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This is to certify that the

thesis entitled

THE EFFECT OF APPLE CULTIVAR, HARVEST MATURITY AND STORAGE
DURATION ON THE CARBON DIOXIDE EVOLUTION RESPONSE OF
FRUIT DAMAGED BY BRUISING

presented by

MICHAEL LEE PARKER

has been accepted towards fulfillment
of the requirements for '

 

 

 

M.S. degree in HORTICULTURE
is» MAW
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TIIE EFFECT 0!" APPLE CULTIVAR, HARVEST NAN/Rm MD STORAGE DURATION

ON T!!! CW! DIOXIDE EVOUITIDI RESPONSE OF

FRUIT DAMAGED DY BRUISING

By

Ali chael tee Parker-

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIENCE

Department of Horticulture

1985

ABSTRACT

THE EFFECT OF ARPLE GULTIVAR, HARVEST UMVURITY AMD STORAGE DURATION
OW TUE CARROU DIOXIDE EVOLUTION RESPONSE OF
FRUIT DANMGED DY BRUISING

Br

Ilicbael Lee Parker

Carbon dioxide evolution was examined as a possible measure for
determining relatively small quantities of bruise damage to apples.
Experiments were conducted to determine tbe effect of apple cultivar,
barvest maturity and storage duration on the carbon dioxide damage
response of the fruit to varying degrees of damage.

Application of impact bruise damage ranging from sligbt to severe
to ’Racspur’, ’Empire’ and ’Rome Deauty’ apple fruit resulted in highly
significant increases in carbon dioxide evolution over non-damaged
fruit. Tbis carbon dioxide response significantly increased with
increased degrees of damage; however, it was not affected by other
factors observed. The damage treatments also resulted in bigbly
significant increases in the percentage of total fruit tissue damage as
assessed by visual means. Tbe amounts were significantly affected by

cultivar, time of harvest and storage duration.

ACKNOWLEDGEMENTS

I would like to express my gratitude to my major professor,
Dr. Donald H. Dewey, for giving me the opportunity to continue
my education, for his instruction and guidance through my research,
and for his help in reviewing my thesis. I would also like to
thank my guidance committee, Dr. J. Flore, Dr. A. Cameron and
Dr. J. Lee, for their participation in my graduate work and research
and for their helpful reviewing of my thesis.

The help of James Brown and Carol (Zurawski) Westerhof was
very much appreciated in setting up the research and in data
collection.

And most of all I would like to express my gratitude to God
and my family and close friends. For the love, patience, wisdom
and endurance I received I will always be grateful.

This research was supported by a grant from the United States-

Israel Binational Agricultural and Deve10pment Fund (BARD).

ii

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LIST OF TABLES............................................... iv
LIST OF FIGURES................................... ........... v
INTRODUCTION............ .......... ....... .................... I
LITERATURE REVIEW................................. ........... 3
MATERIALS AND HETHDDS........................................ 9
RESULTS...................................................... 15
DISCUSSION................................... ................ 55
SUMMARY AND CONCLUSIONS.. ............ .... .................... 61

LITERATURE CITED... .......................................... 62

iii

 

TABLE PAGE

 

l.The internal ethylene, flesh firmness and starch rating
of lo apples at one day following harvest (0-day storage

duration)0......0.00......OOOOOOOOOOOOOOOOOOO0.0000IOOOOOOOO l6

2.The internal ethylene, flesh firmness and starch rating
of lo apples at lOO days following harvest (loo-day storage

dur.ti°n)0.00I..0.0...OOOOOOOOOOOOOOOOOOOOOOOOOOOOOIOOOOOOOO l7

3.The internal ethylene, flesh firmness and starch rating
of IO apples at l99 days following harvest (199-day storage
duration)0.0.0....00......OOOOOOOOOOOOOOOOOOOOOOOO0.0.0.0... 18

h.Analysis of variance mean square values for rate of carbon
dioxide evolution of apples determined at one to six hours
post-treatmentOOOO00.0.0...O00......OOOOOOOOOOOOOOO...COO... ‘9

5.Analysis of variance mean square values for rate of carbon
evolution of apples determined at one to four hours post-
treatment rates minus pre-treatment rates. The data for
'Hacspur' at the 0-day storage duration are not included.... 20

6.The rate of CO evolution of apples at the fifth
hour following damage treatment............................. 2h

7.Analysis of variance mean square values for percent of
bruised tissue based on the total weight of the fruit.
The control treatment data (zero values) were excluded
in the analysis............................................. 25

8.Nean percentages of excised bruised tissue according to
damage treatmntOO00......OOOOOOOOOOOO0.00000000000000000000 26

9.Correlation values between carbon dioxide evolution rates
and percentage of bruised tissue at different hours of
observation. Data for all cultivars, harvests and storage
durations were included along with several individual
interactions.................................................27

iv

LIST 9; FIGURES

FIGURE PAGE

 

l.Pendulum apparatus constructed for application of force to
produce bruising to apple fruit. A fruit is placed on the
platform and the pendulum released to strike the fruit,
which is deflected into the padded catching container.......... ll

2.The effect of damage to apples of all cultivars, storage
durations and harvests on the rate of CD evolution
measured at one to six hours following saplication of the

du.gel.0.0.0.0000...OOOOOOOOOOOCOOOOOOOOOCOOOOOOOOOOO'...O...O. 22

3.The effect of damage treatment and storage duration on the
percent of bruised tissue for three apple cultivars............ 29

h.The effect of cultivar and storage duration on the percent
of bruised tissue for the levels of damage applied to apples... 3l

5.The effect of damage treatment and apple cultivar on the
percent of bruised tissue for three storage durations.......... 33

6.The effect of time of harvest of apples for all cultivars,
storage durations and damage treatments on the rate of
CD evolution at one to six hours following the
apalication of the damage...................................... 37

7.The effect of storage duration of apples for all cultivars,
damage treatments and harvests on the rate of CO
evolution measured at one to six hours following application
of the damage.................................................. ho

8.The effect of cultivar of apples for all damage treatments,
storage durations and harvests on the rate of CO2
evolution measured at one to six hours following the
application of damage.......................................... h2

9.The effect of apple cultivar and time of harvest on the rate
of C0 evolution measured at the fifth hour following
the dgmage treatment of apples at three storage durations...... Ah

l0. The effect of time of harvest and storage duration on the
rate of CD evolution measured at the fifth hour
following Ehe damage treatment of apple cultivars............. £6

ll. The effect of cultivar and storage duration on the rate
of CO evolution measured at the fifth hour following
the d mage treatment for time of harvest....................... h8

INTRODUCTION

Damage to produce during handling, transportation and marketing
results in an estimated 9-l7 percent loss of produce at a value of
l.O-2.0 billion dollars annually (32). Damage to apples destined
for fresh market apples approximates a loss of l3.h million dollars
with up to a 5 percent less in fruit volume (32). Unpublished data
of Schoorl (37) shows a l,OOO-mile truck transport with six handling
operations to bruise lO-Zh percent of the apples. It has been shown
that apples destined for processing lose approximately 2.8 percent
total weight due to the bruising that occurs through all operations
after harvest (#6). Mechanical injury during handling is the major
cause of produce damage (l0,37) which results in bruising for apples
(:2). ' A

Bruising occurs in all operations and has an accumulative
effect on a commodity (23), increasing the amount of damage at each
operation. Citrus fruit in Japan became more severely bruised as it
passed through sorting equipment along a packinghouse line (3).
Similarily. apples moving through marketing channels in New York
City resulted in progressively increased bruising (l0). The
accumulation of bruising adversely affects the quality of fruit and
even slight. barely visible bruising may result in considerable
reduction in quality as the fruit is much more susceptible to decay
as with sweet cherries (26). Improving the protective quality of
the packages in which the product is marketed can reduce damage. A

practical method for determining and comparing the protective

2
quality of packages would be useful in selecting a superior package.
A suitable method to detect bruising must be fast. accurate and
objective.

Research with a wide range of fruit has indicated that an
increase in carbon dioxide evolution of fruit following damage
offers passibilities as a damage indicator. Carbon dioxide is a
natural product evolved from the fruit and is easily and accurately
measured with current technology (l8.22).

The objective of this research was to determine the reliability
of carbon dioxide as a damage indicator for relatively small
quantities of bruising and to determine how this damage response is

affected by cultivar. fruit maturity and storage duration.

LITERATURE REVIEW

Bruising is a term applied to injury, distortion or bursting of
cells resulting from a physical force applied to fruit tissues. In
apples, browning of the tissue is a consequence of cell sap
oxidation (ll,27.h0) which is mediated by concentrations of
chlorogenic acids and flavanol in the fruit tissue (l6). The amount
of bruised apple tissue is highly correlated to the energy absorbed
by the tissue during the applied force (ll). Physical injury to
apples by slowly applied compression forces results in more bruise
damage to the tissue than a rapid impact of the same energy level;
for a given quantity of bruising. an impact force must be twice as
great in order to yield the same amount of damaged tissue as a
compression force (11,27).

Much fruit bruising could be eliminated or greatly reduced by
improving the protective qualities of the packages in which fruit
are handled and marketed. ‘ Klein (l8) has proposed that apple
package performance could be based on carbon dioxide evolution of
the fruit as a damage indicator. There are many factors which may
affect bruising and the physiological response of the fruit to
bruising that need to be understood before this method can be
developed. particularily fruit cultivar and maturity and the length
of time the fruit has been stored (27,38,h0,h6). Apple fruit
maturity significantly affects the degree of bruising since fruit
maturation results in softening of tissue and greater susceptibility
to damage (£6). Apple tissue becomes less elastic so that the
energy absorption through temporary cell deformation is decreased
and the cells absorb less energy before becoming permanently damaged

3

(27). It has been demonstrated that bruise diameter and depth
increase with advancing maturity of the fruit (lA,38,A6). The same
is likely true for cherries since the most mature fruit yielded
higher weight losses than less mature fruit following physical
injury (26). Tomato fruit at the breaker stage of maturity with red
color initially evident were found to be four times more readily
bruised than fruit at the mature green stage and eight times more
readily bruised than at the immature stage (23). Also, the degree
of bruising in peaches is related to maturity. with the most mature
fruit receiving the most damage (A3).

The extent of bruising and the degree and rate of browning of
the damaged tissue varies by cultivar of apple (lA,l6.A0.Al,A6).
Schoorl and Holt (A0) found 'Jonathan', 'Delicious' and 'Granny
Smith' apples after 5 months of storage to have highly significant
differences in the volume of bruised tissue resulting from a
standard applied force. At this time 'Jonathan' was the most
severly bruised followed by 'Delicious' and 'Granny Smith'.
respectively. Hennergren and Lee (A6) reported varietal differences
in the bruising of 'Golden Delicious', 'York' and 'Stayman' fruit.
Bruising, measured as percent of bruised flesh showed 'York' fruit
to be the most resistant to bruising followed by Golden Delicious'
and then 'Stayman'. Hyde and lngle (IA) report the order of
increased resistance of apples to bruising as 'Stayman', 'Hclntosh',
'Delicious', 'Rome', 'Golden Delicious' and 'Jonathan'.

Duration of the storage period also affects the susceptibility
of fruit to bruise damage. Schoorl and Holt (A0) reported that the

longer the fruit are in storage prior to bruising, the greater the

amount of bruised tissue results when a given force is applied.
Fruit ripening and deterioration increase with aging and is
accompanied by decreases in flesh firmness during the storage period
(2). 0n the other hand Hyde and lngle (lA) showed no increase in
the degree of bruising for 6 cultivars of apples stored at 2°C for a
maximum of nine weeks. but rather a decrease in bruising for fruit
stored for longer storage periods.

Visual assessment of mechanical damage to fruit to determine
the extent and source of injury in the handling chain is time
consuming and generally impractical for evaluating large quantities
of fruit. Various methods for detection of the actual occurance of
damage have been considered. An artificial potato containing
sensors and transmitters to measure and record impacts and other
damaging forces was developed for analyzing handling systems (3l).
Another type of device was used for 'Red Delicious' and 'Nclntosh'
apples (7), whereby computer image analysis of bruising was made as
the fruit passed a detector in a packinghouse line. The extent of
damage in apples can also be measured by decreases in the electrical
resistance of fruit tissue when cell sap is released upon rupture of
the cell walls and membranes (8).

A major criterion for predicting the amount of fruit damage
that may occur in a package is the amount of energy absorbed by the
package and its contents (ll.l3,37,38). Holt and Schoorl (ll).
found a high correlation between the quantity of bruised tissue and
the amount of energy absorbed by the apple. One method predicts the
percentage of bruised fruit in various packages for both apples and

pears (38). The percentage of fruit damage in packages is predicted

by an equation based on a relationship of fruit cultivar, drop
height and number of drops (37,38). Prediction of where the most
fruit damage is likely to occur in multilayered packages (39) and
the types of packages that should provide the most protection for
the fruit have also been reported (37). Holt et. al.(l3,Al)
recently proposed the analysis of packaging and handling systems as
determined by the energy absorbed by a package which could be used
for predicting the amount of bruising the fruit receives. This
method relates the bruise volume to the energy absorbed by the
tissue, considering bruise resistance of the fruit, the equation of
the motion of the drop surface and the mass and rebound height of
the package (Al).

Various methods have been used to bruise fruit for experimental
purposes. Robitaille and Janick (3A) dropped a l90 gram stainless
Isteel weight, through a guidance tube unto an apple resting in a
cork ring. Schoorl and Holt (ll,A0) bruised apple halves firmly
fixed in place by use of a specially designed device which used a
spring loaded projectile to strike the fruit. Hyde and Ingle (lA)
used a free swinging weight, much like a pendulum, to bruise fruit.
Greenham (8) used a thumb tip to apply pressure bruises to apples.
Other methods employed are drops of varying heights and numbers,
lnstron type equipment and various devices to damage fruit for
- experimental purposes. No standard or generally accepted method has
been used.

The physiological responses of fruit to physical injury has
been suggested by many as a possible means for quantifying damage

for many fruit. Cutting of cantaloupe doubled the carbon

dioxide(C02) evolution with a ten fold increase in ethylene(C2Hh)
evolution (2A). Avocados increased in respiration, as measured by
C02 evolution and oxygen(02) uptake upon shaking (l). The dropping
of cranberries caused an increase in C02 evolution with a greater
number of drops yielding higher increases in CO2 evolution (22).
The bruising of tart cherries produced an increased CO2 evolution
(33). Vibration forces applied to sweet cherries yielded an
increase in CD2 evolution, but variable results in CZHA evolution
(26). Tomatoes exposed to vibration, impacts and dropping had
increased rates of CD2 evolution (20,2l,29) and tomatoes damaged by
cutting or dropping had increases in CZHA evolution (20,25). Citrus

fruit yield a marked increase in C0 evolution following physical

2
injury. Damage as a result of dropping, rolling, compression and
vibration increased the CD2 evolution of the fruit with greater
levels of damage (3.6.l5,i7,AA,A5). Ethylene increases in citrus
were variable, with 'Satsuma' mandarin exhibiting no increase in
evolution of CZHA after dropping (15), whereas 'Harsh' grapefruit
showed intermediate increases in CZHA as a result of dropping (A5).
Injury to apples at the preclimacteric stage of fruit
development, prior to the production of endogenous CZHA’ caused an
increase in CZHA evolution, whereas damage to fruit at a later stage
of maturation in which endogenous CZHA was being produced resulted
in a decrease in CZHA evolution by the fruit (l8,19,3A). This
variability would make the use of CZHA as an indicator of damage in
apples dependent upon a knowledge of the specific stage of maturity

of the fruit. Studies involving numerous cultivars of apple

indicate consistent increased CO2 output responses as a result of

dropping (7,l8,A2). Apples in which the tissue is cut also
exhibited higher levels of CD2 evolution (9,30). The extent of the
CD2 response in apples is not the same for all cultivars, for

example 'winter Banana' apples exhibited an increased C0 evolution

2
following damage from dropping, whereas 'Rome Beauty' apples had no

significant increase in CO2 evolution due to damage (A2).

MATERIALS A_N_l_l_ METHODS

Three cultivars of apples were selected to provide a range in
fruit bruising characteristics. 'Macspur', a strain of 'Mclntosh',
an early fall harvested apple with flesh that softens fairly rapidly
after harvest and fruit with a normal storage life expectancy of
three to six months, depending on the method of storage. 'Empire'
was selected as representative of a cultivar harvested midseason
which retains good flesh firmness for four to seven months in cold
storage. A late season harvested apple which retains its flesh
firmness during long term storage. up to nine months, was
represented by 'Rome Beauty'('Rome'). The 'Macspur' fruit was
harvested from trees with M7 rootstocks. 'Empire' from trees on M26
with an interstem. and the 'Rome' from trees with either MMlll or M2
rootstocks. 'Macspur' and 'Empire' were harvested from trees
approximately 9 years old and the 'Rome' were harvested from trees
approximately 20 years old. All were grown in experimental plots at
the Michigan State University Graham Station.

Fruit of each cultivar were harvested at three harvest
maturities. one week before the estimated optimum harvest date for
long-term storage, at optimum and one week after optimum for each
cultivar for each of the three storage durations. The optimum

harvest date was based on predicted harvest dates recommended by

Michigan State University for the cultivars of 'Mclntosh',
'Jonathan' and 'Delicious' calculated from bloom data and
temperature conditions following bloom for 30 days. internal

ethylene production of the individual fruit and the accumulation of

ethylene from ten fruit sealed in a container as developed by

IO

Dilley (5).

For each cultivar at each harvest date approximately 500 fruit
of uniform size were carefully hand harvested and placed in foam
tray packs. Trays were randomly placed in corrugated tray pack
boxes to minimize tree and handling variation. The boxes were lined
with a polyethylene film to prevent excessive moisture loss, yet
loosely closed to provide sufficient aeration to allow normal
respiration during storage. The fruit were stored in air at
approximately l°C for lOO or l99 days after harvest. The fruit for
the 0-day storage treatment were not refrigerated.

impact damage to the fruit was applied using the pendulum shown
in Fig. l. The arm of the pendulum was constructed with l/2-inch
steel rod and the striking surface of the pendulum constructed with
a piece of 3-inch angle iron covered with teflon sheeting to prevent
abrasion of the cuticle and epidermis of the fruit. The pendulum
was designed to strike the fruit at the base of the pendulum arc
before reaching the pendulum stop. The fruit was transferred by the
force of the pendulum into a padded catching container to prevent
further damage to the fruit. The pendulum provided a standard force
that could be delivered to any selected location on the fruit, for
this experiment on the cheek of the apple. Since the amount of
tissue bruised by impact varies with different tissues of the apple
(A), it was important to place the bruise in approximately the same
location each time. The pendulum also offered a means to reproduce
an impact force and to vary the impact force in the increments
desired. With the pendulum, a bruise was applied to one side of the

fruit without damaging any other part of the fruit. Five damage

 

Figure l. Pendulum apparatus constructed for application of force to
produce bruising to apple fruit. A fruit is placed on the platform
and the pendulum released to strike the fruit, which is deflected into

the padded catching container.

'A
‘ v

«I

I...

l2

per fruit, 2 small bruises per fruit on opposite sides, A small
bruises per fruit equally spaced around the equator of the fruit and
1 large bruise approximately equivalent to 3 small bruises in
respect to quantity of damaged tissue. The small bruises were
applied with the pendulum swinging through an arc of AO.5O with an
arm radius of 52 cm which produced a bruise approximately equivalent
to dropping the fruit l5 cm onto a hard surface. The one large
bruise was applied with the pendulum swinging through an arc of
90° at an arm radius of 52 cm which yielded a bruise approximately
equivalent to a drop of 50 cm.

A factorial experimental design consisting of A factors
completely randomized was selected with the factors analysed being
cultivar, harvest date, storage period and damage level. An
experimental run was made at each of the 3 storage periods using 3
cultivars, 3 harvest dates and 5 damage levels. There were 5
replications of each damage level.

Approximately lOO fruit were removed from cold storage the day
before the start of each experimental run except for the 0-day
storage period which was made the day after harvest. This time
period allowed the fruit to equilibrate to the laboratory
temperature of approximately 2l°C to ensure uniform bruising results
(27.36). Ten fruit were randomly selected for measurement of
internal ethylene. flesh firmness. and relative starch content using
the procedure outlined by Saltveit (35).

Three fruit, free of visible defects, were randomly placed in

each of 25 plastic airtight freezer containers of 2.5 liter

13

capacity. Each fruit sample was weighed in the containers and (the
containers were randomly assigned to treatments the day before
starting an experiment. Following the bruising treatment the
containers were closed with snap-on airtight lids fitted with serum
caps to facilitate sampling of the internal atmosphere with
syringes. The headspace atmosphere of the containers was analyzed
for CO2 content after closure at hourly intervals up to six hours.
The procedure was modified after the initial run of 'Macspur' at the
O-day storage period in order to obtain greater accuracy in
measuring the CD2 response. Instead of immediately treating the
fruit at the start of the experimental run, a pretreatment analysis

of the C0 evolution of each sample of fruit was made hourly for

2

four hours to provide a base level of CO evolution for each sample.

2
The containers were then opened and completely aerated prior to the
damage treatment of the fruit. After all treatments were applied
the containers were resealed and the atmosphere of the containers
analyzed hourly as in the original method.

The C0 content of the container atmosphere was determined by

2
removing a l ml sample with a syringe for injection into a Carle
8700 (BOA-B) gas chromatograph with silica gel molecular sieve
columns in parallel with a differential thermal conductivity
detector. using helium as the carrier gas. The gas chromatograph
output was recorded with a Hewlett-Packard 3390A integrator which
was calibrated to give readings as percentage of C02.
After the 6-hour reading the sealed containers remained sealed

on the laboratory bench for approximately 2A hours to allow the

brown discoloration of the bruised tissue to occur (l6,Al). Upon

lA

opening, the browned tissue and skin was excised from each apple and
weighed. This was done by cutting the skin around the perimeter of
the visible bruise on the surface of the fruit. Then all of the
brown tissue and skin was removed by using a small scoop to remove
only the brown tissue.

The CD2 evolution data was calculated in units of ml/kg hr,
taking into account fruit weight, headspace of the containers and
the length of time the containers were sealed. The data were
analyzed for CO2 evolution for each hour and for the net response as
determined by subtracting the posttreatment CD2 hourly readings of
each sample from the pretreatment readings where the latter data
were obtained. The weights of bruised tissue were expressed as
percent of total fruit weight for evaluation by analysis of
variance. A correlation of C02 evolution and percent of fruit
weight bruised for the entire experiment was determined for the
fourth and fifth hour post-treatment. Correlations were also made
with modifications in the data for CO2 evolution and percent of
fruit weight bruised. A correlation of the entire experiment,
omitting the data for the control, to eliminate zeros in the data.
was determined for the data at the fourth and fifth hour
post-treatment. The correlation between the increase in C02 from
pre-treatment over post-treatment CO2 mean percent bruised tissue
weight per treatment was determined. Several correlations between

CO2 and percent of bruised tissue for individual cultivars,

maturities and storage durations were determined also.

RESULTS

The physical and physiological characteristics of the fruit at
the beginning of each experimental run are presented in Tables l,2
and 3. Fruit internal ethylene values increased with the later
harvest dates for each cultivar, internal ethylene values also
increased as the fruit were stored from 0 days to I99 days. Flesh
firmness values decreased markedly after lOO days of storage(Table
2). The fruit stored for I99 days(TabIe 3) had similar
characteristics to those stored for lOO days. 'Macspur‘ fruit had
the least flesh firmness, whereas 'Rome' fruit were usually the most
firm throughout the experiment. Starch content decreased for
'Macspur' and 'Empire' at the second and third harvests and upon
storage for lOO days(Table l) and I99 days(Table 3). The starch
rating of 'Rome' fruit at the earliest harvest (Table l) was high,
indicating a low starch content. ‘The subsequent changes due to
later harvests and longer storage periods. therefore, were small.

Since hourly readings were similar, singular hourly C02 data
for main effects and interactions were utilized. Two methods were
evaluated to determine the most suitable time and method of data
collection. The first method utilized the rate of total CO2
evolved at l to 6 hours after the fruit treatment, whereas, the
second method utilized the difference in rate of CO2 before
treatment and the rate of CO2 after the damage treatment at l to A
hours. The mean square values and their level of significance as
determined by analysis of variance are presented in Tables A and 5.

The occurance of highly significant mean square values were similar

for the two methods. The selected hour and method of data

15

1G

 

 

 

 

 

 

Table 1. The internal ethylene, flesh firmness and starch rating
of 10 apples at one day following harvest (0-day storage
duration}.
Internal Ethylene
Ippml
Mban Mean
less 1.0 Flesh Starch
Harvest harvest Cultivar than and Firmness Rating
Data 0.5 1.0 greater (kg/
(1983/ (no. of fruit/
1 9-12 Abcspur 10 0 0 7.9 2.7
1 10-03 Empire 8 1 1 8.1 2.0
1 10-12 Rome 9 0 1 9.1 7.7
2 9-19 Mbcspur 9 0 1 7.1 3.9
2 10-10 Empire 8 2 2 7.9 3.7
2 10-19 Rome- 1 0 9 8.8 7.9
3 9-28 Abcspur 0 0 10 8.7 7.0
3 10-17 Empire 8 0 2 7.5 5.3
3 10-28 Rome 10 0 0 8.8 8.8

 

17

Table 2. The internal ethylene, flash firmness and starch rating
of 10 apples at 100 days following harvest l100-day storage

 

 

 

 

 

 

 

 

duration].
Internal Ethylene
{ppm}
Aban Aban
less 1.0 Flash Starch
harvest harvest Cultivar than and Fineness Rating
Data 0.5 1.0 greater {kg}
{1983/ (no. of fruit)
1 9-12 Abcspur 0 0 10 3.3 8.2
1 10-03 Empire 0 0 10 5.1 9.0
1 10-12 Rome 0 1 9 4.7 8.8
2 9-19 Abcspur 1 0 9 3.7 8.5
2 10-10 Empire 0 0 10 4.1 3.5
2 10-19 Rome 0 0 10 4.8 8.8
3 9-28 Abcspur 0 0 10 4.0 9.0
3 10-17 Empire 0 0 10 4.8 8.5
3 10-28 Rome 0 0 10 4.8 8.8

 

18

Table 3. The internal ethylene, flesh firmness and starch rating of
10 apples at 199 days following harvest (199-day storage

 

 

 

 

 

 

duration).
Internal Ethylene
(ppm)
Mean hban
less 1.0 Flash Starch
harvest harvest Cultivar than and Firmness Rating
Date 0.5 1.0 greater (kg/
(1983/ (no. of fruit/
1 9-12 hacspur 0 0 10 4.3 9.0
1 10-03 Empire 0 0 10 4.7 9.0
1 10-12 Rome 0 0 10 5.2 9.0
2 9-19 hbcspur 0 0 10 3.8 10.0
2 10-10 Empire 0 0 10 3.7 9.0
2 10-19 Rome 0 0 10 5.5 9.0
3 9-28 hbcspur 0 0 10 3.4 10.0
3 10-17 Empire 0 0 10 3.9 9.0
3 10-28 Rome 0 0 10 5.4 9.0

 

19

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collection must detect a clear separation of the damage treatments
to facilitate analysis of the data and the method must be easily
executed. The method employing post-treatment data only, therefore
was selected for use since the complete set of data was available
and this method was much quicker as pretreatment readings were not
needed. Data collected at the fifth hour were selected for use
because the F test(mean square value divided by error term) was
greatest at the fifth hour than at the others due to a small error
value and high mean square value. Furthermore, all main effects of
cultivar, damage treatment. time of harvest and storage duration
were highly significant at all hourly observations (Table A).

The 002 response due to damage treatment decreased in rate of
evolution and was consistent for all damage treatments throughout
(Fig. 2). By the fifth hour there were significant differences(.05
level of probability) in CO2 evolution rates between all damage
treatments except one and two small bruises per fruit and two small
bruises and one large bruise per fruit (Table 6). Four small
bruises per fruit had the highest rate of C02 evolution with the
differences being highly significant from all other damage
treatments. including one large bruise per fruit. The latter was
the second highest in rate of 002 evolution.

The only significant interaction for CO2 evolution involving
damage treatment was for the second hour reading. This interaction

of damage treatment X storage duration (Table A) was significant at

the .05 level of probability.

22

Figure 2. The effect of damage to apples of all cultivars, storage

durations and harvests on the rate of CD2 evolution measured at one to

six hours following application of the damage.

23

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Table 6: The rate of 002 evolution of apples at
the fifth hour following damage treatment, for
all cultivars. storage durations and harvest
maturities.

Damage Treatment C0 Evolution
(ml/kg hr)
2
0 Small Bruises/Fruit l3.85 a
l Small Bruise/Fruit lA.A0 b
2 Small Bruises/Fruit lA.6A bc
A Small Bruises/Fruit l5.2A d
l Large Bruise/Fruit lA.83 c

LSD at .Ol I 0.3A

Letters signify difference by LSD at .0l level
of probability.

The mean square values for the amount of tissue damage are
presented in Table 7. This analysis excluded the non-bruise
treatment because of the absence of damaged tissue, and therefore
zero values. All main effects were highly significant. As shown in
Table 8, highly significant differences occured between amounts of
bruised tissue for all damage treatments. As true for

CO evolution, the greatest tissue damage occured for the treatment

2

of four small bruises per fruit. However, the second most damaging
treatment as measured by bruised tissue was one large bruise. This
was unlike the CO2 response(Table 6) in which the rate of

002 evolution for one large bruise was similar to that of two small

bruises per fruit.

25

Table 7. Analysis of variance mean square values for percent of
bruised tissue based on the total weight of the fruit.
The control treatment data (zero values/ were excluded in
the analysis.

 

 

 

3253:.
Source df Values
Cultivar 2 2.3**
harvest 2 4.0**
Storage Duration 2 41.9**
Damage Treatment 3 174.0**
Cultivar x harvest 4 0.5**
Cultivar x Storage Duration 4 4.5**
harvest x Storage Duration 4 0.4**
Cultivar x Damage Treatment 8 0.2*
harvest x Damage Treatment 8 0.2*
Storage Duration x Damage Treatment 8 2.7**
Cultivar x harvest x Storage Duration 8 1.1**
Cultivar x harvest x Damage Treatment 12 0.08
Cultivar x Storage Duration x Damage Treatment 12 0.3**
harvest x Storage Duration x Damage Treatment 12 0.1
Error 454(2/a 0.08

 

a
The value in parentheses is the number of missing values.

* Indicates significance at the .05 level of probability and **
indicates significance at the .01 level of probability as
determined by the F test.

26

Table 8. Hean percentages of excised bruised tissue
according to damage treatment.

Percent ot total fruit

Treatment weight bruised
................................ ;-------
l Small Bruise/Fruit 0.88 a
2 Small Bruises/Fruit l.73 b
A Small Bruises/Fruit 3.A0 d
1 Large Bruise/Fruit 2.88 c

LSD at .01 ' 0.09

Letters indicate highly significant differences
at the .0l level of probability.

Various correlation relationships of CO2 evolution and percent
of bruised tissue were found to account for no more than I3 percent
of the variance, see Table 9. Correlations of all the data at the
fourth and fifth hours post-treatment account for 7.5 and 8.0
percent of the variance, respectively. Correlations in which the
control values (0 values) were removed at the fifth hour account for
ll.6 percent of the variance. A correlation of the 'Empire' fruit
at the fourth hour post-treatment accounted for l3.0 percent of the
variance. this correlation accounted for the most variance of all

the correlations(TabIe 9).

27

Table 9. Correlation values between carbon dioxide evolution
rates and percentage of bruised tissue at different hours of
observation. Data for all cultivars, harvests and storage
durations were included along with several of the individual
interactions. '

Cultivar, Harvest and
Storage Duration of df 2
Fruit in Correlation R Value

A hr Post-treatment
All Data 666 .077

A hr Index (Pre/Post)
All Data 9A .096

A hr Post-treatment

All Macspur

lst Harvest

lst Storage Duration 2l .055

A hr Post-treatment
All Empire 22l .l3A

A hr Post-treatment

All Empire

2nd Harvest

All Storage Durations 72 .082

A hr Post-treatment

All Empire

2nd Harvest

2nd Storage Duration 22 .0l6

5 hr Post-treatment
All Data 669 .082

5 hr Post-treatment
All Data w/o Control 53A .ll8

28

FACTORS AFFECTING TISSUE DAMAGE

There was a highly significant three way interaction of tissue
damage as determined by brown discoloration involving cultivar,
storage duration. and damage treatment.see Table 7. This three way
interaction is illustrated by the histograms of Figs. 3,A and 5.

Each histogram presented in Figure 3 is for a cultivar relative
to the damage treatments applied following each storage duration.
For 'Macspur' there were highly significant differences at each
damage treatment between fruit stored 0 days and those stored for
l00 and I99 days, with no significant difference between the damage
response of fruit stored for l00 and I99 days. 'Empire' fruit had
highly significant differences between all three storage durations
at all damage treatments, with the least damage at 0 days and the
most at I99 days. 'Rome' fruit responded similar to 'Macspur' in
that there were highly significant differences in the damage
response between fruit stored 0 days and those held for 100 and I99
days, but not between fruit stored for l00 and I99 days. As true
for 'Empire', the least damage to 'Macspur' and 'Rome ' occured at 0
days of storage.

The separate histograms of Figure A are for the damage
treatments. The application of I smell bruise per fruit. upper
left. resulted in no significant difference between cultivars for
both the 0 and l00-day storage durations. The only significant
difference occured at the I99-day storage duration; it was between
the two cultivars of 'Empire' and 'Rome'. Two small bruises per

fruit, upper right histogram, resulted in significant differences

29

Figure 3. The effect of damage treatment and storage duration on the

percent of bruised tissue for three apple cultivars.

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Figure 3.

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Figure A. The effect of cultivar and storage duration on the percent

of bruised tissue for the levels of damage applied to apples.

32

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33

Figure 5. The effect of damage treatment and apple cultivar on the

percent of bruised tissue for three storage durations.

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35

between all three cultivars at the 0-day storage duration. After
the l00-day storage duration 'Macspur' had highly significantly
greater amounts of bruised tissue both 'Empire‘ and 'Rome'. For the
I99-day storage duration, there were highly significant differences
between all three cultivars with 'Empire' receiving the most damage
followed by 'Macspur' and 'Rome'. For four small bruises per fruit,
lower left histogram, no significant differences occured between
'Macspur' and 'Empire' at the 0-day storage duration. There were
highly significant differences between the response of 'Macspur' and
'Empire' to that of 'Rome', with 'Rome' being the most severely
bruised. At the l00-day storage duration there were highly
significant differences between the response of 'Macspur' and that
of 'Empire' and 'Rome', with 'Macspur' being the most severely
bruised with 'Empire' and 'Rome' responding similarily. Fruit
stored for I99 days had highly significant differences between the
response of all three cultivars with 'Empire' being the most
severely bruised followed by 'Macspur' and 'Rome', respectively.
For one large bruise per fruit. shown in the lower right histogram,
there were significant differences at the 0-day storage duration
only between the response of 'Macspur' and 'Empire', with 'Macspur'
and 'Rome' being bruised similarily. above that for 'Empire'. For
the l00-day storage duration there were highly significant
differences between all three cultivars, with 'Macspur being the
most severely bruised followed by 'Rome' and 'Empire', respectively.
There were highly significant differences between the response of

all cultivars for the I99-day storage duration with 'Empire' being

36

the most severely damaged followed by 'Macspur' and ’Rome',
respectively.

The effects of damage and cultivar on the amount of bruised
tissue for the storage durations are illustrated in Fig. 5. At the
time of harvest (0-day storage duration) there were highly
significant differences for 'Macspur' and 'Empire' between all
damage treatments except four small bruises per fruit and one large
bruise per fruit. For 'Rome' fruit there were highly significant
differences between all damage treatments with four small bruises
being the most severe treatment followed by one large bruise, two
small bruises, and one small bruise, respectively. After storage
for l00 and I99 days, all cultivars had highly significant
differences between all damage treatments. Four small bruises per
fruit was the most severe followed by one large bruise, two small

bruises and one small bruise,_respectively.

FACTORS AFFECTING CARBON DIOXIDE EVOLUTION

The 002 evolution of the fruit over time for the main effect

of harvest date is shown in Fig. 6. As characteristic of the curves
for damage treatment(Fig. 2). there was a decline in the rate of

CO evolution with the passage of time at which the CO was

2 2

analyzed. Fruit harvested one week after optimum maturity (third

harvest) had the highest rate of C0 evolution, being approximately

2

l.3 times greater than the other two harvests at all times of
measurement (Fig. 6). Fruit harvested at optimum maturity (harvest

two) had the next highest rates of CO evolution followed closely by

2

fruit harvested one week earlier. The difference between fruit of

37

Figure 6. The effect of time of harvest of apples for all cultivars,

storage durations and damage treatments on the rate of 002 evolution

at one to six hours following the application of the damage.

38

Figure 6 e

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39

harvest three and harvest one or harvest two was highly significant,
whereas the differences between harvest one and harvest two were
highly significant at all readings except the fifth hour.

The effect of storage duration on the rates of C02 evolution
for all cultivars, harvests and damage treatments is depicted in
Fig. 7. Fruit stored for l99 days had the highest rates of
002 evolution followed by fruit stored for l00 days. The lowest
rates of C02 evolution were for fruit treated at harvest (0-day
storage duration). The differences between all three storage
durations were highly significant at all readings. The rates for
fruit stored for 100 days were approximately seventy percent of
those for fruit stored for I99 days; whereas at 0 days the rates
were about sixty percent of those for fruit stored for I99 days.

The reponse of the cultivars, including all harvest dates.
storage durations and damage treatments, is illustrated in Fig. 8.
'Empire' had the highest rate of 002 evolution followed by
'Macspur'. then 'Rome'. There were highly significant differences
in the rate of C02 evolution between all three cultivars. All
curves followed the decreasing pattern with time of sampling as
observed in Figs. 6 and 7.

There was no significant interaction involving damage treatment
on CO2 evolution at any time of sampling (Table A). All two way and
three way interactions involving harvest date, storage duration
and/or cultivar were highly significant (Table A). The three way

interaction of harvest date x storage duration x cultivar was highly

significant as well, with the trends graphically shown in Figs. 9,l0.

and 11.

A0

Figure 7. The effect of storage duration of apples for all cultivars,

damage treatments and harvest on the rate of 002 evolution measured at

one to six hours following application of the damage.

41

Figure 7.

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Figure 8. The effect of cultivar of apples for all damage treatments,
storage durations and harvests on the rate of C02 evolution measured

at one to six hours following the application of damage.

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Figure 8.

 

 

 

 

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Figure 9. The effect of apple cultivar and time of harvest on the

rate of CD evolution measured at the fifth hour following the damage

2

treatment of apples at three storage durations.

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Figure 9.
18‘ ° “W tora e DuratIon I
1 o Empire 0 Day 3 9 I
16 a A _ Rome Beauty 1
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A6

Figure l0. The effect of time of harvest and storage duration on the

rate of 002

treatment of apple cultivars.

evolution measured at the fifth hour following the damage

A7

 

 

 

 

 

 

 

 

 

 

 

 

 

Figure 100
2". MACSPUR I
22-: 1
20: L30 I.05 .4
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0 100 199
STORAGE DURATION (Days)
241 EMPIRE 1
: L30 1
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O 100 199
STORAGE DURATION (Days)
171‘ LSD Ins 1
e . I o, :
1: . - .
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7 l l I
O 100 199

STORAGE DURATION (Days)

A8

Figure ll. The effect of cultivar and storage duration on the rate of

C02 evolution measured at the fifth hour following the damage

treatment for time of harvest.

A9

 

 

 

 

Figure 11.
20 o Hocspur 1
i a Em," 1 st HARVEST J
13.: a Rome Beauty .1
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STORAGE DURATION (Days)

 

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STORAGE DURATION (Days)

50

All cultivars of the 0-day storage duration had increasing
rates of 002 evolution with each succeeding harvest (Fig. 9).
’Empire' fruit had the highest rate of 002 evolution for harvests
one and two, yet the lowest rates at harvest three, similar to that
of 'Rome'. 'Macspur fruit of the first and second harvests had the
lowest rates of C02 evolution at the 0-day storage duration, with a
highly significant increase at harvest three to the highest rate cf
C02 evolution. 'Rome' of harvests one and two had intermediate
levels of €02 evolution between that of 'Empire' and 'Macspur' with
fruit of harvest three being similar to that of 'Empire' after a
highly significant increase.

After l00 days of storage(Fig. 9), all three cultivars had a
decrease in the rate of CO evolution at the second harvest.

2

'Macspur' had the highest rates of CD2 evolution of all three
cultivars at the first harvest. A significant decrease to the
lowest rate of all cultivars occured at the second harvest with a
highly significant increase at the third harvest which was
intermediate in rate between the other two cultivars. 'Empire'
fruit had an intermediate rate of 002 evolution at the first harvest
with a significant decrease at the second harvest, but this level
was the highest of all three cultivars at the second harvest. Hith
'Empire' there was a highly significant increase in the rate of

C0 evolution at the third harvest to the highest rate of all fruit

2
of the loo-day storage duration. 'Rome' fruit had the lowest rate
of all three cultivars at the first harvest with a non-significant

decrease, at the .05 level of probability, at the second harvest. A

51

slight but significant increase was seen at the third harvest, this
rate was the lowest of all three cultivars at the third harvest.

All three cultivars after I99 days of storage had similar
patterns of CO2 evolution relative to sequence of harvest (Fig. 9).
'Empire' and 'Macspur' of harvest one had similar rates of
C02 evolution. For fruit of the second and third harvests. 'Empire'
had the highest rates of 002 evolution. 'Macspur' was intermediate
and 'Rome' was lowest. 'Rome' fruit of the first and second
harvests had similar rates of C02 evolution, but there was a
significant increase for harvest three. All rates of C02 evolution
from ‘Rome' were highly significant lower than the levels of
002 evolution of 'Macspur' and 'Empire' from all three harvests.

The graph for each cultivar in Fig. l0 illustrates the effects
of harvest date and storage duration on 002 evolution. 'Macspur'
fruit of harvest three had the highest levels of CD2 evolution for
all three storage durations. There was a decrease at l00 days for
harvest three to a level similar to harvest one and then a large
increase for fruit stored 199 days, as was true for all harvests.

Harvest two fruit had intermediate levels of CO at the 0-day

2
storage duration with the lowest rates of all three cultivars at the
l00-day storage duration after a small significant increase in
CO2 evolution. There were highly significant increases for harvest
two fruit with successive storage durations to the I99-day storage
duration at a level similar to that of harvest one. Harvest one

fruit had the lowest rates of C0 evolution at the 0-day storage

2

duration. With a highly significant increase at the l00-day storage

duration to a level equivalent to harvest three fruit. Then there

52

was highly significant increase at the l99-day storage duration
which was similar to harvest two, still significantly below the
harvest three fruit.

'Empire' fruit of harvest three (Fig. l0) had significantly
higher rates of C02 than fruit of harvests one and two. The rate
increased significantly with each increase in storage duration.
Harvest one fruit also increased with each increase in storage
durations with the lowest rates of the thre; harvests at the 0 and
I99-day storage duration and between harvest two and harvest three
at the l00-day storage duration. Harvest two fruit had levels of
002 evolution intermediate between harvest one and three at the 0
and l99-day storage durations. 'Empire' from harvest two had the
lowest rate at the l00-day storage duration with no increase in the
rate of C02 evolution between the 0 and l00-day storage durations.

Harvest three of the 'Rome' fruit (Fig. l0-bottom) had the
highest rates of CO2 evolution for all harvest dates with similar
rates at the 0 and l00-day storage durations with a large increase
at the I99-day storage duration. Harvest one fruit had an increase
in the rate of C02 evolution with each successive storage duration,
with the lowest rates of all three cultivars at the 0-day storage
duration. Intermediate levels of C02 evolution occured at the l00
and l99-day storage durations were similar to those of harvest two.
Harvest two fruit had intermediate levels of C02 evolution at the
0-day storage duration with no significant increase after the 0-day
storage duration and a highly significant increase after the l00-day
storage duration.

The effect of storage duration on the cultivar response of

53

fruit from different harvests are presented In Fig. ll. For harvest
one fruit there was an increase in CO2 evolution for all three
cultivars at each successive storage duration. 'Empire' fruit had
the highest rate of 002 evolution at the 0-day storage duration, it
was intermediate at the l00-day storage duration, and similar to
'Macspur' at the I99-day storage duration. The 'Rome' fruit had an
intermediate rate of CO2 evolution at the 0-day storage duration,
and the lowest of the 3 cultivars at the loo and l99-day storage
durations. 'Macspur' fruit, although with the lowest rate of
C02 evolution at the 0-day storage duration, increased greatly to
the highest at the.l00-day storage duration.

The CD2 response of harvest two fruit (Fig. ll) of all
cultivars was similar for fruit stored for 0 and l00 days. 'Empire'
fruit had the highest rate of C02 evolution following all three
storage periods with no significant change between the 0 and l00-day
storage durations with a highly significant between the loo and the
199-day storage durations. 'Rome' fruit had intermediate levels of
C02 evolution for the 0 and l00-day storage durations, with no
significant change between the two. The increase at the I99-day
storage was highly significant, but this fruit had the lowest rate
of C02 evolution of all three cultivars at the I99-day storage
duration. 'Macspur' fruit had the lowest rates of 002 evolution for
the 0 and l00-day storage durations, with a highly significant
increase between the two but smaller in magnitude than other
increases for the cultivars of that harvest. A much higher rate for

'Macspur' occured at the I99-day storage duration than at the

loo-day reading. It was intermediate in rate of C02 evolution at

5A
the I99-day storage duration.

The lower graph in Fig. ll depicts a varied response of the
three cultivars from harvest three. 'Macspur' had the highest rate
of CD2 at the 0-day storage duration, it decreased to an
intermediate level at the l00-day storage duration and remained
intermediate in spite of a highly significant increase at the
I99-day storage duration. 'Empire' and 'Rome' fruit had similar
rates of 002 evolution at the 0-day‘ storage duration, but
significantly below that of 'Macspur'. 'Empire' fruit significantly
increased in CO2 output at both the loo and I99-day storage
durations to the highest rate of all three cultivars. 'Rome', on
the other hand, did not significantly change at the lOO-day storage
duration and showed a highly significant increase at the I99-day
storage duration. 'Rome' had the lowest rates of C02 evolution of
all three cultivars at the loo and I99-day storage durations.

Fruit of harvest three generally had the highest rates of

002 evolution for all three cultivars. The harvest one 'fruit had
intermediate levels of C02 evolution at the 0 and I99-day storage
durations and the lowest rate when sampled after l00 days of
storage. Harvest one fruit were lowest in C02 evolution at the

0-day storage duration, intermediate at the l00-day storage

duration. and variable at the I99-day storage duration.

55

DISCUSSION

The similar and significant increases in C02 evolution and
percentage of bruised tissue that resulted from the damage
treatments substantiate the findings of Klein (l8) for a 002 damage
response of apples. The relatively small amounts of bruising
applied which yielded measurable increases in the rate of
C02 evolution indicate that the CD2 damage response is suitable for
determining small differences in the protective qualities of
packages used for apple handling. One small bruise, approximately
equivalent to dropping a fruit from a height of IS cm yielded a
significant increase in 002 evolution. One large bruise,
approximately equivalent to dropping fruit from a height of 50 cm,
yielded a lower increase in 002 evolution than four small bruises.
Drop heights of 50 cm are not likely to occur during normal handling
practices, whereas, numerous small drops of IS cm may occur.

Neal and Hulme (30) and Klein (l8) concluded that the increased
C02 evolution was due to physical damage resulting from the
decarboxylation of malic acid released from cells upon rupture of
the cellular membrane, as a result of the physical stress. Marks
and Varner (2i) reported this same process to be the source of the
increased 002 damage response in tomatoes. The almost complete
absence of significant interactions of damage treatment with
cultivar, harvest time and/or storage duration on CO2 evolution

suggests that there was a consistent damage response, so that

regardless of variations in other fruit characteristics, there is a

56

consistent measurable increase in C02. Previous studies (Dewey,
Klein and Parker-Unpublished data) have shown that this consistent
damage response occurs for numerous cultivars and for apples of
various maturities and storage backgrounds. The 002 damage response
was found by Klein (l8) to occur even in apple fruit harvested five
weeks after ontogeny.

The significant three way interaction involving cultivar.
storage duration and damage treatment on the amount of bruised
tissue substantiates the findings of Hyde and ingle(lA) that changes
in fruit characteristics are of effect on the amount of damage. It
is widely accepted that apple fruit cultivars, times of harvest and
storage durations have variable maturation, ripening and senescent
responses, all of which result in variable responses in
susceptability to bruising (38,A0,A2,A6). All three cultivars
responded similarly at harvest (Fig. 5), but the effect of fruit
maturation and softening was observed at the loo-day storage
duration with 'Macspur' being bruised to an amount highly
significant above that of 'Empire' and 'Rome' for all treatments
except i small bruise per fruit. Yet after I99 days of storage,
'Empire' was more severly bruised than 'Macspur' and 'Rome'. This
was likely due to the earlier onset of senescence for 'Macspur'
since barely 75 sound fruit were available from the approximately
200 harvested for this experimental run. It is unlikely that the
'Macspur' used at this time were representative of the fruit at
harvest in that they were probably the hardiest and firmest fruit of
that harvest.

As would be expected. the other interactions showed that the

57

fruit receiving the most severe damage treatments had the greatest
amount of bruised tissue. 'Rome' fruit had the least amount of
bruised tissue probably as a result of greater inherent flesh
firmness at harvest and thereafter.

The significant interaction of harvest x cultivar x storage
duration on CO2 evolution was a likely result of fruit maturation,
ripening and senescence. The increased rates of CO2 evolution
between harvest one and harvest three are accountable in that fruit
from harvest three would be well into the climacteric rise so as to
be producing and evolving large quantities of 002 at the time of
harvest. Harvest two fruit should be lower in rates of
002 evolution if picked at the optimum time prior to the initiation
of ripening. Harvest one fruit would have a low rate of CO2 because
the fruit had not entered the climacteric phase of respiration
associated with maturation and the coinciding increase in 002 and
CZHA evolution. Observing the 'Macspur' maturity response, which
was the most consistent. (Table I) it is likely that the fruit at
harvest one were picked before the climacteric rise, whereas harvest
two was just before the climacteric rise and harvest three fruit
were already in the climacteric rise.

'Rome Beauty'. a long-term storage cultivar, had the lowest
rates of CO2 evolution, probably because of the lower rates of
respiration characteristic for this cultivar with maturation and
ripening (39). The main effect of cultivars on the rate of
£02 tended to be misleading showing 'Empire' significantly above
'Macspur' in terms of rate of CO2 evolution at all observations.

According to the three way interaction, 'Macspur' and 'Empire'

58

fluctuated with similar rates of CO2 evolution. Possibly the
similarity in rates of CO2 evolution for these two cultivars is
related to the inherent genetic characteristics of the fruit because
'Empire' was developed from a cross of 'Mclntosh' and 'Red
Delicious'.

The causes for fruit stored for the longest duration to have

the highest rates of CO with the fruit stored for l00 days

2.
intermediate. are probably related to the maturity and ripening
response of the fruit. During cold storage the low temperature
decreases the rate of respiration considerably and the fruit matures
at a slower rate. But when the fruit are brought to room
temperature the respiration rate increases drastically. Fruit
stored for l99 days with the highest rate of CO2 evolution was most
likely a maturity response of the fruit at the end of ripening and
entering the senescent stage of maturity.

CO2 evolution differences due to the respiration response to
temperature are probably minimal since all experiments were
conducted under similar temperature conditions. The temperatures
observed for each experimental run by means of a maximum-minimum
thermometer showed fluctuations no greater than 3°C for the 2A hr
period. Since the fruit was harvested and transported under
variable conditions of temperature and were placed in cold storage
at different times, these factors would tend to increase
experimental variation.

The decrease in the rate of CO evolution of the fruit during

2

the time period in which the CO2 response was measured is a typical

response of the fruit due to handling. The maximum concentration of

59

CO2 in the static system used was observed to be 3.2 percent at the
sixth hour, with a range in the containers from 0.l3 to 0.9A at the
first hour and 0.A3 to 3.2 at the sixth hour. It is unlikely that
at the fifth hour, which was selected as the most suitable time,
that feed back inhibition of CO2 was a factor on respiration.
Oxygen concentrations did not drop below l9 percent within the
containers sealed for 2A hours (data omitted). it is more likely
that the decrease was due to a decrease of the fruit handling
response rather than to an inhibition of respiration by low oxygen
or high carbon dioxide in the containers.

The lack of meaningful correlations between CO2 response and
the amount of damaged tissue was not unexpected due to the complex
interactions observed. There were numerous highly significant
interactions of damage treatment on the amount of damaged tissue
which did not occur for the C02 response. The subjective selection
process required for excising damaged tissue might result in
considerable judgmental error in making a distinction between
damaged and nondamaged tissue, which is based primarily on the
discoloration of damaged tissue as a result of the oxidative
browning reaction. Hith maturation and ripening the ensuing
decrease in flesh firmness could result in numerous undamaged cells
being masked by the discoloration of a relatively small amount of

damaged cells. The studies of Klein (18) that show the CO response

2
to result from only ruptured cells would be a more reliable measure
of actual damage and truly an objective measure of this damage.

Some changes in methods may decrease the variation observed in

this study. This could best be corrected by increasing the number

60

of fruit per sample, which was not feasible for this research
because of the numerous factors studied. The use of more apples per

sample would decrease the effect of natural fruit variations.

6l

SUMMARY Mg CONCLUSIONS

Apple fruit varying widely in stages of maturation and ripeness
as a result of cultivar. time of harvest and storage duration were
subjected to degrees of physical damge by bruising ranging from
slight to severe. Each factor was of significant influence upon the
amount of damaged tissue and the increase in carbon dioxide
evolution brought about by the damage treatment. The two measures
of damage, however, were not correlated, probably because of the
differing influence of these factors on the two methods of damage
assessment. Four of the six interactions involving damage treatment
were significant for the amount of damaged tissue, whereas. none of
the six were significant for carbon dioxide evolution.

The significant differences between all damage levels ranging
from no bruising to four small bruises to one large bruise indicate
that the carbon dioxide response was a reliable measure of damage,
regardless of the cultivar, time of harvest and storage duration of
the fruit. It is suggested that the carbon dioxide response is a
more suitable method of assessing damage than the amount of damaged

tissue as determined by visual examination.

LITERATURE CITED

‘00

ll.

12.

13.

62

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