Modeling the kinetics of flavour release during drinking

Novel mathematical models for flavour release during drinking are described, based on the physiology of breathing and swallowing. Surprisingly, we conclude that most flavour molecules arriving in the nose are extracted from liquid left in the throat, after swallowing. The models are fit to real time flavour release data obtained using APCI-mass spectrometry. Before modelling, raw data are corrected for the effects of varying airflow rate, using the signal from acetone in exhaled air. A simple equilibrium batch extraction model correctly describes flavour release during the first breaths after swallowing a flavoured liquid. It shows that for eight volatiles, whose in vitro air-water partition coefficients vary by a factor of 500, the apparent in vivo air-saliva partition coefficients vary only by a factor of five. To interpret the kinetics of flavour release longer after swallowing, diffusion of flavour into the throat lining is included. This is done using a three-layer model for mass transfer in the throat. An analytical solution of this model gives good fits to typical data. These models de-couple the physiological and physico-chemical aspects of flavour release, clarifying the effect of behaviour on in-vivo flavour release.     

In vitro study of the influence of physiological parameters on dynamic in-mouth flavour release from liquids

Influences of shear rate (surface extension), airflow, in-mouth headspace volume, synthetic saliva and human epithelial cells (modelling mucosa) on the initial dynamic flavour release from liquids were analysed. Simulating physiological mouth parameters, initial dynamic flavour release experiments over a time period of 30 s were carried out using a proven mouth model apparatus. Flavour compounds of different chemical classes were dissolved in water or in aqueous starch hydrolysate in concentrations typically present in food ( micro g/l to mg/l). Forced by increasing shear rates the enlargement of the gas-liquid interface (vortex formation) caused an increased release of flavour molecules. The release of less soluble compounds was reduced by increasing shear forces due to an improved dissolution. Increasing volumetric airflow rates resulted generally in higher release rates and in a change of pattern of release kinetics. Maximum flavour release was found at a ratio of 1:1 for in-mouth headspace and liquid volume. Neither addition of saliva alone nor the combination of saliva and mucosa showed significant influence on in-mouth flavour release from liquids in the model mouth.     

Retro-nasal aroma release is correlated with variations in the in-mouth air cavity volume after empty deglutition

We hypothesized that interindividual differences in motor activities during chewing and/or swallowing were determining factors for the transfer of volatile aroma from the in-mouth air cavity (IMAC) toward the olfactory mucosa. In our first experiment, we looked for changes in IMAC volume after saliva deglutition in 12 healthy subjects. The mean IMAC volume was measured after empty deglutition using an acoustic pharyngometer device. Based on the time course of the IMAC volume after swallowing, we discerned two groups of subjects. The first group displayed a small, constant IMAC volume (2.26 mL ±0.62) that corresponded to a high tongue position. The second group displayed a progressive increase in IMAC (from 6.82 mL ±2.37 to 22.82 mL ±3.04) that corresponded to a progressive lowering of the tongue to its resting position. In our second experiment, we investigated the relationship between IMAC volume changes after deglutition and the level of aroma release at the nostril. For this purpose, the release of menthone was measured at the nostril level in 25 subjects who consumed similar amounts of a mint tablet. The subjects were separated into two groups corresponding to two levels of menthone release: high (H) and low (L). The mean volume of IMAC was measured during and after empty deglutition. Group H displayed a small, constant amplitude of IMAC volume change after deglutition, while Group L displayed a progressive increase in IMAC. It is likely that Group H continuously released the aroma through the veloglossal isthmus as the mint was consumed, while Group L trapped the aroma in the oral cavity and then released it into the nasal cavity upon swallowing. These results show that the in vivo aroma release profile in humans depends closely on the different motor patterns at work during empty deglutition.     

Mechanistic mathematical model for in vivo aroma release during eating of semiliquid foods

The paper describes a mechanistic mathematical model for aroma release in the oropharynx to the nasal cavity during food consumption. The model is based on the physiology of the swallowing process and is validated with atmospheric pressure chemical ionization coupled with mass spectrometry measurements of aroma concentration in the nasal cavity of subjects eating flavored yogurt. The study is conducted on 3 aroma compounds representative for strawberry flavor (ethyl acetate, ethyl butanoate, and ethyl hexanoate) and 3 panelists. The model provides reasonably accurate time predictions of the relative aroma concentration in the nasal cavity and is able to simulate successive swallowing events as well as imperfect velopharyngeal closure. The most influent parameters are found to be the amount of the residual product in the pharynx and its contact area with the air flux, the volume of the nasal cavity, the equilibrium air/product partition coefficient of the volatile compound, the breath airflow rate, as well as the mass transfer coefficient of the aroma compound in the product, and the amount of product in the mouth. This work constitutes a first step toward computer-aided product formulation by allowing calculation of retronasal aroma intensity as a function of transfer and volatility properties of aroma compounds in food matrices and anatomophysiological characteristics of consumers.     

On the mechanism of air ventilaton in anabantoids (Pisces: Teleostei)

Summary Air ventilation in most Anabantoid species is diphasic, consisting of exhalation and inhalation. Exhalation is the release of air from the accessory breathing organs (suprabranchial chambers) through the mouth either into the water near the surface (e.g.,Ctenopoma) or directly into the atmosphere (e.g.,Osphronemus goramy). Inhalation, i.e., taking in fresh air through the mouth at the surface, immediately follows exhalation. X-ray films show (Figs. 5 and 6) that evacuation of the suprabranchial chambers during exhalation is total or nearly total. This, together with the fact that these chambers can contract at most to a very small extent, led to the conclusion that gas is replaced by water entering the chambers during exhalation and that this water is replaced by fresh air during inhalation. Further analysis of films, including conventional films showing the behavior of the opercular apparatus during air ventilation (Fig. 7), leads to a theory of a double-pumping mechanism responsible for air ventilation. This mechanism consists of the buccal apparatus and the opercular apparatus. It is suggested that both of these structures are able to act as both suction and pressure pumps, and thus air ventilation may be explained as the result of alternating activity of these two pumps.In the monophasic air ventilation characteristic of (adult)Anabas testudineus, there is no exhalation phase comparable to that of other Anabantoids. Therefore, no water enters the suprabranchial chambers, which remain filled with gas during the whole ventilation process (Fig. 10). Ventilation is limited to one phase comparable to inhalation in other Anabantoids.The structure of the accessory breathing organs (Fig. 1) and its progressive complication with growth (Fig. 4) were studied inOsphronemus goramy. The arrangement of the labyrinthine plates is in accordance with the requirements of transport of water and gas through the suprabranchial chambers. One plate (the inner plate, Fig. 1) separates these chambers into atrium, ventro-caudal, and dorso-caudal compartments, each with its own opening (valve). This organization seems essential for the transport of gas and water through the suprabranchial chambers and ensures that during exhalation, water flows into the chambers from above, so that while water is filling these chambers displaced gas can be sucked through the deep-lying pharyngeal openings into the expanding buccal cavity.Supported by the Deutsche Forschungsgemeinschaft     

Sensory perception is related to the rate of change of volatile concentration in-nose during eating of model gels.

The relationship between perceived aroma and the volatile concentration measured in-nose was investigated during eating of a model food. Sensory ranking and time-intensity analysis (TI) were used to measure perceived aroma, while in-nose volatile concentration was monitored by atmospheric pressure ionization mass spectrometry, which produced time release data. A gelatine-sucrose gel with a range of gelatine concentrations (2-8% w/w) and flavoured with furfuryl acetate was used as the model food. Sensory scaling showed decreased flavour intensities and TI showed a decrease in the flavour perceived over time, as the gelatine concentration increased. Studies in model systems and in people demonstrated that the different rates of release observed for different gelatine concentrations were not due to binding of volatile to protein in the gel, nor to mucous membranes, but were due to different rates of gel breakdown in-mouth. There were no significant differences in the maximum in-nose volatile concentrations for the different gelatine concentrations, so the amount of volatile present did not correlate well with the sensory analysis. However, the rates of volatile release were different for the different gels and showed a good correlation with sensory data.     

Lung ventilation in salamanders and the evolution of vertebrate air-breathing mechanisms

Functional analysis of lung ventilation in salamanders combined with historical analysis of respiratory pumps provides new perspectives on the evolution of breathing mechanisms in vertebrates. Lung ventilation in the aquatic salamander Necturus maculosus was examined by means of cineradiography, measurement of buccal and pleuroperitoneal cavity pressures, and electromyography of hypaxial musculature. In deoxygenated water Necturus periodically rises to the surface, opens its mouth, expands its buccal cavity to draw in fresh air, exhales air from the lungs, closes its mouth, and then compresses its buccal cavity and pumps air into the lungs. Thus Necturus produces only two buccal movements per breath: one expansion and one compression. Necturus shares the use of this two-stroke buccal pump with lungfishes, frogs and other salamanders. The ubiquitous use of this system by basal sarcopterygians is evidence that a two-stroke buccal pump is the primitive lung ventilation mechanism for sarcopterygian vertebrates. In contrast, basal actinopterygian fishes use a four-stroke buccal pump. In these fishes the buccal cavity expands to fill with expired air, compresses to expel the pulmonary air, expands to fill with fresh air, and then compresses for a second time to pump air into the lungs. Whether the sarcopterygian two-stroke buccal pump and the actinopterygian four-stroke buccal pump arose independently, whether both are derived from a single, primitive osteichthyian breathing mechanism, or whether one might be the primitive pattern and the other derived, cannot be determined. Although Necturus and lungfishes both use a two-stroke buccal pump, they differ in their expiration mechanics. Unlike a lungfish (Protopterus), Necturus exhales by contracting a portion of its hypaxial trunk musculature (the m. Iransversus abdominis) to increase pleuroperitoneal pressure. The occurrence of this same expiratory mechanism in amniotes is evidence that the use of hypaxial musculature for expiration, but not for inspiration, is a primitive tetrapod feature. From this observation we hypothesize that aspiration breathing may have evolved in two stages: initially, from pure buccal pumping to the use of trunk musculature for exhalation but not for inspiration (as in Necturus); and secondarily, to the use of trunk musculature for both exhalation and inhalation by costal aspiration (as in amniotes).     

Electrophysiologic study of the cerebral control of swallowing

Swallowing dominant, created by serial pouring of water into rabbit's mouth cavity was tested in after-action from water supply. As a result of summation in the dominant focus of excitations elicited by stimuli of various modalities, swallowing took place. The study of swallowing dominant revealed by its stimulus which elicits its own unconditioned reflex (blinking), has shown that temporary feedback is characteristic for the dominant as well as for the conditioned reflex. Summation in the swallowing center during dominant testing by stimulation of the eye cornea takes place without conjugated inhibition of the blinking center. During discrete pouring of water into the animal's mouth, evoked potentials appear in the cortical-subcortical structures of the swallowing center. Identical potentials appear in corresponding structures before the summation swallowing. Appearance of these potentials in the electric activity of the dominant focus testifies to its readiness for summation.     

Real-time monitoring of nasal mucosal pH during carbon dioxide stimulation: implications for stimulus dynamics

Carbon dioxide is a commonly employed irritant test compound in nasal chemesthetic studies because it is essentially free of olfactory stimulus properties. CO(2) is thought to act via hydration to H(2)CO(3) and dissociation to H(+) in nasal mucus, with resulting activation of acid sensors. However, transient changes in nasal mucosal pH have not been documented during CO(2) stimulation in humans. We placed a small pH probe on the floor of the right anterior nasal cavity during CO(2) stimulation in eight human subjects with historically high (>30%) and low (< or =20%) CO(2) detection thresholds. Three second pulses of CO(2) (15-45% v/v) paired with air in random order (12-15 s inter-stimulus interval; 60 s inter-trial interval) were administered by nasal cannula at 5 l/min. in an ascending series. For each subject, both a CO(2) detection threshold and suprathreshold psychophysical ratings [psi; labeled magnitude scale] were generated. All subjects showed phasic drops in pH associated with CO(2) stimulation (DeltapH). For all subjects combined, a positive correlation was apparent between applied [CO(2)] and both DeltapH and psi, as well as between DeltapH and psi themselves (P < 0.0001 for each comparison). Subjects with historically low CO(2) thresholds showed steeper dose-response curves for psi as a function of both applied [CO(2)] and DeltapH, but not for DeltapH as a function of applied [CO(2)]. For the six of eight subjects with measurable pH changes at threshold, DeltapH was positively related to log [CO(2) threshold] (P < 0.01). These data imply that variability in CO(2) detection thresholds and suprathreshold rating may derive from intrinsic differences in neural sensitivity, rather than differences in stimulus activation to hydrogen ion.     

Simulation of swallowing dysfunction and mechanical ventilation after a Montgomery T-tube insertion

The Montgomery T-tube is used as a combined tracheal stent and airway after laryngotracheoplasty, to keep the lumen open and prevent mucosal laceration from scarring. It is valuable in the management of upper and mid-tracheal lesions, while invaluable in long and multisegmental stenting lesions. Numerical simulations based on real-patient-tracheal geometry, experimental tissue characterization, and previous numerical estimation of the physiological swallowing force are performed to estimate the consequences of Montgomery T-tube implantation on swallowing and assisted ventilation: structural analysis of swallowing is performed to evaluate patient swallowing capacity, and computational fluid dynamics simulation is carried out to analyze related mechanical ventilation. With an inserted Montgomery T-tube, vertical displacement (Z-axis) reaches 8.01 mm, whereas in the Y-axis, it reaches 6.63 mm. The maximal principal stress obtained during swallowing was 1.6 MPa surrounding the hole and in the upper contact with the tracheal wall. Fluid flow simulation of the mechanical ventilation revealed positive pressure for both inhalation and exhalation, being higher for inspiration. The muscular deflections, considerable during normal breathing, are nonphysiological, and this aspect results in a constant overload of the tracheal muscle. During swallowing, the trachea ascends producing a nonhomogeneous elongation. This movement can be compromised when prosthesis is inserted, which explains the high incidence of glottis close inefficiency. Fluid simulations showed that nonphysiological pressure is established inside the trachea due to mechanical ventilation. This may lead to an overload of the tracheal muscle, explaining several related problems as muscle thinning or decrease in contractile function.     

Flow simulation in the human upper respiratory tract

Computer simulations of airflow patterns within the human upper respiratory tract (URT) are presented. The URT model includes airways of the head (nasal and oral), throat (pharyngeal and laryngeal), and lungs (trachea and main bronchi). The head and throat morphology was based on a cast of a medical school teaching model; tracheobronchial airways were defined mathematically. A body-fitted three-dimensional curvilinear grid system and a multiblock method were employed to graphically represent the surface geometries of the respective airways and to generate the corresponding mesh for computational fluid dynamics simulations. Our results suggest that for a prescribed phase of breath (i.e., inspiration or expiration), convective respiratory airflow patterns are highly dependent on flow rate values. Moreover, velocity profiles were quite different during inhalation and exhalation, both in terms of the sizes, strengths, and locations of localized features such as recirculation zones and air jets. Pressure losses during inhalation were 30-35% higher than for exhalation and were proportional to the square of the flow rate. Because particles are entrained and transported within airstreams, these results may have important applications to the targeted delivery of inhaled drugs.     

Metallic taste and retronasal smell

A series of experiments investigated the nature of metallic taste reports and whether they can be attributed to the development of a retronasal smell. Two studies showed that the metallic sensation reports following oral stimulation with solutions of FeSO4 were reduced to baseline when the nose was occluded. No such reduction was seen for CuSO4 or ZnSO4, which were more bitter and astringent, respectively, and less metallic. A discrimination test based on weak but equi-intense levels of FeSO4 and CuSO4 showed that FeSO4 could be discriminated from water with the nose open but not when occluded, but that discrimination of CuSO4 from water was not impaired by nasal occlusion. A discrimination test demonstrated that the headspace over solutions of FeSO4 was not different from water, although some subjects could discriminate FeSO4 solutions from water in the mouth when the nose was occluded, perhaps by tactile or astringent cues. These results confirm that metallic taste reports following oral stimulation with FeSO4 are likely due to development of a retronasal smell, possibly following a lipid oxidation reaction in the mouth. However, metallic taste reports may arise from different mechanisms with copper and zinc salts.     

Observation of the swallowing process by application of videofluoroscopy and real-time magnetic resonance imaging-consequences for retronasal aroma stimulation.

The process of eating and drinking was observed in vivo by application of videofluoroscopy, a dynamic X-ray technique, as well as real-time magnetic resonance imaging. The study was aimed at elucidating the timing and performance of the physiological organs involved in mastication and swallowing, mainly the tongue, the pharynx and the soft palate (velum palatinum). It was shown for the first time that effective physiological barriers do exist during food consumption that are capable of retaining volatiles such as helium within the oral cavity. These barriers allow the access of odorants to the nasal cavity only at certain times during the eating process. Their effectiveness is related to the texture of the food as well as the amount of food material present in the oral cavity and, thereby, directly influences retronasal aroma perception.     

Volatile emission by contest losers revealed by real-time chemical analysis

Animal interactions often involve chemical exchange but simultaneous evaluation of chemistry and behaviour has been problematical. Here we report findings from a novel method, atmospheric pressure chemical ionization-mass spectrometry (APCI-MS) coupled with manipulation of molecular-mass achieved by rearing organisms on deuterium-enhanced nutrients. This allows real-time monitoring of the occurrence and quantity of volatile chemicals released by each of two interacting individuals, in tandem with behavioural observations. We apply these methods to female-female contests in the parasitoid wasp Goniozus legneri. We show that this species emits the spiroacetal 2-methyl-1,7-dioxaspiro[5.5]undecane. Chemical release is most common in more behaviourally aggressive contests, which occur when prior resource owners successfully resist take-over by similar-sized intruder females. Volatiles released during contests are always emitted by the loser. Aggression in contests is reduced after spiroacetal release. We suggest that the spiroacetal functions as a weapon of rearguard action. We anticipate that APCI-MS, which is rapid, non-intrusive and relatively inexpensive to operate, will be widely applied in studies linking chemistry and behaviour.     

Axon mis-targeting in the olfactory bulb during regeneration of olfactory neuroepithelium

During development, primary olfactory axons typically grow to their topographically correct target zone without extensive remodelling. Similarly, in adults, new axons arising from the normal turnover of sensory neurons essentially project to their target without error. In the present study we have examined axon targeting in the olfactory pathway following extensive chemical ablation of the olfactory neuroepithelium in the P2-tau:LacZ line of mice. These mice express LacZ in the P2 subpopulation of primary olfactory neurons whose axons target topographically fixed glomeruli on the medial and lateral surfaces of the olfactory bulb. Intraperitoneal injections of dichlobenil selectively destroyed the sensory neuroepithelium of the nasal cavity without direct physical insult to the olfactory neuron pathway. Primary olfactory neurons regenerated and LacZ staining revealed the trajectory of the P2 axons. Rather than project solely to their topographically appropriate glomeruli, the regenerating P2 axons now terminated in numerous inappropriate glomeruli which were widely dispersed over the olfactory bulb. While these errors in targeting were refined over time, there was still considerable mis-targeting after four months of regeneration.     

An analysis of a specialized feeding mechanism of the spined loach, Cobitis taenia (L.), and a description of the related structures

Aquarium observations and gut analyses of the spined loach, Cobitis taenia , suggested the use of a technique by which small items of food could be removed from an almost constant stream of substrate, taken in through the mouth and passed out through the gills. The small amount of indigestible substrate present in the gut contents indicated a very high efficiency level for the process. Photographic investigation revealed a two phased feeding cycle based on the respiratory pumps, consisting of an induction/separation phase and an expulsion/swallowing phase. A morphological study showed large concentrations of mucus secreting goblet cells on the roof of the anterior buccopharyngeal cavity. Other morphological adaptations were noted which resulted in a narrowing of the anterior buccal cavity and a widening of the posterior part along with an increase in the distance between the gill rakers. In the light of this information a method of separation based on differential densities of particles being trapped in a constant stream of mucus is postulated.     

Morphology and function of the feeding apparatus of the lungfish, Lepidosiren paradoxa (Dipnoi)

The feeding mechanism of the South American lungfish, Lepidosiren paradoxa retains many primitive teleostome characteristics. In particular, the process of initial prey capture shares four salient functional features with other primitive vertebrates: 1) prey capture by suction feeding, 2) cranial elevation at the cranio-vertebral joint during the mouth opening phase of the strike, 3) the hyoid apparatus plays a major role in mediating expansion of the oral cavity and is one biomechanical pathway involved in depressing the mandible, and 4) peak hyoid excursion occurs after maximum gape is achieved. Lepidosiren also possesses four key morphological and functional specializations of the feeding mechanism: 1) tooth plates, 2) an enlarged cranial rib serving as a site for the origin of muscles depressing the hyoid apparatus, 3) a depressor mandibulae muscle, apparently not homologous to that of amphibians, and 4) a complex sequence of manipulation and chewing of prey in the oral cavity prior to swallowing. The depressor mandibulae is always active during mouth opening, in contrast to some previous suggestions. Chewing cycles include alternating adduction and transport phases. Between each adduction, food may be transported in or out of the buccal cavity to position it between the tooth plates. The depressor mandibulae muscle is active in a double-burst pattern during chewing, with the larger second burst serving to open the mouth during prey transport. Swallowing is characterized by prolonged activity in the hyoid constrictor musculature and the geniothoracicus. Lepidosiren uses hydraulic transport achieved by movements of the hyoid apparatus to position prey within the oral cavity. This function is analogous to that of the tongue in many tetrapods.     

Flavor release of diacetyl and 2-heptanone from cream style dressings in three mouth model systems

The release of volatile compounds from a cream style dressing, which consisted of a thickening agent dispersed in the water phase of an oil in water (o/w) type of emulsion, was studied by the purge-and-trap (PT), dynamic head space mastication (DHM) and dynamic headspace (DH) model systems for diacetyl and 2-heptanone as two volatile compounds. Big differences were detected in the quantity of volatiles released by the three models for both diacetyl and 2-heptanone: PT released the most, followed by DHM and DH. Nitrogen gas bubbling in PT and plunger up-and-down motion in DHM mimic mouth movements and promoted volatile release more than DH. The quantity of volatiles released depended on the nitrogen gas flow rate and isolation period with both the PT and the DHM model. Static headspace measurements indicated that no interaction occurred between the volatiles and the dispersion thickening agent, nor between the volatiles and protein of saliva.     

Desaturation of exhaled air in camels

We have found that camels can reduce the water loss due to evaporation from the respiratory tract in two ways: (1) by decreasing the temperature of the exhaled air and (2) by removal of water vapour from this air, resulting in the exhalation of air at less than 100% relative humidity (r.h.). Camels were kept under desert conditions and deprived of drinking water. In the daytime the exhaled air was at or near body core temperature, while in the cooler night exhaled air wat at or near ambient air temperature. In the daytime the exhaled air was fully saturated, but at night its humidity might fall to approximately 75% r.h. The combination of cooling and desaturation can provide a saving of water of 60% relative to exhalation of saturated air at body temperature. The mechanism responsible for cooling of the exhaled air is a simple heat exchange between the respiratory air and the surfaces of the nasal passageways. On inhalation these surfaces are cooled by the air passing over them, and on exhalation heat from the exhaled air is given off to these cooler surfaces. The mechanism responsible for desaturation of the air appears to depend on the hygroscopic properties of the nasal surfaces when the camel is dehydrated. The surfaces give off water vapour during inhalation and take up water from the respiratory air during exhalation. We have used a simple mechanical model to demonstrate the effectiveness of this mechanism.     

Sound Production in the Salientia: Mechanism and Evolution of the Emitter

Frog sounds involve expulsion of air through the larynx. Inmating, release, rain, and territorial calls, the air vibratesvocal cords and/or arytenoid cartilages. Sound is amplifiedand radiated by the distended buccal cavity and vocal sacs.Distress calls are emitted with open mouth, with minimum laryngealmodulation. The trunk is filled by inflation cycles, but airis driven out by synchronized contractions of the body wallmusculature. The pressure levels are more than five times thoseduring ventilation. In the release call of Bufo valliceps the dilatators and constrictorsof the larynx fire simultaneously keeping the larynx closed.As the pulmonary pressure reaches a peak they cease firing.The arytenoids then separate and vibrate, as do the vocal cords.The dilatators terminate the sound pulse by pulling vibratorsout of the air stream, hence the very sharp termination. Prolongedrelease call sequences include interpulse Teinflations thatreturn air from buccal cavity to lung. Frogs apparently evolved from amphibians too small to use aspirationbreathing. Vocalization represented a critical factor in theirsocial organization and its importance locked these animalsinto reliance upon pulse-pumping rather than the more efficientaspiration breathing.