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Carbon Dioxide Effects on Fruit Respiration ' Carbon Dioxide Effects on Fruit Respiration '. II. Response of Avocados, Bananas, & Lemons Roy E. Young, Roger J. Romani, & Jacob B. Biale Department of Plant Biochemistry, University of California, Los Angeles ,Most studies concerning the effects of the gas activity under air decreased slightly during storage. composition of the atmosphere surrounding fruit have Reduction of oxygen in the atmosphere surrounding been directed toward the response of the combined the fuit reduced the rate of respiration in proportion changes of oxygen an(d carbon dioxide. This was a to the oxygen concentration in the range of 21 to result of the economic need for methods of prolonging 5 % oxygen. Carbon dioxide evolution increased at the storage life by the use of modified atmospheres at oxygen levels below 5 %, indicating the similarity of temperatures above those which cause chilling injury. the behavior of lemons with other fruits characterized Several studies have been concerned with the effects by a critical oxygen concentration. The storage life of decreased oxygen alone. Parija (15) obtained was extended and the (lecomposition of chlorophyll minimum carbon dioxide prodluction by apples at 5 % in mature green lemons was delayed by lowered oxygen. Kidd and WVest (12) found an acceleration oxygen. of the onset of the climacteric in apples by oxygen The effect of added carbon dioxide on respiratorv tensions higher than in air. Singh et al. (19) re- activity of fruits at a particular oxygen concentration ported for mangos a critical oxygen level of 9.2 % has been studied little, largely due to technical diffi- below and above which carbon dioxide evolution in- culties. Limited data are available on the banana. creased. The studies of Claypool and Allen (6, 7) By the use of the katharometer method Gane (9) ob- with apricots, plums, peaches, and pears indicated a served a suppression of the climacteric and reduction decreased rate of CO., output at oxygen tensions be- of respiratory activity in an atmosphere of 10 % low air. For a more comprehensive treatment of this carbon dioxide and 10 % oxygen. Wardlaw (20) subject the recent review by Biale (3) is suggested. subjected unripe bananas to clifferent combinations of For the avocado, a fruit which exhibits a climac- oxygen and carbon dioxide. On the basis of gas teric pattern of respiration, it was shown by Biale analysis he concluded that there was a 50 X redluction (2) that low oxygen reduced the respiratory activity in rate of respiration over a wide range of 02 and during the preclimacteric period andl the (luration of CO2 concentrations as compared with air. His (le- the preclimacteric period was prolonged. Within the terminations were limited to green fruit andiwere range of 2.5 to 21 % oxygen the time required to done in a closed system in which the gaseous composi- reach the climacteric peak was extended in proportion tion could not be kept constant. The use of analytical to the decrease in oxygen tension and the intensity of methods described in the first paper (21) of this series respiration at the peak was reduced. No significant has enabled us to study the effect of carbon dioxide stimulation of respiration wvas observed by concentra- at several oxygen levels on the respiratory activity tions of oxygen above 35 %. However, the cumula- and storage behavior. The responses to CO., of tive carbon dioxide production from the time of pick- avocados, lemlons and bananas are reported at this ing until the climacteric peak was not changed by time. the treatment at any level of oxygen. In the case of the banana, another fruit with a climacteric pat- Materials & Methods tern, it was found by Kidd and West ( 11 ) that storage of the Fuerte variety were in in 2.5 and 5.0 % oxygen did not materially decrease Avocados picke(d the the rate of ripening. On the other hand, Leonard orchard of the Plant Biochemistry department at the of Los or were ob- (13) observed a reduction in CO, liberation by fruit University California, Angeles, stored in oxygen concentrations lower than air and tained from commercial orchards in southern Californ- no effect in concentrations higher than air. ia. Fruit was always picked in the morning and put Lemons did not exhibit the climacteric pattern of under treatment on the same day. Stems were cut close to the and respiration after picking when stored in air or at fruit those with detached stemls were oxygen tensions below that of air, as shown in the not used. Fruit was selected for uniformity of size sttudies of Biale and Young (5). The respiratory and 10 to 25 specimens were used per treatment. Bananas of the Gros Michel variety grown in Central America were obtained with the cooperation of the Consolidated Fruit Co. Fruit was picked in the three-quarter full stage, shipped to Los Angeles 1 Received revised manuscript Dec. 20, 1961. by boat, and were still dark green when received. 416 YOUNG ET AL.-C04 & FRUIT RESPIRATION 417 The transit period was normally 12 days, during about three days. Thereafter there was a gradual which the fruit was subjected to a temperature of rise to a peak of respiration which was 40 % below about 15 C and a relative humidity of about 90 %. the air control and was reached after 21 days com- Single bananas were separated from the hand and pared to 8 days for the air treatment. The peak was 15 to 20 bananas were tested per treatment. They followed by a decline and on the 23rd day the fruit were treated within one day after arrival of the boat. were transferred to air. There was no definite in- Lemons were obtained from commercial packing crease in oxygen uptake after the transfer, indicating houses in Los Angeles County. They were picked that the climacteric had really occurred and these on the basis of size rather than color, and dark green fruit softened in the normal manner. When fruit was fruit was selected for all experiments. Ordinarily treated with a gas mixture containing 10 % carbon one to two days elapsed between picking and the start dioxide and 21 % oxygen, there was essentially no of treatment. Fruit was washed, waxed, and sized change in respiratory activity for 23 days and the at the packing house. In one experiment fruit waxed rate was the same as that of the initial value of the with water wax containing 2,4-D was used. To check control. Upon transfer to air there was an increase the effect of the 2,4-D treatment a sample of the same in respiration, indicating that the climacteric had not pick as used in the experiment was removed from the occurred under the carbon dioxide treatment but had belt ahead of the waxer. Respiratory activity and occurred after transfer to air when the fruit softened color changes were unaffected by the 2,4-D wax treat- normally. ment. Button retention was somewhat better after The effect of increased carbon dioxide at the 10 % 2,4-D treatment. Composite samples of 30 to 50 oxygen level on the respiratory trend of avocados is fruit, uniform as to size and color, were selected for illustrated in figure 2. Fruit for this experiment was each treatment. from the same lot as that used for the experiment in Fruit was put in respiration jars described by figure 1. Treatment of 10 % oxygen alone reduced Biale and Shepherd (4), and gas mixtures passed the climacteric peak value to about 65 ml/kg/hr as continuously as described in paper I of this series compared to 85 ml/kg/hr for air, but delayed the peak (21). Respiratory activity was measured as ml oxy- only very slightly. When 5 % carbon dioxide with gen absorbed per kg fresh weight per hour. Oxygen 10 % oxygen was used, there was again a very grad- uptake was measured on the Model G-2 oxygen ana- ual rise in respiratory activity to a peak value of 48 lyzer as described previously (21). Oxygen uptake ml/kg/hr on the 21st day, which was the same as the measurements were made every 2 to 8 hours during peak value for 21 % 02 with 5 % CO2. The rise after the experimental period, depending upon the number transfer to air was probably due to the fact that one of experimental samples attached to the automatic fruit was infected with a fungal rot. It should be sampling system. Where daily values are reported, noted that the rise in respiration started only about the average of not less than four separate measure- two days after the 10 % oxygen control, but the peak ments is given. was delayed from 9 days for the control to 21 days For the chemical analysis of the lemons, three for the carbon dioxide treatment. Again 10 % car- random samples of seven fruit each were taken from bon dioxide in 10 % oxygen produced no change from each treatment. Juice of seven was pooled for analy- the preclimacteric minimum of the control and showed sis and the average of the three aliquots is reported. no change in respiration up to the 23rd day. Ripen- Ascorbic acid was determined by the method of ing was normal after transfer to air. Fruit under all Ramsey and Colichman (17). treatments was of good flavor, according to informal taste panels at the end of the storage period. There Results was some decay, particularly at the stem end of fruits - Fuerte Avocado. Fruit used for the experimental stored under carbon dioxide. We believe this to be data given in figures 1 and 2 was picked on May 20 a function of the length of storage rather than due and kept at 15 C throughout the experiment.
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