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Ingestive Behavior and Neurosciences

Yeast Proteins Enhance Satiety in Rats^1,2

³ Institut National de la Recherche Agronomique, Unité INRA-INAPG de Physiologie de la Nutrition et du Comportement Alimentaire, Institut National Agronomique Paris-Grignon, F75231 Paris Cedex 05, France and ⁴ Bio-Springer, 94704 Maisons Alfort Cedex, France

^* To whom correspondence should be addressed. E-mail: fromenti{at}inapg.fr.

	ABSTRACT

TOP ABSTRACT Introduction Materials and Methods Results Discussion LITERATURE CITED

 
This study was designed to characterize the suppressant effect of yeast protein and purified peptides on energy intake. For this purpose, 5 experiments were carried out using adult male Wistar rats. Rats that consumed ad libitum a standard yeast protein diet ate significantly less and were leaner over 21 d than rats that consumed ad libitum a standard milk protein diet (Expt. 1). Moreover, rats fed a high yeast protein load reduced their next meal and daily energy intake more than rats fed any other well-balanced, amino acid, high protein load (soy, total milk protein, or wheat gluten) and more than those fed a wheat starch diet (Expt. 2). Purified peptides from the yeast protein extract produced similar effects on subsequent energy intake (Expt. 3). Study of the behavioral satiety sequence showed that rats consuming P14-y or P55-y diets ad libitum did not acquire a conditioned food aversion (Expt. 4). Finally, a preliminary study of gastric emptying in rats fed yeast protein loads showed that yeast protein was emptied more rapidly through the pylorus than total milk protein during a meal, which may induce satiety (Expt. 5). Taken together, these experiments show that yeast proteins enhance satiety in rats more than other proteins.

	Introduction

TOP ABSTRACT Introduction Materials and Methods Results Discussion LITERATURE CITED

	Materials and Methods

TOP ABSTRACT Introduction Materials and Methods Results Discussion LITERATURE CITED

 
    Animals and diets. Adult male Wistar rats from Harlan were housed in individual cages at 22 ± 2°C, under a 12-h reverse light/dark cycle (1000, 2200; with lights on at 2200). All experimental procedures used during the study complied with the guidelines issued by the French National Animal Care Committee and were approved by the Regional (Ile de France Sud) Animal Care and Ethics Committee. Standard protein diets were modified versions of the AIN-93M diet. (Table 1) (7). Instead of casein and cystine, the standard protein diet contained 140 g total milk protein/kg of diet (P14) and the standard yeast diet contained 140 g of yeast protein/kg of diet (P14-y). High protein diets were also based on the AIN-93M diet and contained 55% of P/E from milk (P55). The high protein yeast diet contained 55% P/E from yeast (P55-y). The addition of protein replaced equivalent amounts of sucrose and starch. During Expt. 2, three other diets were used as an intra-oral load: a protein-free wheat starch diet (WS), a high-protein diet containing soy protein (P55-s), and a high-protein diet containing gluten (P55-g). Amino acid analysis of the yeast protein extract demonstrated a well-equilibrated profile (Supplemental Table 1). Except when otherwise specified, all diets were moistened (P55 and P14 diets, 1:2 and 1:1 ratios of powder to water, respectively) to minimize spillage. Food intake was determined by the difference in food cup wt before and after each experimental period, corrected for spillage, the amount of water added and evaporation. Food containers were refilled daily at 1000. All rats had free access to water during the entire experimental period. For the first 10 d of the experiments (prefeeding period), rats were adapted to laboratory conditions, and consumed ad libitum a P14 diet (baseline food for all experiments), except for those under Expt. 3, where the rats were conditioned to a 3-meal pattern. The last day of the prefeeding period is referred to as d 0.

    Experiment 2: Satiety induced by high protein diets. On d 3 of the prefeeding period, rats were surgically equipped with an intra-oral cannula according to the procedure described by Philips and Norgren (8). Rats were allowed 1 wk for postoperative recovery. During this recovery period, the rats were infused with increasing volumes of water (10 mL) to ensure that the cannula remained patent and to habituate the rats accustomed to receiving the intra-oral load. Loads were delivered at a rate of 1 mL/min (9,10) by an infusion pump via polyethylene tubing (ID 1.14 mm) connected to the oral cannula (Biotech IPC-S, 12 channels).

During the entire period (16 d), rats received 3 separate meals/d to reproduce human feeding patterns. They were administered the load via the intra-oral catheter to guarantee complete consumption. The first meal was limited to 3 g and was made available between 1000 and 1015 (breakfast). For the other 2 meals, rats ate ad libitum for 1 h between 1500 and 1600 (lunch) and then between 2000 and 2100 (dinner). This schedule allowed the rats to adapt to eating their meals promptly while ensuring an adequate daily food intake (25 g). All rats received the P14 diet during the 3 meals. After this period of acclimatization and for 4 consecutive d (load period), rats were administered different intra-oral loads as a function of their group for 10 min, 1 h before dinner. We ensured that none of the infused loads were rejected during the infusion period. After the recovery period, 64 rats, weighing 175–190 g, were divided into 5 groups named according to the special load they were offered each day: a protein-free WS load (n = 14) and different high-protein loads containing 40% WS (as starch energy) and 55% protein differing as a function of the group: P55 (total milk protein, n = 12); P55-y (yeast protein, n = 8), P55-s (soy protein, n = 15) and P55-g (wheat gluten, n = 15). These loads were moistened (1:2), and were isovolumic and isocaloric (10 mL, 4.3 MJ/L). Food intake was measured after breakfast, lunch, loads (if any), and dinner. To account for between-day differences in daily food intake, the basal food intake was the mean meal intake during the last 2 d of the adaptation period (d 11 and d 12). To account for between-rat differences of food intake/meal, the amount eaten during each meal was expressed as a percentage of the basal meal food intake. This allowed each rat to serve as its own control. The effects of oral loads on basal food intake were calculated from 3 ratios: 1) the dinner ratio [(load + dinner) mean energy intake during the 4 load days x 100 / (dinner) energy intake during the baseline period)]; 2) the lunch ratio (lunch mean energy intake during the 4 load days x 100 / lunch energy intake during the baseline period); 3) the total next-day energy intake ratio (total next-day mean energy intake during the 4 load days x 100 / total next-day energy intake during the baseline period). These ratios were calculated with the means of the data for loads on d 13, 14, 15, and 16.

    Experiment 3: Effect of a yeast peptide load. A mixture of peptides (ranging from 1 kD to 10 kD) was extracted from the native yeast (used in all yeast diets described above) by Bio-Springer (Maisons-Alfort). These peptides were used as the unique protein source to develop a diet called P14-pep, containing 14% of P/E as peptides. The effect of this diet was compared with the effect of P14 and P14-y diets. Thus, after the prefeeding period, 21 rats weighing 206.4 ± 1.9g were divided into 3 groups: P14, P14-y and P14-pep. At the beginning of the night cycle, rats were presented with an isocaloric (42 kJ) and isovolumic load of 2 g (moistened 1:2) for 30 min. One-half hour after this load, rats were given free access to the P14 diet for the rest of the day, and the amount of P14 diet eaten was measured. Each experimental day (test day) was preceded by a "basal day" when the load in all groups was composed of the P14 diet. Food intakes were measured after 1, 3, and 20 h.

    Experiment 4: Behavioral satiety sequence. The experimental chamber, video equipment, and analysis of video tape were utilized as previously described in Bensaid et al. (7). After the prefeeding period, 18 rats weighing 190–220 g were assigned to 3 groups (P14, P14-y, and P55-y) and received the corresponding diets. The experimental period lasted for 14 d. For 2 d prior to the test day (d 14), the rats were habituated to their test video chamber between 0745 and 1000 (just before food presentation). The behavior of animals was video recorded for the first hour of food presentation (between 1000 and 1100) and food was consumed ad libitum. The food was removed between 0600 and 1000 every day to ensure that rats would eat during the video session. For this analysis, animal behavior was categorized into 4 types, as described by Bensaid et al. (7). The total 1-h food intakes were also measured, as were the parameters concerning each eating bout (number and duration). The mean eating rate was therefore calculated for each rat by dividing the total 1-h energy intake by the total 1-h bout duration (11). Rats were returned to their home cages with their food cup after each behavioral recording session.

    Experiment 5: Gastric emptying kinetics. After the prefeeding period, 60 rats weighing 180–200 g were divided into 3 groups named P14, P55, and P55-y. During the experimental period, the daily feeding schedule was divided into 2 parts: at 1000 (end of light cycle) rats were given a load of 2 g (moistened 1:2) for 30 min, and then ate a P14 diet (moistened 1:1) ad libitum between 1230 and 1800. Every day (except for the last 2 d of the experiment, d 9 and 10), the loads consisted of the P14 diet, but otherwise the diet corresponded to the name of the group. On the last day of the experiment (d 10), the rats were killed using sodium pentobarbital at different time points after the end of the load ingestion period (0, 30, 60, and 180 min). The stomachs were quickly ligatured and removed, weighed, and emptied, and the contents were also weighed. The contents were subsequently freeze-dried and the nitrogen rate determined by mass spectrometry.

    Statistics. Results are expressed as means ± SEM. Differences between groups were determined by 2-way ANOVA (Proc GLM, SAS, version 6.11), and a diet effect was tested except when specified. The level of significance was set at P < 0.05, but when a significant effect was revealed by ANOVA, differences between individual means were determined using a post hoc test (Tukey-Kramer or the Dunnet test when one diet was considered a control diet, usually the P14-fed group). When only 2 groups were involved, data comparisons were performed using Student's t test. In Expt. 2, energy intakes (expressed as a percentage of the mean basal-day energy intakes to account for between-day and between-rat differences) were compared with a baseline of 100% (using a paired Student's t test) to clarify differences between the diets. Because a time effect was involved in Expts. 1 and 3, differences between groups were determined using mixed models for repeated measurements to test the effects of diet (or behavior in Expt. 3), time, and time x diet (or behavior) (Proc Mixed, SAS, version 6.11). Multiple comparaisons were made using ad hoc contrast (Expt. 3) under the mixed models.

	Results

TOP ABSTRACT Introduction Materials and Methods Results Discussion LITERATURE CITED

 
    Energy intake, body weight gain, and body composition in rats fed a standard yeast protein diet (Expt. 1). Until d 0, there were no differences in food intake or body weight gain between the P14 and P14-y groups, and the P14 energy intake was 407 ± 9 kJ/d. As early as d 1, a decrease in daily energy intake was observed in rats receiving the P14-y diet (412.5 ± 10.4 kJ/d and 258.2 ± 14.3 kJ/d in the P14 and P14-y groups, respectively (diet effect, P < 0.001). The reduction in energy intake occurred at the same time as a deceleration in body weight gain over the whole experimental period (diet effect, P < 0.001; Fig. 1A, B). In contrast, body weight gain did not differ between rats in the P14-y and P14-pf groups (P14-pf data are not shown in Fig. 1A, B because of too close similarities with P14-y data). Rats receiving the P14 diet had a higher final body weight than those in the other 2 groups (diet effect, P < 0.001). All organ variables measured were significantly lower in rats receiving P14-y than in those eating the P14 diet ad libitum (except for white adipose tissue in the P14-y group), whereas rats fed the P14-y diet and those that were pair fed (P14-pf group) did not differ (Table 2).

View larger version (14K): [in this window] [in a new window]   Figure 1  Energy intake (A) and body weight gain (B) of rats fed P14 or P14-y diets for 21 d (Expt. 1). Values are means ± SEM, n = 8/group. ***Mixed procedure using repeated measurements with diet effect, P < 0.001.

    Behavioral satiety sequence in rats eating standard or high protein diets containing yeast proteins (Expt. 4). In the P14 group, the 4 behaviors appeared in a well-defined sequence: eating, followed by grooming and/or activity, followed by resting (Fig. 2). Most eating occurred during the first 10 min after P14 diet presentation. The resting phase became predominant after 20 min (contrast of resting vs. other behaviors, P < 0.001) and its duration was the longest of the various behavioral satiety sequence (BSS) parameters. BSS parameters were not disturbed in the P14-y and P55-y groups. Behavioral profiles and parameters did not differ among the 3 groups. However, grooming was more important and resting was less important in the P55-y group compared with the P14-y group (diet effect, P < 0.05).

View larger version (12K): [in this window] [in a new window]   Figures 2  Temporal profile of BSS at d 14 of rats fed P14 (A), P14-y (B) and P55-y (C) diets (Expt. 4). Values are means ± SEM, n = 6/group. ***Contrast of resting vs. other behaviors with a mixed procedure using repeated measurements with behavior effect (data after 10 min), P < 0.001.

View larger version (13K): [in this window] [in a new window]   Figures 3  Kinetics of total (A), dry (B) and protein contents (C) of rat stomachs after P14, P55, and P55-y loads (Expt. 5). Total and dry contents are expressed as relative measurements for 1 g of dry diet ingested. Values are means ± SEM, n = 5. Means at a time without a common letter differ ,P < 0.05.

	Discussion

TOP ABSTRACT Introduction Materials and Methods Results Discussion LITERATURE CITED

Interestingly, yeast proteins more potently inhibited energy intake than other well-balanced amino acid proteins. Various proteins have been used during appetite-suppressing experiments and it appears that not only the protein source but also the experimental conditions (such as the time and quantity of food eaten) can interfere with the feeding response (3,6,12,13). When offered a standard protein diet (14% P/E), adult rats eating ad libitum a diet that contained yeast protein ate less, grew less, and were leaner than rats fed the standard protein (control) diet. When compared with rats pair-fed the control diet, body weight gain and relative body composition did not differ, which suggests that yeast protein acts as an appetite inhibitor and induces a reduction of body weight gain and body fat composition by depressing daily energy intake; yeast proteins, therefore, do not seem to increase energy expenditure. Yeast proteins in a high protein load induced the most marked, and a soy protein load the least marked, depression in food intake. During the various appetite-suppressing experiments described in the literature, it is still not clear what protein properties significantly affect the feeding response. During our experiment (except for yeast), the results did not differ with such biochemically contrasting proteins as soy, gluten, and total milk protein, as previously found in the literature (12,13).

One possibility for this result is that the yeast protein appetite-suppressive effect was partially induced by a low palatability. During Expt. 1, the diets differed only in terms of their oro-sensory properties due to the type of protein used and not because of the content in other macro- or micronutrients, and it can indeed be hypothesized that the yeast diet was less palatable than the control diet. Some decades ago, this question was studied by modifying the oro-sensorial properties of experimental foods by adding flavors (14), unpleasant flavors [citral or eucalyptol (15)], or unpleasant substances [such as quinine or sucro octa acetate (16,17)]. To our knowledge, after a transient decline dependent on the substance used, all groups chronically consuming ad libitum the low palatable diets ate as much as the control group, so that a putative low palatability of yeast was probably not the principal mechanism causing a reduction in energy intake.

Moreover, there are several reasons why the more marked appetite-suppressing activity of the yeast protein extract was not induced by a conditioned food aversion. First of all, the rate of ingestion was not decreased, whereas this would have been the case (after the habituation period) in the event of a typical conditioned food aversion (18,19). This characteristic was confirmed by analysis of the BSS. Indeed, when eating the normal protein diet, the 4 main behaviors appeared in the well-defined sequence of eating and grooming and/or activity and resting (7,11), and this sequence was not modified by a normal or high yeast protein diet. This sequence differed completely from that observed in the event of illness-induced anorexia, such as with LiCl injections (20). However, the short-term (1-h) energy intake of rats chronically fed with P14, P14-y or P55-y diets did not differ (unlike in Expt. 1) despite differences in the total daily energy intake. These results suggest that the decrease in food intake occurred after the first hour of the night period, and that changes occurred to the control of feeding after chronic ingestion of the yeast diets.

Our results show that the gastric kinetics of the yeast load exhibited 2 characteristics that might induce satiety signals (21): 1) a higher stomach volume (Fig. 3A), with mechanical receptors activating vagal afferents, 2) and mainly a higher dose of energy and/or specific nutrients attaining the duodenum more rapidly (Fig. 3B) and potentially sending a strong satiety signal. Indeed, rats fed loads containing yeast proteins or total milk protein exhibited different patterns of gastric emptying. The increase in gastric volume induced by yeast proteins was probably due to an increase in gastric secretions and/or the amount of water drunk (as shown in Expt. 3). This could be due to the particular content of protein extracts [mainly composed of hydrolyzed proteins, small peptides (2–5%) and free amino acids (>80%)], increasing osmotic pressure and hence, stomach secretions. Even though gastric emptying is regulated by feedback signals from the duodenum to ensure a constant flow of energy through the pylorus after a meal (22), an early, short emptying phase before initial closure of the pylorus, does not precisely follow this pattern of regulation. The pylorus remains open during the early stage of a meal, so that it can partly empty directly into the duodenum without energy regulation until the stomach volume stabilizes (23). This emptied portion has been estimated at 15–25% of the complete meal (24), which corresponds to what was obtained with the P14 and P55 loads. In contrast, half of the dry content of the yeast protein load was emptied during the meal. One hypothesis is that, during our experiment, the increase in stomach volume delayed pyloric closure so that more of the load was emptied from the stomach. This mechanism may have been triggered by a larger amount of water being consumed during the meal. This possible higher water intake with a yeast load may have changed its energetic density and therefore influenced the stomach volume of the rats and to a lesser extent the rate of emptying. However, the rate of ingestion did not differ between rats fed the control diet or the diets containing yeast, as shown in Expt. 4. The putative strong duodenal satiety signal may have been enhanced by the fact that the protein content was almost completely hydrolyzed and therefore immediately available for absorption and/or action (in the case of a peptide). Nevertheless, hydrolysis alone was not responsible for the satiety effect of this load because hydrolyzed and total proteins usually have the same effect on energy intake (13). The portion of the yeast diet that emptied very slowly after 30 min was mainly composed of partially hydrolyzed yeast fibers. These fibers, when added to a regular P14 meal, did not affect body weight gain in the rats (data not shown), suggesting that this part of the yeast extract was not responsible for the satiety properties of this diet.

Apart from these experiments, a small proportion of the yeast extract (mainly containing peptides from 1 to 10 kD) had been purified industrially. This fraction reduced energy intake, but only when the peptides had been consumed several times, unlike the yeast extract load that reduced food consumption (as from the first intake). This delayed effect suggests that this fraction may not be the only satiety agent in the yeast extract. Such a latency period tends to trigger properties related to intestinal receptors, the regulation of which results from weekly cell regeneration (25,26). Repeated intakes may therefore stimulate intestinal cells until sufficient specific receptors are available to send a signal strong enough to decrease energy intake (27). Moreover, use of a standard yeast protein load failed to reduce food intake, suggesting the involvement of a dose effect. The amount of peptides in a 2-g P14-pep load was indeed similar to a 3-g P55-y load and produced similar effects on food intake, unlike a 2-g P14-y load that contained much less peptide (10%).

The appetite-suppressant effect of the yeast extract still needs to be explained (e.g., presence of a specific amino acid or peptide acting on gastric kinetics). To further investigate the mechanisms underlying the satiety properties of yeast protein extract and peptides, we will in the future focus on the major processes associated with energy-intake depression, such as 1) vagus-mediated signals produced by increased stomach distension, higher protein, or amino acid levels in the gut, and increased amino acid metabolism in the liver, and 2) elevated peripheral plasma free amino acid or related metabolite concentrations directly monitored by specific areas of the brain. Moreover, the different brain areas probably involved in regulating energy intake (brainstem and hypothalamic area) should be investigated to determine whether yeast diets enhance their activation. Finally, digestive behavior requires further study, especially with respect to the duodenal absorption of peptides.

	FOOTNOTES

 
¹ This work received grants from the Institut National de la Recherche Agronomique (Paris, France), Institut National Agronomique Paris-Grignon (Paris, France), and Bio-Springer (Maisons-Alfort, France).

² Supplemental Table 1 is available with the online posting of this paper at jn.nutrition.org.

⁵ Abbreviations used: BSS, behavioral sequence of satiety; P/E, percentage of protein energy; P14, P14-y, and P14-pep, 14% P/E from milk, yeast, and yeast peptide protein diets, respectively; P14-pf, P14 pair-fed; P55, P55-y, P55-g and P55-s: 55% P/E from milk, yeast, gluten, and soy protein diets, respectively; WS, 100% P/E from wheat starch diet.

Manuscript received 2 January 2006. Initial review completed 8 February 2006. Revision accepted 26 May 2006.

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