Comparison of the composition and sensory characteristics of goose fatty liver obtained by overfeeding and spontaneous fattening1.

Spontaneous liver steatosis can be experimentally induced in domestic Greylag geese by combining a short photoperiod with a sequence of feed restriction followed by ad libitum corn feeding. This could offer an alternative to the conventional "foie gras" production system based on overfeeding. The present work aimed at comparing the compositional characteristics, sensory profile, and acceptability by a consumer panel of fatty livers obtained by overfeeding and spontaneous fattening. In all, 210 male geese were used: 125 geese were raised over a 31-wk period to produce fatty liver without overfeeding ("alternative livers") and 85 were raised using conventional methods with overfeeding ("conventional livers"). Mean liver weight was over 1 kg (1,102 g) in the conventional group and 445 g in the alternative group. The characteristics of the livers were studied in 2 subpopulations: 44 conventional livers representative of the experimental population (mean liver weight 1,064 g) and 42 alternative livers weighing more than 400 g (mean 702 g). Compared with the alternative livers, livers from the conventional group showed significantly (P < 0.05) higher dry matter and lipid contents, lighter color (L*), and lower yellowness (b*). The neutral lipids of alternative livers contained significantly less triglycerides and free fatty acids and significantly more cholesterol and cholesterol esters than those from conventional livers. Detailed analysis of the fatty acid composition of triglycerides showed that the proportion of mono- and polyunsaturated fatty acids was significantly higher in the alternative livers. However, covariance analysis suggested that these differences in lipid composition were mainly due to differences in lipid content between both types of livers. The evaluation of cooked livers by a trained expert panel revealed significant differences in the sensory profile between the conventional and alternative livers. The acceptability by a consumer panel was significantly lower in alternative compared to conventional livers. This difference was not related to weight and/or lipid content since livers of similar weight range (800 to 1,000 g) were compared and showed clear-cut differences for hedonic scores.

Improving residual feed intake of mule progeny of Muscovy ducks: genetic parameters and responses to selection with emphasis on carcass composition and fatty liver quality.

In France, about 95% of the fatty liver production comes from mule ducks, an infertile hybrid cross between female common ducks (Anas platyrhynchos) and Muscovy drakes (Cairina moschata). As a hybrid, genetic improvement of the performance of mule ducks is achieved by selection of the parental populations. Feed represents two-thirds of the total costs of duck production; thus, the breeders' main concern is to improve the birds' feed efficiency without impairing their capacity to be overfed after the growing period. To obtain insight into this economically important trait, we designed a divergent selection experiment for Muscovy sires on the basis of the residual feed intake (RFI) of their male mule progeny. Residual feed intake represents the fraction of feed intake that cannot be explained by maintenance and production requirements. Data comprised records from generations 0 to 2 with 227 Muscovy sires of 2,016 mule ducks. Growth and fat deposition were measured individually for all mule ducks. Half-sib mule ducks were placed together in 1 small pen, and the feed consumption was recorded for each pen; all half-sibs therefore had the same feed intake estimates. Heritability of RFI was high (0.83 ± 0.42), and after 2 generations of selection, the differences between lines in terms of RFI (contrast of 9.13 g/d, representing 0.76 phenotypic standard deviation; P < 0.0001) and feed conversion ratio (contrast of 0.08, representing 0.28 phenotypic standard deviation; P = 0.0003) were marked. In regard to carcass traits, the low-RFI line (high-efficiency animals) had heavier legs (476 vs. 463 g; P < 0.0001) and magret muscles (268 vs. 262 g; P = 0.001) than the high-RFI line (low-efficiency animals). No differences were observed for either the liver weight or the melting rate between the 2 lines (P > 0.46). Selection on RFI thus improves the feed efficiency of animals without impairing their capacity to produce fatty liver.

Is ability to hepatic steatosis influenced by age at the beginning of the overfeeding period in Muscovy and Pekin ducks?

The aim of this study was to analyse the effects of species (Muscovy and Pekin ducks) and age at the beginning of the overfeeding period on fatty liver production, carcass composition and lipid and moisture content of the liver and breast muscle. We reared four groups of 40 ducks per species for the study, starting at 2-week intervals in order to have four different ages together at the beginning of the overfeeding period (10, 12, 14 and 16 weeks). At the end of the overfeeding period, all ducks were slaughtered. Our results confirmed the high levels of difference in carcass composition and lipid content in the plasma, liver and breast muscle between Muscovy and Pekin ducks at all ages. Pekin ducks were not able to develop a high degree of hepatic steatosis, but had increased lipid storage in peripheral adipose and muscle tissues than Muscovy ducks. However, the fatty liver weight of Pekin ducks increased with age, with lipid deposition in the liver and peripheral tissues. The ability of Muscovy ducks to produce fatty livers remained unchanged with age in line, with lipid deposition in the liver and peripheral tissues. The sites of lipid deposition thus depend on species and not on the physiological maturity of ducks.

Detection of QTL controlling metabolism, meat quality, and liver quality traits of the overfed interspecific hybrid mule duck.

The mule duck, an interspecific hybrid obtained by crossing common duck (Anas platyrhynchos) females with Muscovy (Cairina moschata) drakes, is widely used for fatty liver production. The purpose of the present study was to detect and map single and pleiotropic QTL that segregate in the common duck species, and influence the expression of traits in their overfed mule duck offspring. To this end, we generated a common duck backcross (BC) population by crossing Kaiya and heavy Pekin experimental lines, which differ notably in regard to the BW and overfeeding ability of their mule progeny. The BC females were mated to Muscovy drakes and, on average, 4 male mule ducks hatched per BC female (1600 in total) and were measured for growth, metabolism during growth and the overfeeding period, overfeeding ability, and the quality of their breast meat and fatty liver. The phenotypic value of BC females was estimated for each trait by assigning to each female the mean value of the phenotypes of her offspring. Estimations allowed for variance, which depended on the number of male offspring per BC and the heritability of the trait considered. The genetic map used for QTL detection consisted of 91 microsatellite markers aggregated into 16 linkage groups (LG) covering a total of 778 cM. Twenty-two QTL were found to be significant at the 1% chromosome-wide threshold level using the single-trait detection option of the QTLMap software. Most of the QTL detected were related to the quality of breast meat and fatty liver: QTL for meat pH 20 min post mortem were mapped to LG4 (at the 1% genome-wide significance level), and QTL for meat lipid content and cooking losses were mapped to LG2a. The QTL related to fatty liver weight and liver protein and lipid content were for the most part detected on LG2c and LG9. Multitrait analysis highlighted the pleiotropic effects of QTL in these chromosome regions. Apart from the strong QTL for plasma triglyceride content at the end of the overfeeding period mapped to chromosome Z using single-trait analysis, all metabolic trait QTL were detected with the multitrait approach: the QTL mapped to LG14 and LG21 affected the plasma cholesterol and triglyceride contents, whereas the QTL mapped to LG2a seemed to impact glycemia and the basal plasma corticosterone content. A greater density genetic map will be needed to further fine map the QTL.

Duration of transport and holding in lairage at constant postprandial delay to slaughter--effects on fatty liver and breast muscle quality in mule ducks.

The present study was designed to evaluate the effect of preslaughter transport (30 vs. 150 min) and holding of mule ducks in lairage in their transport crates (15 vs. 120 min) on the quality of the meat and fatty liver. A total of 120 birds were allocated in a 2 × 2 factorial design with a constant postprandial delay to slaughter (8 h), to avoid the confounding between the effects of the experimental treatments and those of fasting duration. Under such conditions, extending the transport or holding duration did not induce a loss in preslaughter live weight or liver weight. Similarly, breast muscle glycogen stores were not affected by the treatments, nor was the kinetics of postmortem pH decline affected. The mechanical resistance of raw meat obtained by the compression test significantly increased with holding duration. The gross chemical composition of the livers did not differ significantly among the preslaughter treatments. Residual blood in the liver, as indicated by heme pigment concentration, was enhanced with a longer transport, but this effect was more pronounced after the longest holding duration, as shown by a significant interaction. This, however, did not significantly affect the incidence of appearance defects or the commercial grading of the livers. The percentage of fat loss during the cooking of canned livers was significantly reduced when the transport duration was increased. This effect could not be explained on the basis of the current knowledge for determining the technological quality of fatty liver. The identification of biological markers of liver quality is currently underway in our laboratory. Further investigations studying the differential expression of these biological markers according to preslaughter conditions would provide a better understanding of the effect of transport duration on liver processing yield.

Genetic parameters of product quality and hepatic metabolism in fattened mule ducks.

Genetic parameters of traits related to hepatic lipid metabolism, carcass composition, and product quality of overfed mule ducks were estimated on both parental lines of this hybrid: the common duck line for the maternal side and the Muscovy line for the paternal side. The originality of the statistical model was to include simultaneously the additive genetic effect of the common ducks and that of the Muscovy ducks, revealing a greater genetic determinism in common than in Muscovy. Plasma metabolic indicators (glucose, triglyceride, and cholesterol contents) were heritable, in particular at the end of the overfeeding period, and heritabilities increased with the overfeeding stage. Carcass composition traits were highly heritable in the common line, with values ranging from 0.15 for liver weight, 0.21 for carcass weight, and 0.25 for abdominal fat weight to 0.32 for breast muscle weight. Heritabilities of technological outputs were greater for the fatty liver (0.19 and 0.08, respectively, on common and Muscovy sides for liver melting rate) than for the pectoralis major muscle (between 0.02 and 0.05 on both parental sides for cooking losses). Fortunately, the processing industry is mainly facing problems in liver quality, such as too high of a melting rate, than in meat quality. The meat quality appraisal criteria (such as texture and cooking losses), usually dependent on pH and the rate of decline of pH, were also very lowly heritable. This study demonstrated that genetic determinism of meat quality and ability of overfeeding is not similar in the common population and in the Muscovy population; traits related to fattening, muscle development, and BW have heritability values from 2 to 4 times greater on the common line than on the Muscovy line, which is relevant for considering different selection strategies.

Is there a relationship between the kinetics of lipoprotein lipase activity after a meal and the susceptibility to hepatic steatosis development in ducks?

The difference in the ability of Pekin and Muscovy ducks to develop hepatic steatosis could result from a different peripheral lipoprotein lipase (LPL) activity, which hydrolyses triacylglycerol secreted by the liver. We studied the kinetics of plasma LPL activity in response to a meal at different ages in Pekin and Muscovy ducks. For that purpose, blood samples were taken at 5, 9, 12, 13, and 14 wk of age just before and 1, 2, 4, and 8 h after a meal. To release LPL into general circulation, an i.v. injection of heparin (400 IU/kg of BW) was administered 10 min before blood collection. For that reason, different ducks per genotype were used for each point of measurement (n = 6). Plasma LPL activity measured before the meal was negatively correlated with the weight of the fatty liver measured in the same ducks at 14 wk of age (r = -0.58, P < 0.001). Plasma triacylglycerol level measured before the meal was negatively correlated with plasma LPL activity measured in the same ducks (r = -0.31, P = 0.025) and was negatively correlated with plasma LPL activity measured in the same ducks for each age and each timing (r = -0.39, P < 0.001). At 14 wk of age for Muscovy and Pekin ducks, we observed that a high plasma LPL activity (>200 IU/L of plasma) corresponded to a relatively low development of fatty liver (190 g) induced by overfeeding, whereas a low plasma LPL activity (<150 IU/L of plasma) corresponded to a high propensity to develop fatty liver (470 g). In conclusion, plasma LPL activity measured just before the meal during the rearing period could be used as a marker of hepatic steatosis development during the overfeeding period.

Comparison of nonlinear and spline regression models for describing mule duck growth curves.

This study compared models for growth (BW) before overfeeding period for male mule duck data from 7 families of a QTL experimental design. Four nonlinear models (Gompertz, logistic, Richards, and Weibull) and a spline linear regression model were used. This study compared fixed and mixed effects models to analyze growth. The Akaike information criterion was used to evaluate these alternative models. Among the nonlinear models, the mixed effects Weibull model had the best overall fit. Two parameters, the asymptotic weight and the inflexion point age, were considered random variables associated with individuals in the mixed models. In our study, asymptotic weight had a greater effect in Akaike's information criterion reduction than inflexion point age. In this data set, the between-ducks variability was mostly explained by asymptotic BW. Comparing fixed with mixed effects models, the residual SD was reduced in about 55% in the latter, pointing out the improvement in the accuracy of estimated parameters. The mixed effects spline regression model was the second best model. Given the piecewise nature of growth, this model is able to capture different growth patterns, even with data collected beyond the asymptotic BW.

Behavioral and adrenal responses to various stressors in mule ducks from different commercial genetic selection schemes and their respective parental genotypes.

The mule duck, a hybrid produced by crossing a Muscovy drake and a Pekin female, is reported to express inappropriate behavior such as collective avoidance of people, the resulting distress and physical consequences potentially compromising their welfare. The present study was carried out to characterize the responses of mule duck strains from different commercial selection schemes to various stressful conditions and to confirm previous data on the genetic cross effects observed in a specific genotype. Three independent experiments were conducted with ducks from 3 French breeding companies (A, B, and C). Each experiment compared 2 mule genotypes sharing one common parental origin (paternal for ducks from company A or maternal for ducks from companies B and C). Mule duck males from the 2 genotypes and their respective parental genotypes (Pekin and Muscovy) were subjected to a set of social and stressful physiological and behavioral tests. Previously reported differences in genetic cross effects on fear responses between the parental genotypes and the corresponding hybrid were confirmed in these commercial crosses. Both mule duck and Pekin genotypes showed more active physiological and behavioral responses to stress than Muscovy genotypes. The new finding of this study is that mule genotypes appear to be more sensitive to the social environment than both respective parental genotypes. Few differences were observed between the 2 mule genotypes from A and C. On the other hand, several traits of the 2 mule genotypes from B differed. In addition, A and C mule genotypes were characterized by the same adrenal and behavioral traits but contrasting responses. The B mule genotypes were characterized by a different set of behavioral traits, and only 1 of the 2 B mule ducks was characterized by a group of adrenal traits.

Influence of lipoprotein-lipase activity on plasma triacylglycerol concentration and lipid storage in three genotypes of ducks.

The lipoprotein-lipase (LPL) hydrolyses the triacylglycerols (TG) secreted by the liver and, thus, allows the storage of lipids onto the extrahepatic tissues. The LPL activity has been studied by injection of LPL antibodies in three genotypes of ducks (Muscovy (Cairina moschat), Pekin (Anas plathyrhynchos) and Mule (hybrids of male Muscovy ducks and female Pekin ducks)) under overfeeding condition. The results show a similar weight gain between injected and control animals. A higher liver steatosis is observed in Mule ducks (616+/-18 g; 8.79% of body mass (BW)) and Muscovy ducks (514+/-13 g; 7.05% BW) compared to Pekin ducks (353+/-21 g; 5.89% BW, p<0.05). Pekin ducks showed a much marked extrahepatic fattening of abdominal and subcutaneous adipose tissues. The LPL activity was evaluated by comparing the evolution of the plasma TG concentrations after injections of saline (control animals) or injections of specific LPL-antibodies. Inhibition of LPL activity performed by intravenous injections of LPL-antibodies showed a spectacular increase in the plasma TG concentrations in the three genotypes. That increase was considerably higher in Pekin ducks (98+/-10 g/L) compared to Muscovy ducks (35+/-2 g/L, p<0.01) and Mule ducks (30+/-4 g/L, p<0.01). Those data suggest that a high export of lipids synthesized in liver and a high LPL activity occur in overfed Pekin ducks, which can favour the extrahepatic fattening to the detriment of the liver steatosis, and conversely in overfed Muscovy and Mule ducks.

Expression of the glucokinase gene in mule duck liver and glucokinase activities in chicken and mule duck livers.

The presence of glucokinase (GK), a critical enzyme controlling glucose homeostasis, particularly liver glucose utilization in mammals, has long been a matter of debate in avian species because a number of investigators have failed to detect GK activity in the livers of chickens and several other avian species. In this study, we cloned a partial GK cDNA from mule duck livers and measured GK-like activity in the livers of mule ducks and broiler chickens under 2 nutritional states. Liver samples from 5-wk-old meal-fed male broiler chickens (Ross) were obtained from overnight-fasted chickens (BC) and 5 h after an oral saccharose load (6 mL/kg of BW of a 50% saccharose solution) given just before the meal (BS). Liver samples from 15-wk-old mule ducks were collected after an overnight fast (DC) and 12 h after the last overfeeding meal (DO). A partial cDNA ( approximately 600 bp) was obtained from duck livers. It presented 99% identity with chicken partial GK cDNA (gi 44888789) and 82% identity with human GK (gi 15967158). Chicken liver weights represented 1.8 and 3.3% of BW, respectively, for BC and BS (n = 8, P < 0.05). Glucokinase and low-Michaelis constant hexokinase (HK) activity levels were similar in BC (respectively, 0.88 and 1.00 mU/mg of protein). In response to the meal load, GK activity increased significantly (+57%), whereas HK decreased (-46%) in BS. Duck liver weights represented 1.4 and 7.6% of BW, respectively, for DC and DO (n = 8, P < 0.05). In DC livers, GK activity was significantly higher than HK activity (respectively, 1.76 and 0.63 mU/mg of protein). Both activities were significantly increased in DO (2 times, n = 8, P < 0.05). In conclusion, GK is present in ducks as well as chickens, and it is nutritionally regulated in avian species as well as in mammals. Further work will determine whether the higher liver GK activity and GK:HK ratio in DC compared with BC is related to age or BW or linked to the high lipogenic capacity of the duck liver.

Do age and feeding levels have comparable effects on fat deposition in breast muscle of mule ducks?

The effects of age (from 1 day post-hatch to 98 days of age) and feeding levels (feed restriction followed by overfeeding v. ad libitum feeding) on lipid deposition in breast muscle (quantity and quality, localisation) of mule ducks were determined in relation to muscle energy metabolism (glycolytic and oxidative), plasma levels of lipids, glucose and insulin, and muscle capacity for lipid uptake (characterised by lipoprotein lipase (LPL) activity). Two periods were defined for age effects on intramuscular lipids in breast muscle: - 1 to 42 days of age when lipids (mainly phospholipids and cholesterol provided by egg yolk) stored in the adipocytes during embryonic life were transferred to the muscle fibres and used for growth and energy requirements, - 42 to 98 days of age when the muscle again stored lipids (mainly triglycerides provided by liver lipogenesis), first in fibres and then in adipocytes.Plasma glucose and insulin levels were not affected by age. Plasma levels of lipids and LPL activity in breast muscle were high at 1 and 14 days of age and then decreased, remaining stable until 98 days of age. Energy metabolism activity in the breast muscle (mainly glycolytic activity) increased with age.Feed restriction, corresponding to 79% of ad libitum intake, applied between 42 and 75 days of age only resulted in decreases in plasma insulin concentration and total lipid content of breast muscle, mainly affecting triglyceride and mono-unsaturated fatty acid (MUFA) levels. Overfeeding increased plasma levels of insulin and lipids while glycaemia remained stable. LPL activity and total lipid levels increased in breast muscle, mainly induced by deposition of triglycerides and MUFA occurring particularly during the 2nd week of this period. Glycolytic energy metabolism decreased.In response to age or feeding levels, muscle lipid levels and composition reflect plasma lipid levels and composition and high muscle lipid levels stimulate oxidative energy metabolism.

Effects of intramuscular fat levels on sensory characteristics of duck breast meat.

We conducted a study to evaluate the effects of intramuscular fat levels on the sensory characteristics of duck breast meat. Combining duck genotypes (Muscovy, Pekin, and their crossbreed hinny and mule ducks) and feeding levels (overfeeding between 12 and 14 wk of age vs. ad libitum feeding) enabled us to obtain a wide range of lipid levels in breast muscle. The average values were between 2.55 and 6.40 g per 100 g of muscle. Breast muscle from overfed ducks showed higher lipid and lower water levels than breast muscle from ducks fed ad libitum. Muscle from the overfed ducks was also paler in color and exhibited greater yellowness and cooking loss values. Juiciness was judged lower and flavor more pronounced in overfed ducks. Muscovy ducks exhibited higher breast weight and lower lipid levels than the other genotypes. At the other extreme, Pekin ducks exhibited the highest lipid levels and the lowest breast weights; values for these criteria were intermediate in hinny and mule ducks. Breast muscle of Muscovy ducks was paler, less red, and more yellow than that of other genotypes. Breast muscle of Pekin ducks exhibited the lowest values for lightness, yellowness, and energy necessary to shear meat, as well as the highest cooking loss values, and was judged more tender, juicy and less stringy than that of other genotypes. In contrast, scores for breast muscle of Muscovy ducks were the lowest for tenderness, juiciness, and flavor, and the highest for stringiness. Breast muscle of hinny and mule ducks scored the highest values for redness. Hinny ducks also scored the highest values for flavor. Genotype exerted a higher effect on the sensory quality of breast muscle than did feeding levels. Finally, increasing lipid levels in breast muscle increased lightness, yellowness, cooking loss, tenderness, and flavor, with correlation coefficients of 0.49, 0.47, 0.54, 0.43, and 0.28, respectively. However, breast meat color and tenderness were mainly influenced by genotype.

Effects of dietary fructose on liver steatosis in overfed mule ducks.

Overfeeding of some waterfowl species results in obesity, which is mainly characterized by a dramatic hepatic steatosis induced by strong accumulation of lipids synthesized from dietary glucose in the liver. In mammals, fructose is known to be able to raise plasma triacylglycerol concentrations significantly; consequently, this may induce obesity. The aim of this study was to assess the effect of partial replacement of dietary glucose provided by corn starch with fructose on metabolism and fatty liver production in the Mule ducks. On the basis of 9.5 kg maize (132,920 kJ) given twice a day for 14 days, a supplementation of 9,800 kJ was provided in form of glucose, sucrose or high fructose corn syrup (HFCS: 50 % glucose, 42 % fructose and 8 % other saccharides). Fatty liver weight in ducks fed with glucose supplementation was 499 +/- 21 g. Sucrose or HFCS supplementation brought about a significant increase in liver weight (+ 18.7 % and + 16.3 % vs. glucose supplementation respectively, p < 0.05). These results suggest that the dietary fructose favors the liver steatosis by increasing hepatic lipogenesis. Postprandial plasma insulin concentrations were similar in ducks fed diets with or without fructose, suggesting that the effect of fructose on liver steatosis is not mediated by insulin.