Chapter 37

Application of Polyamines to Maintain Functional

Properties in Stored Fruits
Marı́a Serrano and Daniel Valero

Abstract
Polyamines are natural compounds involved in many growth and developmental processes with ubiquitous
presence in all cells. Research in fruits has been developed to get a better understanding of the role of
polyamines, both endogenous and exogenous, especially during the ripening and senescence processes.
However, in recent years and given the relationship between fruit consumption and human health, the
study of antioxidant compounds responsible for these beneficial effects is of increasing interest.
This chapter focuses on the role of polyamines on the content of bioactive compounds with antioxidant
activity as well as in the activities of the main antioxidant enzymes in fruits.

Key words Postharvest, Putrescine, Spermidine, Spermine, Antioxidants, Phenolics, Anthocyanins,

Carotenoids, Antioxidant enzymes

1 Introduction

The polyamines have a long history, and the first evidence was
obtained in 1678 by Antonie van Leuwenhoek which described
the presence of crystalline substances in the human semen, but was
in 1878 when these crystals were identified as spermine by Lauden-
burg and Abel [1]. In 1924 putrescine (Put), spermidine (SPd),
and spermine (Spm) were synthesized [2]. During the mid-
twentieth century the experiments with polyamines were carried
out in bacteria while experiments with animals appeared in the
1970s with special interest in the association between polyamines
and diseases and largely focused on cancer cells leading to a synthe-
sis of the inhibitor of polyamine synthesis DFMO (α-difluoro-
methylornithine). In plants, the first evidence to the occurrence
of Put was dated in 1911 but was in 1971 [3] when it was con-
cluded that Put, Spd, and Spm are ubiquitous organic cations in
higher plants.
Polyamines are involved in many aspects of plant development
and considered as important molecules associated with both abiotic

Rubén Alcázar and Antonio F. Tiburcio (eds.), Polyamines: Methods and Protocols, Methods in Molecular Biology,
vol. 1694, DOI 10.1007/978-1-4939-7398-9_37, © Springer Science+Business Media LLC 2018

449
450 Marı́a Serrano and Daniel Valero

and biotic stresses, the most studied in fruits being chilling injury
(CI). Thus, polyamines have been reported to stimulate cell divi-
sion, dormancy breaking, germination, development of flower
buds, fruit set and growth [4–8]. However, one of the main aspects
of the relationship between polyamines and fruit are those related
to fruit ripening. Generally, there is an inverse relationship between
the content of endogenous polyamines and ethylene production
which is attributed to the fact that both share the common precur-
sor S-adenosylmethionine (SAM) in their biosynthesis pathway [6].
This relationship has justified the large body of knowledge about
the application of polyamines, either at pre-harvest (during fruit
growth and ripening on tree) or after harvest (postharvest treat-
ments) to delay the postharvest ripening process and maintain fruit
quality attributes such as colour, firmness, acidity, and total soluble
solids during postharvest storage, either at ambient temperature or
under cold storage [6, 9, 10]. However, the role of these treat-
ments on the content of bioactive compounds and antioxidant
activity has been poorly studied. In this chapter we present the
recent literature about the effect of polyamines as pre- or posthar-
vest treatments on the content of phytochemicals with antioxidant
properties in fruit, at harvest and during storage, given the health
beneficial effects attributed to these compounds.

2 Fruit and Health Beneficial Effects

Nowadays, it is widely accepted that nutrition has a great influence

on health and increasing evidence suggests that diets rich in fruits
and vegetables may prevent a wide range of diseases, mainly cardio-
vascular diseases (by reducing risk factors such as diabetes, hyper-
tension, and hyperlipidemia, platelet aggregation, blood pressure,
insulin resistance index, and obesity), several kind of cancers, such
as colon, oesophagus, oral cavity, stomach, pancreas, prostatic,
breast, and ovary (by regulating gene expression in cell proliferation
and apoptosis), neurodegenerative diseases, brain and immune
dysfunction, and even against bacterial and viral diseases [11–13].
These effects are due to phytochemical compounds with antioxi-
dant activity, the most important being phenolics, including antho-
cyanins, carotenoids, vitamins (C and E), and glucosinolates
[14–18]. In addition, it is important to note that whole fruit intake
provides more health beneficial effects than one of their constitu-
ent, because of additive and synergistic effects [13, 14].
Dietary polyamines from plant origin are considered very
important for human health. The body pool of polyamines in
human body is maintained by three sources: endogenous or de
novo biosynthesis, produced by the gut microbiota, and exogenous
intake through the diet [19]. From these sources, the polyamines
derived from the diet are the most important from the point of view
 Application of Polyamines to Maintain Functional Properties in Stored Fruits 451

of quantity, since the capacity to synthetize polyamines decreases

with age. Some of the health-beneficial effects of dietary polya-
mines are those related to protection against oxidative stress, main-
tenance of gut integrity, modulation of inflammation and immune
functions, among others [20]. It seems that polyamine antioxidant
activity leads to reduction in both cell membranes and DNA [21].
The average intake of dietary polyamines is different depending
on the country. In Europe, the highest intake of total polyamines
(700 μmol per day) has been estimated in the Mediterranean
regions compared with northern Europe. This difference could
justify the protective role attributed to the Mediterranean diet.
Among the different food groups, fruit provides the highest
amount of Put (500 μmol/kg) while cheese contains over
600 μmol/kg of Spd [19–21].

3 Polyamines, Bioactive Compounds and Antioxidant Activity in Fruits

3.1 Carotenoids Carotenoids are a group of natural-lipophilic pigments with a gen-

eral structure of C40 tetraterpenoids responsible for yellow to red
colour in fruits. The carotenoids are classified as carotenes (ß-
carotene, lycopene, etc.) and xanthophylls. In addition, carotenoids
can be acyclic (e.g. lycopene), monocyclic (γ-carotene), or dicyclic
(α- and ß-carotene). The most studied carotenoids have been ß-
carotene, lycopene, lutein and zeaxanthin which exhibit a potent
antioxidant activity. The antioxidant ability of these carotenoids
(carotenes and xanthophylls) follows the sequence from high to
low: lycopene > ß-cryptoxanthin
ß-carotene > lutein
zeaxan-
thin > α-carotene > canthaxanthin [6].
The first evidences regarding the effects of polyamines on
carotenoids content in fruits were obtained by Malik and Singh
[22] who found that mango tree treatments by foliar spraying with
Put, Spd, or Spm (0.01, 0.1, and 1 mM) at final fruit set stage
increased total carotenoids in the pulp at harvest time as compared
with fruits from control trees, the main effect being observed with
Put treatments (95%) followed by Spd (33%). More recently, Mehta
et al. [23] reported that ripe fruits from transgenic tomato plants
having the yeast SAMDC gene had higher Spd and Spm concentra-
tions and threefold more lycopene concentration than did the red
fruits from the parental lines. Accordingly, higher levels of poly-
amines and lycopene were found in transgenic tomatoes overex-
pressing human SAMDC gene. The increase in lycopene
concentration was attributed to enhanced levels of gene transcripts
involved in lycopene biosynthesis in transgenic tomatoes [24].
Similarly, plants overexpressing mouse ODC gene yielded tomato
fruits with higher polyamine concentration and higher concentra-
tion of lycopene as compared with fruits from control plants [25].
These effects of PAs are of special significance since carotenoids,
452 Marı́a Serrano and Daniel Valero

including lycopene, are bioactive compounds having antioxidant

capacity and proved beneficial effects in human health [11, 12, 26].
On the contrary, pre- and postharvest Put application to “Angel-
ino” plum led to a linear reduction in the levels of carotenoids,
which were more pronounced with increased concentrations of Put
and storage periods [27]. This effect could be due to the inhibition
of ethylene production and postharvest ripening process as a con-
sequence of Put treatments, since carotenoids increase during rip-
ening and during postharvest storage in plum fruits [6, 16].

3.2 Phenolic Phenolic or polyphenolics are secondary plant metabolites that

Compounds exhibit a wide range of physiological roles in plants such as pigmen-
tation, growth, and resistance to pathogens, among many other
functions. The main phenolics in fruits and vegetables are classified
according to their basic skeleton as C6-C1 (phenolic acids), C6-C3
(hydroxycinnamic acids), C6-C2-C6 (stilbenes), and C6-C3-C6 (fla-
vonoids). Phenolics as a group represent the strongest antioxidants
in plant foods, although the antioxidant activity of individual phe-
nolic compounds may vary depending on their chemical structure.
The antioxidant activity of phenolics is attributable to the electron
delocalization over the aromatic ring and their high redox poten-
tial, which allows them to act as reducing agents, hydrogen donors,
and singlet oxygen quenchers. Generally, total phenolics increase as
ripening advanced for most fruits, although concentration depends
on several factors including species, cultivar, growth conditions,
and environmental factors [6, 16, 28].
Different effects of polyamine treatment on phenolic content
of fruits have been reported depending on the applied polyamine,
concentration, moment of application, and fruit species. Thus, in
table grape postharvest Spm treatment at 0.5 and 1 mM maintained
total phenol concentration over the controls along 75 days of cold
storage, while Spm at 1.5 mM reduced significantly the berry
phenolic content [29]. On the other hand, postharvest Put treat-
ment at 2 mM of table grapes maintained higher concentration
(1.5-fold with respect to controls) of total phenolics during low-
temperature (0  C) storage [30]. These authors also measured the
individual concentration of catechin (flavonoid) and observed that
it was maintained at higher concentration (twofold) after 60 days of
cold storage +5 days of shelf life, which was correlated with the
higher total antioxidant activity obtained at the end of the experi-
ment. In other two table grape cultivars, polyamines have been also
applied as pre-harvest treatments (40 and 20 days before harvest) at
1 or 2 mM, and results revealed that treated berries showed higher
antioxidant activity and total phenolics at harvest and during cold
storage, although the effectiveness of the treatments was affected
by cultivar, polyamine type, and concentration [31].
In mango, total phenolics and total antioxidant capacity
increased during ripening at ambient temperature and during
 Application of Polyamines to Maintain Functional Properties in Stored Fruits 453

cold storage at 11  C although these increases were higher in fruits

treated with Put at 0.5, 1, 1.5, and 2 mM before storage [1]. By
other hand, in kiwifruit total phenolics decreased during storage at
room temperature, although the application of Spd or Spm (0.5, 1
and 1.5 mM), by dipping treatment previously to storage, led to
retention of total phenolic concentration along storage, the main
positive effect being found for Spm at 1.5 mM [33]. Similar results
were observed in pomegranate as a consequence of Put or Spd
postharvest treatments, Spd treatment being more effective than
Put treatment when applied as vacuum infiltration [34]. On the
contrary, Koushesh-saba et al. [35] reported lower phenolic con-
tent in Put- and Spd-treated apricots than in controls. Thus, the
effects of polyamines on total phenolic content (TPC) remain
elusive.

3.3 Anthocyanins Anthocyanins are the water-soluble pigments responsible for the
red, blue, and purple colour of fruits, and have been described as
potent antioxidants. Anthocyanins are located in the vacuole and
classified as flavonoids with glycosilated derivatives of the 3,5,7,30 -
tetrahydroxyflavylium cation. The free aglycones (anthocyanidins)
are highly reactive with sugars to form the glycosides and all antho-
cyanins are O-glycosilated. The main aglycones found in fruits are
pelargonidin, cyanidin, peonidin, delphinidin, petunidin, and mal-
vidin while the most relevant sugars are D-glucose, L-rhamnose, D-
galactose, D-xylose, and arabinose [36]. Anthocyanins have shown
higher antioxidant activity than other phenolic compounds, with
cyanidin being the most common anthocyanidin and the 3-
glucoside the most active anthocyanin with antioxidant activity [6].
Pre-harvest treatment with Put or Spd (at 1 or 2 mM) of
“Olhoghi” and “Rishbaba” table grape cultivars led to berries
with higher content of anthocyanins at harvest, especially for
2 mM Put [31]. These authors also found that after 55 days of
cold storage, anthocyanin concentrations had decreased in control
berries while remained at significant higher levels in all Put- or Spd-
treated ones, probably due to the effect of polyamines on delaying
the postharvest ripening process. In table grape “Flame Seedless”
cultivar, it has been reported that postharvest Spm treatment (0.5,
1 and 1.5 mM) induced a progressive increase in total anthocyanins
during prolonged storage, while in control samples there was an
initial increase (during 45 days of storage) but a sharp decrease
occurred during next 30 days (end of the experiment) of storage.
Interestingly, the effect of Spm treatment on increasing berry
anthocyanin concentration was dose-dependent [29].
In addition, in “Mollar de Elche” pomegranate, postharvest
treatments with Put or Spd at 1 mM (either by dipping or infiltra-
tion under low pressure) maintained the concentration of aril
anthocyanins during cold storage with respect to controls [34]. In
other pomegranate cultivar (“Mridula”) treatment by immersion
454 Marı́a Serrano and Daniel Valero

with 2 mM Put retained also higher anthocyanin during posthar-

vest storage leading to arils with higher antioxidant activity [37].
The mechanism by which Put and Spd induce these effects is still
unknown, although they may be related to their antisenescent
effects related to the suppression of membrane lipid peroxidation
and maintenance of the integrity of membranes [6, 38].

3.4 Vitamins Vitamins are a group of nutrients necessary for human body due to
their biochemical and physiological functions. Vitamins are classi-
fied into lipid-soluble and water soluble, the vitamins A, D, E, and
K being lipophilic, while C and B are hydrophilic. Tocopherols
(vitamin E) are the major lipid-soluble antioxidant vitamins in
fruits, while vitamin C is the major hydrophilic antioxidant vitamin,
although recent evidences indicate that vitamin D could also have a
role as antioxidant [39].
With respect to the relation and/or effect of polyamines in
ascorbic acid content of fruits, it has been found that in transgenic
tomato plants, overexpressing mouse ODC gene, tomato fruits had
higher polyamine concentration than those of the wild type, which
was correlated with an increase in ascorbic acid content as com-
pared with fruits from control plants [25]. Similarly, genetic modi-
fication of tomato fruit ripening by overexpressing human-SAMDC
led to higher endogenous levels of polyamines and enhancement of
ascorbic acid content [24]. This increase in ascorbic acid content in
transgenic tomatoes was attributed to their lower ethylene produc-
tion with respect to the wild type, since ascorbic acid is used as a
cofactor for 1-aminociclopropane-1-carboxilic acid oxidase, the last
enzyme in the ethylene biosynthesis pathway.
By other hand, in two tomato cultivars, postharvest treatments
with Put or Spd (at 1 or 2 mM) did not affect ascorbic acid
concentration, although the combination of both polyamines sig-
nificantly increased the level of ascorbic acid during 15 or 25 days of
storage at 2  C, especially for the combination of Put
1 mM + Spd2 mM [40].
In pomegranate, treatment with Spd alone or in combination
with calcium chloride showed a net increase in ascorbic acid content
in the arils during 4 months of storage at 2  C [41]. Accordingly,
postharvest treatment of pomegranates with 2 mM Put alone or in
combination with carnauba wax revealed that ascorbic acid declin-
ing trend was much pronounced in control as compared to that
found in treated fruit arils, either in pomegranates stored at chilling
temperature (3  C) or at safe temperature (5  C). Among the
applied treatments, the combination of Put and carnauba wax
gave the best results in terms of ascorbic acid retention at 3 and
5  C storage temperatures [37]. Postharvest treatments with Put or
Spd at 1 mM (either by immersion or vacuum-infiltration) in
“Mollar de Elche” pomegranate induced higher content of ascorbic
acid which was detected immediately 1 day after treatments and
 Application of Polyamines to Maintain Functional Properties in Stored Fruits 455

lasted along storage, the effect being higher in those fruits treated
under vacuum-infiltration [34].
These effects of PAs increasing or maintaining high ascorbic
acid content during storage are of special interest, since ascorbic
acid is a bioactive compound with antioxidant capacity and proved
beneficial effects in human health [11, 12].

4 Polyamines and Antioxidant Enzymes

In plant cells, the reactive oxygen species (ROS), such as superoxide

radical (O2 #), peroxide radical (O2 2#), hydrogen peroxide
l l

(H2O2), and hydroxyl radical (OH #), are randomly generated as

l

a consequence of normal metabolism, mainly in reactions catalyzed

by oxidase and lipoxygenase and in ß-oxidation of fatty acids. The
ROS content in plant cell is dependent on their producing systems
and scavenging mechanism, both enzymatic and non-enzymatic
ones [42]. As commented above, non-enzymatic antioxidant com-
pounds are reduced forms of ascorbate and glutathione, tocopher-
ols, phenolics, alkaloids, and carotenoids. In addition, at
physiological concentrations, polyamines are potent scavengers of
hydroxyl radicals while Spd and Spm are also able to quench both
singlet oxygen and hydrogen peroxide [43]. By other hand, enzy-
matic scavenging mechanisms include mainly superoxide dismutase
(SOD), catalase (CAT), peroxidase (POD), and ascorbate peroxi-
dase (APX). SOD detoxifies O2 # free radicals by converting them
l

to O2 and H2O2, which is further converted to H2O and O2 by

CAT, APX, and POD. CAT catalyzes the decomposition of hydro-
gen peroxide to water and oxygen, while APX uses ascorbate and
H2O2 as substrates producing water and dehydroascorbate as pro-
ducts, the last one being converted to ascorbate by glutathione
reductase enzyme. In addition, H2O2 can be also reduced to
water by POD, by using organic molecules such as phenols as
electron donor [44].
In this sense, it has been reported that treatment of cherry
tomatoes with arginine, the amino acid precursor of polyamines,
resulted in increased SOD, CAT, and APX activities compared with
the control, which were associated with the reduction of chilling
injury development in treated fruit [45]. Moreover, in tomato
plants under drought stress, an increase in Put and Spd concentra-
tions were found in tomato fruit, as well as in the activities of SOD
and CAT, the latter being responsible for the reduction in the
oxidative damage induced by the stress conditions [46].
In mango, the activity of SOD, POD and CAT increased
during ripening at ambient temperature and during cold storage
at 11  C although these increases were higher after treatments with
Put at 0.5, 1, 1.5, and 2 mM [32]. Similarly, higher activities of
CAT, POD and SOD were found in two apricot cultivars
456 Marı́a Serrano and Daniel Valero

(“Bagheri” and “Asgarabadi”) during storage at 1  C plus 2 days at

20  C treated with 1 mM Put or Spd than in control fruits, which
were related to a higher resistance to stress and a longer commercial
life [35]. These studies reveal that postharvest polyamine treatment
increases the ability of fruit tissues to eliminate ROS and contri-
butes to alleviate the chilling injury (CI) symptoms observed in
control fruits.
Thus, the effects of exogenous application of polyamines on
increasing the antioxidant enzyme activities would lead to preserve
membrane integrity and to reduce accumulation of ROS and in
turn to alleviate fruit stress, especially chilling injury, as well as to
delay the postharvest ripening process with additional benefits on
maintaining fruit quality properties during postharvest storage.

5 Conclusions

In this chapter we provide the recent information regarding the

effect of polyamines on maintaining functional properties in stored
fruits. With respect to the non-enzymatic antioxidant systems, the
exogenous application of polyamines, as pre- or postharvest treat-
ments, generally resulted in increases on the content of antioxidant
compounds such as phenolics (including anthocyanins), carote-
noids, and vitamins, especially ascorbic acid. On the other hand,
polyamines are also effective on increasing the activity of several
antioxidant enzymes, such as SOD, CAT, POD, and APX. Both
systems could contribute to cleaning the ROS generated during the
postharvest ripening process, and in turn to delay the postharvest
ripening and senescence processes and extending the shelf life of
fruits. Nevertheless, the precise physiological and molecular
mechanisms by which polyamines increase both antioxidant sys-
tems remain elusive and deserve further research.

References

1. Bachrach U (2010) The early history of poly-
amine research. Plant Physiol Biochem
48:490–495
2. Rosenheim O (1924) The isolation of sper-
mine phosphate from semen and testis. Bio-
chem J 18:1253–1263
3. Smith TA (1971) The occurrence, metabolism
and functions of amines in plants. Biol Rev
Camb Philos Soc 46:201–204
4. Alburquerque N, Egea J, Burgos L, Martı́nez-
Romero D, Valero D, Serrano M (2006) The
influence of polyamines on apricot ovary devel-
opment and fruit set. Ann Appl Biol
149:27–33
5. Groppa MD, Benavides MP (2008) Polya-
mines and abiotic stress: recent advances.
Amino Acids 34:35–45
6. Valero D, Serrano M (2010) Postharvest biol-
ogy and technology for preserving fruit quality.
CRC/Taylor & Francis, Boca Raton
7. Asadi R, Ardebili ZO, Abdossi V (2013) The
modified fruit quality by application of differ-
ent kinds of polyamines in apricot tree (Prunus
armeniaca). J Appl Environ Biol Sci 3:28–31
8. Tiburcio AF, Altabella T, Bitrián M, Alcázar R
(2014) The roles of polyamines during the life-
span of plants: from development to stress.
Planta 240:1–18
 Application of Polyamines to Maintain Functional Properties in Stored Fruits
9. Valero D, Martı́nez-Romero D, Serrano M
(2002) The role of polyamines in the improve-
ment of the shelf life of fruit. Trends Food Sci
Technol 13:228–234
10. Serrano M, Zapata PJ, Martı́nez-Romero D,
Dı́az-Mula H, Valero D (2016) Polyamines as
an eco-friendly postharvest tool to maintain
fruit quality. In: Siddiqui MW (ed) Eco-
friendly technology for postharvest produce
quality. Elsevier Inc., London, UK
11. Martin C, Zhang Y, Tonelli C, Petroni K
(2013) Plants, diet, and health. Annu Rev
Plant Biol 64:19–46
12. Nile SH, Park SW (2014) Edible berries: bio-
active components and their effect on human
health. Nutrition 30:134–144
13. Rodriguez-Casado A (2016) The health poten-
tial of fruits and vegetables phytochemicals:
notable examples. Crit Rev Food Sci Nutr
56:1097–1107
14. Grosso G, Galvano F, Mistretta A, Marventano
S, Nolfo F, Calabrese G, Buscemi S, Drago F,
Veronesi U, Scuderi A (2013) Red orange:
experimental models and epidemiological evi-
dence of its benefits on human health. Oxid
Med Cell Longev. Article ID 157240. doi:10.
1155/2013/157240
15. Sharma P (2013) Vitamin C rich fruits can
prevent heart disease. Ind J Clin Biochem
28:213–214
16. Valero D, Serrano M (2013) Growth and rip-
ening stage at harvest modulates postharvest
quality and bioactive compounds with antioxi-
dant activity. Stewart Postharvest Rev 3:5
17. Woodside JV, McGrath AJ, Lyner N, McKinley
MC (2015) Carotenoids and health in older
people. Maturitas 80:63–68
18. Zanotti I, Dall’Asta M, Mena P, Mele L, Bruni
R, Ray S, Del Rio D (2015) Atheroprotective
effects of (poly) phenols: a focus on cell choles-
terol metabolism. Food Funct 6:13–31
19. Ali MA, Poortvliet E, Strömberg R, Yngve A
(2011) Polyamines in foods: development of a
food databse. Food Nutr Res 55:5572. doi:10.
3402/fnr.v55i05572
20. Hunter DC, Burritt DJ (2012) Polyamines of
plant origin – an important dietary consider-
ation for human health. In: Rao V (ed) Phyto-
chemical as nutraceuticals – global approaches
to their role in nutrition and health.
Chapter 12. InTech, Croatia. doi:10.5772/
2375
21. Kalač P (2014) Health effects and occurrence
of dietary polyamines: a review for the period
2005-mid 2013. Food Chem 161:27–39
22. Malik AU, Singh Z (2006) Improved fruit
retention, yield and fruit quality in mango
457
with exogenous application of polyamines. Sci
Hortic 110:167–174
23. Mehta RA, Cassol T, Li N, Ali N, Handa AK,
Mattoo AK (2012) Engineered polyamine
accumulation in tomato enhances phytonutri-
ent content, juice quality, and vine life. Nat
Biotechnol 20:613–618
24. Madhulatha P, Gupta A, Gupta S, Kumar A, Pal
RK, Rajam MV (2014) Fruit-specific overex-
pression of human S-adenosylmethionine
decarboxylase gene results in polyamine accu-
mulation and affects diverse aspects of tomato
fruit development and quality. J Plant Biochem
Biotechnol 23:151–160
25. Pandey R, Gupta A, Chowdhary A, Pal RK,
Rajam MV (2015) Overexpression of mouse
ornithine decarboxylase gene under the control
of fruit-specific promoter enhances fruit quality
in tomato. Plant Mol Biol 87:249–260
26. Friedman M (2013) Anticarcinogenic, cardio-
protective, and other health benefits of tomato
compounds lycopene, α-tomatine, and tomati-
dine in pure form and in fresh and processed
tomatoes. J Agric Food Chem 61:9534–9550
27. Khan AS, Singh Z, Abbasi NA, Swinny EE
(2008) Pre- or post-harvest applications of
putrescine at low temperature storage affect
fruit ripening and quality of ‘Angelino’ plum.
J Sci Food Agric 88:1686–1695
28. Serrano M, Dı́az-Mula HM, Valero D (2011)
Antioxidant compounds in fruits and vegeta-
bles and changes during postharvest storage
and processing. Stewart Postharv Rev 1:1
29. Harindra Champa WA, Gill MIS, Mahajan
BVC, Bedi S (2015) Exogenous treatment of
spermine to maintain quality and extend post-
harvest life of table grapes (Vitis vinifera L.) cv.
Flame Seedless under low temperature storage.
LWT Food Sci Technol 60:412–419
30. Shiri MA, Ghasemnezhad M, Bakshi D, Sari-
khani H (2013) Effect of postharvest putres-
cine application and chitosan coating on
maintaining quality of table grape cv. ‘Shah-
roudi’ during long-term storage. J Food Proc
Preserv 37:999–1007
31. Mirdehghan SH, Rahimi S (2016) Pre-harvest
application of polyamines enhances antioxi-
dants and table grape (Vitis vinifera L.) quality
during postharvest period. Food Chem
196:1040–1047
32. Razzaq K, Khan AS, Malik AU, Shahid M,
Ullah S (2014) Role of putrescine in regulating
fruit softening and antioxidative enzymes sys-
tems in ‘Samar Bahisht Chaunsa’ mango. Post-
harvest Biol Technol 96:23–32
33. Jhalegar MJ, Sharma RR, Pal RK, Rana V
(2012) Effect of postharvest treatments with
458 Marı́a Serrano and Daniel Valero
polyamines on physiological and biochemical
attributes of kiwifruit (Actinidia deliciosa) cv.
Allison Fruits 67:13–22
34. Mirdehghan SH, Rahemi M, Serrano M, Guil-
lén F, Martı́nez-Romero D, Valero D (2007)
The application of polyamines by pressure or
immersion as a tool to maintain functional
properties in stored pomegranates arils. J
Agric Food Chem 55:755–760
35. Koushesh sab M, Arzani K, Barzegar M (2012)
Postharvest polyamine application alleviates
chilling injury and affects apricot storage abil-
ity. J Agric Food Chem 60:8947–8953
36. Castañeda-Ovando A, Pacheco-Hernández
ML, Páez-Hernández ME, Rodrı́guez JA,
Galán-Vidal A (2009) Chemical studies of
anthocyanins: a review. Food Chem
113:859–871
37. Barman K, Asrey R, Pal RK, Kaur C, Jha SK
(2014) Influence of putrescine and carnauba
wax on functional and sensory quality of pome-
granate (Punica granatum L.) fruits during
storage. J Food Sci Technol 51:111–117
38. Lester GE (2000) Polyamines and their cellular
anti-senescence properties in honey dew musk-
melon fruit. Plant Sci 160:105–112
39. Asensi-Fabado MA, Munné-Bosh S (2010)
Vitamins in plants: occurrence, biosynthesis
and antioxidant function. Trends Plant Sci
15:582–592
40. Javanmardi J, Rahemi M, Nasirzadeh M
(2013) Post-storage quality and physiological
responses of tomato fruits treated with polya-
mines. Adv Hort Sci 27:173–181
41. Ramezanian A, Rahemi M, Maftoun M, Bah-
man K, Eshghi S, Safizadeh MR (2010) The
ameliorative effects of spermidine and calcium
chloride on chilling injury in pomegranate
fruits after long-term storage. Fruits
65:169–176
42. Apel K, Hirt H (2004) Reactive oxygen spe-
cies: metabolism, oxidative stress, and signal
transduction. Annu Rev Plant Biol
55:373–399
43. Mozdzan M, Szemra J, Rysz J, Stolarek RA,
Nowak D (2006) Anti-oxidant activity of sper-
mine and spermidine re-evaluated with oxidiz-
ing systems involving iron and copper ions. Int
J Biochem Cell Biol 38:69–81
44. Tareen MJ, Abbasi NA, Hafiz IA (2012) Post-
harvest application of salicylic acid enhanced
antioxidant enzyme activity and maintained
quality of peach cv. ’Flordaking’ fruit during
storage. Sci Hortic 142:221–228
45. Zhang X, Shen L, Li F, Zhang Y, Meng D,
Sheng J (2010) Up-regulating arginase contri-
butes to amelioration of chilling stress and the
antioxidant system in cherry tomato fruits. J Sci
Food Agric 90:2195–2202
46. Sánchez-Rodrı́guez E, Romero L, Ruı́z JM
(2016) Accumulation of free polyamines
enhances antioxidant responses in fruits of
grafted tomato plants under water stress. J
Plant Physiol 190:72–76