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Journal of Food Engineering 87 (2008) 116123 www.elsevier.com/locate/jfoodeng

Respiration rate of banana fruit under aerobic conditions at dierent storage temperatures
S.D. Bhande a, M.R. Ravindra b, T.K. Goswami a,*
a

Department of Agricultural and Food Engineering, Indian Institute of Technology Kharagpur, West Bengal 721 302, India b NDRI Southern Campus Adugodi, Bangalore 560 030, India Received 10 August 2007; received in revised form 15 November 2007; accepted 20 November 2007 Available online 28 November 2007

Abstract The underlying principle behind storage techniques like controlled atmosphere storage and modied atmosphere packaging involves manipulation of respiration rate of the stored produce. However, since respiration rate is dependent on factors like storage temperature and composition of storage atmosphere, a mathematical approach to predict the respiration rate under given conditions would be an immense help in both design and process control of such storage systems. Experimental data were generated at temperatures 10, 15, 20, 25 and 30 C for banana fruit using the closed system method. The generated data were used to develop two dierent models based on regression analysis and enzyme kinetics, respectively. Both models were tested for its validity at 12 C. The models showed good agreement with the experimentally estimated respiration rate, though the model based on enzymatic kinetic with Arrhenius type temperature dependence was found to have a closer agreement than the other model studied. 2007 Elsevier Ltd. All rights reserved.
Keywords: Banana; Respiration rate; Enzyme kinetics; Regression coecient; Modeling

1. Introduction Fresh fruits always enjoy good market demand. However, fruits are perishable commodity, which generally have short shelf-life. The deterioration of fruit is associated with the physiological and biochemical activities and starts right from the moment and they are separated from the mother plant (Kader, 1986; Kays, 1991). Living cells of harvested plant products respire continuously, utilizing oxygen (O2) from the surrounding environment and releasing carbon dioxide (CO2). Respiration involves a series of oxidation reduction reactions where a variety of substances found within the cells are oxidized to CO2 (Kays, 1991). It is a major factor contributing to the post harvest losses of perishables. The post harvest respiratory response of fresh produce depends on the storage air temperature and its composition in terms of O2, CO2 and ethylene. The diminution of enzymatic activities by providing low temperature,
*

Corresponding author. Tel.: +91 3222 283122; fax: +91 3222 255303. E-mail address: tkg@agfe.iitkgp.ernet.in (T.K. Goswami).

low O2 and slightly high CO2, in general, reduces utilization rate of substrates (i.e. carbohydrates, organic acid and other reserves) and increases post harvest life of the fruit beyond its normal span (Kader, 1986; Saltveit, 2004). Hence, the storage temperature and storage air composition are major external factors, which can be maneuvered to keep the fruit in pristine condition as far beyond its normal season as practical. Measurement of respiration rate of produce at a particular storage temperature and gas composition is time consuming and needs special equipments for gas analysis. Various mathematical models have been developed to correlate the respiration rate with dierent storage parameters such as gas composition, i.e. O2 and CO2 and temperature. India is one of the leading producers of fruits accounting for about 10.4% of all fruits and nearly 40% of tropical fruits produced globally (FAO, 2005), with many indigenous varieties of mango, banana and other tropical fruits (Singh, 1993). However, not many references are present in literature regarding the respiration rate of these varieties. Knowledge of its respiration rate would go a long way in

0260-8774/$ - see front matter 2007 Elsevier Ltd. All rights reserved. doi:10.1016/j.jfoodeng.2007.11.019

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Nomenclature a b E Ea GCO2 GO2 Kic Kio Kmc Kmo N R RCO2 regression coecient regression coecient mean relative deviation modulus, % activation energy, kJ g1 mol1 carbon dioxide concentration, decimal oxygen concentration, decimal inhibition constant for CO2 evolution, % CO2 inhibition constant for O2 consumption, % CO2 MichaelisMenten constant for CO2 evolution, % O2 MichaelisMenten constant for O2 consumption, % O2 number of respiration data points universal gas constant, 8.314, kJ g1 mol1 K1 respiration rate, ml [CO2] kg1 h1 Rm RO 2 Rp T Tabs t Dt Vfr Vmc Vmo W model parameter for MichaelisMenten equation respiration rate, ml [O2] kg1 h1 respiration pre-exponential constant factor storage temperature, C absolute temperature, K storage time, h time dierence between two gas measurements, h free volume of the respiration chamber, ml maximum respiration rate for CO2 evolution, ml kg1 h1 maximum respiration rate for O2 consumption, ml kg1 h1 mass of fruit, kg

proper planning and implementation of programmes for storage, handling and transportation of these fruits. Banana (Musa paradisiaca L.) is a general term embracing a number of species or hybrids in the genus Musa of the family Musaceae (Zhang et al., 2005). Bananas are an important staple food that is critical to the nutritional and economic well being of millions of people throughout the developing world and are grown in about 120 countries (Olorunda, 2000). India is the worlds largest producer of banana with an annual production of 16.8 MMT (FAOSTAT, 2005). The more recent approach for modeling respiration rate by employing MichaelisMenten type equation is based on enzyme kinetics (Lee et al., 1991). This provides a simple description of respiration, based on the assumption that diusion and solubility of O2 and CO2 in plant tissue regulate reactions catalysed by allosteric enzymes. McLaughlin and OBeirne (1999) described the eect of CO2 on dry coleslaw mix respiration rate in three ways in competitive inhibition, uncompetitive inhibition and non-competitive inhibition. The uncompetitive model of inhibition tted best with the experimental results. However, due to the specicity of the model parameters, the same need to be identied and quantied for each fruit and vegetable product. The present study was undertaken with the specic objectives of (a) determination of respiration rate of banana (Musa paradisiaca cv martaman) by using closed system respirometer at dierent temperatures and (b) to develop and test a suitable mathematical model to predict the respiration rate of the fruit as a function of O2 and CO2 concentrations and the storage temperature. 2. Materials and methods 2.1. Banana Green and fresh banana fruit (Musa paradisiaca L.) of local popular variety martaman were obtained from com-

mercial sources. Fruits were washed to remove adhering dirt, and used for the investigations. Attention was paid to ensure that the fruit were of uniform size and mass and the maturity index was established using starch iodine index test (Dadzie and Orchard, 1997). The fruit samples used for the study varied from 0.12 to 0.16 m in length and 0.09 to 0.11 m circumference along the middle of the fruit length. The weight of the individual banana fruits chosen for the study was 0.110 0.015 kg. The physico-chemical parameters evaluated for the fruit samples are presented in Table 1. About 15 fruits from the harvested lot were randomly chosen for this evaluation. Firmness of the pulp was measured destructively using a texture analyzer (Model TA- XT2i, Stable Microsystems Ltd., UK) tted with a standard penetrometer probe (SS, 5 mm diameter). pH of the pulp was determined using a pH meter (Model lpH system 361, Systronics). The TSS was determined using a hand refractometer (Model No. PAL 1, ATAGO, Japan). The total sugar content and titratable acidity were measured using the method described in Ranganna (1987). The volume of banana fruit was determined by volume displacement method. About 200 fruits were used for the generation of respiration data and the fruit lot was divided across ve temperature levels, three experiments at each temperature.

Table 1 Physico-chemical parameters estimated for the green mature banana Parameter Total sugar (%) TSS (%) Firmness, N Titratable acidity (%) pH Value 1.1 0.3 17.7 0.2 5.07 0.51 0.4 0.05 5.7 0.2

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2.2. Gas exchange measurement Closed systems can provide a convenient way of characterizing respiration of fresh produce a single set of experiment (Hong and Kim, 2001). In this process the changes in O2 and CO2 concentrations within a sealed container, which result from respiration, could be measured directly. Air-tight respirometer chambers of size 0.1420 174 0.229 m made up of acrylic (Perspex) sheet was used. A NiCr thermocouple was inserted into one side of the respirometer and used to measure the storage temperature. Fruits were kept in respirometer from the open topside and were closed with the lid while inserting neoprene gasket in between. Lid was closed with nuts and bolts provided on the respirometer to make it airtight. A schematic sketch of the system used for generation of the respiration data is represented in Fig. 1. The average values obtained from the two sampling ports were considered as data. The system was kept in humidity control chamber (Digital System, Kolkata, India) which was maintained at the desired temperature with a tolerance limit of 0.2 C. Gas composition of respirometer was analyzed at regular intervals depending on the storage temperature of banana. Typically the interval chosen were 2 h for the temperatures at 30 and 25 C, 4 h at 20 and 15 C, 8 h at 10 C. Gas analysis was done till the CO2 concentration reached to 18%, till aerobic respiration (Hagger et al., 1992). Changes in the concentration of O2 and CO2 over a certain period of time were measured and used to estimate respiration rates. After an interval of time, gas samples of the container were analyzed for O2 and CO2 concentration. Mass of fruits taken during the experiment is shown in Table 2. Free volume of respirometer was the total volume of respirometer minus volume occupied by its content. The free volume of the chamber was measured by water displacement method. All measurements were conducted in triplicate and the data are represented as mean and standard deviation.

2.3. Gas analysis Storage gas sample was analyzed quantitatively for O2 and CO2 concentrations using a gas analyzer (MAP check combi, PBI-Dansensor). In this instrument, a zirconium sensor is used for O2 determination and an infrared detector is used to detect CO2. During spot measurement of gas composition inside the respirometer, the sampling needle was inserted through a silicon disc on the respirometer. After about 30 s, screen displays the CO2 and O2 percentages, which was developed due to respiration of banana. 2.4. Modeling and data analysis The experimental respiration rate was calculated from the concentration dierence, mass of produce and free volume of chamber. The respiration rates in terms of O2 and CO2 at a given temperature are calculated using the following equations as given by Kays (1991):   GO2 t GO2 t1 V fr 1 R O2 Dt W   GCO2 t1 GCO2 t V fr RCO2 2 Dt W where RO2 is the respiration rate, ml [O2] kg1 h1, RCO2 is the respiration rate, ml [CO2] kg1 h1, GO2 and GCO2 are the gas concentrations for O2 and CO2, respectively, t is the storage time in hours, Dt the time dierence between two gas measurements, Vfr is the free volume of the respiration chamber, ml and W is the mass of the fruit in kg. Two dierent approaches were attempted to model the respiration rate based on the experimental data as outlined below. 2.4.1. Model 1 A regression function is often used to t the data of gas concentration versus time, and the respiration rate at given time is determined from the rst derivative of the regression function (Cameron et al., 1989; Hagger et al., 1992; Mahajan and Goswami, 2001). By using the generated experimental respiration data, a two parameter, non-exponential equation similar to the Peleg model proposed for moisture sorption curves (Peleg, 1988) was tted to O2 concentration and CO2 concentration at dierent storage periods using regression analysis (Systat, SPSS, Chicago). A similar model has been applied for respiration data of apples by Mahajan and Goswami (2001). The resultant regression equations for O2 consumption and CO2 evolution are shown in Eqs. (3) and (4) to determine the values of the coecients:   t GO2 0:21 3 at b t 4 GCO2 at b

To gas analysis

Points of sampling of headspace gas Gas tight chamber for housing the fruit

to temperature sensor

Fig. 1. Schematic sketch for the closed system method for generation of respiration data.

S.D. Bhande et al. / Journal of Food Engineering 87 (2008) 116123 Table 2 Free volume of respirometer and weight of banana taken for generating the respiration dataa Storage temperature (C) 10 15 20 25 30 Weight of fruits (W) (kg) 1.430 1.433 1.437 1.434 1.409 (0.07) (0.11) (0.12) (0.11) (0.02) Free volume of respirometer (VfR) (ml) 4076.78 4073.25 4068.16 4071.67 4101.88 (72.46) (121.71) (133.21) (121.21) (22.10)

119

this study. The respiration data for O2 and CO2 were obtained by the closed system respirometer kept at 12 C. Free volume of respirometer and mass of banana taken for verication of model were 3239 ml and 1.384 kg, respectively. The experimental respiration rates for banana at dierent combinations of O2 and CO2 at 12 C temperature were determined using Eqs. (1) and (2), respectively. 2.6. Goodness of t This parameter is widely adopted throughout the literature to evaluate the goodness of t of mathematical expression. Moduli below 10% are indicative of reasonably good t, 1020% fairly good t and 2030% not satisfactory t for all practical purposes. The goodness of t between predicted and experimental respiration rates was obtained by calculating the mean relative percentage deviation modulus (McLaughlin and OBeirne, 1999) as given in Eq. (10). In general, lower the modulus better is the agreement between experimental values and predicted values: E
N 100 X jRexp Rpre j N 1 Rexp

Value in parenthesis is the standard deviation. a Values are average of three replications.

The rate of change of gas concentration was determined from the rst derivative of the regression functions as outlined in Eqs. (5) and (6): dGO2 atat b2 at b1 dt dGCO2 2 1 atat b at b dt 5 6

At any given time, the respiration rate of the sample was then calculated by substituting the values of dG/dt obtained from Eqs. (5) and (6) in Eqs. (1) and (2), respectively. The temperature dependence of the model coecients a and b were estimated by linear interpolation between the two temperatures. 2.4.2. Model 2 A model was developed using principles of enzyme kinetics with uncompetitive inhibition (Lee et al., 1991; Peppelenbos and Vant Leven, 1996) for predicting the rate of respiration of banana fruits. The model has three parameters viz., Vm, Km, and Ki for both O2 consumption and CO2 evolution as shown in Eqs. (7) and (8). The model parameters were determined using the experimental respiration data. The model parameters obtained for banana were then correlated at dierent temperatures using Arrhenius plot as shown in Eq. (9): R O2 V mo GO2 h i GCO2 K mo 1 K G O2 io 7 8

10

where E is the mean relative deviation modulus in %; N is the number of respiration data points; Rexp is the experimental respiration rate in ml kg1 h1 and Rpre is the predicted respiration rate in ml kg1 h1. 3. Results and discussion The respiration data corresponding to dierent temperatures indicated that as the temperature increased the respiration progressed at a faster rate. As an example, a typical data set for 20 C is presented in Fig. 2 Also, for each temperature, the respiration rate appeared to slow down as time progressed. The same trend is clearly reected in Fig. 3 by the gradual reduction in the slope of the data points with time. This could be attributed to the decreasing O2 and increasing CO2 concentration in the gaseous environment with time. At 10 C, the initial respiration rate recorded was about 15.81 and 15.37 ml kg1 h1 for RO2 and RCO2 , respectively.

Gas Concentration, %

V mc GO2 h i GCO2 K mc 1 K G O2 ic   E a Rm Rp exp RT abs RCO2 2.5. Verication of the model

25 20 15 10 5 0 0 10 20 30 40 50 60 70
GO2 GCO2

Model predicted respiration rates of banana were veried with experimental respiration rates at 12 C storage temperature. This temperature was chosen as it is the commonly recommended temperature for storage of banana (Kader, 2003; Tan, 2006) and the temperature was within the range of the experimental temperature levels used for

Time, h

Fig. 2. Experimental data for O2 consumption and CO2 evolution at 20 C.

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1.5

Comparatively, the initial respiration rates at 30 C estimated were 36.85 and 40.85 ml kg1 h1, respectively. Also, the gas concentrations reached their upper limit (CO2 > 18%) after 88 and 68 h at 10 and 15 C, respectively. While the same gas concentration was achieved within 48 and 32 h for experiments at 25 and 30 C, respectively. This trend is expected as temperature has been identied as the most important factor inuencing respiration behavior of fruits (Saltveit, 2004). The respiratory quotient (RQ), as estimated as the ratio of CO2 evolved to the O2 consumed, was found to vary with time for all the temperature levels studied. The average RQ was also found to vary as the temperature increased as indicated in Fig. 4. A similar temperature dependence of RQ has been reported for mangoes by Nakamura et al. (2004). 3.1. Parameter estimation for the dierent models 3.1.1. Model 1 The regression coecients a, and b of Eqs. (3) and (4), and correlation coecients (R2) at dierent storage temperatures are shown in Table 3. The above regression functions described the experimental data very well (R2 > 0.992) for banana. It can be inferred from the values of the regression coefcients a and b as shown in the Table 3 that both the parameters were inuenced by the storage temperature and that coecient b was more inuenced by temperature than coecient a. From its analogy to the Peleg model, a, which is the rate constant could physically relate to the

RCO2 / RO2, decimal

1.25 1 0.75 0.5 0.25 0 5 10 15 20 25

o

30

35

Temperature, C

Fig. 4. Changes in value of respiration quotient with temperature.

consumption or evolution rate of the gases at the very beginning, i.e. when t = 0. Where as, b, by the same analogy, is the capacity constant of the model. The value of b would imply the respective gas contents attainable by the system when time t is equal to innity. The values of a and b obtained in this study are similar to the ranges for the parameters reported for respiration of apples by Mahajan and Goswami (2001) For any unknown temperature, the value of a and b can be determined by linear interpolation of the known values. The corresponding values of a and b when substituted in Eqs. (5) and (6) estimate dGO2 /dt and dGCO2 /dt, respectively. The respiration rates for the given temperature can then be determined using Eqs. (1) and (2). 3.1.2. Model 2 By tting the experimental data at each temperature the model parameters of Eqs. (7) and (8) were determined using multiple regression analysis. The regression constants were then used to yield the values of the model constants, which are given in Table 4 along with the corresponding R2 value. The values in Table 4 show that the model parameters were dependent on the storage temperature. The coecients of determinations was found to be more than 0.972 indicating that the relationship between respiration rate O2 and CO2 concentrations tted well with the uncompetitive inhibition enzyme kinetics model. Dependence on temperature as estimated by the Arrhenius relation Eq. (9) was determined by plotting the log values of the model parameters against the inverse of corresponding temperature in absolute units. The resulting linear plot is shown as Fig. 5. A close observation of the plot showed that the plot actually appeared to have two regions. Similar results were also observed by Song et al. (1992) and Jaime et al. (2001). This was attributed to a change in the rate limiting enzymatic action, as each enzyme may be aected dierently by changes in temperature. Although, an exact transition temperature is dicult to establish from Fig. 6, it appears to be in agreement with the observation of Song et al. (1992) that the transition takes place in the vicinity of 15 C The values for the activation energy and the pre-exponential factor for each parameter as determined as the

30 25
Experimental Predicted Model 1 Predicetd Model 2

RO2 , ml/kg h

20 15 10 5 0 0 10 20 30

40

50

60

Time, h
30 25

RCO2, ml / kg h

20 15 10 5 0 0 10 20

Experimental Predicted Model 1 Predicted Model 2

30

40

50

60

Time, h

Fig. 3. Average of experimentally estimated and predicted respiration rates for banana at 20 C.

S.D. Bhande et al. / Journal of Food Engineering 87 (2008) 116123 Table 3 Values of regression coecients for model 1a
ln(Vm) (ml/kg h)
4.5
Vm(O2)

121

Storage temperature of banana (C)

Respiration expression in terms of

Regression coecients a b 203.88 (6.25) 196.84 (9.87) 145.78 (5.86) 134.82 (12.95) 161.65 (4.16) 110.65 (0.80) 93.23 (4.28) 82.68 (15.06) 71.47 (3.08) 63.04 (7.25)

R2

4 3.5 3 2.5 2 1.5 0.0032

Vm(CO2)

10

O2 consumption CO2 evolution

3.25 (0.19) 3.23 (0.15) 3.52 (0.25) 3.48 (0.11) 3.63 (0.20) 3.52 (0.13) 3.43 (0.12) 3.44 (0.04) 3.39 (0.15) 3.43 (0.09)

0.997 0.997 0.994 0.992 0.992 0.992

15

O2 consumption CO2 evolution

0.0033

0.0034

0.0035

0.0036

1/T (K )
2.9 2.7

-1

20

O2 consumption CO2 evolution

ln(Km) (O2 %)

2.5 2.3 2.1 1.9 1.7 1.5 0.0032 0.0033 0.0034 0.0035 0.0036
-1

25

O2 consumption CO2 evolution

0.998 0.994 0.999 0.999

Km(O2) Km(CO2)

30

O2 consumption CO2 evolution

Value in parenthesis is the standard deviation. a Values are average of three replications.
2.7 2.5
ln(Ki) (CO2 %)

1/T (K )

Table 4 Model parameters for uncompetitive inhibition enzyme kinetics for dierent storage temperaturesa (model 2) Storage temperature (C) 10 Respiration expressed in terms of O2 CO2 15 O2 CO2 20 O2 CO2 25 O2 CO2 30 O2 CO2 Vm (ml kg1 h1), 16.36 (2.09) 26.13 (2.43) 21.41 (1.65) 34.22 (2.45) 26.17 (2.34) 38.57 (4.25) 31.18 (2.36) 44.31 (4.01) 43.76 (3.06) 56.43 (1.83) Km, % O2 6.4 (0.94) 10.42 (1.29) 7.32 (2.3) 11.9 (2.32) 8.42 (1.89) 12.8 (1.99) 8.79 (1.71) 13.16 (0.51) 8.92 (1.78) 14.12 (2.05) Ki, % CO2 10.11 (0.54) 12.75 (1.51) 8.51 (1.05) 11.31 (1.37) 8.13 (2.54) 9.36 (0.85) 5.51 (1.25) 10.03 (1.37) 6.13 (0.55) 7.79 (1.23) R2

2.3 2.1 1.9 1.7

Ki(O2) Ki(CO2)

0.970 0.975 0.982 0.997 0.960 0.970 0.990 0.982 0.981 0.987

1.5 0.0032

0.0033

0.0034
1/T (K )
-1

0.0035

0.0036

Fig. 5. Arrhenius relation for dierent model parameters of enzyme kinetics.

6 imply that the inhibition by CO2 varied negatively to temperature increase. Also, the high values of Vm and Km should indicate the strong inuence of temperature on the respiration kinetics. By using these constants the values of the model parameters for the MichaelisMenten equation (Eqs. (7) and (8)) can be estimated at any temperature, in the range of this study and then the respiration rate at the same temperature predicted using Eqs. (7) and (8). 3.2. Verication of the models Since the respiration rates were generated at temperature between 10 and 30 C with the step of 5 C, it was necessary to verify whether the developed models were capable of predicting the respiration rates at any temperature within this domain of experimental temperatures. Accordingly, the respiration rates predicted by the models model

Value in parenthesis is the standard deviation. a Values are average of three replications.

slope and Y-axis intercept of the respective plots are shown in Tables 5 and 6. The negative values of Ki given in Table

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16

14 12 10 8 6 4 2 0 0 20 40

Experimental Predicted Model 1 Predicted Model 2

60

80

dence based on Arrhenius law against the experimentally determined respiration rate at 12 C is depicted in Fig. 6. The mean relative deviation moduli (Eq. (10)) between the respiration rates of banana at 12 C predicted by regression analysis (model 1) and that obtained through experiments were 10.43% and 11.57% for O2 consumption and CO2 evolution, respectively, while the same value by the model based on enzyme kinetics (model 2) were found to be 8.38% and 7.58% for O2 consumption and CO2 evolution, respectively, indicating a better agreement between the latter model and the experimental results. 4. Conclusion

Respiration rate, ml O 2 / kg h

Time, h
18

Respiration rate ml CO2 / kg h

16 14 12 10 8 6 4 2 0 0 10 20 30 40 Time, h

Experimental Predicted Model 1 Predicted Model 2

50

60

70

80

Fig. 6. Experimentally estimated and predicted respiration rates for banana at 12 C.

were veried with the experimental respiration rate at 12 C storage temperature. For model 1, the values of the regression coecient a and b at 12 C were estimated by the linear interpolation of the values of the same coecients at 10 and 15 C shown in Table 3. By using activation energy and pre-exponential factor from Table 6, the model parameters for enzyme kinetics at 12 for both O2 consumption and CO2 evolution were calculated and the respiration rate at 12 C was predicted using Eqs. (7) and (8). The comparison of regression analysis (model 1) and enzyme kinetics model (model 2) with temperature depen-

Respiration rates determined by closed system method were found to vary inversely with increase in storage temperature in the range of 1030 C for the variety of banana studied. This could be attributed to the diminishing concentrations O2 and proportional increase in CO2 concentrations in the respirometer as the storage time progresses. The relationship between the concentration of O2 and CO2 concentrations in the storage space with storage time was expressed as a regression function at the dierent temperatures with R2 values >0.992 for the t. The eect of the gas concentrations on the respiration rate was also found to t the uncompetitive inhibition enzyme kinetics for all the temperature levels studied. The parameters for this model were found to follow the Arrhenius relation for temperature dependence. Both models exhibited a trend, which was in good agreement with the experimentally determined respiration rate. Validation of the models at 12 C yielded mean relative deviation moduli values of 10.43% and 11.57% for O2 consumption and CO2 evolution, respectively, for the model based on regression analysis. While, the model based on enzyme kinetics resulted in mean relative deviation moduli values of 8.38% and 7.58% for O2 consumption and CO2 evolution, respectively. The results demonstrate that the models applied well to predict the respiration rate of banana (cv martaman), in the temperature range studied.

Table 5 Slope (Ea/R) and Y-axis intercept (ln Rp) of Arrhenius relation for dierent model parameters of enzyme kinetics Vm O2 Slope Y-axis intercept R2 3504.9 15.26 0.985 CO2 2534.8 12.14 0.975 Km O2 2167 9.45 0.905 CO2 1243.4 7.01 0.975 Ki O2 2441.9 6.32 0.934 CO2 1831.8 3.92 0.944

Table 6 Activation energy and pre-exponential factor of Arrhenius type equation for dierent model parameters of uncompetitive inhibition Parameters for Arrhenius equation Vm O2 Ea kJ g1 mol1 Rp 28.15 4.2 106 CO2 20.08 1.8 105 Km O2 17.03 1.2 104 CO2 11.35 1.2 103 Ki O2 21.29 1.7 103 CO2 14.25 1.8 102

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