BIOCHEMICAL CHANGES IN

FRUIT RIPENING
Mantana BUANONG
 LEARNING OBJECTIVES
1. To identify the plant organs that are subject to
continuous changes after harvest.
2. To understand biological factors involved in
deterioration.

2
https://vignette.wikia.nocookie.net/powerlisting/images/2/2b/Apple-stages-of-decay.jpg/revision/latest?cb=20151102010213
PLANT ORGANS

Vegetative organs
1. Root
2. Stem
3. Leaf
Vegetative organs
❑Root is an organ of a plant body, typically
growing through gravity direction.

❑The major function of roots is absorption of water

and inorganic nutrients. Many roots often function
in storage of food.

❑Dietary roots are such as carrot, radish, beetroot,

etc.
❑ Stems are the organ supporting for the elevation
of leaves, flowers and fruits, transporting of fluids
between the roots and the shoots in the xylem
and phloem, and storing of nutrients.

❑Dietary stem are such as bamboo shoot,

asparagus, ginger, etc.
❑ Leaves containing chlorophyll are the main site
for photosynthesis that would stored some
nutrient and water. Leaves are prominent in the
human diet as leafy vegetables.

❑ Plants using leaves for food include leafy herbs,

leafy vegetables, lectuce, etc.
 PLANT ORGANS
Reproductive organs
4. Seed
5. Flower
6. Fruit
Reproductive organs
❑ Monocotyledons (monocots) and dicotyledons
(dicots) make up the two large groups of
flowering plants, differentiated by their seed
structures.

Coconut – the biggest plant seed used for fresh market.

▪ Flowering plants as angiosperms, which are more
widespread than any other group of plants.
▪Many plants produce highly visible flowers that have a
distinctive size, colour, or fragrance.

▪Flowers are used for food such as broccoli,

cauliflower, etc.
FRUIT: a matured ovary containing one or more seeds.
The three fruit layers are:
1. Exocarp: the outermost layer often consisting of
only epidermis.
2. Mesocarp: or middle layer which varies in
thickness.
3. Endocarp: shows variation from one species to
another.

https://home.csulb.edu/~rodrigue/geog330/fruitsseeds.html
TYPES OF FRUIT
1. Based on the floral origin.

2. Based on floral structure.

3. Based on physical characteristics.

BASED ON FLORAL ORIGIN

Derived from more than

one pistil on one flower:
Strawberry, raspberry
Derived from a single Derived from several
pistil of one flower: flowers, (inflorescence):
Pea, tomato, lily, apple Pineapple, fig
and cucumber https://www.backyardnature.net/frt_aggr.htm
12
BASED ON FLORAL STRUCTURE
1. True fruits
- Composed of only the ovary wall.
- Pea, tomato, lily
2. Accessory fruits
- Composed of the ovary wall plus other flower parts.
- Apple, cucumber, strawberry, fig
BASED ON PHYSICAL CHARACTERISTICS
Flesh fruit
1. Drupe
- One carpel with a single seed, fleshy outer and
inner stony layer (peach, plum).
BASED ON PHYSICAL CHARACTERISTICS
Flesh fruit
2. Berry
- Consist of one or more carpels with few to many
seeds - the ovary wall fleshy (tomato, grape and
banana).
BASED ON PHYSICAL CHARACTERISTICS
Flesh fruit
3. Hesperidium
- Tough rind, numerous carpels separable as
‘sections’ (orange, grapefruit, lime).
BASED ON PHYSICAL CHARACTERISTICS
Flesh fruit
4. Pepo
- Leather-skinned, one carpel with 3-4 lines of
ovules (melon).

17
BASED ON PHYSICAL CHARACTERISTICS
Flesh fruit
5. Pome
- Fleshy accessory fruit (a fleshy fruit composed of
a mature ovary along with other enlarged parts of
the flowers), with cartilaginous core (apple, pear).

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 VEGETABLE

All plant parts could

be used for food using
horticultural prospect
from the roots to the
shoots. Vegetables
can be grouped into
three main categories:

• Bulbs, roots and tubers

• Flowers, buds, stems and
leaves
• Fruit, seeds and pods

Wills et. al. 2006. Postharvest: An Introduction to the Physiology and Handling of Fruit, Vegetable and Ornamentals. (5ed.) UNSW Press, Sydney.
❑ Fresh fruits, vegetables and flowers are living
tissues that are subject to continuous changes
after harvest!
❑ Some of these changes are desirable, but most
are not wanted. The main goal of postharvest
research is to slow these changes as much as
necessary.

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❑ After harvest – fruits and vegetables are
detached from the mother plant and do not
‘enjoy’ anymore from continuous supply of water
and nutrients.

❑ Therefore, after harvest, fruit and vegetables

depend on their own carbon and water reserves
and become perishable – they loose water and
dry matter!

21
 BIOLOGICAL FACTORS INVOLVED IN
DETERIORATION
❑ Respiration
❑ Ethylene production
❑ Compositional changes
❑ Water loss
❑ Physical damage
❑ Physiological breakdown
❑ Pathological breakdown

22
https://www.ift.org/iftnext/2020/may/how-ripe-is-your-produce
 MAJOR DEVELOPMENTAL CHANGES DURING
TOMATO FRUIT DEVELOPMENT AND RIPENING
http://www.plantcell.org/content/16/suppl_1/S170

Cell division Carotenoid

Ethylene/respiration
Cell expansion Softening

Relative changes in cell division, cell expansion, respiration, ethylene

synthesis, fruit softening, and carotenoid accumulation are shown over
the course of fruit development. The time from anthesis (a) to mature
green (MG; fully expanded unripe fruit with mature seed), breaker (BR;
first visible carotenoid accumulation), and red ripe (RR) can vary
23
substantially among cultivars (dpa, days after anthesis).
24
25
Respiration is the process by which stored organic
materials (carbohydrates, proteins, fats) are broken
down into simple end products with a release of
energy. During respiration, oxygen (O2) is used and
carbon dioxide (CO2) is produced. 26
Respiration involves degradation of food reserves,
especially sugars, in order to produce chemical energy
(in the form of ATP and NADH) needed to maintain
cellular metabolic activity.

27
When glucose is used as a substrate, the equation
for respiration is:

Respiration results in loss of food reserves, loss of

flavor, especially sweetness, loss of salable dry
weight, and release of heat that increases the costs
of refrigeration, and release of CO2 thus, requiring
extensive ventilation.

28
We can determine the rate of development and
aging by measuring the rate of CO2 production.
❑Rate of deterioration α rate of respiration.
❑Faster respiration, shorter life.
❑Respiration increases
exponentially with T.

Respiratory output of CO2 can undergo dramatic

change as fruits ripen. Early research on apple and
pear led to a classic model of a postharvest
climacteric rise associated with ripening and linked
in time with ethylene production. Studies with
tropical fruits such as avocado and banana then
revealed characteristic 'waveforms' of even wider
amplitude (Biale (1950) Annu. Rev. Plant Physiol. 1,
29
183-206).
▪ The respiration rate of a product determine how
fast the chemical process occurs.
▪ The higher the rate the faster glucose will be
transformed and the faster all supplies will be
depleted.
▪ Therefore, when it
comes to storage of
fruits and vegetables
we want to keep the
respiration rates as
low as possible
without killing the
produce.
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RESPIRATION RATE OF SOME FRESH COMMODITIES

31
 FACTORS AFFECTING RESPIARTION RATE

1. Oxygen
2. Temperature
3. Plant type / organ
4. Surrounding threat

Factor 1: OXYGEN

• Low levels of O2 → reduce respiratory rate

But very low levels of O2 could accelerate
breakdown of sugars to ethanol in the cells!!!!
32
Factor 2: TEMPERATURE

Temperature directly affects the respiration rates of fresh

produce. Respiration is a complex process involving many
enzymatic reactions.
TEMPERATURE QUOTIENT OF RESPIRATION: Q10

• Q10 = rate of process at one temperature divided

by the rate at 10C lower temperature.
• Q10 for respiration is 2.0 - 2.5 at the storage
temperatures between 5 and 35C.

Respiration rate increase double for every 10 °C

increase in temperature. Temperature above 20°C
can be cause abnormal physiological phenomena33
Factor 3: PLANT TYPE/ORGAN

• Greater metabolic activity = greater respiratory rates.

• Plants can be divided into 2 groups by their
respiration.

1. Climacteric Fruit
– Sharp increase in rate immediately
before fruit ripening = climacteric rise
in respiration
– Huge increase in ethylene production

2. Non-climacteric Fruit
– No peak of respiration
– Insensitive to ethylene
34
 3

1 : PRECLIMACERIC PERIOD 3 : CLIMACTERIC PEAK PERIOD

2 : CLIMACTERIC RISE PERIOD 4 : POSTCLIMACTERIC PERIOD 35
 CLIMACTERIC FRUITS NON-CLIMACTERIC FRUITS
Apple Muskmelon Blackberry Lychee
Apricot Nectarine Cacao Okra
Avocado Papaya Carambola Olive
Banana Passion fruit Cashew apple Orange
Biriba Peach Cherry Peas
Blueberry Pear Cucumber Pepper
Breadfruit Persimmon Date Pineapple
Cherimoya Plantain Eggplant Pomegranate
Durian Plum Grape Prickly pear
Feijoa Quince Grapefruit Raspberry
Fig Rambutan Jujube Strawberry
Guava Sapodilla Lemon Summer squash
Jackfruit Sapote Lime Tamarillo
Kiwifruit Soursop Longan Tangerine/Mandarin
Mango Tomato Loquat Watermelon 36
Factor 3: PLANT TYPE/ORGAN

❑ Root tips, developing buds and meristematic

regions in general have higher respiration rates

❑ Seeds - low respiratory rates, dormant,

desiccation results in slowdown of respiration

37
 HORTICULTURAL COMMODITIES CLASSIFIED
ACCORDING TO THEIR RESPIRAION RATES
(Wilson 1995. available online http://ces.ncsu.edu/depts/hort/hil/hil-800.html)

Range at 5°C
Class Commodities
(mg CO2/kg-hr)*
Very low <5 Dates, dried fruits and vegetables, nuts
Apple, beet, celery, citrus fruits, garlic, grape,
Low 5-10 honeydew melon, kiwifruit,onion, papaya,
persimmon, pineapple, potato (mature), watermelon
Apricot, banana, blueberry, cabbage, cantaloupe,
Moderate 10-20 carrot (topped), celeriac, cherry, cucumber, fig,
lettuce (head), mango, nectarine, olive, peach, pear,
plum, potato (immature), tomato
Avocado, blackberry, carrot (with tops), cauliflower,
High 20-40 leeks, lettuce (leaf), lima bean, radish (with tops),
raspberry
Artichoke, bean sprouts, broccoli, Brussels sprouts,
Very high 40-60 cut flowers, endive, green onions, okra, snap bean,
watercress
Extremely high >60 Asparagus, mushroom, parsley, peas, spinash,
sweet corn 38
Factor 4: SURROUNDING THREAT

• Mechanical damage
- fresh cut produce induces higher respiration

• C2H4 in atmosphere

• Disease infection

Postharvest strawberry fruit decay

caused by the fungi Penicillium spp.
(circular blue and white molds) and
Botrytis cinerea (fluffy gray mold) (A),
Rhizopus stolonifer (B), Colletotrichum
spp. (C), and Mucor spp. (D).
Feliziani, E. and Romanazzi, G. ‘Postharvest Decay of Strawberry Fruit:
Etiology, Epidemiology, and Disease Management’. 1 Jan. 2016 : 47 – 39
63.
FRUIT RIPENNING IS INDUCED BY ETHYLENE

40
CLASSIFICATION OF FRUITS ACCORDING TO THEIR
MAXIMUM ETHYLENE PRODUCTION RATE

41
CHANGE IN CARBOHYDRATES
Structure of carbohydrate
❑ Cellulose is made up of hundreds of glucose sugars
joined together to form a long chain.
❑ Hemicellulose are also long chains of sugars including
different types of sugar, such as glucose, xylose,
galactose, and mannose and instead of being linear are
branched structures

42
❑ Pectin are also long branched chains of sugars;
galacturonic acid, rhamnose, galactose, and arabinose

The primary cell wall is constructed of a web of cellulose micro-fibrils,

hemicellulose polysaccharides, and the hetero-polysaccharide pectin

43
The major pectic enzymes classified based on their
type of action involved in degradation of pectins.
1. Polygalacturonase (PG): hydrolytic enzyme
playing an important role in pectin termination.
▪ Endo-PG randomly catalyzes cleavage of α-1,4-
glycosidic linkages randomly in pectin.
▪ Exo-PG releases galacturonic acid by catalyzing
the glycosidic bonds hydrolysis from the non-
reducing end of the pectic substances.

44
The major pectic enzymes classified based on their
type of action involved in degradation of pectins.
2. Pectin methyl esterase (PME): causes de-
esterification by catalyzing the pectin methyl group
hydrolysis.
3. Pectin lyase (PL): act at methyl-ester bonds
leading to beta-elimination of glycosidic bonds by
formation of double bonds in galacturonic acids.
https://www.ukessays.com/essays/sciences/role-pectic-enzymes-fruit-ripening-6817.php

45
 Firmness of mango fruit var.
“Dashehari” during different stages of
fruit ripening, measured using
penetrometer.

46
CHANGE IN CARBOHYDRATES
Non-structure of carbohydrate

▪ Conversion of starch to sugars - Not desirable in

potato but very desirable in apples, bananas.
▪ Conversion of sugars to starch - Not desirable in
sweet corn, pea but very desirable in potato. 47
 CARBOHYDRATES IN DIFFERENT
FRUITS/VEGETABLES (100g)
Fruit/Vegetable Carbohydrates
Apple 10.5 g
Banana 26 g
Avocado 2g
Guava 4.4 g
Mango 9.5 g
Pineapple 12 g
Mangosteen 5.6 g
Asparagus 2g
Carrot 8g
Cucumber 1.5g
Mushroom 0.5g
Okra 3g
Potato 15-25g
Tomatoes 3g
48
http://www.weightlossforall.com/carbohydrates-vegetables.htm
DIETARY FIBER
Dietary fiber - includes all parts of plant foods that
human body can't digest or absorb. Fiber is often
classified into two categories: those that don't
dissolve in water (insoluble fiber) and those that do
(soluble fiber).

49
INSOLUBLE FIBER
o A natural laxative and includes cellulose and
lignin which occur in whole grains (especially
wheat bran), and hemicellulose (partly soluble)
found in whole grains, nuts, seeds, fruits and
vegetables.

o This type of fiber increases stool bulk, so it can be

of benefit to those who struggle with
constipation or irregular stools.

50
SOLUBLE FIBER

o This type of fiber dissolves in water to form a gel-

like material.

o It can help lower blood cholesterol and glucose

levels.

o Types of soluble fiber include pectin, which

occurs in fruits (apples, strawberries, citrus fruits);
beta-glucans, found in konjac, oats, barley and
rye; gums, found in beans, cereals (barley, oats,
rice), seeds and seaweed.

51
52
The chemical structure of Konjac consists, mainly, in
mannose and glucose in the ratio 8:5 linked by
beta (1-->4) glycosidic bonds.

GLUCOMANNAN

Molecular weight is between

200,000-2,000,000 Daltons. 53
Inulin: a natural dietary fiber found in over 35,000
common foods like artichokes, asparagus, onions,
garlic.

beta(2→1) fructosyl-fructose 54
PROTEINS

▪ Very low levels of proteins in fresh produce (<2%).

▪ Nuts, beans 8-20%.
▪ Contains essential amino acids: leucine,
isolucine, valine, lysine, tryptophan,
phenylalanine, tyrosine and threonine.

55
56
LIPIDS

Wax or cuticle
o Fatty acid ester, mono hydroxyalcohol.
o Produce more during maturation in fruits.

Mango fruit at the ripe (A) and overripe (B) stages. Cuticles
stained with Oil Red from ripe (C) and overripe (D). 57
LIPIDS

Stored lipid
o Triglyceride, fatty acids.
o Essential fatty acids.
o Unsaturated fatty acids
- linoleic (18:2),
- linolenic acids (18:3)

58
STRUCTURE LIPID

o Phospholipid

Changes of plant lipid affecting the quality

o Membrane transition -> disorder
o Phospholipid signaling
o Lipid peroxidation
- Beta oxidation
- LOX pathway

59
Membranes are the primary site of cold-induced injury, leading to a
cascade of cellular processes with adverse effects on the plant. When
exposure to low temperature is brief, the effects may be transitory and
plants survive. However, the plant will exhibit necrosis or die if exposure
60
is
maintained.
LIPID PEROXIDATION
▪ Lipid peroxidation: the oxidative
degradation of lipids.
▪ It is the process whereby free
radicals “ steal” electrons from
the lipids in cell membranes,
resulting in cell damage.
▪ This process proceeds by a free
redical chain reaction
mechanism. It most often
affects polyunsaturated fatty
acids (PUFA).
▪ In addition, end products of
lipid peroxidation may be
mutagenic and carcinogenic.
Main molecules involved in the initiation and propagation of lipid peroxidation.
LH: lipid molecule; LOO●: lipid peroxyl radical; L●: lipid radical; LOOH: 61lipid
hydroperoxide. https://www.mdpi.com/2076-3921/9/3/231/htm
COMPOSITIONAL CHANGES
Compositional changes that occur during ripening
and continue after harvest:

plants accumulate these

molecules in their flower petals
to attract pollinators, or in their
fruits to attract mammals or
birds.

❑ Carotenoids generally make yellow, orange, and red

colors.
❑ Flavonoids can make yellow and red but also purple and
bluish colors. 62
❑ Betalains, which usually make purple colors.
Morphological changes during the lifespan of petals in intact plants. (a) Different
stages during the lifespan of corolla in intact plants (a typical observation, n=5);
(b) Chlorophyll (green color) and anthocyanin (red color) contents in buds and
petals (mean ±SEM, n=5); (c) petal movement capability (yellow color) and
fresh weight (FW) (purple color) of petals (mean ±SEM, n=5).

Azad A K et al. J. Exp. Bot. 2008;59:2085-2095

© 2008 The Author(s).
63
Morphological changes during the lifespan of petals in intact plants.
LOSS IN CHLOROPHYLL CONTENT
o Loss of chlorophyll (green color) – is desirable in
fruit but not in vegetables.
o During ripening ethylene promotes the
degradation of chlorophyll .

64
LOSS IN CHLOROPHYLL CONTENT
o Replacement of MG atom in the chlorophyll by
hydrogen atom under acidic condition with the
formation of phaeophytin.
Phaeophytin formation: a color change from bright
green to dull olive green

65
LOSS IN CHLOROPHYLL CONTENT
o Hydrolysis of chlorophyll to chlorophyllide and
phytol
- Catalyzed by chlorophyllase
- Followed by a replacement of Mg atom with
hydrogen
- Resulting in the formation of pheophorbide

COLORLESS PRODUCT
66
LOSS IN CHLOROPHYLL CONTENT
o Bleaching of chlorophyll by the action of
lipoxygenase, peroxidase and catalase.
- Degradation of fatty acid hydro peroxidase.
- Oxidation-reduction reaction in the presence of
oxygen.

67
DEVELOPMENT OF CAROTENOIDS
o Development of carotenoids (yellow and orange
colors) - desirable in various fruits, such as
apricots, peaches, citrus, tomatoes, etc.
o Carotenoids isoprene compounds compose of
isoprene joined head to tail to form a system of
conjugated double bonds.
- Carotenes
- Lycopene
- Xanthophylls

68
SYNTHESIS OF XANTHOPHYLL
o Occurs in banana.
o After treating unripened bananas (green) with
ethylene, they become ripened (yellow) due to
the synthesis of xanthophyll.

69
DEVELOPMENT OF ANTHOCYANINS
o Development of anthocyanins (red and blue
colors) - desirable in various fruits, such as
cherries, strawberries, etc.

o Anthocyanin are flavonoid pigments whose

structure is based on the phenyl propanoid
molecules and is water-soluble pigments formed
during ripening process.

70
SYNTHESIS OF SUGAR AND ORGANIC ACID
o Typical taste of fruits is determined by the
content of sugars and organic acids.
o Also phenolic compounds and tannins may
affect the taste.
o During ripening starch is converted into sucrose,
glucose and fructose.
o Starch hydrolysis is a major change during
ripening of climacteric fruits.

71
SYNTHESIS OF SUGAR AND ORGANIC ACID
o The concentration of organic acids also reach to
a maximum during growth and development of
fruit on tree.
o There is a decrease during storage highly
depending on temperature.
o Citric and malic acids from Krebs cycle.
o Other includes ascorbic acid and oxalic acid.

72
SYNTHESIS OF VOLATILE COMPOUNDS
o Aroma is produced by volatiles synthesized during
ripening including aldehydes, esters, lactones,
terpenes, and sulfur compounds.
o Volatiles originate from proteins, carbohydrates,
lipids, and vitamins.
o Taste is provided by many nonvolatile components
including sugars and acids present in fruits.

73
74
WATER LOSS
Water loss is a main cause of deterioration because
it results in:
▪ Direct loss of salable weight
▪ Loss in appearance (wilting and shriveling)
▪ Loss of textural quality (softening, crispness)Small
fruit have large surface-to-volume ratios, and
especially suffer from water loss!

75
76
TRANSPIRATION LOSSES FOR FRUITS & VEGETABLES
STORED AT VARIOUS RELATIVE HUMIDITIES

file:///F:/Postharvest%2
0Technology/organic-
stored-fruit-veg-
NYSIPM.pdf 77
WAYS TO REDUCE WATER LOSS AFTER HARVEST:
o Low temperatures
o High RH
o Prevent surface injuries
o Application of waxes or other coatings
o Wrapping with plastic films

78
PHYSICAL DAMAGE
o Physical damage (surface injuries, bruising,
vibration damage) is a main contributor to
deterioration.
o The damaged areas become brown (because
leakage of phenolic compounds), accelerate
water loss, stimulate ethylene production, and
provide sites for pathogen invasion.

79
80
PHYSIOLOGICAL BREAKDOWN
Physiological disorders may develop following
storage under undesirable conditions or as a result
of improper preharvest management leading to
‘weak’ fruit with nutritional imbalances.

Physiological disorders caused by improper storage

conditions:
o Chilling injuries
o Freezing injuries
o Heat damage
o Low humidity
o Low O2 injuries
o High CO2 injuries
81
Symptoms of chilling injury in cantaloupe melons. A) Initial pink
discoloration of injured epidermal areas; B) Progressive darkening to red
of injured areas; C) Final brown to black color of injured areas; D)
Appearance of slight depressions on epidermis; E) Large depressions at
the end of the holding period; F) Lattice of depressions following outline
of discolored areas; G) Small, sharply defined depressions or pitting of
the epidermis; H) Whitening of the corky net on discolored areas; I)
Superficial nature of injuries: damaged epidermis above the intact 82
mesocarp.
83
PATHOLOGICAL BREAKDOWN
❑ One of the most obvious symptoms of
deterioration is growth of pathogens.

❑ Healthy fruit are mostly resistant to pathogens,

but senesced and damaged fruit become
susceptible to infection.

❑ Infection by pathogens became a very serious

problem in postharvest handling in recent years,
since health authorities consistently reduced the
permitted residue limits (MRL’s) for chemical
fungicides.

84
(A) Bitter Rot,
(B) Blue mold postharvest decay of
apple caused by the fungus
Colletotrichum spp. and
Penicillium expansum
respectively
(C) Decay of nectarine fruitcaused
by P. expansum
(D) Brown Rot of peach caused by
Monilinia fructicola
(E) Black Mold on garlic caused by
Aspergillus niger
(F) Green mold caused by P.
digitatum on citrus fruits
(G) Anthracnose of pepper fruit
caused by Colletotrichum sp.
(H) Decay of table grapes caused
by Rhizopus stolonifer and
Aspergillus niger

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THANK YOU FOR YOUR ATTENTION