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We performed coproparasitological testing of free-living golden-headed lion tamarins, Leontopithecus chrysomelas, using the Hoffmann-Pons-Janner method. In total, we collected 118 samples from ten groups: four living in Federal Protected Area and six living in Non-Protected Areas of cocoa farms. Eggs from parasites of the Acanthocephala phylum and Spiruridae, Ancylostomatidae, Ascarididae and Oxyuridae families were identified, as well as the genus Strongyloides (Nematode: Strongyloididae) and phylum Apicomplexa. This is the first description of infection with coccidian, Trichuridae family and Strongyloides spp. in L. chrysomelas. A total of 48% (n= 57) of the animals were infected and the highest prevalence (37.2±SD 8.72, n = 44) was for Acanthocephalidae, followed by Spiruridae (8.5±SD 5.03, n = 10). There was no difference in parasite prevalence by age classes or sex. However, we found higher diversity and prevalence of parasites in animals living in the Federal Protected Area. These results suggest that intestinal parasites may be influenced by environmental factors, such as the management of the areas where the animals live, in addition to the feeding behavior of L. chrysomelas and distinct transmission strategies of parasites. The combination of ecological and demographic data combined with parasitological studies may contribute to conservation programs for this species.

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Abstract We performed coproparasitological testing of free-living golden-headed lion tamarins, Leontopithecus chrysomelas, using the Hoffmann-Pons-Janner method. In total, we collected 118 samples from ten groups: four living in Federal Protected Area and six living in Non-Protected Areas of cocoa farms. Eggs from parasites of the Acanthocephala phylum and Spiruridae, Ancylostomatidae, Ascarididae and Oxyuridae families were identified, as well as the genus Strongyloides (Nematode: Strongyloididae) and phylum Apicomplexa. This is the first description of infection with coccidian, Trichuridae family and Strongyloides spp. in L. chrysomelas. A total of 48% (n= 57) of the animals were infected and the highest prevalence (37.2±SD 8.72, n = 44) was for Acanthocephalidae, followed by Spiruridae (8.5±SD 5.03, n = 10). There was no difference in parasite prevalence by age classes or sex. However, we found higher diversity and prevalence of parasites in animals living in the Federal Protected Area. These results suggest that intestinal parasites may be influenced by environmental factors, such as the management of the areas where the animals live, in addition to the feeding behavior of L. chrysomelas and distinct transmission strategies of parasites. The combination of ecological and demographic data combined with parasitological studies may contribute to conservation programs for this species.

Resumo Foram realizados testes coproparasitológicos de micos-leões-dourados de vida livre, Leontopithecus chrysomelas, usando-se o método de Hoffmann-Pons-Janner. No total, foram coletadas 118 amostras de dez grupos: quatro grupos residentes em Área de Conservação Federal e seis grupos em Área não protegida de fazendas de cacau. Ovos de parasitas do filo Acantocephala e das famílias Spiruridae, Ancylostomatidae, Ascarididae, Oxyuridae foram identificados, bem como o gênero Strongyloides (Nematoda: Strongyloididae) e o filo Apicomplexa. Esta é a primeira descrição de infecção de coccídeos, família Trichuridae e Strongyloides spp. em L. chrysomelas. Um total de 48% (n = 57) dos animais estavam parasitados e a maior prevalência (37,2 ±DP 8,72, n = 44) foi para Acanthocephalan, seguido por Spiruridae (8,5±DP 5,03, n = 10). Não houve diferença na prevalência do táxon de parasita por idade ou sexo. No entanto, foi encontrada maior diversidade e prevalência de parasitas em animais que vivem na Unidade de Conservação Federal. Esses resultados sugerem que os parasitas intestinais podem ser influenciados por fatores ambientais, como o manejo das áreas, além do comportamento alimentar de L. chrysomelas e distintas estratégias de transmissão dos parasitas. A combinação de dados ecológicos e demográficos com estudos parasitológicos podem contribuir para programas de conservação dessa espécie.

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Abstract Both anthropogenic actions and abiotic parameters, such as rainfall, temperature and photoperiod, can affect fruit and flower availability for animals, which consequently affects nutritional status and thus animals' health. Herein, we investigated whether abiotic factors are related to changes in fruit availability that can lead to changes in feeding behavior and, consequently, in endoparasite load and general health status in two groups of golden-headed lion tamarins (Leontopithecus chrysomelas) living in degraded fragments of Atlantic forest in Southern Bahia, Brazil. We detected that there was a high variation in availability of ripe fruits throughout the year, with lower availability occurring at the end of spring and beginning of summer. Despite this, there was no difference in tamarins' general health status, body mass and blood counts between seasons. This is probably because during native fruit scarcity, the tamarins eat cultivated species, such as banana (Musa spp.) and jackfruit (Artocarpus heterophyllus). Temperature and daylength were negatively correlated with golden-headed lion tamarin endoparasite loads. Contrary to our expectations, endoparasite loads are not linked to fruit scarcity and consequent changes in feeding behavior. Nevertheless, we found higher parasite diversity in the group of golden-headed lion tamarins that occupied the smallest home range. The smaller the area available, the greater the contact with parasites the animal will have, as they are forced to travel constantly along the same routes in the forest, increasing infection risk and re-infection rates. Our results highlight how animals' health is associated with environmental health as well as the need for constant monitoring to ensure the effective conservation of endangered species, such as the golden-headed lion tamarin.

Resumo Parâmetros abióticos, como precipitação, temperatura e fotoperíodo, podem afetar a disponibilidade de frutos e flores para os animais, o que consequentemente afeta o estado nutricional e a saúde dos animais. Neste estudo, investigamos se fatores abióticos estão relacionados com alterações na disponibilidade de frutos, o que pode levar a mudanças no comportamento alimentar e, consequentemente, na carga de endoparasitas e estado de saúde geral em dois grupos de mico-leão-de-cara-dourada (Leontopithecus chrysomelas) que vivem em fragmentos degradados de Floresta Atlântica no sul da Bahia, Brasil. Detectamos que houve grande variação na disponibilidade de frutos maduros ao longo do ano, com menor disponibilidade no final da primavera e início do verão. Apesar disto, não houve diferenças no estado geral de saúde, na massa corporal ou nas contagens de células sanguíneas dos animais entre as estações do ano. Isto provavelmente ocorreu porque durante a escassez de frutos nativos, os micos comem espécies cultivadas, tais como a banana (Musa spp.) e jaca (Artocarpus heterophyllus). A temperatura e a duração do dia foram negativamente correlacionadas com a carga de endoparasitas de mico-leão-de-cara-dourada. Contrário ao previsto, a carga de endoparasitas não está ligada à escassez sazonal de frutos e consequentes mudanças no comportamento alimentar. Entretanto, encontramos maior diversidade de endoparasitas no grupo de mico-leão-de-cara-dourada que usou uma área de vida menor. Quanto menor a área disponível, maior o contato com parasitas, porque os micos são forçados a se deslocar constantemente pelas mesmas rotas na floresta, aumentando o risco de infecção e as taxas de reinfecção. Nossos resultados destacam como a saúde dos animais está associada à saúde ambiental, bem como a necessidade de monitoramento constante para a conservação eficaz das espécies ameaçadas de extinção, como o mico-leão-de-cara-dourada.

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Plant species models are among the available tools to predict the future of ecosystems threatened by climate change, habitat loss, and degradation. However, they suffer from low to no inclusion of plant dispersal, which is necessary to predict ecosystem evolution. A variety of seed dispersal models have been conceived for anemochorous and zoochorous plant species, but the coupling between vegetation models and seed dispersal processes remains rare. The main challenge in modelling zoochoric dispersal is simulating animal movements in their complex habitat. Recent developments allow straightforward applications of hidden Markov modelling (HMM) to animal movements, which could ease generalizations when modelling zoochoric seed dispersal. We tested the use of HMM to model seed dispersal by an endangered primate in the Brazilian Atlantic forest, to demonstrate its potential simplicity to simulate seed dispersal processes. We also discuss how to adapt it to other species. We collected information on movement, fruit consumption, deposition, and habitat use of Leontopithecus chrysomelas. We analysed daily trajectories using HMM and built a deterministic Model Of Seed Transfer (MOST), which replicated, with good approximation, the primate's movement and seed deposition patterns as observed in the field. Our results suggest that the dispersal behaviour and short daily-trajectories of L. chrysomelas restrict the species' role in large-scale forest regeneration, but contribute to the prevalence of resource tree species locally, and potentially maintaining tree diversity by preventing local extinction. However, it may be possible to accurately simulate dispersal in an area, without necessarily quantifying variables that influence movement, if the movement can be broken down to step-length and turning angles, and parametrised along with the distribution of gut-transit times. For future objectives, coupling MOST with a DVM could be used to test hypotheses on tree species survival in various scenarios, simulating regeneration and growth at regional scales by including data on main dispersal agents over the area of interest, distribution of tree species, and land use data. The principal advantage of the MOST model is its functionality with data available from the literature as the variables are easy to parametrise. We suggest using the coupled model to perform experiments using only available information, but varying the numbers and species of seed dispersers, or modifying land cover or configuration to test for possible thresholds preventing the extinction of selected tree species.

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Individual behavioral differences may influence how animals cope with altered environments. Depending on their behavioral traits, individuals may thus vary in how their health is affected by environmental conditions. We investigated the relationship between individual behavior of free-living golden-headed lion tamarins (Leontopithecus chrysomelas) responding to a novel object (to assess exploration-avoidance), and their habitat use and health status (endoparasitism; clinical measures: biometric data, heart rate, respiratory frequency, and temperature; fecal glucocorticoid metabolites). As parasite transmission can be affected by individual variation in social contact and social grooming, we also evaluated whether more sociable individuals show higher endoparasite loads compared with less sociable animals. Four groups living in landscapes with different levels of human disturbance were investigated: two in degraded forest fragments in an agricultural matrix (DFAM-higher disturbance), and two in a cocoa agroforestry system (cabruca-lower disturbance) in the Atlantic forest of South Bahia, Brazil. Using a subjective ratings approach, highly correlated adjective descriptors were combined to produce z-score ratings of one derived variable ("confidence"), which was selected to characterize the tamarins' exploration/avoidance responses during a novel object test. The higher the confidence score, the longer female tamarins spent foraging for prey independent of landscape, and the greater their body mass independent of sex and landscape. Only DFAM individuals showed intestinal parasite infection. Endoparasite loads were positively correlated with the number of grooming partners, suggesting an association between social grooming and transmission (more groomers = more endoparasites). Individual behavior, including in a test situation, may thus have some predictive value for behavior in a free-living context, and for its health consequences.

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The current extinction and climate change crises pressure us to predict population dynamics with ever-greater accuracy. Although predictions rest on the well-advanced theory of age-structured populations, two key issues remain poorly explored. Specifically, how the age-dependency in demographic rates and the year-to-year interactions between survival and fecundity affect stochastic population growth rates. We use inference, simulations and mathematical derivations to explore how environmental perturbations determine population growth rates for populations with different age-specific demographic rates and when ages are reduced to stages. We find that stage- vs. age-based models can produce markedly divergent stochastic population growth rates. The differences are most pronounced when there are survival-fecundity-trade-offs, which reduce the variance in the population growth rate. Finally, the expected value and variance of the stochastic growth rates of populations with different age-specific demographic rates can diverge to the extent that, while some populations may thrive, others will inevitably go extinct.

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This study evaluated the genetic structure of wild populations of the endangered primate, Leontopithecus chrysomelas. We tested the assumption that populations of L. chrysomelas, given their larger population size and a higher degree of habitat continuity, would have higher genetic diversity and less genetic structuring than other lion tamarins. We used 11 microsatellites and 122 hair samples from different locations to assess their genetic diversity and genetic structure, and to make inferences about the isolation by distance. The overall expected heterozygosity (0.51 ± 0.03) and the average number of alleles (3.6 ± 0.2) were relatively low, as is the case in other endangered lion tamarins. Genetic clustering analyses indicated two main clusters, whereas the statistical analyses based on genotype similarities and Fst suggested further substructure. A Mantel test showed that only 34% of this genetic differentiation was explained by the linear distance. In addition to linear distance, structural differences in the landscape, physical barriers and behavioural factors may be causing significant genetic structuring. Overall, this study suggests that these populations have a relatively low genetic diversity and a relatively high population genetic structure, putting in question whether the presence of agroforest systems (known locally as cabruca) is enough to fully re-establish functional landscape connectivity.

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In a landscape fragmented by agriculture, the extent to which forest-dwelling primates can use the matrix between fragments can be critical for their long-term survival. So far, the golden-headed lion tamarin (Leontopithecus chrysomelas), an endangered primate inhabiting the Atlantic Forest of south Bahia, is only known to use shaded cacao (Theobroma cacao) agroforests within the matrix. We report on the use of a rubber plantation by a group of golden-headed lion tamarins between August 2013 and January 2014. The group used the rubber plantation on 16 of the 22 observation days (73 %), and we recorded behaviours such as eating, grooming and sleeping, consistent with the use of the area as a home range. We also observed associations with Wied's marmosets (Callithrix kuhlii). The locations of group sightings were not uniformly spread across the entire area of the rubber plantation, suggesting preferred use of certain areas. The presence of resources such as jackfruits (Artocarpus heterophyllus) and epiphytic bromeliads may be attracting both species to these plantations. In addition to shaded cacao plantations, rubber plantations with the appropriate structure may be a viable option for increasing forest connectivity for both species in south Bahia, reconciling economic rubber production with primate conservation.

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Parasite prevalence and abundance are important factors affecting species' conservation. During necropsies on a free-living golden-headed lion tamarin ( Leontopithecus chrysomelas ) and two Wied's marmosets ( Callithrix kuhlii ) in the Atlantic Forest of southern Bahia, Brazil, we collected a large number of adult intestinal parasites that we identified as Prosthenorchis elegans. This parasite is pathogenic for neotropical primates. Prosthenorchis spp. infestation is influenced by diet with increased risk of exposure from ingesting invertebrate intermediate hosts. The biological similarities and sympatric nature of these two nonhuman primates support that they may harbor similar infectious and parasitic agents.

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Little is known about frugivory of epiphytes. We investigated diurnal frugivores consuming Aechmea depressa, an endemic species from Southern Bahia, Brazil, through 67 hours of observation on three fruiting individuals. Infructescences were visited for 2 percent of the time and only by golden-headed lion tamarins (GHLTs; Leontopithecus chrysomelas). Aechmea depressa invests on physical defense to retain fruits for prolonged periods, however, GHLTs used their small size and dexterity to remove rigid bracts and extract fruits from the infructescence. Prior studies indicate GHLTs disperse Aechmea seeds. Given the low visitation rates and probability that few frugivores successfully manipulate A. depressa fruits, we suggest that GHLTs are extremely important to maintaining the populations of this bromeliad species.

Pouco se sabe sobre a frugivoria em plantas epífitas. Nós investigamos os frugívoros diurnos de Aechmea depressa, uma espécie endêmica do Sul da Bahia através de 67 horas de observação dos frutos de três indivíduos. As infrutescências foram visitadas em 2 por cento do tempo somente por micos-leão-da-cara-dourada (MLCD; Leontopithecus chrysomelas). Aechmea depressa investe na defesa física para reter seus frutos por períodos prolongados entretanto, o MLCD utiliza seu pequeno tamanho e extrema destreza para remover as brácteas rígidas e extrair os frutos da infrutescência. Estudos prévios indicam que os MLCDs dispersam sementes de espécies de Aechmea. Baseando-se na baixa taxa de visitação e na probabilidade de que poucos frugívoros manipulem adequadamente os frutos de A. depressa, nós sugerimos que os MLCD sejam extremamente importantes para manter as populações desta espécie de Bromeliaceae.

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The stability of lycopene in an olive oil/tomato emulsion during thermal processing (80-140 °C) was studied. Initially, the degradation of total lycopene (all-E plus Z-forms) occurred quickly at temperatures above 100 °C. However, a nonzero plateau value, depending on the processing temperature, was attained after longer treatment times. Besides degradation, the isomerization of total-Z-lycopene as well as the individual isomerization of all-E-, 5-Z-, 9-Z-, and 13-Z-lycopene was studied in detail. After prolonged heating, the isomer conversion reached a temperature-dependent equilibrium state. The degradation of total lycopene and the isomerization could be described by a fractional conversion model. The temperature dependency of the corresponding reaction rate constants was quantified by the Arrhenius equation. The activation energy of degradation was estimated to be 28 kJ/mol, and the activation energy of overall (all-E and total-Z) isomerization was estimated to be 52 kJ/mol.

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The effect of high pressure-high temperature (HPHT) processing on the formation of acrylamide and other Maillard-type reaction compounds was investigated in order to elucidate the impact of HPHT conditions on the different stages of the Maillard reaction. This study was performed in equimolar asparagine-glucose model systems that were treated at various HP/HT conditions (100-115 °C, 400-700 MPa, 0-60 min), and, for comparison, the model system was also heat-treated at ambient pressure. On the treated samples, the concentration of acrylamide, reactants, hydroxymethylfurfural, organic acids, and melanoidins was determined and the pH prior to and after treatment was measured. Based on the measured responses, the retarding effect of high pressure on the overall Maillard reaction was demonstrated; no or little differences were observed between 400 and 700 MPa. The study was conducted in two types of buffer, i.e. phosphate and MES buffer. In case of acrylamide, aspartic acid and browning, a higher concentration was generated in the MES buffer system, but these differences with the phosphate buffer system could be ascribed to pH changes resulting from the application of combined high pressure and high temperature. Based on the results, acrylamide formation is not expected to pose a major hazard to HPHT-treated products.

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When gliadin, a mixture of wheat storage proteins containing only intramolecular disulfide (SS) bonds, is heated at high temperatures and preferably at alkaline pH, the SS bonds are cleaved by ß-elimination reactions leading to decreased cystine levels and the generation of dehydroalanine (DHA) and free sulfhydryl (SH) groups. DHA and the free SH group of cysteine can further react to form the irreversible cross-link lanthionine (LAN). The kinetics of this reaction were studied by heating model systems containing gliadin at different pH values (pH 6.0, 8.0 and 11.0) at temperatures up to 120 °C. Multiresponse modeling was applied to simultaneously describe the course of the reaction partners, intermediates and products. The estimated kinetic parameters indicate that the reaction rate constant for the elimination reaction increases with temperature and pH. Moreover, the predominant reaction consuming the intermediary DHA is the cross-link with cysteine to form LAN following second-order reaction kinetics. The corresponding reaction rate constant is less dependent on temperature and pH. Use of the proposed kinetic model to estimate reaction product concentrations in cereal-based foods allowed us to conclude that the ß-elimination reaction may be less important during, e.g., bread making, but may well contribute to gluten network formation during the production of soft wheat products. It may also well be relevant in the production of bioplastics made from gluten.

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The effect of thermal processing on the stability of beta-carotene in carrot puree was investigated in a broad temperature range (80-150 degrees C). Heat induced changes in the stability of beta-carotene resulting in the conversion into its cis-isomers until an equilibrium state was reached after prolonged heating. By using nonlinear one-step regression analysis, the overall isomerization of all-trans-beta-carotene and the formation of individual cis-isomers could be modeled with a fractional conversion model. The Arrhenius equation was used to describe the temperature dependence of the reaction rate constants. As indicated by the low activation energies for all compounds (11 kJ mol(-1)), the isomerization rate constants showed little sensitivity toward the treatment temperature. The temperature dependence of the equilibrium concentration values after prolonged heating (C(f)) varied for the different compounds, but in all cases, a linear relation between the C(f) values and the treatment temperature could be noted. Although isomerization was observed as a result of thermal processing, it could be concluded that during industrially relevant heating processes, the retention of all-trans-beta-carotene in plain carrot puree was relatively high, which is most likely due to the presence of the protecting food matrix.

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The influence of water activity on the formation and elimination reactions of acrylamide was examined by means of multiresponse modeling on two different levels of complexity: basic equimolar asparagine-glucose systems and equimolar potato-based asparagine-glucose systems. To this end, model systems were first equilibrated to initial water activities in the range of 0.88-0.99 (corresponding roughly to the moisture gradient observed in French fries) and then heated at temperatures between 120 and 200 degrees C during different reaction times. For each sample, the concentration of acrylamide, glucose, asparagine, and aspartic acid was measured, as well as the extent of browning. A mechanistic model was proposed to model the five measured responses simultaneously. For both types of model systems, the model prediction was quite adequate, with the exception of the extent of browning, especially in the case of the potato-based model system. Moreover, the corresponding estimated kinetic parameters for acrylamide formation and elimination did not change significantly (based on a 95% confidence level) within the range of water activities tested, nor between the systems in the absence or presence of the potato matrix. The only remarkable difference was observed for the activation energy of acrylamide elimination, which was lower in the presence of the potato matrix, although not always significant. In general, these results confirm the generic nature of the model proposed and show that the influence of different moisture levels on acrylamide formation and elimination is minimal and that the addition of a potato matrix has little or no influence on the kinetic model and corresponding kinetic parameters.

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The parasite prevalence and infection intensity in primate wild populations can be affected by many variables linked to host and/or parasite ecology or either to interparasite competition/mutualism. In this study, we tested how host sex, age, and place of origin, as well parasitic concomitant infections affect the structure of golden lion and golden-headed lion tamarins parasite community, considering Trypanosoma cruzi and intestinal helminths infection in these primates. A total of 206 tamarins from two Atlantic Coastal rain forest areas in Brazil were tested during 4 years for prevalence of T. cruzi infection and helminth prevalence. Three intestinal helminth groups showed high prevalences in both tamarin species: Prosthenorchis sp., Spiruridae, and Trichostrongylidae. An association between presence of T. cruzi infection and higher intestinal helminth prevalence was found in both tamarin species. Two explanations for this association seem to be plausible: (1) lower helminth-linked mortality rates in T. cruzi-infected tamarins and (2) lower elimination rates of helminths in such tamarins. A higher frequency of T. cruzi-positive blood cultures was significantly correlated to female tamarins and to the presence of Trichostrongylidae infection. The possibility of an increase in the transmissibility of T. cruzi and the three analyzed helminths in lion tamarins with concomitant infections is discussed.

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The influence of water activity on the kinetics of acrylamide formation and elimination reaction was investigated using low-moisture equimolar asparagine-glucose model systems, which were heated at temperatures between 120 and 200 degrees C for variable heating times. To determine the water content corresponding to the water activities tested, a sorption moisture isotherm was constructed experimentally. The acrylamide concentrations measured at different water activities could be modeled on the basis of a reaction scheme including not only acrylamide formation and elimination reactions but also an alternative Maillard reaction between both reactants. The corresponding rate constants and activation energies were estimated using nonlinear regression analysis. Whereas the rate constant for acrylamide formation varied only slightly with the initial water activity of the model system, the elimination rate constant showed a clear minimum around a water activity of 0.82. The opposite trend, namely, a maximum at a water activity of 0.82, was found for the Maillard reaction rate constant as a function of water activity, which confirms data from literature. The activation energies for the different reactions changed in a comparable way as the corresponding rate constant with water activity.

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The effect of pH on acrylamide formation and elimination kinetics was studied in an equimolar (0.1 M) asparagine-glucose model system in phosphate or citrate buffer, heated at temperatures between 120 and 200 degrees C. To describe the experimental data, a simplified kinetic model was proposed and kinetic parameters were estimated by combined nonlinear regression and numerical integration on the data obtained under nonisothermal conditions. The model was subsequently validated in a more realistic potato-based matrix with varying pH. By increasing acidity, the reaction rate constants at T(ref) (160 degrees C) for both acrylamide formation and elimination can significantly be reduced, whereas the temperature dependence of both reaction rate constants increases. The introduction of a lyophilized potato matrix (20%) did not affect the acrylamide formation reaction rate constant at reference temperature (160 degrees C) as compared to the asparagine-glucose model system; the elimination rate constant at T(ref), on the contrary, was almost doubled.

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The kinetics of acrylamide (AA) was analyzed by heating a simple model system consisting of asparagine and glucose, fructose, or sucrose (0.01 M, pH 6) at temperatures between 140 and 200 degrees C. The AA concentration appeared to be the net result of simultaneous formation and elimination. A general kinetic model describing the AA yield was identified, and kinetic parameters were obtained by nonlinear regression on the nonisothermally derived data. On the basis of kinetic parameters, the AA formation appeared to proceed faster and to be more temperature sensitive in the asparagine-glucose than in the asparagine-fructose model system. The AA elimination kinetics, on the other hand, was similar. Significantly less AA was formed in the asparagine-sucrose model system as compared to the model systems with glucose or fructose.

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The effect of amino acids other than asparagine on acrylamide (AA) formation/elimination kinetics was studied in an asparagine-glucose model system (0.01 M, pH 6) heated at temperatures between 140 and 200 degrees C. Addition of cysteine or lysine to the model significantly lowered the AA yield, whereas addition of glutamine had a strong promoting effect and of alanine a rather neutral effect on the AA formation. This was also reflected by AA formation/elimination kinetics, which for all model systems studied could be modeled by two consecutive first-order reactions. The ratio of the elimination to the formation rate constant increased from the systems to which glutamine or alanine was added, over the control model system, to the model systems that contained lysine or cysteine.

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