BLBS102-c27 BLBS102-Simpson March 21, 2012 13:25 Trim: 276mm X 219mm Printer Name: Yet to Come
546 Part 5: Fruits, Vegetables, and Cereals
Ethylene
receptor
C 2 H 4
Gene
expression
Phospholipid
Outside
Inside
Outside
Inside
Ca
Ca
Ca
Ca
Ca
Ca
Ca
Ca
Ca
Ca
H
H
H
H
H
H H
H
Ca
Ca
PLD
PLD
PLD
PLD
Phospholipase D
Increased cytosolic
Ca2+, H+
Phosphatidic acid
Diacylglycerols
Phosphatidate
phosphatase
Free fatty acids
Fatty aldehydes
Calmodulin
Lipoxygenase
Alkanes
Peroxidized fatty
Free radicals acids
Gel phase
formation, reduced
membrane fluidity
Leakage
Damage to Ca2+
-H+ATPase
Lipolytic acyl
hydrolase
Ca^2
Autocatalytic
Figure 27.5.Diagrammatic representation of the autocatalytic pathway of phospholipid degradation that occur during fruit ripening/harvest
stress in horticultural produce.
PLD to the membrane where it can initiate membrane lipid
degradation. The precise relation between the stimulation of the
ethylene receptor and PLD activation is not fully understood,
but could involve the release of calcium and migration of PLD
to the membrane, formation of a metabolising enzyme complex
(metabolon) with other lipid degrading enzymes of the pathway
as well as calmodulin. PLD alpha appear to be the key enzyme
responsible for the initiation of membrane lipid degradation in
tomato fruits (Pinhero et al. 2003). Antisense inhibition of PLD
alpha in tomato fruits resulted in the reduction of PLD activity
and consequently, an improvement in the shelf life, firmness,
soluble solids and lycopene content of the ripe fruits (Whitaker
et al. 2001, Pinhero et al. 2003, Oke et al. 2003, Paliyath et al.
2008a). There are other phospholipid degrading enzymes such
as phospholipase C and phospholipase A 2. Several roles of these
enzymes in signal transduction processes have been extensively
reviewed (Wang 2001, Meijer and Munnik 2003).
LOX exists as both soluble and membranous forms in tomato
fruits (Todd et al. 1990). Very little information is available on
phosphatidate phosphatase and lipolytic acyl hydrolase in fruits.
Proteolysis and Structure Breakdown
in Chloroplasts
The major proteinaceous compartment in fruits is the chloro-
plast which is distributed in the epidermal and hypodermal lay-
ers of fruits. The chloroplasts are not very abundant in fruits.
During senescence, the chloroplast structure is gradually disas-
sembled with a decline in chlorophyll levels due to the degra-
dation and disorganisation of the grana lamellar stacks of the
chloroplast. With the disorganisation of the thylakoid, globular
structures termed as plastoglobuli accumulate within the chloro-
plast stroma, which are rich in degraded lipids. The degradation
of chloroplasts and chlorophyll result in the unmasking of other