Temperature and Respiration of Ripening Bananas
Author(s): Albert Jackson Olney
Source: Botanical Gazette , Dec., 1926, Vol. 82, No. 4 (Dec., 1926), pp. 415-426
Published by: The University of Chicago Press
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TEMPERATURE AND RESPIRATION OF
RIPENING BANANAS
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 3 61
ALBERT JACKSON OLNEY
(WITH ONE FIGURE)
Introduction
The appearance of the banana as an important fruit on the
American market is relatively recent, although it has been cultivated
and used from earliest times. The first shipment to the United
States (25) was made in i804, and another in i830, but it was not
until i866 that bananas were imported regularly, and then only in
modest quantities to seaboard towns. During the last twenty-five
years the banana trade has been extended throughout the United
States until now it represents an important industry.
Bananas must be picked while still green, in order to withstand
the long journey from the tropical countries to the markets. A week
to ten days are required for transportation to points in the United
States. If ripening begins in transit the fruit deteriorates before it
reaches its destination, and results in heavy losses. The problem of
chief concern, therefore, is that of finding improved methods of re-
tarding the ripening in transit and storage so that the fruit may reach
the market in a condition of high quality and attractive appearance.
The degree of maturity when harvested is determined by the growers
in a -purely empirical way, the degree of maturity when picked de-
pending on the distance to be shipped. Those going to Europe are
harvested somewhat greener than those for the United States, and
the greener lots are said to be poorer in quality than those picked
when more mature (13). Shipments are made in specially con-
structed steamships providing reduced temperatures by ventilation
or refrigeration. Naturally ventilated steamships are commonly used
for shipments from Central America or Jamaica to the Gulf ports.
Distribution is then made to the central portions of the country in
refrigerator cars. Refrigeration shipments are held as near i6' C.
as possible.
4IT5 [Botanical Gazette, vol. 82
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4i6 BOTANICAL GAZETTE [DECEMBER
After arriving at their destination the bunches are hung in curing
rooms at I 7-20' C., providing 82-86 per cent humidity. The trade
considers i50 C. the lowest safe temperature for green bananas, and
140 C. as the minimum for ripe ones. Chilling is said to occur at
lower temperatures, injuring the quality and preventing the de-
velopment of attractive appearance. Cargoes sometimes arrive at
the ports too ripe for general distribution, while other shipments ap-
parently handled in the same manner arrive in good condition. The
causes for such occurrences are not well understood by importers.
There is a general belief that a few ripe or ripening bananas among
a lot of green ones greatly accelerate the ripening of the cargo.
The first noticeable development of the green fruit is the gradual
change of color, accompanied by a softening of the pulp, due to the
hydrolysis of pectose to pectin (I7). The change from green to
yellow takes place in 3-5 days at curing temperatures after arriving
at their destination. Two or three days after the bright yellow stage,
brown specks develop through the yellow, and the pulp reaches its
highest flavor, aroma, and quality. This stage is short; the pulp
soon becomes too soft, loses flavor, and begins to decay. When green
the peel is filled with a white viscous fluid and the pulp is acid, bitter,
and astringent. GERBER (14) cites BOUSSINGAULT and COREN-
WINDER (9, io) as attributing the acidity of bananas to malic aci
but he was able to show only the presence of citric acid. As the ripen-
ing continues the peel becomes dry and leathery, and the pulp in-
creases in sweetness until the fully ripe stage is reached.
A number of investigators have studied various phases of the
ripening process of bananas. The Jamaica Department of Agricul-
ture (27) investigated the gases given off by oranges and bananas.
The carbon dioxide liberated was thought to retard the ripening,
but unknown gases or emanations which they thought were given
off by the oranges were believed to induce premature ripening. The
practical deduction was drawn that separate storage was desirable
for the two sorts of fruit during sea transportation. The chemical
changes taking place in ripening bananas have been reported by
GORE (i6), MARCANO and MUNTZ (2I), RICCIARDI (24), DOHERTY
(I2), COLBY (8), BALLAND (4), ATWATER and BRYANT (I), CHACE,
TOLMAN and MUNSON (7), PRINSEN GEERLIGS (22), YOSHIMURA
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g926] OLNEY-TEMPERATURE AND RESPIRATION OF BANANAS 4I7
(28), REICH (23), TALLARICO (26), and BAILEY (23). Most of these
analyses were made on green and ripe fruit. GORE (i6) makes a
sweeping criticism of the early analyses because their results were
invariably expressed in terms of the percentage of the pulp when
analyzed, so that the data are on a constantly shifting basis, since
the peel continually loses weight while the pulp increases in weight
during ripening.
LANGWORTHY and MILNER (20) studied the ripening of bananas
in a respiratory calorimeter. They found that the volume of oxygen
consumed was practically equal to the volume of carbon dioxide
liberated. Their determinations show from the thermal quotient and
carbon dioxide produced that the respiration was due entirely to the
combustion of carbohydrates.
Materials and methods
The bananas used for the following experiments were obtained
from the United Fruit Company through the kindness of Mr. J. W.
LEATHERS, to whom thanks are gratefully accorded. Green bananas
were obtained as soon as possible after arrival in Chicago. Two lots
were received from Honduras, one ventilated and one refrigerated,
and a ventilated and a refrigerated lot from Jamaica.
JOHNSTONE'S modification of Magness' respiration apparatus
(ig) was used to determine the amount of oxygen consumed and
carbon dioxide liberated during the ripening. The apparatus was
maintained at 200 C. in a Freas water bath. One hour was allowed
for adjustments of temperature of the apparatus to that of the bath
at the beginning of each experiment, and one-half hour between the
daily runs when fresh quantities of NaOH were inserted for absorp-
tion of carbon dioxide. A Beckman thermometer was used to check
the uniformity of the temperature of the bath. The temperature
variation ranged from o.oS5 to 0.20? C. The oxygen absorbed by the
bananas was considered equal to the volume of water drawn into the
cylinder. The amount of carbon dioxide liberated by the bananas
was determined by titrating the NaOiH, using the double indicator
method (5). The individual bananas were cut apart and weighed be-
fore being put into the respiratory chamber. Later weighings were
made at the end of each day's run. Four refrigerators operated at
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4I8 BOTANICAL GAZETTE [DECEMBER
20, 70, I2? and I70 C. were used for determining the effect of low
temperatures on ripening.
Results
The results obtained from respiration studies are given in tables
I, II, III, and IV. At the end of each experiment evidences of initial
decay were usually apparent, although the pulp was in good condi-
tion. The maturity of the different lots was not uniform when re-
go
8o~~~~~~~~~
70
40_
30
20
24 48 72 96) 120 144 I6)8
Hours
FIG. I.-A, Honduras refrigerated 20?; B, Jamaica refrigerated 200; C, Jamlaica
venltilated 200; D, Honduras ventilated 200; E, Jamaica refrigerated 120 C.
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1926] OLNEY -TEMPERATURE AND RESPIRATION OF BANANAS 4I9
ceived, even when account is made for the method of cooling in ship-
ment. The Jamaica ventilated lot appeared most mature, while the
Jamaica refrigerated and the Honduras ventilated were similar in
apparent maturity, and the Honduras refrigerated was greener than
the others and not as ripe at the end of the experiment. Table V
shows the daily loss in weight and production of carbon dioxide at
I2' C. Fig. i indicates the nature of the respiration of the different
lots at 20? C. and at I20 C. Table VI shows the effects of low tem-
peratures on ripening.
The Jamaica ventilated bananas (table I) ripened very rapidly,
and by the end of six days were becoming soft, showing initial decay,
TABLE I
JAMAICA VENTILATED; RESPIRATION AT 200 C.
PER-
WEIGHT CENT- CALOR- CC. CO. CC. 0 CO./O.
DATE AND TIME OF RUN (GM.) AGE KGMHR IES PER PER PER PER REMARKS
LOSS IN KGMHR KGMHR KGMHR KGMHR
WEIGHT
MarchI4,23hours.. 325.0 o.o6 7.475 0.22 44-9444.I5I 1.OI Slightly
yellow
March I5, 25 hours. . 324.8 o.i8 8.I20 0.27 53.7I 49. 26i i.09 Mostly
yellow
March i6, 23.5 hours 324.2 0.27 7.6i8 0.3i 6i. 57 55. I3I II Yellow
March I7, 23.5 hours 323.3 o.o6 7.597 0.32 62. 97 53. 3II i.i8 Yellow
March i8, 23.5 hours 323-I o.i8 7.593 0.3i 60. 50 50.04I I.20 Brown
March I9, 23.5 hours 322.5 0.28 7.578 0.30 58. 80 46. 22I I.24 Very ripe
and were too ripe for handling on the market; however, the pulp
was sound and had a rich aromatic flavor. The fruit inclosed in the
respiratory chamber seemed to decay more rapidly than samples of
the same lot stored in the laboratory, due to higher humidity. The
loss in weight during this experiment was very low, about 6o per
cent being due to liberation of carbon dioxide. The kilogram hours
were determined by multiplying the kilograms of bananas by the
number of hours in the run. The calories of heat produced were cal-
culated from LANGWORTHY and MILNER'S (20) determinations,
showing that 2.58 calories of heat are produced for each gram of
carbon dioxide liberated. The volume of carbon dioxide given off
and oxygen consumed were nearly equal in the daily runs, the re-
spiratory quotient gradually increasing from i.oi for the first run
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420 BOTANICAL GAZETTE [DECEMBER
to I.24 at the end. There was a gradual increase in the volume of
carbon dioxide produced until the bananas were fully yellow, fol-
lowed by a slight decline.
The Jamaica refrigerated lot appeared green when received but
ripened rapidly. The daily loss in weight was fairly uniform, and
averaged about 0.4 per cent of the weight of the fruit per day, of
TABLE II
JAMAICA REFRIGERATED; RESPIRATION AT 200 C.
PER-
WEIGHT CENT- CALOR- CC. CO C CC. OX COVO,
DATE AND TIME OF RUN EGHT AGE KGMIHR IES PER PER PER PER REMARKS
(G.) SS IN KGMHR KGM11R KGAMHR KGMHR
WEIGHT
March 28, 24 hours.. 770. I 0.53 i8. 48 0.38 75. 98 77. 65 O0 97 Green
March 29, 23 hours.. 766.0 0.45 I7.6i 0.44 86. 42 83. i9I. 03 Yellowing
March 30, 23.5 hours 762. 5 0.43 I7. 90 0.43 85. I3 76. 8i I. IO Mostly
yellow
March 3I, 23 hours.. 759. 2 O. 4I i8. 20 0.42 82. 47 76. 37 I.07 Yellow
April I, 24 hours.. 756. I O. 4I I8.40 0.42 83. 72 74. 97 I . II Yellow
April 2, 23 hours.. 752. 9 0.42 I7. 30 0.36 70.48 6o. II I . I7 Brown
April 3,23 hours.. 750. 3 0.34 I7.20 O. 3o 6.oo 59.30 I.OI Veryripe
TABLE III
HONDURAS VENTILATED; RESPIRATION AT 200 C.
PER-
WEIGHT CENT- CALOR- CC. CO. CC. O0 COWO/
DATE AND TIME OF RUN WE I.) AGE KGIrHR IES PER PER PER PER REMARKS
(f. LOSS IN KGMHR KGAMHR KGAMHR KGM11R
WEIGHT
April II, 23 hours.. 688.4 0.40 I5. 83 0.32 63. 2i 64. IO O. 99 Light green
April I2, 23 hours. . . 685. 6 0.42 I5.76 0.36 7I- 34 62. 78 I. I3 Ycllowing
April I3, 23 hours... 682. 7 0.38 I5.70 0.33 64.45 62.4I I. o8 Mostly
yellow
April I4, 23 hours. . . 68o. I 0.35 I5. 64 0. 29 56. 52 53. o6I. o6 Yellow
April I5, 23 hours ... 677. 7 O. 37 I5. 58 0. 27 54. 27 49. o8 i.io Yellow
April I6, 23 hours. . . 675. 2 0.34 I5. 53 0. 26 52.48 49.58 I.05 Browning
April I7, 23 hours.. 672. 9 0.33 I5. 47 0. 25 48. 82 47. 86 I.02 Very ripe
which 3I per cent was due to loss of carbon dioxide. The volume of
carbon dioxide produced was considerably greater per kilogram per
hour than that produced by the Jamaica ventilated lot. The greatest
production of carbon dioxide occurred on the second day, followed
by a gradual decline to the end of the yellow stage, and then a rapid
falling off. The respiratory quotient was slightly more than unity
except for the first day.
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I926] OLNEY-TEMPERATURE AND RESPIRATION OF BANANAS 42I
The Honduras ventilated bananas ripened at approximately the
same rate as the Jamaica refrigerated, and the daily loss in weight
was similar in amount. The largest production of carbon dioxide
occurred on the second day and gradually declined to the end of the
experiment. The respiratory quotient was slightly above unity in
most cases.
TABLE IV
HONDURAS REFRIGERATED; RESPIRATION AT 200 C.
PER-
WEIGHT CENT- CALOR- CC. CO2 CC. 02 CO2/02
DATE AND TIME OF RUN (GHT AGE KGMHR IES PER PER PER PER REMARKS
(GM.) LOSS IN KGMHR KGMHR KGMHR KGMHR
WEIGHT
April i8, 23.5 hours. I54-5 0.50 3.630 0.32 62. 80 63.35 0 99 Green
April I9, 23.5 hours. I53-7 0.50 3.6I2 0.37 72.65 66.44 i. o8 Slightly
yellow
April 20, 23.5 hours. I52.9 0.39 3.593 0.38 74-59 73-75 I.OI Yellowing
April 2I, 23.5 hours. I52.3 0.46 3.579 0.44 87.79 89.4I 0.99 Yellow
April 22, 23.5 hours. I5I. 6 0.39 3.562 0.43 85.29 84.22 I.OI Yellow
April 23, 23.5 hours. I5I.0 0.39 3.548 0.37 72.71 66.38 I. 09 Slightly
brown
April 24, 23.5 hours. I50.4 0.33 3.534 0.34 67.68 67.9I o.99 Ripe
TABLE V
JAMAICA REFRIGERATED; RESPIRATION AT I20 C.
PERCENT-
DATE AND TIME OF RUN WEIGHT Loss
AGE IN
KGMHR PER KGMHR REMARKS
PRKAH
WEIGHT
March 28, 24 hours.8 5.0 0.29 I4.04 22.27 Green
March 29, 23 hours.... 583.3 0.25 I3.42 29.96 No change
March 30, 22 hours...... i. 8 0. 22 I3. 8o 29. I3 No change
March 3I, 22 hours....8.o. 58 0. 24 I2. 77 29. 73 No change
April I, 23 hours ... 579.0 0.34 I3.3I 26. 85 Slight change
The Honduras refrigerated bananas were less mature when re-
ceived than any of the other lots. The loss in weight for the first day
was o.5 per cent of the weight of the fruit, and declined to 0.3 per
cent for the last day of the experiment. About 25 per cent of this
loss was due to liberation of carbon dioxide and the remainder to
transpiration. The volume of carbon dioxide was relatively high per
kilogram per hour, and higher than the Honduras ventilated, the
maximum occurring on the fourth day. The respiratory quotient was
more nearly unity than any of the other lots.
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422 BOTANICAL GAZETTE [DECEMBER
The respiration of the Jamaica refrigerated lot at I 2? C. is shown
in table V. The daily loss in weight was reduced to nearly one-half
and the volume of carbon dioxide produced to about one-third that
at 20 C.
The effect of low temperatures on the ripening of bananas is
shown in table VI. The lots held at 2', 70, and a part of those at
I2? C. were transferred April 5 to the I7? refrigerator, and the re-
mainder of those held at I2? were continued at that temperature.
The lot held at I7' ripened in six days, while those held at the lower
TABLE VI
EFFECT OF LOW TEMPERATURE ON RIPENING (PUT IN REFRIGERATORS MARCH 28)
DATE OF 20 C - C 2 C 17 C
EXAMINATION 2c c I2c17c
March 30... Green Green Green Quite yellow
April Slightly No change No change Yellow
darker
April 3 . Same Same Same Ripe (removed)
April 5. Same Same Slightly yellow
Put at I7? Put at I7O Part put at I7O and part continued at 120
April 7 . Slight Slight Little
change
April 7-I4.. Gradual change to a dull yellowish Gradual change to yellowish
brown with black and dark brown green with black areas;
areas; no bright yellow stage; pulp remained hard; not eatable
became soft, poor quality, lacking
aroma; no marked differences in the
three lots
temperatures showed little change during this time, and when trans-
ferred to ripening temperature failed to ripen normally. After eleven
days they had developed a dull unattractive appearance and lacked
quality and aroma. Those held continuously at I 20 C. were still hard
and green at the end of the experiment, although they were begin-
ning to show signs of decay.
Discussion
The changes taking place during ripening of bananas are rapid,
and the fully ripe period is very short, usually lasting about two or
three days at ordinary temperatures. The rate of respiration at 20?
C. was found to increase rapidly during the first two or three days
and then gradually declined. The refrigerated lots, tables II and IV,
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I926] OLNEY-TEMPERATURE AND RESPIRATION OF BANANAS 423
gave a high respiration rate followed by a rather sharp decline; while
the ventilated lots, tables I and III, did not respire at so high a rate
and declined more gradually. This difference in respiratory activity
may have been due to an accumulation of sugars at the low temper-
atures, so that a more rapid respiration occurred when brought into
ripening temperatures. The volumes of carbon dioxide liberated and
oxygen absorbed were practically equal, as shown in the tables. In
the combustion of carbohydrates the volume of carbon dioxide pro-
duced is exactly the same as that of oxygen consumed; that is, the
respiratory quotient is i.oo. It appears therefore that the oxidation
of carbohydrates is the principal process involved. This is also con-
firmed by inspection of analyses of the banana (i6), which show that
carbohydrates and water comprise about 96 per cent of the whole
fruit, with less than i per cent of fats or proteins. Further to the
point, LANGWORTHY and MILNER determined that the thermal
quotient for normal ripening bananas indicates that the carbon
dioxide produced is due solely to the complete combustion of carbo-
hydrates. They found that 2.58 calories of heat are produced for
each gram of carbon dioxide eliminated. Calculations on this basis
show that approximately 0.3 calorie is produced per kilogram per
hour during ripening at 20? C. This means the production of IO-I5
calories of heat per hour from a single bunch of bananas at maximum
respiration, and shows the possibility of a considerable rise in
temperature in the ship or car during transportation. This may be
an important factor in causing the occurrence of occasional ship-
ments which become over-ripe in transit. It is important, therefore,
not only that the surrounding temperature should be carefully regu-
lated, but that proper ventilation should be provided to carry away
the heat formed by the bananas themselves. It has been suggested
that the heat liberated by the bananas during ripening may be due
in part to bacterial action, but bacteriological studies by BAILEY
(3) indicate that the inner part of the pulp is practically sterile,
while the inner portion of the peel is sparsely inhabited by bacteria
which he found were practically inactive until the fruit became ripe.
The loss in weight during ripening was relatively low, due to the
highly cutinized surface of the peel and the high humidity main-
tained throughout the experiments. About one-fourth of the loss
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424 BOTANICAL GAZETTE [DECEMBER
was due to escape of carbon dioxide, and the remainder to transpira-
tion of water. During ripening the pulp increases considerably in
water content, due partly to absorption of water from the peel
because of increased osmotic pressure as starch is changed to
sugar, and partly from the combustion of glucose according to the
equation C6H1206+602->6CO2+6H20. The amount of carbon
dioxide liberated at I2? was reduced approximately to one-third that
at 20? C. The volume of oxygen absorbed at this temperature was
not determined, and consequently the respiratory quotient cannot
be shown, but the efficacy of reduced temperatures in retarding
ripening is clearly indicated.
On March 28, samples of green bananas were put in refrigerators
at 20, 70, I2', and I70 C. Those held at I70 were fully ripened in six
days, while those held at the lower temperatures were still very
green. After eight days those held at 2? and 70, and a part of those
at I20 were put in the I 70 refrigerator and held there for eleven days.
It was soon strikingly apparent that the pre-refrigeration at 20, 70?
and I20 had made a decided change in the character of ripening
when placed at I7f C. All lots ripened slowly. The change in color
was very gradual and the bright clear yellow failed to develop. At
the end of the experiment the appearance of all lots was dull and
unattractive, yellowish brown in color, spotted with blackened areas,
and showing evidences of initial decay. The pulp had softened and
appeared to be in good condition. The quality was almost as good
as some often obtained on the market, but was noticeably flat and
decidedly lacking in the aroma so chracteristic of properly ripened
bananas. These results are in agreement with the earlier work of
GERBER (14). This indicates that a temperature of I20 or lower not
only retards ripening but permanently affects the fruit, so that
normal ripening does not occur when brought into ripening temper-
atures.
Summary
i. At 20? C. green bananas ripen rapidly, becoming fully ripe in
5-7 days after arrival in Chicago.
2. The respiratory quotient is approximately unity, indicating
that the oxidation consists solely of the combustion of carbo-
hydrates.
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g926] OLNEY -TEMPERATURE AND RESPIRATION OF BANANAS 425
3. The rate of respiration increases rapidly at the beginning of
ripening and falls off gradually.
4. Refrigerated bananas had a higher rate of respiration than
those naturally ventilated, and declined more rapidly as they became
ripe.
5. Respiration at I20 was reduced to about one-third that at
200 C.
6. The daily loss in weight was relatively low under the condi-
tions of these experiments.
7. Approximately 0.3 calorie of heat is produced per kilogram per
hour by ripening bananas at 26? C.
8. Refrigeration at I 2? C. or lower not only retarded ripening in
storage, but permanently prevented normal ripening when trans-
ferred later to a ripening temperature. The low temperatures pre-
vented the development of bright yellow color, high quality, and
aroma.
Grateful acknowledgment is made to Professor C. A. SHULL for
suggestions and criticisms, and to Dr. S. V. EATON for reading the
manuscript.
UNIVERSITY OF KENTUCKY
LEXINGTON, Ky.
[Accepted for publication January I8, 1926]
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