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Selection Of Tomato Landraces With High Fruit Yield and Nutritional Quality Under Elevated Temperatures


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Background: Global warming and extreme or adverse events induced by climatic fluctuations, are an important threat for plants growth and agricultural production. Adaptability to environmental changes prevalently derives from a large set of genetic traits affecting physiological and agronomic parameters. Therefore, the identification of genotypes that are good yield performer at high temperatures is becoming increasingly necessary for future breeding programs. Here, we analyzed the performances of different tomato landraces grown under elevated temperatures in terms of yield and nutritional quality of the fruit. Finally, we evaluated the antioxidant and anti-inflammatory activities of fruit extracts from the selected tomato landraces. Results: Tomato landraces here analyzed in a hot climate differed in terms of yield performance, physicochemical parameters of fruit (pH, titratable acidity, °Brix, firmness), bioactive compounds (ascorbic acid, carotenoids and polyphenols) and anti-inflammatory potential. Three of these landraces (named E30, E94 and PDVIT) showed higher fruit quality and nutritional value. An estimated Evaluation Index allowed to identify PDVIT as the best performer in term of yield and fruit quality under high temperatures. Conclusion: The analyses here performed highlight the possibility to identify new landraces that can combine good yield performances and fruit nutritional quality at high temperatures, useful for future breeding programs. This article is protected by copyright. All rights reserved.
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Research Article
Received: 31 October 2019 Revised: 30 January 2020 Accepted article published: 5 February 2020 Published online in Wiley Online Library: 29 February 2020
( DOI 10.1002/jsfa.10312
Selection of tomato landraces with high fruit
yield and nutritional quality under elevated
Aurelia Scarano,aFabrizio Olivieri,bCarmela Gerardi,aMarina Liso,c
Maurizio Chiesa,dMarcello Chieppa,c,e Luigi Frusciante,bAmalia Barone,b
Angelo Santinoa
and Maria Manuela Riganob
BACKGROUND: Global warming and extreme or adverse events induced by climatic uctuations are an important threat for
plants growth and agricultural production. Adaptability to environmental changes prevalently derives from a large set of
genetic traits affecting physiological and agronomic parameters. Therefore, the identication of genotypes that are good yield
performer at high temperatures is becoming increasingly necessary for future breeding programs. Here, we analyzed the per-
formances of different tomato landraces grown under elevated temperatures in terms of yield and nutritional quality of the
fruit. Finally, we evaluated the antioxidant and anti-inammatory activities of fruit extracts from the tomato landraces selected.
RESULTS: The tomato landraces analyzed here in a hot climate differed in terms of yield performance, physicochemical param-
eters of fruit (pH, titratable acidity, degrees Brix, rmness), bioactive compounds (ascorbic acid, carotenoids, and polyphenols),
and anti-inammatory potential. Three of these landraces (named E30, E94, and PDVIT) showed higher fruit quality and nutri-
tional value. An estimated evaluation index allowed identication of PDVIT as the best performer in terms of yield and fruit
quality under high temperatures.
CONCLUSION: The analyses performed here highlight the possibility to identify new landraces that can combine good yield per-
formances and fruit nutritional quality at high temperatures, information that is useful for future breeding programs.
© 2020 The Authors. Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of
Chemical Industry.
Supporting information may be found in the online version of this article.
Keywords: tomato; high temperature tolerance; yield performance; fruit quality; nutritional value
One of the most important challenges facing us today is deal-
ing with global warming, which can greatly impact on crop
production and food accessibility. Together with water stress
and high salinity, temperatures and light uctuations during
plant growth are the most important abiotic stresses that
plants have to face.
Tomato is one of the major cultivated crops worldwide and a
model system to study plant response to the changing environmen-
tal conditions. The optimal range of temperatures for tomato cultiva-
tion is between 25 and 30 °C during the day and 20 °Cduringthe
However, tomato varieties or landraces can exhibit individual
differences in terms of yield performances, depending on the genetic
background and the adaptation to the environment in specicgeo-
graphic areas. In this context, the exploration of the natural variation
and the screening of genotypes and landraces that are good yield
performers at high temperatures may help to understand the mech-
anisms underlying high-temperature tolerance and can provide
agronomic traits and genetic diversity useful for breeding.
Increases in the average temperatures and UV radiation can
have a signicant impact on plant growth and, consequently, on
crop yield and fruit quality. Indeed, different biochemical mecha-
nisms, including plastid biogenesis and pigments/secondary
*Correspondence to: A Santino, ISPACNR, Institute of Science of Food
Production, CNR Unit of Lecce, via Provinciale Lecce-Monteroni, 73100
Lecce, Italy. E-mail:
aISPACNR, Institute of Science of Food Production, CNR Unit of Lecce, Lecce, Italy
bDepartment of Agricultural Sciences, University of Naples Federico II, Naples, Italy
cNational Institute of Gastroenterology S. De Bellis, Institute of Research, Bari,
dBiotecgen S.r.l., Lecce, Italy
eDepartment of Immunology and Cell Biology, European Biomedical Research
Institute of Salerno (EBRIS), Salerno, Italy
© 2020 The Authors. Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.
This is an open access article under the terms of the Creative Commons Attribution-NonCommercial License, which permits use, distribution and
reproduction in any medium, provided the original work is properly cited and is not used for commercial purposes.
metabolites synthesis, can be activated in plant cells in response
to abiotic stresses.
Since thermo-tolerance requires the mod-
ulation of biochemical pathways involved in reactive oxygen spe-
cies (ROS) detoxication, antioxidant compounds, including
carotenoids, ascorbic acid (AsA) and polyphenols, are accumu-
lated in response to heat stress.
Carotenoids can also contribute
to membranes uidity and permeability in response to tempera-
ture uctuations.
Indeed, these molecules can exert a protec-
tive role in photosynthetic membranes and play important roles
in structural stabilization, light harvesting, and photoprotection.
The increase in the biosynthesis of polyphenolic compounds
and avonoids has also been reported following the exposure of
many plant species to UV radiation, making their accumulation
one of the most evolutionarily conserved responses to this type
of abiotic stress.
Polyphenolic compounds, AsA, and carotenoids are also rele-
vant for human health, since antioxidant and anti-inammatory
properties have been described following their in vitro or in vivo
In particular, phenolic compounds may have
therapeutic roles in inammation-based diseases and various
type of cancers.
Furthermore, ascorbic acid shows signicant
antioxidant and electron donor capability and is able to protect
DNA from oxidation-induced damage.
Finally, it has been
demonstrated that carotenoids possess an apoptotic-inducing
effect in cancer cells and reduce oxidized low-density lipoprotein
cholesterol levels.
The health-promoting properties of these
bioactive compounds contribute to the nutritional value of crops
such as tomato and represent a parameter of preference in con-
sumer choice of food products. Indeed, some of these bioactive
compounds, such as AsA and carotenoids, cannot be synthesized
by animals and thus have to be introduced with the diet.
these reasons, the daily consumption of plant-derived food
enriched in these compounds has been highly promoted in the
last decade.
In this study, we analyzed the performances, in terms of nal
yield and fruit quality on a set of ten tomato landraces showing
good performances when grown under elevated temperatures.
Their productivity and eld performances under restrictive envi-
ronmental conditions were estimated. Moreover, the bioactive
compounds content and antioxidants/anti-inammatory proper-
ties have been evaluated in extracts from ripe fruits from different
tomato landraces, with the nal aim of identifying the best ones in
terms of both yield and nutritional fruit quality. Altogether, this
study highlights the possibility to select tomato landraces com-
bining desirable agronomic and nutritional traits, such as good
yield performances and high nutritionally active phytonutrients
content, for future applications in breeding programs.
Plant materials
Plant material consisted of ten tomato indeterminate landraces
(listed in Table 1) and the control variety Moneymaker. In 2017
they were grown in Apulia (Pulsano, 40°2300300 N latitude,
17°2101700 E longitude), a region of southern Italy greatly devoted
to tomato cultivation and usually characterized by high tempera-
tures during the growing season. Seeds were sown in plateau
under a plastic-house in April, and seedlings were then trans-
planted in open eld in June. Plants were distributed following a
complete randomized block design, with three replicate plots
per landrace and ten plants per replicate. For fruit quality and
nutritional traits analyses, ten fruits at the red ripe stage from at
least three plants from each replicate plot were collected at the
same time and pooled. Some traits were evaluated on fresh fruit,
and others on fruit frozen in liquid nitrogen and stored at 80 °C
until analysis. For yield determination, fruits at red ripe stage were
collected on the same day from all the plants for each replicate
plot. Total fruit number and fresh weight (FW) were measured to
allow yield evaluation per plant.
Quality traits evaluation
Physicochemical traits were evaluated on fresh fruit. The determi-
nation of pH was carried out by using a pH meter (Mettler-Toledo,
Milan, Italy), and the total acidity was determined by titrating
Table 1. List of the plant materials tested for production, quality, and nutritional traits under high temperature
Genotype Source Accession no. Common name
of origin Collection site
destination Fruit size Fruit shape
Corbarino PC04 Italy Nocera (Salerno) Processing Small (2530 g) Ovate
Corbarino PC05 Italy Sant'Antonio
Abate (Salerno)
Processing Small (2025 g) Elliptic
Pantano Romanesco Italy Fondi (Latina) Fresh market Big (200250 g) Flattened
Sel PC07 Italy Pagani (Salerno) Processing Small (1520 g) Ovate
Sel PC16 Italy Nocera (Salerno) Fresh market Small (2050 g) Ovate
E36 Campania
Vesuvio Foglia Riccia Italy S. Vito (Naples) Fresh market/
Small (2530 g) Ovate
E53 TGRC LA0147 Honduras Tegucigalpa
Fresh market Medium (80100 g) Oblate
E76 TGRC LA4449 Black plum URSS Processing Small (2025 g) Ovate
E94 NPGS PI272890 1404 Guatemala Quetzaltenango,
nd Small (4050 g) Irregular
Caramella Italy Scafati (Salerno) Fresh market/
Small (1015 g) Elliptic
MoneymakerTGRC LA2706 Moneymaker Great Britain Fresh market Medium (5060 g) Circular
Germplasm collections maintained at Italian public institution. A Scarano et al. © 2020 The Authors.
Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.
J Sci Food Agric 2020; 100: 27912799
10 mL of tomato juice with a solution of 0.1 mol L
hydroxide. A few drops of tomato juice were also used to estimate
the total soluble solids (degrees Brix) by adding them to the prism
plate of a refractometer (Hanna Instruments). Finally, the rmness
was measured by a penetrometer (PCE-PTR200 penetrometer,
Capannori, Italy) using an 8 mm tip. All extracts were from three
biological replicates, and three technical assays were carried out
on each biological repetition.
Extraction and detection of polyphenols
Whole tomato fruits were cut and frozen in liquid nitrogen, freeze-
dried, and nely ground. Samples of powder (200 mg) were
extracted twice in methanol:water 80:20 (v/v). The extracts were cen-
trifuged, and the supernatants were combined, ltered through a
0.22 μmlter, and stored at 20 °C until use. Polyphenols were
detected at 320 nm by reversed-phase high-performance liquid
chromatography with diode array detector (RP-HPLC DAD) (Agilent
1100 HPLC system). Separation was performed on a C18 column
(5 UltraSphere, 80 Å pore, 25 mm), with a linear gradient from 20%
to 60% acetonitrile, in 55 min, with a ow of 1 mL min
at 25 °C.
Concentrations were obtained by referring to calibration curves,
and results were expressed in micrograms per gram or milligrams
per gram (in the case of rutin and chlorogenic acid) of dried weight.
Carotenoids content
Freeze-dried tomato powder (50 mg) was added to 2 mL of 60%
potassium hydroxide, 2 mL of absolute ethanol, 1 mL of 1%
sodium chloride (NaCl), 5 mL of 0.05% butylated hydroxytoluene
in acetone. The mix was incubated at 60 °C for 30 min. A 1% solu-
tion of NaCl (15 mL) was added to the mix, and extractions were
performed with 15 mL hexane:ethyl acetate 9:1 (v/v). Extracts
were centrifuged, evaporated using a rotary evaporator, and col-
lected in 1 mL of ethyl acetate. Analyses were performed using
RP-HPLC DAD (Agilent 1100 HPLC system) according to the
method previously described.
AsA determination
AsA determinations were carried out by a colorimetric method with
modications reported by Rigano et al.
Briey, 500 mg of frozen
powder from tomato fruits was extracted with 300 μLof6%tri-
chloroacetic acid (TCA). The mixture was vortexed, incubated on
ice for 15 min, and centrifuged at 15 700×gfor 20 min. To 20 μL
of supernatant were added 20 μLof0.4molL
phosphate buffer
(pH 7.4), 10 μL of double-distilled water, and 80 μL of color reagent
solution. This last solution was prepared by mixing solution A (31%
(w/v) phosphoric acid, 4.6% (w/v) TCA, and 0.6% (w/v) ferric chlo-
ride) with solution B (4% (w/v) 2,20-dipyridyl). These mixtures were
incubated at 37 °C for 40 min and measured at 525 nm using a
UVvisible spectrophotometer (NanoPhotometer; Implen). All
extracts were from three biological replicates, and three technical
assays were carried out on each biological repetition. The concen-
tration was expressed in micrograms per gram FW.
Determination of antioxidant activity
2,20-Azino-bis(3-ethylbenzothiazoline-6-sulfonic acid) diammo-
nium salt (ABTS; Sigma-Aldrich) radical cations were prepared by
mixing an aqueous solution of 2.45 mmol L
potassium persul-
fate (nal concentration) with an aqueous solution of 7 mmol L
ABTS (nal concentration) and allowed to stand in the dark at
room temperature for 1216 h before use. The ABTS radical cation
solution was diluted in phosphate-buffered saline (PBS; pH 7.4) to
an absorbance of 0.40 at 734 nm. Trolox was used to prepare a
standard calibration curve (016 μmol L
). After the addition of
200 μL of diluted ABTS to 10 μL of Trolox standard or extracts diluted
in PBS, in each well of a 96-well plate (Costar), the absorbance was
read at 734 nm 6 min after initial mixing using an Innite200Pro plate
reader (Tecan). All extracts were from three biological replicates, and
three technical assays were carried out on each biological repetition.
The percentage absorbance inhibition at 734 nm was calculated as a
function of concentration of Trolox, and the Trolox equivalent antiox-
idant capacity (TEAC) value was expressed as Trolox equivalent (TE,
μmol) using Magellan v7.2 software.
Culture of dendritic cells and enzyme-linked
immunosorbent assay
Bone-marrow-derived dendritic cells (BMDCs) were obtained
from C57BL/6 mice, in agreement with national and international
guidelines, approved by the authors' institutional review board
(Organism for Animal Wellbeing OPBA).
BMDCs were harvested as previously described
and plated in
Roswell Park Memorial Institute 1640 medium supplemented with
fetal bovine serum, antibiotics, recombinant mouse granulocyte
macrophage stimulating factor and recombinant mouse interleu-
kin (IL)-4 at 37°C in a humidied 5% carbon dioxide atmosphere.
BMDCs were treated with tomato methanol extracts (100 mg
lyophilized powder per milliliter, 1:25 nal dilution), after adminis-
tration of lipopolysaccharide (LPS; 1 μgmL
) at day 8 for 24 h.
BMDCs culture media were analyzed for IL-6 and IL-12 in triplicate
using an enzyme-linked immunosorbent assay (ELISA) kit as
described by the manufacturer (R&D Systems).
Data analysis
Values are expressed as mean plus/minus standard deviation (SD).
Group differences were analyzed and compared by paired two-
tailed Student's t-tests. Yield differences among the genotypes ana-
lyzed were determined using SPSS Package 6, version 15.0. Signi-
cant different yields were determined by comparing mean values
through a factorial analysis of variance with Duncan post hoc test
at a signicance level of 0.05. Spearman correlations were calcu-
lated to analyze co-occurrence and associations among all traits
measured. The P-values obtained for multiple tests were corrected
using the Benjamini and Hochberg false discovery rate (FDR). In
order to identify the landraces with a desirable combination of
traits, an evaluation index (EI) was estimated by assigning a score
to each trait, which was a maximum of 11 to a minimum of 1 des-
cending from the highest to the lowest value for all traits except
for pH, IL-6, and IL-12, where the maximum score (11) was assigned
to the lower value and the minimum score (1) to the highest value.
Agronomic performances of tomato landraces under harsh
temperature conditions
A group of tomato landraces was previously selected for yield per-
formances under high temperatures in two regions of southern
Italy, Campania and Apulia, in 2016.
In the present study, we
decided to test them again under high temperatures in an open
eld in the Apulia region in 2017. Plants were transplanted with
a 1 month delay with respect to the usual agronomical practice
of the area, exposing them to higher temperatures during the crit-
ical stages of owering and fruit setting. Figure S1 reports the
values of mean, maximum, and minimum temperatures recorded
over the growing season, together with the average relative
humidity of the whole day. As shown, more than 40 days (38.5%
Selection of tomato landraces with high fruit yield
J Sci Food Agric 2020; 100: 27912799 © 2020 The Authors.
Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.
of the whole growing season, from the end of May to the begin-
ning of September) reached temperatures over 32 °C, which is
considered the critical temperature affecting the reproductive
stage of tomato, as well as of other species.
On 16 days
(15.4%), a temperature higher than 35°C was even recorded. In
addition, very high temperatures were also recorded in the night,
and this is considered a critical point for pollen maturation.
Based on the evaluation of production per plant (Table 2), four
genotypes were classied as good (E8, E32, E36, Moneymaker),
ve as medium (E7, E17, E53, E76, PDVIT), and two as low pro-
ducers (E30, E94) under high-temperature conditions. Indeed,
the rst group exhibited a yield per plant higher than 4 kg, the
second group yield ranged from 2 to 4 kg per plant, whereas
the third group produced less than 2 kg per plant. As a whole,
yield performances conrmed previous data and allowed a reli-
able classication of the genotypes in the aforementioned three
Assessment of main quality parameters
With the aim of identifying those landraces simultaneously exhi-
biting good productivity and good nutritional quality under high
temperatures, and thus worth proposing as resilient varieties, we
evaluated the main quality parameters of the ten landraces. As
already reported in a previous study,
we chose the indetermi-
nate variety Moneymakeras a control genotype, considering
that it generally exhibits stable yields and fruit quality traits under
different environmental conditions and in different years. Con-
cerning the main fruit quality traits, the pH values ranged from
4.3 (landraces PDVIT and E76) to 4.6 (E17), with a mean value of
4.4, whereas the titratable acidity ranged from 0.38 g to 0.55 g
of citric acid per 100 mL of tomato juice in E36 and E76 respec-
tively (Table 3). The soluble solid content varied from 3.6 °Bx
(E7) to 8.1 °Bx (E8). Finally, E94 showed the lowest level of rmness
(8.7 kg cm
) and PDVIT the maximum (18.9 kg cm
). Notewor-
thy is that the three landraces E8, E30, and E76 showed high
degrees Brix levels (>7), and two of these (E30 and E76) also had
a titratable acidity value higher than 0.5. Altogether, these traits
affect consumer taste, who generally prefer rm, sweet, and acid
and consequently the commercial value of tomatoes.
In recent years, more attention has been paid to the nutritional
value of food products, focusing on the benecial effects of fresh
fruit and vegetables. To verify if the selected landraces could also
provide a good nutritional value when grown under high temper-
atures, the main tomato phytonutrients contents, such as AsA,
polyphenols, and carotenoids, were determined. These com-
pounds have been associated to health benets and the reduc-
tion of inammatory and aging-related diseases.
the dietary intake of these compounds is highly recommended.
The AsA levels in the red ripe fruit of the ten tomato landraces
and in the control cv Moneymaker are reported in Fig. 1. Most
landraces exhibited values of approximately 350 μgg
which was statistically different from the value recorded in Mon-
eymaker(232 μgg
FW), whereas two of them (E30 and E94)
reached a mean value higher than 400 μgg
FW. The landrace
PDVIT was the best performer, with a mean value over 500 μgg
FW. These values are in line with the values previously reported
for tomato genotypes, where AsA ranged from 100 to 880 μgg
FW, even though commercial cultivars are generally characterized
by lower contents (from 100 to 400 μgg
FW), probably due to
the breeding process.
However, the AsA content in fresh tomato
fruits is also dependent on genotype, climatic conditions, fruit
development, and maturation.
Carotenoids accumulation has been reported following abiotic
stresses, such as high or low temperature and high light, upon
which the homeostasis of ROS metabolism is challenged. For
Table 2. Yield (kg) per plant measured on ten landraces and the
control cv Moneymaker during 2017
Genotype Yield (kg)/plant Production level
E7 3.43 ±0.57
E8 4.15 ±1.12
E17 2.71 ±0.29
E30 1.70 ±039
E32 4.93 ±0.83
E36 5.67 ±0.58
E53 3.67 ±0.25
E76 2.59 ±0.41
E94 1.67 ±0.71
PDVIT 2.98 ±0.51
Moneymaker4.85 ±0.60
Following Duncan's post hoc test, the level of production under high
temperature was also reported. Values followed by the same letters
are not signicantly different.
Table 3. Qualitative traits (mean and standard error) parameters measured on red ripe fruit. Statistical analysis was carried out by comparing values
with those recorded in the cv Moneymaker (M)
Genotype pH Soluble solid content (°Bx) Titratable acidity (g citric acid/100 mL juice) Firmness (kg cm
E7 4.45 ±0.05 3.62 + 0.26** 0.43 + 0.01 14.31 + 1.11
E8 4.41 ±0.02 8.13 ±0.01** 0.46 + 0.01 15.16 + 0.12
E17 4.65 + 0.01* 4.29 + 0.17** 0.49 + 0,02 14.02 + 2.10
E30 4.42 + 0.04 7.88 + 0.21** 0.53 + 0.04 13.21 + 1.34
E32 4.49 + 0.02 6.58 + 0.24 0.39 + 0.01 15.03 + 0.18
E36 4.40 + 0.05 3.60 + 0.45** 0.38 + 0.03 14.05 + 2.51
E53 4.40 + 0.04 5.92 + 0.20 0.47 + 0.02 11.17 + 1.45
E76 4.32 + 0.02* 7.74 + 0.26* 0.55 + 0.03* 10.91 + 0.98*
E94 4.36 + 0.04 6.78 + 0.12 0.48 + 0.03 8.68 + 0.01**
PDVIT 4.31 + 0.06 5.92 + 0.01 0.50 + 0.02 18.90 + 1.50
Moneymaker4.47 + 0.04 6.17 + 0.27 0.43 + 0.02 15.18 + 1.05
*P<0.05, **P<0.01 (Student's t-test). A Scarano et al. © 2020 The Authors.
Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.
J Sci Food Agric 2020; 100: 27912799
these reasons, carotenoids are implicated in the thermo-tolerance
of different plant species, including tomato.
Therefore, herein,
we analyzed the carotenoids content in the fruits of the tomato
landraces considered here. Concerning the main carotenoids
(i.e. lycopene, -carotene, and lutein), our data showed signi-
cantly higher levels for these compounds in E30, E36, and E94
compared with cv Moneymaker (Fig. 2). Noteworthy is that a
higher -carotene content compared with the control cultivar
was also evidenced in PDVIT. This compound is considered a pro-
vitamin, which can be converted into retinol, a phytochemical
essential for vision. -Carotene is also known to act as a strong
antioxidant and is the best quencher of singlet oxygen.
The -carotene/lycopene ratio (Fig. S2) showed a different trend
among the landraces tested, with higher levels recorded in E17,
E30, and E32 (0.76, 0.68, and 1.16 respectively) compared with
cv Moneymaker (0.27) and other landraces. The differences in
the -carotene/lycopene ratio can be attributable to a possible
different genetic background of the landraces tested, which
might affect either the -carotene or lycopene biosynthesis.
Regarding this point, further molecular analyses are needed to
better elucidate possible genetic variations, which could explain
the differences in the amounts of these phytochemicals. In addi-
tion, harsh environmental conditions, such as high temperature,
can also impact lycopene accumulation in tomato fruits.
Brandt et al.,
for example, reported decreased lycopene biosyn-
thesis under high temperatures in the tomato F
Flavonoids, such as quercetin-3-rutinoside (rutin), and hydroxy-
cinnamic acids (such as chlorogenic, ferulic, and caffeic acids) are
the most representative phenolic compounds of tomatoes.
These compounds are characterized by the presence of phenolics
rings and hydroxyl groups in their structure that can scavenge
free radicals,
thus inhibiting the generation of ROS. In this
study, no signicant differences were detected in the amount of
Figure 2. Content of the main carotenoids in tomato fruits of different landraces and the cv Moneymaker (M). Analyses were carried out by HPLC on fruit
extracts. Statistical analysis was carried out by comparing the content of each compound with that recorded in the cv Moneymaker (M). Data are mean
plus/minus SD (n=3). *P<0.05 (Student's t-test).
Figure 1. AsA content in the red ripe fruit of the ten tomato landraces and the control variety Moneymaker. Data are showed as mean ±SD (n=3).
*P<0.05, **P<0.01 (Student's t-test).
Selection of tomato landraces with high fruit yield
J Sci Food Agric 2020; 100: 27912799 © 2020 The Authors.
Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.
chlorogenic acid among the ten landraces and the commercial cv
Moneymaker (Fig. 3). However, signicantly higher rutin levels
were detected in E36, E94, and PDVIT, compared with the control
cultivar (Fig. 3). A lower rutin level was instead detected in E32. In
the majority of the landraces analyzed in this study, higher levels
of kaempferol-O-rutinoside, one of the most common derivatives
of kaempferol in tomato fruit, were detected in E7, E17, E30,
E32, E36, E94, and PDVIT in comparison with Moneymaker
(Fig. 4). A higher kaempferol-O-glucoside content was also
detected in E94. Low levels of other phenolic compounds, such
as naringenin, were overall detected in the majority of land-
races (Fig. 4).
The higher polyphenols levels detected in some of the
tomato landraces analyzed could also be explained as a
stronger response to abiotic stresses, including heat
stress and exposure to UV radiation. Indeed, changes in
the polyphenols content have already been reported
in tomato following UV exposure.
In fact, genes implicated
in polyphenol biosynthesis are activated by light exposure,
and a sunscreenfunction has been proposed for these phy-
tochemicals to protect the tissues from possible damage gen-
erated by UV radiation.
Some authors reported that high
temperatures and light exposure stimulate the production of
phenolic acids and other avonoids.
Indeed, heat stress
positively modulates the activity of the enzyme phenylalanine
ammonia-lyase and affects the total phenols content by acti-
vating their biosynthesis and inhibiting their oxidation in
tomato plants.
Figure 4. Polyphenols content in tomato fruits from different landraces and the cv Moneymaker (M). Quantication was carried out by HPLC using
methanolic extracts of fruits. Statistical analysis was carried out by comparing the content of each compound with that recorded in the cv Moneymaker
(M). Data are mean plus/minus SD (n=3). *P<0.05, **P<0.01 (Student's t-test).
Figure 3. Chlorogenic acid and rutin content in fruits of different landraces and the cv Moneymaker (M). Quantication was carried out by HPLC using
fruit methanolic extracts. Statistical analysis was carried out by comparing the content of each compound with thatrecorded in the cv Moneymaker (M).
Data are mean plus/minus SD (n=3). *P<0.05, **P<0.01 (Student's t-test). A Scarano et al. © 2020 The Authors.
Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.
J Sci Food Agric 2020; 100: 27912799
Antioxidant/anti-inammatory properties of tomato
The antioxidant capacity of methanol extracts from fruits of
the ten tomato landraces is reported in Fig. 5(a), using the
TEAC assay. Previous studies reported that rutin and phenolic
acids can contribute to the overall antioxidant capacity of
tomato, neutralizing free radicals by acting as electron donors
or inhibiting the activity of enzymes involved in the produc-
tion of free radicals.
Our results indicated a slight, not signif-
icant, higher antioxidant activity in E32, followed by E17 and
E30. Conversely, a signicant lower antioxidant activity was
evidenced in E53 and E76, which also showed a lower polyphe-
nols content (see Fig. 4). E76 also showed a lower AsA content
than in the other landraces. Indeed, it is well known that the
antioxidant property of a food matrix is dependent on the
presence and levels of different compounds comprising phe-
nolic species and AsA.
To assess the anti-inammatory potential and the ability to trig-
ger an in vitro immune-modulating response, tomato extracts
were incubated with murine dendritic cells. LPS was used as an
inammatory stimulant. Figure 5(b) shows the effects of the
administration of tomato extracts on the production of pro-
inammatory IL-6 and IL-12. Signicantly lower levels of IL-6
were detected in the presence of E7 and E8 extracts. Both these
landraces showed increased levels of AsA, which could be
related to the decreased levels of the pro-inammatory activity
of IL-6. E8, together with E94 and PDVIT extracts, was also able
to decrease the secretion of the pro-inammatory IL-12 (Fig. 5
(c)) even though the differences observed were not signicant.
Some studies have reported the reduction of pro-inammatory
interleukins in dendritic cells mediated by the aglycone querce-
tin, thus demonstrating an anti-inammatory activity for this
In this context, rutin, which is the most
abundant phenolic compound in our quantication, could be
responsible, at least in part, for the anti-inammatory activity
detected in the tomato extracts. However, our data did not
establish a clear correlation between the content in phenolic
compounds and the anti-inammatory activity in all the land-
races tested. Indeed, the role of avonoids has been generally
described using chemically pure standards, only partially reect-
ing the real anti-inammatory activity that can be exerted by a
whole fruit or vegetable extracts.
Combining yield and quality parameters
To examine the possible co-occurrence of both yield and quality
features in the landraces tested, we calculated Spearman correla-
tions considering all the traits included in this study (Fig. 6(a)).
Positive correlations, signicant in some cases, were found among
the secondary metabolites analyzed (AsA, carotenoids, phenolic
acids, and avonoids), indicating that the observed changes
within the plant secondary metabolism involve the cross-talk
among different classes of phytochemicals and thus different bio-
chemical pathways. Furthermore, some of these metabolites pos-
itively correlated to physicochemical parameters such as acidity,
degrees Brix, and fruit rmness, indicating an association between
secondary metabolites and traits inuencing fruit organoleptic
An estimated EI was also calculated considering the quality
and nutritional traits analyzed all together. EI varied from a
minimum of 123 for E17 to a maximum of 175 for PDVIT, with
a mean value of 141.2. The distribution of the ten landraces
according their EI value and yield is shown in Fig. 6(b). From
this analysis, it is evident that three landraces (E30, E94,
PDVIT) exhibit better performances in terms of fruit quality
(EI >160), though two of them (E30 and E94) showed low
yield (less than 2 kg per plant) under high temperatures. All
the better performing landraces in terms of yield (classied
as high or medium producers), with the exception of PDVIT,
exhibited lower EI levels, thus evidencing lower values of
quality traits.
In this paper, one landrace (PDVIT) was selected as a good com-
promise between yield performances and good fruit quality and
nutritional traits when growing under high temperature. Addi-
tional molecular and physiological analyses in other environ-
ments are in progress in order to further characterize this
selected landrace. Indeed, this landrace shows a level of adapt-
ability that can be useful in adverse conditions, making it a suit-
able candidate for breeding programs, since it can be
Figure 5. Antioxidant and anti-inammatory activities of tomato extracts
from different landraces. (a) Antioxidant capacity proles measured by
TEAC. Data are mean μmol TE ±SD (n=3), *P<0.05 (Student's t-test).
Levels of pro-inammatory (b) IL-6 and (c) IL-12 in BMDCs stimulated with
LPS and treated with methanolic tomato extracts from fruits of different
tomato landraces. Concentrations of cytokines were determined by ELISA
test. Data are expressed as mean plus/minus SD (n=3); *P<0.05
(Student's t-test).
Selection of tomato landraces with high fruit yield
J Sci Food Agric 2020; 100: 27912799 © 2020 The Authors.
Journal of The Science of Food and Agriculture published by John Wiley & Sons Ltd on behalf of Society of Chemical Industry.
considered as a balanced landrace
in terms of stress resilience
and fruit nutritional quality.
This work was in part founded by the European Union's Horizon
2020 research and innovation program through the TomGEM pro-
ject under grant agreement no. 679796 and by the Apulia region
through the SICURA project (KC3U5Y1). We acknowledge Mr
Leone D'Amico for technical assistance.
The authors declare no conict of interest.
Supporting information may be found in the online version of this
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... CO 2 concentrations can influence the quantity of nutrients in food products (vitamin B, protein, and micronutrients). [49,[79][80][81] 12 ...
... Unlike most of them, some of these tomato landraces presented both good fruit size and setting and increased levels of phytonutrients. Such data indicate that the combination of tolerance to high temperatures with medium-high yield performances and fruit nutritional quality can be considered in the selection and breeding programs of local tomato landraces [79]. ...
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Plants are often exposed to different types of abiotic stress conditions such as salt stress, drought, low‐ and high‐temperature stress, heavy metal stress, and UV‐light stress. These stress conditions induce several morphological and physio‐biochemical alterations in plants that ultimately result in the significant reduction in growth and yield of plants. Abiotic stress conditions are principal inducers of oxidative stress in plants. In this process, free oxygen radicals overcome the free radical neutralization system resulting in the oxidative damage in the cell. In the oxidative stress conditions, huge increment in reactive species (ROS) occur. ROS and reactive nitrogen species (RNS) are produced in a systematic manner to ensure homeostasis at cell to tissue level and both serve as secondary messenger molecules in signaling pathways. ROS ( 1 O 2 , O 2 •− , OH •− , H 2 O 2 ) and RNS (ONOO − , NO) activate antioxidant defense system of plants, which plays role in scavenging of oxygen species by synthesizing enzymatic and nonenzymatic antioxidants resulting in the alleviation of damaging effects of these oxygen species. Different metallic micronutrients, such as Cu, Mg, Fe, Zn, Ni, and Mn, play a pivotal role as inhibitors, activators, or components of enzymes, whereas K, Ca, and Cl also showed the potential capability to participate in ROS metabolism. In this chapter, we considered important aspects of nutrients in ROS metabolism; however, limited information is available in the literature, and there is a dire need to explore more about the role of nutrients in ROS metabolism.
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The study was undertaken to evaluate the fruit quality parameters of six contrasting tomato genotypes including three susceptible (Arka Abha, IIHR-2627 and IIHR-2914) and three tolerant (IIHR-2202, IIHR-2745 and IIHR-2841) to high temperature stress. After flower initiation, tomato genotypes were grown under polytunnel to expose them to high temperature (40±2°C) till the time of harvesting. Uniformly ripen healthy fruits were harvested and analyzed for fruit quality parameters comprises total phenols, total flavonoids, antioxidant capacity in terms of FRAP (Ferric Reducing Antioxidant Potential) and DPPH (2,2-Diphenyl-1-picrylhydrazyl), total carotenoids, lycopene, total sugars, vitamin C, Total Soluble Solids (TSS) and Titrable Acidity (TA). Results revealed that there were significant differences between the genotypes in tomato fruit quality parameters. High temperature reduced total carotenoids and lycopene content in tomato fruits and it was decreased significantly in Arka Abha and IIHR-2914 compared to other genotypes. IIHR-2202 recorded very high antioxidant capacity in terms of FRAP, DPPH radical scavenging ability under high temperature stress. TA and TSS increased significantly in all the genotypes under high temperature stress. Tolerant genotypes recorded higher total phenols and total flavonoids content both under control and high temperature stress conditions. Genotypic variations were observed in the above stated biochemical parameters to high temperature stress. Based on these results, two genotypes namely IIHR-2841 and IIHR-2202 were found to be good at maintaining the all quality parameters under high temperature stress compared to Arka Abha and IIHR-2914.
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Background Tomato is an economically important crop with fruits that are a significant source of bioactive compounds such as ascorbic acid and phenolics. Nowadays, the majority of the enzymes of the biosynthetic pathways and of the structural genes controlling the production and the accumulation of antioxidants in plants are known; however, the mechanisms that regulate the expression of these genes are yet to be investigated. Here, we analyzed the transcriptomic changes occurring during ripening in the fruits of two tomato cultivars (E1 and E115), characterized by a different accumulation of antioxidants, in order to identify candidate genes potentially involved in the biosynthesis of ascorbic acid and phenylpropanoids. Results RNA sequencing analyses allowed identifying several structural and regulator genes putatively involved in ascorbate and phenylpropanoids biosynthesis in tomato fruits. Furthermore, transcription factors that may control antioxidants biosynthesis were identified through a weighted gene co-expression network analysis (WGCNA). Results obtained by RNA-seq and WGCNA analyses were further confirmed by RT-qPCR carried out at different ripening stages on ten cultivated tomato genotypes that accumulate different amount of bioactive compounds in the fruit. These analyses allowed us to identify one pectin methylesterase, which may affect the release of pectin-derived D-Galacturonic acid as metabolic precursor of ascorbate biosynthesis. Results reported in the present work allowed also identifying one L-ascorbate oxidase, which may favor the accumulation of reduced ascorbate in tomato fruits. Finally, the pivotal role of the enzymes chalcone synthases (CHS) in controlling the accumulation of phenolic compounds in cultivated tomato genotypes and the transcriptional control of the CHS genes exerted by Myb12 were confirmed. Conclusions By using transcriptomic analyses, candidate genes encoding transcription factors and structural genes were identified that may be involved in the accumulation of ascorbic acid and phenylpropanoids in tomato fruits of cultivated genotypes. These analyses provided novel insights into the molecular mechanisms controlling antioxidants accumulation in ripening tomato fruits. The structural genes and regulators here identified could also be used as efficient genetic markers for selecting high antioxidants tomato cultivars. Electronic supplementary material The online version of this article (10.1186/s12864-019-5428-4) contains supplementary material, which is available to authorized users.
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High temperature is one of the most detrimental abiotic stresses in tomatoes. Many studies highlighted that even small increases in temperature can alter the plant reproductive system, causing a significant reduction in tomato yield. The aim of this study was to exploit the phenotypic and genomic variations of a tomato landrace collection grown at high temperatures. Fifteen genotypes were selected as the best performing in two experimental fields. The selection was based on six yield-related traits, including flower earliness, number of flowers per inflorescence, fruit set, number of fruit per plant, fruit weight and yield per plant. In order to identify markers targeting traits that could be highly influenced by adverse climate conditions, such as flowering and fruit setting, an association mapping approach was undertaken exploiting a tomato high-throughput genomic array. The phenotypic variability observed allowed us to identify a total of 15 common markers associated with the studied traits. In particular, the most relevant associations co-localized with genes involved in the floral structure development, such as the style2.1 gene, or with genes directly involved in the response to abiotic stresses. These promising candidate genes will be functionally validated and transferred to a cultivated tomato to improve its performance under high temperatures
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Salt stress generally induces important negative effects on tomato (Solanum lycopersicum) productivity but it may also cause a positive effect improving fruit quality, one of the greatest challenges in nowadays agriculture. Because of the genetic erosion of this horticultural species, the recovery of locally adapted landraces could play a very important role in avoiding, at least partially, production losses and simultaneously improving fruit quality. Two tomato landraces endemic of the Spanish Southeast area, characterized by the harsh climatic conditions of the Mediterranean basin, have been selected: Negro Yeste (NY) characterized by its dark-red colored fruits and Verdal (V), which fruits did not achieve the characteristic red color at ripening. Here the agronomic, physiological, and metabolic responses of these landraces were compared with the reference tomato commercial cv. Moneymaker (MM), in plants grown without salt (control) and with salt stress (100 mM NaCl) for 70 days. The higher salt tolerance of both landraces was mainly reflected in the fruit number, as NY only reduced the fruit number in salt stress by 20% whereas in MM it was reduced till 43%, and in V the fruit number even showed an increase of 33% with salt stress. An important fruit quality parameter is soluble solids content, which increases induced by salinity were significantly higher in both landraces (60 and 78% in NY and V, respectively) compared with MM (34%). Although both landraces showed a similar response in relation to the high chlorophyll accumulation detected in their fruits, the fruit metabolic profiles were very different. Increased carotenoids levels were found in NY fruits, especially lycopene in ripe fruit, and this characteristic was observed in both control and salt stress. Contrarily, the carotenoid biosynthesis pathway was disrupted in V ripe fruits, but other metabolites, such as Ca2+, mannose, formate, and glutamate were accumulated. These results highlight the potential of tomato landraces to improve nutritional fruit quality and maintain fruit yield stability under salt stress.
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Flavonoids represent a wide group of plant secondary metabolites implicated in many physiological roles, from the attraction of pollinators to the protection against biotic or abiotic stresses. Flavonoids are synthetized in a number of horticultural crops that are important components of our daily diet. In the last decades, the consumption of vegetables rich in antioxidants has been strongly promoted from the perspective of prevention/protection against chronic diseases. Therefore, due to their nutritional importance, several attempts have been made to enhance flavonoid levels in species of agronomic interest. In this review, we focus on the flavonoid biodiversity among the major horticultural species, which is responsible of differences among closely related species and influences the qualitative/quantitative composition. We also review the role of flavonoids in the nutritional quality of plant products, contributing to their organoleptic and nutritional properties, and the main strategies of biofortification to increase their content.
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This narrative mini- review summarizes current knowledge of the role of polyphenols in health outcomes—and non-communicable diseases specifically—and discusses the implications of this evidence for public health, and for future directions for public health practice, policy, and research. The publications cited originate mainly from animal models and feeding experiments, as well as human cohort and case-control studies. Hypothesized protective effects of polyphenols in acute and chronic diseases, including obesity, neurodegenerative diseases, type 2 diabetes, and cardiovascular diseases, are evaluated. Potential harmful effects of some polyphenols are also considered, counterbalanced with the limited evidence of harm in the research literature. Recent international governmental regulations are discussed, as the safety and health claims of only a few specific polyphenolic compounds have been officially sanctioned. The implications of food processing on the bioavailability of polyphenols are also assessed, in addition to the health claims and marketing of polyphenols as a functional food. Finally, this mini-review asserts the need for increased regulation and guidelines for polyphenol consumption and supplementation in order to ensure consumers remain safe and informed about polyphenols.
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Constant global warming is one of the most detrimental environmental factors for agriculture causing significant losses in productivity as heat stress (HS) conditions damage plant growth and reproduction. In flowering plants such as tomato, HS has drastic repercussions on development and functionality of male reproductive organs and pollen. Response mechanisms to HS in tomato anthers and pollen have been widely investigated by transcriptomics; on the contrary, exhaustive proteomic evidences are still lacking. In this context, a differential proteomic study was performed on tomato anthers collected from two genotypes (thermo-tolerant and thermo-sensitive) to explore stress response mechanisms and identify proteins possibly associated to thermo-tolerance. Results showed that HS mainly affected energy and amino acid metabolism and nitrogen assimilation and modulated the expression of proteins involved in assuring protein quality and ROS detoxification. Moreover, proteins potentially associated to thermo-tolerant features, such as glutamine synthetase, S-adenosylmethionine synthase and polyphenol oxidase, were identified.
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Climate change has become a serious threat for crop productivity worldwide. The increased frequency of heat waves strongly affects reproductive success and thus yield for many crop species, implying that breeding for thermotolerant cultivars is critical for food security. Insight into the genetic architecture of reproductive heat tolerance contributes to our fundamental understanding of the stress sensitivity of this process and at the same time may have applied value. In the case of tomato (Solanum lycopersicum), germplasm screenings for thermotolerance have often used yield as the main measured trait. However, due to the complex nature of yield and the relatively narrow genetic variation present in the cultivated germplasm screened, there has been limited progress in understanding the genetic basis of reproductive heat tolerance. Extending the screening to wild accessions of related species that cover a range of climatic conditions might be an effective approach to find novel, more tolerant genetic resources. The purpose of this study was to provide insight into the sensitivity of individual reproductive key traits (i.e. the number of pollen per flower, pollen viability and style protrusion) to heat-wave like long-term mild heat (LTMH), and determine the extent to which genetic variation exists for these traits among wild tomato species. We found that these traits were highly variable among the screened accessions. Although no overall thermotolerant species were identified, several S. pimpinellifolium individuals outperformed the best performing cultivar in terms of pollen viability under LTMH. Furthermore, we reveal that there has been local adaptation of reproductive heat tolerance, as accessions from lower elevations and higher annual temperature are more likely to show high pollen viability under LTMH.
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Dietary polyphenols are associated with a wide range of health benefits, protecting against chronic diseases and promoting healthy aging. Dietary polyphenols offer a complementary approach to the treatment of inflammatory bowel diseases (IBDs), a group of common chronic intestinal inflammation syndromes for which there is no cure. Tomato is widely consumed but its content of polyphenols is low. We developed a tomato variety, Bronze, enriched in three distinct classes of polyphenols: flavonols, anthocyanins, and stilbenoids. Using Bronze tomatoes as a dietary supplement as well as Indigo (high anthocyanins and flavonols), ResTom (high stilbenoids) and wild-type tomatoes, we examined the effects of the different polyphenols on the host gut microbiota, inflammatory responses, and the symptoms of chronic IBD, in a mouse model. Bronze tomatoes significantly impacted the symptoms of IBD. A similar result was observed using diets supplemented with red grape skin containing flavonols, anthocyanins, and stilbenoids, suggesting that effective protection is provided by different classes of polyphenols acting synergistically.
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The increase of L-Ascorbic Acid (AsA) content in tomato (Solanum lycopersicum) is a common goal in breeding programs due to its beneficial effect on human health. To shed light into the regulation of fruit AsA content, we exploited a Solanum pennellii introgression line (IL12-4-SL) harbouring one quantitative trait locus that increases the content of total AsA in the fruit. Biochemical and transcriptomic analyses were carried out in fruits of IL12-4-SL in comparison with the cultivated line M82 at different stages of ripening. AsA content was studied in relation with pectin methylesterase (PME) activity and the degree of pectin methylesterification (DME). Our results indicated that the increase of AsA content in IL12-4-SL fruits was related with pectin de-methylesterification/degradation. Specific PME, polygalacturonase (PG) and UDP-D-glucuronic-acid-4-epimerase (UGlcAE) isoforms were proposed as components of the D-galacturonate pathway leading to AsA biosynthesis. The relationship between AsA content and PME activity was also exploited in PMEI tobacco plants expressing a specific PME inhibitor (PMEI). Here we report that tobacco PMEI plants, altered in PME activity and degree of pectin methylesterification, showed a reduction in low methylesterified pectic domains and exhibited a reduced AsA content. Overall, our results provide novel biochemical and genetic traits for increasing antioxidant content by marker-assisted selection in the Solanaceae family.