1. Principles of Postharvest Physiology
Dr. Ron Porat
Dept. of Postharvest Science of Fresh Produce
ARO, The Volcani Center, Bet Dagan, Israel

2. It is estimated that about one-third of the fresh produce harvested worldwide is lost at some point between harvest and consumption.

3. According to the UN Food and Agriculture Organization (FAO), the annual world production of fruits and vegetables is about ~ 1,500,000,000 tons!
That means an average loss of ~ 500,000, 000 tons of produce each year!

4. The goal of this lecture is to provide the basic biological information needed to understand why fruits, vegetables and cut flowers deteriorate after harvest, and then to learn how to apply appropriate postharvest operation techniques in order to maintain quality and reduce losses.

5. First, a few basic principles:
1. 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.

6. 2. 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!

7. 3. Fresh horticultural crops are diverse in morphological structure (roots, stems, leaves, flowers, fruit, etc.), in composition, and general physiology.
Therefore, optimal postharvest requirements vary among commodities!

8. Biological factors involved in deterioration
 Respiration
Ethylene production
Compositional changes
Water loss
Physical damage
Physiological breakdown
Pathological breakdown

9. Respiration
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.

10. 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.

11. When glucose is used as a substrate, the equation for respiration is:
C6H12O6 + 6 O2 → 6 CO2 + 6 H2O kcal heat
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.

12. In general, the rate of deterioration of harvested commodities is proportional to their respiration rate:
Commodities with high respiration rates will have short potential storage lives.
Commodities with low respiration rates will have long potential storage lives.

13. Horticultural products classified according to their respiration rates

14. According to their respiration behavior during maturation and ripening, fruits are classified to either climacteric or non-climacteric.
Climacteric fruits – show a large increase in respiration and ethylene production during ripening.
Non-climacteric fruits – show no change in their generally low respiration and ethylene production rates during ripening.

15. Respiratory phases of climacteric and non-climacteric fruit

16. Fruits classified according to respiratory behavior

17. Magnitudes of respiration of climacteric fruit

18. Climacteric respiration during ripening of tomato and banana fruit

19. At last, respiration rates (and deterioration) varies among different types of commodities:
Storage organs (nuts, tubers) – have low respiration rates.
Meristem tissues (asparagus, broccoli) – have very high respiration rates.
Commodities harvested during active growth (spinach, green onions, bean sprouts) – have high respiration rates.
Mature fruit (citrus, grape) – have low respiration rates.

20. Ethylene
Ethylene (C2H4) is a simple gaseous organic molecule.
As a plant hormone, it is involved in regulation of growth, ripening, senescence and abscission processes.
Ethylene is naturally synthesized in plants, but also evolves from engines and fires. It is biologically active at very low concentrations in the range of ppm levels.

21. Ethylene biosynthesis pathway
MET
(methionine)
SAM synthase
Methylation
Polyamine synthesis
SAM
(S-adenosyl-methionine)
ACC synthase
ACC
(1-aminocyclopropane-1-carboxylic acid)
ACC oxidase
Ethylene
ACS and ACO are the key regulatory enzymes in ethylene biosynthesis

22. Autocatalytic production of ethylene
In climacteric fruit, the natural increase in ethylene production during ripening stimulates its own synthesis – a phenomenon called “autocatalytic production of ethylene”.

23. In some climacteric fruits, such as avocado, banana, melons, pears and tomatoes, the autocatalytic positive feedback may increase ethylene production rates by 1,000 x fold during ripening.
Exposure to external ethylene also enhances autocatalytic production of ethylene and ripening!

24. Regulation of ethylene biosynthesis
Generally, ethylene production increases:
 with maturity at harvest
after physical damage (dropping, wounding)
in decayed fruit
at increasing temperatures up to 30C
during stresses (water stress, chilling, etc.)
On the other hand, ethylene is reduced by:
low storage temperatures
reduced O2 levels (below 8%)
elevated CO2 levels (above 2%)

25. Atmospheric ethylene
Beside its own biosynthesis, harvested produce may be exposed to atmospheric ethylene.
Sources for atmospheric ethylene include exhaust from trucks and forklifts, pollution from industrial activity and burning of fuels, fires, and from nearby ripening climacteric fruit.

26. Beneficial uses of ethylene in postharvest
Commercial postharvest applications of ethylene are used to:
Promote full and uniform ripening of bananas, avocado, mango and green-harvested tomatoes.
Promote color change (degreening) in citrus fruit.

27. Optimal conditions to promote ripening
Considerations to achieve uniform ripening:
 Temperature between 18-25C
RH about 90-95%
Ethylene concentration between ppm
Duration of treatment between 1-5 days
Adequate air circulation in ripening room
Ventilation and air exchange to prevent excess accumulation of CO2

28. Ripe and “ready to eat” mango fruit

29. Sources of ethylene
Ethylene could be applied to ripening rooms from:
 Industrial gas cylinders (including explosive-proof ethylene mixtures in inert gases without presence of oxygen)
Ethylene generators (produce ethylene from dehydration of ethanol by heating)
C2H5OH – H20  C2H4
Ethylene releasing agents (‘Ethepon’)
Use of ripen fruit

30. * It should be cautious that mixtures of ethylene gas in air may be explosive at concentrations of %!

31. Undesirable effects of ethylene
As the main ‘ripening’ and ‘senescence’ hormone, ethylene has detrimental effects on postharvest storage and quality.
Accelerates leaf senescence.
Accelerates ripening and fruit softening.
Accelerates flower wilting
Accelerates abscission
Stimulates sprouting of tomato
Stimulates lignification and toughening of asparagus
Causes leaf and peel disorders

32. Leaf yellowing and senescence

33. Yellowing and change in texture

34. Flower wilting

35. Peel disorders caused by ethylene

36. Preventing the deteriorative effects of ethylene
Measures to reduce ethylene effects:
1. Eliminate the sources of ethylene:
● Do not hold stocks of ripen fruit near ethylene sensitive commodities
● Use electric powered forklifts
● The truck loading area should be isolated from handling and storage areas.
● Remove decayed pellets of fruit
● Do not smoke or burn fires in the packinghouse area

37. 2. Ventilation: harvested produce synthesize ethylene
2. Ventilation: harvested produce synthesize ethylene. Therefore, simple ventilation (using an intake fan and a passive exhaust) can effectively remove excess ethylene.
3. Ethylene absorbers: ethylene may be absorbed from storage rooms by using chemical scavengers, such as potassium permanganate and charcoal.
4. Inhibition of ethylene action: use of 1-MCP.
1-MCP was developed 10 years ago, and is a very effective inhibitor of ethylene. It is now being licensed for use in fruit and vegetables.

38. Commercial products of 1-MCP
EthyBlock® – for use with ornamentals
SmartFresh® – for use with fruit and vegetables

39. Compositional changes
Compositional changes that occur during ripening and continue after harvest:
● Changes in pigments:
Loss of chlorophyll (green color) – is desirable in fruit but not in vegetables.
Development of carotenoids (yellow and orange colors) – desirable in various fruits, such as apricots, peaches, citrus, tomatoes, etc.
Development of antocyanins (red and blue colors) - desirable in various fruits, such as cherries, strawberries, etc.

40. Loss of chlorophyll after harvest

41. ● Changes in metabolites:
Loss of sugars
Loss of acids
Loss of amino acids
Loss of lipids
Loss of vitamin content

42. Metabolic changes during ripening of tomato

43. Metabolic changes in broccoli after harvest
Chlorophyll
Carotenoids
Vitamin C
Sugars

44. 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!

45. Water loss and shriveling
Apple
Peach

46. Ways to reduce water loss after harvest:
Low temperatures
High RH
Prevent surface injuries
Application of waxes or other coatings
Wrapping with plastic films

47. Reduction of water loss and shrinkage using plastic liners

48. Effects of wax and seal packaging on weight loss of mandarins

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

50. Bruising damage in apple, nectarine and cherry

51. Bruising and skin abrasion in guava and banana

52. 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.

53. Physiological disorders caused by improper storage conditions:
- Chilling injuries
- Freezing injuries
- Heat damage
- Low humidity
- Low O2 injuries
- High CO2 injuries

54. Chilling injuries
Tropical and subtropical fruits are sensitive to low non-freezing temperatures below 15C.
The most common symptoms of chilling injuries include surface and internal browning, pitting, water soaked areas, uneven ripening or failure to ripen, development of off flavors, and increased decay incidence.

55. Cold storage chart Chilling sensitive commodities are stored at 8-13C
Non-chilling sensitive commodities are stored at 0-3C
(Postharvest Technology, 2002)

56. List of fruit and vegetables tolerant or sensitive to chilling
(chilling tolerant)
(chilling sensitive)

57. Chilling injuries in citrus and avocado

58. Chilling injuries in yam, peach and mango

59. Chilling injuries in banana

60. Chilling injuries in pineapple and guava

61. Freezing injuries
Storing produce below their freezing point causes immediate collapse of the tissue and complete loss.

62. Heat injury
Caused by exposure to direct sun light or excessively high temperatures. The symptoms include bleaching, surface burning, uneven ripening and softening.

63. Low humidity (desiccation)
Storage under low humidity conditions may enhance disorders related to desiccation and senescence.

64. Nutritional imbalances
Certain disorders may develop after harvest because of nutritional deficiencies. For example, bitter pit and water core symptoms in apples are related to deficiency in calcium.
Bitter pit
Water core

65. 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.

66. P. expansum
Rhyzopus
Botrytis

67. Colletotrichum
Phytopthora
Alternaria
P. digitatum

68. Summary of postharvest physiology changes
 Respiration
Ethylene production
Compositional changes
Water loss
Physical damage
Physiological breakdown
Pathological breakdown