An Overview of Calcium’s Role in Lychee Fruit Cracking

X. Huang, H.C. Wang, J. Li., W. Yuan, J. Yin

J. Lu and H.B. Huang Dongguan Agricultural Research Center
College of Horticulture Dongguan, Guangdong
South China Agricultural University China
Guangzhou
China

Keywords: calcium deficiency, pericarp, anions, cations, ontogeny, kinase, cell wall

Abstract
This paper overviews the studies of calcium in relation to lychee fruit cracking.
Evidences support the following conclusions about calcium’s role: (i) Deficiency of
calcium causes severer cracking, (ii) calcium contributes to cracking resistance
through its structural role in the cell walls of the pericarp, (iii) there are a number of
barriers that hinder the calcium applied to become a structural part in the cell walls,
(iv) the effect of calcium applied may vary with time of application and its combined
anions, (v) availability of calcium in the early stage of fruit ontogeny is important for
cracking resistance, and (vi) calcium contributes only part of cracking resistance and
application of the element is not the whole answer towards the problem.

INTRODUCTION
Fruit cracking is a major physiological disorder in lychee being common in China
(Chen and Huang, 2001) and India (Mitra and Ghosh, 1991), the two largest lychee
producers in the world. It also occurs in Thailand (Sethapakee, 2002), Nepal (Budathoki,
2002) and South Africa (Joubert, 1970, 1983). In China, the annual loss caused by fruit
cracking in ‘Nuomici’ (Fig. 4) alone is equivalent to US$ 2.5 million (Li et al., 1992). In
India, one third of the crop may be lost in susceptible cultivars (Kanwar et al., 1972b).
Many studies have examined the roles of temperature, water, plant growth
regulators and mineral nutrients in fruit cracking. These studies have yielded valuable
information concerning the causes of lychee fruit cracking. However, it remains a tough
problem that awaits effective tackling measures.
Although climatic factors such as drought, high temperature and excessive rains
were found to induce cracking (Kanwar et al., 1972a and b; Kanwar and Nijjar, 1984a and
b; Sharma and Ray, 1987; Chande and Sharma, 1992; Li et al., 2001), cracking incidence
varies significantly among trees within the same orchard that have encountered the same
climatic conditions, which indicates that internal factors in the trees are also affecting cracking
incidence. Tree nutrient status is the prime concern among all of the internal factors; and
among the mineral nutrients, calcium is the most extensively studied. Calcium being one of
the essential macro-elements in plants; its deficiency in fruit is associated with a wide
range of physiological disorders including fruit cracking (Shear, 1975).

ROLES OF CALCIUM IN FRUIT CRACKING

The roles of calcium in fruit cracking have been examined by: (i) comparing
calcium contents in cracked and normal fruits; (ii) comparing calcium contents in
cracking-resistant and susceptible cultivars; (iii) comparing calcium status in trees with
different cracking incidence; (iv) comparing calcium availability in soils in orchards with
different cracking incidence; and (v) studying the effects of calcium application.

Comparisons among Fruit, Trees and Orchards

Cracking rate was reported to be unrelated to calcium concentration in nectarines
(Fogle and Faust, 1976). In lychee however, the pericarp of cracked fruit had significantly
lower concentrations of calcium than those of the unaffected fruit within the same cultivars
(Sanyal et al., 1990; Li and Huang, 1995; Li et al., 1999; Lin, 2001). Trees with lower

Proc. 2nd IS on Lychee, Longan, Rambutan & 231

Other Sapindaceae Plants
Eds. N. Chomchalow and N. Sukhvibul
Acta Hort. 665, ISHS 2005
cracking rates had higher calcium level, which was significantly and negatively correlated
to cracking rate (P<0.05; Fig. 1). In addition, orchards with high cracking rates had lower
concentrations of exchangeable calcium in the soil. Calcium absorbed through roots is
transported with the transpiration stream in the vessels. Fruits usually have a lower
transpiration rate and consequently lower calcium concentration than leaves. Li and
Huang (1995) found that under drought conditions calcium uptake by lychee fruit was
reduced and the transpiring leaves might draw calcium out of the fruit, and this was
paralleled with higher fruit cracking rates.

Differences among Susceptible and Resistant Cultivars

Sanyal et al. (1990) found no relationship between cracking and concentration of
calcium in the pericarp of different lychee cultivars. However, the tested cultivars with
cracking rate ranging from 0.7 to 12% were different in maturation season, and thus fruit
development might have encountered different climatic regimes, which was inadequate to
give a significant comparison. By studying ‘Huaizhi’ and ‘Nuomici’ cultivated in the
same location, both being late maturing but differing significantly in cracking rates (<1%
vs. >20%), our results showed that the calcium concentration in the pericarp of highly
cracking-resistant ‘Huaizhi’ was significantly higher than in that of susceptible ‘Nuomici’
(Fig. 2).

Effects of Calcium Application

Reports have shown that calcium application is effective in reducing fruit cracking
in sweet cherries (Verner, 1938; Tukey, 1984; Callan, 1986; Ruper et al., 1997; Lang et al.,
1998; Demirsoy et al., 1998a; Richardson and Ystaas, 1998; Fernandez et al., 1998;
Howard et al., 1998; Marshall and Weaver, 1999), apples (Moon et al., 1999), peaches
(Sun and Liu, 1997), oranges (Xu et al., 1994; Zou et al., 1995), pears (Raese, 1987), prunes
(Cline and Tehrani, 1973) and figs (Aksoy et al., 1994), although no significant effect has
been reported in watermelon (Ali et al., 1998), grapes (Combrink et al., 1982; Nam et al.,
1996) and sweet cherries (Hermann and Feucht, 1984; Looney, 1985; Koffmann et al.,
1996). There are some studies suggesting that the effects of calcium in conjunction with
other elements like boron in muskmelon (Combrink et al., 1995) and copper in sweet
cherry (Brown et al., 1995) were better than calcium applied alone. Cracking of detached
sweet cherries could be induced when immersed in distilled water, but significantly
reduced when the fruits were pre-dipped in the solution containing calcium (Alani, 1980;
Hermann and Feucht, 1984; Glenn and Poovaiah, 1989). In contrast, treatments with
calcium-chelating agent EDTA (ethylenediamine tetraacetate) or EGTA (ethyleneglycol-bis-
(beta-aminoethyl ether) N, N, N, N-tetraacetic acid) which removes calcium from fruit
tissues resulted in severer cracking (Alani, 1980; Glenn and Poovaiah, 1989).
Immersion test of detached ‘Nuomici’ lychee made by Ou (1988) showed that
treatment of calcium hydroxide at 0.4% increased the critical cracking turgor (0.25 vs.
0.17 MPa) measured with pressure chamber method (see Huang et al., 1999) and pericarp
tensile strength (111 g.m-2 vs. 78 g.m-2). In field experiments, effects of the application of
calcium salts either alone or combined with other nutrients thus far reported have been
positive. Peng et al. (2000) sprayed three times with 100 mmol.L-1 Ca(NO3)2 at full
bloom, 19 and 39 days after full bloom (DAFB) reduced fruit cracking in ‘Nuomici’ from
14.4 to 10.4%. Li et al. (1999) treated ‘Nuomici’ trees with a chelated calcium solution
(containing 180 mmol.L-1 Ca) and brought the cracking rate down to 17.5% from the
27.7% in the control. Trees treated with a high-Ca (132 mg.L-1) nutrient solution
produced 15.4% cracked fruit, while untreated trees produced 47.2% (Qiu et al., 2000).
Our two years’ results (Table 1 and 2) also showed that calcium application generally
reduced cracking, but the effect varied with the time of application and calcium
formulations. As cracking incidence differed greatly among individual trees, the effects
were generally not significant on a statistical basis. It seems that the effect of calcium
application to alleviate cracking is limited.
The above studies suggest that calcium as a necessary element play an important

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role in lychee cracking, and deficiency of the element tends to lead to severer cracking.
However, the limited effect of calcium application on reduction of cracking necessitates a
better insight towards calcium’s acting modes in the fruit.

HOW CALCIUM ACTS

Cracking, Cell Walls and Calcium

Calcium serves as an intracellular “secondary message” by activating enzymes
such as protein kinases, alone or in complex with calmodulin. The activated kinases
covalently modify other target enzymes, and consequently modulate physiological
responses (Zocchi and Mignani, 1995). The function of “secondary message” is fulfilled
at a very low calcium concentration (at the micromolar level).
Calcium also plays an important structural role in the cell walls and cell
membranes. In these cases a millimolar calcium concentration level is required for
calcium to serve as the cementing materials in the cell walls and to maintain the integrity
of the membranes (Zocchi and Mignani, 1995). As the structural role requires a three
digital higher calcium concentration than its role of “secondary message”, calcium
deficiency related disorders are therefore mainly associated with the failure of its
structural role.
Fruit cracking happens when the stress exerted on the fruit skin from the
expanding aril exceeds the mechanical strength (the strain) of the skin, which is largely
contributed by the cell walls. The load-carrying materials in the cell walls include
cellulose integrated into microfibril networks, and hemicellulose and pectin within the
wall matrix (Brett and Waldron, 1990). Calcium is the cementing material in the matrix
phase as it bridges pectin molecules at carboxylic groups through ionic bonds, and
therefore contributes to the mechanical strength of the cell walls. In this way, calcium
may contribute to fruit cracking resistance (Alani, 1980; Sekse and Ystaas, 1998;
Richardson and Ystaasm, 1998; Huang et al., 1999, 2001). In sweet cherry,
autoradiographs of 45Ca2+ showed the epidermal region to be the site of Ca2+ action in
altering fruit cracking (Glenn and Poovaiah, 1989). Removal of calcium from cell walls
with chelating agents such as EDTA and EGTA increased solubility of the wall pectin and
induced severe cracking in sweet cherry (Alani, 1980; Glenn and Poovaiah, 1989).
To reveal the detailed association between structural calcium in cell walls of the
pericarp and cracking in lychee, it is necessary to develop techniques in quantifying the
concentration of structural calcium.

Methodology in Studying Structural Calcium in the Cell Walls

Calcium concentration in plant tissue can be determined with ashed samples by
means of atomic absorption method. But this method does not discriminate between free
and bound calcium. Free calcium (Ca2+) can be analyzed by a number of methods
including selective microelectrodes (Felle, 1989), fluorescent probe (Grynkiewicz et al.,
1985; Bush and Jones, 1990) and antimonite precipitation (Peng and Iwahori, 2001).
However, there are some technical difficulties in quantifying the concentration of
structural calcium in the cell walls. Localized and in situ semi-quantitative microanalysis
of calcium in the cell walls can be made with X-ray electron microprobe (scanning
electron microscope plus energy dispersive spectrometer) (Glenn and Poovaiah, 1990;
Huang et al., 1999). By using this technique, we revealed that the concentration of
calcium in the cell walls of the endocarp, mesocarp and epicarp of cracking-resistant
‘Huaizhi’ was significantly higher than in susceptible ‘Nuomici’ (Huang et al., 1999).
Quantitative determination of calcium concentration in cell walls can be made by
stepped extraction of tissue calcium in conjunction with atomic absorption method (Chen
and Uemoto, 1976). Procedure of sequential extraction of different forms of calcium in
plant tissues is shown in Fig. 3. Acetic acid is capable of dissolving calcium in both pectate
and phosphate. Since the concentration of calcium pectate is by far higher than calcium
phosphate in plant tissues (Chen and Uemoto, 1976; Gong et al., 1992; Dong et al., 2003),

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we used calcium in acetic acid soluble fraction to represent the structural calcium in
lychee pericarp, which was higher in ‘Huaizhi’ than in ‘Nuomici’ (Huang et al., 2001).
Our recent study based on X-ray microanalysis and sequence extraction suggested
that endocarp was likely the major site of calcium action in increasing cracking
resistance (results to be published).

FACTORS AFFECTING THE EFFECT OF CALCIUM APPLICATION

To make the application effective, calcium cations must enter the pericarp tissues.
As it is very difficult for calcium to penetrate the cuticle layer, uptake of calcium applied
to the surface of the fruit is mainly through the stomata. Our observations with scanning
electron microscope showed that stomata were very sparse on lychee epidermis,
indicating that only small part of calcium adhering to the fruit surface could be absorbed.
Weather conditions such as humidity and temperature, which influence stomata behavior
may have significant effect the uptake of applied calcium.
Even after the entry to the pericarp, calcium must be translocated to the cell walls.
Calcium as a less mobile element, the mechanism for its translocation is still unclear.
Moreover, applied calcium may be precipitated with organic acids present in the pericarp
such as oxalic acid. X-ray microanalyses revealed the epidermal cells of ‘Nuomici’
pericarp contained calcium-rich solids in the cytoplasm (Huang et al., 2001), which
proved the occurrence of calcium precipitation that reduces calcium availability to the cell
walls. Finally, calcium binding with the pectin depends upon the availability of negatively
charged carboxylic groups (uronic group), which might be blocked by methyl
esterification (Zocchi and Mignani, 1995). Therefore, synthesis and metabolism of pectin
in the cell walls influence the formation of structural calcium.
Because of the barriers mentioned above, only a small portion of applied calcium
could be used for the cell wall construction, and thus produces limited result in cracking
control. To overcome these barriers, perhaps calcium should be applied at proper time and
in proper formulation.
Callan (1986) found Ca(OH)2 more effective than CaCl2 in reducing cracking in
sweet cherry. In our study, under the same molar concentration of calcium, Ca(NO3)2
solution seemed more effective than CaCl2 solution (Table 2) in controlling lychee
cracking. The results indicate that the anions of calcium salts may play some roles.
Whether they influence cracking through their direct effect or through their effect on
calcium absorption needs clarification. Although addition of chelating agent (citric acid)
and NAA into the CaCl2 solution showed some effect on lowering cracking rate in 2001
(Table 1), more investigations should be made to ascertain the real effect.
Results (Table 2) of single spray of different formulations of calcium at three
developmental stages showed that the application of calcium at early fruit development
stage (two weeks after anthesis) was most effective, followed by the application prior to
aril expansion. Poorest effect was obtained in the application at five weeks after anthesis.
The results suggested that the supply of calcium in the early stage and prior to rapid aril
growth was important for cracking resistance.

FURTHER CONCERNS IN CALCIUM STUDY

Since calcium contributes to cracking resistance through its structural role in the
cell walls, mechanism and regulation of structural calcium formation need to be
understood. A good approach is to study the metabolism of pectin that bonds calcium
with its negatively charged carboxylic groups (uronic group). The amount of calcium
bound depends upon the amount of pectin, the number of uronic groups and the degree of
methyl esterification.
Calcium availability is reduced when it is precipitated with organic acids such as
oxalate. Production, distribution and metabolism of organic acids in the pericarp is
involved in calcium action and remains to be studied in relation to fruit cracking. Ways to
prevent applied calcium from being precipitated needs to be found.
The time effect of calcium application on fruit cracking necessitates a closer look

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into the pattern of calcium uptake by fruit throughout the development. As the effect of
calcium application can be influenced by the accompanying anions, improved
formulations of calcium can be obtained by selecting the better combinations.

CONCLUSION
The contribution to cracking resistance by calcium is related to its structural roles
in the cell walls. Applied calcium may not become the structural part of cell walls
because there are barriers in calcium uptake and mobilization, and blockages in calcium
binding sites in the cell walls. It seems that the availability of calcium during early fruit
development is important for cracking resistance and the effect of applied calcium may be
influenced by its combined anions. There are still many aspects concerning calcium’s role
in fruit cracking that remain to be understood.
Lychee cracking is caused by multiple factors including genetic background,
physiological status of fruit and environmental conditions. Although there are good
evidences to prove the close relationship between calcium and fruit cracking, calcium is
only a part of the answer to the problem, viz., we cannot expect ‘Nuomici’ as cracking
resistant as ‘Huaizhi’ through supplying it with calcium. But we have no way out until
‘Nuomici’ is to be replaced by other cultivars as elitist and appealing as it is. Increase in
calcium availability is effective in reducing cracking only when there is deficiency of the
element. Controlling cracking should be approached through integrated management.

ACKNOWLEDGEMENTS
Our studies relevant to this overview were supported financially by the National
Natural Science Foundation of China (Project No. 30000104) and Natural Science
Foundation of Guangdong Province (Project No.990687).

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Zou, H.Q. and Xu, J.K. 1995. Studies on the relation between peel anatomical structure
and fruit-cracking in ‘Hongjiang’ sweet orange. J. South China Agri.Univ. 16:90-96.

Tables

Table 1. Effects of different calcium solutions on cracking in ‘Nuomici’ lychee. Calcium

compounds were sprayed to fruit 4, 6 and 8 weeks after full bloom (WAFB); different
treatments are conducted on the same trees with four replicates. Concentrations: CaCl2
20 mmol.L-1; CA (citrate acid), 13.3 mol.L-1; NAA, 40 mg.L-1 (result of 2001).

Treatment CK CaCl2 CaCl2+CA CaCl2+ CA +NAA

Cracking rate (%) 21.5±11.4 16.6±2.4 21.7±10.2 13.2±2.6

Table 2. Effects of single spray of different calcium spray at different stages on cracking rate
(%) in ‘Nuomici’ lychee. Different treatments were done within the same trees with
three replicates. Concentration: CaCl2 40 mmol.L-1; CA (citrate acid) 27 mmol.L-1;
NAA 40 mg.L-1 (results of 2002).

Treatment Treatment at 2 Treatment at 5 Treatment at 8

WAFB WAFB WAFB
CK 33±7.56 49.8±17.5 34.8±2.4
CaCl2 30±10.7 34.1±18.7 31.8±8.2
Ca(NO3)2 19±8.0 42.4±23.2 15.6±1.8
CaCl2+NAA 21±15.1 45.5±7.6 27.6±15.6
CaCl2+CA 20±12.2 53.7±21.1 34.1±0.8
CaCl2+CA+NAA 22±15.1 72.7±26.7 26.6±4.9

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Figures

50

40

Cracking rate (%)

30

20

10 2
R = 0.8145
0
10 10
.5 11 11
.5 12
Ca content in leaf (mg/g dw)

Fig. 1. Correlation between cracking rate and calcium concentration in the leaf (Li et al.,
1992).

10.0
9.0
Ca concentration (mg/g DW

8.0
7.0
6.0
5.0
4.0
3.0
2.0
1.0
0.0
5/5 5/12 5/19 5/26 6/2 6/9 6/16 6/20

1.60

1.20
Cal concentration
(mg/g fw)

0.80

0.40

0.00
4.21 5.50 5.19 6.20 6.16

Date

Fig. 2. Comparison of calcium (extractable with 1 mmol.L-1 HCl) concentration in

pericarp of ‘Nuomici’ lychee (■) and ‘Huaizhi’ (□) A: results of 2000 on dry
weight basis; B: results of 2002 on fresh weight basis.

239
 Plant tissue
⇓
Grounded in ion-free water set for 2 hr
⇓
Centrifuge ⇒ supernatant – water soluble calcium
⇓
Precipitation suspended with 1 mol/l NaCl solution for 2 hr
⇓
Centrifuge ⇒ supernatant – calcium pectate
⇓ (wall-bound)
Precipitation suspended with 2% acetic acid solution for 2 hr
⇓
Centrifuge ⇒ supernatant – calcium phosphate
⇓
Precipitation suspended with 0.6 mol/l HCl solution for 2 hr
⇓
Centrifuge ⇒ supernatant – calcium oxalate
⇓
Precipitation – Calcium silicate etc.

Fig. 3. Procedure of sequential extraction of different forms of calcium in plant tissues

(Chen and Uemoto, 1976).

Fig. 4. Severe cracking in ‘Nuomici’ lychee.

240