The salivary proteome reflects some traits of dietary habits in diabetic and non-diabetic older adults.

PURPOSE: Objective markers of usual diet are of interest as alternative or validating tools in nutritional epidemiology research. The main purpose of the work was to assess whether saliva protein composition can reflect dietary habits in older adults, and how type 2 diabetes impacted on the saliva-diet correlates. METHODS: 214 participants were selected from 2 European cohorts of community-dwelling older adults (3C-Bordeaux and Seniors-ENRICA-2), using a case-control design nested in each cohort. Cases were individuals with type 2 diabetes. Dietary information was obtained using the Mediterranean Diet Adherence Screener (MEDAS). Saliva was successfully obtained from 211 subjects, and its proteome analyzed by liquid chromatography-tandem mass spectrometry. RESULTS: The relative abundance of 246 saliva proteins was obtained across all participants. The salivary proteome differed depending on the intake level of some food groups (especially vegetables, fruits, sweet snacks and red meat), in a diabetic status- and cohort-specific manner. Gene Set Enrichment Analysis suggested that some biological processes were consistently affected by diet across cohorts, for example enhanced platelet degranulation in high consumers of sweet snacks. Minimal models were then fitted to predict dietary variables by sociodemographic, clinical and salivary proteome variables. For the food group «sweet snacks¼, selected salivary proteins contributed to the predictive model and improved its performance in the Seniors-ENRICA-2 cohort and when both cohorts were combined. CONCLUSION: Saliva proteome composition of elderly individuals can reflect some aspects of dietary patterns.

Longitudinal analysis of the salivary metabolome of breast-fed and formula-fed infants over the first year of life.

INTRODUCTION: The salivary metabolome has been increasingly studied over the past ten years due to the potential of saliva as a non-invasive source of biomarkers. However, although saliva has been studied in relation to various diseases, its dynamic evolution during life is not known. This is particularly true for the first months of life. Infancy is indeed a critical period during which numerous behavioural and physiological events occur, such as dietary transitions and tooth eruption, which can lead to important biological modifications in the oral cavity. OBJECTIVES: The aim of this work was therefore to study the evolution of the salivary metabolome during the first months of life by 1H NMR. METHODS: Saliva of 32 infants with different milk feeding histories (breast vs formula) was collected at 6 stages, including 3 months old, 15 days before the onset of complementary feeding (CF), approximately 15 days after the onset of CF, approximately 21 days after the onset of CF and at approximately 11 and 15 months, and analysed. RESULTS: The longitudinal analysis showed a significant modification of the profiles of 18 metabolites over time; 14 presented an increase in abundance whereas 4 presented a decrease. These modifications seemed to be linked, for the most part, to an increase in oral microbial metabolism. Milk feeding history during the first months of life had no effect on metabolites. CONCLUSION: This work shows that the salivary metabolome should be considered when studying the changes occurring during infancy.

The bacterial species profiles of the lingual and salivary microbiota differ with basic tastes sensitivity in human.

Taste perception is crucial and impairments, which can be linked to pathologies, can lead to eating disorders. It is triggered by taste compounds stimulating receptors located on the tongue. However, the tongue is covered by a film containing saliva and microorganisms suspected to modulate the taste receptor environment. The present study aimed to elucidate the links between taste sensitivity (sweetness, sourness, bitterness, saltiness, umami) and the salivary as well as the tongue microbiota using shotgun metagenomics. 109 bacterial species were correlated with at least one taste. Interestingly, when a species was correlated with at least two tastes, the correlations were unidirectional, indicating a putative global implication. Some Streptococcus, SR1 and Rickenellaceae species correlated with five tastes. When comparing both ecosystems, saliva appears to be a better taste predictor than tongue. This work shows the implication of the oral microbiota in taste and exhibits specificities depending on the ecosystem considered.

Alterations in the Oral Microbiome Associated With Diabetes, Overweight, and Dietary Components.

The Mediterranean diet (MedDiet) represents the traditional food consumption patterns of people living in countries bordering the Mediterranean Sea and is associated with a reduced incidence of obesity and type-2 diabetes mellitus (T2DM). The objective of this study was to examine differences in the composition of the oral microbiome in older adults with T2DM and/or high body mass index (BMI) and whether the microbiome was influenced by elements of a MedDiet. Using a nested case-control design individuals affected by T2DM were selected from the Seniors-ENRICA-2 cohort concurrently with non-diabetic controls. BMI was measured, a validated dietary history taken, and adherence to a Mediterranean diet calculated using the MEDAS (Mediterranean Diet Adherence Screener) index. Oral health status was assessed by questionnaire and unstimulated whole mouth saliva was collected, and salivary flow rate calculated. Richness and diversity of the salivary microbiome were reduced in participants with T2DM compared to those without diabetes. The bacterial community structure in saliva showed distinct "signatures" or "salivatypes," characterized by predominance of particular bacterial genera. Salivatype 1 was more represented in subjects with T2DM, whilst those with obesity (BMI ≥ 30 kg/m2) had a predominance of salivatype 2, and control participants without T2DM or obesity had an increased presence of salivatype 3. There was an association of salivatype 1 with increased consumption of sugary snacks combined with reduced consumption of fish/shellfish and nuts. It can be concluded that the microbial community structure of saliva is altered in T2DM and obesity and is associated with altered consumption of particular food items. In order to further substantiate these observations a prospective study should be undertaken to assess the impact of diets aimed at modifying diabetic status and reducing weight.

Impact of Very Hot Drink Consumption Habits, Age, and Sex, on Taste Sensitivity.

The temperature range for consuming hot drinks includes temperatures that can damage cells on the tongue. We hypothesized that the consumption of very hot drinks can lead to a decrease in the ability to perceive low concentrations of tastants. We evaluated the ability to perceive low concentrations of five prototypical sapid compounds in 42 women and 40 men aged 18-65. A questionnaire made it possible to collect the usual frequencies (number of unit/day) and consumption temperature levels (medium hot/very hot) for four very common hot drinks (coffee, tea, herbal infusions, and hot chocolate). Our results showed that subjects who consumed very hot drinks (versus medium hot) were less sensitive to sweet (p = 0.020) and salty (p = 0.046) tastes. An aggravating effect of high consumption frequencies was only shown for sweet taste (p = 0.036). Moreover, our data also showed that women were more sensitive than men to sour, bitter, and umami tastes (p values < 0.05), as well as that taste sensitivity decreases with age, especially after 50 years old (all tastes; p values < 0.05). These findings strengthen our knowledge about the influence of sex and age on taste sensitivity, and they provide knowledge on the influence of consumption habits related to hot drinks on taste sensitivity.

The role of overweight in the association between the Mediterranean diet and the risk of type 2 diabetes mellitus: a mediation analysis among 21 585 UK biobank participants.

BACKGROUND: There is growing evidence that the Mediterranean (Medi) diet may lower the risk of type 2 diabetes mellitus (T2DM). Whether this association is due to the Medi diet by itself or is mediated by a diet-associated lower rate of overweight is uncertain. Our aim was to disentangle these relationships among UK adults. METHODS: Based on 21 585 participants from the UK Biobank cohort, the adherence to the Medi diet (high fruits, vegetables, legumes, cereals, fish, olive oil; low meat, dairy products; and intermediate alcohol intakes) was assessed (range 0-18). Data on diabetes were self-reported, and overweight was defined as a body mass index (BMI) ≥ 25 kg/m². A mediation analysis was implemented to disentangle the role of overweight in the Medi diet-T2DM relationship. RESULTS: The average baseline Medi diet score was 8.8 [standard deviation (SD) 2.6]. During a mean follow-up of 6.1 years, 473 individuals developed T2DM. A higher adherence to a Medi diet (+1 point) was associated with 14% decreased risk of T2DM [hazard ratio (HR): 0.86, 95% confidence interval (CI): 0.82-0.90]. This association split into an indirect effect of 10%, mediated by lower odds of overweight (HR: 0.90, 95% CI: 0.87-0.92), and a direct effect of the Medi diet of 4% (HR: 0.96, 95% CI: 0.93-0.99), regardless of the effect mediated by overweight. CONCLUSIONS: Considered as a single mediator, reduced overweight mainly contributes to the association between greater Medi diet adherence and lower risk of T2DM on this British subsample. However, the direct effect of the diet on the risk of T2DM, even weaker, should not be overlooked.

Effects of sucrose on salivary flow and composition: differences between real and sham intake.

Human saliva contains numerous salivary components that are fundamental for a healthy oral environment and the oral processing of foods. To study a possible differential influence of orosensory stimulation and metabolic activation on salivary composition, human parotid salivary flow, pH, A(280), and alpha-amylase activity were measured before, during and after real or sham (sip-and-spit) sucrose intakes. Variations in these salivary characteristics were related to perceived satiety. Sucrose, as either real or sham intake, increased salivary flow and pH and decreased A(280) before returning to pre-intake levels. Increased salivation was dependent on the sucrose concentration and was accompanied with a higher pH and lower A(280). After sucrose ingestion, the salivary alpha-amylase activity increased, while no increase occurred after sham sucrose intake. Similarly, rated satiety increased with real but not by sham sucrose intake. This indicated that salivary alpha-amylase is associated with perceived satiety controlled by caloric perception downstream of the oral cavity.

Biological films adhering to the oral soft tissues: Structure, composition, and potential impact on taste perception.

The role of free-flowing saliva in taste perception is increasingly recognized, but saliva is also present in the mouth as films intimately associated to soft or hard tissues. On mucosal surfaces, particularly on the tongue, the structure and composition of such films (including its microbial constitutive part) may play a particular role in the sense of taste due to their proximity with the taste anatomical structures. This review compiles the current knowledge on the structure of biological films adhering to oral mucosae and on their biochemical and microbiological composition, before presenting possible implications for taste perception. PRACTICAL APPLICATIONS: The understanding of the role of oral biological films on taste perception may provide new avenues of research and development for the industry or academia interested broadly in chemosensation.

How are macronutrient intake, BMI, ethnicity, age, and gender related to the composition of unstimulated saliva? A case study.

This study investigated how macronutrient intake, BMI, ethnicity, age, and gender are related to the composition of unstimulated saliva. First, two groups of Caucasian, Dutch subjects varying in daily intake of carbohydrate, fat, and protein were selected. The daily intake of macronutrients differed by two- to threefold between the low (n = 14) and high (n = 16) macronutrient intake groups. The same subjects were divided into two groups based on BMI: normal weight (n = 14, 22.5 ± 2.0 kg/m2 ) and overweight (n = 16, 28.1 ± 3.4 kg/m2 ). Second, one group of Caucasian, Dutch (n = 15) and one group of Asian, Chinese (n = 15) subjects were selected. Unstimulated saliva was collected from all groups. Protein concentration, amylolytic activity, lipolytic activity, and saliva flow rate were determined. None of the salivary parameters varied according to macronutrient intake and BMI. An effect of ethnicity on protein concentration was observed (p < .01; η2 = 0.142), with Asians having a 45% higher protein concentration in unstimulated saliva than Caucasians. Age had a significant effect on all salivary parameters. Protein concentration (p < .01; η2 = 0.256), amylolytic activity (p < .01; η2 = 0.234), and lipolytic activity (p < .05; η2 = 0.207) increased with age, while saliva flow rate decreased (p < .01; η2 = 0.262). Gender had a significant effect on saliva flow rate (p < .01; η2 = 0.130), with male subjects having a 32% higher flow rate than females. Age was the factor that had the greatest impact on the characteristics of unstimulated saliva. As the modulation of saliva composition according to diet has been reported previously, the extent to which macronutrient intake can affect saliva composition needs to be further investigated. PRACTICAL APPLICATIONS: Saliva plays an important role in food oral processing. From the breakdown of food structures to the binding of flavor compounds and the formation of a swallowable bolus, saliva is essential for the perception and appreciation of foods. Identifying the factors that affect saliva composition is, therefore, necessary to understand the differences in eating behavior, food perception, and preference across different consumer groups. This article aims to highlight the importance of considering saliva variability when designing food products that meet the needs of specific consumer groups.

The Associations between Biochemical and Microbiological Variables and Taste Differ in Whole Saliva and in the Film Lining the Tongue.

The objective of this work was to investigate whether the biological film lining the tongue may play a role in taste perception. For that purpose, the tongue film and saliva of 21 healthy subjects were characterized, focusing on microorganisms and their main metabolic substrates and products. In parallel, taste sensitivity was evaluated using a test recently developed by our group, and the links between biological and sensory data were explored by a correlative approach. Saliva and tongue film differed significantly in biochemical composition (proportions of glucose, fructose, sucrose, and lactic, butyric, and acetic acids) and in microbiological profiles: compared to saliva, tongue film was characterized by significantly lower proportions of Bacteroidetes (p<0.001) and its main genus Prevotella (p<0.01) and significantly higher proportions of Firmicutes (p<0.01), Actinobacteria (p<0.001), and the genus Streptococcus (p<0.05). Generic taste sensitivity was linked to biological variables in the two compartments, but variables that appeared influent in saliva (flow, organic acids, proportion of Actinobacteria and Firmicutes) and in tongue film (sugars and proportions of Bacteroidetes) were not the same. This study points to two interesting areas in taste research: the oral microbiome and the specific characterization of the film lining the tongue.

The basal free fatty acid concentration in human saliva is related to salivary lipolytic activity.

Fat perception during eating is a complex sensation that involves various sensory modalities, such as texture, aroma and taste. Taste is supported by the discovery of fatty acid receptors in the tongue papillae. Dietary fat is mainly composed of esterified fatty acids, whereas only free fatty acids can bind to taste receptors. Some authors have mentioned the necessity and efficiency of salivary lipolytic activity to hydrolyse the esterified fatty acids present in foods and enable fat perception. Our hypothesis is that salivary lipolytic activity is also involved in regulating the basal level of salivary fatty acids in humans. To test this hypothesis, total fatty acid (TFA) and free fatty acid (FFA) concentrations and selected salivary characteristics (such as lipolytic activity) were analysed in the resting saliva of 54 subjects. The results show differences in the TFA and FFA profiles, with TFA and FFA concentrations of 8.99 and 3.56 µg/mL of saliva, respectively. Interestingly, lipolytic activity had a significant positive correlation with FFA concentration (0.51, p < 0.01). This result highlights a possible physiological role of salivary lipolytic activity in the regulation of the basal FFA concentration. This regulation could be involved in fat taste sensitivity.

Associations between food consumption patterns and saliva composition: Specificities of eating difficulties children.

Identifying objective markers of diet would be beneficial to research fields such as nutritional epidemiology. As a preliminary study on the validity of using saliva for this purpose, and in order to explore the relationship between saliva and diet, we focused on clearly contrasted groups of children: children with eating difficulties (ED) receiving at least 50% of their energy intake through artificial nutrition vs healthy controls (C). Saliva of ED and C children was analyzed by various methods (targeted biochemical analyses, 2-D electrophoresis coupled to MS, 1H NMR) and their diet was characterized using food frequency questionnaires, considering 148 food items grouped into 13 categories. Complete datasets were obtained for 16 ED and 16 C subjects (median age 4.7y and 5.0y, respectively) and the statistical link between salivary and dietary characteristics was studied by Multiple Factor Analysis (MFA). Overall, ED children showed as expected lower consumption frequency scores and higher food selectivity. The two groups of children differed in "diet/saliva" associations. Some distinctive salivary variables were common to both groups of children. For example, carbonic anhydrase 6 and the consumption frequency of biscuits & sweets and drinks were positively associated with the MFA axis 1 in C children, but oppositely associated in ED children. Specifically for ED children, abundant salivary proteins (cystatins, amylase, amylase fragments) and some metabolites (amino acids, galactose, lactate) correlated with axis 1, together with the consumption frequency of sauces & seasonings, bread & cereal products, ready-to-eat meals, fish, biscuits & sweets, drinks and potatoes. Specifically for C children, several proteins (serum albumin, haptoglobin, Igκ, apolipoprotein A-1, α-1 antitrypsin) correlated with axis 1, together with the consumption frequency of biscuits & sweets, milk & dairy products, drinks, fruit, meat and vegetables. This study demonstrates that the qualitative aspect of diet is linked to saliva composition, and that the associations between dietary consumption and salivary composition differ between groups of subjects with contrasted diets.

Salivary Composition Is Associated with Liking and Usual Nutrient Intake.

Salivary flow and composition have an impact on flavor perception. However, very few studies have explored the relationship between saliva, individual liking and usual dietary intake. The aim of our study was to evaluate the association of salivary flow and composition with both a liking for fat, saltiness and sweetness and the usual nutrient intake in an adult French population. Liking for fat, saltiness, and sweetness were inferred from liking scores obtained during hedonic tests on 32 food products among 282 French adults participating in the Nutrinet-Santé Study. Before assessing liking, resting saliva was collected. Standard biochemical analyses were performed to assess specific component concentrations and enzymatic activities. Dietary data were collected using three web-based 24 h records. Relationships between salivary flow and composition, sensory liking and nutrient intake were assessed using linear regression. Total antioxidant capacity was positively associated with simple carbohydrate intake (ß = 31.3, 95% CI = 1.58; 60.99) and inversely related to complex carbohydrate consumption (ß = -52.4, 95% CI = -87.51; -19.71). Amylolysis was positively associated with both total (ß = 0.20, 95% CI = 0.01; 0.38) and simple carbohydrate intake (ß = 0.21, 95% CI = 0.01; 0.39). Salivary flow was positively associated with liking for fat (ß = 0.14, 95% CI = 0.03; 0.25). Proteolysis was positively associated with liking for saltiness and for fat (ß = 0.31, 95% CI = 0.02; 0.59; ß = 0.28, 95% CI = 0.01; 0.56, respectively). Amylolysis was inversely associated with liking for sweetness (ß = -10.13, 95% CI = -19.51; -0.75). Carbonic anhydrase 6 was inversely associated with liking for saltiness (ß = -46.77, 95% CI = -86.24; -7.30). Saliva does not substantially vary according to a usual diet, except for carbohydrate intake, whereas the specific association between salivary flow/composition and sensory liking suggests the influence of saliva characteristics in food acceptance.

Multi-omics profiling reveals that eating difficulties developed consecutively to artificial nutrition in the neonatal period are associated to specific saliva composition.

Prolonged enteral or parenteral nutrition in neonatal periods sometimes results in eating difficulties persisting for years, with reduced food intake through the oral route and thereby reduced stimulation of the oral cavity. Aiming at describing the consequences on oral physiology, saliva of 21 children with eating difficulties (ED) was compared to that of 23 healthy controls, using various omics and targeted methods. Overall, despite heterogeneity within the groups (age, medication etc.), the three spectral methods (MALDI-TOF, SELDI-TOF, (1)H NMR) allowed discriminating ED and controls, confirming that oral stimulation by food intake plays a role in shaping the composition of saliva. Saliva of ED patients exhibited a lower antioxidant status and lower levels of the salivary protease inhibitors cystatins. Other discriminant features (IgA1, dimethylamine) may relate to modified oral and/or intestinal microbial ecology. Finally, salivary profiles of ED patients were partly comparable to those of subjects with exacerbated gustatory sensitivities, in particular with reduced abundance of cystatin SN and higher abundance of zinc-alpha-2-glycoprotein. Whether this translates taste hypersensitivity and contributes to the eating difficulties deserves further attention.

Relating ionisation of calcium chloride in saliva to bitterness perception.

Saliva plays a role in the perception of bitter, sour and salty tastes that are presumed to be derived from the concentration of free cations or anions ions dissolved in saliva. The role of ionisation of calcium in bitter taste was studied by determining binding in vitro mixture of saliva and protein solutions and in spit. In vitro, the addition of whey to calcium chloride solutions increased the calcium binding, pH and viscosity. The addition of saliva to these mixtures, the increased calcium binding and the induced small changes in viscosity and pH were thought not to contribute significantly to bitterness perception. Nonstimulated saliva, at pH 7.5, contained about 5 mM calcium, of which about one third was ionised. The bitter threshold of fully ionised calcium chloride in water varied between 1 and 15 mM among individuals. In spit, after tasting whey, ionised calcium was found to have increased at low, but decreased at high, calcium concentrations and varied 30% among individuals. Bitterness was related, on average, to the concentration of ionised calcium and not to the total concentration of calcium in spit. A general explicative model based on the composition of bulk saliva is discussed in relation to perception threshold and the likely importance of saliva from von Ebner's gland.

NaCl and sugar release, salivation and taste during mastication of salted chewing gum.

Salt perception impacts on food acceptability and nutrition and depends upon salt release from foods that was determined in situ during mastication of chewing gum with up to 10% (1800 mmol/kg) added NaCl. The mechanical action of chewing increased salivation, which was further increased by the presence of salt, particularly above 180 mmol NaCl/kg gum or above 100 mM NaCl in saliva. The average resting salivary flow rate was 1 ml/min, increasing to 4 and 6 ml/min with gums containing low and high salt, respectively. Thus, stimulation of salivation by salt occurred at a concentration well above the taste threshold of 20 mM NaCl. NaCl concentration in nonstimulated saliva was about 10 mM and increased to 500 mM after 30 s chewing of the 10% NaCl gum and returned to near nonstimulated levels after 4 min chewing. Changes in pH of saliva were more gradual, increasing to a maximum at about 2 min and remaining elevated after 4 min. Salty taste was related to the free chloride ion concentration in saliva irrespective of the initial salt concentration in the gum with an indication of adaptation after 3 min chewing. During chewing, salty taste increased ahead of the increase in salivary conductivity and the salt concentration in the sublingual saliva varied in a cyclic fashion about every 20 s. This is consistent with a cyclic swallowing of saliva and replacement with newly secreted saliva of low salt content and mastication releasing further salt from the gum.

Role of saliva in oral food perception.

Saliva is the first fluid that comes into contact with food during oral processing. Because saliva is the medium that bathes the taste receptors, is the fluid through which taste and aroma compounds are released into the oral cavity and is mixed continuously with food during bolus formation, it is an essential actor in oral chemosensory perception. The complexity of saliva composition, with compounds originating from different salivary glands, from gingival crevicular fluid, from micro-organisms and from food debris, together with its variable nature increases the possibilities for interactions with food compounds and for different roles in perception. These factors are increasingly being taken into account in current research on food perception. The aim of this paper is to review the principal roles of saliva in oral perception, with particular focus on chemosensory perception. These include the protection of taste buds, the effects of flow rates, salivary hormones, electrolytes and organic compounds, and finally the impact of perception on salivary secretions.

Nutri-metabolomics applied to taste perception phenotype: human subjects with high and low sensitivity to taste of fat differ in salivary response to oleic acid.

Saliva has different functions in the mouth and is involved, for example, in taste perception. Saliva composition can also be modified rapidly by taste stimulation. It remains unclear, however, whether the perceived intensity of a tastant may modulate this response. Based on increasing evidence that fat can be perceived by the taste system and that fat taste perception may be associated with fat intake, the aim of this work was to study if stimulation by a fatty acid (oleic acid) modifies saliva composition differently in subjects highly (sensitive+) or weakly (sensitive-) sensitive to that taste. For that purpose, saliva of two groups of subjects was collected after stimulation by either a control emulsion or an emulsion containing 5.61 mM oleic acid. Saliva was analyzed by 2D electrophoresis and (1)H NMR spectroscopy. The results show that sensitive+ and sensitive- subjects differ in their salivary response in terms of proteome and metabolome composition. Oppositely to sensitive- subjects, sensitive+ subjects responded to oleic acid by increased abundance of polymeric immunoglobulin receptor, rab GDP dissociation inhibitor beta, and organic acids, and decreased abundance of metabolites characteristic of mucins. The results highlight that modification of saliva composition by taste stimulation may be modulated by taste perception.

Variability of human saliva composition: possible relationships with fat perception and liking.

Saliva is the medium that bathes the taste receptors in the oral cavity and in which aroma and taste compounds are released when food is eaten. Moreover saliva contains enzymes and molecules that can interact with food. To date, little research has been devoted to the intra- and inter-individual variabilities of these components and their inter-relationships. The first aim of this work was to study intra- and inter-individual variabilities over time in the composition of molecules likely to interact with food in the mouth, with particular focus on molecules that might interact with fat. The second aim was to try to relate this composition to a liking for fat and its perception. Stimulated and unstimulated saliva from 13 subjects was collected in the morning and afternoon on three occasions at 4-month intervals. Saliva characteristics such as flow, protein concentration, lipolysis, proteolysis, amylolysis, lipocalin concentration, lysozyme activity, total antioxidant status and uric acid concentrations were measured, as well as the liking for and perceived fattiness of a fat solution. The results showed that for most of the measured characteristics, intra-subject variability in saliva composition was smaller than inter-subject variability, with remarkable stability over time (8 months) in terms of flow, lypolysis, proteolysis and total antioxidant status. Relationships were found between some of these characteristics (lipolysis, lipocalin and flow) and fat-liking or perception, showing that the composition of saliva may play an important role in fat perception and liking.

Proteomic analysis of human whole and parotid salivas following stimulation by different tastes.

Whole and parotid salivas, collected after stimulation with tastants, were analyzed by 2D electrophoresis and mass spectrometry. In whole saliva, the number of proteins affected by taste stimulation increased in the order sweet < umami < bitter < acid. Annexin A1 and calgranulin A, involved in inflammation, were over-represented after umami, bitter, and sour stimulations. Their low abundance or absence in parotid saliva after bitter stimulation suggested that they originated from other oral glands or tissues.