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Whole Fat and Free Fatty Acid Contribution to the Taste of Fat
Chelsea Grimsley
Fall 2003
A critical literature review submitted in partial fulfillments of the Senior Research Thesis.
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Gustatory Detection of Free Fatty Acids in Rats: A Review - Prof. David W. Pittman, Papers of Psychology
An in-depth review of studies examining the gustatory detection of free fatty acids in rats. The research explores the role of whole fats, specifically corn oil and vegetable oil, and the effects of free fatty acids on taste receptor cells. Behavioral and human evidence is presented to support the idea that rats and humans have a gustatory cue for fat.
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Whole Fat and Free Fatty Acid Contribution to the Taste of Fat
Chelsea Grimsley Fall 2003
A critical literature review submitted in partial fulfillments of the Senior Research Thesis.
Abstract The increasing obesity rates in human society have prompted research in the taste of fat. This review addresses how dietary fat, in particular, free fatty acids, interacts with the gustatory system. Evidence drawn from whole fat and free fatty acid studies indicates a definite gustatory component for fat. Whole fat studies reveal that rats increase the intake of oil based on its attractiveness to the sense of taste. Removal of both pre- ingestive and post-ingestive effects supports the presence of a component in the gustatory system. Post-ingestive cues such as satiety have displayed significant effects upon the consumption of fat. However, orosensory cues such, as olfaction and texture are not contributors to the intake of fat. Early exposure to a high-fat diet results in a future preference for whole fats and a recovery period fails to alter their attraction to these foods. Whole fats are broken down into free fatty acids in the oral cavity. Saliva enzymes such as lingual lipase contribute to this reduction. Molecular and behavioral evidence reinforces gustatory activation of free fatty acids in rats. The identification of a fatty acid transporter (FAT) indicates molecular evidence that free fatty acids stimulate this sensory system. The inhibition of delayed-rectifying potassium (DRK) channels in taste receptor cells by free fatty acids indicates a transduction mechanism for fat in the gustatory system. Human studies show that the ability to taste PROP correlates with the capacity to detect free fatty acids in high-fat foods. These tasters have a higher density of fungiform papillae, which increases their susceptibility to free fatty acids. All of this evidence verifies the possibility of a gustatory component for fat in the taste system. This review will examine proof from whole fat studies using corn oil and vegetable oil and
Introduction Over the past decades, obesity has become an increasing threat the human race. Statistics show a rise in both childhood and adult obesity (Kamphius et al., 2001). A selective appetite for foods high in dietary fat is a characteristic that leads to many human obesity syndromes (Drewnowski, 1997). In the past decades a multitude of studies have been conducted to identify the factors underlying the human attraction to fat. Researchers have used animal models such as rats, to study the causes and effects of obesity. If experimenters can distinguish why rats are attracted towards foods containing fat, then they could possibly suggest that the taste system has a gustatory cue for fat. In turn, this would also indicate that fat has a taste. All fat found in food is termed dietary fat. Whole fats include oils, such as corn oil, vegetable oil, and triglycerides. Studies have proven that these fats increase intake in rat studies, prompting further research of their specific components. Free fatty acids are the basic chemical components of fat and represent chemicals that could potentially activate the gustatory system. Linoleic, linolenic and oleic acid are the three principle free fatty acids in corn oil. This review examines the potential for whole fats and free fatty acids to act on the taste system of rats and humans. Fats are responsible for the sensory properties in most foods and contribute to eating pleasure. Pre-ingestive and post-ingestive effects are two main factors that influence the intake of dietary fat. Pre-ingestive orosensory cues include olfaction, gustation, and texture. Post-ingestive effects include feedback signals and satiety, which can either promote or suppress intake. Eliminating either of these two ingestive effects can reveal the recognition of a possible gustatory component for fat.
The taste system has evolved mechanisms that enable the recognition of many different compounds. Taste reception begins when chemical stimuli react with receptors in papillae of the tongue. The papillae contain taste buds, which contain up to fifty taste receptor cells. Transduction occurs when sapid molecules interact with the taste receptor cells. The four basic taste categories: salt, sour, bitter, sweet, and most recently introduced, umami, have different types of transduction mechanisms. Metabotropic bitter and sweet receptors activate G-proteins, which increase the amount of calcium in the taste receptor cell and stimulate neurotransmitter release. The transduction of sodium chloride (salty) and acids (sour) begins when sodium ions and hydrogen protons pass into the cell. This causes sodium chloride and acids to directly permeate ion channels and release neurotransmitters (Gilbertson, 2003). Human taste thresholds are relative to differences in transduction abilities. People who have the ability to taste phenylthiocarbamide (PTC) and 6-n-propylthiouracil (PROP) are more sensitive to bitter tastes than non-PROP tasters. In clinical studies, PROP tasters have displayed more fungiform papillae and an enhanced perception of fatty foods. The genetic predisposition for PROP tasters to detect fat could possibly be related to obesity. Further evidence supporting the correlation is explored later in this review. This paper examines the potential taste of fat in both rat and animal studies. Evidence of whole fats and free fatty acids suggest there is a gustatory cue that increases the consumption of fat. Molecular, behavioral and human research more clearly defines the correlation between this component and fat intake.
gastric cannulas and the food drains out of the esophagus or stomach through a tube. A removable screw cap closes the cannulas to prevent drainage during the non-testing period. Because this device does not allow for the rat to experience post-ingestive effects, it is widely used in many whole fat studies to assess the behavioral aspects of consumption based on orosensory cues alone. Several studies have compared sensory and post-ingestive factors with sham- feeding. In one study, the sham-feeding of oil is compared in rats on a high fat diet (HF) and a high carbohydrate diet (HC) (Reed et al., 1990). If sensory factors were important for the intake of oil by HF rats, then they should sham feed more oil than the HC rats. Assuming that post-ingestive factors are important, both groups should consume approximately the same amount. However, this assumption was not evident. The results from this study showed that the HF rats consumed more corn oil than the HC rats, but both did not differ over the whole testing period. This confirmed that sensory factors were important in the intake of fat. Past studies have proved that HF rats under normal conditions, no sham feeding, consume significantly more. Evidence from this study suggests that some metabolic or physiological result of being on a high fat diet causes rats to prefer sensory properties or corn oil however, they are not enough to maintain a difference. Another study investigated the interaction between both orosensory and post- ingestive influences on the consumption of corn oil in thirty-minute test (Davis et al., 1995). The results were based upon the microstructure of the rats’ licking behavior and licking rates. The accumulation of ingested fluid gave a negative feedback signal preventing further intake at the intermediate concentrations and decreased intake at the
three highest concentrations. The initial rate of licking increased in a monotonic fashion with concentration and then rose rapidly with further increases in the oil concentration. This suggests that the ingestion of low concentrations is primarily controlled by stimulation of the orophyrangeal nerve with post-ingestive factors playing a minimal part. At the higher concentrations, post-ingestive feedback of the gastrointestinal tract plays a more important role in determination of volume ingested, which in turn correlates with licking rates. In conclusion, it was determined that even when the rats’ intake was the same for different concentrations of corn oil, their motives for ingesting the same amounts is different. Mindell et al. performed a similar study in which the sham-feeding responses of deprived and non-deprived rats to corn oil and mineral oil were observed. The results from this study found that the orosensory effects of corn oil and mineral oil are enough to stimulate ingestion when the post-ingestive effects have been minimized. Although the sensory component responsible for the rats’ discrimination of the two oils was not identified in this experiment, several possibilities were identified. The authors suggest that olfactory, gustatory, tactile, and/or temperature cues could have contributed. Whatever this mechanism is, it is strong enough to discriminate corn oil versus mineral oil and is increased by food deprivation (Mindell et al., 1990). Studies have been performed examining rats’ preference for whole fats after an intervention such as a sensory altering surgery. Takeda et al. (2001) examined the contribution of olfaction in normal mice and anosmia mice that had undergone an olfactory blockade procedure. The contribution of olfactory stimuli in a conditioned place preference test (CPP) was investigated. The control group preferred corn oil over
Warwick et al. (1990) showed that rats bred on a high fat diet consumed more fat and more of the highest fat mixtures than rats breed on a low fat diet. It was also shown that even when those animals on a high fat diet were switched to a low fat diet, their preference maintained towards the higher fat substances. The animals were exposed to either diet for four weeks and then their preferences were observed. The fat level of each diet was manipulated by the addition of peanut oil to peanut butter across three concentrations. The high fat rats mainly consumed the mixture containing more peanut oil where as the low fat rats consumed more of the mixture with less peanut oil. These data prove that early consumption of a high fat diet results in a shift in food preference that is not reversible by conditioning with a low fat diet. The fact that the rats could not change their preference after a diet alteration has similarities of humans’ struggle with dieting and weight loss. This study also suggests that the potential gustatory component for fat cannot be altered. A comparison of the intake of oil at different concentrations and under different conditions such as free food or chronic food deprivation has also been studied (Kimura and Fujimoto, 2003). It proved that the rats fed ad libitum chose oil over the water however their intake was very small. The food-deprived rats consumed more oil and generally preferred higher concentrations than the ad libitum rats. The results also showed that their intake became stable at mid-range concentrations. This proposes that there is a volumetric limit for the intake of the oil emulsion relative to deprivation. This study also displayed the effects recovery in the food-deprived rats. The results indicated that the rats still preferred the oil emulsion but the intake was decreased from previous testing conditions. This study shows that when rats are deprived of food, they ingest
more oil than rats on a free food diet and their preference cannot be changed by a recovery period. The results from post-ingestive studies indicate that the increase in oil and fat intake must be due to a gustatory component. Smith et al. (2001) indicated that a significant component of a rat’s macronutrient diet might be driven by taste-based factors. This was the first report of using brief-access tests to dissociate orosensory influences from post-ingestive factors during intake. Preference thresholds, licking activity and consumption of corn oil and sucrose were observed in two inbred mouse strains. AKR/J mice prefer a higher amount of energy from fat sources where as SWR/J mice prefer a higher amount of carbohydrate. This carbohydrate preference changes across different paradigms and suggests that SWR/J mice have a greater responsivity to some orosensory or post-ingestive factors. The SWR/J mice displayed lower preference thresholds and consumed more of the solutions. In the brief-access tests, the SWR/J mice licked more of sucrose and corn oil at increasing concentrations. These results indicated that both motor and sensory factors influence lick responses to tastants. The tendency for AKR/J mice to select high fat diets was more dependent on post-ingestive factors because they showed indifference for sucrose in both short and long-term tests. It was also found that flavor did not play a role in this strain. The SWR/J mice used in this study possess the genetic predisposition to taste bitter substances and show avoidance to PROP. In addition to these perceptions, this study showed that this strain of mice has a greater responsivity to sucrose and corn oil. These factors combine to suggest that SWR/J mice can taste fat.
Molecular Evidence Saliva enzymes break fat down into free fatty acids. Lingual lipase and amylase are two digestive enzymes that are secreted from serous glands of the rat tongue. Lingual lipase is the enzyme responsible for the first step in fat digestion. It hydrolyzes triacylglycerols into fatty acids and partial acylglycerols in the stomach. This enzyme facilitates production of emulsions that are needed in fat digestion by pancreatic lipase. The only known source of lingual lipase is in lingual serous glands between the circumvallate and foliate papillae. While lingual lipase is present at birth, amylase is present in insufficient amounts in the immature animal. Hamosh and Hand (1978) first studied the structural development of serous cells of the rat tongue using light and electron microscopy and compared it to the accumulation of lingual lipase. The structural development of lingual serous glands was viewed in three stages. Lipase activity was first seen in twenty-day fetuses and then increased fourteen fold by birth. It was noticed that activity decreased in half during the first suckling period because of the release of enzymes. Activity returned to birth levels around the second postnatal day. The accumulation of amylase followed the same schedule. This study found that lingual lipase is secreted during the first suckling period and enzyme activity increases significantly throughout the neonatal stages. Pancreatic lipase was seen to decrease rapidly after birth and recovered after weaning. It is concluded that lingual lipase substitutes for the digestive function of pancreatic lipase. Milk and gastric mucosa also contributed to fat digestion. The presence of free fatty acids and partial glycerides with similar properties of lingual lipase in young rats
indicates that lingual lipase is active in the gastric digestion of fat. It is also seen that lingual lipase initiates the digestion of dietary lipids in the human stomach. Field et al. (1989) simultaneously purified lingual lipase and amylase from rat lingual serous glands. Both were purified from the same serous glands by the same procedure. The lipase and amylase were co-purified until the last hydrophobic chromatographic step. This report gave a scheme for isolation and simultaneous purification of lingual lipase and amylase. Similarities and differences between known amylases and lipases were revealed. Further investigation of antibody production and genetic analysis of these enzymes is needed to draw further conclusions. A fatty acid transporter (FAT) has been identified in vallate taste cells. This mechanism transports free fatty acids to the taste receptor cells in the tongue. Preliminary reports regarding this mechanism state that this transporter possibly precedes neurotransmitter release in the gustatory system. This also foreshadows that there may be multiple mechanisms that respond to dietary fat in the form of free fatty acids. It is presumed that if the fatty acid transporter or other proteins are involved in the sensation of fat in the oral cavity, then the transporter should be apparent in the taste organs. A study that examined the distribution of fatty acid transporter mRNA in taste buds of circumvallate papillae in rats. Only small amounts of this transporter were found in the papillae and none in the anterior or posterior epithelium layers of the tongue. This suggested involvement of a fatty acid transporter in the taste sensation of circumvallate papillae. A chemical analysis of the tongue was then performed using anti-fatty acid transporter serum to identify the site of the transporter in the papillae. A western blot analysis failed to detect this transporter in either the circumvallate papillae or the anterior
extracellular level. The authors found that free fatty acids produced from lingual lipase after ingestion of fat directly affect taste receptor cells. This study was the first evidence that there was a gustatory cue for rats produced by fat (Gilbertson et al., 1997). Gilbertson’s experiments are the only studies that have examined the molecular contribution for the detection of fat in the gustatory system. Despite the few neurophysiological studies, there is behavioral evidence that clearly supports the taste of fat. Behavioral Evidence There are many studies that provide behavioral evidence that free fatty acids activate the gustatory system in rats. Some of this evidence is achieved through conditioned taste aversion tests and preference tests. A conditioned taste aversion test (CTA) is a common technique used to measure malaise of a particular agent assuming that when a new substance is paired with an aversive substance; animals should avoid it in the future. Preference studies have been conducted to prove the role of free fatty acids. Preference testing can determine the partiality of a substance across many different concentrations. The preference of rats towards fat can be determined by concentration and physical form. One study proved that rats not only prefer long chain fatty acids to a vehicle but they prefer them to their derivatives as well (Tsuruta et al., 1999). Of the three long chain fatty acids (LCFA) used, rats preferred linoleic acid to oleic acid and linolenic acid to linoleic acid. One possible conclusion for the rats’ selectivity of these fatty acids could be the number of unsaturated bonds involved. The derivative may be combined to be more like a whole fat, therefore having a higher volatility and increasing
sensitivity. The preference of LCFA to LCFA derivatives reveals that both the carbon chain and the carboxylate group play a part in the recognition of these fatty acids. This evidence helps prove that rats have a potential receptor molecule for fatty acids in their olfactory and gustatory systems. Conditioned taste aversions reveal that rats can detect linoleic acid (McCormack et al., 2003). This study identified the detection threshold of linoleic acid in rats to be ≥66μΜ. When this concentration of linoleic acid was paired with a LiCl injection, the rats exhibited future avoidance of the acid. The rats did not show a conditioned taste aversion to 44μΜ of linoleic acid. These results led to the identification of gustatory neural pathway involved in this detection. Transection of the chorda tympani nerve eliminated a conditioned taste aversion to linoleic acid. This indicated that rats can detect the presence of linoleic acid at higher concentrations and that the chorda tympani nerve is responsible for transmitting neural information in conditioned taste aversion tests. Linoleic acid modifies the licking responses of rats to sweet, sour and salt tastants (Herzog et al., 2003). In this study, linoleic acid was added to different concentrations of sucrose, NaCl, citric acid and QHCl. The results showed that licking responses increased when linoleic acid was added to sucrose and citric acid solutions. Linoleic acid reduced the licking responses across most NaCl solutions and had no effect on QHCl solutions. It was also observed that the sucrose and NaCl solutions were perceived as more intense with the addition of linoleic acid. These data prove that linoleic acid causes inhibition of basolateral K+^ channels thus delaying the depolarization of taste receptor cells in response to taste stimuli.
Subjects were identified as PROP supertaster, medium tasters or nontasters by comparing the taste-intensity function of PROP to NaCl for each individual. Supertasters showed significantly higher densities of fungiform papillae than medium tasters. In turn, medium tasters had higher densities than nontasters. Capsaicin is a chemical irritant that produces heat sensations when small amounts are present. Tasters have lower pain thresholds for these chemical irritants. Capsaicin ratings showed that both medium and supertasters perceived an oral burn from intake of capsaicin than the nontasters. These perceptions supported the categorization of PROP tasters. More significantly, these two tasters had an increased ability to discriminate differences in fat content of salad dressings. The oiliness ratings from this study did not prove to be statistically significant however; fat content ratings were very strong. Tasters employ oiliness perception in order to judge fat content in salad dressings; this evidence helps support the hypothesis that PROP tasters perceive more fat in food due to their sensitivity to its texture and taste (Tepper and Nurse, 1997 & 1998). Since PROP tasters have an enhanced ability to perceive fat in food, their ability to detect free fatty acids in high-fat foods can be viewed as well. One study tested whether the ability to detect linoleic acid in ice cream is related to the ability to taste PROP (Nasser et al., 2001). Because PROP tasters are more sensitive to oral cues, it is proposed that they might be more sensitive to fatty acids. High-fat vanilla ice cream was mixed with an ethanol-based linoleic acid solution and the detection capabilities in tasters and nontasters were observed. Eight out of ten PROP tasters reported a higher fat content in the experimental ice cream versus the control ice cream where as only one out of six nontasters reported a higher fat perceptions. This data showed that PROP tasters not only
have the capability to detect fat in foods, but can detect free fatty acids in high-fat foods as well. Differences in subjects’ linoleic acid taste perceptions were then analyzed. Linoleic acid tasters increased their ability to discriminate between low-energy ice cream containing linoleic acid and low-energy ice cream containing oleic acid over linoleic acid nontasters. It was also seen that linoleic acid tasters based their intake of ice cream on satiety. The tasters decreased intake rates earlier than the non-tasters reporting feelings of “fullness.” There was a strong relationship between the amount consumed and satiety (Kamphuis et al., 2003). This study shows that linoleic acid taster status might affect food intake regulation. In addition, the ability to detect this free fatty acid could possibly explain the basis for terminating a meal. The evidence from these human studies proves that people with the ability to taste 6-n-propylthiouracil (PROP) have a greater density of fungiform papillae. This allows for more trigeminal nerve stimulation and an increase in oral texture perception. These characteristics of PROP tasters yield an amplified perception of fat. PROP tasters also have the ability to detect free fatty acids in high-fat foods. In particular, linoleic acid has an effect on food intake regulation. These studies support the hypothesis that there is a component in the human gustatory system that allows for the recognition of fat based on taste. Conclusion This literature review examines how dietary fat, in particular free fatty acids may activate the gustatory system. All of the whole fat studies are based upon the fact that