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Orange peel disorder in sweet cherry: Mechanism and triggers

Authors:
  • Lehr- und Versuchsanstalt für Gartenbau und Arboristik e.V. - Obstbauversuchsstation Müncheberg

Abstract

Skin shrivelling of sweet cherry (Prunus avium L.), referred to here as ‘orange-peel’ disorder, compromises fruit appearance and thus market value. The objectives were to establish a protocol to describe and quantify orange peel disorder and to identify the mechanism and factors determining its incidence. Fruit was stored for 28 to 33 d at 2 °C and 76% RH and orange peel disorder was quantified as a topographical roughness using an interferometer or by rating the fruit for orange peel using a four-step scoring scheme. Fruit with orange peel disorder had a skin topography similar to that of a citrus fruit − just on a finer scale. Under the conditions of the test, orange peel was first visible after ∼7 d and continued to increase in severity thereafter. Orange peel was most severe on the shoulder and in the equatorial and distal regions of the fruit. There was no relationship between the distribution of orange peel and that of stomata or of microcracking. At a microscopic level, the depressions in the fruit surface were markedly larger than the periclinal areas of individual epidermal cells, but similar in size to the mesh formed by the network of minor veins visible just beneath the skin. Susceptibility to orange peel differed among cultivars. Least susceptible were ‘Dönissens Gelbe’ and, ‘Gil Peck’, intermediate were ‘Sam’, ‘Kordia’, ‘Merchant’, and the sour cherry ‘Ungarische Traubige’, and most susceptible were ‘Adriana’, ‘Regina’, and ‘Hedelfinger’. Incidence of orange peel during storage was negatively related to relative humidity (more at lower humidities) but also developed at 100% RH in the absence of transpiration. Submerging fruit in water for 2 d partly reversed orange peel. There was no significant difference in the permeance of the skins or in turgors of cells of the outer mesocarp between fruit without and with orange peel. There was a significant difference between the osmotic potential of the flesh (more negative) and that of the skin (less negative). During storage, the osmotic potentials of flesh and skin both decreased slightly, but the difference between them remained constant. The results show water loss from the skin is causal in orange peel disorder. The water loss from the skin occurs both by two routes: (1) transpiration to the atmosphere and also (2) by osmotic dehydration to the flesh.
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('&'CD'$%$'''$'%$$C D('
($'''Q
(&&$'%$)'$'$'$'$%$'
'
1. Introduction
3'$''&$&'($$$$((
44'%>'%$'$'4''$'
%+4&$L''')$'$'
'$')%'4R''$('$)$&
'L$'4%'$(&+$'$4
H$'(4'$'$'$$$%''4'4'$
$4'$)'$%)%'$)'$$'$%
$(>$$L$'%%$$C<'..@
$%%'$ !! $$>S$'$ !!:+) !:$'$ !:D
+'H$)&'E'>%$'F'$$''(4
%'4>H$'(4'4A%'$'
)%%A%'$'H)(4%H$'(
$>%4'4'C6('$..@D+%'$)'%'4'H$'(
>$'4&'(5'')$&
:
$%(&'4'%$$'4''&$'%%(+%$''
'$%$'4'''$'($$('''4'C$$$'
$ !!-6'$ !!''$ !?D3$'''($'
(')($')'%''$4'4'C'$ !?D'
4'4'N4%''$(%$'%$$
%$'$''$C$$$'$ !!-$$$SN
!@D5$('$'4'$)%$&)$(
4'$E$%FE1$FE$$'FE%FCSN
%$$'SN'$ !-D
+$4'$'4'$'(''4$4$'
4'3'$$'1($'$'$$4%$)$')'$
L(E%'F$%%$$+$''%$%($%$($4
CNQ''D&'$'4'4'FE4F
$%%$$+$'$4'C&&4''$orange
peelD$'4$'$N'')'($&CSN'$ !-D
'''$4'$$4$''''$%
$%H'4)%''$''$'$
)'($%(4$''
+%%4''(&$CD'$$'1$H$''$')(
'$%$C D''4('($CD$4$'
$N''$)'(
2. Materials and methods
2.1. Plant materials
?
&'(CPrunus avium 6T$$TTOPTT
TTLTT$TT$'TTK$TTK$TT$TD$(
CPrunus cerasus6T7$+$TD$4'E$?F''CPrunus
cerasus 6AP. canescens *#3D&')$'$''$'4
'$'4'%'$$$''L$''
''$$'$'4'617)'($)$'K'C? I:TU
.I:.TDU%$'$''44$4C<$>%$(D'%'
C$'>$$''D$$'4(4'4$%%'&
$%%5'&%$($'$$''(4$4'
''%$$%$4$)&'$'%$%
'&$'''$$'(3'$4'&'44'(4
$1(4''$)$%4'>44$&$$
+'&$'' '+&$''$$14'
'$%$''&')$C''$ !?D7'&%L4'
&'$$$''$'$' I<4%',,$(
+K&$$G'(4'$)$$'CU$<D(CM-@JKA
..?D3)$4'$&$'$''$4''$
%C ,?>#<$'P''$(D
2.2. Quantifying orange peel
#$%(%'&H$'L'4'($$
%4$$4'$'$%4 -',,$' I<5'&$
$)($CM-JKDU$<CM-@JKD$)&$'CM!!JKD
5&'$4'&$H$'$'$''%$'4$
%4,!
@
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*U$4$O)+S7D$$$$(4'&$CV
:?:*D+>$$4'4$&$'(
)&'4$4'$'&'(4''4'H$'$
$3W !C*D%$ !>4$L$'''
$L$''$$4'4'4$$&$,A, $&
$'$4L)4'%'$''
3)$$&'CE''FD'%$$4,A,
 4'4'4$+A''4$%&$H$'L$E$$4$
F&(%(,O'($'$C'
$H$'*$'( !,D+''%$%&4''$$
4$$roughness*Q($E)'$$4$FC'X!D
$$'$'4$%$'''&CE)$(FD$$)
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'4'$H$%'4)$($'4%$$')''
)'$$4$C'X!D3'%'$''''$'')'$$
'C'X!D'%''4$4'4'4
$4'4$'4$%3'%()$$)'$$'&'
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'4$44'&'1&$'$N'C$'$'
&%$'$D
3$'''4'($4'%&$'H$'4(
')'(4$%+! $,%'4'&''
(%'&'''$')(%'%')(7
-
%L'&$%&$$'4'F'
H$'$%$
2.3. Experiments
2.3.1. Establishing the protocol
+'$A%'4'$$%'4$$
%4!E$F4'%'$')4'$'!',&
'$H'(G'''4'('$$''$'
%<$$'$&'$$$%CS!
6$('V''1$$(DH%%&'$'$
$$CO-,#(%O'$$$(D$%(
$$$(CO-#(%D
2.3.2. Morphological characterization
5H(''4'$)4)$($%$
&'$$'N'$'K&$$$'$'
$'%'&$')4)$($%$H$'L
+'4$$$%)%'&$'$
$' I<$-@JKE$F&'(#$%&$H$'L(
'4'($('$)+''4
$%(%'N'4'4'4$&$'+4'
&$)'$%A$C'$)'(DH$'$$$'$C'($$D1
"
$%'4$%'4'C4'D''&
4'$'4'$''C$D
''$$'%'&'$'$'($'4%$
$)$(''4$4'4$&'$%(%'&'
'$'$('4'(#'$'$'$$$(
''$('''4('%$4$$''$'$
$%'''+'4''$'$&$'$'$4
4$''$'$%')'$'$%'
'%$$$')'$%'')$(+$$(&$$'
T$$TTLTT$TT$'TTK$T$T$T4
L)4'C$$,A,YD&$'%')$
+$'$''14'%4'$%'
%$4$'4'4'%$$%$$$4%$'1
4'$'4''$%$'&'4'
)'&''Q&H$'L'&%%$(
''$'$'$$%4$%4'4''$
$1$'&%'('%($'Z:!C*W>@!#(%D
$%'$%CO-$O-,#(%D+%$$%$$$4
%$&H$'L($$$(CO:
#(%D+144''$%$' !!'?!!['$4
'4$&$H$'L4&'$&'$Q&'C!JD
+$4''&)&'$$)%$)&$
$)'$&'4&'($$$($
''$$$$)4'$$'$$%$
$%$$L'$%%A$'
.
+H$'4('&'4'>)'&'&
%%$$$)$$'$''$%'$\.!
66]C$1(%Q0
U)1(^*$)$O$D_$$\?66]C<A6
U)1(^D_
N$'%:!'$C?!D+'U$U,&$$$'$L$
'$'4,!'%)'$&'4' :'%'$
'$''$'$$Q$)4$'4')
'&+4$'&')$Q
'$4'$%)($)%$%'(
%(C*W>@!#(%D+$$(')'&&
H$'L($$$(CO:#(%D
2.3.3. Factors a"ecting orange peel
%''(4T$$TTOPTTTTLTT$T
T$'TTK$TTK$T$T$T&'(')$$'E7$
+$F(&$)$$'4&'$$' I<$-@JK'&
&$'4'($
+N'4''$4%$%)%'&$'
E$F4'K%'$&$)$$''<+356><$'
C<'+H3'%45''6`$5$$','
@D5'&'$' I<$-@JK$')$$'
!
+N'4'$%$'&$)'$'(EK$F4'$' I<
%('(A$)$'$'$'CA..?D&$'4' "
'$$')'&!!JKC&$'D.JKC<D--JK
CU$<D::JKC<$< D$-JKC($DK&$'(
'4'(5'&'$&'($'4:$&$'>)$%
$'$'$'%$' I<'A%'&$%$'$H'
$'&."JKC&$'D""JKC<D-?JKCU$<D:@JK
C<$< D$!JKC($D3'A%''4'&$($'(
$'1&$'$' I<4 
+'4(&'N%$'('&$'C'$%$'D
$'4'4$%'%$4'4
E$$F$EK$F4'&''$&'$%(%'&$'
%$CD&A$$1$$''4$
'''%$%+&''$'N
$>)$$C*$(>$'*$(6)$(D
'$''4$4'&$A%'LC-$'D4'
NC'$ !!!D+N&L&'1
&$'H'('&$&'$)'$%C'$5a+$>
#N#N$(D+'&$''''
A%'L<&$'%>&'$'''4$
'$$&$'(%$'&4N
$@'%+$'4&$'CF>D&$$$'$'%4$
$'$%'4$)'+%$CP>D4'
'&$')$%&$$$'4
P=F
A ∙ ∆ C

3'H$'AC D'$$4'L4'N$
∆ C
C>,D'$'&$')$%'$''&'4'
NC
Ci
D$'$'%''$C
Co
D
∆ C =CiCo

'
Co
$)($%$'$(1C($P.""D
')44'$%$''
Ci
+
Ci
''$(
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%')'%$'C.::>,$' I<U...D+A%'&$
$'&'?%$'
+4'$Q&$'%''$')%'4$
%(%'&'('$'&$'%''$4
$Q$$''$%4 .O)%'4$%
(%'&$4&(H$'4('4'($('
$)+'%''$4'Q&$'
(&$')$%%'(GA'$'4E$F4'$
$%+&$'%''$&$'(%$('
%($C !?D'4'%
'%$(%$$(&%%$$$1$
%$$%$Q&''4$'
$4>??'!$4',!'&%'('
%(C*W>@!#(%D<$$'$&'$$'$$$
CO-#(%D+4%$(&$'$A%$$
%'$4'''$4%'
''$$'%'&''$'4$%
&)'$'4($)'E$F$4'4 "
'$$'*(''4'$)%$%(%'<
'&$''%%C'..,D$'
%'$'$C !:D$$'$C !:D
*Q(4''$%&%'$$$%$(L&'
C:$<$(D7%%H$'4
'%%&''$)'$%$(&
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'%&''4'$H''''$%$(+
$$$''&$%$$($4'!'?''
%'$%'$'''%4'$%$(($4%$'4'
'&$'%'''$)4'%'&$
$''+%$%'&$'$'
%''$'&$'&' !%$'
2.4. Data analysis and presentation
O$'$$%'L$'$$$''$$'$$
$$'&'(&$'$'$'$(
K$$(&$'''$''$4'&$%$$
C).,3'''3<$(U<DL$4'R'4
'$'$''!!?!!$!!!)$'(222$222
%')(
3. Results
,
5'&'$%$$$%$()''4$
%)$((>%'$'$%%$$%$
+14)$($%$&)$$$$('(&%%
%$'C5*D4$$'$('4'($'
',O$4'4'4$&5<>5+$'
)'(4'$%(%'$A(''$)'
'4'4$&'($%$('$(3'4'(
)$$'($'4$$(&''$($%C5<
!D+L'''$(%'%''$&5O
CD+4'4$$$%$('')$)$($
'%$)%'4$%%)$($%$$
%'$'C5 D+$&55Q''L$
'$4(%')%'C,D+)$($%$&%%
$'(4'''$%$('4$$%%$'$'
4$$'$L$
')'(4$%$$A($'
$)4)$($%$$C5 >OD&')
$%$%)$($%$'$&''&'(
(%'+$A)$(%'$)$?![$'$A
%$'@?[+'4H'()$($%$$%'$
'4$'![C5 *D+)4)$(CYX!.-2225 D$
4%$CYX!.?2225 5D&($'''$$
+L'4$%&$%$(''$$4'-4
'$'''$$)$$'!,%4'C5,'D
O'$')'(4(%'$%' $A('
:
$('C5,D+&$4'$)'(4
'(%''& $ "4'$'''$$$)$
$'C5,'D)'(4(%'N'&4'4'
4$C5,*D(%'&')'4'4''
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+$',$4!'-['4'$4
%',!C5,<D
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)$(+&)&$''$'&$)$'%
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%$$'%G'$$44''$%$''$''
?
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$':$'$'$'C5@*<D)($($'
&$)$$'(%$)%'$'&$($44'
'$^'3''$$( ![
'A%'&$%$'H$'$')($H$''$')($$'$
&'$
''$4' "$$''()$$$')
$'%$'$%''($'$'(C5-D
@
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4'&$$'$&$'>)$%$'$'$'%+'4(&'
$%(%'&)4'&($'$4'$'
'$($'$&$'>)$%$'$'$'%4:C5-D
(1&$'4 C5-*DK($'$&$'>)$%
$'$'$'%$''N'')'(4$%C5-D
&)&4'&$&$')'(4$%$
+$)'($A(&$$&$%&$
)&)$%(%'$&$($)$'$)
)$4' C5-*D
+$%$'4'A44'&''$&'$%
(%'$$(&''C5"D+&$L$'N
%$'&'&''$&'$%C5"'D
$(''A%'&''$'4''&$N$%
'&4'&'$&'''C$'$'&D
+4'Q$&$'%''$$%&$'3
'$4L$'''&$'%''$$'$(H$''
'%''$4Q$C$ !?D3$'4'
'$$'''$%%$$4$%&$
$$$'!?H)$''$-J$C5.D+&$'
%''$&$H$'L($'''4$
'%''$'%''$$C$$')D'
%'$4%$(1$C5.*D5'$
$'%''%''$4$(%''$$''(
?!J%$(&$$$'+H$'$)$&$'%''$4'
-
+F&$'%''$$'(''$%$(
''%''$4'Q$'(&$')$%'(
$$$'($')'$&)'
NCQD'%''$'&'C$')D
$'QC$')D'$'$$$'$'
$'$'!-$C5.<D
+&$N''&4'&''$&'$%
C+$ D+')$'$&$&$((&$')''
'%''$4'4'FA%G
4. Discussion
#'('$'4&%'$'LCD#$%
$$'1($'%'(%4$'%$%('$'$(H$'L
%($'4'(C D+$''$
%N%''(4')$'$%C,D#$%
$(($'
4.1. Orange peel is characterized by an increase in surface roughness that may be
quanti(ed using interferometry
+'4'&$'H$'4($%3'%)'$
)$'''$'$&%$$H$'L$'4
*$%''4'H$)$'4$)'$
$4$''$')$L$$+$$
"
'%$$'44'4($H$'4()$$
4$'%$%('$'$(%$'''$(
+)$($%$%4$%$'''$'4&$
&4H(''+'4H')$($%$$)$
%'$'4$'!['$A)'$4!! 
,$$$)4:-Z!:[,CH)$$'
:?[5:*D')4$)$(&H)$''$': -
4'$$$'4$4''Q$)$)4@
Z!@[,CH)$'::[5:*D($)H)$'4$',
Q+$$)H)$''$',C4($%DQ
&R''%$%''(%$4$%
4.2. Orange peel develops during storage in a cultivar-speci(c manner
#$%)%$$('$$')$>%L$
'4&A%''L'(%'4$%L'$%%$$4'$'
&4'$$' I<$-@J$')'(EK$F&$$'$A%'
'$''(%'&$$(''$$'$)'&'4'&$'
$''$'''7'4'('&$%''4('%4
')$N'%'''$%EOPF$E
F$'%'E$FE$'F$E$F'$'
%''($E$$FELF$EK$F'%'
C+$5?D+N&'''&$$''$'
%''('$%$''$'
4.3. Development of orange peel results from dehydration of the fruit skin
.
#$%'4&$'4'4'+$
$'$'$'%''$%$''$$$'$'$'%
''($'4'('(Q)4'L'
%'4%')$'%'&'KL'4'$'%
'$'$'4&$'C$D$')'(4
$%$A(CD5'&4'
&'$%&$$'&$'4&'$'%$'$(
)+$'')'(4$%$'$%
(%'''($%%$
)4'4($'C''($'4
'D4')$''$''$%')%
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')$4%'4'$'&31&
4'4'&$'%$'''')4'$'&$'
&CK&'$..-D$H*'$C !:D$'
$$$4&$'$''4'4$'$'$($'$
'%$$(&$''$%')$')
'$'('$%$'5'N'$%''(4&4'
')$')$($''N&$''(&''4'
C*'$ !:D3')$&&$'4($(&$''$%
4'4$$%(%$$'$''3'$'
'4'&4&$'''4$($'$')
'%'%%')&'('&)&4
&$&$'(%'4$'&'(C*0&'$
!-D+&'$'$'&$&&'
$'4$&$''%$''''&$$$'%'
&
3$%)$(>'$^
A'%4($'+($$$($
,
'''$')$%4&'(L)
$%$%$'&'(''''4$$4
$'$$$%4'$%$''(
C*$$$ ! D+)$(%'$N4'
$$'&'$%&'(
5. Conclusion
#$%'4$($'4'4$'&'(4'
$(&$''CD'$'%$^'C D'Q$$'4'
$')'%''$+4$'4$4$$$'1(
)$($%$'4'%$'$''('''4
&$'%($('$%^%$'$$%4$%CM,^ @D4
4''$%%'$'%4%'A(
)'$')$>QA%$'&$(4N$^)''4
$($''%$$^&$'$5$
%$'$%'4)&'%'$)'$44'
'$%$'C$'&'%$'$$>'()'D$
'4$%'''$''%+$'
'%''$'&'C$')D$'QC$')D
$$)44($')'%'$4
'$%$''$'(%$(''&$'>)$%$'$'
)'$L$'%%$$+(''$'
$'%$%$''$$'
$'&4'4')$&'A'&%''(''$%
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References
'+# !?'$%$'&'(4'
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 !!@$''4$$$)$'%$
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 !:$'&'$)&'
',.,/ 
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6%%*( !!%$($'
%'$)'44&''$)'*+?",?/:
6(U'O%4$..@$)'$$&'
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(('$7<*3'$'$<$7%%:/::
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U...($$)'$%$'%(($
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4''$'$''4'$'($4$&'''.-
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[$:,"[44'4:$' I<-@JKCOD -.[$@:-[
44'4 "$' I<-@JKCD$, ,[$.?.[C5D4
4'4 "$' I<-@JK
Fig. 25H(''4C*D$)C<OD4)$(C<D
$%$C*ODN'%'$'$4E$F&'(&''
$&'$%(%'$$$4'%%
)'(4(%'&$)$4'4!' "$' I<$-'::J
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 '$'(%'$,)(%'O$'$%'
$d'$$C5DK$'%'&$)4
)$(CD)4%$C5D44'&''$&'$%O$'$
(%')$4'
Fig. 3CD+4$C'D$)%'4$%
E$F&'($A($$'4'
,
$$4'%'$$' I<$-@JK
!4'(%'4$%'(%'
 '$'(%'$,)(%'C*D)'(4
$%N'4'4$$A('$'%
#$%&$H$'L%A$H$'$$'$4'4'
4$''CDC<D$CD5'&$'$' I<
$-@JK4 "',,C<DK$'%'&$$'4
$$4&'(')$N%''('$%CE$$F
EOPFEFELFE$FE$'FEK$FEK$F
E$FD$$(')$CE7$+$FDC<D4)$4'
&'$)&'(')$CE$F'DO$'$%'$d'$$

Fig. 4CDO''4'$'$'&)$($%$'4$44'
4N'')$&'$%(%'4& "', '$$' I<
$-@JKV$($$'($')'%$(%')'
+A%'&$(('$)$%4'$$$'4
'$'$$)$(C*D<$')4H(''4%$$$4
%$4''$%$$4)$($%$44'&'
$%(%'$4'$$(''&4)
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'$''$)$($%$&$$$$'$%+
$)$($%$&4%$$1$'$$('
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,,
Fig. 5K$'%4'4$%4C%(D$$4'
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... Pitting and bruising are expressed as sunken areas in the surface of the fruit that occur when cells in the epidermis-hypodermis collapse due to mechanical damage at harvest or during postharvest handling [17]. On the other hand, pebbling (i.e., orange-peel disorder) is a minor physiological disorder characterized by skin roughness caused by water loss from the fruit skin [18,19]. ...
... Our pebbling results for 'Santina' were lower than those reported by Cliff et al. [47] (i.e., 80% with 'Skeena' sweet cherries after 28 d at 0.5 • C) but similar to those reported by Zoffoli et al. [12] with 'Santina', which is highly susceptible to this disorder. Water loss from the skin has been proposed as the main cause of pebbling, but the problem was not ameliorated under saturated storage conditions such as modified atmosphere packaging [19]. More research is required in order to identify the preharvest practices that affect the fruit epidermal properties controlling skin conductance. ...
Article
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The use of protective covers, such as high tunnels, is recognized as an effective technology to reduce rain-induced fruit cracking in sweet cherries; however, there is a lack of information concerning the effects of this production system on the fruit’s mineral concentration, quality, and postharvest life. This study assesses the feasibility of using high tunnels on ‘Santina’ sweet cherries under the Mediterranean climate of the Central Valley of Chile to obtain earlier harvests of high-quality fruit with long storage life. The study included two plots: Plot 1 during the 2018/2019 growing season, and Plot 2 during the 2019/2020 growing season. High temperatures and relative humidity inside the high tunnels during bloom and fruit set decreased fruit yield, particularly in Plot 1. On average, trees inside the high tunnels were harvested 11 days earlier than those in the open. Fruit from covered trees were significantly larger (13%) and softer (10%) than those from the outside. Fruit quality characteristics, such as soluble solids concentration and titratable acidity, were not affected by high-tunnel-protected cultivation. Fruit from covered and uncovered trees maintained the firmness differences obtained at harvest between treatments, but showed similar postharvest quality after 45 days at 0 °C and a further 3 days at 20 °C on the other characteristics. The covered fruit had lower Ca concentrations (7.7 mg 100 g−1) and higher K:Ca, Mg:Ca, and N:Ca ratios. Significant relationships were found between Ca or K:Ca and fruit firmness at harvest. Lower Ca concentrations in the fruit may explain the lower firmness of fruit grown under plastic covers. There were no differences between covered and uncovered cherries in either cracking susceptibility or induced pitting. ‘Santina’ cherries were very sensitive to pitting damage, but this is not associated with the fruit’s Ca concentration. The results obtained show that high tunnels influenced fruit yield, development, and quality, and emphasize that the fruit’s Ca concentration under this growing condition plays a significant role in the firmness of ‘Santina’ sweet cherries.
... This unexpected behaviour indicates the osmotic potential of the exocarp is less negative than that of the mesocarp-the expressed juice from a cherry is predominantly from the mesocarp 13 . More surprisingly, the osmotic potential difference between exocarp and mesocarp is remarkably stable with time; for example, the difference does not decline when the fruit are stored for quite lengthy periods 14 . Moreover, the magnitude of the difference does not depend on the transpiration history of the fruit 11 . ...
... Incidentally, the water potential difference between the skin (less negative) and the flesh (more negative) represents the driving force for an internal re-distribution of water from skin to flesh that occurs postharvest. This redistribution of water causes the 'orange peel' disorder in sweet cherry 14 . Also, a major fraction of fruit cracks in the stylar scar region which is consistent with the more negative osmotic potential measured in this study 23,24 . ...
Article
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A fleshy fruit is commonly assumed to resemble a thin-walled pressure vessel containing a homogenous carbohydrate solution. Using sweet cherry (Prunus avium L.) as a model system, we investigate how local differences in cell water potential affect H2O and D2O (heavy water) partitioning. The partitioning of H2O and D2O was mapped non-destructively using magnetic resonance imaging (MRI). The change in size of mesocarp cells due to water movement was monitored by optical coherence tomography (OCT, non-destructive). Osmotic potential was mapped using micro-osmometry (destructive). Virtual sections through the fruit revealed that the H2O distribution followed a net pattern in the outer mesocarp and a radial pattern in the inner mesocarp. These patterns align with the disposition of the vascular bundles. D2O uptake through the skin paralleled the acropetal gradient in cell osmotic potential gradient (from less negative to more negative). Cells in the vicinity of a vascular bundle were of more negative osmotic potential than cells more distant from a vascular bundle. OCT revealed net H2O uptake was the result of some cells loosing volume and other cells increasing volume. H2O and D2O partitioning following uptake is non-uniform and related to the spatial heterogeneity in the osmotic potential of mesocarp cells.
... European plum is not the only fruitcrop species where preharvest shrivel symptoms have been reported. Shrivel has also been reported in sweet cherry [3], kiwifruit [4] and grapes [5,6]. ...
Article
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Neck shrivel is a quality disorder of European plum ( Prunus × domestica L.). It has been suggested that backflow in the xylem (from fruit to tree) could contribute to the incidence of neck shrivel in plum. The objective was to quantify rates of xylem, phloem and of transpiration flow in developing plum fruit. Using linear variable displacement transducers, changes in fruit volume were recorded 1) in un-treated control fruit, 2) in fruit that had their pedicels steam-girdled (phloem interrupted, xylem still functional) and 3) in detached fruit, left in the canopy (xylem and phloem interrupted). Xylem flow rates were occasionally negative in the early hours after sunrise, indicating xylem sap backflow from fruit to tree. Later in the day, xylem flows were positive and generally higher in daytime and lower at night. Significant phloem flow occurred in daytime, but ceased after sunset. During stage II (but not during stage III), the rates of xylem flow and transpiration were variable and closely related to atmospheric vapor pressure deficit. The relative contribution of xylem inflow to total sap inflow averaged 79% during stage II, decreasing to 25% during stage III. In contrast, phloem sap inflow averaged 21% of total sap inflow during stage II, increasing to 75% in stage III. Our results indicate that xylem backflow occurs early in the day. However, xylem backflow rates are considered too low to significantly contribute to the incidence of neck shrivel.
... For strawberry fruit, water loss through the fruit skin is likely to be a critical factor in shriveling and compromised appearance. For sweet cherry, shrivel-type phenomena such as 'orange peel' disorder are caused by skin dehydration, with the latter exacerbated if fruit are allowed to transpire excessively [11][12][13]. ...
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Water movements through the fruit skin play critical roles in many disorders of strawberry ( Fragaria × ananassa Duch.) such as water soaking, cracking and shriveling. The objective was to identify the mechanisms of fruit water loss (dry skin, transpiration) and water uptake (wet skin, osmosis). Fruits were held above dried silica gel or incubated in deionized water. Water movements were quantified gravimetrically. Transpiration and osmotic uptake increased linearly with time. Abrading the thin cuticle (0.62 g m ⁻² ) increased rates of transpiration 2.6–fold, the rates of osmotic uptake 7.9-fold. The osmotic potential of the expressed juice was nearly the same for green and for white fruit but decreased in red fruit stages. Fruit turgor was low throughout development, except for green fruit. There was no relationship between the rates of water movement and fruit osmotic potential. The skin permeance for transpiration and for osmotic uptake were both high (relative to other fruit species) but were two orders of magnitude greater for osmotic uptake than for transpiration. Incubating fruit in isotonic solutions of osmolytes of different sizes resulted in increases in fruit mass that depended on the osmolyte. The rate of osmotic uptake decreased asymptotically as molecular size of the osmolyte increased. When transpiration and osmotic uptake experiments were conducted sequentially on the same fruit, the rates of transpiration were higher for fruit previously incubated in water. Fluorescence microscopy revealed considerable microcracking in a fruit previously incubated in water. Our findings indicate that the high permeance for osmotic uptake is accounted for by an extremely thin cuticle and by viscous water flow through microcracks and along polar pathways.
... In contrast, fruit transpiration was enhanced as storage temperature increased, and relative humidity decreased in sweet cherry fruit (Athoo et al., 2015). Relative humidity management at cold storage strongly affected fruit fresh weight loss and the incidence of orange peel in cold-stored 'Regina' sweet cherry (Schlegel et al., 2018). The incidence and severity of physiological disorders additionally enhanced fruit water loss along with the increase in fruit surface area (Moggia et al., 2017). ...
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Fig fruit are highly perishable after harvest. To elucidate the relationship between fruit physiological performance and major targeted metabolites during cold storage, fig fruit of three major cultivars, ‘Masui Dauphine’, ‘Bongresi’, and ‘Banane’, were stored at 0.5 °C for up to 30 d, followed by 1 d at 20 °C. Fruit fresh weight loss increased and fruit length and diameter decreased during cold storage, regardless of cultivars. Lightness and chroma from peel and cortex tissues decreased, although differences were detected among cultivars. Hue angle was differently responsive according to cultivars, fruit regions, and tissues. Soluble solids content (SSC), titratable acidity (TA), and the levels of glucose and fructose increased, irrespective of fig cultivars but individual organic acids remained unchanged during cold storage. The contents of aspartic acid and glutamic acid increased, but the levels of other free amino acids decreased, regardless of cultivars during cold storage. The overall heatmap responses were highest at 30 d, compared with the other removal times during cold storage. Principal component analysis (PCA) scores plot. indicated the biochemical and physiological fruit responses of the three fig cultivars were clearly separated and diverged based on storage times and cultivars. Based on the results of the PCA loading plot, SSC, TA, glucose, fructose, and fruit fresh weight loss were closely linked with glutamate and aspartate. Therefore, the results indicated the physiological and biochemical responses of fig fruit quality attributes and major targeted metabolites could be affected either by storage duration or by fig cultivar.
... Shrivel symptoms are also reported in grapes [Vitis vinifera L. (Bondada and Keller, 2012)], kiwifruit [Actinidia chinensis Planch. (Burdon et al., 2014)], and sweet cherries [Prunus avium L. (Schlegel et al., 2018)]. ...
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Neck shrivel is a physiological disorder of european plum ( Prunus × domestica L.) fruit, characterized by a shriveled pedicel end and a turgescent stylar end. Affected fruit are perceived as of poor quality. Little is known of the mechanistic basis of neck shrivel, but microcracking of the cuticle has been implicated. The objective of our study was to quantify transpiration through the skin surfaces of european plums with and without symptoms of neck shrivel. Cumulative transpiration increased linearly with time and was greater in the susceptible european plum cultivar Hauszwetsche Wolff with neck shrivel, compared with fruit of the same cultivar but without neck shrivel and compared with fruit of the nonsusceptible unnamed clone P5-112. Cumulative transpiration of epidermal skin segments (ES) excised from symptomatic ‘Hauszwetsche Wolff’ from near the pedicel end exceeded that from ES excised from near the stylar end. The permeance of ES from near the pedicel end of ‘Hauszwetsche Wolff’ with neck shrivel (12.4 ± 2.6 × 10 ⁻⁴ m·s ⁻¹ ) exceeded that of ES from near the stylar end (2.9 ± 0.4 × 10 ⁻⁴ m·s ⁻¹ ) 4.3-fold. However, in the clone P5-112, the same difference was only 1.6-fold (1.3 ± 0.8 × 10 ⁻⁴ m·s ⁻¹ vs. 0.8 ± 0.3 × 10 ⁻⁴ m·s ⁻¹ ). Microscopy revealed numerous microcracks near the pedicel end of symptomatic ‘Hauszwetsche Wolff’ fruit but markedly fewer microcracks near the stylar end. The microcracks near the pedicel end were oriented parallel to the pedicel/style axis, whereas those near the stylar end were randomly oriented. Juices extracted from near the pedicel end of susceptible cultivars had consistently more negative osmotic potentials [ψ S (e.g., for Doppelte Hauszwetsche −5.1 ± 0.1 MPa)] than those from near the stylar end (e.g., for Doppelte Hauszwetsche −4.0 ± 0.1 MPa) or that from fruit without symptoms of neck shrivel (e.g., for pedicel end and stylar scar regions of Doppelte Hauszwetsche −3.8 ± 0.1 vs. −3.3 ± 0.1 MPa, respectively). Our results indicate that increased transpiration through microcracks near the pedicel end may contribute to neck shrivel but that the causes of neck shrivel are likely more complex.
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Main conclusion: Xylem flow is progressively shut down during maturation beginning with minor veins at the stylar end and progressing to major veins and finally to bundles at the stem end. This study investigates the functionality of the xylem vascular system in developing sweet cherry fruit (Prunus avium L.). The tracers acid fuchsin and gadoteric acid were fed to the pedicel of detached fruit. The tracer distribution was studied using light microscopy and magnetic resonance imaging. The vasculature of the sweet cherry comprises five major bundles. Three of these supply the flesh; two enter the pit to supply the ovules. All vascular bundles branch into major and minor veins that interconnect via numerous anastomoses. The flow in the xylem as indexed by the tracer distribution decreases continuously during development. The decrease is first evident at the stylar (distal) end of the fruit during pit hardening and progresses basipetally towards the pedicel (proximal) end of the fruit at maturity. That growth strains are the cause of the decreased conductance is indicated by: elastic strain relaxation after tissue excision, the presence of ruptured vessels in vivo, the presence of intrafascicular cavities, and the absence of tyloses.
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Main conclusion: Cell wall swelling, fracture mode (along the middle lamellae vs. across cell walls), stiffness, and pressure at fracture of the sweet cherry fruit skin are closely related. Skin cracking is a common phenomenon in many crops bearing fleshy fruit. The objectives were to investigate relationships between the mode of fracture, the extent of cell wall swelling, and the mechanical properties of the fruit skin using sweet cherry (Prunus avium) as a model. Cracking was induced by incubating whole fruit in deionised water or by fracturing exocarp segments (ESs) in biaxial tensile tests. The fracture mode of epidermal cells was investigated by light microscopy. In biaxial tensile tests, the anticlinal cell walls of the ES fractured predominantly across the cell walls (rather than along) and showed no cell wall swelling. In contrast, fruit incubated in water fractured predominantly along the anticlinal epidermal cell walls and the cell walls were swollen. Swelling of cell walls also occurred when ESs were incubated in malic acid, in hypertonic solutions of sucrose, or in water. Compared to the untreated controls, these treatments resulted in more frequent fractures along the cell walls, lower pressures at fracture (p fracture), and lower moduli of elasticity (E, i.e., less stiff). Conversely, compared to the untreated controls, incubating the ES in CaCl2 and in high concentrations of ethanol resulted in thinner cell walls, in less frequent fractures along the cell walls, higher E and p fracture. Our study demonstrates that fracture mode, stiffness, and pressure at fracture are closely related to cell wall swelling. A number of other factors, including cultivar, ripening stage, turgor, CaCl2, and malic acid, exert their effects only indirectly, i.e., by affecting cell wall swelling.
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Key message Flow in the phloem increased in developing sweet cherry, whereas xylem flow continuously decreased resulting in a fruit water potential that was independent from the tree at fruit maturity. Abstract Rain cracking of sweet cherry fruit is associated with water uptake through the skin, but probably also through the vasculature of the pedicel. The aim of this study was to quantify the xylem, phloem, and transpiration contributions to the overall water balance of developing sweet cherry fruit. Using linear variable displacement transducers, changes in fruit diameter were monitored in untreated control fruit, fruit whose pedicels had been steam-girdled, and fruit whose pedicels had been cut through (these fruits were detached, but remained in situ in the canopy). Fruit volume changes with time were inferred from the measured diameter changes. Pedicel xylem and phloem sap-flow rates, and fruit transpiration rates, were inferred from fruit volume changes. Daily phloem flows were low during stage II (phase of pit development), but markedly increased in stage III (phase of cell enlargement in flesh). Xylem flows exceeded phloem flows in stage II, but decreased continuously in stage III to nearly zero at harvest. Transpiration flow essentially mirrored xylem flow in stage II, but exceeded xylem flow in stage III. Transpiration flow was closely related to the water vapour pressure deficit. Phloem flow was linearly related to the increase in fruit dry mass per-unit time. The data reveal a decrease in xylem sap flow throughout stage III resulting in a water-potential isolation of the mature fruit from the tree. Such isolation would prevent uncontrolled osmotic uptake of xylem water by the sugary flesh and a putative backflow in the xylem from fruit to tree.
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The cuticular membrane (CM) represents the primary barrier to water uptake into sweet cherry (Prunus avium L.) fruit and thus has a central role in rain-induced cracking. The objective was to quantify CM properties potentially relevant to cracking and to estimate variance components and broad-sense heritabilities for these traits in selected sweet cherry cultivars. Within the scion cultivars, CM mass per area ranged from 0.85 g·m-2 in 'Rainier' to 1.61 g·m-2 in 'Kordia'. Wax mass accounted for one-fourth of CM mass and ranged from 0.21 g·m-2 in 'Burlat' to 0.42 g·m-2 in 'Zeppelin'. Biaxial elastic strain of the CM averaged 76.7% across cultivars and ranged from 56.6% in 'Namosa' to 97.0% in 'Oktavia'. Strain was a linear function of fruit mass (r2 = 0.33, P < 0.0001). Partitioning total variance into variance components revealed that fruit mass, CM, and wax mass and strain of the CM had a high genotypic variance and a low residual error variance. Stomatal density ranged from 0.12 stomata/mm2 in 'Adriana' to 2.13 stomata/mm2 in 'Namosa'. The heritability of stomatal density was 67.5%. Across cultivars and years, mean densities of microcracks were of similar orders of magnitude as those of stomata, but ranges were larger and the heritabilities of microcrack density lower. Permeability for transpiration was lowest in 'Flamingo Srim' and highest in 'Nadino'; that for osmotic water uptake was lowest in 'Adriana' and highest in 'Hedelfinger'. Heritability estimates for permeabilities were low. Based on these data, breeding strategies for less cracking susceptible fruit should focus on genotypes that maintain an intact CM throughout development. This may be achieved by selecting for low CM strain and high CM thickness because thicker CM have more "reserve" for thinning. Finally, genotypes that deposit cutin and wax also during Stage III would be most interesting but were not found among the cultivars investigated.
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The skin is the primary load-bearing structure in a sweet cherry fruit (Prunus avium L.). Failure of the skin in rain cracking is considered to be related to water uptake. Little is known of the skin’s water potential, its osmotic potential (ΨΠS), and turgor. The objective here was to quantify ΨΠS relative to the osmotic potential of the flesh (ΨΠF). Spatial resolution was achieved by monitoring plasmolysis in epidermal cells in tissue sections, incubated in selected osmotica using a light microscope method. Decreasing the osmotic potential [ΨΠ (more negative)] of the incubation medium increased the proportion (percent) of plasmolyzed epidermal cells. The pattern of increasing plasmolysis was sigmoidal with increasing osmolyte concentration. The value of ΨΠ for 50% of cells plasmolyzed, depended to some extent on the osmolyte used. The value of ΨΠ became slightly less negative for the osmolytes tested in the order: 1) mannitol, 2) sucrose, and 3) artificial cherry juice (a solution comprising the five major osmolytes of sweet cherry juice in the appropriate proportions and concentrations). There was little difference in the value of ΨΠat 50% plasmolysis between the cultivars Hedelfinger, Sam, and Sweetheart. In all three cultivars, the value of ΨΠF (measured for expressed juice using an osmometer) was markedly more negative than that of ΨΠS (measured for 50% plasmolysis). Incubating skin segments in juice from the same fruit resulted in the plasmolysis of most (85.7% to 96.4%) of the epidermal cells. As fruit development progressed from stage II [27 day after full bloom (DAFB)] to the fully mature stage III (97 DAFB), plasmolysis occurred for increasingly more negative values of ΨΠ. Moreover, the difference between the osmotic potential values recorded for the flesh ΨΠF and for the skin ΨΠS increased. Plasmolysis of epidermal cells was accompanied by a marked swelling of their walls. The results indicate a marked difference in the osmotic potential of flesh (ΨΠFtrended more negative) and skin cells (ΨΠS trended less negative). © 2015 American Society for Horticultural Science. All rights reserved.
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When mature sweet cherries (Prunus avium L.) came into contact with sweet cherry juice, cracking dramatically increased. The objectives of our study were: 1) to quantify the cracking of fruit in cherry juice, 2) to determine which constituent(s) of the juice especially promote cracking and, 3) to establish its/their mode of action in promoting cracking. Artificial juice was made up as an aqueous solution of the same five pure chemicals and at the same relative concentrations as the five major osmolytes of real sweet cherry juice. Artificial and real juice was used at half-isotonic concentrations as the real juice from that batch of fruit. Cracking of sweet cherries placed in either artificial or real juice was more rapid and occurred for lower net water uptakes than of fruit placed in half-isotonic polyethylene glycol 6000. The crack-promoting component in sweet cherry juice was malic acid. Further tests with malic acid, and other organic acids, and with different concentrations of malic acid, with and without pH control, and with the enantiomers of malic acid, showed the effects were primarily related to the pH of the incubation solution. Leakage of anthocyanin from discs of flesh was increased in the presence of malic acid and greater in hypotonic than hypertonic solutions, suggesting that malic acid increases the permeability of the plasma membrane and tonoplast and weakens the cell walls. Malic acid may be an important link (amplifier) in a reaction chain that begins with the bursting of individual epidermal cells and ends with the formation of macroscopic skin cracks. © 2015, American Society for Horticultural Science. All rights Reserved.
Chapter
This book contains 20 chapters and is divided into 4 parts focusing on genetic resources and improvement, ecophysiology and production, pests and diseases and their management and harvesting, processing and utilization of sweet and sour cherries.
Book
The new edition of Physicochemical and Environmental Plant Physiology uses elementary chemistry, physics, and mathematics to explain and develop key concepts in plant physiology. In fundamental ways, all physiological processes that occur in cells, tissues, organs, and organisms obey such relations. Topics include diffusion, membranes, water relations, ion transport, photochemistry, bioenergetics of energy conversion, photosynthesis, environmental influences on plant temperature, and gas exchange for leaves and whole plants. This new edition maintains the unparalleled commitment to clear presentation and improves upon the user friendliness of the previous versions. * All illustrations have been redrawn, many in two-color * New material includes: 14 new figures, 100 new references, 20 new equations and considerable new and revised text * Extensive cross-referencing with a simpler system for chapter sections and subsections * Easy-to-use format including major equations being presented at the beginning of each chapter, and calculations presented outside of the chapter text. Physicochemical and Environmental Plant Physiology, 3rd edition, establishes a new standard of excellence in the teaching and quantitative understanding of plant physiology.
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The rheological properties and structure of the flesh tissue at harvest-ripe fruit of the sweet cherry Prunus avium L. cultivars Bing, Lapins, Regina, Santina, Sweetheart and Van were measured using a textural analyzer equipment and described by light microscopy, respectively. The rheological measurements inflicted reproducible levels of mechanical damage (pitting). The structural and rheological properties of the mesocarp and epidermis correlate with their susceptibility to mechanical damage. Epidermal cell width and area of external mesocarp cells were negatively associated with susceptibility to mechanical damage while cell number, quantified in 1 mm² of external mesocarp, was positively associated. Fruit of Bing and Regina are the least susceptible to mechanical damage among the cultivars under study. The least sensitivity of Regina fruit was associated with its wide epidermal cells and high value of modulus of elasticity, while in Bing with its low number of cells as well as high values of stress and strain at bioyield point. The high susceptibility to mechanical damage of Sweetheart and Lapins fruit was associated with their large number of cells in the external mesocarp tissue and with the lowest values of strain at bioyield point, therefore with the lowest deformation capacity of the tissue.
Chapter
The effects of surfactants on water permeability of isolated astomatous cuticles from Citrus leaves were measured. Rates of water loss across cuticles were determined gravimetrically prior to and after application of surfactants using coverages ranging from 0.015 to 25 g/m². Sodium dodecylsulfate did not affect water permeability. Polyoxyethylene p-t-octylphenol and polyoxyethylene nonylphenol in the HLB range of 4 to 16 increased water permeability by up to 2.1 fold. Polyoxyethylene alkylether in the HLB range of 5 to 13 increased water permeability by seven- to eightfold. The cationic surfactant dodecyltrimethylammonium chloride (HLB 18.5) increased water permeability very effectively by up to twentytwofold.