Scientia Horticulturae 216 (2017) 12–21

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Scientia Horticulturae
journal homepage: www.elsevier.com/locate/scihorti

Review

An overview of preharvest factors affecting vitamin C content of citrus

fruit
Lembe Samukelo Magwaza a,∗ , Asanda Mditshwa b , Samson Zeray Tesfay b ,
Umezuruike Linus Opara c,d
a
Department of Crop Science, School of Agricultural, Earth and Environmental Sciences, University of KwaZulu-Natal, Private Bag X01, Scottsville, 3209,
Pietermaritzburg, South Africa
b
Department of Horticultural Sciences, School of Agricultural, Earth and Environmental Sciences, University of KwaZulu-Natal, Private Bag X01, Scottsville,
3209, Pietermaritzburg, South Africa
c
Postharvest Technology Research Laboratory, South African Research Chair in Postharvest Technology, Department of Horticultural Science, Stellenbosch
University, Stellenbosch, 7602, South Africa
d
Postharvest Technology Research Laboratory, South African Research Chair in Postharvest Technology, Department of Food Science, Stellenbosch
University, Stellenbosch, 7602, South Africa

a r t i c l e i n f o a b s t r a c t

Article history: Citrus fruits are an important diet for human health by providing some nutritional and antioxidant com-
Received 13 September 2016 ponents, especially ascorbic acid (AsA). As the major antioxidant compound, the concentration of AsA is
Received in revised form an important nutritional quality indicator for citrus fruit and its derivative products. Although the empha-
20 December 2016
sis is on vitamin C content of harvested citrus fruit, this review of literature focuses on preharvest factors
Accepted 22 December 2016
affecting this nutritional quality variable. The literature reviewed in this study showed that the content
of AsA in citrus fruits is influenced by numerous preharvest factors including genotype, cultural practices,
Keywords:
climatic conditions, fruit maturity and postharvest management. To better understand the mechanism
Ascorbic acid
Biosynthesis
underlying the vitamin C content of different citrus fruits, the introduction of this paper will first provide
Nutritional quality an overview of major proposed biosynthetic pathways and genes encoding for enzymes involved in AsA
Antioxidant biosynthesis and oxidation. The review will then concentrate on the current state of knowledge regard-
Maturity ing inherent genetic, environmental, fertilization, maturity and harvesting factors affecting AsA of citrus
Canopy position fruits. Some suggestions for future research are also highlighted.
© 2016 Elsevier B.V. All rights reserved.

Contents

1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
2. Preharvest factors affecting AsA content . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
2.1. Cultivar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
2.2. Rootstocks . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
2.3. Rainfall and irrigation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
2.4. Light and temperature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
2.5. Fertilization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
2.5.1. Macronutrients . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
2.5.2. Micronutrients . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
2.5.3. Organic vs inorganic fertilizers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
2.6. Hormones and plant growth regulators . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
3. Harvesting factors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

∗ Corresponding author.
E-mail addresses: magwazal@ukzn.ac.za, lembe.magwaza@gmail.com (L.S. Magwaza).

http://dx.doi.org/10.1016/j.scienta.2016.12.021
0304-4238/© 2016 Elsevier B.V. All rights reserved.
 L.S. Magwaza et al. / Scientia Horticulturae 216 (2017) 12–21 13

3.1. Time of harvest (Maturity) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

4. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

1. Introduction and biotic stimuli (Davey et al., 2000, 2006; Tsaniklidis et al.,
2014).
Citrus fruit, juices and other derivative products are vastly The biological roles of AsA have been studied extensively
consumed worldwide and their consumption has increased con- and are well documented, however, the molecular mechanism
siderably over the past decade, particularly due to the association regulating its metabolism, biosynthesis, and levels in plant tis-
between a higher levels of the fruit in the diet and minimized sues is still not well understood (Smirnoff, 1996; Alós et al.,
threat of non-communicable diseases such as neurodegenerative 2014; Lado et al., 2015). Although biosynthesis is considered
and cardiovascular diseases, stroke, cataractogenesis, cancer and the leading source responsible for the accumulation of AsA in
diabetes (Cardeñosa et al., 2015). The benefits of citrus in the human plants tissues, similar to most biochemical compounds, its com-
diet are linked to high levels of bioactive components and antioxi- ponents and levels are controlled by the balance among its
dants capacity contributed by numerous groups of phytochemicals, biosynthesis, catabolism, and recycling (Ishikawa et al., 2006;
polyphenols, and vitamins in citrus fruit (Ladaniya, 2008). However, Yang et al., 2011). Several studies have evaluated the pathway
citrus products with potentially beneficial levels of phytochemi- for AsA biosynthesis in different plants and the general agree-
cals such as phenolics and flavonoids cannot currently be marketed ment is that there are four effective biosynthetic routes. The
based on antioxidant content. Only two vitamins, namely vitamin four accepted biosynthesis pathways are L-galactose, L-gulose,
C and carotenoids are commonly found in citrus fruits can be effec- the myoinositol and D-galacturonic acid pathways, named accord-
tively used in health-related marketing. ing to the molecules that act as precursors (Fig. 1). In general,
Ascorbic acid (AsA) also referred as vitamin C is among some AsA biosynthesis in citrus is mostly by the L-galactose pathway,
of the vital water-soluble vitamins, naturally occurring in various where AsA is produced from D-mannose-1-phosphate through a
vegetables and fruits and cannot be synthesized by human bodies sequence of enzymatic reactions catalyzed by enzymes comprising
(Chebrolu et al., 2012; Valente et al., 2014). Due to its high content of GDP-d-mannose pyrophosphorylase (GMP or VTC1); GDP-d-
of vitamin C and a large volume of per capita consumption, citrus mannose-3 -5 -epimerase (GME), GDP-l-galactose phosphorylase
fruit is undoubtedly the main contributor in human nutrition. As (GGP or VTC2), L-galactose-1-phosphate phosphatase (GPP or
such, citrus fruits are a significant source of AsA in the human diet. VTC4), L-galactose dehydrogenase (GalDH), and L-galactono-1,4-
As the major antioxidant compound, the concentration of vitamin lactone dehydrogenase (GalLDH) (Zhang et al., 2015). However,
C is an important nutritional quality indicator of citrus fruit and recent studies have also suggested AsA biosynthesis in citrus
its derivative products (Gardner et al., 2000; Zerdin et al., 2003). through the D-galacturonic acid pathway, which involves the activ-
The current recommended daily allowance (RDA) for AsA intake is ity of D-galacturonic acid reductase (GalUR). As shown in Fig. 1, AsA
100–120 mg/day to accomplish cellular requirement and reduced can also be regenerated by NADPH-dependent monodehydroascor-
risk of cardiovascular and neurodegenerative diseases, cancer and bate reductase (MDHAR) and dehydroascorbate reductase (DHAR)
stroke in healthy humans (Naidu, 2003). For instance, one glass of via the ascorbate–glutathione cycle (Agius et al., 2003; Ishikawa
orange juice (200 mL) have been found to deliver between 30 and et al., 2006; Xu et al., 2013; Alós et al., 2014; Lado et al., 2015;
80% of RDA of vitamin C (Gliszczyńska-Świgło et al., 2004). Zhang et al., 2015).
The common name “Vitamin C” is trivial for two biologically The content of AsA in citrus fruits can be influenced by numerous
active compounds, being ascorbic acid and its first oxidation prod- factors including genotype, preharvest cultural practices, climatic
uct, dehydroascorbic acid (DHA) (Gokmen et al., 2000). Therefore, conditions, fruit maturity, harvesting procedures, and postharvest
the total vitamin C content of citrus fruit is the sum of these management (Klein and Perry, 1982; Lee and Kader, 2000; Porat
two biologically active compounds, AsA and DHA (Nováková et al., et al., 2004; Marcilla et al., 2006). AsA is the least chemically inert
2008; Valente et al., 2014). Most physiological roles of AsA are nutrient, susceptible to postharvest loss, being subject to oxidative
linked to its capability to be a reducing agent in a biochemical and enzymatic degradation DHA and latter a permanent oxidation
reaction (Tsaniklidis et al., 2014). AsA is a low molecular weight to diketogulonic acid (DKG) with no vitamin C activity (Johnston
secondary compound, contributing to the antioxidant capacity of et al., 2007; Odriozola-Serrano et al., 2007). Thus, vitamin C of cit-
plant tissues, especially during different stressful environments rus fruit depends not only on the AsA content at harvest but also
(Fotopoulos et al., 2008). As an antioxidant, the primary func- on the changes occurring during postharvest handling, storage, pro-
tion of AsA is to detoxify reactive oxygen species (ROS) and free cessing, and preparation (Lee and Kader, 2000; Kalt, 2005; Rickman
radicals such as hydroxyl radical (OH− ), superoxide (O2 − ), sin- et al., 2007). Although the emphasis of this review is on vitamin C
glet oxygen (O2 − ) and hydrogen peroxide (H2 O2 ) produced during content of harvested citrus fruit, preharvest conditions significantly
metabolic processes of photosynthesis (Smirnoff, 2011). AsA is influencing this nutritional quality variable will be the main focus.
known as a cofactor for enzymes and also plays a role in the Since levels of vitamin C vary widely due to preharvest produc-
regulation of genes involved in defense mechanism and signal- tion conditions, this review will concentrate on the current state
ing in hormone pathways (Smirnoff, 1996; Lado et al., 2015). Like of knowledge regarding inherent genetic, environmental, fertiliza-
in many plants, its metabolism in citrus plays an important role tion, maturity and harvesting factors affecting AsA of citrus fruits.
in several physiological processes such as photosynthesis, growth To better understand the fundamental mechanism of the vitamin
regulation, and senescence. In many crops, the enhancement of C content of different citrus fruits, the introduction section of this
AsA concentration also extends shelf-life. Its accumulation in dif- review provides an overview of major proposed biosynthetic path-
ferent tissues and organs is interrelated with oxidative stress ways and regulation of genes coding for enzymes involved in AsA
defense mechanism and is altered by physiological phenomena biosynthesis and oxidation. Finally, this review provides some sug-
such as cell expansion, growth, senescence and, several abiotic gestions for future research are also highlighted.
14 L.S. Magwaza et al. / Scientia Horticulturae 216 (2017) 12–21

Fig. 1. A simple schematic diagram of ascorbic acid biosynthetic, catabolic and recycling pathways in plants. Four pathways have been proposed, namely, 1) the myoinositol, 2)
L-gulose, 3) L-galactose, and 4) D-galacturonic acid pathways according to the precursor molecules. The major L-galactose pathway is highlighted in green and the alternative
proposed D-galacturonic acid pathways highlighted in blue. The major enzymes catalyzing the reactions in these proposed pathways are shown in red, namely, phospho-
glucose isomerase (PGI); hexokinase (HXK); phosphomannomutase (PMM); phospho- mannose isomerase (PMI); GDP-d-mannose-3 -5 -epimerase (GME); GDP-l-galactose
phosphorylase (GGP); L-galactose-1-phosphate phosphatase (GPP); L-galactose dehydrogenase (GaLDH); L-galactono-1,4-lactone dehydrogenase (GalLDH); myoinositol oxy-
genase (MIOX); L-gulono-1,4-lactone oxidase (GLOase); D-galacturonic acid reductase (GalUR); monodehydroascorbate reductase (MDHAR), dehydroascorbate reductase
(DHAR); ascorbate peroxidase (APX); and ascorbate oxidase (AO). Adapted from Alós et al. (2014) and Lado et al. (2015). (For interpretation of the references to colour in this
figure legend, the reader is referred to the web version of this article.)

2. Preharvest factors affecting AsA content
There is enough literature suggesting that appropriate posthar-
vest management practices of citrus fruit can decrease postharvest
nutritional losses and enhance fruit quality. However, several stud-
ies have indicated that fruit biochemical composition and quality
are mostly affected by preharvest factors, such as scion-rootstock
combination, carbohydrate content (Goldschmidt, 1999), crop load
and fruit position within the tree canopy (Volz et al., 1993), light
and temperature (Ferguson et al., 1999). Genetic make-up, cultural
practices, and environmental conditions that result in an increase
in fruit ROS will also influence AsA content at harvest (Davey et al.,
 L.S. Magwaza et al. / Scientia Horticulturae 216 (2017) 12–21 15

Table 1 Table 2
Vitamin C concentration of different citrus types and cultivars. Vitamin C content in different edible tissues of citrus fruit of four species.

Citrus species Cultivar Vitamin C (mg/100 mL) Reference Citrus species Edible tissues Vitamin C (mg/100 g FW)

Mandarin Kinnow 18.2 Sharma C. unshiu Juice sack 25.4

Nagpur 16.2 et al. Segment membrane 32.6
Cleopatra 20.2 (2006) Segment 26.1
Kaula 19.5
C. reticulata Juice sack 45.3
Safor 29.0 Sdiri Segment membrane 14.2
Garbí 21.2 et al. Segment 38.9
Fortune 29.8 (2012)
C. sinensis Juice sack 32.9
Kara 25.5
Segment membrane 35.2
Murcott 23.5
Segment 35.2
Fortune 28.3 − 31.6 Lafuente et al. (2011)
C. changshanensis Juice sack 41.6
Sweet oranges Mosambi 22.2 Sharma
Segment membrane 15.7
Pineapple 22.8 et al.
Segment 32.7
Blood Red 20.8 (2006)
Valencia Late 18.9
Jaffa 21.5
Table 3
Lemon Eureka 39.6 Sharma Ascorbate contents in peel and pulp different citrus types and cultivars.
Lisbon 42.3 et al.
Kagzi Kalan 40.8 (2006) Pulp Peel
−1
Lime Kagzi 46.2 Sharma et al. (2006) mg mL
Lima orange 46.1 43.2 Barros et al. (2012)
Grapefruit Foster 23.6 Sharma Pera orange 68.1 24.3
Duncan 21.2 et al. Tahiti lime 41.4 6.84
Marsh 22.8 (2006) Sweet lime 60.2 22.6
Ponkan mandarin 41.1 47.6
Pummelo Local 19.2 Sharma
Kaoopan 21.6 et al. ␮mol g−1 FW
China 21.0 (2006) Egan No.2 orange 11.33 2.03 Yang et al. (2011)
Guoqing No.1 orange 8.76 1.56
Tangelo Thornton 19.2 Sharma et al. (2006)
Newhall mandarin 10.52 3.69
Dream mandarin 8.76 3.44

2000; Lee and Kader, 2000). This indicates the complexity of the
relationship between preharvest factors and biochemical quality
of harvested fruit.
2.1. Cultivar
The chemical and nutritional attributes of citrus fruit, including
carotenoids and AsA content, are mainly determined by multigenic
inheritance (Fanciullino et al., 2006). Different studies have shown
that the levels of AsA in citrus fruit juice sacs are highly variable,
ranging from 16.2 to 46.2 mg/mL juice depending on species and
cultivar (Table 1) (Sharma et al., 2006; Lafuente et al., 2011; Sdiri
et al., 2012).
Mandarins (Citrus unshiu Marc.) and oranges (Citrus sinensis
Osb.) are the two commercial species which have been shown
to have a significant difference in AsA levels (Nagy, 1980). The
AsA composition and content in the pulp of oranges is generally
higher than in mandarins. The lower contents of AsA in man-
darins compared to oranges were observed by Dhuique-Mayer
et al. (2005) and recently confirmed in an analysis by Yang et al.
(2011) which showed that the average AsA contents in fruit juice
sacs of mandarin and orange at the maturity stage were 27.3 and
48.9 mg/100 mL juice, respectively. Variances in AsA concentra-
tion between mandarins and oranges have been suggested to be
linked to the differentials in the expression of genes encoding
for enzymes involved in the L-galactose pathway during ripening
and the difference in the activity of AsA oxidation and recycling
enzymes (Yang et al., 2011 Alós et al., 2014). In general, Martí et al.
(2009) and Escobedo-Avellaneda et al. (2014) showed that oranges,
generally have the highest AsA levels (29 and 82 mg/100 mL
of juice), followed by lemons (30–50 mg/100 mL), grapefruits
(30–60 mg/100 mL) and mandarins (20–60 mg/100 mL).
Even more significant variations were found among citrus geno-
types and different edible tissues, where AsA content of juice sack
and segments of all tested species and cultivars ranged from 25.4
to 45.3 mg/100 g FW with juice segments having higher AsA than
juice sack (Table 2; Abeysinghe et al., 2010). Barros et al. (2012)
compared four citrus species (C. latifolia Tanaka cv. Tahiti; C. limet-
tioides Tanaka cv. Sweet lime; C. sinensis, cvs. Pera and Lima and
C. reticulate, cv. Ponkan) grown in Brazil in relation to the ascor-
bic acid of pulps and peels. In general, these authors reported the
pulps of most species to have significantly higher contents of AsA
except for ‘Ponkan’ mandarin, which had the highest AsA content
in the pulp. Yang et al. (2011) observed that total AsA in fruit pulp
was approximately one-fourth of that in the peel (Table 3). This
was in accordance with a previous report by Nagy (1980). Differ-
ences in AsA content between flavedo, albedo, and pulp tissues have
also been detected by Cano et al. (2008) and Bermejo et al. (2011)
and this has led to the generalization that the flavedo (the orange
coloured part of the peel) is the richest tissue compared to albedo
(the spongy white part of the peel) or fleshy pulp. The reason for
the differences in AsA between these fruit tissues is still not known.
However, it has been hypothesized that higher AsA contents in
the peel tissues may be attributed to the elevated requirement for
antioxidant compounds to counteract ROS generated by environ-
mental stresses which the fruit peel is directly exposed compared
to the pulp, located in the inner part of the fruit (Tausz et al., 2004;
Yang et al., 2011; Lado et al., 2015).
The ‘Pera’ oranges had the highest AsA content (68 mg/100 mL)
compared to the lowest levels recorded in ‘Tahiti’ lime and ‘Ponkan’
mandarin with 41.4 and 41.1 mg/100 mL, respectively (Barros et al.,
2012). These values fall within the range reported earlier in a
survey by Nagy (1980) who showed that AsA contents of the
major citrus cultivars from several countries ranged from 14 to
88 mg/100 mL of juice for sweet oranges, 15–45 mg/100 mL for
limes and 15–55 mg/100 mL for mandarins. However, Sharma et al.
(2006) showed that lemons and lime have the highest concentra-
tion of AsA.
Oranges are regarded as a significant source of vitamin C
(Klimczak et al., 2007). A study by Gliszczyńska-Świgło et al. (2004)
analyzed 14 commercial orange juices from different producers and
16 L.S. Magwaza et al. / Scientia Horticulturae 216 (2017) 12–21
reported that vitamin C content ranged from 150 to 450 mg L−1 of
juice. Cano et al. (2008) observed that sweet orange cultivars had
the highest levels of total vitamin C compared to clementines man-
darins and mandarin hybrids. ‘Navel’ oranges had the highest levels
while ‘Lane late’ and ‘Fukumoto’ had the lowest concentration. In
mandarin, the ‘Hernandina’ and ‘Oronules’ (Clementines group)
had the highest concentrations of AsA (46.0 and 47.4 mg/100 g,
respectively) whereas in Satsume group ‘Owari’, ‘Okitsu’, ‘Avasa
Pri-10 and ‘Avasa Pri-19 had the lower values of 20.0, 21.3, 24.9
and 25.5 mg/100 g, respectively. The results by Cano et al. (2008)
were in accordance with Lee and Kader (2000) and Dhuique-Mayer
et al. (2005) who reported the higher concentration of AsA in orange
cultivars than mandarin cultivars. In a later study Sdiri et al. (2012)
showed that total vitamin C content of different mandarin cultivars
ranged from 21.19 to 29.80 mg/100 mL of juice (Table 3), with ‘For-
tune’ and ‘Safor’ mandarins having the highest content, followed by
‘Kara’ mandarin and the lowest concentration observed in ‘Garbí’
fruit.
Cano et al. (2008) further subjected vitamin C concentration of
different citrus cultivars to principal component (PC) analysis (PCA)
and showed a high load of total vitamin C on PC2 (R2 = 0.882), and
hesperidin and narirutin on PC1 (R2 = 0.965) appeared, a classifica-
tion based on vitamin C and flavanoinds (hesperidin and narirutin)
can be established, where the cultivars of mandarins and oranges
were completely separated. This indicated that the concentration
of vitamin C and flavonoids in citrus fruit are important variables
to group the different cultivars. This is an important nutritional
aspect demonstrating higher concentration of vitamin C should be
one of the important factors to choose cultivars with a high poten-
tial as nutraceutical source. However, it should be kept in mind
that the wide variation in vitamin C levels of different citrus fruit
reported by most investigators is also because of many climatic and
horticultural factors.
2.2. Rootstocks
Biochemical characteristics, including vitamin C content of vari-
ous citrus cultivars, are significantly influenced by genetic influence
arising among different genotypes. Furthermore, the variation may
also be brought about by the effect of rootstocks, which are known
to invariably influence the fruit size and nutritional quality vari-
ables (Singh, 2001; Sharma et al., 2006). There is some evidence of
the effects of rootstock on the production of vitamin C in citrus fruit.
For instance, the effects of rootstock on fruit quality was shown
on ‘Temple’ oranges, where fruit grown on sweet orange rootstock
had the lowest content of AsA while those growing on rough lemon
rootstock had the highest content (Nagy, 1980). Hifny et al. (2013)
also showed that AsA concentrations of ‘Valencia’ oranges fruit har-
vested from trees budded on ‘Sour’ orange rootstock were superior
to those budded on ‘Volkamer’ lemon (C. volkameriana) rootstock.
Similar results were reported by El-Sayed et al. (2007), who found
that Washington navel orange budded on ‘Volkamer’ lemon root-
stock produced fruits with highest concentration of ascorbic acid.
The positive effect of ‘Volkamer’ lemon rootstock on vitamin C has
also been reported on ‘Marsh’ and ‘Ruby red’ grapefruit (Ramin and
Alirezanezhad, 2005).
A recent study by Cardeñosa et al. (2015) evaluated ‘Lane Late’
orange grafted on six different citrus rootstocks (Carrizo, Cleopatra,
Foner-Alcaide n◦13, Forner-Alcaide n◦41, Forner-Alcaiden◦5, and
Macrophylla) to characterize their antioxidant activity and quantify
their vitamin C content. These authors reported higher levels of AsA
in ‘Cleopatra’ rootstock while ‘Forner-Alcaide 41’ rootstock showed
significantly lower levels. These results proved that the differences
of AsA compound are strongly influenced by the type of rootstock.
‘Cleopatra’ rootstock had higher antioxidant activity and vitamin
C in both seasons of the study, indicating that if higher vitamin C
concentration is the goal, an increase in its use might be a good
solution in sweet orange production.
Similar results were reported earlier by Bassal (2009) who eval-
uated vitamin C content of ‘Marisol’ clementine grafted on four
different rootstocks, namely, ‘Carrizo’ citrange, ‘Cleopatra’ man-
darin, ‘Sour’ orange and ‘Swingle’ citrumelo under the Egyptian
conditions. These authors showed that the content of AsA was
considerably affected by rootstock with fruits from trees grafted
on ‘Sour’ orange having lower AsA compared to those grafted on
‘Swingle’ citrumelo in the first season and on ‘Cleopatra’ man-
darin and ‘Carrizo’ citrange in the second one. The results by Bassal
(2009) and Cardeñosa et al. (2015) collectively demonstrated that
if higher vitamin C content is the goal, ‘Cleopatra’ is the best root-
stock to choose. However, Hifny et al. (2012) reported contrasting
results where ‘Navel’ orange fruits from trees grafted on ‘Sour’
orange rootstock had a significantly higher vitamin C than those
on ‘Volkamer’ rootstock. This contrast is an indication that there is
no perfect rootstock even for a specific production system. Accord-
ing to Ramin and Alirezanezhad (2005), the nutritional quality of
the citrus fruit juice is directly linked to the effects of rootstock
to the plant-water relation regulating sucrose transport. Therefore,
the interaction between rootstocks and cultivars should be consid-
ered carefully as these may be a thought-provoking approach for
the adaptation of citrus cultivars to diverse climatic conditions. Fur-
thermore, other tree characteristics and fruit quality factors should
be considered, mainly due to the unacceptability of deciding about
rootstocks using a single factor. A decision to use a particular root-
stock should mostly depend on the most important limiting factors.
For instance, if the limiting factor is vitamin C content, based on
the literature reviewed above, it may be suggested to substitute
‘Sour’ orange rootstocks of with ‘Volkamer’ lemon and ‘Cleopatra’
mandarin.
2.3. Rainfall and irrigation
The accumulation of vitamin C has been shown to be negatively
correlated with irrigation and rainfall (Toivonen et al., 1994). In
a study by Navarro et al. (2015), the concentration of AsA in the
juice of ‘Star Ruby’ grapefruits was not affected by water stress
treatments. A similar trend was also observed by Buendía et al.
(2008).
However, some studies have reported that reducing water avail-
ability to 50% of the crop requirement significantly increased the
AsA content in mandarin (Navarro et al., 2010; Panigrahi et al.,
2014) as well as tomato fruits (Favati et al., 2009). Navarro et al.
(2010) showed that if deficit irrigation treatment was imposed dur-
ing phase II of fruit growth, the concentration of AsA was 15% higher
than the control. From these observations, the increase in concen-
tration could either result from biosynthesis of AsA, in response to
water stress, or the lower fruit water content, resulting from the
lower plant water potential (Navarro et al., 2010). Water deficit
during the stage II and III of fruit growth increase soluble sugar
accumulation in ‘Satsuma’ mandarin fruit (Yakushiji et al., 1996).
This may be supported by the fact that sugar is the primary precur-
sor for AsA biosynthesis (Wheeler et al., 1998) and therefore can
play a significant role in the signaling and regulating gene expres-
sion of enzymes involved in the metabolism of AsA (Nishikawa
et al., 2005). However, recent studies have shown that AsA con-
tent is not strongly linked to the metabolism of sugars and their
content in the tissue (Massot et al., 2010; Yang et al., 2011).
2.4. Light and temperature
Light interception and temperature are some of the principal
environmental factors determining the level of AsA (Massot et al.,
 L.S. Magwaza et al. / Scientia Horticulturae 216 (2017) 12–21
2013). Previous results revealed that temperature and sunlight
have a significant effect on AsA content of various horticultural
products (Klein and Perry, 1982). This was supported by Lee and
Kader (2000) who also reported that light intensity and availabil-
ity during development have a strong effect on the amount of
AsA synthesized. In general, Lee and Kader (2000) suggested that
sun-exposed fruit and sides of fruit receiving a higher amount of
sunlight during growth have higher levels AsA than shaded fruit or
sides.
Shading citrus fruit with the net has been reported in earlier
studies to decrease canopy temperature, air temperature, fruit tem-
perature, greatly affecting growth and quality of many citrus fruits
(Jifon and Syvertsen, 2001; Lee et al., 2015; Lado et al., 2015). The
AsA pool was also increased in leaves grown at high light intensity
and at low temperature (Smirnoff, 1996). A study by Magwaza et al.
(2013) assessed the effect of canopy position and shading on AsA
on the peel of ‘Nules Clementine’ mandarin (C. reticulata Blanco)
fruit. The results showed that AsA content was higher in the peel of
sun-exposed outside fruit compared to the shaded fruit inside the
canopy. These findings suggest the possibility that ascorbic acid
concentration in the peel could be used as a potential biochemi-
cal marker of fruit position within the canopy. These results were
similar to those reported by Izumi et al. (1990) who showed that
exterior canopy fruit had higher AsA than interior canopy fruit.
Similar results were recently reported by Lado et al. (2015)
who observed that light avoidance by fruit bagging resulted in
a remarkable decrease in the content of AsA in the flavedo of
‘Navelina’ orange and ‘Clemenules’ mandarin compared to light-
exposed fruits. These results demonstrated that intensification of
radiation interception is important to have citrus fruit with ele-
vated levels of AsA and antioxidant capacity. However, the amount
of AsA in the pulp of fruit from both citrus species by the same
authors (Lado et al., 2015) did not show differentials in response to
fruit shading, indicating that the influence on the accumulation of
AsA may be restricted to the flavedo, which is exposed to light.
Direct exposure of fruit on the outside position of the canopy
to sunlight may have led to an up-regulation of the AsA biosyn-
thetic and recycling pathways (Ma and Cheng, 2004). Considering
that carbohydrates produced during photosynthesis are precursors
of AsA synthesis, it can also be hypothesized that lower AsA in the
peels of sun-shaded fruit may have resulted from reduced photo-
synthesis due lower photosynthetically active radiation on shaded
portions of the tree canopy. As a potent antioxidant in citrus fruit
(Mathur et al., 2011; Sdiri et al., 2012), a higher concentration of AsA
observed in the peel of fruit harvested from the sun-exposed posi-
tion of the canopy may also suggest a defense mechanism against
preharvest stresses.
According to the discussion in the Introduction section (Section
1) of this review, AsA is synthesized from carbohydrates precur-
sors (Fig. 1). Therefore, a lower concentration of sugars in shaded
fruit inside the canopy could contribute to lower levels of AsA com-
pared to sun-exposed outside fruit (Valpuesta and Botella, 2004;
Zhan et al., 2013). A positive correlation between AsA and sucrose
and glucose, by Magwaza et al. (2013) confirmed the role of these
sugars, especially glucose as a primary substrate for AsA synthesis
pathway (Valpuesta and Botella, 2004).
2.5. Fertilization
2.5.1. Macronutrients
The effect of tree nutrition on internal fruit quality and stor-
age ability of citrus fruits are significant and well known. High
nitrogen (N) fertilization of citrus fruit trees has been reported to
be associated with significant reduction in levels of AsA (Lee and
Kader, 2000). Increasing N and phosphorus (P) supply to citrus trees
have been shown to results in lower AsA content in fruit while
17
increasing potassium (K) fertilization increased AsA content (Nagy,
1980). In acid limes (C. aurantifolia Swingle), Prasad and Putcha
(1979) reported that applying 800 g nitrogen/plant/year increased
the retention of AsA content during postharvest storage at room
temperature for 12 days. ‘Ray Ruby’ grapefruits receiving higher
amount of potassium had reduced content of AsA while increased
the concentration of DHA. Findings that higher N reduces AsA con-
tent were also confirmed by Hifny et al. (2013) who observed that
adding 50% of total recommended N/tree/year at either fruit cell
division stage or fruit cell expansion stages showed an increase
in vitamin C content compared to full recommended dosage of
400 g N/tree/year.
Reduced vitamin C content in plants receiving high N rates may
be due to increased vegetative growth and large fruit, generally
improved by the N fertilization leading to the occurrence of a dilu-
tion effect in the plant tissues (Lee and Kader, 2000). Furthermore,
reduction in AsA as a result of N fertilization may results from
reduced light intensity and accumulation of AA in shaded parts
due to increased foliage (Lee and Kader, 2000). However, Chen
et al. (1999) reported an increase in Ascorbic acid content in fruit
receiving higher N fertilizer. Similarly, increasing N fertigation to
‘Balady’ mandarin trees from 400 to 1000 g N/tree/year was shown
to be associated with improved vitamin C content and the recom-
mended dose in this respect was 800 g N/tree/year (Wassel et al.,
2007). Splitting N application had a positive effect on vitamin C con-
tent where splitting N to 23 and 27 doses resulted in higher ascorbic
acid than 21 doses. These results are in line with findings of El-Saida
(2006), who found that application of N at (1200 g/tree) to ‘Balady’
mandarin budded on SO increased vitamin C content. Chen et al.
(1999) hypothesized that an increase in AsA could be attributed to
the increase in fruit size resulting from the increase in shoot and
leaf area caused by N-application. The best results regarding phe-
nological stages were gained when 50% of total given N/tree/year
was applied at either fruit cell division stage or fruit cell expansion
stage as they caused the highest values of other fruit biochem-
ical characteristics in comparison to control or other nutritional
treatments.
Phosphorus is well known to influence fruit internal quality
(Nagy, 1980). Bar-Akiva et al. (1967) reported a positive correlation
between phosphorus and vitamin C content of grapefruit. Contrast-
ing results were reported by Dou et al. (2005) who showed that
in grapefruit, vitamin C was the highest when trees received with
optimum K (186 kg/ha) without P. Optimal P (48 kg/ha) application
without K significantly reduced juice vitamin C concentration. Sim-
ilar results were reported by Mann and Sandhu (1988) who found
that in ‘Kinnow’ mandarins, vitamin C was increased by K, but not
by P application.
Potassium is regarded as an essential element required for a
consistent production of high quality citrus fruits. Nagy (1980)
reported that K effects on citrus fruit quality are more important
than its effect on yield. Several studies have shown that high lev-
els of K fertilization increase vitamin C in lemons (Embleton and
Jones, 1966), oranges (Reitz and Koo, 1960), and grapefruit (Smith
and Rasmussen, 1960; Dou et al., 2005). This was supported by
Patil and Alva (2002) on a study to determine the effect of differ-
ent levels of K on vitamin C on ‘Ray Ruby’ grapefruit. The study
by the later authors showed that application of higher amounts of
K reduced the content AsA while increased the concentration of
DHA compared to fruit from untreated control. These authors also
showed that total concentration of vitamin C was not affected by
the application of varying levels of K. Although the total concentra-
tion of AsA was not influenced by varying levels of K, application of
K at 93 kg/ha/year had higher DHA and reduced AsA concentration
compared to those fruit from untreated trees. These findings were
in accordance with those previously reported where it was shown
that optimum supplies of K increase the concentration of vitamin
18 L.S. Magwaza et al. / Scientia Horticulturae 216 (2017) 12–21
C in various citrus fruit types (Smith, 1966; Jones et al., 1973; Nagy,
1980; Hearn, 1993). Since the metabolism of sugars is significantly
affected by the availability of K in the tissue, fertilizing citrus trees
with optimum K is likely to have a beneficial effect on the concen-
tration of AsA (Embleton and Jones, 1966; Smith and Rasmussen,
1960).
2.5.2. Micronutrients
The concentration of AsA is not uniquely affected by macronu-
trients. Several studies have reported the effect of micronutrients
on the improvement of ascorbic acid of citrus fruit. Mishra et al.
(2003) evaluated the effect of zinc (Zn) and boron (B) on yield and
quality ‘Kinnow’ mandarins. Fruit harvested from trees receiving
Zn (0.5%) + B (0.4%) micronutrient treatments had higher AsA. Sim-
ilar results were observed by Babu et al. (1984) who reported higher
AsA concentration in ‘Kagzi’ limes (C. aurantifolia Swingle) when Zn
(0.6 percent) and 2,4-D (20 ppm) were applied during spring and
summer flushes.
However, a foliar application of B as the boric acid was reported
to significantly reduce the content of AsA in ‘Kinnow’ mandarin fruit
(Ullah et al., 2012). B plays an important role in the biosynthesis of
auxin in the plant meristem and its deficiency leads to increase
in the level of bound auxin and reduction in the indole acetic acid
(IAA)-oxidase activity. As a result of increased levels of bound auxin,
the ascorbic acid content of the fruit was increased. Therefore the
increased ascorbic acid content of the control trees might be due to
increased level of bound auxin which is produced in the absence of
B.
Several studies have indicated that application of Zn enhances
AsA content in fruit juice (Nakhlla, 1998; Abd El-moneim et al.,
2007). A study carried out in Egypt on ten years old ‘Washing-
ton’ navel orange trees (C. sinensis L. Osbeck) budded to ‘Volkamer’
lemon (C. volkamerina L.) rootstock revealed that chelated Zn spray
at 0.4% increased vitamin C content (Abd El-moneim et al., 2007).
From the results by Ashraf et al. (2013), it was concluded that foliar
application of Zn + K is effective in the enhancement of AsA concen-
tration. These authors showed that application of sulfate of potash
at a rate of 75 kg K2 O ha−1 was more beneficial than muriate of
potash at the same rate because it enhanced appreciably the ascor-
bic acid content. Similarly, ‘Kinnow’ mandarin trees sprayed with
0.6% zinc sulfate exhibited the highest increase in ascorbic acid con-
tents compared to the untreated control (Razzaq et al., 2013). The
Zn has an important role in the biosynthesis of auxins and increased
production of auxins has been previously shown to increase the
content of AsA in ‘Kinnow’ mandarin (Nawaz et al., 2008). The
results by Razzaq et al. (2013) were similar to those previously
reported, where the exogenous application of Zn sprays increased
the AsA contents in ‘Balady’ mandarin (El-Menshawi et al., 1997).
Similarly, in ‘Navel’ oranges, Nakhlla (1998) reported that Zn sprays
improved vitamin C content in fruit.
2.5.3. Organic vs inorganic fertilizers
Organic agriculture has become one of the important sectors and
there is a substantial increase in consumer interest in organically
produced food products. Organic food products often fetch 15% to
20% price premium over conventionally grown crops (O’Connell
et al., 2012; Kleemann, 2013). Due to price differences and the
importance of vitamin C in citrus fruit, research efforts have been
made to compare AsA content of organically and conventionally
grown citrus products.
The effect of the production system on vitamin C has been stud-
ied in several citrus species (Tarozzi et al., 2006; Lester et al., 2007;
Duarte et al., 2012; Çandır et al., 2013). Some authors have found
evidence suggesting that organically produced citrus fruit accrue
higher content of vitamin C than conventionally produced fruit
(Rapisarda et al., 2005). A higher AsA content in fruit from organi-
cally produced compared to conventional production systems was
reported in ‘Tarocco’ oranges (Tarozzi et al., 2006), ‘Newhall’ and
‘Navelina oranges (Raigón, 2007), “Valencia late” and ‘Baia’ oranges
(Duarte et al., 2010), ‘Rio Red’ grapefruit (Lester et al., 2007),
‘Nules’ and ‘Okitsu’ mandarin (Raigón, 2007). However, other stud-
ies also show results that are either opposite or show no differences
between the two farming systems (Elhassan et al., 2011; Roussos,
2011). For instance, Tarozzi et al. (2006) reported that in ‘Baía’
and ‘Valencia late’ oranges, AsA content was significantly higher
in organically grown fruit, but in other orange cultivars (‘Newhall’,
‘Lane late’, ‘Dalmau’, and ‘Rohde’), there were no significant differ-
ences observed between fruits from organic and inorganic farming
systems. Similarly, some mandarins (‘Fina’, ‘Fortune’, ‘Fremont’,
‘Hernandina’, Ortanique’ and ‘Tangera’), the juice had higher AsA
concentration in fruit harvested from organic farming (Duarte et al.,
2012). The differences in results obtained by different authors in
studies discussed above do not contradict each other but show that
higher content of AsA is not always linked with organic farming,
but depends significantly on the citrus type and cultivar. This sup-
ports a statement by Winter and Davis (2006) who concluded that
it is impossible to have a guarantee from a nutritional perspective,
that organically grown fruit and vegetables are superior to those
produced using the conventional agricultural system.
2.6. Hormones and plant growth regulators
Application and influence of plant growth regulators (PGRs)
such as gibberellic acid (GA3 ) and cytokinins for production of qual-
ity fruit is well documented for mandarins (Garcia-Luis et al., 1985;
Pozo et al., 2000; Khalid et al., 2012), grapefruit (El-Zeftawi, 1980),
and oranges (Fidelibus et al., 2002). The beneficial effects of hor-
mones such as GA3 on fruit quality was reported by investigators
such as Morton (1981) who found that GA3 either applied at full
flowering or at small fruit stage, significantly increased the number
of harvested ‘Washington’ navel fruits.
Similarly, Nodiya and Mikaberidze (1991) reported that Sat-
suma mandarin fruits treated with 60 ppm of gibberellic acid before
the fruit showed signs of yellowing had a higher accumulation of
vitamin C content. This is consistent with the statement by Eassa
et al. (2012) who reported that improved fruit vitamin C con-
tent may be due to its role in enhancement the nutritional status
reflected on yield and fruit quality. Kuiper (1993) suggested that
this is mostly because the sink strength is established and regulated
by plant growth regulators. Plant hormones such as cytokinins and
gibberellic acid have been found to increase movement of photo-
synthates to fruit and regulate several rate-limiting constituents
involved in carbon partitioning (Ozga and Dennis, 2003). Brenner
and Cheikh (1995) argued that PGRs may either stimulate mobiliza-
tion of nutrients through the phloem, adjust the sink strength of the
tissue by enhancing its growth and increase the capacity for unload-
ing of sugars from the phloem or they may act on metabolism and
compartmentalization of sugar and its secondary metabolites.
Some scientists reported the inconsistent effect of PGRs on
ascorbic acid contents of citrus fruit (Lima and Davies, 1984). Using
young (3–5 years) trees, Khalid et al. (2012) investigated the effect
of PGRs, including benzyladenine (BA), kinetin and GA3 applied
at different growth stages on the quality of ‘Kinnow’ mandarins.
These authors reported that the PGRs had a significant nega-
tive influence on juice ascorbic acid with maximum AA contents
(58.45 mg 100 mL−1 ) of juice observed on the control treatment
while minimum AA contents (34.88 mg 100 mL−1 ) were observed
with 20 mg L−1 kinetin at FS stage. These results suggested (val-
idated) that AA contents of fruit from young ‘Kinnow’ mandarin
trees could not be improved by the exogenous application of
kinetin. It was deduced that the decrease in AsA contents in fruit
from trees treated with PGRs might be due to their positive influ-
 L.S. Magwaza et al. / Scientia Horticulturae 216 (2017) 12–21
ence on sink strength (reproductive growth) which was supported
by more juice mass (%) and TSS in fruit from PGRs treated trees
compared to the untreated control. In contrast to findings by Khalid
et al. (2012), Saleem et al. (2008) found higher AsA content in blood
oranges from GA3 treated trees compared to the control. These dif-
ferences in response to GA3 treatment between these studies might
be due to the difference in cultivar and age of the plant.
Furthermore, there has been reports on the potential involve-
ment of ethylene in AsA biosynthesis for various fruit. It was
also reported that AsA is also co-factor of 1-aminocyclopropane-
1carboxylic acid oxidase (ACC oxidase) in ethylene biosynthesis
(Ververidis et al., 1992; Liu et al., 1999). However, in some climac-
teric fruit, for example, Kiwi fruit, the ethylene production declines
during ripening towards maturity (Li et al., 2010). These findings
prove that the correlation between AsA and ethylene is complex, it
depends on plant species.
3. Harvesting factors
3.1. Time of harvest (Maturity)
Vitamin C contents in different fruits are known to be modulated
during fruit development. In tissues of citrus fruit, AsA content and
concentration is uneven, varying with the stage of fruit develop-
ment (Yang et al., 2011). In fruit such as oranges, tangerines and
grapefruit, Ladaniya et al. reported vitamin C content to decrease
with maturity, indicating that early harvest or mid harvest fruit
have more AsA than late season fruit. Yang et al. (2011) analyzed
AsA contents in fruits of ‘Satsuma’ mandarin and two commercial
‘Navel’ orange cultivars during phases II and III of fruit develop-
ment. These authors showed that AsA content in pulps of fruit had
different changing patterns between mandarin and orange, indicat-
ing that the ultimate concentration in the pulp tissue of citrus fruit
is highly dependent on both cultivar and the stage of development.
This is also supports the statement by Nagy (1980) who reported
that concentration of AsA usually decreases as fruit matures and
ripens. Furthermore, the study by Yang et al. (2011) suggested
that the pattern of AsA accumulation is also under the influence
of different cultural practices and environmental stresses during
fruit growth and maturation, partially explaining the inconsistent
profile of AsA in different citrus types and cultivars.
4. Conclusion
The nutritional quality of citrus fruit and juices is related pri-
marily to the content of AsA and dehydroascorbic acid, the sum
of which is known as vitamin C. The present review of litera-
ture demonstrated that the variance among species, cultivars, and
fruit tissues is regulated by genes encoding for different enzymes
involved in the two major AsA biosynthesis pathways, L-galactose,
and D-galacturonic acid pathways. Although, there seem to be a
consensus among various research groups on biosynthetic path-
ways and molecular mechanism for AsA in citrus fruit, however,
a general understanding of the variations in the expression of
the multiple genes and isoforms involved in the main metabolic
pathways controlling AsA concentration including biosynthesis,
degradation, and recycling pathways, during the development and
ripening of citrus fruit, is still limited and should constitute the
objective of future studies. Evaluation of the current knowledge
of genome and transcriptome provided collective insights into
the origin of sweet orange and the genomic basis of vitamin C
metabolism and provided a rich resource of genetic information for
citrus breeding and genetic improvement. Future studies compar-
ing the developed sweet orange genome with other citrus cultivars
including existing and new genotypes should provide new insights
19
into citrus biological and evolutionary questions. AsA was observed
among reviewed literature that it is highly sensitive to various
preharvest factors, including scion-rootstock relationship, climatic
conditions, cultural practices, hormones, light, and temperature. In
general, the largest differences in vitamin C composition observed
are mostly attributed to genetic factors, mostly species, cultivars
and rootstock-scion interaction. This confirms that the selection of
genotype with a potential to increase amounts of vitamin C is a
more important factor than climatic conditions and cultural prac-
tices. To break existing bottlenecks about the interaction between
AsA and other compounds contributing to the antioxidant capac-
ity of citrus fruit, their synergism, and/or antagonism in citrus fruits
should be more explored. Although a significant amount of research
have been reported on the effect of PGRs on AsA content, there is no
clear biological mechanism on how they affect the AsA biosynthesis
in fruit.
Acknowledgements
This work is based upon research supported by the Postharvest
Innovation Fund of the Department of Science and Technol-
ogy (Project Number: 2/2014), Citrus Research International and
National Research Foundation (Thuthuka Project Number: 94022).
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