The Respiration of Bananas in Presence of Ethylene

Author(s): R. Gane
Source: The New Phytologist, Vol. 36, No. 2 (Apr. 24, 1937), pp. 170-178
Published by: Wiley on behalf of the New Phytologist Trust
Stable URL: https://www.jstor.org/stable/2428502
Accessed: 27-07-2020 12:57 UTC

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 [I70 ]

THE RESPIRATION OF BANANAS IN

PRESENCE OF ETHYLENE

BY R. GANE, M.Sc., PH.D.

Low Temperature Research Station, Cambridge

(With 7 figures in the text)

THE banana fruit of commerce is always gathered immature and

green and ripened after transport. There are not yet available
the results of any complete study of the temperature-respiratory
activity-time relationships of immaturely gathered fruit. Such results
as have been obtained with post-transport fruit (I936) suggest that
the onset of ripening is accompanied by a sharp rise in respiratory
activity similar in general character to that occurring in the apple
80

AA

9~ 60 1

20 i

z
5 JO L~~5 20 25
Days

Fig. i. Rate of production of carbon dioxide by bananas at I50 C. in air and

in air containing I part per million of ethylene. Bananas II in fresh air
throughout. At A, bananas I ventilated with air containing I p.p.m. of
ethylene.

and pear as described by Kidd & West and termed by them the
climacteric. In the case of the apple these authors have shown that
ethylene administered before the climacteric stimulates its immediate
onset and that administration after the climacteric has no effect.
A series of experiments has been carried out in which post-ship-
ment bananas have been treated with ethylene in the pre-climacteric
phase of low respiratory activity. In all cases the action of ethylene
has been the stimulation of the immediate onset of the rise in
respiratory activity associated with ripening.
Fig. i presents the results of one such experiment. Comparable
samples of fruit were obtained by using the two halves of a hand, one

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 Respiration of Bananas in Presence of Ethylene I7I
of which was ventilated with air and the other with the gas mixture.
The gas mixture was made by introducing 2-i c.c. of ethylene in a
partially evacuated steel cylinder and then adding 75 cu. ft. of air
from a similar cylinder containing I50 cu. ft. of air under pressure.
Fig. 2 presents the result of an experiment in which ethylene was
administered in the post-climacteric phase. At this stage it is without
effect on respiratory activity.
An interesting extension of this type of experiment is one in which
the application of ethylene is made after the upward tendency in
respiratory activity has already begun. The results obtained in this
case are shown in Fig. 3. It will be seen that the rate of rise is
accelerated.

,40

20
X20 4 8
Days

Fig. 2. Rate of production of carbon dioxide by bananas at I50 C., in air and
in air containing ethylene. At A, I c.c. of ethylene was added to the air
stream.

Now in the case of apples the present author (1935) has shown that
with the onset of the climacteric rise in respiratory activity ethylene
is produced by the fruit itself, while Kidd & West have shown that
the volatiles from post-climacteric apples stimulate the climacteric
in unripe fruit and also that the same is true in the case of bananas.
Confirmation of their observation with regard to bananas has been
obtained.
Three samples of bananas were obtained from one hand of fruit,
sample I was ventilated with air from a cylinder, sample II was
similarly ventilated but o i c.c. of ethylene added to the air stream,
while sample III was ventilated with air that had previously been
used to ventilate ripe bananas and then passed over moist soda lime
to remove carbon dioxide.

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 I72 R. GANE

The course of respiratory activity of all three samples is shown in

Fig. 4. Readings of the respiratory activity were obtained every

80'

"A 601
b~

40

s 20 5

Days

Fig. 3. Rate of production of carbon dioxide by bananas at I5? C., in air and
in air containing I part per million of ethylene. Bananas I in air through-
out. At A, bananas II ventilated with air containing I p.p.m. of ethylene.

Days

Fig. 4. Effect of volatile products from ripe bananas on the production of

carbon dioxide by unripe bananas (at I5? C.). Banana I in fresh air
throughout. At A, air from a ripe banana was passed over banana III.
At B, banana III was returned to fresh air. At C, O-I C.C. of ethylene was
added to air stream over banana II.

4 hours, and it will be seen that in samples II and III the respiration
started to increase after 8 hours from the time dosing started.1
A germinating-seed test for the production of traces of ethylene
by bananas has been applied. Peas germinating in a closed vessel in

1 In this particular series, the peak of respiratory activity was reached at

almost the same time by all three samples, and although the respiration was
increased by the treatment it was observed that ripening in this experiment did
not appear to be greatly accelerated by the ethylene treatment.

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 Respiration of Bananas in Presence of Ethylene I73
the presence of ripe bananas at 25? C. with the minimum ventilation
required to keep the carbon dioxide below io per cent exhibited the
abnormalities in growth which are described as typical of the effects
of ethylene (Crocker, I929). Such effects, however, were only pro-
duced by restricting ventilation and at a high temperature. Similar
effects are produced by apples with ventilation rapid enough to keep
the carbon dioxide below o*5 per cent and at lower temperatures.
When unripe green bananas are used these effects are not produced.

80 IAI: IBIo IJ 1D
so

60

0
405 10 15

Days

Fig. 5. Production of carbon dioxide by bananas in air at I5? C. with con-

tinuous and discontinuous ventilation. Bananas II, continuous ventila-
tion throughout. Bananas I, no ventilation at periods A, B, C and D).

In parallel with these germinating-seed tests for the presence of

ethylene, the effects upon the subsequent respiratory activity of
bananas were studied after periods in which ventilation was restricted
to allow the accumulation of 1O per cent carbon dioxide. The data
shown in Fig. 5 refer to two samples from the same hand of fruit, one
sample was ventilated continuously and the other discontinuously.1
There is no sign of the action of any stimulating substance in the
pre-climacteric phase, but as soon as the fruit passes into the climac-
teric phase, a period of restricted ventilation accelerates the comple-
tion of the climacteric.
There is a strong probability therefore that bananas behave like
apples and begin to produce ethylene with the onset of the climacteric.
1 There is, of course, an initial high value due to the accumulated carbon
dioxide, but this is soon lost.

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 I74 R. GANE
The fact that treatment of the fruit with ethylene (i part
per million) or a temporary restriction of ventilation accelerales the
climacteric rise is interesting. In their work on apples and pears
Kidd & West (933) have shown on the one hand that restricted
ventilation or ethylene treatment will stimulate a climacteric change
even in very immature fruit, and on the other hand that if single
fruits are used (pears in this case), gathered sufficiently immature and
ventilated sufficiently rapidly, no climacteric ever occurred. They
argue that (i) ethylene or a substance with similar action is being
produced prior to the climacteric; (2) there must be a threshold value
of ethylene concentration in the cells necessary for stimulation;
(3) probably the rate of production of ethylene rises with age while
the threshold value for stimulation falls. The results of the experi-
ments described above suggest that the cells of the fruit do not all
reach the point of auto-stimulation simultaneously. If so, it should be
expected that at the stage when some have reached the critical point
and others are near it, an artificial raising of the concentration of
ethylene would cause an accelerated rise in the respiratory activity of
the whole fruit.
The active substance from ripe bananas is destroyed, or inacti-
vated, by ozone. A stream of air was passed over ripe bananas, and
then divided into two halves, each of which was led over a similar
sample of green bananas; into one stream a small oxonizer introduced
ozone at a concentration of from 70 to go parts per million. The
bananas exposed to this gas ripened at a much later date than the
other sample.
In this experiment the lenticels of the fruit exposed to ozone
became discoloured and dark green unfiltrated areas were formed
round them, which later became brown, and finally black. The
experiment was repeated in a modified form in which the comparison
instituted was between fruit ventilated with fresh air in the normal
way and fruit ventilated with air passed first over ripe bananas, then
through an ozonizer, and finally through a deozonizer.1 The behaviour
of the two samples was identical. The ozone had clearly removed
stimulating substances while the deozonizer had eliminated the
damaging effects of ozone.
With a view to surveying the sensitivity of the banana fruit to
ozone comparable samples of bananas were stored at I50 C. and
ventilated with air ahd with air containing a low concentration of
ozone. The ozone was produced by a small ozonizer built into the
1 A glass tube filled with scrap rubber.

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 Respiration of Bananas in Presence of Ethylene I75
inlet tube of the fruit container. The quantity of ozone produced
could be varied by altering the resistance in the mains leads of the
high tension transformer. The concentration of ozone is approxi-
mately constant over a range of air speeds. Any ozone in the air
stream issuing from the fruit container was removed by passing
through a glass tube containing scrap rubber.
Even the lowest concentration of ozone used, I 5 parts per million,
caused injury to the peel. A new observation was also made, namely,
that one of the effects of ozone is a retardation of ripening. This
retardation is probably due to a plugging of the stomata of the fruit
by the decomposition products formed by the action of ozone. The
data are plotted in Fig. 6. A similar retardation of ripening is shown
by fruit treated with hydrogen peroxide, dilute aqueous iodine and
coated with vaseline, and the conclusion has been drawn that the
effect is due to a blocking of the channels through which the gaseous
exchange takes place and consequently to an increased carbon dioxide
and diminished oxygen tension within the fruit.
Certain suggestions of possible value in the handling of fruit arise
from the facts established above. The first is that the problem of
" ships ripes " in banana-carrying vessels may find its partial solution
in the combined use of ozonizers and deozonizers for the treatment of
the air circulated through the holds and over the coolers by fans. The
second is that great care should be taken to avoid storing ripe fruit
or fruit that has been treated with ethylene, as is often the case with
early gathered oranges, in the same air with unripe fruit. The effect
upon bananas of the volatile products of a ripe apple (Bramley
Seedling) is shown in Fig. 7. Sufficient ethylene escapes from one
apple ventilated at the rate of 25 c.c. per min. to start the ripening
processes of bananas immediately. In view of the work of Kidd &
West (I932) on the ripening of apples, it is probable that premature
ripening of bananas also can be induced by the volatile products from
pears, peaches and tomatoes, but not by fully coloured oranges or
grapes.
The opinion is held in the banana-carrying trade that oranges and
citrus fruits in general are potential sources of danger and liable to
cause the bananas to ripen. Laboratory experiments do not confirm
this, though it should be remembered that, when citrus fruits have
been treated with ethylene, this gas would dissolve in the tissues of
the fruit and escape from solution again after the fruit was removed
from the conditioning rooms. There is no evidence as to the time
required for all the ethylene to be lost from these tissues.

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I76 R. GANE

40 40 p.p.m.
OZONEOO

20 AIR
AI

60 25-30 ppIm.

_40
420

40
0

20

5 10 15 20
Days

Fig. 6. The effect of ozone on the production of carbon dioxide

by bananas at I5? C.

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 Respiration of Bananas in Presence of Ethylene 177
Laboratory tests suggest that no ethylene-like substance is
produced by rotting banana stems, rotting bananas, yeast growing in
a culture solution, or by onions. None of these produce any substance
which acclerates the ripening of bananas. Using a different indicator,
either tomato shoots or potato shoots, Denny, Miller, Crocker,

80

b4

B6O
0
b~

40

A I
5 10 15
Days

Fig. 7. Effect of the volatile products from an apple on the production of

carbon dioxide by bananas at I50 C. Bananas II in fresh air throughout.
At A, air was passed over an apple (Bramley Seedling) and then over
bananas I. At B the apple was removed from the air stream.

Hitchcock & Zimmerman (I935) have shown that ethylenc is

produced by a variety of tissues including parts of flowers, e.g. petal
anthers; immature fruits; leaves; roots and tubers; but data is not
available to show whether the quantity produced is sufficient to
affect bananas.
SUMMARY

Evidence is given to show that ripe bananas produce ethylene.

The effect on seedlings of Pisum sativum is paralleled by low con-
centrations of ethylene.
An acceleration of ripening of unripe bananas by ethylene at a
concentration of I part per million is similar to that produced by

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 I78 R. GANE
the products of metabolism of ripe bananas. It would appear that
ethylene is a normal product of metabolism during the climacteric
when it acts as an autocatalyst.
Ethylene can be removed from air by ozone.
Ozone causes a retardation of normal ripening, probably by
gumming up the stomata. Similar retarding effects are shown by
fruit treated with hydrogen peroxide, iodine and vaseline.

REFERENCES

CROCKER, W. (i929). A delicate method of detecting illuminating gas in a

greenhouse. Prof. Pap. Boyce Thompson Inst. 1, No. ii, p. 8i.
CROCKER, W., HITCHCOCK, A. E. & ZIMMERMAN, P. W. (I935). Similarity in the
effects of ethylene and the plant auxins. Contr. Boyce Thompson Inst. 7,
23I.
DENNY, F. E. (1935). Testing plant tissue for emanations causing leaf epinasty.
Contr. Boyce Thompson Inst. 7, 34I.
DENNY, F. E. & MILLER, L. P. (I935). Production of ethylene by plant tissue
as indicated by the epinastic response of leaves. Con/i. Boyce Thompson
Inst. 7, 97.
GANE, R. (I935). The formation of ethylene by plant tissues, and its significance
in the ripening of fruits. J. Pomol. 13, 35I.
(I936). A study of the respiration of bananas. New Phytol. 35, 383.
KIDD, F. & WEST, C. (I932). Effects of ethylene and of apple vapours on the
ripening of fruits. Rep. Food Invest. Bd., Lond., p. 55.
(I933). The influence of the composition of the atmosphere upon the
incidence of the climacteric in apples. Rep. Food Invest. Bd., Lond., p. 5I.

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