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The location and utilization of quantitative trait loci (QTL) and candidate genes with significant effects on economically important traits are becoming increasingly important in livestock breeding programs. The porcine cholecystokinin type A receptor (CCKAR) is a candidate gene for performance traits, due to its known role in the physiological control of feed intake, satiety, and obesity. We investigated the association of CCKAR polymorphisms with feeding, growth, and efficiency traits in an F2 population derived from a cross between Meishan and Large White founder animals and in lines of Large White pigs that had been divergently selected on the basis of lean growth efficiency traits. In the F2 population, CCKAR genotype was significantly associated with daily feed intake and average daily gain. The effects of the polymorphisms were then assessed in a larger-scale analysis of segregating commercial lines. A newly discovered single-nucleotide polymorphism (SNP) within the 5′-untranslated region (5′-UTR) had highly significant effects on feed intake, average daily gain, and days to 110 kg, which were not seen for a previously reported SNP within the CCKAR gene. Furthermore, we provide evidence that the novel SNP disrupts the binding of the YY1 transcription factor, which raises the possibility that it is the causal variant. The 5′-UTR SNP could be utilized as a molecular genetic test for increased feed intake, faster lean growth, and reduced days to market weight in segregating commercial lines.

Associations between productivity and a range of immune traits were tested by multiple regression analysis on 128 Large White pigs (62 male, 66 female). Daily weight gain, daily feed intake, and efficiency (i.e. weight gain/feed intake), assessed from 14 to 24 weeks of age, were the productivity traits. Total and differential white blood cell count and leukocyte sub-sets positive for CD4, CD8, CD11R1, gamma delta T cells, B cell, and monocyte markers, all measured at 18 and 24 weeks, were the immune traits. At 24 weeks of age, higher percentages of monocytes were associated with a decrease in daily weight gain. Higher percentages of B cells were associated with a decrease in daily feed intake. Higher percentages of CD11R1 positive cells were associated with a decrease in daily weight gain and a decrease in efficiency. Relationships at the age of 18 weeks were similar, but slightly less significant. Associations between the numbers of monocytes, MIL-4 positive cells and B cells and certain performance traits were also observed at the age of 24 weeks. Overall, these results indicate an association between productivity and certain immune traits. We suggest that the observed associations between these immune traits and performance could be due to the impact of sub-clinical infections.

Amplified fragment length polymorphic (AFLP®) markers were used to discriminate between lines of pigs, divergently selected over seven generations for components of efficient lean growth rate. A total of 270 animals with 30 animals per line were genotyped for 239 polymorphic AFLP markers. Canonical variate analysis identified linear combinations of the AFLP marker scores that grouped animals by selection line with no overlap between selection lines. Cluster analysis of AFLP marker scores identified 10 groups of animals with 226 of the 270 animals clustered into nine groups, each consisting of animals from only one selection line. AFLP marker genotyping, using the EcoRI and TaqI restriction enzymes, provided an effective means of discriminating between animals of different selection lines that have arisen from one base population.

Summary Divergent selection over seven generations for daily food intake (DFI) and for two lean growth rate (LGA and LGS) selection objectives in pigs produced correlated responses in serum insulin-like growth factor-1 (IGF-1) concentration. The LGA and LGS selection objectives were for lean growth, but with performance testing on ad libitum (LGA) or on restricted (LGS) feeding regimes. There were 342 animals in the study, with an average of 46 pigs in each selection line and 72 pigs in the control line. All animals were performance-tested from 30 to 90 kg on an ad libitum feeding regime. Selection on DFI increased IGF-1 at 6 weeks of age (158 versus 104 μg/l, SED 21; control 136 μg/l) and selection for high LGA increased IGF-1 at 30 kg liveweight (202 versus 161 μg/l, SED 14; control 185 μg/l). At 6 weeks of age and at 30 kg of weight, there was no response in IGF-1 with selection on LGS (128 versus 129 and 167 versus 153 μg/l, respectively) suggesting that IGF-1 may not be a reliable physiological predictor of genetic merit for such a selection objective, which combined rate and efficiency of lean growth. IGF-1 measured immediately prior to the start of performance test may be useful in determining which animals to performance test for a lean growth selection objective, whereas IGF-1 measured at 6 weeks of age may be a useful physiological predictor of genetic merit for daily food intake during performance test. Zusammenfassung Serumkonzentrationen des insulinahnlichen Wachstumsfaktors-1 bei Schweinen, die entweder auf tagliche Futteraufnahme oder auf hohe Wachstumsrate fur Magerfleischanteil selektiert wurden Die Selektion von Schweinen uber sieben Generationen einerseits auf tagliche Futteraufnahme (DFI) und andererseits auf zwei Wachstumsraten fur Magerfleischanteil (LGA, ad-libitum-Futterung; LGS, rationierte Futterung) ergab korrelierende Konzentrationen am insulinahnlichen Wachstumsfaktor (IGF-1). An der Studie waren 342 Tiere mit einer durchschnittlichen Tierzahl von 46 Schweinen pro Selektionsgruppe und 72 Schweinen als Kontrollgruppe beteiligt. Alle Tiere wurden zwischen 30 und 90 kg bei der ad-libitum Futterung leistungsgepruft. Die Selektion auf tagliche Futteraufnahme (DFI) erhohte die IGF-1 Konzentrationen in der 6. Lebenswoche von 104 μg/l auf 158 μg/l (Kontrollgruppe 136 μg/l) und die Selektion auf LGA bei der ad-libitum-Futterung bei einem Lebendgewicht von 30 kg von 161 μg/l auf 202 μg/l (Kontrollgruppe 185 μg/l). In der 6. Lebenswoche und bei 30 kg Lebendgewicht gab es keinen Anstieg der IGF-1 Konzentrationen in der Selektionsgruppe LGS mit rationierter Futterung (128 μg/l bzw. 129 μfg/l in der 6. Lebenswoche; 167 μg/l bzw. 153 μg/l bei 30 kg),was darauf hinweisen konnte, dass IGF-1 wahrscheinlich kein verlasslicher, physiologischer Marker fur den genetischen Erfolg fur ein solches Selektionsziel, welches die Wachstumsrate und die Effizienz beim Magerfleischwachstum kombiniert, darstellt. Die direkt vor Prufungsbeginn ermittelten IGF-1 Konzentrationen konnten bei der Auswahl der Tiere, die an einer Leistungsprufung fur die Wachstumsrate des Magerfleischanteils als Selektionsziel herangezogen werden sollten, hilfreich sein. Die IGF-1 Konzentrationen, die in der 6. Lebenswoche gemessen werden, sind wahrscheinlich ein hilfreicher physiologischer Marker fur den genetischen Erfolg fur DFI wahrend einer Leistungsprufung.

The sensitivity to dietary lysine: energy content of pigs from lines divergently selected over seven generations for components of efficient lean growth rate was quantified by the within-selection line regression of performance test traits on diet class. Large White pigs were performance tested over three 14-day test-periods starting at 30, 50 and 75 kg and offered, to appetite, isoenergetic diets differing in dietary total lysine: energy (0•59, 0•69, 0•81, 0•91, 1•01, 1•12 and 1•23 g/MJ digestible energy (DE)). Within each litter, full-sibs were performance tested on different diets. Pigs were also performance tested on a diet-choice procedure using diets with total lysine: energy of 0•69 and 1•12 g/MJ DE to determine the correlation between performance on phase-feeding and diet-choice.The study consisted of 578 animals with 459 pigs tested with phase-feeding and 119 tested on diet-choice procedures. The study detected significant selection line and dietary effects on performance test traits, but no significant between-selection line differences in sensitivity to dietary total lysine: energy. When combinations of performance test traits were transformed into predicted lysine and energy utilization traits there were significant selection line effects on predicted nutrient allocation, but not in responses to increasing dietary total lysine: energy. The lack of between-selection line differences in sensitivity to dietary total lysine: energy indicated that the consequences of changing dietary total lysine: energy will be broadly similar across a range of genotypes, as spanned by the selection lines of the study.Genetic correlations between performance traits of phase-fed and diet-choice pigs suggested that predictions of genetic merit will be similar with animals tested on either procedure between 30 and 75 kg but post 75 kg predicted genetic merit for growth rate and lysine intake should be estimated separately for performance on diet-choice or for performance on a given diet. In contrast, predicted genetic merit for fat deposition with performance testing on diet-choice will be highly correlated with predicted genetic merit with testing on a single diet.

Correlated responses in physiological traits to divergent selection on components of efficient lean growth identified physiological traits for use as predictors of genetic merit and provided a biological explanation for the between-selection line differences in protein and lipid deposition. Responses (differences between high (H) and low (L) selection lines) in protein metabolism during ad-libitum feeding were associated with divergent selection for daily food intake (DFI) (reduced serum creatinine concentration (H = 1·30 v. L = 1·56, s.e.d. 0·08 mg/dl)) and for lean growth rate on an ad-libitum feeding regime (LGA) (increased serum urea (H = 48 v. L = 36, s.e.d. 4 mg/dl) and creatinine (H = 1·74 v. L = 1·45 mg/dl)) concentrations, but not with selection for lean growth on a restricted feeding regime (LGS). Following 24-h fasting, responses in lipid metabolism, in the form of higher serum non-esterified fatty acid concentrations, were detected with divergent selection for both LGA (399 v. 248, s.e.d. 66 µmol/l) and LGS (H = 361 v. L = 107 µmol/l). The high LGS line appeared to ‘preserve’ protein to a greater extent than the high LGA line and similarly there was greater maintenance of lipid depots by the low LGS line compared with the low LGA line. A tentative ranking of the two pairs of lean growth selection lines on the basis of ‘importance’ of protein deposition would be high LGS, high LGA, low LGA and low LGS.Coheritabilities for serum creatinine concentration with predicted lysine balance and lysine required for protein deposition (-0·17 and 0·17, s.e. 0·08) indicated that serum creatinine concentration may usefully be included in breeding value prediction for lysine requirement and protein deposition to increase the accuracy of predicted genetic merit.Fasting did not increase the coheritabilities for serum creatinine concentration, so inclusion of serum creatinine concentration in a selection criterion for dietary lysine requirement or protein deposition does not require withdrawal of food before blood sampling animals.

Serum urea concentrations were measured in Large White pigs from lines divergently selected for components of efficient lean growth rate and performance tested over three 14-d test periods starting at 30, 50, and 75 kg. Two methods of performance testing were used. Phase-fed pigs were fed to appetite isoenergetic diets differing in total lysine:energy ratio (0.58, 0.69, 0.81, 0.91, 1.01, 1.12, and 1.23 g/MJ of digestible energy), whereas diet-choice pigs were offered a choice of the 0.69 and 1.12 lysine:energy diets. Between test periods, all animals were fed one diet: 0.91 g of lysine/ MJ of digestible energy. The study consisted of 230 boars and gilts with 150 pigs performance tested on phase-feeding and 80 pigs on diet-choice. The line selected for high lean food conversion had lower urea concentrations on each diet than the line selected for high lean growth rate, despite similar predicted lysine balances. Efficiency of lean growth rather than the rate of lean growth may be a better selection strategy in the context of nitrogen excretion. Urea concentrations at the end of each test period were correlated with lysine intake (0.33, 0.48 and 0.65; standard error, 0.08) and predicted lysine balance (0.39, 0.44, and 0.64), but were uncorrelated with predicted lysine for protein deposition (0.01, 0.08, and 0.08) and maintenance. Urea concentration at the end of a test period was not a useful predictor of protein deposition, even after accounting for pretest variation in urea concentration and food intake during test. The expected response pattern of serum urea concentration to diets differing in total lysine:energy would be nonlinear, with the point of inflection occurring at the required dietary total lysine:energy for each genotype. However, there was no evidence of such an inflection point such that the prediction of lysine requirement from urea concentration was not possible for the selection lines in the study.

Lactational performance was measured in Large White gilts from lines that had been divergently selected over seven generations for daily food intake (DFI), lean food conversion ratio (LFC), lean growth rate with ad-libitum (LGA) or restricted (LGS) feeding during performance test. Control line gilts were included in the study. During the lactation period of 21 days, gilts were given to appetite five isoenergetic diets differing in ileal digestible lysine: energy (0·40, 0·58, 0·76, 0·94 and 1·12 g/MJ digestible energy). The study consisted of 223 gilts with a similar number of animals in each selection line.Live-weight loss was greater in the low LFC and LGA lines than in the high lines and food intake was significantly lower in the low LGA line than in the high line. Litter-weight gain of the low LFC and high LGA lines were greater than in the complementary selection lines. Prediction equations for nutrient utilization were used to express the effect of diet and selection line in terms of energy and protein inputs and outputs. Selection on DFI or LGS resulted in gilts that did not mobilize lipid during lactation as sufficient energy for milk and maintenance was provided by dietary intake. In contrast, there was insufficient dietary energy with selection on LFC or on low LGA so lipid mobilization was required to achieve energy balance. The energy required to excrete excess protein and energy from lipid mobilization increased as the dietary lysine energy ratio increased, but there were no other dietary effects on energy and protein utilization. Genotype with nutrition interactions were detected for energy intake and lipid mobilization, which were due to the lines selected for low DFI and LGA. The general absence of genotype with nutrition interactions for lactational performance in gilts selected for components of efficient lean growth and the lack of significant dietary effects on energy utilization indicated that the consequences of changing nutritional inputs will be broadly similar for genotypes within the set of genotypes studied. Selection strategies which result in reduced food intake during lactation should be avoided if lipid mobilization is then required to attain energy balance and there are resultant negative effects on subsequent reproductive performance.

The sensitivity of pigs selected for high daily food intake (DFI), low lean food conversion (LFC) and high lean growth rate (LGS) to dietary change of plasma fatty acids was assessed. The difference between the two diets was effectively a substitution of palmitic (C16:0), oleic (C18:1 n-9) and linoleic (C18:2 n-6) with linolenic (C18:3 n-3) fatty acids. Fatty acid compositions of plasma free fatty acids, neutral lipids and phospholipids were measured in 90 kg animals fed a base or high linolenic (C18:3 n-3) fatty acid diet, based on whole linseed, for four days. There were 24 animals from each selection line and 24 animals from an unselected control line, with boars and gilts represented equally in each line. Half of the selection line animals were fed the base diet and half were fed the high C18:3 diet, but all control animals were fed the base diet. Prior to slaughter, animals were fasted for 18 h. The fatty acids primarily affected by dietary change were C18:3 n-3 and its products, C20:5 n-3 (EPA) and C22:6 n-3 (DHA). The sensitivity of a selection line to dietary change was parameterised by the relative shift in fatty acid composition through changing from the base diet to the high C18:3 diet. In neutral lipids, the sensitivities of C18:3 n-3 in the DFI and LFC lines were similar but greater than in the LGS line (3.0 v. 1.8, S.E.D. 0.15), while, for phospholipid and free fatty acids, the sensitivity of the DFI line was greater than in the LFC and LGS lines (2.3 v. 1.8 and 2.0 v. 1.4, respectively). For C20:5 n-3 and C22:6 n-3, the DFI and LFC lines were more sensitive to dietary change than the LGS line (total lipid : 2.3 v. 1.9 and 1.5 v. 1.2). In general, the DFI line was most sensitive to dietary change and the LGS line was the least sensitive. The difference in sensitivities of the lipid classes to the high C18:3 diet between the selection lines could result from differences in body fat content and may explain the general lack of genotype with nutrition interactions in post-1990 genotypes in comparison with fatter circa-1970 genotypes.

The effects of genotype, diet and the genotype with diet interaction on fatty acid composition of neutral lipid and phospholipid of intramuscular fat in pigs were examined. Pigs from four selection lines and a control line were fed isoenergetic diets, which differed in ileal digestible lysine:digestible energy (A: 0.40, C: 0.76 and E: 1.12 g lysine/MJ DE). The selected lines resulted from seven generations of selection for high daily food intake (DFI), lean food conversion ratio (LFC) and lean growth rate on ad-libitum (LGA) or restricted feeding regimes in a Large White population. During performance test, from 30 to 90 kg, pigs were fed on either ad-libitum or restricted (0.75 g/g ad-libitum daily food intake) feeding regimes. A trained sensory panel assessed intensities of pork flavour and abnormal flavour and the hedonic characteristics of flavour liking and overall acceptability. The proportion of neutral lipid in the LGA line was lower (0.64 v 0.75, s.e.d. 0.03) while the polyunsaturated:saturated (P:S) ratio of the LFC line (0.53 v 0.41, s.e.d. 0.05) was higher than the control line and other selection lines. The DFI line had a higher phospholipid n-6:n-3 ratio than other selection lines (14 v 11, s.e.d. 1.0). Diets C and E resulted in lower neutral lipid:phospholipid ratio (0.7 v 0.8, s.e.d. 0.01), n-6:n-3 ratios for both neutral lipid (5 v 13, s.e.d. 0.8) and phospholipids (7 v 20, s.e.d. 0.6) and a higher P:S ratio (0.5 v 0.3, s.e.d. 0.03) for neutral lipid than diet A. Neutral lipid fatty acids C18:2 n-6, C18:3 n-3, C20:3 n-6, C20:4 n-6, C20:5 n-3, C22:5 n-3 and C22:6 n-3 were negatively correlated with pork flavour (-0.30, s.e.d. 0.10), flavour liking (-0.33) and overall acceptability (-0.30). In contrast, the fatty acids C16:1, C18:1 ω9 and C18:1 ω11 were positively correlated with pork flavour (0.36), flavour liking (0.39) and overall acceptability (0.40). However, correlations for fatty acids in the phospholipid class were positive for C18:2 n-6, C20:4 n-6 and C22:4 n-6 with pork flavour (0.33), flavour liking (0.23) and overall acceptability (0.23). The proportion of neutral lipid had non-significant correlations with flavour traits; pork flavour (0.01), abnormal flavour (0.08), flavour liking (-0.03) and overall acceptability (0.03). The selection lines responded in a similar manner to the different diets, such that there was little evidence for genotype with nutrition interactions for fatty acid concentrations of neutral lipids and phospholipids. Selection for high lean growth will reduce intramuscular fat, but the increased relative amount of phospholipid and concentration of polyunsaturated fatty acids are unlikely to reduce eating quality. Nutritional effects on intramuscular fat characteristics were greater than genetic effects, such that nutritional approaches to feeding pigs will provide effective methods of reducing the n-6:n-3 fatty acid ratio of human dietary fat from pigmeat and improving human health.

Ages at first behavioural oestrus and at elevated plasma progesterone concentration were measured in three selection groups, after seven generations of divergent selection for lean growth rate (LGA), lean food conversion (LFC) and daily food intake (DFI) in a population of Large White pigs. First physiological oestrus was defined to have occurred when a blood plasma progesterone concentration of at least 1 μg/l was detected from weekly sampling of gilts. The study consisted of 146 gilts, which were given 0·75 , 0·81 , 0·88 , 0·94 or 1·0 g/g of daily ad-libitum food intake during performance test and then 1.9, 2.05, 2.2, 2.35 or 2.5 kg/day, respectively, until conception, to determine if there were differences between selection lines in their sensitivity to changes in nutritional inputs.Responses in oestrus and performance test traits were dependent on selection group. First physiological oestrus was later with selection for high LFC than for low LFC (234 v. 215, s.e.d. 9.1 days) but there was no significant response within each of the LG A (224 v. 226 days) and DFI (218 v. 206 days) selection groups. The probability of exhibiting oestrous behaviour signs at first physiological oestrus was significantly lower in the high LG A line (0·62 v. 0·93 or 0·5 v. 2.5, s.e.d. 0·75 on the logit scale) than in the low line but there were no responses in the LFC and DFI groups. For animals exhibiting oestrous behaviour signs at first physiological oestrus, there were no significant responses in oestrous behaviour score for the three selection groups. Live weight at first physiological oestrus in the LFC and LG A selection groups was greater in the high lines than in the low lines (120 v. 109 and 123 v. 112, s.e.d. 4.3 kg) but not in the DFI selection group (116 v. 111 kg). Responses in ultrasonic backfat (-7.3, -8.2 and 5.0, s.e.d. 1.5 mm) and muscle depth (4.9, 6.1 and -3.5, s.e.d. 1.4 mm) at first physiological oestrus were of similar magnitude in the LGA, LFC and DFI selection groups.Increasing the ration (amount of food offered) did not have a linear effect on performance test traits and reproductive development, such that ration had to be included in the model as a fixed effect, rather than a covariate. There was no significant effect of ration or of selection line with ration interaction for traits associated with first oestrus.Selection for lean growth rate had no adverse effect on reproductive development, unlike selection for lean food conversion. Detection of first oestrus with oestrous behaviour signs combined with physiological assessment may be required in genotypes selected exclusively for lean growth rate, rather than relying only on observed behavioural signs of oestrus.Ages at first behavioural oestrus and at elevated plasma progesterone concentration were measured in three selection groups, after seven generations of divergent selection for lean growth rate (LGA), lean food conversion (LFC) and daily food intake (DFI) in a population of Large White pigs. First physiological oestrus was defined to have occurred when a blood plasma progesterone concentration of at least 1 μg/l was detected from weekly sampling of gilts. The study consisted of 146 gilts, which were given 0·75 , 0·81 , 0·88 , 0·94 or 1·0 g/g of daily ad-libitum food intake during performance test and then 1.9, 2.05, 2.2, 2.35 or 2.5 kg/day, respectively, until conception, to determine if there were differences between selection lines in their sensitivity to changes in nutritional inputs.Responses in oestrus and performance test traits were dependent on selection group. First physiological oestrus was later with selection for high LFC than for low LFC (234 v. 215, s.e.d. 9.1 days) but there was no significant response within each of the LG A (224 v. 226 days) and DFI (218 v. 206 days) selection groups. The probability of exhibiting oestrous behaviour signs at first physiological oestrus was significantly lower in the high LG A line (0·62 v. 0·93 or 0·5 v. 2.5, s.e.d. 0·75 on the logit scale) than in the low line but there were no responses in the LFC and DFI groups. For animals exhibiting oestrous behaviour signs at first physiological oestrus, there were no significant responses in oestrous behaviour score for the three selection groups. Live weight at first physiological oestrus in the LFC and LG A selection groups was greater in the high lines than in the low lines (120 v. 109 and 123 v. 112, s.e.d. 4.3 kg) but not in the DFI selection group (116 v. 111 kg). Responses in ultrasonic backfat (-7.3, -8.2 and 5.0, s.e.d. 1.5 mm) and muscle depth (4.9, 6.1 and -3.5, s.e.d. 1.4 mm) at first physiological oestrus were of similar magnitude in the LGA, LFC and DFI selection groups. Increasing the ration (amount of food offered) did not have a linear effect on performance test traits and reproductive development, such that ration had to be included in the model as a fixed effect, rather than a covariate. There was no significant effect of ration or of selection line with ration interaction for traits associated with first oestrus. Selection for lean growth rate had no adverse effect on reproductive development, unlike selection for lean food conversion. Detection of first oestrus with oestrous behaviour signs combined with physiological assessment may be required in genotypes selected exclusively for lean growth rate, rather than relying only on observed behavioural signs of oestrus.

Genotype with nutrition interactions in carcass composition and meat quality traits were examined by testing pigs from four selection lines and a control line on isoenergetic diets, which differed in lysine: energy content. The selected lines resulted from seven generations of selection for high daily food intake, lean food conversion ratio (LFC) and lean growth rate onad-libitumor restricted (LGS) feeding regimes in a Large White population. There were 128 pigs in the study, with 24 pigs per selection line and 32 pigs from a control line. During performance test, 30 to 90 kg, pigs were offered one of three isoenergetic diets, 14·0 MJ digestible energy (DE) per kg dry matter, which differed in ileal digestible lysine: digestible energy (A: 0·40 , C: 0·76 and E: 1·12 g lysine per MJ DE) onad-libitumor restricted (0.75 g/gad-libitumdaily food intake) feeding regimes.For the majority of performance test, carcass composition and meat quality traits there was no evidence of a genotype with diet or genotype with feeding regime interaction. The selection line with feeding regime interactions for average daily gain, daily food intake and rates of lean and subcutaneous fat deposition were primarily due to no feeding regime effect for the LFC selection line. Selection for high LFC had reducedad-libitumdaily food intake to such an extent that it was not significantly different from daily food intake on a restricted feeding regime, unlike other selection lines in the study. A selection line with feeding regime interaction was detected for muscle рH24hand muscle reflectance, which resulted from the LGS selection line. LGS pigs offered food ad libitum had higher muscle рH24hand lower muscle reflectance than LGS pigs given food at a restricted level, while there was no effect of feeding regime for the other selection lines.The general absence of genotype with nutrition interactions for traits measured in the study indicated that the ranking of genotypes for performance test traits, carcass composition and meat quality traits will not be dependent on diet or feeding regime. Genotype specific nutritional inputs will also not be required for identification of pigs of high genetic merit, within a genotype. However, diet and feeding regime had significant effects on carcass composition and meat quality traits, such that the estimated mean value of a genotype will be dependent on the diet or feeding regime used to evaluate the genotype.

Responses in carcass composition and meat quality after seven generations of selection for components of lean growth rate were examined in a population of Large White pigs. There were four selection groups in the study, with divergent selection for lean growth rate on ad libitum (LGA) or restricted (LGS) feeding regimes, lean food conversion ratio (LFC) and daily food intake (DFI). In generations six and seven, two offspring from each of 10 sires, within each selection line, were to be allocated for half-carcass dissection and measurement of meat quality. There were 320 animals in the study, with 40 animals from each of the high and low selection lines and, within each selection line, two offspring per sire.Responses in carcass composition were similar in the three selection groups given food ad libitum, but rates of lean and fat growth rate differed between selection lines. Intramuscular fat content was reduced with selection for high LGA and high LFC but was increased with selection for low DFI (-1·7 and -3·2 v. 2·7 (s.e.d. 0·7) mg/g), which was unexpected given the higher carcass fat content of the low DFI line, relative to the high line (249 v. 190 (s.e.d. 7) g/kg). Muscle colour was darker, as measured by trained sensory panel assessment, in selection lines which reduced the rate of fat deposition to achieve a leaner carcass (high LFC and low DFI) but there was no response in muscle colour with selection for LGA (0·4 and 0·3 v. 0·0 (s.e.d. 0·1)). Responses in muscle shear force (5·3 v. 4·4 (s.e.d. 0·4) kg) and flavour liking (4·0 v. 4·3 (s.e.d. 0·12)) were limited to the LGA and LFC selection groups, respectively. There were no significant responses in muscle moisture content, muscle pH or myofibrillar fragmentation index, nor were there any responses in meat quality with selection on LGS. Therefore, decreasing the rate of fat deposition was associated with darker meat and increasing the rate of lean growth was associated with higher shear force. There were selection strategy specific responses in the fatty acid composition of intramuscular fat, which may have contributed to the responses in eating quality.In general, responses in meat quality were small, such that incorporation of meat quality traits in selection objectives, which are primarily focused on increasing the efficiency of lean meat production, may not be necessary. However, it would be pertinent to evaluate periodically genotypes of breeding companies for muscle quality traits.

The responses in traits measured at the start of the mating period and at farrowing were examined after seven generations of divergent selection for daily food intake (DFI), lean food conversion (LFC), lean growth rate (LGA) on ad-\ibitum feeding and lean growth on scale feeding (LGS). Weight and backfat depth at mating were measured on 330 gilts and 74 boars, which were selected on the basis of performance test traits. Farrowing information was available on 259 gilts. At the start of the mating period, gilts selected for high DFI, LGA or LGS had similar live weights (135, 137 and 137 (s.e.d. 4·5) kg) but different backfat depths (20·3, 14·0 and 11·3 (s.e.d. 1·3) mm) while the corresponding low lines had different live weights (129, 117 and 124 kg), but similar backfat depths (17·5, 17·8 and 17·8 mm). Gilts selected for high LFC had lower mean live weight and backfat depth (114 kg and 10·7 mm) than gilts in the other selection lines. Conception rates of gilts selected for low DFI or high LGS were similar (0·62 and 0·64, s.e.d. 0·12) and lower than for the alternative selection strategies (0-78), but the low DFI gilts were significantly older at farrowing than gilts selected for high LGS (424 v. 408 (s.e.d. 5·5) days). Responses in live weight, backfat depth, age at mating and particular reproduction traits were selection strategy dependent, such that identification of relationships between growth and reproduction traits will require measurement of additional growth traits at an earlier age than in the current study

Genetic and phenotypic relationships between performance test and reproduction traits were estimated, after five generations of divergent selection for components of efficient lean growth, in a population of Large Wltite pigs. On ad-libitum feeding, a total of 4334 pigs were performance tested, of which 884 selected gilts had measurements of reproduction traits. On a restricted feeding regime, 1558 pigs were tested, which included 336 selected gilts with reproduction records. For pigs given food ad libitum, genetic correlations between litter weights at birth and weaning with daily food intake (0·48 and 0·42, s.e. 0·16) and with growth rate on test (0·65 and 0·52) were positive, but correlations with backfat depths were not significantly different from zero. For pigs given food at a restricted level, litter birth weight was positively genetically correlated with growth rate (0·50, s.e. 0·18) and negatively correlated with backfat depths (-0·48, s.e. 0·16). Phenotypic and environmental correlations between performance test and reproduction traits were all less than 0·10 in magnitude, for pigs tested on either feeding regime. The variation in backfat depth enabled detection of a non-linear relationship between predicted breeding values for litter weight at birth with predicted breeding values for average backfat depth of farrowing gilts performance tested on ad-libitum feeding, but not for gilts tested on restricted feeding. The positive genetic correlations between growth rate and daily food intake with litter traits suggested that selection strategies which change growth and daily food intake may result in relatively greater genetic changes in piglet growth rate than in litter size.

Carcass composition was measured after six generations of divergent selection for lean growth rate on ad-libitum and restricted feeding, lean food conversion and daily food intake in populations of Large White (LW) and Landrace (LR) pigs. There were 161 half-carcass dissections in LW pigs and for LR pigs, a double sampling procedure combined information from 53 half-carcass and 53 hand joint dissections. The performance test started at 30 kg and finished at 85 kg with ad-libitum feeding and after 84 days with restricted feeding, and pigs were slaughtered at the end of the test.In the LR population, selection for lean growth on restricted feeding increased carcass lean content (605 v. 557 (s.e.d. 19) g/kg), but there were no significant responses in carcass lean content with the selection strategies on adlibitum feeding. Selection for lean food conversion and high lean growth on restricted feeding reduced carcass fat content (201 v. 241 (s.e.d. 14) and 150 v. 218 (s.e.d. 18) g/kg), but selection for high lean growth rate with adlibitum increased carcass fat content (212 v. 185 (s.e.d. 11) g/kg). Responses in carcass composition were not significant with selection on daily food intake.In the LW population, selection for high lean food conversion or low daily food intake increased carcass lean content (539 v. 494 and 543 v. 477 (s.e.d. 11) g/kg) to a greater extent than selection on lean growth rate (509 v. 475 g/kg). Responses in carcass fat content were equal and opposite to those in carcass lean content. Selection on lean growth rate with ad-libitum feeding increased lean tissue growth rate (LTGR) (491 v. 422 (s.e.d. 23) g/day), but there was no change in fat tissue growth rate (FTGR) (206 v. 217 (s.e.d. 15) g/day). In contrast, FTGR was reduced with selection on lean food conversion (169 v. 225 g/day), but LTGR was not significantly increased (520 v. 482 g/day). Selection for lean growth rate with restricted feeding combined the desirable strategies of lean growth rate on adlibitum feeding and lean food conversion, as LTGR was increased (416 v. 359 (s.e.d. 12) g/day) and FTGR decreased (126 v. 156 (s.e.d. 7) g/day). The preferred selection strategy may be lean growth rate on restricted feeding, which simultaneously emphasizes rate and efficiency of lean growth.For ad-libitum fed LW pigs, coheritabilities for growth rate, daily food intake and backfat depth with carcass lean content were negative (-0·12, -0·22 and -0·50 (s.e. 0·05), but positive with carcass subcutaneous fat content (0·22, 0·24 and 0·50), when estimated from six generations of performance test data and carcass dissection data in generations 2, 4 and 6.

The genotype with feeding regime interaction was examined by testing pigs from four selection groups on both ad libitum and restricted feeding regimes. Within each selection group, there were high, low and control lines, which had been selected for lean food conversion ratio (LFC), daily food intake (DFI) or lean growth rate on ad libitum (LGA) or on restricted (LGS) feeding, in Large White-Edinburgh (LW) and Landrace-Wye (LR) populations. There were 1187 LW pigs and 768 LR pigs in the study, with 344 LW and 133 LR pigs tested on the alternative feeding regime.In the LW population, pigs in the high LGS line grew significantly faster than the high LGA and LFC lines with ad libitum feeding (919 v. 847 and 786 (s.e.d. 31) g/day), but had similar food conversion ratios and backfat depths. The high LGS and high DFI lines were similar for growth rate, daily food intake and food conversion ratio, but backfat depth was significantly lower in the high LGS line than in the high DFI line (12·0 v. 25·9 (s.e.d. 0·7) mm). On restricting feeding, the rankings of the selection lines for growth rate, food conversion ratio and mid-back fat depth were broadly similar to those with ad libitum feeding, except for the high LFC line. In the LR population, the high LGS, LGA and LFC lines did not differ significantly in growth rate, daily food intake, food conversion ratio and backfat depth within either the ad libitum or restricted feeding regimes. Growth rate of the high LGS line was similar to the high DFI line on ad libitum feeding (828 v. 836 (s.e.d. 40) g/day), but significantly higher on restricted feeding (704 v. 636 (s.e.d. 23) g/day). On both feeding regimes, food conversion ratio and backfat depth were significantly lower in the high LGS line compared with the high DFI line.In the LW population, the genetic correlation between feeding regimes for growth rate was significantly less than one (0·4 (s.e. 0·20)), but was not significantly different from unity (0·8 (s.e. 0·14)) for backfat depths. Based on the performance test results, selection for lean growth with testing on a restricted feeding regime may be preferable to testing animals on an ad libitum feeding regime.

Adipocyte dimensions of subcutaneous fat, sampled by biopsy at the 13th rib, were measured in 20-week-old rams from lines of Texel-Oxford and Scottish Blackface sheep, both divergently selected for carcass lean content. A total of 163 animals were measured, with close to equal numbers per breed-line combination. In both breeds, the high (lean) selection line had significantly lower backfat depths (0·71 mm in the Texel-Oxford and 0·83 mm in the Scottish Blackface, s.e.d. = 0·14 and 0·13 mm, respectively), but body weight did not differ between the lines. The ultrasonic fat depth differences between the Texel-Oxford selected lines were accompanied by increases in adipocyte diameter, area, diameter standard deviation within each sample and implied cell number, calculated as the ratio of ultrasonic fat depth to average adipocyte diameter. In the Scottish Blackface sheep there were no selection line differences in adipocyte dimensions, but there was an increase in implied cell number in the line selected for increased fatness. Across breeds, ultrasonic fat depth was correlated with both adipocyte diameter and implied adipocyte number (r = 0·58 and 0·75, respectively), but these latter two measurements were uncorrelated with each other.

Responses to four generations of divergent selection in pigs for lean growth rate (LGS) with restricted feeding were studied. The selection criterion was designed to obtain equal correlated responses in growth rate and carcass lean content, measured in phenotypic s.d. Animals were to be performance tested in individual pens with a mean starting weight of 30 kg for a period of 84 days. Daily food intake was equal to 0·75 gig of the daily food intake for pigs offered food ad-libitum. In the high, low and control lines, there ivere 1250 Large White-Edinburgh (LW) pigs and 875 British Landrace-Wye (LR) pigs. Each selection line consisted of 10 sires and 20 dams, with a generation interval ofl year.After four generations of selection, cumulative selection differentials were 5·9 and 4·8 phenotypic s.d. for LW and LR populations, with similar responses, 1·8 (s.e. 0·17) phenotypic s.d. Mean weight at the end of test, growth rate and backfat depths at the shoulder, mid back and loin were 89 kg, 712 g/day, 26,13 and 13 mm for LW and for LR pigs were 87 kg, 683 g/day, 28,10 and 10 mm. High line pigs were heavier at the end of test (4·3 (s.e.d. 1·4) kg and 4·0 (s.e.d. 1·6) kg) for LW and LR populations, with corresponding responses in growth rate (54 (s.e.d. 16) g/day and 47 (s.e.d. 18) g/day). Responses in the three backfat depths were −4·1 (s.e.d. 1·2) mm, −2·6 (s.e.d. 0·7) mm and −2·9 (s.e.d. 0·7) mm for LW and −2·2 (s.e.d. 0·05) mm, −2·2 (s.e.d. 0·4) mm and −2·4 (s.e.d. 0·5) mm for LR populations. Responses in weight off test and backfat depths were symmetric about the control lines.Heritabilities for LGS were 0·34 and 0·28 (s.e.d. 0·5) for the LW and LR populations, when estimated by residual maximum likelihood. Common environmental effects for LGS were 0·11 (s.e. 0·03) for LW and 0·17 (s.e. 0·04) for LR. Heritabilities for growth rate and average backfat depth were similar for LW and LR populations (0·17 and 0·29, s.e. 0·05), as were common environmental effects (0·10 s.e. 0·04). Average phenotypic and genetic correlations between growth rate and backfat, for LW and LR populations, were small (0·15 (s.e. 0·03) and −0·06 (s.e. 0·16), respectively).Responses to selection and genetic parameter estimates demonstrate that there is substantial genetic variation in growth and fat deposition when pigs are performance tested on restricted feeding.

Responses to four generations of divergent selection for lean groivth rate with ad-libitum feeding (LGA), for lean food conversion (LFC) and for daily food intake (DFI) in Large White pigs were studied. The LGA (LFC) selection criterion was designed to obtain equal correlated responses in growth rate (food conversion ratio) and carcass lean content, measured in phenotypic s.d. The selection criteria had phenotypic s.d. of 27, 29 and 253 units, respectively, and results are presented in s.d. units. There was a total of 3537 pigs, with an average of 40 boars and 40 gilts performance tested in each of the high, low and control lines per generation and the lines consisted of 10 sires and 20 dams. The generation interval was equal to 13·5 months. Animals were performance tested in individual pens with mean starting and finishing weights of 30 kg and 85 kg respectively.Cumulative selection differentials in the three selection groups were 5·8, 3·6 and 3·3 phenotypic s.d. for LGA, LFC and DFI respectively. Direct responses to divergent selection were 1·7, 1·3 and 1·2 (s.e. 0·17) for LGA, LFC and DFI. The correlated response in LFC (1·6 (s.e. 0·18)) with selection on LGA was greater than the direct response in LFC. Conversely, the direct response in LGA was greater than the correlated response (1·1 (s.e. 0·18)) with selection on LFC. The response in LFC (–1·1 (s.e. 0·17)) with selection on DFI was similar in size but opposite in sign to the direct response in LFC. Responses were asymmetric about the control, as the high LGA and LFC responses were proportionately smaller (0·74 and 0·58) than low line responses. In contrast, the difference between the high DFI and control was four times greater than the difference between low line and control.Heritabilities of LGA, LFC and DFI were 0·38, 0·35 and 0·29 (s.e. 0·03), when estimated by residual maximum likelihood, with common environmental effects of 0·09 (s.e. 0·02). Genetic correlations for LGA with LFC and DFI were positive, 0·76 (s.e. 0·03) and 0·23 (s.e. 0·07), but the genetic correlation between DFI and LFC was negative, –0·45 (s.e. 0·06). The experiment demonstrated that substantial responses to selection can be achieved in LGA, LFC and DFI. Selection on LGA resulted in larger direct and correlated responses than selection on LFC.

Genetic and phenotypic parameters and correlated responses in performance test traits were estimated for populations of Large White (LW) and British Landrace (LR) pigs tested in Edinburgh and Wye respectively, to four generations of divergent selection for lean growth rate (LGA), lean food conversion (LFC) and daily food intake (DFI) with ad-libitum feeding.There were differences between the two populations in genetic parameters, as LW heritabilities for growth rate, daily food intake and backfat depths were higher and the correlation between growth rate and backfat was positive for LW, but negative for LR. However, heritabilities, genetic and phenotypic correlations were generally comparable between selection groups, within each population. Genetic and phenotypic correlations indicated that animals with high daily food intakes were faster growing, had positive residual food intakes (RFI), were fatter with higher food conversion ratios. RFI was highly correlated with daily food intake and food conversion ratio, but phenotypically independent of growth rate and backfat, as expected.Selection for LGA, in LW and LR populations, increased growth rate (54 and 101 g/day), but reduced backfat (−3·9 and −2·0 mm), food conversion ratio (−0·23 and −0·25) and total food intake (−11·8 and −12·6 kg). There was no change in daily food intake in LW pigs (−19 g/day), but daily food intake increased in the LR pigs (69 g/day). With selection for LFC in LW and LR populations, there was no response in groivth rate (9 and 9 g/day), but backfat (−4·1 and −2·1 mm), total (−6·6 and −11·8 kg) and daily food intake (−90 and −172 g) were reduced, as animals had lower food conversion ratios (−0·13 and −0·22). LW and LR pigs selected for DFI ate more food in total (6·8 and 5·9 kg) and on a daily basis (314 and 230 g), grew faster (94 and 51 g/day) and had higher food conversion ratios (0·12 and 0·13). Backfat was increased in LW pigs (3·7 mm), but not in the LR population.In general, efficiency of lean growth was improved by increasing groivth rate, with little change in daily food intake from selection for LGA, but was primarily due to reduced daily food intake with selection on LFC.

Responses to divergent selection for lean growth rate with ad-libitum feeding (LGA), for lean food conversion (LFC) and for daily food intake (DFI) in Landrace pigs were studied. Selection was practised for four generations with a generation interval ofl year. A total of 2642 pigs were performance tested in the high, low and control lines, with an average of 37 boars and 39 gilts performance tested per selection line in each generation. The average within-line inbreeding coefficient at generation four was equal to 0·04. There was one control line for the DFI and LFC selection groups and another control line for the LGA selection group. Animals were performance tested in individual pens with mean starting and finishing weights of 30 kg and 85 kg respectively with ad-libitum feeding. The selection criteria had phenotypic s.d. of 32, 29 and 274 units, for LGA, LFC and DFI, respectively, and results are presented in phenotypic s.d.Cumulative selection differentials (CSD) were 5·1, 4·5 and 5·5 phenotypic s.d. for LGA, LFC and DFI, respectively. Direct responses to selection were 1·4,1·1 and 0·9 (s.e. 0·20) for LGA, LFC and DFI. In each of the three selection groups, the CSD and direct responses to selection were symmetric about the control lines. The correlated response in LFC (1·1, s.e. 0·19) with selection on LGA was equal to the direct response in LFC. In contrast, the direct response in LGA was greater than the correlated response (0·7, s.e. 0·18) with selection on LFC. There was a negative correlated response in DFI (-0·6, s.e. 0·18) with selection on LFC, but the response with selection on LGA was not significant (0·2, s.e. 0·16).Heritabilities for LGA, LFC and DFI ivere 0·25, 0·25 and 0·18 (s.e. 0·03), when estimated by residual maximum likelihood, with common environmental effects of 0·12 (s.e. 0·02). Genetic correlations for LFC with LGA and DFI were respectively positive (0·87, s.e. 0·02) and negative (-0·36, s.e. 0·09), while the genetic correlation between DFI and LGA was not statistically different from zero, 0·13 (s.e. 0·10). Selection on components of efficient lean growth has identified LGA as an effective selection objective for improving both LGA and LFC, without a reduction in DFI.

Fatty acid synthetase and lipoprotein lipase activities, lipid content of adipose tissue and the fatty acid composition of subcutaneous fat, sampled by biopsy at the 13th rib, were measured in 20-week-old rams from lines of Texel-Oxford (TO) and Scottish Blackface (SB) sheep, both divergently selected for carcass lean content. A total of 150 animals were measured, with close to equal numbers of animals per selection line-breed combination.In both breeds, the high (lean) selection lines had significantly lower backfat depths (TO : 0·5 mm and SB : 0·6 mm, s.e.d. 0·2) than the low (fat) lines. The lipid content of subcutaneous fat was 65 mg lipid per g fat tissue wet weight (s.e.d. 24) greater in TO rams than in SB rams. The TO low line had a higher lipid content than the high selection line (426 v. 448 (s.e.d. 36)) and although the SB selection lines did not differ, the selection line with breed interaction was not significant. SB rams had higher fatty acid synthetase activity (3·1 v. 2·6 (s.e.d. 0·3) on a log scale) but there were no differences between selection lines. Lipoprotein lipase activities were similar between breeds and selection lines. The lower concentration of myristic acid (C14:0) of TO rams compared with SB rams (0·9 (s.e.d. 0·3)) was the only breed or selection line difference which was statistically significant for fatty acid composition of subcutaneous fat.Lipid content of subcutaneous fat and lipoprotein lipase activity were highly correlated and both were positively correlated with performance test traits, especially with backfat depth. The correlation between backfat depth and fatty acid synthetase activity was not different from zero. Performance test traits, lipid content of subcutaneous fat and lipoprotein lipase activity were positively correlated with the unsaturated fatty acids, with the exception of C18 :1 when correlations were negative.

A divergent selection experiment for carcass leanness in Texel-Oxford sheep was established to examine the differences between genetically lean and fat animals derived from the same base population. The selection criterion was designed to change body composition without a corresponding change in live weight, using an index of ultrasonic backfat depth and live weight at 20 weeks of age. The index was constructed using estimates of genetic and phenotypic parameters, which were available at the start of the experiment in 1985. The difference between the high and low lines, after 3 years of selection, for the selection index, live weight at 20 weeks of age, ultrasonic backfat and muscle depths was 0·59,1·30 kg, -0·76 mm and 0·88 mm respectively. The estimated difference for carcass lean and fat weight was 0·72 kg and -0·04 kg respectively with 13·5 g/kg and -13·8 g/kg for carcass lean and fat proportion. The realized heritability for the selection index, estimated from the regression of cumulative response on cumulative selection differential was 0·42, with a standard error of 0·25, after taking account of genetic drift. The selection index heritability, estimated with residual maximum likelihood (REML) methodology, was 0·46 (s.e. 0·14). REML heritability estimates for live weight at 20 weeks of age, ultrasonic backfat and muscle depths were 0·20 (s.e. 0·13), 0·35 (s.e. 0·14) and 0·43 (s.e. 0·14) respectively.

Correlated responses in slaughter and carcass traits to divergent selection for high or low carcass lean content in Texel-Oxford sheep were measured in the crossbred progeny of 23 rams. Rams were selected using the selection index: -0·995 FATD + 0·206 WT20, where FATD and WT20 are the ultrasonic backfat depth and live weight at 20 weeks of age, with both traits standardized to have zero mean and unit phenotypic variance. A total of 329 crossbred lambs were slaughtered at 16 weeks of age and 250 lambs were slaughtered at fixed weight, 38 to 40 kg for castrated males and 36 to 38 kg for ewe lambs. Progeny from high-line rams grew faster, as indicated by the heavier slaughter weight of lambs slaughtered at fixed age (04 (s.e.d. 0·5) kg) and the lower age at slaughter for lambs slaughtered at fixed weight (-5 (s.e.d. 3) days). Subcutaneous and internal fat scores and the Meat and Livestock Commission carcass appraisal and conformation scores were all lower in the high line than in the low line and the magnitude of the selection line differences for lambs slaughtered at fixed weight was essentially double that of lambs slaughtered at fixed age (0·8 v. 0·5 score units).Shoulder joints of 254 lambs, slaughtered at fixed age, were dissected and half carcasses were dissected on 66 of the lambs. Carcass and shoulder joint information was combined with double-sampling methodology, using multiple regression equations to predict carcass composition. For each tissue type, viz. lean, subcutaneous fat, internal fat and bone, the correlations between carcass content and shoulder joint content were at least 0·90 and the proportions of variation in carcass tissue content accounted for by the prediction equations were also at least 0·90. Progeny of high-line rams had higher carcass lean weight, (402 (s.e.d. 140) g) than progeny from the low line, with no difference in carcass fat weight, such that carcass lean content was higher, (10 (s.e.d. 5) g/kg), and carcass fat content was lower, (-10 (s.e.d. 7) g/kg). Based on the slaughter and carcass traits of these crossbred progeny, divergent selection for high and low carcass lean content has established lines of animals with different rates of lean growth but similar rates offat deposition.

Duroc and halothane negative British Landrace boars and gilts were performance tested on ad libitum or restricted feeding regimes, with like-sexed non-littermate groups of one, two, three or four pigs per pen. There was a total of 320 pigs with 20 litter groups per breed with four boars and four gilts per litter group. Within each feeding regime, a boar and a gilt from each litter were tested on one of two diets in 1986, for each of the four combinations, and pigs were tested similarly for two other diets in 1987. Carcass composition was determined by half-carcass dissection of 160 pigs allocated between treatments.There was a breed × sex interaction for growth and performance traits for pigs fed ad libitum. Duroc boars were faster growing and more efficient than Landrace boars, but Duroc gilts grew more slowly and were less efficient than Landrace gilts. On restricted feeding, Duroc pigs were more efficient than Landrace pigs.At constant slaughter weight, Duroc pigs had less subcutaneous fat but more intermuscular fat. Although they had less separable fat in the carcass, Duroc pigs were not leaner as weights of bone, skin, head, feet and tail were heavier than for Landrace pigs.Group penning and group feeding of pigs may have enhanced competition effects resulting in positive genetic and phenotypic correlations between growth rate and backfat depths on both feeding regimes.The positive genetic correlation between growth rate and fat deposition resulted in a negative genetic correlation between growth rate and carcass lean content and a lower genetic correlation with lean tissue growth rate than in other studies.

The melanocortin-4 receptor (MC4R) has a vital role in the control of energy balance and the genetic basis of obesity. A polymorphism, which results in the replacement of aspartic acid with asparagine at position 298 of the porcine MC4R gene, within the seventh transmembrane domain, has previously been described. In the current study, allele frequencies for this Asp298Asn polymorphism were investigated in lines of Large White pigs which had been divergently selected for seven generations based on lean food conversion (LFC), lean growth with ad libitum feeding (LGA), lean growth with restricted feeding (LGS) and daily feed intake (DFI). The association of the Asp298Asn polymorphism with performance traits in these lines was assessed. The frequency of Asp298 was higher (P < 0.001) in the LFC high line (0.48) than the low line (0.00), while the frequency of Asn298 was higher (P < 0.01) in the LGA high line (0.22) than the low line (0.04). When analysed across all lines, the Asp298Asn polymorphism was significantly associated with ultrasonic backfat depth, average daily gain and daily feed intake (P < 0.05). Asp298 homozygous animals had mean values of 13.3 mm, 733 g and 1933 g for backfat, average daily gain and daily feed intake respectively, compared with 14.7 mm, 805 g and 2098 g for Asn298 homozygotes. Therefore, the data support a role for the MC4R Asp298Asn polymorphism in the genetic basis of economically important traits in Large White pigs.

Serum urea concentrations were measured in Large White pigs from lines divergently selected for components of efficient lean growth rate and performance tested over three 14-d test periods starting at 30, 50, and 75 kg. Two methods of performance testing were used. Phase-fed pigs were fed to appetite isoenergetic diets differing in total lysine:energy ratio (0.58, 0.69, 0.81, 0.91, 1.01, 1.12, and 1.23 g/MJ of digestible energy), whereas diet-choice pigs were offered a choice of the 0.69 and 1.12 lysine:energy diets. Between test periods, all animals were fed one diet: 0.91 g of lysine/MJ of digestible energy. The study consisted of 230 boars and gilts with 150 pigs performance tested on phase-feeding and 80 pigs on diet-choice. The line selected for high lean food conversion had lower urea concentrations on each diet than the line selected for high lean growth rate, despite similar predicted lysine balances. Efficiency of lean growth rather than the rate of lean growth may be a better selection strategy in the context of nitrogen excretion. Urea concentrations at the end of each test period were correlated with lysine intake (0.33, 0.48 and 0.65; standard error, 0.08) and predicted lysine balance (0.39,0.44, and 0.64), but were uncorrelated with predicted lysine for protein deposition (0.01, 0.08, and 0.08) and maintenance. Urea concentration at the end of a test period was not a useful predictor of protein deposition, even after accounting for pretest variation in urea concentration and food intake during test. The expected response pattern of serum urea concentration to diets differing in total lysine:energy would be nonlinear, with the point of inflection occurring at the required dietary total lysine:energy for each genotype. However, there was no evidence of such an inflection point such that the prediction of lysine requirement from urea concentration was not possible for the selection lines in the study.

Responses in log transformed serum neuropeptide Y (NPY) concentration and leptin concentration after six generations of divergent selection on components of efficient lean growth in pigs were measured. From an animal breeding perspective, serum NPY and/or leptin concentrations could be used as physiological predictors of genetic merit if there were significant responses to selection. At 90 kg liveweight, log transformed serum NPY concentrations were increased with divergent selection for low food conversion ratio (LFC) (6.31 versus 5.72, SED 0.09 log(pmol/L)) or for high lean growth rate (LGA) (5.80 versus 5.37 log(pmol/L)) but not with selection on daily food intake (DFI) (6.26 versus 6.14 log(pmol/L)). Selection for high DFI was associated with increased serum leptin concentration (3.06 versus 2.45, SED 0.21 ng/mL human equivalent (HE)) as was selection for low LFC (3.04 versus 2.46 ng/mL HE). Correlations between leptin and predicted lipid weight increased with stage of test (0.13, 0.34 and 0.43, SE 0.08 at 30, 50 and 75 kg). The high correlations between successive serum NPY concentrations (0.80, SE 0.11) suggest that changes in body composition with time would not be reflected in serum NPY concentrations. Serum NPY and, to a lesser extent, serum leptin concentrations were insensitive to dietary differences in total lysine: energy and indicated that studies using a genetic resource population of animals may be more powerful than nutritional studies using isoenergetic diets differing in lysine content to examine aspects of function of serum concentrations of NPY and leptin in pigs.

Selection for high or low daily food intake (DFI) in Large White pigs resulted in higher serum leptin concentration, fat deposition and food intake in the high DFI line. The response in serum leptin concentration indicated that the higher fat deposition of the high DFI line was not due to insufficient leptin production, as in the Lepob/Lepob mouse. Serum leptin was more highly correlated with fat deposition than with food intake indicating that the response in serum leptin was primarily due to increased fat deposition rather than to higher energy intake per se. The low correlations between serum leptin measured at 30 kg and performance test traits indicate that serum leptin would not be efficient for selection of animals prior to performance test. However, the consistent positive correlations between serum leptin and a measure of fat deposition suggest that serum leptin could usefully be incorporated in selection criteria for genetic improvement of carcass lean content in pigs.

Genetic and nutritional effects on the boar taint traits of androstenone, skatole and indole and the eating quality traits associated with boar taint were examined by testing animals from four selection lines and a control line on isoenergetic diets, which differed in ileal digestible lysine: digestible energy (0.40, 0.76 and 1.12 g lysine/MJ DE). The selected lines resulted from seven generations of selection for high daily food intake, lean food conversion ratio (LFC) and lean growth rate on ad libitum (LGA) or restricted (LGS) feeding regimes in a Large White population. During performance test, from 30 to 90 kg, boars were fed on either ad libitum or restricted (0.75 g/g ad libitum daily food intake) feeding regimes. A sensory panel assessed heated fat samples for androstenone odour, skatole odour and abnormal odour. There were no significant differences between the selection and control lines or diets for log transformed fat content of androstenone, skatole, indole. The significant diet with feeding regime interaction for log transformed fat content of skatole and indole were essentially due to significantly higher log transformed fat contents with ad libitum feeding of the high lysine diet compared to restricted feeding (skatole: −1.94 vs −3.06, s.e.d. 0.43; indole: −3.44 vs −4.22, s.e.d. 0.28), as differences between feeding regimes on diets A and C were not significantly different from zero. There were no significant differences between selection and control lines for sensory panel score for abnormal odour or androstenone odour, but the LFC and LGA selection lines had a significantly higher skatole odour score than the LGS selection line. Neither diet nor feeding regime had any significant effect on sensory panel assessment of odour. Log transformed fat content of androstenone and skatole were significantly correlated with sensory panel score for skatole odour (0.37 and 0.46, s.e. 0.12), but not with sensory panel score for androstenone odour (0.06 and 0.09), such that they would not be useful predictors of androstenone odour.

Testing several genotypes on one diet may impose nutritional constraints on growth of one or more of those genotypes. Use of a diet choice procedure may enable pigs to ‘choose’ the appropriate combination of the diets to attain the growth determined by their genotype. In an animal breeding context, the diet choice procedure offers the potential to increase the accuracy of predicting an animal’s genetic merit for performance test traits. In the few diet choice studies made with pigs heavier than 35 kg (e.g. Bradford and Gous, 1991), the animals have been of one genotype. Therefore, animals from the Edinburgh lean growth selection experiment (Cameron, 1994) provided the experimental resource to determine if between-selection line differences were enhanced by testing pigs on a diet choice procedure compared with testing on a single diet.

Productivity in pigs can be improved by continued selection, however the impact of such selection on immune responses and resistance towards infectious challenges is not known. A risk is that this method may lead to a correlated reduction in the immune response and disease resistance. To estimate the effect of selection for performance traits upon immune responses, we compared levels of immune traits between divergent lines of Large White pigs selected for either lean growth under restricted feeding or feed intake.

An in vivo procedure to determine nutrient requirements for growing pigs would have substantial cost advantages over serial slaughter and furthermore repeated measurements would allow estimation of protein requirements at each stage of growth. Serum urea concentration (Urea) has been proposed as a candidate indicator of protein requirement as over-supply of protein will increase Urea (Chen et al., 1995). The study estimated genetic variation in Urea at different stages of performance test and the relationships between Urea and performance traits.

Insulin-like growth factor-1 has been positively associated components of growth in pigs, such as protein deposition. Therefore, serum insulin-like growth factor-1 concentration (IGF-1) was measured in lines of pigs divergently selected for daily food intake (DFI) or for efficient lean growth rate (LGS) to determine if there was a non-zero genetic association between IGF-1 and the selection criteria. If a genetic association did exist, then IGF-1 could be used as a physiological predictor of genetic merit in a breeding programme. The presence of a major gene effect on IGF-1 was also examined in the study.

A delay in the onset of puberty in gilts with intensive selection for efficient lean growth would have substantial implications for pig breeding programmes due to later first parity matings. Therefore the current study determined the responses in gilt traits at puberty to selection for components of efficient lean growth rate.

Responses in gilt live weight, backfat depth and food intake during lactation and in pre-weaning piglet growth rate were examined after seven generations of divergent selection for daily food intake (DFI), lean food conversion (LFC) or lean growth rate (LGA) on ad-libitum feeding or lean growth on restricted feeding (LGS). There were 252 Large White gilts in the study. Selection for low DFI resulted in gilts with less backfat (25·7 v. 30·7 (s.e.d. 2·21) mm) at farrowing and a substantially lowerfood intake (129 v. 146 (s.e.d. 5) kg) during lactation, but similar reductions in live weight (42 (s.e.d. 6) kg) and backfat depth (8·4 (s.e.d. 1·7) mm) than with selection for high DFI. Therefore, the lower piglet growth (167 v. 295 (s.e.d. 11) g/day) with selection for low DFI compared with selection for high DFI was primarily due to lower food intake of the gilts, as energy for milk production from food was reduced. In contrast, selection for high LFC resulted in relatively smaller changes in live weight (37 v. 48 (s.e.d. 5) kg) and backfat depth (7·6 v. 8·9 (s.e.d. 1·3) mm) than selection for low LFC, which combined with a lower food intake (132 v. 148 (s.e.d. 4) kg) during lactation, resulted in lower piglet growth (181 v. 200 (s.e.d. 11) g/day). The higher food intake of high LGA gilts (137 v. 121 (s.e.d. 4) kg) compensated for the relatively lower reductions in live weight (41 v. 46 (s.e.d. 5) kg) and backfat depth (5·5 v. 6·7 (s.e.d. 1·3) mm) during lactation compared with the low LGA line, such that piglet growth was similar (195 v. 289 (s.e.d. 11) g/day) in the two selection lines. In the high and low LGS selection lines, piglet growth was similar (195 v. 186 (s.e.d. 11) g/day) as was gilt food intake (125 v. 227 (s.e.d. 5) kg) and the changes in live weight (39 v. 41 (s.e.d. 6) kg) and backfat depth (8·1 v. 7·7 (s.e.d. 2·2) mm) during lactation. An examination of the daily energy used in litter gain and the energy available from gilt food intake and mobilization of body lipid indicated that one equation to predict the amount of body lipid mobilized during lactation was not appropriate for different genotypes. Responses in gilt food intake and the changes in live weight and backfat during lactation were selection strategy dependent. However, in general, the selection strategies which reduced gilt voluntary food intake during lactation or resulted in lower live weight and backfat depth at farrowing were detrimental to piglet growth rate.

Genetic and phenotypic relationships between performance test and reproduction traits were estimated in a population of Large White pigs, after five generations of divergent selection for components of efficient lean growth rate. The parameters are required to evaluate alternative selection strategies in pig breeding programmes.

This study determined the correlated responses in carcass composition after four generations of divergent selection for components of efficient lean growth rate in Large White (LW) and Landrace (LR) pigs.

Divergent selection lines for lean growth on a restricted feeding regime, in Large White and Landrace pigs, were established to complement the lean growth selection lines on ad-libitum feeding. This study estimated the direct and correlated responses after four generations of selection and the corresponding genetic and phenotypic parameters.The selection objective for lean growth on restricted or scale (LGS) feeding was to obtain equal correlated responses in growth rate and carcass lean content, measured in phenotypic s.d. The selection criterion included measurements of growth rate and ultrasonic backfat depth.Large White (LW) and Landrace (LR) boars and gilts were purchased from eight British nucleus herds and boars from national artificial insemination centres in 1982. Homozygous or heterozygous halothane positive pigs were not included in the experiment. The base populations consisted of 31 LW and 19 LR sires and 57 LW and 67 LR dams. Within each population, there were high and low selection lines with a control line, each consisting of 10 boars and 20 gilts, with a generation interval of 13.5 and 12 months for LW and LR pigs. Animals were performance tested in individual pens from 30±3 kg for a period of 84 days and fed a high energy (13.8 MJ DE/kg DM) and high protein (210 g/kg DM crude protein) pelleted ration. Daily food intake was equal to 0.75 g/g of the daily food intake for ad-libitum fed pigs and the total food intake was 134 kg for LW pigs and 150 kg for LR pigs. On average, 3 boars and 3 gilts were tested per litter. The total number of pigs tested per line and average inbreeding coefficients at generation four, by population are given below.

Correlated responses in reproductive performance to five generations of divergent selection for daily food intake (DFI), lean food conversion (LFC), lean growth rate on ad–libitum feeding (LGA), and lean growth rate on scale feeding (LGS) were studied. Litter traits were measured on 1220 Large White gilts. Mean litter weights at birth and weaning were 12·9 kg and 63·5 kg, with average litter sizes of 10·3 and 7·9. Responses to selection in the high and low lines for litter size in the DFI and LFC selection groups were 1·9 and –1·5 (s.e.d. VI) at birth and 0·9 and –1·8 (s.e.d. 1·2) at weaning. Responses in litter birth weights were respectively positive and negative for DFI and LFC (3·0 and –2·8 (s.e.d. 1·4) kg) and the response in LGS (3 kg) was greater than in LGA (–0·1 kg). Selection line differences in litter weaning weight followed a similar pattern to birth weight for DFI and LFC (17·5 and –17·3 (s.e.d. 10·1) kg). Responses in litter weights were a result of selection line differences in both litter sizes and piglet weights. The relationships between litter size, litter weights and piglet weights at birth and weaning were essentially linear. An extra piglet at birth and weaning corresponded to an increase of 1·0 (s.e. 0·02) kg and 6·9 (s.e. 0·1) kg in litter weights. Piglet birth and weaning weights were decreased by 0·03 (s.e. 0·003) kg and 0·19 (s.e. 0·02) kg. A uterine constraint on piglet growth was implied, but there was no evidence for a limit to uterine capacity. Heritabilities for litter size, weight and piglet weight at birth of 0·06, 0·11 (s.e. 0·04) and 0·16 (s.e. 0·02) respectively were similar to those at weaning. Common environmental effects on piglet weights at birth and weaning were substantially higher than the heritabilities (0·38 and 0·45, s.e. 0·01). The study indicated that selection for lean growth on either an ad–libitum or restricted feeding regime did not significantly affect reproductive performance, but the high lean food conversion ratio and low daily food intake selection lines had impaired reproductive performance.

Selection was carried out in a line of pigs for increased growth rate of lean tissue. The selection criterion was weight of lean in the ham predicted from live backfat and weight measurements after a 12-week performance test commencing at 25 kg live weight. All pigs were given the same total amount of food over the test period. The scale was set to about proportionately 0·85 of predicted ad libitum intake. Boars selected with an intensity of 1/12 were used for 6 months and sows selected with an intensity of 1/4 were kept for two farrowings. An unselected control line was maintained concurrently.After five generations, performances of selected and control line pigs were compared on ad libitum and scale feeding as they grew to 85 kg. Responses in the selected line on scale feeding were +51 g/day for growth rate (GR), −0·16 for food conversion ratio (FCR), −2·2 mm for backfat (F) and +0·47 kg for ham lean (HL). On ad libitum feeding, responses were much higher in the selected line, giving rise to line × food interactions. Responses were +128 g/day for GR, −0·27 for FCR, −2·3 mm for F, +1·01 kg for HL and +0·15 kg/day for food intake (FI). Estimates of the heritability of HL from variance components were 0·43 (s.e. 0·15) on scale feeding and 0·28 (s.e. 0·19) on ad libitum feeding. The realized heritability of HL on scale feeding was 0·29 (s.e. 0·04) and its co-heritabilities with the other traits on both feeding levels were of similar magnitude to its heritability. Scale feeding exposed genetic variation in the partitioning of food between lean and fat deposition and appeared to be a suitable selection regimen for performance on ad libitum feeding.

The accuracy of six methods to estimate carcass leanness in young rams was studied in 36 Texel-Oxford rams, measured at 5 months of age. The rams were slaughtered and dissected. Plasma non-esterified fatty acid and very low density lipoprotein concentrations, sampled before, during and after fasting, showed no relationship with carcass leanness. The same was true for food conversion efficiency measured from 6 to 16 weeks of age, and for measurements taken with the Scanogram ultrasonic machine. The Vetscan and Danscanner ultrasonic machines gave repeatable measurements of fat depth (0·41 and 0·46 respectively). The Vetscan was the best predictor of carcass leanness by proportionally reducing the residual mean square by about 0·20, corresponding to a correlation of −0·45 between ultrasonic fat depth and carcass leanness. Inclusion of average daily gain and ultrasonic fat depth in a selection index would allow appreciable improvements in both traits.

The performance of crossbred ewes sired by Border Leicester (BL), Bluefaced Leicester (BFL) and ABRO Damline (DL) crossing sires was compared. The dams of the crossbred ewes were of three hill breeds (Scottish Blackface, Swaledale and Welsh Mountain). Contemporary comparisons were made on 18 commercial and college farms throughout England and Wales from 1977 to 1981. A total of 1 277 crossbred ewes were involved with data on up to three years' lambings per ewe, giving a total of 3 522 mating records. The analyses were by least squares, fitting constants for terminal ram breed × farm × year of birth of ewe groups, sire breed, dam breed, age of crossbred ewe and all two-way interactions.The DL crossbred ewes were lighter at mating than the other crossbreds at each age. The proportion of 1-year-old crossbred ewes lambing was highest for the DL (0·78), compared to BFL (0·73) and BL (0·62), indicating earlier sexual maturity. Over all ages, the DL crossbreds were more prolific with 1·79 lambs per ewe lambing compared to BFL (1·63) and BL (1·53) and more lambs were weaned (1·33, 1·29 and 1·19 respectively). The total litter weight at both birth and weaning of the DL crossbreds was greater than the BL crossbreds but less than the BFL crossbreds at each age. For prolificacy and early sexual maturity, the DL cross is superior to both breeds. In overall production, the DL cross had no advantage over the BFL cross, as the litters of the DL crossbred ewes were lighter than the BFL litters at both birth and 10 weeks of age

Blood samples from 4,427 cows in over 300 commercial dairy herds were examined as part of a health monitoring scheme. Preliminary evidence is presented indicating that marginal concentrations of serum magnesium are related to below mean levels of blood urea nitrogen in such a manner as to suggest that marginal magnesium deficiency is a possible cause of reduced voluntary feed intake in cows managed in commercial dairy farming systems.

Performance records for the progeny of matings of Oxford, Texel, Texel-Oxford, Charollais, Charmoise and Meatlinc rams with crossbred ewes were analysed. The lambs, born from 1980 to 1982, were randomly allocated to three slaughter groups: (1) slaughter at fixed weights of 36 and 38 kg for ewe and castrated lambs; (2) slaughter according to estimated fat cover; (3) slaughter at a fixed age of 4·5 or 5·5 months with half-carcass dissection. The analyses were by least squares and effects were fitted for terminal sire breed, breed of maternal grand sire, year of birth, age of dam at lambing, birth-rearing type and sex of lamb and all two-way interactions. The results followed a well-established pattern where weight for age rankings were constant from birth, through weaning, to slaughter. Progeny of Oxford rams were the heaviest, followed by Texel-Oxford, Meatlinc and Charollais, Texel and Charmoise in that order. Lambs sired by rams of low mature weight reached a fixed weight at a later age than those sired by rams of high mature weight. They also reached a similar degree of fat cover at an earlier age and at lighter weights than the larger breeds. The carcass traits indicated that Texel crosses had a higher lean and lower fat proportion than would be expected from their growth and mature weight.The comparative performance of crossbred lambs from 15 terminal sire breeds was assessed, at the same proportion of carcass subcutaneous fat, by combining the results of three experiments.

The precision of estimates of genetic variances and covariances obtained from multivariate selection experiments of various designs are discussed. The efficiencies of experimental designs are compared using criteria based on a confidence region of the estimated genetic parameters, with estimation using both responses and selection differentials and offspring-parent regression. A good selection criterion is shown to be to select individuals as parents using an index of the sums of squares and crossproducts of the phenotypic measurements. Formulae are given for the optimum selection proportion when the relative numbers of individuals in the parent and progeny generations are fixed or variable. Although the optimum depends on "a priori" knowledge of the genetic parameters to be estimated, the designs are very robust to poor estimates. For bivariate uncorrelated data, the variance of the estimated genetic parameters can be reduced by approximately 0.4 relative to designs of a more conventional nature when half of the individuals are selected on one trait and half on the other trait. There are larger reductions in variances if the traits are correlated.

The interaction of sire with feeding regime (ad libitum and restricted) was assessed in the first stage of a selection experiment. There were 79 sires (59 Large White (Edinburgh) and 20 British Landrace (Wye)) with 1549 progeny (745 at Edinburgh and 804 at Wye) in the study. Progeny within each litter were allocated at random to each feeding regime. Pigs started test at 30 (± 3) kg and finished test at 85 (± 5) kg for ad libitum fed pigs or after 84 days on test for restricted fed pigs. All pigs were individually penned. At the end of test, food intake, weight off test and ultrasonic backfat depths were recorded. Variance components were estimated using restricted maximum likelihood and a weighted analysis took account of the different residual variances between feeding regimes, within each breed.There were differences in some genetic parameters due to feeding regime within breeds. No evidence of a sire with feeding regime interaction for average daily gain or food conversion ratio was detected in Large White pigs (genetic correlations between ad libitum and restricted feeding were 1·21 (s.e. 0·17) and 0·89 (s.e. 0·19) respectively). Similar conclusions could not be made for the sample of British Landrace pigs due to the large standard errors of the genetic correlation estimates. For ultrasonic backfat depths, the negative estimates of sire variances under restricted feeding prevent calculation of the genetic correlations and conclusions to be made about the sire with feeding regime interaction.