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18.1 Fruits 845

CO 2 production decreases. This provides an

explanation for the change in RQ during the

climacteric stage. CO 2 production increases

more rapidly than O 2 uptake, thus the RQ is

greater than 1. The shift from the citric acid cycle

to malate degradation in apples is also reflected

by the effect of citrate and malate on succinate

production. As ripening proceeds, production

of succinate from citrate drops to zero. An

increase in succinate content after addition of

malate in the initial stage of ripening is probably

a feedback reaction. In this case, a decrease

is also observed later on, suggesting a greater

change in metabolic patterns.

18.1.3.3 Changes in Individual Constituents

18.1.3.3.1 Carbohydrates

During ripening of fruits, significant changes occur

in the carbohydrate fraction. For example, between

picking and onset of decay in apples about

20% of the available carbohydrates have been utilized.

During the growth of apples on trees, the starch

content rises and then drops to a negligible level

by the time of harvest. This drop appears to be

related to the increase in climacteric respiration.

Contrary to starch, the sugar content rises. Other

sources in addition to starch should be available

for conversion to sugars. A decrease in hemicelluloses

suggests that they are a possible source.

Organic acids may also be an additional source of

sugars.

A marked decrease in starch in bananas parallels

an increase in the contents of glucose, fructose

and saccharose. Biosynthesis of the latter occurs

by two pathways:

1) UDPG + Fru-6-p → UDP + Sac-6 F -P

→ Sac + P in

2) UDPG + Fru → UDP + Sac (18.42)

The content of hemicelluloses drops from 9% to

1–2% (relative to fresh weight), hence they act as

a storage pool in carbohydrate metabolism. There

is also a drop in the sugar content in bananas during

the post-climacteric stage.

Differences in various fruits can be remarkable.

In oranges and grapefruits the acid content drops

during ripening while the sugar level rises. In

lemons, however, there is an increase in acids.

Decreases in arabinans, cellulose and other

polysaccharides are found in pears during

ripening. Cellulase enzyme activity has been

confirmed in tomatoes.

Remarkable changes occur in the pectin fractions

during ripening of many fruits (e. g., bananas,

citrus fruits, strawberries, mangoes, cantaloupes

and melons). The molecular weight of pectins

decreases and there is a decrease in the degree

of methylation. Insoluble protopectin is increasingly

transformed into soluble forms. Protopectin

is tightly associated with cellulose in the cell

wall matrix. Its galacturonic acid residues are

acetylated at OH-groups in positions 2 and 3 or

are bound to polysaccharides as lignin (R 1 =H,

CH 3 , polysaccharide: arabinan, galactan and possibly

cellulose; R 2 =H,CH 3 CO, polysaccharide,

lignin):

(18.43)

Soluble pectins bind polyphenols, quench their

astringent effect and, thus, contribute to the mild

taste of ripe fruits.

After prolonged storage there is a decrease in soluble

pectins in apples. This drop is associated

with a mealy, soft texture. Similar events occur

in pears, but much more rapidly and with more

extensive demethylation of pectin. Generally, the

degree of pectin esterification drops from 85%

to about 40% during ripening of pears, peaches

and avocados. This drop is due to a remarkable

increase in activities of polygalacturonases and

pectin esterases. The rise in free galacturonic acid

is negligible; therefore it appears that the release

of uronic acid is associated with its simultaneous

conversion through other reactions.

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