Acta Physiol Plant (2015) 37:1718
DOI 10.1007/s11738-014-1718-2
REVIEW
Physiological and genetic factors influencing fruit cracking
Abdollah Khadivi-Khub
Received: 29 August 2014 / Revised: 3 November 2014 / Accepted: 4 November 2014
 Franciszek Górski Institute of Plant Physiology, Polish Academy of Sciences, Kraków 2014
Abstract One of the main disorders that widely limit
fruit quality and quantity is fruit cracking or splitting that is
observed on the fruit skin and flesh in the preharvest phase.
Besides, cracking can occur during postharvest in some
fruits, mostly attributable to the environmental conditions
of storage. Value of cracked fruits is reduced and these
fruits are not marketable because of the poor fruit quality.
Many fruits such as apple, sweet cherry, grape, plum,
pomegranate, grape, persimmon, litchi, avocado, pistachio,
citrus, banana as well as tomato can crack or split. There
are many factors that influence fruit cracking. In this work,
genetic, morphological, environmental and physiological
aspects of fruit cracking are reviewed. Under the same
environmental conditions, fruits from different cultivars
show differences in cracking susceptibility. Some correla-
tions have been observed between susceptibility of fruit
cracking and some fruit traits (fruit shape, fruit size, fruit
firmness; anatomy and strength of the fruit skin, stomata in
fruit skin, cuticular properties, osmotic concentration,
water capacity of the fruit pulp and growth stage of the
fruit). Also, orchard management (such as irrigation and
nutrition) and environmental condition (such as tempera-
ture, wind and light) can influence fruit cracking. Besides,
fruit cracking is quantitative trait and is controlled by
several genes. The best way to reduce fruit cracking at
present would be a suitable orchard management that takes
into account and try to minimize stress of the water,
nutrition and physiological factors that contribute to fruit
Communicated by A. K. Kononowicz.
A. Khadivi-Khub (&)
Department of Horticultural Sciences, Faculty of Agriculture and
Natural Resources, Arak University, 38156-8-8349 Arak, Iran
e-mail: akhadivi@ut.ac.ir; a-khadivi@araku.ac.ir
cracking. Also, the most resistant cultivars to fruit cracking
that have desirable fruit quality can be selected for
cultivation.
Keywords Fruit cracking
Cultivar
Genetic

Environment
Irrigation
Nutrition
Introduction
Fruit cracking is a considerable disorder that causes neg-
ative effect in fruit marketability mainly because splits
reduce the fruits quality as they cause poor appearance,
decrease shelf life and even render the fruit unmarketable
due to fungal infection (Peet 1992). Fruit cracking may
occur during fruit growth, development and ripening. Also,
cracking can occur during postharvest in some fruits,
mostly attributable to the environmental conditions of
storage. Fruit cracking is a physical failure of the fruit peel
that manifests as fractures in the peel or cuticle of certain
fruits, or splitting, a more extreme form of splitting that
penetrates deep into the pulp (Opara 1996). Cracks reduce
marketability and provide entry for insects and fungi,
causing significant income loss in the fresh market and
processing industries. Cracked fruits are susceptible to
disease of storage and their storage and shelf life are
shorter (Khadivi-Khub 2009). Fruit cracking disorder is
commonly found in sweet cherry, plum, apricot, apple,
litchi, pomegranate, citrus, banana, avocado, grape, per-
simmon, peach, tomato, pistachio etc. (Opara 1996; Simon
2006; Khadivi-Khub 2009).
Genetic differences in cracking susceptibility between
cultivars of a species have been correlated with fruit size
and shape, fruit growth and development, fruit cuticle and
sugar level (Emmons and Scott 1998). Also, environmental
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Fig. 1 Fruit cracking in different species. a Pomegranate (http://
www.lafujimama.com/2010/11/from-blossom-to-bottle-a-pomegranate-
story/). b Grape (http://www.csu.edu.au/__data/assets/pdf_file/0003/
393456/NWGIC-fs1-fruit-splitting.pdf). c Sweet orange (www.page
glance.com/harald-hoffmann.com). d Sweet cherry (http://extension.
conditions especially temperature and humidity influence
fruit cracking. There are many types of fruit cracking based
on species: longitudinal or burst cracking; ring or con-
centric cracking; crazing or russeting; star or radial
cracking; lenticilar cracking; and core failure. Radical
cracking is developed from the radical section of stem
towards the fruit center. Concentric cracking is appeared on
fruit that leads to cracking in a single fruit which is known
as concentric e.g., sweet cherry (Peet 1992). Although fruit
cracking has been investigated since the 1930s, and
breeders have incorporated crack resistance into many
fruits, minor advances have been made in understanding
the physiology of fruit cracking in any of the crops. In turn,
this has made it difficult to recommend preventive
measures.
Part of the problem is that fruit cracking is episodic in
nature, ranging from severe in some years in certain
locations to virtually unknown in others. Fruit cracking is
difficult to study, even in controlled conditions, because we
lack experimental methods to induce cracking. Growers
and researchers have associated a number of physiological,
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Acta Physiol Plant (2015) 37:1718
oregonstate.edu/umatilla/mf/sites/default/files/WA_State_Cherry_
Cracking_Kaiser_Dec_07.pdf). e Persimmon (http://ucce.ucdavis.
edu/files/…/391-472.pdf). f Apple (http://www.plantmanagementnet
work.org/pub/php/research/2010/apple/)
biochemical, environmental, cultural, anatomical and
genetic factors with high incidence of fruit cracking. This
review concentrates including advances in understanding
the causes of cracking in some fruits including sweet
cherry, apple, wax apple, grape, citrus, pomegranate, per-
simmon, litchi, pistachio, avocado, banana as well as
tomato. Pictures of fruit cracking in several species are
shown in Fig. 1.
The nature of cracking in some fruits
Sweet cherry
Three types of fruit cracking in sweet cherry are known:
1—semicircular or circular cracks around stem end on the
cavity; 2—similar small splits at the fruit apical end; and
3—many often deep splits on the fruit sides and it is known
as lateral splitting (Simon 2006; Khadivi-Khub 2009). The
small and fine splits at the base and apical end of cherry
fruit may occur at very early stages, many days before fruit
Acta Physiol Plant (2015) 37:1718
maturity and harvest. These fine cracks at an early stage of
fruit development decrease the market value of fruits. In
contrast, the large, deep, side cracks provide a point of
entry for the fungi (brown rot and gray mold) and are
frequently followed by severe attacks. Fungal attacks will
affect fruit quality and render them unmarketable. Factors
influencing fruit cracking in sweet cherry are:
Fruit size
It is commonly assumed that large fruits in sweet cherry are
more susceptible to cracking than smaller ones. Several
studies (Sekse 2008; Khadivi-Khub et al. 2007) have
shown that cultivars having large fruits show higher trend
to crack than small-fruited cultivars. Way (1967) showed
that fruits of heavy crop-loaded trees crack less than light
cropping trees for the same sweet cherry cultivar.
Fruit firmness
It is a generally accepted opinion that firm-fleshed cherry
cultivars have higher trend to fruit cracking than soft-fleshed
cultivars. Fruit cracking in sweet cherry is caused by excess
water uptake resulting in bursting of the skin; it seems
dialectical that firm-fleshed fruit cherries are more prone to
fruit cracking than soft-fleshed cultivars (Christensen 1996;
Yamaguchi et al. 2004; Khadivi-Khub 2007). Iranian cul-
tivars such as ‘Arak’, ‘Mashhad’, ‘Meshkinshahr’, ‘Mehali-
Karaj’ and ‘Hamedan’ have soft flesh fruit and do not show
any cracking, but ‘Soorati-Lavasn’, ‘Rafat’, ‘Ghazvin’,
‘Maremot’ and ‘Siah-Daneshkade’ with firm flesh fruit
show high cracking (Khadivi-Khub et al. 2007).
Fruit shape
Fruit shape is very important factor of fruit cracking in
sweet cherry (Beyer et al. 2002; Sekse et al. 2005; Sekse
2008; Khadivi-Khub 2007). They thought that fruits can
crack more easily when they have right angle to the cur-
vature of the shape. The fruit shape can play a significant
role in cherry fruit cracking. Cultivars with kidney or heart
fruit shape such as ‘Rafat’ and ‘Ghazvin’, Iranian culti-
vars, have deeper stem cavity and therefore the rain drops
may remain there for a long time giving possibility for
higher water uptake through the fruit skin (Khadivi-Khub
et al. 2007). It means that if a cultivar is genetically
determined (based on structure and thickness of the cuticle
and sugar content) for moderate susceptibility to splitting
and it has kidney, blocky or heart shape, maybe it will
have stem end type splitting (Simon 2006; Khadivi-Khub
2007).
Page 3 of 14 1718
Fruit development
The generally accepted opinion is that susceptibility to
cracking is increased with advanced maturity in most sweet
cherry cultivars, but there are cases of deviation from this
relationship. The reason of this deviation is due to climatic
conditions influencing the fruit’s susceptibility to cracking
before the beginning of fruit sampling or picking (Chris-
tensen 1996; Yamaguchi et al. 2004).
Stomata in the fruit skin
Water absorption in leaves happens mostly through epi-
dermal cells and the absorption through stomata is signif-
icant (Moing et al. 2004). Sawada (1931) found that at least
a part of the water absorbed into cherry fruit comes through
the stomata. The density of stomata does not appear to have
any significant influence on the amount of water taken up
before cracking occurred (Christensen 1972).
Cuticular properties
Glenn and Poovaiah (1989) noted that cuticle was sepa-
rated from the cell wall because of water absorption
through the fruit skin and the associated swelling in the
epidermal cell wall section resulted in cuticular fractur-
ing, which commonly preceded fruit splitting. Belmans
et al. (1990) observed a high correlation between thick-
ness of cuticle and the resistance of 13 cultivars. Knoche
and Peschel (2006) demonstrated that exposure of the
fruit surface to liquid water or high concentration of
water resulted in formation of micro-cracks in cuticular
membrane.
Osmotic concentration
Tucker (1934) showed a correlation between the sugar
content and cracking tendency of cultivars. But, Verner
(1957) reported that fruit cracking was directly affected by
the osmotic concentration of the fruit juice.
Water-retaining capacity of the fruit pulp
Kertesz and Nebel (1935) and Beyer et al. (2002) found
that the water-retention capacity and swelling of this col-
loidal pulp are important in determining cracking suscep-
tibility. It is hypothesis that dwarfing rootstocks, because of
smaller roots, are exposed to stress and cause cracking
(Knoche and Peschel 2002; Hovland and Sekse 2004;
Simon et al. 2004; Khadivi-Khub 2009).
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Genetic
It has been reported that some SSR and RAPD markers are
significantly associated with fruit cracking so that BPPCT-
004197 SSR locus and TIBMBA-09521 RAPD marker have
high association with fruit cracking (Khadivi-Khub 2007,
2011). Also, investigations for quantitative trait locus
(QTL) detection were conducted for cracking resistance in
sweet cherry (Quero-Garcia et al. 2012). They reported that
QTL of cracking resistance at each fruit zone were found
on distinct linkage groups. This result confirms that dif-
ferent genetic mechanisms are responsible for cracking in
each region of fruit. In total, QTL observed for pistillar end
and stem end cracking resistance are significant than QTL
detected for fruit side cracking resistance (Quero-Garcia
et al. 2012).
Environment
Temperature has an important function in fruit cracking
rate. In total, there is a linear increasing in fruit cracking
with temperature increasing from 10 to 40 C. Also, tem-
perature influences many other agents such as the cell
walls’ permeability and cells’ biochemical processes. It is
very important to know the influence of temperature in fruit
cracking if we want to find some measure against fruit
cracking, because usually the temperature is higher
underneath the screen than outside (Simon 2006). Fur-
thermore, the distribution and quality of rainfall during the
maturation and ripening stages can affect the susceptibility
of sweet cherry to fruit cracking (Christensen 1996; Simon
2006).
Apple
Some commercial apple cultivars show high fruit cracking.
‘Golden Russet’, ‘Fuji’ and ‘Gala’ are three commercial
cultivars, but have susceptibility to fruit cracking (Sada-
mori et al. 1963; Opara 1996; Andrews et al. 1999;
O’Rourke et al. 2003). Factors influencing the fruit
cracking in apple are:
Fruit development
Fruit cracking in apple is divided into two kinds according
to the symptoms. One of them is ‘Internal Ring-Cracking’
(IRC) that arises at the fruit stem joint and another is
‘Stem-End Splitting’ (SES) emanating in the stem cavity
(Opara 1996). As IRC is found in all fruits in which SES
happened (Opara 1996), IRC is an early symptom for fruit
cracking followed by SES. Study into the influence of
orchard management works indicated that the ratio of fruits
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Acta Physiol Plant (2015) 37:1718
with SES or IRC was increased with frequent irrigation
compared to fruits without irrigation (Opara et al. 2000).
Thus, cell enlargement induced by water absorption can
have promoted IRC or SES. The imbalance of cell
enlargement in a fruit can cause minute splits among cells
and induce IRC. As IRC occurs just below of epicarp, an
imbalance among the cell enlargement for mesocarp and
epicarp appears to trigger it (Opara 1996). Also, apples
grown detached from the tree and exposed to the sun are
most likely to crack (Verner 1957).
Gene expression
Researchers have found that some genes related to cell wall
have a role in fruit cracking of apple. Expansins are pro-
teins in cell wall that can promote wall loosening and
weakening during cell expansion under expandable stress
(McQueen-Manson and Cosgrove 1994; Cosgrove 2000).
Expansins isolated from cucumber hypocotyls (Cosgrove
1999), tomato (Brummell et al. 1999) and peach (Hayama
et al. 2001) showed correlation with fruit development.
Also, six expansin genes have been isolated from apples;
the expression patterns of these genes were distinguished
during fruit development (Wakasa et al. 2003). Out of
them, MdEXPA3, previously known as MdEXP2, is mainly
appeared during fruit enlargement (Wakasa et al. 2003).
Since expansins enhance extensibility of cell wall and
promote cell expansion, they may have a role in fruit
cracking. Kasai et al. (2008) reported that MdEXPA3
transcripts observed in the mesocarp at 30 days after full
bloom (DAFB) gained a maximum at 95 days after full
bloom and afterward decreased, therefore paralleling the
rate of fruit growth. In contrast, the level of transcript in the
epicarp was below the detection limit until 50 days after
full bloom, and then increased until 109 days after full
bloom to remain high until the end of observation. As IRC
started to occur just before the increase of MdEXPA3
transcript levels in the epicarp, the differential expression
in epicarp and mesocarp can be related to the initiation of
IRC. Bagging decreased the level of IRC and SES to one-
eighth without influencing fruit enlargement, and per-
suaded MdEXPA3 expression at earlier phase in the epicarp
but not in mesocarp. These findings suggest that persuaded
accumulation of MdEXPA3 mRNA in epicarp decreases
the prone of fruit cracking. Therefore, early symptoms of
fruit cracking coincide with situations in which MdEXPA3
expression in the mesocarp exceeds that in the epicarp. In
such situations, epicarp cells can be unable to follow the
expansion of mesocarp cells because of insufficient rate of
growth promoting expansins. If so, IRC is appeared as a
consequence of the imbalance of expansin-dependent tis-
sue growth levels (Kasai et al. 2008).
Acta Physiol Plant (2015) 37:1718
Wax apple
Wax apple (Syzygium samarangense) fruit is an economi-
cally valuable fruit in Southeast Asia. The main problem in
the production of this crop is cracking which reduces its
market value. Lu and Lin (2011) reported that the contents
of total soluble sugars and total titratable acid were both
20 % higher in cracked fruits than in non-cracked fruits of
wax apple, and the osmotic potential was 40 % lower;
water potential was similar; turgor pressure was 60 %
higher, and specific activity of polygalacturonase was
131 % higher. The increase in total soluble sugars and total
titratable acid during fruit maturation leads to decreased
tissue osmotic potential. Water absorption in response to
this decrease in osmotic potential can cause cells to swell,
which increase the turgor pressure resulting in rupture of
the cells and tissues. The increase in polygalacturonase
activity weakens the cell walls. Those combined factors
result in fruit cracking (Lu and Lin 2011).
Grape
In certain years, table grape growers report substantial
amounts of berry cracking in sound clusters as harvest
approaches. Personal observation indicates that this
cracking occurs most often on the fruit’s blossom end and
sides; splitting near the base of the stem is less common.
Berry cracking jeopardizes profitability by increasing har-
vest costs and decreasing yield. Fruit quality and storage
life both on the vine and in cold storage are also reduced
due to the threat of bunch rot; flexibility in marketing is
therefore lost.
Among the table grape cultivars, ‘Flame Seedless’ is
considered quite sensitive to berry cracking. Exotic can
crack if there is a heavy crop load and the bunches are not
properly thinned. ‘Ribier’ is also sensitive, particularly to
blossom end cracking. Cultivars considered moderately
sensitive to cracking include ‘Thompson Seedless’, ‘Ruby
Seedless’ and ‘Cardinal’. ‘Emperor’ would be considered
resistant due to its thick skin (Smilanick et al. 2000). Field
and laboratory researches have attempted to understand the
cause of berry cracking in grape. Factors influencing fruit
cracking in grape are:
Berry skin changes during fruit growth
The grape berry, like many other fruits, has three distinct
growth stages (Peynaud and Ribereau-Gayon 1971).
Critical to understanding the factors associated with berry
cracking are the changes taking place in the grape skin
during fruit development. Meynhardt (1999) showed that
the structure of the berry skin influenced the grape’s
ability to resist cracking. Considine and Kriedemann
Page 5 of 14 1718
(2000) stated that the cracking of sound berries in the
field is evidence that the skin limits growth. They sup-
ported this with the observation that peeled grape berries
imbibed twice as much water as unpeeled berries without
showing any signs of cracking. Considine (1982) and
Yamamoto and Satoh (1994) also demonstrated that skin
fractures developed through cell walls rather than the
simple separation of two cells whose walls were con-
nected by adhesion.
Further understanding of the grape skin’s role in berry
cracking can be found in reviewing some of what is pres-
ently known about the physiological changes it undergoes
during fruit growth. Important to this discussion are the
concepts of elasticity and plasticity. Elasticity is the ability
to recover from having been stretched. A balloon is a
suitable example of elasticity when its air is released.
Plasticity is the ability to retain a new shape. Together
these terms describe the berry skin extensibility during
growth. Matthews et al. (1987) showed that skin extensi-
bility remained relatively constant during the first stage of
fruit growth when cell division is occurring. The same was
also true for Stage II when growth is minimal. Observa-
tions by Considine and Knox (1999) indicated that the
berry skin’s cell walls are thickening at this time. Matthews
et al. (1987) further reported a marked increase in exten-
sibility at the beginning of Stage III, the period of rapid cell
enlargement. At this same time, Considine and Kriede-
mann (2000) observed that the berry skin cell walls became
thinner, perhaps as a result of enzyme activity. Towards the
end of Stage III (2–3 weeks before harvest), a rapid
decrease in berry skin extensibility was observed by Mat-
thews et al. (1987). This ‘‘hardening-off’’ of the skin tissue
may predispose it to cracking should the berry experience a
sudden change in water status.
Although the regulation of grape berry enlargement is
not well understood, present evidence suggests that the
berry skin limits growth and undergoes complex chemical
changes during the course of fruit development. It is con-
ceivable that environmental conditions could have a sig-
nificant impact on these processes and affect the skin’s
ability to resist cracking near harvest.
The role of turgor, berry size and sugar content
Considine (1982) studied the amount of turgor needed to
crack the skin of mature berries. For susceptible cultivars,
they observed that 50 % of the berries were cracked when
the cell turgor was 15 atmospheres. Resistant cultivars
required 40 atmospheres turgor to achieve 50 % cracking.
They believed that mature fruit could contain sufficient
sugar content to absorb enough water to crack the skin
given the right environmental conditions. These were high
soil moisture and a period of low water use by the vine.
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Cool, humid days with little wind, following irrigation,
would be ideal for cracking-susceptible cultivars having
moderate sugar content. Considine (1982) also reported
reduced berry cracking with application of gibberellic acid.
Susceptibility to fruit cracking was not necessarily corre-
lated to berry size or seededness. Direct evidence for the
importance of fruit soluble solids is presented in the
observation of Considine and Kriedemann (2000), who
found that splitting of grapes occurs when they are sub-
merged in solutions of low osmotic potential.
Temperature
Cracking often peaks at the time of peak temperatures and
light. High temperatures in general and sudden high tem-
peratures in midafternoon cause berries to crack. An
explanation for these temperature effects is given by recent
studies with grapes. Raising fruit temperature dramatically
increase the pressure exerted by the pulp on the skin and at
the same time decrease skin stiffness and strength,
increasing the incidence of splitting (Lang and Düring
1990).
Dilute sprays after color break, ethrel, girdling and late
irrigations coincided with a sudden cool period increase
moisture on the berry surface and cause berry crack-
ing (InChang et al. 2007). Calcium treatments can develop
thicker epidermal and sub-epidermal layer of cell wall in
grape and the strengthened berry skin of calcium treat-
ments effectively decrease berry cracking rate under criti-
cal turgor pressure (Choi 2010).
Citrus
Fruit cracking in citrus can be started in mid-summer,
although it mostly occurs in autumn. Fruits of ‘Navel’
oranges are the most susceptible to cracking, followed by
‘Valencia’, ‘Hamlin’, ‘Murcott’, ‘tangelos’ and some tight-
skinned kinds of mandarins. Grapefruit, ‘Dancy’ tanger-
ines, and ‘Temple’ fruits have a lower incidence of split-
ting. The cracks in citrus are usually started at the fruit
blossom end, which is the weakest part in the skin. The
split can be wide and deep or may be shallow and short
(Garcia-Luis et al. 1994). Two kinds of cracking including
radial and transverse have been reported for fruit cracking
in citrus. Radial cracking is more prevalent than transverse.
Besides, partial cracking is more common, while cracking
down to inner center is rather scarce. Organisms such as
Aspergillus, Alternaria, Fusarium, and Penicillium can
infect the cracked area of the fruit and cause disease that
leads to partial rotting and early fruit dropping from trees
(Agusti et al. 2002; Khadivi-Khub et al. 2010). Factors
influencing fruit cracking in citrus are:
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Acta Physiol Plant (2015) 37:1718
Tree age
Fruit of young trees is more prone to cracking than older
ones. The cracking provides a suitable breeding back-
ground for fruit fly, thus the cracked fruits should be
removed from tree. When roots transport sugars and water
to shoots for fruit ripening, the skin may be unable to
expand rapidly enough, thus fruit cracking occurs (Agusti
et al. 2002).
Environment
High fluctuations in temperature, soil moisture, humidity
and fertilizer amounts are closely associated with fruit
splitting. It is concept that the fruit cracking occurs by a
composition of the mentioned factors in comparison to a
single agent. Fruit cracking is usually appeared when
growth conditions become erratic such as drought stress,
insufficient fertilization, sudden rainfalls and hot and cold
temperatures (Garcia-Luis et al. 1994). Thus, sufficient
water and nutrients should be always provided for trees
(Garcia-Luis et al. 1994).
Growth regulators
Gibberellic acid (GA3) influences fruit cracking based on
application time. Application of GA3 at flowering increases
cracking in citrus fruit, but decreases it shortly after the end
of the June drop (Agusti et al. 2002). The application of a
mixture of GA3 and 2,4-D resulted the best findings
(Monselise and Costo 1985; Almela et al. 1994) so that this
treatment significantly reduced fruit splitting. Josan et al.
(1998) reported that GA3 and NAA treatments decreased
fruit cracking in lemon, while ABA content was higher in
control fruits, which controls had higher fruit cracking
ratio. Also, Monselise and Costo (1985) observed that
sprays calcium nitrate at the beginning of cell enlargement
highly decreased the cracking proportion in citrus fruit.
Pomegranate
The main physiological disorder of pomegranate is fruit
cracking. It is resulted from the pressure of the quickly
expanding arils on the stretched peel (Yilmaz and Özgüven
2006). ‘Wonderful’ is a good cultivar with great juice
amount and quality. This cultivar is resistant to fruit
cracking after rainfall on maturing (Karp 2006). ‘Malas-
Saveh’, the famed Iranian cultivar, has moderate suscep-
tibility to fruit cracking. Besides, ‘Jalore Seedless’, ‘Kho-
gand’ and ‘Bedana Bosec’ have comparatively fruit
cracking resistance (Khadivi-Khub 2009). Currently, very
few studies have been carried out aimed at explaining this
Acta Physiol Plant (2015) 37:1718
complex phenomenon physiologically in pomegranate and
suggested some factor influencing it:
Irrigation
Prasad and Mali (2002) reported erratic irrigation or
excessive rainfall during the maturation period causing
fruit cracking in pomegranate. Previously, studies on dif-
ferent fruit species indicated high influences of plant
nutrients and transpiration levels on fruit cracking (Aksoy
and Akyuz 1993). Kumar (1990) reported pomegranate
fruit cracking is increased because of fluctuation of soil
moisture and relative humidity, rain or heavy irrigation
following a dry spell and dry wind. The pomegranate
cultivars have significant differences in susceptibility and
resistance to fruit cracking. Some practices such as
mulching of tree basin and soil working techniques like
crescent bund with open catchment pits have been found to
be useful in many crops for conserving soil moisture
(Sharma and Singh 2008).
Tree age
In young trees, boron deficiency causes fruit cracking.
However, in older trees, this disorder occurs due to mois-
ture imbalance, erratic irrigation, rains and extreme chan-
ges in day and night temperatures. At fruit ripening time, if
the soils become too dry followed by serious watering or
rains, cracking can be occurred. If harvesting of ripened
fruits is delayed for a long time, fruit cracking may be
induced (El-Khawaga 2007).
Growth regulators
Yilmaz and Özgüven (2006) estimated abscisic acid
(ABA), indole 3-acetic acid (IAA) and gibberellic acid
(GA3) contents in fruit peel of pomegranate cultivars. They
reported that the level of ABA was generally lower in the
peel of healthy (non-cracked) fruits than cracked fruits. It
was shown that cracked fruit peel had higher ABA and
lower IAA contents than that of healthy (non-cracked)
fruits. But, no significant differences were detected
between the content of GA3 in cracked and healthy fruit
peels. Growth regulators such as CPPU and nutrients such
as boron also play an important role in improving fruit
quality in different fruits (Sharma and Belsare 2011).
Besides, 5 ppm CPPU in mid-May in combination with
in situ moisture conservation method crescent bund with
open catchment pits (CBOC) decreased fruit cracking and
improved fruit quality in pomegranate cv. Kandhari (Sahu
et al. 2013).
The capacity of water retention in trees should be
increased using organic manures. The trees should be
Page 7 of 14 1718
watered regularly during the fruit development phase.
Application of boron (50 ppm), zinc sulfate, calcium
hydroxide and GA3 (40 ppm) via spraying minimizes the
incidence of cracking on the young fruits (Yilmaz and
Özgüven 2006). Furthermore, Boric acid at 0.20 % under
crescent bund with open catchment pits in mid-May, in-
crease tree vigor, and photosynthetic rate in pomegranate
fruits and decrease fruit cracking (Sahu et al. 2013).
Litchi
Litchi fruit cracking is a serious problem in many com-
mercial orchards. Fruit splitting in litchi is directly corre-
lated with cultivar, watering interval due to hot wind,
endogenous amounts of growth regulators, and higher
levels of GA and ABA in the skin, aril and seed (Menzel
1984). Some reasons for fruit cracking in litchi include:
Gene expression
To know the correlation between fruit splitting and gene
expression, Yong et al. (2006) identified two expansin
genes from fruit of litchi and then measured their expres-
sion profiles in aril and pericarp at different fruit devel-
opment stages, using ‘Huaizhi’ (the cracking-resistant
cultivar) and ‘Nuomici’ (the cracking-susceptible cultivar).
Two full-length cDNAs of 1,087 and 1,010 base pairs
encoding expansin, called LcExp1 and LcExp2, were
extracted from expanding fruit by RT-PCR and RACE-
PCR methods. They reported that the LcExp1 mRNA was
increased and reached to the highest level at the end of
growth phase (80 days after anthesis) in pericarp of
‘Huaizhi’, while the mRNA could be detected at the stage
of rapid fruit growth, and then increased slightly and
finally kept remained almost constant in pericarp of ‘Nu-
omici’. Similar accumulation of LcExp2 mRNA was
observed in fruit aril of ‘Nuomici’ and ‘Huaizhi’, whereas
LcExp2 accumulated only in pericarp of ‘Huaizhi’, but did
not appear in pericarp of ‘Nuomici’. According to their
results, expression of two expansin genes in pericarp of
litchi fruit is closely associated with fruit growth and
cracking.
Fruit development
Cracking is closely related to fruit growth and develop-
ment. The critical period of fruit cracking for ‘Nuomici’
cultivar is from 60 days after bloom until harvest, which is
the rapid growth phase of the aril (Li et al. 1999). Usually,
the increase in fruit diameter exceed 1.00 mm day-1
before the fruit cracked. Fruit usually cracks within
1–2 days after that. This is not to say that fruit growing
abruptly always cracks (Li et al. 2001). The ‘‘spurt of fruit
123
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growth’’ exerts a stress on the peel, which ruptures only if
this exceeds the resistance. This varies with different cul-
tivar and fruit on a tree.
Peel morphology
Mesocarp as the largest portion of litchi pericarp is found
most apt to crack. Litchi cultivars have significant structural
differences. Pericarp with regular mesocarp cells, distinct
and tidy spongy tissues of the lower mesocarp, smaller
intercellular space and clearer endocarp margins have
higher flexibility (Wang 1998; Li et al. 2001). It is con-
firmed that the characteristics of crackles on the peel are
important indices for judging cultivars of different suscep-
tibility to crack. For the easy-to-crack ‘Nuomici’, crackles
are broader and shallower as compared with the resistant
‘Huaizhi’. The tubercles of cracked pericarp are smaller and
sharper than that of the non-cracked peel, indicating the
impeded peel development. The junctions between crackles
of a non-cracked pericarp are tightly and well arranged,
while cracked peel is weakened and less orderly (Li and
Huang 1996; Huang et al. 1999). Rab and Haq (2012)
reported that both skin and pulp traits in fruit of litchi can
characterize susceptibility level of litchi fruit to cracking.
Physio-biochemical characteristics
Peel thickness of ‘Nuomici’ is increased before 62 days
after bloom. It is decreased rapidly when fruits redden,
coinciding with the peak of fruit cracking. This clearly
indicates the reduction of peel thickness leading to weak-
ening of the peel. Using the Threshold Cracking Turgor
(TCT) and Pericarp Tensile Strength (PTT) as described by
Huang et al. (1999), it is shown that both indices were
higher for ‘Huaizhi’ than in crack-susceptible ‘Nuomici’.
Water content, total soluble solids, glucose and fructose in
the aril are higher in cracked fruits, whereas osmotic
potential and sucrose are significantly lower. The decom-
position of disaccharides into monosaccharides might play
a role in fruit cracking (Li and Huang 1996). Also, Sharma
and Dhillon (1998) noted that there were higher ABA
contents in the aril and peel of cracked litchi fruit than that
of non-cracked fruits.
Water relations
The spurt of fruit growth and expansion of aril cells is
caused by water uptake. Drought increases fruit cracking
up to 31 % compared with 23 % in the controls (Li et al.
2001). Drought impedes peel development and reduced
Ca2? import. Defoliation or ABA sprays increase fruit
cracking. The fact that more fruit cracked under low VPD
is attributable to lower leaf transpiration. Furthermore,
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Acta Physiol Plant (2015) 37:1718
increased importation of assimilates and other nutrients
into the aril led to lower osmotic potentials as fruit matured
(Li et al. 2001). The peak of fruit cracking occurs with
respiration during June rain, suggesting a possible link
between respiration and water uptake. This is substantiated
by fruit immersion test. Peak of water uptake occurs at
15 h after fruit being immersed in an ethephon solution,
and is accompanied by a considerable fruit crack, while
water uptake is slow and cracking is avoided in the control.
It is probable that active water uptake by fruit is involved
in fruit expansion (Li et al. 1992).
Girdling and defoliation
It has been reported that girdling and defoliation increased
fruit cracking on 2 and 4 days after these treatments. The
increase of fruit cracking after girdling is attributable to the
increase of sugar in the aril. Besides, defoliation presum-
ably reduces leaf transpiration (Li et al. 2001).
Mineral nutrition
Concentrations of N (NH4? and NO3-) and Ca2? in leaves
of ‘Nuomici’ cultivar are negatively significantly corre-
lated with fruit cracking (r = -0.80), whereas no rela-
tionships with leaf P (H2PO4- and HPO42-), K?, Mg2?
and B (H3BO3) contents. The concentrations of B (H3BO3)
and Ca2? are higher in the normal peel than the cracked
peel and the concentrations of Ca2?, P (H2PO4- and
HPO42-) and Mg2? higher in the aril of a cracked fruit than
in that of normal fruit (Li and Huang 1996; Huang et al.
2001). Exchangeable Ca2? and Mg2? and available P
(H2PO4- and HPO42-), and hydrolyzed N (NH4? and
NO3-) are also lower in the orchard with severe cracking.
It is suggested that Ca2? and B (H3BO3) reduce cracking
(Li et al. 1999).
Environment
Correlations have been found between the daily fruit
cracking rate, and vapor pressure deficit (-0.90), temper-
ature (-0.80), sunshine hours (-0.70) and relative
humidity (0.90) (Li et al. 2001). Occurrence of hot wind
during summer months can cause fruit cracking (Li et al.
2001).
Some commercial orchards now receive Ca2?, B
(H3BO3), Zn2? and K? plus synthetic auxin sprays during
fruit development, organic fertilizer and lime applications
during the winter, and calcium nitrate application before
flowering. Irrigation is also important, along with pruning
and bagging. Spray with boron 0.40 % reduces fruit
cracking at the pit-hardening stage of fruits (Li and Huang
1996).
Acta Physiol Plant (2015) 37:1718
Persimmon
Calyx-end fruit cracking is a type of cracking specific to
Japanese persimmon and is also called calyx separation
(Kitagava and Glucina 1994). Japanese persimmons are
categorized into two kinds including astringent and non-
astringent, on the basis of the absence or presence of
astringency in the fruit at ripening time. However, each
kind is further clustered into two sub-types including
constant and variant types, based on the relation between
presence of seeds and flesh color (Yonemori et al. 2000).
Horticulturally, the cultivars of Japanese persimmon are
currently categorized into the following four types: polli-
nation-variant, astringent (PVA); pollination-constant,
astringent (PCA); pollination-variant, non-astringent
(PVNA) and pollination-constant, non-astringent (PCNA)
(Yonemori et al. 2000). Cultivars exhibiting cracking are
common among PCNA cultivars including leading culti-
vars in Japan such as ‘Fuyu’. However, there are very few
cracking cultivars among the other type cultivars (Yamada
et al. 1988; Yonemori et al. 2000).
There are several causes for calyx-end fruit cracking in
persimmon which include genetic factors, rainfall, high
temperature, water status in fruits, excess nitrogen fertil-
izer, low yield, large fruits and high fruit thinning (Yone-
mori et al. 2000). Two of them have been explained below.
Genetic
Yamada et al. (1988) reported that the inheritance of sus-
ceptibility to calyx-end cracking is controlled quantita-
tively. Cultivars that do not crack are homozygous
recessive, whereas cultivars that crack are heterozygous
(Yamada et al. 1988). Cracking in fruit of PCNA cultivars
has been a serious limitation in breeding programs
(Yonemori et al. 2000). Crosses between cultivars that do
not crack yield mostly non-cracking offspring and the
number of offspring that crack is usually in proportion to
the cracking severity of the parents. Therefore, crosses
between PCNA cultivars inevitably produce a high per-
centage of offspring that crack. The magnitude of cracking
is highly environmentally influenced (Yamada et al. 1988)
and the level of cracking fluctuates greatly from year to
year. Because of large yearly fluctuations, it is very diffi-
cult to precisely evaluate the genotype value of a cultivar
or selection (Falconer 1996). Yamada et al. (2002) esti-
mated environmental variance components for calyx-end
fruit cracking in PCNA persimmons and reported that all
offspring genotypes exhibiting a phenotypic cracking
incidence of less than 20 and 11 % should be selected in
single-year and three-year evaluation, respectively.
Page 9 of 14 1718
Rainfall
The calyx-end cracking may be affected by the time and
quantity of rainfall in relationship to the fruit growth pat-
tern, which is presumably related to the fruit ripening time
(Yamada et al. 1988). The calcium nitrate, calcium chlo-
ride and boron application decrease skin cracking in fruit of
‘Fuyu’ cultivar. The effect of boron is additive when is
used in successive years (Ferri et al. 2008).
Pistachio
Pistachio trees have tolerance to drought stress and may
survive with low water. However, fundamental levels of
water should be provided during the growing season for the
highest yields (Khadivi-Khub 2009). Shells of pistachio
nuts are typically cracked, which are covered by a hull.
But, in many nuts, hull is not cracked so that hull is intact
until after harvesting. This phenomenon distinguishes nuts
of pistachio from walnuts and almonds, because their hulls
are frequently ruptured in the orchard but typically not the
shells (Sherafati et al. 2012). Unfortunately, a small
amount of typical nuts of pistachio, called as ‘‘early split
nuts’’ rupture both their hulls and shells in the orchard and
exposed kernel to invasion by insects and fungi. Another
disorder is that the rupture of the hull usually results in dark
staining of the shell.
The hulls of nuts in pistachio sometimes split, while are
still on the tree. However, this cracking is different from
the hull-cracking of early-split nuts, and may be distin-
guished by the cracking location. For early-split nuts, the
hull is ruptured only when the shell is cracked, and the split
is always along the shell suture or split, whereas hull-
cracking occurred elsewhere and sometimes occurred when
the shell has not split. Another difference is that hull-
splitting in nuts of pistachio only happened very near to
harvesting time.
The hulls in early-split nuts are ruptured throughout a
period beginning in late-July and continued through to
harvest time, usually in September. This long period allows
that molds and infested with the insect navel orangeworm
(Amyelois transitella) decay the kernels of some early-split
nuts. Based on researches, irregular cracking and early
splitting of hull in orchards of the pistachio are the most
important agents that make susceptibly of the pistachio to
aflatoxin contamination (Sherafati et al. 2012). Although
relatively little nuts show early splits, those are very
important, because fungi decay their kernels and they are
frequently contaminated with aflatoxin (Doster and Mic-
hailides, 1995). There are two main reasons inducing early
splits in pistachio including:
123
1718 Page 10 of 14
Irregular irrigation
Water deficiency in orchards in April and May induces
early-split nuts of pistachio in late summer (Doster et al.
2001) or skipping one irrigation in mid-May (during for-
mation of shell) fundamentally increases the amounts of
early splits at harvesting time but do not influence normal
shell splitting (Goldhamer et al. 1985), while excessive
water deficiency in July and August results in decreased
early-split formation. However, water deficiency during
shell hardening, especially in June, does not influence
early-split formation (Doster et al. 2001).
Unbalanced nutrition
Early splitting in pistachio nut can be correlated with
unbalanced nutrition, which induced Aspergillus infection.
Previous studies have shown that K?, P (H2PO4- and
HPO42-) and Fe2? deficiency and Zn2? toxicity are asso-
ciated with early splitting in pistachio nut (Sherafati et al.
2012). Besides, early splitting in pistachio nut has shown
positive and significant correlation with Cu2? content in
leaf and negative significant correlations with K? and Ca2?
content in leaf (Sherafati et al. 2012). Also, it has been
reported that Fe2? and its ratio to other nutrient elements
influenced on early splitting (Hosseinifard and Panahi
2006). Therefore, sufficient irrigation should be provided in
late spring to decrease early splitting in pistachio nuts at
harvesting time. Also, balanced nutrition must be consid-
ered during growth season because it is a main factor for
inhibition of early splits.
Avocado
Fruit cracking can be a problem in some commercial cul-
tivars of avocado such as ‘Zutano’ and ‘Bacon’. Also,
fruits on young trees of ‘Lamb Hass’ cultivar are suscep-
tible to cracking, but cracking is uncommon in the fruit of
‘Hass’ cultivar (Hofshi and Arpaia 2002). There are two
distinct types of cracking in avocado fruit. Stylar end
cracking or breakdown occurs in many cultivars with green
fruit skin and usually begins at the fruit blossom end or
stylar. These splits may be extended nearly to the fruit stem
end (Kadman et al. 1973). Random or side cracking in peel
is observed in fruit of ‘Lamb Hass’ coming from young
trees. It is not known that this is solely a juvenile trait for
‘Lamb Hass’ cultivar or because of several environmental
stresses (Hofshi and Arpaia, 2002). Thus, fruit cracking
may be caused by either environmental or genetic factors.
However, the main causes of fruit splitting in avocado are
unknown, but a supposition is that it may be fluctuations in
soil moisture available to the tree while the fruit is devel-
oping. Depending on cultivar, cracking may or may not be
123
Acta Physiol Plant (2015) 37:1718
accompanied by heavy lenticel corking (Lahav and Whiley
2002). Cultural practices can be potentially modified by
growers to minimize the environmentally induced fruit
cracking (Hofshi and Arpaia 2002).
Banana
Some commercial cultivars of banana such as ‘Mas’ are to
be prone to peel/flesh splitting. Fruit cracking in banana is
an extreme form in which the splits are penetrated deep
into the fruit pulp. Open wounds are provided by splitting
that moisture loss is rapidly facilitated and infection of
opportunistic fungi is permitted (Wo et al. 2005). The
reason of fruit splitting in banana is unclear; however,
periodic water stress and calcium deficiency can increase
this physiological disorder (Nelson et al. 2006). Wo et al.
(2005) reported that moisture level in both flesh and peel of
health (non-cracked) fruits of banana was significantly
lower than those of cracked fruits. Thus, they suggested
that moisture is one of the most important agents causing
fruit splitting in banana.
Tomato
Tomato is one of the most popular commercial vegetables
and its fruit has high susceptibility to cracking. Several
theories are suggested for fruit cracking in tomato that
include:
Genetic differences
Cultivar differences in susceptibility to fruit cracking
clearly exist (Abbott et al. 1986), but the physiological
basis of these differences is far from clear. Although great
strides have been made by breeders in increasing the crack
resistance of modern tomato cultivars, the genetic of crack
resistance is not well understood. Walter (1967) stated that
no researchers have concluded that inheritance of cracking
resistance is simple. Different types of cracking including
concentric, burst, and radial probably are controlled by
different sets of genes. Even when considering only one
type of cracking, direct selection for resistance is difficult
because many genes seem to be responsible, heritability is
low, and environmental influence is high (Cuartero et al.
1981).
Besides, Moctezuma et al. (2003) reported a positive
correlation between cracked fruit number with low levels
of b-galactosidase gene (TBG6 mRNA); so that the strong
suppression causes increasing in the percentage of cracked
fruits. According to TBG6 mRNA expression at the early
steps of fruit growth and the occurrence of cracks related to
decreased amounts of TBG6 mRNA, the cell expansion
may be facilitated with helping TBG6 gene product.
Acta Physiol Plant (2015) 37:1718
Irregular irrigation, especially going from very dry to very
wet conditions
This is the explanation given most often to explain crack-
ing in tomatoes. Modifying in soil moisture during fruit
growth influences strength of skin. Reducing soil moisture
content increases skin strength. In contrast, strength of skin
is reduced when soil moisture content is increased (Ka-
mimura 1977; Peet 1992).
High temperatures and high light
In the greenhouse and in the field, cracking often peaks at
the time of peak temperatures and light. High temperatures
in general and sudden high temperatures in midafternoon in
particular cause red-ripe tomato fruit to crack in the
greenhouse (Frazier and Bowers 1947).
Aspects of fruit anatomy
Skin characteristics necessary for crack resistance have
been characterized in several studies (Cotner et al. 1969;
Kamimura 1977). They showed that wide skin tensile
strength and high elasticity from the turning stage to the
pink stage of ripening were important. Resistant cultivars
all had at least one or both of these characteristics. Certain
fruit shapes and sizes predispose fruit to cracking, as
Considine and Brown (1981) point out in their discussion
of the physics of fruit cracking. As a fruit increases in size,
the physical stress on the enclosing membrane (the skin)
increases, and cracks occur in areas of the fruit where stress
is the greatest. In a tomato fruit, stress is the greatest in the
skin near the calyx (Peet 1992).
Excessively rapid fruit growth causes cracking
Tomato fruit lacks secondary development of the cell walls
and has no way to stop fruit expansion except through
regulation of water pressure (Considine and Brown 1981).
Thus, quickly fruit development can be especially prepared
to splitting. Reduced cracking in cultivars with many fruit
(Young, 1958) could be a result of competition among the
fruits for carbohydrates, thus reducing fruit growth rates. In
greenhouse tomato production, Dutch researchers (Schil-
stra-vanVeelen and Bakker 1985) demonstrated a rela-
tionship between rapid fruit growth and skin crazing or
russeting. Russeting is a form of cracking in which minute
cracks develop all over the surface of the fruit. Fruit that
received more assimilates to enhance growth was more
likely to crack.
Besides, fruit growth rate in tomato during ripening is
highly influenced by environmental factors such as relative
humidity and radiation. This growth rate is as an important
Page 11 of 14 1718
factor which affects fruit cracking because fruit cracking-
susceptible cultivars have a higher growth rate than fruit
cracking-resistant cultivars. Therefore, higher relative
humidity and spring time, which induce higher fruit growth
rate, cause higher fruit cracking percentage. Environmental
factors alter the assemblage of cuticle and its ingredients
during fruit growth and development. High differences in the
cuticle mechanical efficiency in a tomato cultivar are observed
among health (non-cracked) and cracked fruits. Therefore,
cracked fruits are covered with a less deformable cuticle than
health (non-cracked) fruits (Domı́nguez et al. 2012).
High differences between day and night temperatures
Fruit cooling, which occurs at night, would be expected to
contract bulky fruits such as tomatoes (Corey and Tan
1990). This negative pressure could suck in moisture that is
condensed or otherwise present on the fruit surface, and also
could draw in moisture from the plant itself. As tempera-
tures rise during the day and the fruit heats up in the sun,
positive pressures build inside the fruit, stretching the gas-
impermeable tomato skin outward as the fruit expands in
volume (Corey and Tan 1990). Presumably, the greater the
day/night differential, the greater the stress on the skin.
High humidity
High humidity or changes between night and day humidity
are associated with increase cracking. High humidity
increases tomato fruit cracking in the greenhouse, especially
at high temperatures (Byari 1984). The effects of humidity on
tomato fruit cracking may be related to water and gas pres-
sure increases. The tomato skin is 97 % gas-impermeable
(Corey and Tan 1990). As gas and water pressures increase in
the fruit because of a temperature increase or an increase in
water supply, the only way this outward pressure can be
relieved is if the skin expands or if water is drawn out of the
fruit and back into the plant (Peet 1992).
Conclusion
Although fruit cracking has been investigated since the
1930s, and breeders have developed crack-resistant culti-
vars in many species, low advances have been made in
understanding physiology of fruit splitting and cracking in
any of these crops. In turn, this has made it difficult to
recommend preventive measures. There are many causes of
fruit cracking. Genetic and fruit characteristics such as fruit
maturity, fruit size and fruit firmness (Peet 1992; Simon
2006), lack of orchard management such as moisture stress,
nutrient deficiency, insect-pest and disease, bagging and
early picking (Predieri et al. 2003) and environmental
123
1718 Page 12 of 14
factors such as temperature and wind (Simon 2006) are the
main causes influencing fruit cracking.
No cultural practices have been found that can consis-
tently reduce the occurrence of cracking. But, some approach
are suggested that growers can apply for prevent this disor-
der. The best way to reduce fruit cracking at present would be
a suitable orchard management that takes into account and
try to minimize stress of the water, nutrition and physio-
logical factors that contribute to fruit cracking. Also, the
most resistant cultivars to fruit cracking having desirable
fruit quality should be selected for cultivation. Thus, fruit
cracking susceptibility in the selected cultivars should be
evaluated before cultivation. For this propose, the percentage
of fruit cracking for most reviewed fruits here can be eval-
uated by counting the number of fruit cracked after
immersing fruits in water (Rodrigues et al. 2008; Khadivi-
Khub et al. 2008).
Furthermore, choosing cultivation place is a well-effi-
cient way to reduce rain drainage of fruit orchard. The
suitable cultivation place must be with light or no rain
event at near ripening time (Longstroth and Perry 1996).
Furthermore, fruit trees should be covered on the top and
permit free airflow at maturity stage (Meland 2005).
Besides, regular watering should be provided because it is
the most important factor that retains fruit moisture (Prasad
and Mali 2002). Furthermore, spraying of growth regula-
tors such as gibberellins and auxins reduces the occurrence
of fruit cracking in some plants such as litchi, pomegranate,
citrus and sweet cherry (Veera and Das, 1971; Simon
2006). Also, it seems that boron and calcium chloride
solutions are effective in many fruits such as sweet cherry,
apple, citrus, persimmon and citrus to reduce the cracking
of fruits (Wermund et al. 2005; Choi 2010).
Author contribution Dr. Abdollah Khadivi-Khub works
about physiology and breeding of fruit trees. He reviewed
articles and wrote the manuscript.
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