production is expensive (USDA, 2016). The investments start by the use
of certified seeds (Tüzel and Öztekin, 2017). Breeding and seed production are car-
ried out in order to sustain the desires from growers, including desirable traits such
as high yield and disease resistance, but also based on the destination of the cultiva-
tion and on consumer preferences (Carbonell et al., 2018, Yamamoto et al., 2016).

During production, recent technologies are used to maximize yield and product
quality, and to reduce use of pesticides. Use of seedbeds, mulch, branch conduction,
pruning, drip irrigation, and application of specific macro and micro nutrients during
each development phase of tomatoes all contribute towards reaching the potential
yield provided by the certified seeds (Tüzel and Öztekin, 2017). In open field pro-
duction, mechanical harvest is used for fruits destined to industrial purposes and
hand-picked harvesting for fruits destined to fresh consumption (USDA, 2016).

Fresh tomato prices have been found to suffer more from price variation than
processed tomatoes, and the price is clearly dependent on the shipping-point price.
It has been estimated that one-fourth of the final price is paid by the shipment chain
(USDA, 2016).

1.4 Losses from the field to consumers

Food losses, happening from the field to consumers, are greatly affected by the
locality where the product was grown, yearly climate conditions, season of cultiva-
tion and, nowadays, also on climate change (Kader, 2005, Hodges, 2010). Losses
differ also depending on whether the production occurs in a Developed Country
(DC) or a Less Developed Country (LDC). In DCs, the main losses (23%) are con-
centrated to the treatment of the product by the consumers (Kader, 2005). The larg-
est losses in LDCs (up to 50%) are concentrated to the treatment of the product by
the producers (Hodges, 2010). Thus, the losses start in the field (FAO, 1989) and
have various causes, including biological spoilage (Hodges, 2010), awaiting for

9
shipment (Campbell et al., 1986, Ayandiji and Omidiji, 2011), unavailability of stor-
age facilities (Ayandiji and Omidiji, 2011), and injuries (Campbell et al., 1986).

In general, causes of losses can be divided into two groups: primary causes and
secondary causes. Primary causes of losses include biological, microbiological,
chemical, biochemical, mechanical, physical, physiological and psychological
causes of loss (FAO, 1981). For tomato fruits, two classes of injuries have been
described: Injuries during the production, and mechanical injuries at harvest. A
source of tomato losses is related to the over-ripening of the tomatoes on the plant
(Campbell et al., 1986). Secondary causes of loss include unsuitable harvesting tech-
niques, inappropriate containers for transport, unsuitable storage facilities, inappro-
priate transportation, unsuitable refrigeration, inappropriate drying equipment, tra-
ditional processing, legal standards, and lack of knowledge on management and
bumper crops (FAO, 1981). Improper temperature management during harvest and
storage will quickly result in a reduction in value of tomatoes (Hodges, 2010,
Ayandiji and Omidiji, 2011). During the entire value chain, losses occur also at har-
vest time, preparation during extraction of the edible part, preservation, processing,
storage and transportation (Amalendu et al., 2003). Transport procedures may lead
to postharvest losses through damages or injuries on the skin of the tomatoes, re-
sulting in attacks of pathogens during storage (Campbell et al., 1986, Ayandiji and
Omidiji, 2011).

1.5 Globalization and its impact on horticultural

farmers. A social point of view

With increased globalization, horticultural products for exportation have be-

come an alternative, providing job opportunities for small-hold farmers while this
type of operation is rather labor intensive. For example, the export of horticultural
products from the North of Mexico is absorbing workers from surrounding poor
regions causing migration from the South of Mexico (Zabin and Hughes, 1995).
Horticultural production systems are normally more work intensive than agricultural
ones for obtaining the final product, as has been concluded from studies in Vietnam,
Philippines and Bangladesh (Weinberger and Lumpkin, 2007). Thus, export of hor-
ticultural products is reducing poverty, by the provision of new and alternative jobs
to small-scale farmers. (Weinberger and Lumpkin, 2007, Van den Broeck and
Maertens, 2017). However, the farmer plays an important role by having impact on
the quality of the produced tomato. The production is highly affected by the degree
of the farmers’ knowledge and on the support by regional and national institutions.

10
Lack of knowledge and support are known to increase harvest losses (Emana et al.,
2017). Regional efforts to increase tomato production by smallholders in Africa has
shown positive results, e.g. when novel leaf curl disease resistant tomatoes seeds
were released in West Africa (Perez et al., 2017). In addition, regional efforts in the
horticultural production chain is known to benefit women, due to the fact that
women represent 80% of the work-force in this production chain, and increasing
yield and quality of the produce results in more decision power for women within
their communities (Alter, 2009).

11
2 Factors determining the quality of tomato
fruits
Factors that influence product quality can be divided in two groups: Intrinsic factors
(color, shape, and freedom of defects) and internal attributes such as texture, sweet-
ness, acidity, aroma, flavor, shelf life and nutritional value (Hewett, 2006). Growers
are mainly basing their decisions, as regards which cultivars to produce, on intrinsic
factors desired by the buyer (FAO, 1989).
Crop management together with external conditions during the pre-harvest period
will determine the quality of the product (FAO, 1989, Hewett, 2006), and during the
best circumstances, this quality may be kept during the post-harvest period when
quality improvements are not possible (FAO, 1989).
Opportunities to extend the shelf life of perishable commodities have been evaluated
worldwide (Hodges, 2010), although adaptation to technologies prolonging shelf
life is greatly dependent on knowledge of the grower (Kader, 2005).

2.1 Nutritional factors

2.1.1 Carotenoids, including lycopene

Carotenoids are well known as natural compounds involved in reducing the risk of
development of several types of diseases such as diabetes, gastrointestinal and car-
diovascular diseases e.g. by reducing the amount of oxidized low density lipopro-
teins (Hyman et al., 2004, Rao and Rao, 2007, Preedy and Watson, 2008). Carote-
noids are also known to prevent development of degenerative diseases such as blind-
ness, xerophthalmia and degeneration of muscles (Paliyath et al., 2008). In addition,
carotenoids have been correlated to anticancer properties of stomach, lung and pros-
tate cancer (Giovannucci, 1999, Wang et al., 2016). A recent study have shown that

12
high-risk prostate cancer patients, constantly consuming lycopene through intake of
a tomato sauce during a period of 4 years, significantly reduced their risk to contract
prostate cancer. These patients also became low-risk prostate cancer patients after
the 4 years period of lycopene consumption (Wang et al., 2016).

The health benefits of carotenoids have been attributed to their function as natural
antioxidants. Lycopene is the major antioxidant carotenoid in tomato followed by β
carotenoid (Hyman et al., 2004). Furthermore, α-carotene, β-carotene and β-crypto-
xanthin are precursors of vitamin A, which is not the case for all carotenoids
(Institute of Medicine, 2000, Apel and Bock, 2009).

Bioavailability of the carotenoids is highly important as related to their impact for

human nutrition. Homogenization and heat treatments are common steps in trans-
forming fresh tomato to other food commodities. Homogenization is normally
breaking down the aggregated structures of the cell by changing the fiber network.
However, homogenization at 90ºC in 30 minutes was shown to not change the fiber
network (Colle et al., 2010). Similarly, another study demonstrated that homogeni-
zation at 100 MPa reduced bioavailability of carotenoids with one third as compared
to that of not treated samples (Panozzo et al., 2013).Thus, homogenization is nor-
mally resulting in a higher stability of the fiber network, thereby leading to signifi-
cantly lower bioavailability of carotenoids (Colle et al., 2010).

Other food processing steps, such as cooking, are instead known to contribute to
breakdown of the cell walls, thereby facilitating bioavailability of carotenoids (Shi
and Le Maguer, 2000). In addition, bioavailability of carotenoids increases with the
presence of lipids, as carotenoids are fat-soluble pigments (Rao and Rao, 2007).
Carotenoids combined with lipids are known to result in an isomerization of the
lycopene. Therefore, through isomerization with corn oil, the bioavailability of ly-
copene from processed tomato combined with corn oil was found to be 3-4 times
higher as compared to that from fresh tomatoes combined with corn oil (Gartner et
al., 1997, Shi and Le Maguer, 2000).

The most accurate method for objective detection and quantification of carotenoids
is to use HPLC (Serino et al., 2009). Thus, accurate detection of lycopene in fresh,
canned or juice samples has been performed applying HPLC methods (Davis et al.,
2003, Hyman et al., 2004). However, spectrophotometric and colorimetric methods
are also commonly used to quantify lycopene. Choice of method and accuracy of
results are highly dependent on qualified labor, time and accuracy needed (Davis et
al., 2003, Hyman et al., 2004).

13
2.1.2 Vitamin C including ascorbic acid

Tomato is an important source for humans of vitamin C, including ascorbic acid and
some, generally low, amounts of dehydroascorbic acid. No significant differences
was found in quantity of ascorbic acid for different types of tomatoes, e.g. salad
tomato versus those for processing. However, ascorbic acid was among the compo-
nents in tomato highly sensitive to thermal degradation (Abushita et al., 2000). Red
tomatoes contain approximately 19 mg of vitamin C (Nunes, 2008) and depending
of the cultivar and growing conditions ascorbic acid contents is around 14.6 mg per
100 g of fresh red tomato (Abushita et al., 2000).

Synthesis and accumulation of ascorbic acid have been correlated with the maturity
stage of tomatoes. Ripening tomatoes under storage conditions was shown to de-
crease the content of ascorbic acid in the tomatoes with up to 50% as compared to
vine ripe tomatoes, demonstrating that accumulation of ascorbic acid is favored if
the fruit reaches the maturity stage on the plant (Nunes, 2008). Salt stress during
growth of tomato enhances the concentration of sugars, organic acids and amino
acids, all characters being well appreciated by consumers (Passam et al., 2007).

The vitamin C content in fresh tomatoes could be preserved under storage condi-
tions if modified atmosphere conditions was applied with a polyolefin films as an
enclosure barrier with enough O 2 permeability to avoid anaerobic respiration
(Passam et al., 2007). Exclusion of such barriers between the tomatoes and the at-
mosphere lead to the disappearance of vitamin C within 3 days (FAO, 1989). Also,
vitamin C disappeared under heat conditions (FAO, 1989).

2.1.3 Polyphenols contents and antioxidant properties

Polyphenols are bioactive natural compounds present in vegetables, fruits and seeds
(Quideau, 2011). A bioactive compound could be essential or non-essential for hu-
mans, but is mostly considered beneficial for the human health (Biesalski et al.,
2009). Polyphenols can also be seen as “secondary metabolites” which presence are
correlated to corresponding genes, although the content is also being influenced by
the environment (Siracusa et al., 2012).

Polyphenol compounds are characterized by their phenolic structure in natural prod-

ucts (Tsao, 2010) and various types are soluble or insoluble in water (Papadopoulou

14
and Frazier, 2004). The most common polyphenols in tomatoes are flavonoids, hy-
droxycinnamic acids and anthocyanins (Siracusa et al., 2012, Verhoeyen et al.,
2002, Bovy et al., 2002), and the polyphenols are predominantly present in the to-
mato peel. In tomatoes, 30 different phenolic compounds have been determined be-
ing present; coffeic acid, caffeic acid-o-hexoside, chlorogenic acid, couma-
roylquinic acid, coumeric acid-o- hexoside, cryptocholorogeic acid,
dicaffeoylquinic acid, eriodictyol-o-hexoside, ferulic acid, ferulic acid -o-hexoside,
feruoylquinic acid -o -hexoside, gallic acid, homovanillic acid, kaempferol-o-ruti-
noside-hexoside, kaempferol-3-o-glucoside, kaempferol-3-o-rutinoside,
naringenin, naringen-in-o-dihexoside, naringenin-o-hexoside, neochlorogenic acid,
o-acetylprunin, phloretin-c-diglycoside, phloridzin-c-diglycoside, protocate-chuic
acid, prunin (naringenin-7-o-glucoside), quercetin, rutin-o-hexoside-pentoside, ru-
tin-o-hexoside, rutin-o-pentoside, rutin (quercetin 3-o-rhamnosyl-glucoside)
(Vallverdú-Queralt et al., 2010).

The physiological role of flavonoids is related to plant growth, protection from UV

radiation due to pigment production, seed coat development, pollen growth and de-
velopment of pathogen resistance (Harbone, 1995). Polyphenols’ role in protecting
the plant from the attack of viruses and other microorganism has also been reported
(Hemingway and Laks, 1992).

Pholyphenol compounds are suggested to influence human health; daily consump-

tion of food containing polyphenols is suggested to reduce the risk of cardiovascular
diseases (Bovy et al., 2002) by inhibition of the activity of kinase proteins, and ara-
chidonic acid metabolism (Vassallo, 2008). Such diet may also prevent against
chronic degenerative diseases (Vallverdú-Queralt et al., 2010), arthrosclerosis and
free radical damage (Stewart et al., 2000). It has been reported that in human cells
polyphenols may inhibit infections by herpes, pox, rabies and other viruses
(Hemingway and Laks, 1992).

Table 1 shows a list of the most abundant phenolic compounds, reported in fresh
tomato and in tomato sauce with or without olive oil. Bioavailability of naringenin
and quercetin when digesting the tomatoes, was found to increase after mechanical
intervention, heating or due to the addition of olive oil to the tomato sauce
(Martínez-Huélamo et al., 2016).

15
Table 1. Levels of phenolic compounds in fresh tomato and tomato sauce with or without olive oil
(Martínez-Huélamo et al., 2016).

Compound Fresh tomato Tomato sauce with Tomato sauce

FW (mg/g) oil (mg/g) without oil (mg/g)
5-CQA 0.0011 0.0008 0.0009

4-Caffeolyquinic acid 0.0011 0.0007 0.0006

CA hexoside I 0.0016 0.0014 0.0013

FA hexoside 0.0031 0.0025 0.0031

Homovanillic hexoside 0.0935 0.1278 0.1383

Niringenin 0.0048 0.0075 0.0066

Quercetin 0.0003 0.0003 0.0003

Rutin 0.0022 0,0054 0.0051

*Sample dose of 500 grams in fresh tomato and 250 grams in sauce; results presented in miligram per
gram (mg/g); FW, Fresh weight; 5-CQA, 5-caffeoylquinic acid; CA, Caffeic acid; FA, Ferulic acid.

2.2 Sensory factors

2.2.1 Textural properties

Textural properties in food are defined as a group of physical properties that con-
sumers can identify and evaluate during mastication (Bourne, 2014). In tomatoes,
texture is generally changing during fruit ripening (Chaıb et al., 2007). A variety of
methods exists, suitable for measurements of texture, and the most common ones
are defined as rheological, sensory and morphological. The most commonly used
rheological method study the flow of matter using an device called penetrometer.
This method has been used to measure resistance to deformation in tomato. Signif-
icant differences of resistance among cultivars and their offspring have been re-
ported (Chaıb et al., 2007).

16
Sensory methods are using trained panels, normally expressing textural properties
by the use of terms such as fresh, firmness, mealiness, juiciness and skin toughness.
How important the texture is for the quality traits can be divided into three catego-
ries: Critical if the texture determines the price of the commodity; important if the
influence of the texture is significant but not a determinant of the price, and minor
when texture is in principal not considered important at all. For vegetables, the tex-
ture is normally classified as important (Bourne, 2014).

Morphological studies have been carried out using microscopy to evaluate textural
properties in tomato studying cellular structure of the pericarp (Chaıb et al., 2007).

2.2.2 Aroma and flavor

Generally, the flavor is a combination of the perception from retronasal aroma, and
the perception from somatic fibers at the level of the trigeminal nerves and taste,
meaning a combination of volatile and non-volatile compounds (Preedy and
Watson, 2008). Flavor-influencing compounds include sugars (mainly sucrose,
fructose, glucose); acids (mainly citric acid and malic acid) and about 400 volatile
compounds. Vine ripe tomatoes have the highest level of those parameters (Preedy
and Watson, 2008).

Several substances determine the final flavor of tomatoes. A key source is sugar.
Changes in sugar content lead to dramatic changes in flavor, sweetness and sour-
ness. The perception of sweetness can also be altered by reduction or the total ab-
sence of citric and malic acid (Preedy and Watson, 2008). Tomato carotenoids in-
fluence aroma and flavor. The composition of carotenoids is impacting the expres-
sion of aroma volatiles and flavor, while carotenoids in the fruit are degraded. Ly-
copene increase lemon-scented monoterpene aldehyde (Lewinsohn et al., 2005).

Postharvest storage of tomatoes under nitrogen was shown to reduce the amount of
C6 volatile compounds in the stored tomatoes, as did storage of the tomatoes at low
temperature. Returning the tomatoes to normal conditions, did not result in a full
recovery of the content of volatile compounds (Boukobza and Taylor, 2002).

17
2.2.3 Total soluble solids and titratable acidity

Total soluble solids (TSS) is mainly composed of a blend of sucrose, hexose, citrate
and malate that all together reach 78% of the total content. The remaining 22% rep-
resents minor solids such as minerals, phenols, soluble pectins, amino acids and
ascorbic acid (Beckles, 2012). To evaluate this parameter it is necessary to use a
refractometer, which is used worldwide due to its accessibility and adequate preci-
sion for commercial purposes (Malundo et al., 1995).

Acidity corresponds to the amount of acids in a solution. Titratable acidity (TA) is

a type of measurement that deals with total acid concentration in any food. The core
unit of acidity is hydrogen ions (H+) (Nielsen, 2017). In fruits juice titration is done
with an alkaline solution until reaching a pH of 8.1(Wills et al., 1990).
.
TSS is generally low in tomato, i.e. less than 5% for green tomatoes. When the
tomato turns red, TSS increases to around 7%. Contrarily, acidity in tomato is high
(0.9 g/100 g fresh tomato) at the green stage, but decrease to 0.4 g/100 g fresh to-
mato or less for red tomatoes (Serrano et al., 2005). The TSS:TA ratio can be used
to predict the sensory taste perception of tomato (Beckles, 2012).

TSS and TA are thus important quality factors to include in breeding programs of
tomatoes, although their measurements need to be carefully evaluated to correspond
to consumers’ desires and to not be biased by the design of the sensory tests
(Mattheis and Fellman, 1999). Furthermore, a high TSS content in tomato is often
associated with reduced yields (Stevens and Allen Rick, 1986), and with low quality
appearance of the tomatoes under field conditions. The TSS values were also found
to decrease under excessive irrigation (Lahoz et al., 2016) which might be related
to the negative correlation between tomato size and TTS content, also being present
between tomato size and sweetness (Beckles, 2012).

2.2.4 Enzyme activity

The main enzymes in tomato being involved in break-down of pectins; (i.e. pecto-
lytic enzymes or pectinases), are polygalacturonase, pectin methylesterase (PME)
and beta galactosidase (Van Dijk et al., 2006). Polygalacturonase is hydrolyzing the
alpha 1,4 glycosidic bonds of the galacturonic acids during the ripening process of

18
tomatoes (Fischer and Bennett, 1991). Thus, the polygalacturonase activity was pre-
viously thought of being the solely responsible factor of the softening of tomatoes
during ripening due to degradation of the pectin although studies on transgenic to-
matoes have shown this not being the case (Brummell and Harpster, 2001). The
PME activity was shown to influence the pectin metabolism and was correlated to
the content of total soluble solid contents in tomato (Tieman et al., 1992). A decay
of PME activity was seen, exponentially over time, although low activity in trans-
genic tomatoes resulted in loss of tissue integrity with no change in fruit firmness
(Tieman and Handa, 1994, Van Dijk et al., 2006).

2.2.5 Volatile organic compounds

Fruits normally contain a large amount of different volatile compounds that are re-
sponsible for the aroma and flavor of the fruit. Thus, a large amount of the different
volatiles are synthesized during the ripening phase of the fruit and the content and
composition of the volatiles are known to change with maturity stages of the fruit
(Baldassarre et al., 2015). Volatile compounds are known to be part of the defense
mechanisms against herbivores in tomatoes. Furthermore, damages by insects or by
mechanical wounding produced changes in the composition and content of volatile
organic compounds. (Raghava et al., 2010). Also, plants under attack of insects have
been found to show higher concentration of volatile compounds compared with con-
trols (Lara et al., 2015). Thus, a tomato plants attacked by Tuta absoluta expelled
an increased amount of volatile compounds (76% to 86%), e.g. β-caryophyllene, ɑ-
pinene and ɑ-phellandrene, and these compounds were found to attract Macrolophus
pygmaeus, a natural enemy of the attacking insects (Lara et al., 2015).

Presence, amount and composition of volatiles compounds in tomato are affected

by growing conditions. Low irrigation levels were found to be positively correlated
with quality of tomatoes, with an increase in soluble solids and volatile compounds,
resulting in improved aroma and lycopene content. However, the low irrigation lev-
els also reduced the yield of production with 16.4% compared to the control (Lahoz
et al., 2016). A treatment with HIPEF (high-intensity-pulsed electric field) on the
tomatoes was found to preserve flavor quality and stability of the fruits (Aguilo-
Aguayo et al., 2010). Furthermore, a treatment with the biocontrol agent, Bacillus
amylo-liquefaciens strain T5, on tomato plants reduced the attack of Ralstonia sol-
anacearum with approximately 85%. This biocontrol treatment promoted the pro-
duction of 13 different volatile compounds when the plant was under attack and
among these 10% had antibacterial activity (Raza et al., 2016).

19
When the tomato is ripe, the role of volatile compounds changes, from a protection
role towards a role of attraction towards insects and other animals that can contribute
with the dispersal of the tomatoes seeds. The main changes that occur in tomato are
in terpenoids, aldehydes, ketones, carotenoids and lipids (Nath et al., 2014 ,
Baldassarre et al., 2015).

The starting molecules for biosynthesis of volatiles in tomato are proteins, carbohy-
drates and lipids. When the starting molecule is a protein, the resulting volatiles are
aldehydes, alcohols, aromatic amino acids, and keto acids, while lipids and carbo-
hydrates results in alcohols, esters, aldehydes, lactones and ketones (Preedy and
Watson, 2008). The most common volatile compounds of three cultivars are pre-
sented in Table 2, but large influence of the cultivar on the quantity of the volatile
compounds has been reported (Preedy and Watson, 2008).

Table 2 Volatile compounds quantified in three tomato varieties (Preedy & Watson, 2008).

Volatile Centera Centera Zucker Zucker Lucy Lucy

compound (average) (%SD) (average) (%SD) (average) (%SD)

3-methyl-butanal 24.6 26.6 92 16.8 179.2 14.2

2-methyl-butanal 10.9 47.7 65.8 5.5 108 16.1

pent-1-en-3-one 155.7 17.8 295.3 24.1 202.6 4.5

pentanal 47.8 21 52.4 14.3 38.1 4

pentan-3-one 33.9 6.5 48 10.8 33.8 8.6

2-methylbut-2-enal 44.9 64.1 246.1 12.3 349.8 7.6

* Values are the average of three replicates in µg/kg fruit and the standard deviation in percentage.

2.2.6 Color

An important feature for a plant is to successfully spreading its seeds. Therefore,

various spreading mechanisms have evolved, including colors and other sensory
traits attracting animals in order to get assistance in spreading of the seeds in nature
(Hiwasa and Ezura, 2014).

20
Certain theories are stressing that food attributes have resulted in humans choosing
food with attractive colors, due to these often being associated with sensory attrib-
utes and maturity stages (Francis, 1995). Based on consumer’s preferences, the
United States Department of Agriculture (USDA) has selected four factors in toma-
toes to determine their quality choice: color 30%, the absence of defects 30%,
weight 20% and wholeness 20% (Gould, 1992). Color of the tomatoes has been
found to influence perception of also other traits in tomatoes when untrained panel-
ists are used to rank the quality of tomatoes. In general, a positive perception of
color resulted in high perception of also other traits such as general appearance,
flavor, aroma, and texture (Stommel et al., 2005 ). For the above reasons, it can be
concluded that consumers use color as a major factor in their decision to which type
of tomato to procure (López Camelo and Gómez, 2004).

Due to the importance of color for human perception of tomatoes, physiological

processes behind the formation of color have reached increasing attention from the
research society. The visible changes of color in tomatoes goes from green to bright
colors as the tomato matures (López Camelo and Gómez, 2004, Paliyath et al.,
2008). The ripening of the tomato can be seen as a combined physiological and
chemical process that occurs in the fruit (Paliyath et al., 2008).

Abscisic acid (ABA) and ethylene are the two main plant hormones involved in the
ripening process of the tomato plants.(Hiwasa and Ezura, 2014, Zhang et al., 2009).
Since tomato is classified as a climacteric plant, the ripening process starts with an
abrupt increment of ethylene production resulting in a dramatic change of color,
normally from green to red.(Hiwasa and Ezura, 2014, Zhang et al., 2009). In addi-
tion, ethylene is accelerating the ripening process of the fruits, simultaneously hav-
ing a negative impact on the shelf life of the tomatoes (Hiwasa and Ezura, 2014).
External applications of ABA have been used to accelerate fruit coloration and sof-
tening (Zhang et al., 2009). Furthermore, the application of an external inhibitor of
ABA (i.e. fluridone or nordihydroguaiaretic acid (NDGA) was found to delay fruit
ripening through the inhibition of ABA production (Zhang et al., 2009).

Besides the plant hormones, temperature is also known to influence the rate of
changes of color in tomato. The development rate of red color in tomato is increas-
ing as the temperature increases until 30°C, while higher temperatures are known to
inhibit red color development (Nunes, 2008). The physiological explanation of the
temperature effect on color change in tomato is reported as an activation and deac-
tivation of certain proteins involved in ripening processes (Neta-Sharir et al., 2005).
One type of such proteins are known as heat shock proteins (HSP), being activated
under high temperatures, thereby accelerating the ripening process resulting in a

21
skin color change. The specific group of HSP in tomato is designated HSP21 (Neta-
Sharir et al., 2005).

Comparing wild type tomatoes with genetically modified ones accumulating a

higher level of HSP21 protein, showed an increased expression of HSP21 in the
modified tomato at room temperature compared to the wild type tomatoes and the
modified type produced HSP21 even at cold storage (2 ºC). The transformed toma-
toes developed an orange-red color with an increase of carotenoids after 9 days,
despite the fact that they exhibited severe cold injury. Control tomatoes neither dis-
played changes of color, nor carotenoid accumulation, in these cold conditions
(Neta-Sharir et al., 2005). The transformed tomatoes were also found to change from
the green stage to the turning stage 7 to 10 days earlier than the control group. How-
ever, at similar maturity stage, no differences in carotenoid concentration between
the two tomato types were recorded. Thus, a role of HSP21 to promote carotenoids
accumulation during fruit maturation was postulated (Neta-Sharir et al., 2005).

The development of color in the tomato fruit during ripening is mainly due to the
synthesis of various types of carotenoid pigments, particularly lycopene (90%) and
β-carotene (5 to 10%), but also of minor carotenoids, such as lutein and phytoene
(Mikkelsen, 2005, Paliyath et al., 2008, Serino et al., 2009). The composition of the
carotenoids in the tomato fruit determines the color of the tomato fruits (Ballester et
al., 2010). Thus, the change of color in the tomatoes has been highly correlated with
the content of lycopene in the red fruit (Stommel et al., 2005 , Preedy and Watson,
2008, Hwang et al., 2016). The β- carotene content is the major determinant of the
orange color of tomatoes (Hwang et al., 2016). Pink tomatoes have a transparent
peel and is the result of a shortage of naringenin and chalcone flavonoids (Ballester
et al., 2010).

Color measurements to understand the skin pigmentation in tomatoes is used as an

objective indicator to validate the ripening stage of the tomato fruits. Based on pre-
sent knowledge it is recommended to validate the color maturity index relationship
for each cultivar to succeed in harvesting tomatoes at the optimum level of quality
(Beckles, 2012). Traditionally color charts have been applied to determine ripening
in tomatoes (López Camelo and Gómez, 2004). However, modern instrument has
been introduced and nowadays mainly colorimeters are used to measure color on a
numerical base applying the Commission Internationale d’Eclairage L*a*b*
(CIELAB) color sphere. CIELAB express color in three coordinates, where L de-
scribes the brightness through a scale from white to black, a* measures colors from
green (-a*) to red (+a*) and b* those from blue (-b*) to yellow (+b*). For tomatoes
the a* is the most representative parameter as a negative value represent green color

22
and an immature tomato, while a positive value rep-resent red color and a mature
tomato (Preedy and Watson, 2008). In one investigation, the three color parameters,
L*, a* and b*, showed significant changes for tomatoes during ripening, although,
some commonly used color components such as chroma (intensity of color), value
(darkness of lightness) and hue angle (certain angle in a 360º wheel color), calcu-
lated from these parameters, was not suitable for determination of tomatoes ripeness
(López Camelo and Gómez, 2004).

Changes of tomato color during ripening and carotenoids content at similar maturity
stages have been found highly associated. At the breaker stage of tomato, the a* and
b* values are null or negative and carotenoids contents also remain low. At mature
red stage, in the tomatoes collected in the field, the a* values increased and β caro-
tene and lycopene synthesis also increased. Finally in ripe red tomatoes, the a* val-
ues reached their maximum levels simultaneously as the β-carotene synthesis cease.
At this point of ripening, lycopene represent 95% of the carotenoids in all colored
tomatoes (Nunes, 2008).

23
3 Factors affecting quality parameters

3.1 Effects of environmental conditions on fruit quality

during development of fruits

3.1.1 Temperature

Temperature affects directly the final size and color of the tomato. The temperature
has a large impact on production and accumulation of energy through photosynthe-
sis in the tomato plant that is later translocated to the developing fruits. To reach
optimal development of the tomato plant and its fruits, different temperatures are
recommended during various stages of the plant development. At the germination
stage, the most suitable temperature is 16 ºC to 29ºC, while at the growing stage,
the optimal temperature is 18ºC to 28ºC and at fruit set stage it is 14ºC to 24ºC
(Jones, 2007).

The highest lycopene and red color accumulation was seen at 24ºC during the day
and 14 ºC during the night (Jones, 2007). When the temperature range is between
26º C and 30º C, the TTS content increases due to a switch in enzyme activity and
tomatoes also become sweeter (Beckles, 2012). Under Mediterranean conditions,
temperature has been shown as a limiting factor for fruit set and yield, and the upper
limit being 25ºC to 26ºC (Harel et al., 2014).

In addition, if plants are exposed to temperatures above 30º C, this will induce high
evapotranspiration rate and much less carbohydrate biosynthesis will be dedicated

24
for assimilation and sweetness (Beckles, 2012). Therefore, an inhibition of lycopene
biosynthesis will occur, promoting a change in color of fruits turning to yellow or
orange (Jones, 2007).

3.1.2 Relative Humidity

Relative humidity influences the performance of the tomato plants and fruits. It
should be between 30 to 90% (Schwarz et al., 2014) A low relative humidity results
in firmer and juicier fruits with a minimum of physical disorders (Xu et al., 2007).
Previously, Blossom end rot (BER) has been considered to result from calcium de-
ficiency, however, abiotic stress such as very low relative humidity cause high tran-
spiration rates and accumulation of calcium in the mature leaves, which leads an
increase of reactive oxygen leading a risk of fruit disorders such as BER, where the
end part of a fruit becomes darkens, flattens out and eventually burn; resulting in
cracking (scarfs on tomato skin) and presence of oidium, thrips, spider and mites
(Saure, 2014).

A high relative humidity instead promotes low transpiration rates, thereby inducing
early growth of leaves with less area due to an increase of stomatal conductance and
low calcium accumulation. The latter conditions increase the risk of diseases such
as botrytis, mildew, leaf mold (Atherton and Rudich, 2012, Heuvelink and Dorais,
2005, Jones Jr, 2007).

What is to be considered as ideal humidity conditions change due to the develop-

ment stages. At germination stage, 75 to 88% is desirable; at the seedling stage, 71
to 80% is preferable and from vegetative to termination stages 60 to 80% is benefi-
cial (Jones Jr, 2007). To control humidity, growers use heat and ventilation in con-
trolled environments (Heuvelink and Dorais, 2005) and in arid climates evaporative
cooling is used to control the relative humidity (Abdel-Ghany, 2006). Therefore, a
relative humidity above 85% combined with high temperature over 30º C become
critical for the fruit set. However, to reduce high relative humidity it is recom-
mended to ventilating the greenhouse (Heuvelink and Dorais, 2005).

3.1.3 Light

Light influences performance of tomato plants and fruits, resulting in various quality
of tomatoes produced in different seasons. Tomatoes produced during the winter are

25
known to have low TTS compared with tomatoes produced in summer. In addition,
tomato fruits under indoor production conditions have lower TTS due to poor light
quality as compared with out-door production (Beckles, 2012).

UV light has been shown to influence accumulation of total polyphenol contents in

tomatoes. Coffeic acid, ferulic acid and p-coumaric acid contents were found to be
about 20% higher in tomatoes grown with UV light (290 to 400 nm) as compared
with samples grown under UV restriction (below 380 nm) (Luthria et al., 2006).

3.1.4 Soil nutrients

The tomato plant has a high demand of nutrients. Nitrogen, phosphorus and potas-
sium are the major macronutrients used at tomato plant cultivation. Excess or defi-
ciency of one or more of these nutrients are known to impact negatively on fruit
quality (Beckles, 2012).

Macronutrients: Nitrogen, phosphorus and potassium

According to previous studies, N supply has been found to affect TSS in tomato,
although various impacts have been reported. Thus, reduction of nitrate from 12
mM to 4 mM was found to increase TSS concentration (mainly sucrose, fructose
and glucose), reduce acidity, while yield was not affected. However, studies in
cherry tomato treated with 0 to 36 mM of nitrate, showed instead a high and positive
correlation between nitrate addition and TTS concentration and here nitrate concen-
trations above 9 mM TSS also affected negatively the yield (Beckles, 2012).

Effects of potassium and phosphorus have been also studied. High concentrations
of potassium lead to an increase in TSS concentration. However, this increase may
also be related to an increase in electrical conductivity (EC), due to the fact that
concentration of these nutrients, soil water content and salinity is interacting
(Beckles, 2012). In addition, type of nutrient source, e.g. liquid or solid fertilizer,
may impact absorption by plants (Beckles, 2012). Total amount of carotenoids, and
in particular of lycopene in tomato fruit, is generally positively correlated to the
amounts of potassium available for the tomato plant (Mikkelsen, 2005).

Salinity

26
Salinity in a range of 2.5 to 14 dS m-1 electrical conductivity (EC) has a positive
impact of sugar contents of tomato fruits (Cuartero and Fernández-Muñoz, 1998).
Manipulation of salinity affects the internal metabolism of the tomato plant by im-
pacting the length of time for development and maturation, Thus, increase in salinity
to a certain degree leads to a higher TSS (Beckles, 2012). However, this change has
a yield cost of 10% of reduction for each 1 dS m-1, from 8 dS m-1to 14 dS m-1, and
a higher concentration of salts (EC of 15 dS m-1) reduced the number of fruits as
well (Cuartero and Fernández-Muñoz, 1998).

Nutrient deficiency

Nutrient deficiency in the soil severely reduces tomato quality and yield. Lack of
major nutrients, such as potassium, causes non-uniform ripening color of the tomato
fruit (Haifa, 2014). Furthermore, an increase of photosynthesis was associated with
potassium deficiency, and symptoms such as diameter expansion of the stem could
be detected, although such changes were only noticed late in the plant development
(Kanai et al., 2011) Calcium applications have been found increasing firmness of
the tomato (Magee et al., 2003).

Several studies have compared the performance of organic and inorganic production
systems in order to determine how these affect quality parameters. In general, con-
tradictory results have been presented as to how different production systems are
affecting the TSS. In order to elucidate effects on nutrient contents in the different
systems, further studies are needed (Beckles, 2012).

3.1.5 Size

The size of the tomatoes depends on the genetic background, the cultivation location
of the plant and on cultural practices. During fruit set, cultural practices such as
pruning have a direct influence on the tomato fruit size. Through the reduction of
the number of buds, the number of fruits is reduced, and therefore an increase of
size of the tomatoes left on the plant is likely the result (Beckles, 2012).

3.2 Current post harvest management of tomato

27
 A variety of methods are used to extend the shelf life of tomatoes (Preedy and
Watson, 2008). One major opportunity to impact both edibility and marketability of
the tomatoes is to control the synthesis and action of the plant hormone ethylene.
Methods used to control this hormone includes application of innocuous products
on the surface of the tomatoes, decrease of storage temperature, application of mod-
ified atmosphere packing or combination of these methods (Gross et al., 2016).

3.2.1 Temperature treatments and their implication on shelf life,

appearance, color, firmness, chilling injuries and bioactive com-
pounds

Low temperature treatment of tomatoes contribute to a reduction of the ripening

processes, thereby slowing down changes in texture, aroma, flavor and nutrition
contents, and stopping changes of color. After removing tomatoes from low tem-
perature treatment, light red tomatoes are able to change color to dark deep red
(Paull, 1999). However, immature tomatoes are more sensitive than mature ones to
low temperatures. Thus, immature tomatoes might be harmed by low temperatures
and develop chilling injuries such as decrease of volatile contents, high increase of
softness as a result of loss of cell turgor, and abnormal ripening progress resulting
in non-uniform red color in full ripening stage (Nunes, 2008, Passam et al., 2007).
Besides chilling injuries, tomatoes treated with low temperatures develop less in-
tense color as compared with fully ripe tomatoes in the field (Nunes, 2008).

One opportunity is to combine different temperatures during storage to enhance

quality. Thus, treatment of fully ripe tomatoes with low temperature of 2 ºC to 5 °C
for short periods, before they are transferred to a temperature of 20 ºC, results in an
increase in shelf life for the tomatoes due to a decrease in natural decay processes
of the fruit. Combining temperatures of 9 °C during six days and then change to 20
°C for one day, prevented against pitting in tomato (Nunes, 2008).

Shock temperature treatments have also been used in tomatoes to increase resistance
towards pests and decrease the chilling injuries from later cold storage. One example
of such treatment is a hot water dipping in 52 °C for one minute, which was found
to prevent against chilling injury and botrytis attacks (Fallik et al., 2002). In another
study, a short heat treatment before low temperature storage reduced fruit cracking
under modified atmosphere packing (MAP) and delayed color development in
cherry tomatoes (Ali et al., 2004). However, a too high temperature, or duration of

28
the heat treatment, results in heat injury, with irregular color during ripening, devel-
opment of decay, brown spots and cracking. Tomatoes exposed to 38 °C for 24 hours
showed severe injuries (Nunes, 2008).

3.2.2 Application of organic substances after harvest to improve or extend

shelf life of tomato

Applications of organic compound, such as methyl jasmonate and methyl salicylate,

reduced chilling injury and promoted the gene expression of heat shock proteins.
Both compounds are also related to plant defense response (Ament et al., 2010).

Coating gum arabic on green mature tomatoes increased shelf life up to 20 days by
reducing ripening rate at 20ºC and also maintained the overall quality (Ali et al.,
2010). Amino acid aminoethoxyvinylglycine (AVG) treatment at 30 kPa vacuum
infiltration enhanced firmness and increased shelf life of tomato fruit to 20 days at
12 ºC (Candir et al., 2017). On the other hand, other compounds promote accelera-
tion of ripening, such as application of melatonin (50 uM) in a postharvest treatment,
which increased lycopene level as compared with a control group (Sun et al., 2015).

3.2.3 Application of inorganic substances after harvest to extend shelf life

of tomato

The most common pathogens attacking tomato after harvest are (Gross et al.,
2016):
 Alternaria. alternata
 Colletotrichum spp
 Clavibacter. michiganensis
 Pseudomonas syringae
 Xanthomonas campestris
 Fusarium spp,
 Botrytis cinerea
 Phytophthora. infestans
 Phoma lycopersici
 Phomopsis spp
 Phytophthora spp
 Stemphylium herbarum
 Rhizopus spp

29
  Sclerotium. rolfssi
 Rhizoctonia. solani
 Geotrichum candidum
 Sclerotinia spp.

To prevent against attack from these pathogens and thereby preserve the quality of
the fruits, inorganic substances e.g. chlorine in a concentration of less than 100 ppm,
are used during the packing sanitization process. Chlorine eliminates certain gram-
positive and gram-negative bacteria, yeast and molds, as well as spores and viruses
(Gross et al., 2016).

Currently, research on tomato is based on maturity stages to apply different strate-

gies to extend the shelf life of this commodity. Maturity stages are characterized as
follow (Zhang et al., 2009).

Stage 1: mature green (approximately 40 days after anthesis,

Stage 2: breaker 1 (approximately. 44 days after anthesis)
Stage 3 breaker 2 (approximately 45 days after anthesis)
Stage 4 Turing (approximately 46 days after anthesis)
Stage 5 Pink (approximately 50 days after anthesis)
Stage 6 Mature red (approximately 53 days after anthesis).

A short application of calcium chloride at 1%, combined with boric acid at 0.1%,
on mature green tomatoes then packed in polyethylene film (0.44mm thickness) was
found to slow down the ripening rate, maintain firmness, weight, color, and increase
TSS and TA. Thereby, this treatment extended the shelf life of the tomatoes to 72
days while stored at 28º to 30ºC (Sammi and Masud, 2009).

Active paraffin-based paper packing combined with Barck cinnamon essential oil
(6% w/w) promoted a total inhibition of Alternaria alternata in cherry tomatoes. In
the same study, other parameters such as color and weight did not differ as compared
with the control (Rodriguez-Lafuente et al., 2010).

Granular active carbon combined with palladium (1%) in modified asthenosphere

packaging (MAP) and cold storage, reduced softness, weight loss, delayed tomato
decay and reduced expression of 23 volatile compounds as compared with the con-
trol. In addition, when the packages were opened, the treated tomatoes showed im-
proved odor intensity compared with the control, as ranked by panelists. However,
the use of palladium has been reported to significantly increase allergic reactions in
sensitive humans. Therefore, granular active carbon under MAP conditions have

30
also been evaluated, excluding the treatment with palladium, resulting in only 65%
of decay compared with 100% in the control tomatoes after 28 days. Thus, granular
active carbon without palladium is a suitable method to extend shelf life of tomato
(Bailén et al., 2006, Kielhorn et al., 2002 ).

Artificial applications of organic molecules have diverse effects on postharvest fruit

physiology. Ethylene is an organic compound formed naturally as a plant hormone
in tomato. It is responsible for major changes in physical, chemical, physiological
and biological properties. Those changes affect sensory traits in tomato during the
ripening process, independent if the fruit is attached or not to the plant. Ethylene is
known to affect the firmness, flavor, color and bioactive compound levels (Preedy
and Watson, 2008). Since ethylene accelerate ripening, an inhibitor called 1–
methylcyclopropene (1-MCP) has been tested in order to delay ethylene effects on
tomato fruits. One result from such an evaluation of the use of 1-MCP showed a
change in lycopene concentration at the break stage (Opiyo and Ying, 2005). The
use of 1 µL L-1 1-MCP for 24 hours, delayed fruit ripening, reduced respiration rate
and inhibited ethylene production after 20 days at 20ºC and 85 to 95% of RH (Wang
et al., 2010a).

3.2.4 UV light application on tomato and impact on shelf life

During the last years, use of ultraviolet (UV) light has become an attractive alterna-
tive method to improve tomato quality and shelf life. The suggested reason for the
positive effects reported from this treatment is that it triggers biochemical defense
reactions in the tomatoes, but also that fruits and vegetables are sanitized by the use
of UV light (Ribeiro et al., 2012). The use of Chryptococcus laurentii combined
with UV-C was found to eliminate infection of Botrytis cinerea and Alternaria al-
ternata (Zhang et al., 2013). Further, the same doses of UV-C application delayed
tomato softening due an inhibition of ethylene production in mature green cherry
tomatoes (Bu et al., 2013). Other parameters such as color, pH and Brix did not
show any differences as compared with the control, when UV-light was applied
(Pataro et al., 2015).

Molecular studies performed to understand the genetic molecular mechanism from

application of UV- C revealed that 274 genes were upregulated and 403 were down-
regulated. Upregulated genes were responsible to target the defense response sys-
tem, signal transduction and metabolism, and downregulated genes were involved

31
in photosynthesis, lipid metabolism and cell wall disassembly. These results may
explain the increased disease resistance in treated tomatoes (Liu et al., 2012).

3.2.5 Application of modified or controlled atmosphere using carbon

dioxide, oxygen, nitrogen and ozone treatment and impact on shelf
life

Modified atmosphere (MA) or controlled atmosphere (CA) were both shown to re-
duce ethylene concentration and thereby retard maturation of tomato fruits. Gener-
ally, increased carbon dioxide concentration and decreased oxygen concentration
results in a reduction in respiration rate and ethylene synthesis. The most suitable
levels to extend shelf life of fresh tomatoes are 3-5% oxygen, 1-5% carbon dioxide
and 94-96% nitrogen. Such a modified atmosphere does not affect the TSS content
(Beckles, 2012).

Modified atmosphere packing (MAP) was adopted to ensure modified atmosphere

conditions in smaller scale. Different types of low density polyethylene films are
applied for the MAP, which then constitute a barrier between the general atmos-
phere and the atmosphere in the package. This barrier is made of different types of
low density polyethylene package (Beckles, 2012). Generally, MAP creates an arti-
ficial atmosphere and permits a moderate gas exchange with the outer atmosphere.
By the use of MAP, shelf life of tomatoes increased through a retarded respiration
processes, reduced water loss, less disease attacks, and less bruising (Beckles,
2012).

The use of ozone enriched atmosphere resulted in tomatoes with increased firmness
(Tzortzakis et al., 2007, Rodoni et al., 2010), and additionally the tomatoes have
been judged sweeter as compared to control tomatoes (Beckles, 2012). This treat-
ment did not affect weight loss, antioxidant status, ethylene production and vitamin
C concentration (Tzortzakis et al., 2007). Ozone treatments were also shown to re-
sult in reduced attacks of microorganisms (Beckles, 2012), decreased activity of the
pectin-methyl esterase (PME) enzyme, weight loss and fruit damage as compared
to the control (Rodoni et al., 2010). In addition, ozone reduced solubilization and
depolymerization of pectin polysaccharides (Rodoni et al., 2010).

32
3.2.6 Mechanical technologies in postharvest management of tomato

Novel methods are emerging to be used on tomatoes to increase quality and shelf-
life, e.g. inert treatments such as ultrasounds and thermosonication (Pinheiro et al.,
2016). The use of ultrasound was found to change the microstructure of epidermis,
resulting in brighter and a more orange color of the treated tomatoes (Fava et al.,
2017).

3.3 Effects of postharvest techniques on fruit quality traits

Quite a number of studies have been carried out to understand how post-harvest
conditions are affecting the tomato fruit quality and how the quality can be improved
through different treatments. Below effects of post-harvest treatment on various pa-
rameters are described.

3.3.1 Texture

As mentioned above, a combined treatment applying calcium chloride at 1%, boric

acid at 0.1% and 400 ppm of potassium permanganate on green tomatoes, followed
by packing in polyethylene film, increased shelf life up 32 days and improved and
overall quality characteristics of tomatoes, but this treatment also improved the tex-
ture (Sammi and Masud, 2009). Also, treatment with 7.5 ppm of potassium perman-
ganate preserved texture at 14 to 18 ºC after 21 days (Wabali et al., 2017). Another
study showed that pulse light treatment at 4, 6, and 8 J/ cm2 fluence time did not
affect the texture after 10 days of storing although microbial growth was reduced
(Valdivia-Nájar et al., 2018). Furthermore, hot air treatment during 12 hours at 38ºC
on mature green tomatoes delayed firmness decline, due to slowdown of pectin deg-
radation (Wei et al., 2018).

3.3.2 Flavor and volatile compounds

For postharvest storage of tomatoes, a common method applied is the use of modi-
fied ambient conditions which greatly affect flavor and volatile compounds. Storage
of tomatoes under nitrogen was shown to reduce the amount of C6 volatile com-
pounds in the stored tomatoes, as did storage of the tomatoes at low temperature, 6

33
°C for three days. Returning the tomatoes to normal conditions, did not result in a
full recovery of the content of volatile compounds (Boukobza and Taylor, 2002).
Refrigeration or short treatment with high temperature affected negatively and irre-
versibly the content of volatile compounds (Nunes, 2008). However, according to
recommendations from University of California, chilling injury occurs at tempera-
tures below 10 ºC if held longer than two weeks, and recommended temperatures
are between 7 Cº and 15ºC depending of maturity stage and length of storage time
(Suslow, 1997)

A continuous treatment of green tomatoes with 3% to 5% of oxygen and 0% of

carbon dioxide at 13 ºC for 6 weeks resulted in more acid flavor of the tomatoes
(Parsons et al., 1970). Flavor volatiles are generally affected by storage temperatures
due to the alteration of the volatile synthesis. The content is reduced at low temper-
atures, while partly recovered after transfer to 20 ºC, although the flavor is still re-
duced (Zhang et al., 2016). Coating tomatoes with arabic gum or beeswax plus so-
dium bicarbonate at 2%, resulted in maintained flavor under low temperatures
(Fagundes et al., 2015). Also, the above mentioned application of a solution of boric
acid at 0.1% and calcium chloride at 1% on green tomatoes followed by packing
with polyethylene film resulted in a maintained flavor at storage of 28º to 30ºC
(Sammi and Masud, 2009).

3.3.3 Total soluble solids and titratable acidity

Tomatoes ripened on the vine have higher levels of TSS due to a higher importation
of sugars from the plant to the fruit during this stage (Nunes, 2008). Therefore, it is
highly recommended to harvest during the mature green or breaker stage to reduce
the negative impact on TSS and TA and the consequently lower quality and less
pleasant taste (Beckles, 2012).

Both TSS and TA were found increased after a heat pulse treatment between 37°C
to 42 °C, followed by 14 °C in storage room conditions (Beckles, 2012). A combi-
nation of continuous oriented polypropylene (OPP 30µm thick) and perforated holes
smaller than 2 mm, preserved better TA, TSS and vitamin C concentration in to-
matoes due to reduction, but not inhibition, of CO 2 levels at 20 ºC and ≤ 90% RH
(D’Aquino et al., 2016). Green mature tomatoes exposed at a concentration of 1 µL
L-1 of 1-MCP for 24 hours, followed by storage at 20 ºC and 85 to 95% of RH for
20 days resulted in maintained firmness, TSS and TA (Wang et al., 2010b). Short
enrichment with ozone enhanced sugar content and acidity (Rodoni et al., 2010)

34
3.3.4 Treatments effecting fruit color during the ripening process

A combination of oxygen (O 2 ) and carbon dioxide (CO 2 ) content can be used to

delay ripening and therefore color change of tomato. For red tomatoes 3% CO 2 and
2% O 2 was reported optimal (Wills, 1998) while for mature green tomatoes 3% to
5% of CO 2 and low O 2 was preferential, extending shelf life with 6 weeks without
any off flavors (Parsons et al., 1970).

Ultrasound was found to change the microstructure of epidermis, resulting in that

the tomatoes became brighter and conferred a more orange color compared to un-
treated tomatoes (Fava et al., 2017).

Phospholipase D is an enzyme that activates the deterioration system of the cell

membrane in tomatoes, thereby contributing to the ripening process. Lysophospha-
tidylethanolamine (LPE) is an inhibitor of this enzyme and can be used to increase
shelf life and preserve quality of tomato (Amalendu et al., 2003).

The plant hormone abscisic acid (ABA) is known to promote ethylene synthesis and
thereby induce ripening, resulting in tomatoes turning red faster. The inhibitors flu-
ridone or nordihydroguaiaretic acid (NGDA) are reducing the ABA production, and
thereby delaying fruit ripening and color development of tomato fruits (Zhang et al.,
2009).

The amino acid aminoethoxyvinylglycine (AVG) suppress ethylene production in

plants, and consequently delays fruit ripening (Boller et al., 1979). Furthermore,
AVG enhances fruit firmness and has thus a potential use in postharvest manage-
ment (Rath and Prentice, 2004). AVG was shown to postpone the natural ripening
process by 10 days through a lowering of ethylene production, lycopene color val-
ues, and a* and C* values (Candir et al., 2017).

Temperature during storage is known to influence the color of the tomatoes. Table
3 summarizes the relationship between storage temperature, numbers of days treated
in a certain temperature, change in color related to human perception, and changes
in color, using a colorimeter in fresh tomatoes.

35
Table 3. Relationships between storage temperature and change in color.

Storage Number of Change Change in color Reference

temperature treatment days in color (colorimeter)
(human perception)

10, 15 or 20ºC Unknown Color increased slightly re- (Proulx et al., 2001)
gardless of the temperature

10, 15 or 20ºC Unknown Change L* decreased during storage (Proulx et al., 2001)

20ºC Unknown From turning to L* decreased gradually from (Nussinovitch et al.,

red 50.8 to 42.7 1996)

20ºC 7 L* value decrease after 7 days (Auerswald et al.,

1999)

20ºC 12 Mature-green to Linear increase in a*/b* (Giovanelli et al., 1999)

full ripe

37ºC 12 Yellowish color Increases in L*, b* and (Gnanasekharan et al.,

chroma values and decreases 1992)
in a* and hue

* Hunter’s values a* b* and L*; color index a*/b* (ratio of red to green component).

3.3.5 Change in nutrients and nutrient contents

Treatments of tomatoes with UV-C, with an average of 4 kJ m−2, increased, up to

20%,the concentrations of phenolic compounds, such as gallic acid, chlorogenic
acid, syringic acid, p-coumaric acid and quercetin; enhanced accumulation of anti-
oxidants; organic acids; ascorbic acid; lycopene and total carotenoids. Furthermore,
a sensory panel gave higher rates on taste and quality of treated tomatoes as com-
pared to control (Beckles, 2012, Liu et al., 2012, Charles et al., 2016). Therefore,
the application of this interval of light has been suggested to enhance accumulation
of healthy compounds in tomatoes (Pataro et al., 2015).

36
Also, methyl jasmonate was found to increase antioxidant levels in tomatoes simul-
taneously as the resistance to diseases during cold storage was promoted (Ding et
al., 2002). An application of 1 MCP for 24 hours followed by storage at 20 ºC for
20 days contributed to increased ascorbic acid and soluble phenolic contents (Wang
et al., 2010a).

Furthermore, ozone enrichment was found to enhance phenolic compounds accu-

mulation and antioxidant capacity in tomatoes (Rodoni et al., 2010). Ultraviolet ra-
diation preserved higher quantities of total phenolic content, tomatoes had lower
water loss and showed a decrease in bacterial presence after treatment (Pinheiro et
al., 2016).

3.4 Reduction of losses

Food loss and waste in the food chain occur during agricultural production, post-
harvest handling, storage, processing, and also during distribution and consumption
(Porat et al., 2018).

3.4.1 Reduction of losses by the use of pre-harvest factors

Variation of environmental conditions, such as soil, climate, season, geographic po-

sition, production in greenhouse or open field under different fertilization systems,
are affecting the tomatoes quality parameters e.g. color, size and weight (Beckles,
2012). To avoid losses in the production, several methods have been developed: e.g.
adequate application of pesticides may reduce postharvest losses (FAO, 1989). In
rainy areas, high furrows are a common practice to avoid over watering and flood-
ing.

A careful and well-planned crop management, including the use of fertigation and
proper watering together with the addition of auxin (a plant regulator) contribute to
an increase in fruit production and fruit size, a reduction of losses and a faster rip-
ening process (Oda and Saito, 2006).

Another common strategy to prevent losses is the use of greenhouse production,

which also satisfies the internal demand for fresh tomato consumption in off season.
Also, in field production, plastic tunnels can be used to control ripening during the
low temperature season. In the greenhouse, forcing, semiforcing, and late raising

37
are commonly applied to induce proper growth and maturity of tomatoes (Oda and
Saito, 2006).

3.4.2 Reduction of losses during postharvest handling

To prevent damage during harvest or transport, a complex combination of technol-

ogy, pest management, marketing and distribution systems are used to reduce post-
harvest losses (FAO, 1981). To reduce losses it is important to understand the dif-
ferent sources of the postharvest losses. For example, a comparison between com-
mercial farmers and subsistence farmers elucidated the main sources of post-harvest
losses; i.e. market price stability (29%), logistic control activity (21%), and quality
control activities (23%). Farmers with advanced logistics, high activity and quality
controls combined with a stable market (70%) showed in general lower postharvest
losses as compared with subsistence farmers who work in vulnerable context, use
basic logistic, perform low quality control and have limited access to the formal
market (only 8%) (Macheka et al., 2018).

Simple practices before transportation e.g. washing, sorting and the use of clean
plastic package were shown to reduce aerobic bacterial count (ABC), coliform bac-
terial count, yeast and mold by 2.51%, 32.7% and 29.86% respectively, as compared
with the control (Khadka et al., 2017). During transport, forced-air cooling (FAC)
or pressure cooling (PC) are the most efficient cooling methods in a dry climate
(Sargent, 1998). In general, most of the existing problems related to postharvest
losses in tomatoes can be handled with the current available technology and infor-
mation (Kader, 2005). Thus, a combination of modified atmosphere packages to-
gether with application of aloe gel were found to increase lycopene, phenolics, and
ascorbic acid contents, while microbial counts were reduced at 16 days of storage.
However, a longer period between harvest and consumption resulted in a reduced
natural flavor of the tomatoes (Mirdehghan and Valero, 2017). Parameters related
to nutritional aspects and flavor have in general shorter durability as compared with
parameters traditionally used to decide shelf life (Kader, 2008).

Regarding socioeconomic aspects, application of postharvest management can have

a positive economic impact on growers, reduce their poverty, and provide food se-
curity and a positive effect on sustainable use of resources (Kader, 2005). National
initiatives organized by public agencies may reduce postharvest losses. For example
in Japan, the Ministry of Agriculture released cultivars resistant to fruit malfor-
mation, strips, and less susceptible to cracking (Oda and Saito, 2006). In addition,

38
the use of high technology during selection and well-designed packaging protecting
the final product, which was combined with an organized logistic solution, including
cold chain management and retail packaging, resulted in improved tomato quality
and reduced losses (Porat et al., 2018).

3.4.3 Biochemical changes caused by mechanical damage

Different pre and post-harvest conditions may cause mechanical damage, but par-
ticularly after harvest large injuries may occur, which lead to reduction in TSS and
TA contents (Beckles, 2012). Causes of the mechanical damage in tomato fruits are
impact damage, compression, abrasion and puncture. Impact damage and compres-
sion cause bruising and cracking damage, abrasion affects visual perception and
color, depending of the maturity stages. Tomatoes may be injured by puncturing
objects such as stems from others tomatoes, perforating skin surface during harvest,
storage or packing (Li and Thomas, 2014). At the cellular level this may result in
cell lysis and several undesirable chemical reactions such as accel-erated transpira-
tion, respiration, ethylene production and pathogen infestation (Beckles, 2012). Me-
chanical harvest increased pectin methylesterase (PME) activity and increasing soft-
ness as compared with hand harvest. Tomatoes exposed to 40º C or higher temper-
atures became more damaged and soft. However, these conditions promoted better
juice consistency after processing as compared with tomatoes stored at lower tem-
peratures (Held et al., 2015).

39
4 Genetic studies in tomato

4.1 Genetic studies related to biodiversity using molecular

markers

Tomato genetic biodiversity studies provide information on the genetic identity,

genetic relationship, genetic diversity and genetic distance among accessions of to-
matoes, information obtained primarily through the use of different types of molec-
ular markers (Parmar et al., 2010). Genetic studies are often applied to discriminate
biodiversity within and between commercial and wild species of tomatoes and their
relatives (Passam et al., 2007). Genetic variation can be seen as a natural variation,
which depends on the presence of different alleles and their frequency (Wray, 2008).
Polymorphism for different alleles has been found to be low in cultivated tomatoes
due to self- pollination and a narrow genetic base. On the contrary, genetic distance
and polymorphism of wild tomato relatives (L. peruvianum, L. pennellii and L. hir-
sutum) have been found high. These wild tomatoes thereby constitute an enormous
source to improve quality traits, as well as biotic and abiotic resistance of cultivated
tomatoes (Passam et al., 2007).

By the use of 165 SSR (simple sequence repeat) markers, four main clusters were
found among eight tomato cultivars, indicating the use of SSR markers being a cost
effective method for diversity evaluation studies (Kwon et al., 2009, Todorovska et
al., 2014). Generally, simple sequence repeats (SSR) and retrotransposon based se-
quence specific amplification polymorphisms (SSAP) have been shown to be more
accurate than randomly amplified polymorphic DNAs (RAPDs) to determine ge-
netic variation in tomatoes (Passam et al., 2007). The use of single nucleotide poly-

40
morphisms markers (SNPs) has shown wild tomatoes having a 20–fold higher var-
iation than cultivated crop accessions, indicating genetic erosion among cultivated
tomatoes. In addition, a correlation between geographical location and growth habit
of the tomatoes has been reported (The Tomato Genome et al., 2012). Currently,
next generation technique (NGT) is reported as the most powerful technique to iden-
tify DNA sequence polymorphisms and create molecular markers to use in plant
breeding programs (Devran et al., 2018). Biodiversity studies within populations is
highly important, providing new information to breeders to develop cultivars with
genetic tolerance or resistance to biotic and abiotic problems in tomato (Parmar et
al., 2010).

4.2 Genetic studies related to quality parameters using

molecular markers

The genetic map of tomatoes is a valuable tool, contributing to opportunities for

identification of chromosomal regions responsible for certain quantitative or quali-
tative traits. The understanding of location of various genes can combined with tra-
ditional plant breeding techniques result in the introduction of desirable character-
istics into commercial cultivars (Devran et al., 2018). Important quality traits of to-
matoes can either be controlled by a single gene or by quantitative trait loci (QTL),
were several minor genes are combined to determine the trait. Domestic tomatoes
have a high variability in fruit shape as compared with the wild types and this trait
has been found controlled by QTLs. Actually, tomato traits like shape, color, size,
ripening, organoleptic quality, pollination, field diseases resistance as well as re-
sistance to abiotic factors, such as chilling temperatures, are all known to be con-
trolled by QTLs (Passam et al., 2007).

4.2.1 Nutrient contents and sensory factors

Wild tomatoes provide a potential resource for breeding of tomatoes with in-
creased nutrient content. The Solanum L. cheesmanii is one species naturally hold-
ing high β-carotene and sugar contents, while the Solanum L. peruvianum Mill is a
source of aromatic fragrance (Passam et al., 2007). A study on recombinant inbred
line (RIL) crossing S. lycopersicum and S. pimpellifolium, demonstrated that both
parents contribute to the genetic variation among the progeny for content of calcium,
copper, iron, magnesium, manganese, sodium, phosphorus sulfur, strontium and

41
zinc meaning that a plant breeding program may use this information to seek to
increase contents of those minerals (Capel et al., 2017).
Studies on heritability can be carried out in tomatoes using Restricted maxi-mum
likelihood estimation (REML), where genetic heritability represents similarities be-
tween parents and their offspring, and a mathematical mixed model with fixed ef-
fects and random effects are used for the calculation (Wray, 2008, Hall and Bush,
2016). A recent study on tomato using REML demonstrated that amino acids were
shown to have higher and stable heritability (0.57) as compared with the heritability
of volatile compounds (0.39) during a period of 2 years. In the same study, the phe-
notypic heritability was found ranging between 0.28 and 0.77 while the genetic her-
itability ranged between 0.11 to 0.82. The genetic and phenotypic heritability were
also found to have low correlation. Using genome-wide association studies, eleven
associated traits of amino acids, acids, sugars and volatile compounds were consist-
ently detected through two years in chromosomes 1,4, 6, 8 and 9 (Bauchet et al.,
2017).

An over-expression of malate in transformed tomato plants was found to affect

a number of post-harvest quality traits, including water loss and wrinkling; TSS
content, and resistance to opportunistic pathogen infection (Centeno et al., 2011).

4.2.2 Texture and yield

A disassembly of polysaccharides leads to fruit softening and specific enzyme ac-

tivities are highly correlated with these changes. Through the use of transgenic to-
mato, a gene related to pectatelyase (PL) was silenced, which resulted in a severe
change in tomato fruit texture and softening of the fruits. PL reduction significantly
increased fruit firmness, maintained the fruit integrity longer than in the control but
showed no effect on yield, weight, ethylene biosynthesis, color or TSS. In addition,
an increase of thickness and density of tricellular junction zones as well as a pro-
longed shelf life was observed while the PL was silenced (Uluisik et al., 2016)).

4.2.3 Color

An impressive amount of diversity in color exists naturally in tomatoes, and some

types of tomatoes even retain their green color, meaning that they never change
color to red and ripe. To understand the genetic background for such differences in
processes is important while designing breeding programs for tomatoes. Through

42
the use of real time PCR to evaluate mutants of tomatoes, changes in enzyme activ-
ity associated with cell wall disassembly have been demonstrated as a reason for the
colorless non-ripening behaviour. Expression of cell wall degrading enzymes such
as polygalacturonase, pectinesterase, galactanase and xyloglucan endotransglyco-
sylase was dramatically reduced in mutants unable to change color and maturate.
On the contrary, higher levels of the enzyme pectinesterase were detected in non-
ripening wild cultivars as compared with the levels in the mutant. Thus, a link be-
tween events during ripening controlling cell separation in cell walls of tomato and
those involved in the formation of dehiscence zones in dry fruits, was postulated
(Eriksson et al., 2004).

The table below presents a summary of tomato genes known associated with color
(Rick, 2017). Some genes, such as the chrysanthemum sterile, are affecting or-
ange, red and brown color. Others genes are known to influence color more specif-
ically. For example, the gene tangerine, on the arm L in chromosome 10, controls
the orange color trait (Rick, 2017). The gene CYC-B (77 SNP) is known to confer
orange color to tomatoes and this gene has also been shown to correlate with the
abundance of β-carotenoids (Hwang et al., 2016).

43
Table 4 List of genes associated with color traits in tomato flesh fruit (Rick, 2017).

Gene Allele Gene name Phenotype

chrs Chrysanthemum ster- Red fruit brown sec-

ile tors

hp2 j High pigment 2 Red fruit

B og Beta carotene Orange

chrs Chrysanthemum ster- Orange

ile

Del Delta Orange

Dps Dispyros Orange

Nr Never ripe Orange

T 2 Tangerine Orange

Vo Virescent Orange Orange

ant Aurantia Orange

aur Aurantiaca Orange

K K Yellow

r 2s Yellowflesh Yellow

mic Minuscula Yellow

abi Aubergine Purple

chrs Chrysanthemum ster- Brown color

ile

gf Green flesh Brown color

dg Dark green Dark green

44
5 General Conclusions
The tomato crop is a highly important horticultural product in the world. The
consumption is global and thanks to advances in research, an increasing number of
places are suitable to produce tomato in open fields or in the greenhouse with dif-
ferent climate conditions. In addition, studies on native germplasm have become
increasingly important, as these are contributing to our understanding of lost quality
traits in commercial cultivars, such as resistance to rough climate conditions. Fur-
ther, he majority of the resistant quality traits also influences the yield negatively.

The main quality parameters, such as textural properties, aroma, volatile com-
pounds, flavor, total soluble solids, titratable acidity, and color are used to classify
tomatoes. Most of these parameters are highly affected by temperature, relative hu-
midity, light, soil nutrients and the size of the cultivar during fruit production. A
deeper understanding of the relation among the mentioned traits and their interaction
with the environment will facilitate the development of methods to preserve or im-
prove the quality parameters of tomatoes.

Postharvest management, such as the application of organic and/or inorganic

substances, or modified atmosphere packing, contributes to a pro-longed shelf life,
prolongs the window for distribution s of quality tomatoes and preserves the quality
parameters of the tomato.

Recent studies have clearly shown the importance of reducing post-harvest

losses in tomatoes. Reasonable living conditions of farmers and use or adaptation of
technology, logistics and preservation of quality parameters in all parts of the chain
from production to consumers are important factors to secure low post-harvest
losses
In conclusion, this introductory paper shows the presence of a large amount of
knowledge, based on traditional as well as on modern and powerful technology. A
huge and deep understanding is available of the tomato plant and its interaction with

45
the environment, nutrient conditions, management and effects on quality traits.
However, there are still important gaps of knowledge of the role of certain second-
ary metabolites and their potential impact on the tomato plant or human health.

46
Acknowledgements

I am very thankful to my supervisors Eva Johansson and Marie Olsson for their full
support and feedback to make this introductory paper. I will never forget for all their
patience and encouraging words, guides and enormous contributions to meet this
goal and allow this work to represent my major interests and solve major questions
on tomato crop.
I also want to thank to my supervisor Alberto Centellas for his suggestions and
his support.
Regarding the technical support, I would like to thank the library of SLU because
of providing the request regarding academic books and access to journals. To the IT
department for their computational support in any time.
I am also thankful to my children Paola and Adrian for all their company, pa-
tience and lovely support all the time.
Finally, I would like to thank Marisa and Emilia, for their enthusiasm and kind-
ness always willing to help in any circumstances.

47
References

ABDEL-GHANY, A. M. K., T. (2006). 2006. Cooling Efficiency of Fogging Systems for

Greenhouses. Biosystems Engineering, , 94, 97-109.
ABUSHITA, A. A., DAOOD, H. G. & BIACS, P. A. 2000. Change in Carotenoids and Antioxidant
Vitamins in Tomato as a Function of Varietal and Technological Factors. Journal of
Agricultural and Food Chemistry, 48, 2075-2081.
AGUILO-AGUAYO, I., SOLIVA FORTUNY, R. & MARTIN-BELLOSO, O. 2010. Volatile,
compounds and changes in flavour-related enzymes during cold storage of high - intensity
pulsed electric field- and heat- processed tomato juices. Journal of the Science of Food and
Agriculture, 90, 1597-1604.
ALI, A., MAQBOOL, M., RAMACHANDRAN, S. & ALDERSON, P. G. 2010. Gum arabic as a
novel edible coating for enhancing shelf-life and improving postharvest quality of tomato
(Solanum lycopersicum L.) fruit. Postharvest Biology and Technology, 58, 42-47.
ALI, M. S., NAKANO, K. & MAEZAWA, S. 2004. Combined effect of heat treatment and modified
atmosphere packaging on the color development of cherry tomato. Postharvest Biology
and Technology, 34, 113-116.
ALTER, C. M. 2009. Informalization of Labour Markets Is Formalization the Answer? In: RAZAVI,
S. (ed.) The Gendered Impacts of Liberalization: Towards "Embedded Liberalism"?
.New York Taylor & Francis.
AMALENDU, C., ARUN, S. M., RAMASWAMY & HOSAHALLI, S. 2003. Handbook of
postharvest technology: cereals, fruits, vegetables, tea, and spices, New York, CRC Press
Taylor and Francis Group.
AMENT, K., KRASIKOV, V., ALLMANN, S., REP, M., TAKKEN, F. L. W. & SCHUURINK, R.
C. 2010. Methyl salicylate production in tomato affects biotic interactions. The Plant
Journal, 62, 124-134.
APEL, W. & BOCK, R. 2009. Enhancement of Carotenoid Biosynthesis in Transplastomic Tomatoes
by Induced Lycopene-to-Provitamin A Conversion. Plant Physiology, 151, 59.
ATHERTON, J. & RUDICH, J. 2012. The tomato crop: a scientific basis for improvement.,
Dordrecht, Springer Netherlands.
AUERSWALD, H., PETERS, P., BRÜCKNER, B., KRUMBEIN, A. & KUCHENBUCH, R. 1999.
Sensory analysis and instrumental measurements of short-term stored tomatoes
(Lycopersicon esculentum Mill.). Postharvest Biology and Technology, 15, 323-334.
AYANDIJI, A. & OMIDIJI, D. 2011. Determinant post harvest losses among tomato farmers in
imeko-afon local government area of agun state, Nigeria. Global Journal of Science
Frontier Reserarch, 11, 23-27.
BAI, Y. & LINDHOUT, P. 2007. Domestication and Breeding of Tomatoes: What have We Gained
and What Can We Gain in the Future? Annals of Botany, 100, 1085-1094.
BAILÉN, G., GUILLÉN, F., CASTILLO, S., SERRANO, M., VALERO, D. & MARTÍNEZ-
ROMERO, D. 2006. Use of Activated Carbon inside Modified Atmosphere Packages To

48
 Maintain Tomato Fruit Quality during Cold Storage. Journal of Agricultural and Food
Chemistry, 54, 2229-2235.
BALDASSARRE, V., CABASSI, G. & SPADAFORA, N. 2015. Wounding tomato fruit elicits
ripening-stage specific changes in gene extression and production of volatile compounds.
Journal of experimental Botani, 66, 1511-1526.
BALLESTER, A.-R., MOLTHOFF, J., DE VOS, R., HEKKERT, B. T. L., ORZAEZ, D.,
FERNÁNDEZ-MORENO, J.-P., TRIPODI, P., GRANDILLO, S., MARTIN, C.,
HELDENS, J., YKEMA, M., GRANELL, A. & BOVY, A. 2010. Biochemical and
Molecular Analysis of Pink Tomatoes: Deregulated Expression of the Gene Encoding
Transcription Factor SlMYB12 Leads to Pink Tomato Fruit Color. Plant Physiology, 152,
71.
BAUCHET, G., GRENIER, S., SAMSON, N., SEGURA, V., KENDE, A., BEEKWILDER, J.,
CANKAR, K., GALLOIS, J. L., GRICOURT, J., BONNET, J., BAXTER, C., GRIVET, L.
& CAUSSE, M. 2017. Identification of major loci and genomic regions controlling acid
and volatile content in tomato fruit: implications for flavor improvement. New Phytologist,
215, 624-641.
BAUCHET, G. & MATHILDE, C. 2012. Genetic Diversity in Tomato (Solanum lycopersicum) and
Its Wild Relatives. In: MATHILDE CAUSSE, E. D. M. Ç. (ed.) Genetic Diversity in
Plants. Rijeka: IntechOpen.
BECKLES, D. M. 2012. Factors affecting the postharvest soluble solids and sugar content of tomato
(Solanum lycopersicum L.) fruit. Postharvest Biology and Technology, 63, 129-140.
BEECHER, G. R. 1998. Nutrient content of tomatoes and tomato products. Proceedings of the
Society for Experimental Biology and Medicine. , 218, 98-100.
BERGOUGNOUX, V. 2014. The history of tomato: From domestication to biopharming. Plant
Biotechnology 2013: "Green for Good II". 32, 170-189.
BIESALSKI, H. K., DRAGSTED LO FAU - ELMADFA, I., ELMADFA I FAU - GROSSKLAUS,
R., GROSSKLAUS R FAU - MULLER, M., MULLER M FAU - SCHRENK, D.,
SCHRENK D FAU - WALTER, P., WALTER P FAU - WEBER, P. & WEBER, P. 2009.
Bioactive compounds: definition and assessment of activity. Nutrition, 25, 1202-5.
BLANCA, J., IZARES, J. N. C., CORDERO, L., PASCUAL, L., DIEZ, M. J. & NUEZ, F. 2012.
Variation Revealed by SNP Genotyping and Morphology Provides Insight into the Origin
of the Tomato. PLoS ONE, 7, e48198.
BOLLER, T., HERNER, R. C. & KENDE, H. 1979. Assay for and enzymatic formation of an
ethylene precursor, 1-aminocyclopropane-1-carboxylic acid. Planta, 145, 293-303.
BOUKOBZA, F. & TAYLOR, A. J. 2002. Effect of postharvest treatment on flavour volatiles of
tomatoes. Postharvest Biology and Technology, 25, 321-331.
BOURNE, M. 2014. Food Texture and Viscosity: Concept and Measurement, Academic Press.
BOVY, A., DE VOS, R., KEMPER, M., SCHIJLEN, E., PERTEJO, M. A., MUIR, S., COLLINS,
G., ROBINSON, S., VERHOEYEN, M., HUGHES, S., SANTOS-BUELGA, C. & VAN
TUNEN, A. 2002. High-Flavonol Tomatoes Resulting from the Heterologous Expression
of the Maize Transcription Factor Genes LC and C1. The Plant Cell, 14, 2509-2526.
BRUMMELL, D. A. & HARPSTER, M. H. 2001. Cell wall metabolism in fruit softening and quality
and its manipulation in transgenic plants. Plant Molecular Biology, 47, 311-339.
BU, J., YU, Y., AISIKAER, G. & YING, T. 2013. Postharvest UV-C irradiation inhibits the
production of ethylene and the activity of cell wall-degrading enzymes during softening of
tomato (Lycopersicon esculentum L.) fruit. Postharvest Biology and Technology, 86, 337-
345.
CAMPBELL, D. T., PRUSSIA, S. E. & SHEWFELT, R. L. 1986. Evaluating Postharvest Injury to
Fresh Market Tomatoes. Journal of Food Distribution Research, September/86, 16-25.
CANDIR, E., CANDIR, A. & SEN, F. 2017. Effects of aminoethoxyvinylglycine treatment by
vacuum infiltration method on postharvest storage and shelf life of tomato fruit.
Postharvest Biology and Technology, 125, 13-25.
CANENE-ADAMS, K., K., C. J., ZARIPHEH, S., JEFFERY, E. H. & ERDMAN, J. W. 2005. The
Tomato As a Functional Food. The Journal of nutrition, 135, 1226-1230.
CAPEL, C., YUSTE-LISBONA, F. J., LOPEZ-CASADO, G., ANGOSTO, T., HEREDIA, A.,
CUARTERO, J., FERNANDEZ-MUNOZ, R., LOZANO, R. & CAPEL, J. 2017. QTL

49
 mapping of fruit mineral contents provides new chances for molecular breeding of tomato
nutritional traits. Theoretical and Applied Genetics, 130, 903-913.
CARBONELL, P., ALONSO, A., GRAU, A., SALINAS, F. J., GARCÍA-MARTÍNEZ, S. & RUIZ,
J. J. 2018. Twenty Years of Tomato Breeding at EPSO-UMH: Transfer Resistance from
Wild Types to Local Landraces—From the First Molecular Markers to Genotyping by
Sequencing (GBS). Diversity, 10, 12.
CENTENO, D. C., OSORIO, S., NUNES-NESI, A., BERTOLO, A. L. F., CARNEIRO, R. T.,
ARAUJO, W. L., STEINHAUSER, M. C., MICHALSKA, J., ROHRMANN, J.,
GEIGENBERGER, P., OLIVER, S. N., STITT, M., CARRARI, F., ROSE, J. K. C. &
FERNIE, A. R. 2011. Malate Plays a Crucial Role in Starch Metabolism, Ripening, and
Soluble Solid Content of Tomato Fruit and Affects Postharvest Softening. Plant Cell, 23,
162-184.
CHAıB, J., DEVAUX, M.-F., GROTTE, M.-G., ROBINI, K., CAUSSE, M., LAHAYE, M. &
MARTY, I. 2007. Physiological relationships among physical, sensory, and morphological
attributes of texture in tomato fruits. Journal of Experimental Botany, 58, 915-1925.
CHARLES, M. T., ARUL, J., CHRLEBOIS, D., YAGANZA, E. S., ROLLAND, D., ROUSSEL, D.
& MERISIER, M. J. 2016. Posthaverst UV-C treatment of tomato fruits: Changes in
simple sugars and organic acids contens during storage. LWT - Food Science and
Technology, 65, 557-564.
COLLE, I., VAN BUGGENHOUT, S., VAN LOEY, A. & HENDRICKX, M. 2010. High pressure
homogenization followed by thermal processing of tomato pulp: Influence on
microstructure and lycopene in vitro bioaccessibility. Food Research International, 43,
2193-2200.
CUARTERO, J. & FERNÁNDEZ-MUÑOZ, R. 1998. Tomato and salinity. Scientia Horticulturae,
78, 83-125.
D’AQUINO, S., MISTRIOTIS, A., BRIASSOULIS, D., DI LORENZO, M. L., MALINCONICO,
M. & PALMA, A. 2016. Influence of modified atmosphere packaging on postharvest
quality of cherry tomatoes held at 20°C. Postharvest Biology and Technology, 115, 103-
112.
DAVIS, A., FISH, W. & PERKINS-VEAZIE, P. 2003. A rapid spectrophotometric method for
analyzing lycopene content in tomato and tomato products. Postharvest Biology and
Technology, 28, 425-430.
DEVRAN, Z., KAHVECI, E., HONG, Y., STUDHOLME, D. J. & TÖR, M. 2018. Identifying
molecular markers suitable for Frl selection in tomato breeding. Theoretical and Applied
Genetics, 131, 2099-2105.
DING, C. K., WANG, C. Y., GROSS, K. C. & SMITH, D. L. 2002. Jasmonate and salicylate induce
the expression of pathogenesis-related-protein genes and increase resistance to chilling
injury in tomato fruit. Planta, 214, 895-901.
EMANA, B., AFARI-SEFA, V., NENGUWO, N., AYANA, A., KEBEDE, D. & MOHAMMED, H.
2017. Characterization of pre- and postharvest losses of tomato supply chain in Ethiopia.
Agriculture & Food Security, 6, 3.
ERIKSSON, E. M., BOVY, A., MANNING, K., HARRISON, L., ANDREWS, J., DE SILVA, J.,
TUCKER, G. A. & SEYMOUR, G. B. 2004. Effect of the Colorless non-ripening mutation
on cell wall biochemistry and gene expression during tomato fruit development and
ripening. Plant Physiol, 136, 4184-97.
FAGUNDES, C., PALOU, L., MONTEIRO, A. R. & PÉREZ-GAGO, M. B. 2015. Hydroxypropyl
methylcellulose-beeswax edible coatings formulated with antifungal food additives to
reduce alternaria black spot and maintain postharvest quality of cold-stored cherry
tomatoes. Scientia Horticulturae, 193, 249-257.
FALLIK, E., ILIC, Z., ALKALAI- TUVIA, S., COPEL, A. & POLEVAYA, Y. 2002. A short hot
water rinsing and brushing reduces chilling injury and enhances resistance against Botrytis
cinerea in fresh harvested tomato. Advances in Horticultural Science, 16, 3-6.
FAO. 1981. Food loss prevention in perishable crops [Online]. Rome-Italy: Food and Agriculture
Organization of the United States. Available:
[Link] [Accessed 2017 08 16 2017].

50
FAO. 1989. Prevention of food losses: fuit, vegetale and root crops: A training manual [Online].
Rome - Italy: Food and Agriculture Organization of the United Nations. Available:
[Link] [Accessed].
FAO. 2016. Food and Agriculture Organization of the United Nations [Online]. Rome-Italy: Food
and Agriculture organiation of the United States. Available:
[Link]/search/tomato%20/E [Accessed].
FAOSTAT. 2013. Crops and livestock products [Online]. Food and Agricultural Organization of the
Unided Nations. Available: [Link] [Accessed].
FAVA, J., NIETO, A., HODARA, K., ALZAMORA, S. M. & CASTRO, M. A. 2017. A Study on
Structure (Micro, Ultra, Nano), Mechanical, and Color Changes of Solanum lycopersicum
L. (Cherry Tomato) Fruits Induced by Hydrogen Peroxide and Ultrasound. Food and
Bioprocess Technology, 10, 1324-1336.
FISCHER, R. L. & BENNETT, A. B. 1991. Role of cell wall hydrolases in fruit ripening. Annual
review of plant biology, 42, 675-703.
FRANCIS, F. J. 1995. Quality as influenced by color. Food Quality and Preference, 6, 149-155.
GARTNER, C., STAHL W FAU - SIES, H. & SIES, H. 1997. Lycopene is more bioavailable from
tomato paste than from fresh tomatoes. Am J Clin Nutr., 66, 116.
GEBHARDT, S. E. & THOMAS, G. R. 2002. Nutrivive value of foods, Beltsbille, Maryland, USDA.
GIOVANELLI, G., LAVELLI, V., PERI, C. & NOBILI, S. 1999. Variation in antioxidant
components of tomato during vine and post-harvest ripening. Journal of the Science of
Food and Agriculture, 79, 1583-1588.
GIOVANNUCCI, E. 1999. Tomatoes, tomato-based products, lycopene, and cancer: review of the
epidemiologic literature. Journal of the national cancer institute, 91, 317-331.
GNANASEKHARAN, V., SHEWFELT, R. L. & CHINNAN, M. S. 1992. Detection of Color
Changes in Green Vegetables. Journal of Food Science, 57, 149-154.
GOULD, W. 1992. Tomato production, processing and technology. , Cambridge: Woodhead
Publishing Ltd.
GROSS, K. C., WANG, C. Y. & SALVEIT, M. 2016. The Commercial Storage of Fruits,
Vegetables, and Florist and Nursery Stocks Beltsville Meryland, United States Department
of Agriculture.
HAIFA 2014. Tomato crop guide: Nutrients deficiency symptoms. [Link]
[Link]/knowledge_center/crop_guides/tomato/plant_nutrition/nutrient_deficiency_sym
ptoms/.
HALL, J. B. & BUSH, W. S. 2016. Analysis of Heritability Using Genome-Wide Data: Analysis of
Heritability Using Genome-Wide Data. In: HAINES, J. L., KORF, B. R., MORTON, C.
C., SEIDMAN, C. E., SEIDMAN, J. G. & SMITH, D. R. (eds.) Current Protocols in
Human Genetics. Hoboken, NJ, USA: John Wiley & Sons, Inc.
HARBONE, J. B. 1995. The flavonoids; Advances in researc since 1986. Journal of Chemical
Education, 72, A-73.
HAREL, D., SOFER, M., BRONER, M., ZOHAR, D. & GANTZ, S. 2014. Growth Stage Specific
Kc of Greenhouse Tomato Plants Grown in Semi-Arid Mediterranean Region. .Journal of
Agricultural Science, 6, 132.
HELD, M. T., ANTHON, G. E. & BARRETT, D. M. 2015. The effects of bruising and temperature
on enzyme activity and textural qualities of tomato juice. Journal of the Science of Food
and Agriculture, 95, 1598-1604.
HEMINGWAY, R. W. & LAKS, P. E. 1992. Plant Polyphenols: Synthesis properties, significance,
New York, USA, Plenum Press.
HEUVELINK, E. & DORAIS, M. 2005. Tomatoes, Wallingford, CABI.
HEWETT, E. W. 2006. An overview of preharvest factors influencing postharvest quality of
horticultural products. International Journal of Postharvest Technology and Innovation, 1,
4-15.
HIWASA, K. & EZURA, H. 2014. Climacteric and Non-climacteric Ripening. Fruit ripening:
physiology, signalling and genomics. London: CABI.
HODGES, R. J. 2010. Postharvest losses and waste in developed and less developed countries:
opportunities to improve resource use. Journal of Agricultural Science, 149, 37-45.
HWANG, I., KIM, Y., HAN, J. & NOU, I. S. 2016. Orange color is associated with CYC-B
expression in tomato fleshy fruit. Molecular Breeding, 36, 42.

51
HYMAN, J. R., GAUS, J. & FOOLAD, M. R. 2004. A rapid and accurate methodh for estimating
tomato lycopene content by measuring chromaticity values of fruit puree. Journal of the
American Society for Horticltural Science, 129, 717-723.
INSTITUTE OF MEDICINE, U. P. O. D. A. A. R. C. 2000. Dietary Reference Intakes for Vitamin
C, Vitamin E, Selenium, and Carotenoids. NCBI.
JENKINS, J. A. 1948. The Origin of the Cultivated Tomato. Economic Botany, 2, 379-392.
JONES JR, J. B. 2007. Tomato plant culture: in the field, greenhouse, and home garden, CRC Press.
JONES, R. 2007. Effects of postharvest handling conditions and cooking on anthocyanin, lycopene,
and glucosinolate content and bioavailability in fruits and vegetables. New Zealand journal
of crop and horticultural science, 35, 219-227.
KADER, A. A. 2005. Increasing food availability by reducing postharvest losses of fresh produce.
Acta Hort., 682, 2169-2176.
KADER, A. A. 2008. Perspective flavor quality of fruits and vegetables. Journal of the Science of
Food and Agriculture, 88, 1863-1868.
KANAI, S., REDA, E. M., EL-SHEMYC, H. A., PANIGRAHID, R., PRAVAT, K. M., ITO, J.,
NGUYEN, T. N., SANEOKAA, H. & E, K. F. 2011. Potassium deficiency affects water
status and photosynthetic rate of the vegetative sink in green house tomato prior to its
effects on source activity. Plant Science, 180, 368-374.
KHADKA, R. B., MARASINI, M., RAWAL, R., GAUTAM, D. M. & ACEDO, A. L. 2017. Effects
of Variety and Postharvest Handling Practices on Microbial Population at Different Stages
of the Value Chain of Fresh Tomato (Solanum lycopersicum) in Western Terai of Nepal.
BioMed Research International, 2017, 1-6.
KIELHORN, J., MELBER C FAU - KELLER, D., KELLER D FAU - MANGELSDORF, I. &
MANGELSDORF, I. 2002 Palladium--a review of exposure and effects to human health.
Int J Hyg Environ Health, 205, 417-32.
KWON, Y.-S., PARK, S.-G. & YI, S.-I. 2009. Assessment of genetic variation among commercial
tomato (Solanum lycopersicum L) varieties using SSR markers and morphological
characteristics. Genes & Genomics, 31, 1-10.
LAHOZ, I., PEREZ DEL, C., VALCARCEL, M. & MACUA, J. I. 2016. Effect of water deficit on te
agronomicla performance and quality of processing tomato. Scientia horticulturae, 200,
55-65.
LARA, D. B., CAPARROS, R., FAUCONNIER, M.-L., BROSTAUX, Y., FRANCIS, F. &
VERHEGGEN, F. 2015. Tuta absoluta-induced plant volatiles: attractiveness towards the
generalist predatorMacrolophus pygmaeus. Arthropod-Plant Interactions, 9, 465-476.
LEWINSOHN, E., SITRIT, Y., BAR, E., AZULAY, Y., IBDAH, M., MEIR, A., YOSEF, E.,
ZAMIR, D. & TADMOR, Y. 2005. Not just colors—carotenoid degradation as a link
between pigmentation and aroma in tomato and watermelon fruit. Trends in Food Science
& Technology, 16, 407-415.
LI, Z. & THOMAS, C. 2014. Quantitative evaluation of mechanical damage to fresh fruits. Trends in
Food Science & Technology, 35, 138-150.
LIU, C.-H., CAI, L.-Y., LU, X.-Y., HAN, X.-X. & YING, T.-J. 2012. Effect of Postharvest UV-C
Irradiation on Phenolic Compound Content and Antioxidant Activity of Tomato Fruit
During Storage. Journal of Integrative Agriculture, 11, 159-165.
LÓPEZ CAMELO, A. F. & GÓMEZ, P. A. 2004. Comparação dos índices de cor para maturação do
tomate. Horticultura Brasileira, 22, 534-537.
LUTHRIA, D. L., SUDARSAN, M., KRIZEK, D. T. & KRIZEK, D. T. 2006. Content of total
phenolics and phenolic acids in tomato (Lycopersicon esculentum Mill.) fruits as
influenced by cultivar and solar UV radiation. Journal of Food Composition and Analysis,
19.
MACHEKA, L., SPELT, E. J. H., BAKKER, E.-J., VAN DER VORST, J. G. A. J. & LUNING, P.
A. 2018. Identification of determinants of postharvest losses in Zimbabwean tomato supply
chains as basis for dedicated interventions. Food Control, 87, 135-144.
MAGEE, R. L., CAPORASO, F. & PRAKASH, A. 2003. Effects of Exogenous Calcium Salt
Treatments on Inhibiting Irradiation-Induced Softening in Diced Roma Tomatoes. Journal
of Food Science, 68, 2430-2435.

52
MALUNDO, T. M. M., SHEWFELT, R. L., SCOTT, J. W. & SCOTT 1995. Flavor quality of fresh
tomato (Lycopersicon esculentum Mill.) as affected by sugar and acid levels. Postharvest
Biology and Technology, 6, 103-110.
MARTÍNEZ-HUÉLAMO, M., VALLVERDU, A., DI LECCE, G., VALDERAS-MARTıNEZ, P.,
TULIPANI, S., JAUREGUI, O., ESCRIBANO-FERRER, E., ESTRUCH, R., ILLAN, M.
& LAMUELA-RAVENTO, R. 2016. Bioavailability of tomato polyphenols is enhanced by
processing and fat addition: Evidence from a randomized feeding trial. Mol. Nutr. Food
Res, 00, 1-12.
MATTHEIS, J. P. & FELLMAN, J. K. 1999. Preharvest factors influencing flavor of fresh fruit and
vegetables. Postharvest Biology and Technology, 15, 227-232.
MIKKELSEN, R. L. 2005. Tomato Flavor and Plant Nutrition: A Brief Review. Better Crops, Vol.
89, 14-15.
MIRDEHGHAN, S. H. & VALERO, D. 2017. Bioactive compounds in tomato fruit and its
antioxidant activity as affected by incorporation of Aloe, eugenol, and thymol in fruit
package during storage. International Journal of Food Properties, 20, 1798-1806.
NATH, P., BOUZAYEN, M., MATTOO, A. K. & CLAUDE PECH, J. 2014 Fruit ripening:
physiology, signalling and genomics. , CABI.
NETA-SHARIR, I., ISAACSON, T., LURIE, S. & WEISS, D. 2005. Dual Role for Tomato Heat
Shock Protein 21: Protecting Photosystem II from Oxidative Stress and Promoting Color
Changes during Fruit Maturation. The Plant Cell, 17, 1829.
NIELSEN, S. 2017. Food Analysis. , Springer Science & Business Media.
NUNES, M. C. 2008. Color atlas of postharvest quality of fruits and vegetables "Quality of fruits
and vegetables", United Kingdom, Experimental Agriculture.
NUSSINOVITCH, A., WARD, G. & LURIE, S. 1996. Nondestructive Measurement of Peel Gloss
and Roughness to Determine Tomato Fruit Ripening and Chilling Injury. Journal of Food
Science, 61, 383-387.
ODA, M. & SAITO, T. 2006. Tomato. Horticulture in Japan. Kyoto Japan: The Japanese Society for
Horticulture Science.
OPIYO, A. M. & YING, T.-J. 2005. The effects of 1-methylcyclopropene treatment on the shelf life
and quality of cherry tomato (Lycopersicon esculentum var. cerasiforme) fruit.
International Journal of Food Science & Technology, 40, 665-673.
PALIYATH, G., MURR, D. P., HANDA, A. K. & LURIE, S. 2008. Postharvest Biology and
technology of fruits, vegetables, and Flowers, John Wiley & Sons.
PANOZZO, A., LEMMENS, L., VAN LOEY, A., MANZOCCO, L., NICOLI, M. C. &
HENDRICKX, M. 2013. Microstructure and bioaccessibility of different carotenoid
species as affected by high pressure homogenisation: A case study on differently coloured
tomatoes. Food Chemistry, 141, 4094-4100.
PAPADOPOULOU, A. & FRAZIER, R. A. 2004. Characterization of protein–polyphenol
interactions. Trends in Food Science & Technology, 15, 186-190.
PARMAR, P., OZA, V. P., CHAUHAN, V., PATEL, A. D., KATHIRIA, K. B. &
SUBRAMANIAN, R. B. 2010. Genetic diversity and DNA fingerprint study of tomato
discerned by SSR markers. International Journal of Biotechnology & Biochemistry, 6,
657+.
PARSONS, C. S., ANDERSON, R. E. & PENNEY, R. W. 1970. Storage of mature-green tomatoes
in controlled atmospheres. Journal of the American Society of Horticultural Science, 95,
791-4.
PASSAM, H. C., KARAPANOS, I. C., BEBELI, P. J. & SAVVAS, D. 2007. A review of recent
research on tomato nutrition, breeding and post harvest technology with refrence to fruit
quality. The European Journal of Plant Science and Biotechnology, 1, 1-21.
PATARO, G., SINIK, M., CAPITOLI, M. M., DONSÌ, G. & FERRARI, G. 2015. The influence of
post-harvest UV-C and pulsed light treatments on quality and antioxidant properties of
tomato fruits during storage. Innovative Food Science and Emerging Technologies, 30,
103-111.
PAULL, R. 1999. Effect of temperature and relative humidity on fresh commodity quality.
Postharvest Biology and Technology,, 15, 263–277.
PERALTA, I., KNAPP, S. & SPOONER, D. 2006. Nomenclature for wild and cultivated tomatoes.
Tomato Genetics Cooperative Report. AGRIS.

53
PERALTA, I. E., KNAPP, S. & SPOONER, D. M. 2005. New Species of Wild Tomatoes (Solanum
Section Lycopersicon: Solanaceae) from Northern Peru. Systematic Botany, 30, 424-434.
PEREZ, K., FROIKIN-GORDON, J. S., ABDOURHAMANE, I. K., LEVASSEUR, V., ALFARI,
A. A., MENSAH, A., BONSU, O., HABSATOU, B., ASSOGBA-KOMLAN, F.,
MBAYE, A. A., NOUSSOUROU, M., OTOIDOBIGA, L. C., OUÉDRAOGO, L., KON,
T., ROJAS, M. R., GAMBY, K. T., SHOTKOSKI, F., GILBERTSON, R. L. & JAHN, M.
M. 2017. Connecting smallholder tomato producers to improved seed in West Africa.
Agriculture & Food Security, 6, 42.
PINHEIRO, J. C., ALEGRIA, C. S. M., ABREU, M. M. M. N., GONÇALVES, E. M. & SILVA, C.
L. M. 2016. Evaluation of Alternative Preservation Treatments (Water Heat Treatment,
Ultrasounds, Thermosonication and UV-C Radiation) to Improve Safety and Quality of
Whole Tomato. . Food and Bioprocess Technology, , 9, 924–935.
PORAT, R., LICHTER, A., TERRY, L. A., HARKER, R. & BUZBY, J. 2018. Postharvest losses of
fruit and vegetables during retail and in consumers’ homes: Quantifications, causes, and
means of prevention. Postharvest Biology and Technology, 139, 135-149.
PREEDY, V. R. & WATSON, R. R. 2008. Tomatoes and Tomto Products: Nutritional, Medicinal
and Therapeutic Properties, Boca Raton CRC Press.
PROULX, E., NUNES, M. C. N., EMOND, J. P. & BRECHT, J. K. 2001. Quality curves for tomato
exposed at chilling and non-chilling temperatures. . HortScience, , 36,, 509.
QUIDEAU, S. 2011. Why bother with polyphenols? [Online]. Talence Cedex, Fance: Groupe
Polyphenols. Available: [Link]
polyphenols/ [Accessed 2018 10 09 2018].
RAGHAVA, T., RAVIKUMAR, P., HEGDE, R. & KUSH, A. 2010. Spatial and temporal volatile
organic compund response of select tomato cultvars to herbivory and mechanical injury.
Plant Science, 179, 520-526.
RAO, A. V. & RAO, L. G. 2007. Carotenoids and human health. Pharmacological Research, 55,
207-216.
RATH, A. C. & PRENTICE, A. J. 2004. Yield increase and higher flesh firmness of 'Arctic Snow'
nectarines both at harvest in Australia and after export to Taiwan following pre-harvest
application of ReTain Plant Growth Regulator (aminoethoxyvinylglycine, AVG).
Australian Journal of Experimental Agriculture, 44, 343.
RAZA, W., WANG, J., WU, Y., LING, N., WEI, Z., HUANG, Q. & SHEN, Q. 2016. Effects of
volatile organic compounds produced by Bacillus amyloliquefaciens on the growth and
virulence traits of tomato bacterial wilt pathogen Rastonia solanacearum. Applied
Microbiology and Biotechnology, 100, 7639-7650.
RENDÓN-ANAYA, M. & HERRERA-ESTRELLA, A. 2017. Requirement of Whole-Genome
Sequencing. . In: PÉREZ DE LA VEGA, M., SANTALLA, M. & MARSOLAIS, F. (eds.)
The Common Bean Genome. . Cham: Springer International Publishing.
RIBEIRO, C., J. , CANADA, J. & ALVARENGA, B. 2012. Prospects of UV radiation for
application in postharvest technology. Emirates Journal of Food and Agriculture, 24, 586-
97.
RICK, C. M. 2017. Gene details [Online]. Available:
[Link] [Accessed].
RODONI, L., CASADEI, N., CONCELLÓN, A., CHAVES ALICIA, A. R. & VICENTE, A. R.
2010. Effect of Short-Term Ozone Treatments on Tomato (Solanum lycopersicum L.) Fruit
Quality and Cell Wall Degradation. Journal of Agricultural and Food Chemistry, 58, 594-
599.
RODRIGUEZ-LAFUENTE, A., NERIN, C. & BATLLE, R. 2010. Active Paraffin-Based Paper
Packaging for Extending the Shelf Life of Cherry Tomatoes. Journal of Agricultural and
Food Chemistry, 58, 6780-6786.
SAMMI, S. & MASUD, T. 2009. Effect of different packaging systems on the quality of tomato
(Lycopersicon esculentum var. Rio Grande) fruits during storage. International Journal of
Food Science & Technology, 44, 918-926.
SARGENT, S. A. 1998. Handling and Cooling Techniques for Maintaining Postharvest Quality. ,
Florida, University of Florida.
SAURE, M. C. 2014. Why calcium deficiency is not the cause of blossom-end rot in tomato and
pepper fruit – a reappraisal. Scientia Horticulturae, 174, 151-154.

54
SCHWARZ, D., THOMPSON, A. J. & KLäRING, H.-P. 2014. Guidelines to use tomato in
experiments with a controlled environment. Frontiers in Plant Science, 5.
SERINO, S., GOMEZ, L., COSTAGLIOLA, G. U. Y. & GAUTIER, H. L. E. N. E. 2009. HPLC
Assay of Tomato Carotenoids: Validation of a Rapid Microextraction Technique. Journal
of Agricultural and Food Chemistry, 57, 8753-8760.
SERRANO, M., GUILLÉN, F., MARTÍNEZ-ROMERO, D., CASTILLO, S. & VALERO, D. 2005.
Chemical Constituents and Antioxidant Activity of Sweet Cherry at Different Ripening
Stages. Journal of Agricultural and Food Chemistry, 53, 2741-2745.
SHI, J. & LE MAGUER, M. 2000. Lycopene in tomatoes: chemical and physical properties affected
by food processing. Crit Rev Food Sci Nutr., 40, 1-42.
SIRACUSA, L., PATANÈ, C., AVOLA, G. & RUBERTO, G. 2012. Polyphenols as
Chemotaxonomic Markers in Italian “Long-Storage” Tomato Genotypes. Journal of
Agricultural and Food Chemistry, 60, 309-314.
STEVENS, M. & ALLEN RICK, C. M. 1986. Genetics and breeding. In: ATHERTON, J. G. &
RUDICH, J. (eds.) The Tomato Crop: A scientific basis for improvement. Dordrecht:
Springer Netherlands.
STEWART, A. J., BOZONNET, S., MULLEN, W., JENKINS, G. I., LEAN, M. E. J. & CROZIER,
A. 2000. Occurrence of Flavonols in Tomatoes and Tomato-Based Products. Journal of
Agricultural and Food Chemistry, 48, 2663-2669.
STOMMEL, J., ABBOTT, J. A., SAFTNER, R. A. & CAMP, M. J. 2005 Sensory and objective
quality attributes of beta-carotene and lycopene-rich tomato fruit. Journal of the American
Society for Horticultural Science 130, 244-251.
SUN, Q., ZHANG, N., WANG, J., ZHANG, H., LI, D., SHI, J., LI, R., WEEDA, S., ZHAO, B.,
REN, S. & GUO, Y.-D. 2015. Melatonin promotes ripening and improves quality of
tomato fruit during postharvest life. Journal of Experimental Botany, , Vol. 66, pp.-657–
668.
SUSLOW, M. C. A. T. 1997. Recommendation for Maintaining Postharvest Quality [Online].
Unated States. Available:
[Link]
English/?uid=36&ds=799 [Accessed 2018 12 01 2018].
THE TOMATO GENOME, C., SATO, S., TABATA, S., HIRAKAWA, H., ASAMIZU, E.,
SHIRASAWA, K., ISOBE, S., KANEKO, T., NAKAMURA, Y., SHIBATA, D., AOKI,
K., EGHOLM, M., KNIGHT, J., BOGDEN, R., LI, C., SHUANG, Y., XU, X., PAN, S.,
CHENG, S., LIU, X., REN, Y., WANG, J., ALBIERO, A., DAL PERO, F., TODESCO,
S., VAN ECK, J., BUELS, R. M., BOMBARELY, A., GOSSELIN, J. R., HUANG, M.,
LETO, J. A., MENDA, N., STRICKLER, S., MAO, L., GAO, S., TECLE, I. Y., YORK,
T., ZHENG, Y., VREBALOV, J. T., LEE, J., ZHONG, S., MUELLER, L. A.,
STIEKEMA, W. J., RIBECA, P., ALIOTO, T., YANG, W., HUANG, S., DU, Y.,
ZHANG, Z., GAO, J., GUO, Y., WANG, X., LI, Y., HE, J., LI, C., CHENG, Z., ZUO, J.,
REN, J., ZHAO, J., YAN, L., JIANG, H., WANG, B., LI, H., LI, Z., FU, F., CHEN, B.,
HAN, B., FENG, Q., FAN, D., WANG, Y., LING, H., XUE, Y., WARE, D., RICHARD
MCCOMBIE, W., LIPPMAN, Z. B., CHIA, J.-M., JIANG, K., PASTERNAK, S.,
GELLEY, L., KRAMER, M., ANDERSON, L. K., CHANG, S.-B., ROYER, S. M.,
SHEARER, L. A., STACK, S. M., ROSE, J. K. C., XU, Y., EANNETTA, N., MATAS, A.
J., MCQUINN, R., TANKSLEY, S. D., CAMARA, F., GUIGÓ, R., ROMBAUTS, S.,
FAWCETT, J., VAN DE PEER, Y., ZAMIR, D., LIANG, C., SPANNAGL, M.,
GUNDLACH, H., et al. 2012. The tomato genome sequence provides insights into fleshy
fruit evolution. Nature, 485, 635.
TIEMAN, D. M. & HANDA, A. K. 1994. Reduction in Pectin Methylesterase Activity Modifies
Tissue Integrity and Cation Levels in Ripening Tomato (Lycopersicon esculentum Mill.)
Fruits. Plant Physiology, 106, 429.
TIEMAN, D. M., HARRIMAN, R. W., RAMAMOHAN, G. & HANDA, A. K. 1992. An Antisense
Pectin Methylesterase Gene Alters Pectin Chemistry and Soluble Solids in Tomato Fruit.
The Plant Cell, 4, 667.
TODOROVSKA, E., IVANOVA, A., GANEVA, D., PEVICHAROVA, G., MOLLE, E., BOJINOV,
B., RADKOVA, M. & DANAILOV, Z. 2014. Assessment of genetic variation in

55
 Bulgarian tomato (Solanum lycopersicum L.) genotypes, using fluorescent SSR genotyping
platform. Biotechnology & Biotechnological Equipment, 28, 68-76.
TSAO, R. 2010. Chemistry and Biochemistry of Dietary Polyphenols. Nutrients, 2, 1231-1246.
TÜZEL, Y. & ÖZTEKIN, G. 2017. Tomato In: DUFFY, R. (ed.) Good Agricultural Practices for
greenhouse vegetable production in the South East European countries. Principles for
sustainable intensification of smallholder farms. Rome: Food and Agricultural
Organization of the United States.
TZORTZAKIS, N., BORLAND, A., SINGLETON, I. & BARNES, J. 2007. Impact of atmospheric
ozone-enrichment on quality-related attributes of tomato fruit. Postharvest Biology and
Technology, 45, 317-325.
ULUISIK, S., CHAPMAN, N. H., SMITH, R., POOLE, M., ADAMS, G., GILLIS, R. B., BESONG,
T. M. D., SHELDON, J., STIEGELMEYER, S., PEREZ, L., SAMSULRIZAL, N.,
WANG, D., FISK, I. D., YANG, N., BAXTER, C., RICKETT, D., FRAY, R., BLANCO-
ULATE, B., POWELL, A. L. T., HARDING, S. E., CRAIGON, J., ROSE, J. K. C., FICH,
E. A., SUN, L., DOMOZYCH, D. S., FRASER, P. D., TUCKER, G. A., GRIERSON, D.
& SEYMOUR, G. B. 2016. Genetic improvement of tomato by targeted control of fruit
softening. Nature Biotechnology, 34, 950.
USDA. 2016. Vegetables and pulses Yearbook Tables [Online]. United States Department of
Agriculture. Available: [Link]
[Accessed 2018].
VALDIVIA-NÁJAR, C. G., MARTÍN-BELLOSO, O. & SOLIVA-FORTUNY, R. 2018. Impact of
pulsed light treatments and storage time on the texture quality of fresh-cut tomatoes.
Innovative Food Science & Emerging Technologies, 45, 29-35.
VALLVERDÚ-QUERALT, A., JÁUREGUI, O., MEDINA-REMÓN, A., ANDRÉS-LACUEVA, C.
& LAMUELA-RAVENTÓS, R. M. 2010. Improved characterization of tomato
polyphenols using liquid chromatography/electrospray ionization linear ion trap
quadrupole Orbitrap mass spectrometry and liquid chromatography/electrospray ionization
tandem mass spectrometry. Rapid Communications in Mass Spectrometry, 24, 2986-2992.
VAN DEN BROECK, G. & MAERTENS, M. 2017. Moving Up or Moving Out? Insights into Rural
Development and Poverty Reduction in Senegal. World Development, 99, 95-109.
VAN DIJK, C., BOERIU, C., STOLLE-SMITS, T. & TIJSKENS, L. M. M. 2006. The firmness of
stored tomatoes (cv. Tradiro). 2. Kinetic and Near Infrared models to describe pectin
degrading enzymes and firmness loss. Journal of Food Engineering, 77, 585-593.
VASSALLO, N. 2008. Plyphenols and Health New and recent advances, Nova Publishers.
VERHOEYEN, M. E., BOVY, A., COLLINS, S., MUIR, S. & ROBINSON, C. 2002. Increasing
antioxidant levels in tomatoes through modification of the flavonoid biosynthetic pathway.
Journal of Experimental Botany, 53, 2099-2106.
WABALI, V. C., ESIRI, A. & ZITTE, L. 2017. A sensory assessment of color and textural quality of
refrigerated tomatoes preserved with different concentrations of potassium permanganate.
Food Science & Nutrition, 5, 434-438.
WANG, M., CAO, J., LIN, L. I. N., SUN, J. & JIANG, W. 2010a. Effect of 1-Methylcyclopropene
on nutritional quality and antioxidant activity of tomato fruit (Solanum Lycopersicon L)
during storage. Journal of Food Quality, 33, 150-164.
WANG, Y., JACOBS, E., NEWTON, C. & MCCULLOUGH, M. 2016. Lycopene, tomato products
and prostate cancer-specific mortality among men diagnosed with nonmetastatic prostate
cancer in the Cancer Prevention Study II Nutrition Cohort. International Journal of
Cancer, 138, 2846-2855.
WANG, Y., REN, X., SONG, X., YU, T., LU, H., WANG, P., WANG, J. & ZHENG, X. D. 2010b.
Control of postharvest decay on cherry tomatoes by marine yeast Rhodosporidium
paludigenum and calcium chloride. Journal of Applied Microbiology, 109, 651-656.
WEI, Y., ZHOU, D., WANG, Z., TU, S., SHAO, X., PENG, J., PAN, L. & TU, K. 2018. Hot air
treatment reduces postharvest decay and delays softening of cherry tomato by regulating
gene expression and activities of cell wall-degrading enzymes: Effect of hot air treatment
on cherry tomato. Journal of the Science of Food and Agriculture, 98, 2105-2112.
WEINBERGER, K. & LUMPKIN, T. A. 2007. Diversification into Horticulture and Poverty
Reduction: A Research Agenda. World Development, 35, 1464-1480.

56
WILLS, R., LEE, T. & GRAHAM, D. 1990. An introduction to the physiology and handling of fruits
and vegetables. Postharvest. 1 ed.: Springer US.
WRAY, N. V., P. . 2008. Estimating Trait Heritability. [Online]. Scitable by nature education.
Available: [Link]
[Accessed 2018-10-26] 2018].
XU, H.-L., IRAQI, D. & GOSSELIN, A. 2007. Effect of Ambient Humidity on Physiological
Activities and Fruit Yield and Quality of Greenhouse Tomato. In: ISHS (ed.) XXVII
International Horticultural Congress - IHC2006: International Symposium on Advances in
Environmental Control. Acta Horticulturae
YAMAMOTO, E., MATSUNAGA, H., ONOGI, A., KAJIYA-KANEGAE, H., MINAMIKAWA,
M., SUZUKI, A., SHIRASAWA, K., HIRAKAWA, H., NUNOME, T., YAMAGUCHI,
H., MIYATAKE, K., OHYAMA, A., IWATA, H. & FUKUOKA, H. 2016. A simulation-
based breeding design that uses whole-genome prediction in tomato. Scientific Reports, 6,
19454.
ZABIN, C. F. & HUGHES, S. 1995. Economic integration and labor flows: stage migration in farm
labor markets in Mexico and the United States. Int Migr Rev., 29, 395-422.
ZHANG, B., TIEMAN, D. M., JIAO, C., XU, Y., CHEN, K., FEI, Z., GIOVANNONI, J. J. &
KLEE, H. J. 2016. Chilling-induced tomato flavor loss is associated with altered volatile
synthesis and transient changes in DNA methylation. Proceedings of the National
Academy of Sciences, 113, 12580.
ZHANG, C., CHEN, K. & WANG, G. 2013. Combination of the biocontrol yeast Cryptococcus
laurentii with UV-C treatment for control of postharvest diseases of tomato fruit.
BioControl, 58, 269-281.
ZHANG, M., YUAN, B. & LENG, P. 2009. The role of ABA in triggering ethylene biosynthesis and
ripening of tomato fruit. Journal of Experimental Botany, 60, 1579-1588.

57