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Simultaneous Modelling of the Thermal Degradation Kinetics of Pectin Methylesterase in Lettuce (Lactuca Sativa L.) and Carrot (Daucus Carota L.) Extracts: Analysis of Seasonal Variation

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Simultaneous Modelling of the Thermal Degradation Kinetics of Pectin Methylesterase in Lettuce (Lactuca Sativa L.) and Carrot (Daucus Carota L.) Extracts: Analysis of Seasonal Variation Biosci. Biotechnol. Biochem., 71 (10), 2383–2392, 2007 Simultaneous Modelling of the Thermal Degradation Kinetics of Pectin Methylesterase in Lettuce (Lactuca sativa L.) and Carrot (Daucus carota L.) Extracts: Analysis of Seasonal Variation and Tissue Type y Daniel RICO, Ana B. MARTI´N-DIANA, Catherine BARRY-RYAN, Gary T. M. HENEHAN, and Jesu´sM.FRI´AS Postharvest Technology Unit, Dublin Institute of Technology (DIT), School of Food Science and Environmental Health, Faculty of Tourism and Food, Cathal Brugha Street, Dublin 1, Ireland Received September 5, 2006; Accepted June 28, 2007; Online Publication, October 7, 2007 [doi:10.1271/bbb.60484] The thermal degradation kinetics of pectin methyl- combined with other agents, has also been used to esterase (PME) from carrot and lettuce were studied. prevent loss of quality in fresh-cut vegetables, especially Fresh extracts were exposed to temperatures from 55 to in leafy vegetables (e.g., fresh-cut lettuce). The time of 70 C until the enzyme was inactivated. A model based exposure in these cases is very limited.7–11) Although on the presence of two forms of the enzyme, one active there are studies on carrot PME and the effect of and one non-active, is proposed. The natural variability LTB,6,12) no information on lettuce PME has been of the PME activity was taken into the model in the reported. form of normally distributed random effects. The com- The thermal kinetics of PME inactivation have been mon model parameters obtained (cleavage constant extensively studied, and there are several studies that (0:0395 Æ 0:0062 sÀ1), degradation constant (0:556 Æ propose different models that are being employed. In the 0:112 sÀ1), cleavage energy of activation (469 Æ 23 area of juice processing, thermal inactivation experi- kJ molÀ1) and degradation energy of activation (488 Æ ments on PME have generally yielded an apparent first- 18 kJ molÀ1)) show that the PME degradation kinetics of order kinetic mechanism working with both the juice the two vegetables can be explained with a single set of and the purified enzyme.13–17) However, Collet et al.18) parameters. (orange juice) and Anthon and Barrett19) (carrot and potatoes) have proposed two forms of enzyme with Key words: lettuce (Lactuca sativa L.); carrot (Daucus different thermal inactivation parameters. carota L.); pectin methylesterase; mixed- When working with purified PME, Nguyen et al.20,21) effects modelling; random effect and Castro et al.22) have proposed a similar two fraction biexponential model for the mild thermal and high- The textural changes in fruits and vegetables are pressure inactivation kinetics of purified strawberry, related to certain enzymatic and non-enzymatic proc- carrot and green pepper pectin-esterase. Similar param- esses. The enzymatic degradation of pectins is catalysed eters between the models were found in all studies, by pectin methylesterase (PME) and polygalacturonase leading to the hypothesis that the enzymes of different (PG).1) Pectin is first partially demethylated by PME, species might respond similarly to heat and pressure and later depolymerised by PG to polygalacturonic acid, stresses. The inactivation of commercial PME by a causing a loss of firmness.2) However, the controlled pulsed electric field treatment has been studied by Giner activation of PME results interesting to maintain textural et al.23) who proposed the Weibull model as the most properties, as the partial demethylation of pectins appropriate (over first-order kinetics) to describe the increases the cross-linking with cations.3) This effect is kinetics of the process by using a Bayesian model favoured in the case of carrots, as endogenous PG selection procedure. activity is almost non-existent.4) The stimulation of PME Finally, Tijskens et al.24) have found that the thermal activity at temperatures between 50 C and blanching inactivation kinetics of PME activity in carrot and conditions has been correlated with textural mainte- potatoes samples showed the characteristic biphasic nance.1,5–8) Low-temperature blanching (LTB), alone or shape of enzyme degradation kinetics and proposed a y To whom correspondence should be addressed. Fax: +353-1-4024495; E-mail: [email protected] Abbreviations: PME, pectin methylesterase; pro PME, non-active pro-pectin methylesterase form; LTB, low temperature blanching; ODE, ordinary differential equations; T, temperature of the extract; Tb, bath temperature; Tr, reference temperature; t, time; R, universal gas constant; kcle-ref , reference cleavage constant; kdeg-ref , reference degradation constant; keff , effective thermal conductivity; Ea-cle, cleavage energy of activation; Ea-deg, degradation energy of activation; AIC, Akaike information criterion; BLASTX, Basic Local Alignment Tool 2384 D. RICO et al. two-step reaction mechanism, with an active bound is going to be important when compared with the natural enzyme that was released into an active soluble enzyme, variability of the product. This is of particular impor- and used a two-step nonlinear regression procedure to tance when translating technological advances from the estimate the parameters and overcome the problems laboratory (a low variability scenario) to the retail sector with convergence created by experimental variability. A (a high variability scenario). similar model has been presented by the same authors in Statistical models to estimate components of variance an investigation on the blanching of peach pieces, and in non-linear experiments, namely in the form of non- the seasonal variability of the PME activity showed linear mixed-effect modelling, are tools that have comparable parameters between seasons.25) recently started to become available to the life scientist PME activity has recently been suggested as an working bench.32,33) This modelling technique has re- indicator of thermal processing in fruit and vegetables cently started to be used in the area of food phenomena on the basis of its apparent first-order inactivation kinetic modelling.23,34–37) A non-linear mixed effect kinetics and the fact that it is present in most of those model presents two components, a fixed effects part with products.26) From the foregoing evidence presented, a model that describes the deterministic knowledge of there is still a need to obtain more information on the how the average population behaves in respect to the thermal inactivation of this enzyme and the fundamental responses studied and a random-effect part which governing mechanism and to particularly study similar- represents the stochastic component of the model and ities in the thermal inactivation kinetics of the enzyme accounts for unexplained inter- and intra- individual between different vegetables. variability in the form of statistical distributions. PMEs are extracellular isoenzymes in higher plants The objective here was to study the thermal kinetics that are produced in an inactivated form by the plant cell of PME from carrot and lettuce and to propose a and then excreted to the cell wall where their perform common thermal inactivation kinetic model based on demethylesterification of the pectins. Micheli,27) based physiological assumptions and the high similarity of the on the work of Bordenave28) and Shinde,29) has reviewed two enzymes. The use of mixed-effect modelling to the classification of PME enzymes of different origin describe the thermal degradation of the enzyme from and proposed a model mechanism for PME isoform different tissues, vegetables and seasons, together with excretion that is generally agreed to happen in higher an estimation of the batch-to-batch variability inherent plants. With this mechanism the PME gene encodes not to a natural product was also part of the objective of this only the protein sequence, but also the pre- and pro- work. regions. The pre- region is a signal peptide that is generally cleaved early in the intracellular space. Pro- Materials and Methods PME is then secreted to the apoplasm via the Golgi cisternae, and only active mature PME is found in the Experimental design. The experiments were con- cell wall. It is speculated that the pro-PME sequence is ducted from March to September 2004. Iceberg lettuce not active or has very reduced activity and that the (Lactuca sativa L.) and carrots (Daucus carota L.) were enzyme is only activated with cleavage of the pro- used in the assays as models of leafy and root sequence by extracellular proteases. Under this mecha- vegetables. A study of the distribution of PME in the nism, vegetable extracts produced by tissue lysis and tissues was conducted. PME isolated from photosyn- assayed for PME activity possibly contain the two forms thetic and vascular tissues was measured in lettuce and of PME (pro-PME and mature PME). in the cortex (phloem) and core (xylem) of carrot. Many plants acclimatise in response to changes in temperature. This adaptation process produces physio- Raw material and sample preparation. Iceberg lettuce logical and biochemical changes in the plants.30,31) The (Lactuca sativa L.) and carrots (Daucus carota L.) were predictable variations in initial concentrations of PME obtained from a local supermarket, brought to the between two vegetable, season and tissue type, as well laboratory within 12 hours after the harvest and stored at as the natural product variability, may influence the 4 C before processing. The outer leaves (damaged or behaviour of this enzymatic degradation and affect the wilted) from the lettuce heads were discarded, and the post-harvest treatment efficiency. Tijskens et al.25) have core was excised with a stainless steel knife. The rest of studied the seasonal variation of the enzyme and its the leaves were separated and used. The carrots were variation between batches, and have already proposed peeled with a stainless steel peeler and cut into discs that the thermal degradation of PME in peaches was (5 mm thickness) for easy tissue identification. Distill- governed by a model in which the rate parameters were ed water at room temperature (18–20 C) was used to constant between seasons and only the initial enzyme wash the fresh-cut vegetables by immersion with species concentration in each of the batches varied.
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