Partitioning of genetic effects on lifetime performance of mice

Summary A total of 2,457 lifetime performance records of 29 genetic groups of mice was analyzed using multiple regression of records on the proportion of gene contribution from 6 lines (designated as Lines M_P, m_Q, W_P, w_Q, C_P and c_Q). Genetic effects were partitioned into line additive, line maternal, direct heterosis, maternal heterosis and paternal heterosis effects. The line additive and line maternal effects were expressed as deviations from Line c_Q. Seventeen of 25 line additive effects differed significantly (P<0.05) from Line c_Q whereas only 4 of 25 line maternal effects deviated significantly from Line c_Q. Deviations in line additive effects from c_Q were negative in all lines examined whereas deviations in line maternal effects from c_Q were all positive, indicating a negative relationship between line additive and line maternal effects. Direct heterosis effects were all positive and significant (P < 0.01) except in the M_PxW_P cross which was produced by mating Lines M_P and W_P of the same base population (P). Maternal heterosis effects were significant in 10 of 20 cases whereas paternal heterosis effects were significant in 13 of 20 cases. Although direct heterosis is a major component of total heterosis effects (sum of direct, maternal and paternal heterosis), the results suggest that parental heterosis may need to be considered in producing multiple way crosses. The fitting of line additive, line maternal, direct heterosis, maternal heterosis and paternal heterosis effects in the multiple regression model effectively accounted for all genetic effects in lifetime performance.Animal Research Centre Contribution No. 1326     

Partitioning Average and Heterotic Components of Direct and Maternal Genetic Effects on Growth in Mice Using Crossfostering Techniques

Crossfostering was performed using lines selected for increased 6-week body weight (H₆) and increased 3-to 6-week postweaning gain (M16) and their reciprocal F₁ crosses as nurse dams in the selected crossfostering group, and base population controls (C₂, ICR) and their reciprocal F₁ crosses in the control group. The offspring suckled were H₆, M16 and F₂ crosses in the selected group, and C₂, ICR and their F₂ crosses in the control group. Measurements taken on the individual offspring were body weights at birth (WB) and at 12, 21, 31, 42, and 63 days (W12, W21, W31, W42 and W63, respectively) and weight gains between adjacent ages (GB-12, G12–21, G21–31, G31–42 and G42–63, respectively). Least squares constants fitted to populations of genetic and nurse dams were used to calculate specific linear contrasts. Correlated responses to selection in average direct genetic effects were significant and positive for all traits examined in both H₆ and M16, while the correlated responses in average maternal genetic effects were negative in M16 and negligible in H₆. Selection response was primarily due to average direct genetic effects while the contribution of average maternal genetic effects was of secondary importance. The response in average direct genetic effects was smaller in M16 than in H₆ through weaning (WB, W12 and W21), but was larger in M16 for postweaning weights (W31, W42 and W63). The correlated responses in average maternal genetic effects were consistently smaller in M16 than in H₆. Direct heterosis was significant for all traits except for G12–21 and G42–63 in the control group, whereas maternal heterosis was significant for weight gains at early ages and for body weights. Direct heterosis tended to be larger than maternal heterosis in both selected and control crosses. Percent direct heterosis for body weight was larger in the selected crosses relative to the control crosses through 31 days of age, but the trend was reversed by 63 days. Percent maternal heterosis was consistently larger in the selected crosses.     

Influence of early pregnancy on reproductive rate in lines of mice selected for litter size

Summary The influence of male-induced early puberty on female reproductive rate was determined in three lines of mice differing in litter size and body weight. The lines originated from a single base population and had undergone 20 generations of selection for the following criteria: large litter size at birth (L⁺), large litter size and small 6-week body weight (L⁺W^–), or small litter size and large 6-week body weight (L^–W⁺). Females were paired with a mature intact male of the same line at 3, 5 or 7 weeks of age. Mean mating age, averaged over lines, was 26.5 ± .3, 38.3 ± .3 and 52.7 ± .3 days. Exposure to a mature male accelerated female sexual maturation in each line. When contrasted with their sibs mated at a later age, early-pregnant females from each line exhibited a decline in one or more component of reproductive performance, suggesting that the physiological state of the very young female was not optimum for normal pregnancy. In comparisons of early and later mating ages, all three lines showed a decreased littering rate at first mating, number born alive, and individual birth weight of progeny adjusted for litter size; L⁺ and L⁺W^– mice showed an increased perinatal mortality rate; L⁺ and L^–W⁺ had a reduction in litter size at birth. When the L⁺, L⁺W^– and L^–W⁺ lines were compared with an unselected strain and a line selected for high postweaning gain in similar experiments, a genotype by environment interaction was apparent since all lines did not respond in a similar manner to early mating. The line ranking for litter size at birth for each age at male-exposure was L⁺>L⁺W^–>L^–W⁺, despite the significant line by age interaction. When litter size was adjusted by covariance for body weight at mating, the significant effects of age at male-exposure and line by age interaction were eliminated. All fertile females were remated after they had weaned their first litter to obtain information on litter size in parity two. Line differences in litter size at birth and number born alive were uniform across parities. An age by parity interaction was evident since the decreased fecundity at younger ages of male exposure in the L⁺ and L^–W⁺ litters of parity one was not evident in parity two. Litter feed efficiency during first parity gestation was defined as litter birth weight divided by either cumulative feed intake of the dam from mating to parturition (GEI) or cumulative feed intake from weaning to parturition (GEII). The ranking of lines for GEI and GEH was L⁺> L⁺W^–>L^–W⁺, but when feed efficiency was adjusted for littering rate, L⁺W^– and L^–W⁺ were not significantly different. With regard to age at mating, the ranking for GEI (7 wk > 5 wk > 3 wk) was reversed from GEII (3 wk >5 wk > 7 wk) and these significant differences were maintained after adjustment for littering rate.Paper No. 6302 of the Journal Series of the North Carolina Agricultural Research Service, Raleigh. The use of trade names in this publication does not imply endorsement by the North Carolina Agricultural Research Service of the products named, nor criticism of similar ones not mentionedOn leave at the Institute of Animal Genetics, West Mains Road, Edinburgh EH9 3JN, Scotland. Supported, in part, by an Underwood Fund Fellowship, Agricultural Research Council, and an Edinburgh University Biological Fellowship in the Department of Genetics     

Long-Term Selection Response for 12-Day Litter Weight in Mice

Long-term selection for increased 12-day litter weight in two replicate lines (W₂, W₃) of mice resulted in an apparent selection limit at about 17 generations. Quadratic polynomial and exponential models were fitted to the data in order to estimate the plateaued response and half-life of the selection process. Using the polynomial results, the half-life estimates were 4.5 and 8.6 generations for W₂ and W₃, respectively. The plateaued responses were 5.1 and 5.8 g which, when expressed in phenotypic standard deviation units, became 1.1 and 1.3. The exponential model provides similar estimates. A negative association between 12-day litter weight and fitness was not considered to be an adequate explanation for the plateau since there was no decrease in fertility of the selected lines. Evidence that exhaustion of genetic variability was not the cause of the plateau came from the immediate response to reverse selection. It was proposed that the plateau may be due to a negative genetic correlation between direct and maternal genetic effects, which would be expected to occur after many generations of selection. There were positive correlated responses in both replicates for adult body weight, which was in agreement with the positive genetic correlation between preweaning and postweaning body weight. The expected positive correlated response for number born was realized in only one of the replicates.     

Single-trait and antagonistic index selection for litter size and body weight in mice

Individual selection based on female performance only was conducted in four lines of mice: L⁺ for increased litter size, W⁺ for increased 6-week body weight, L^-W⁺ for a selection index aimed at decreasing litter size and increasing 6-week body weight and L⁺W^- for a selection index aimed at increasing litter size and decreasing 6-week body weight. A fifth line (K) served as an unselected control. All litters were standardized to eight mice at one day of age. Expected heritability was based on twice the regression of offspring on dam (h²_d), which contains additive genetic variance due to direct (σ²_Ao) and maternal (σ²_Am) effects and their covariance (σ_AoAm). Responses and correlated responses were measured either deviated (method 1) or not deviated (method 2) from the control line. Realized heritabilities (h²_R) for litter size were 0.19 ± 0.04 (1) and 0.16 ± 0.03 (2), which were similar to h ²_d of 0.17 ± 0.04. The h² _R for 6-week body weight of 0.55 ± 0.07 (1) and 0.44 ± 0.07 (2) agreed with h²_d of 0.42 ± 0.02. Realized genetic correlations (r*_GR) between litter size and 6-week body weight calculated from the double-selection experiment were 0.52 ± 0.10 (1) and 0.52 ± 0.13 (2), which were not significantly different from the base population estimate of r* _Gd = 0.63 ± 0.14. Divergence (L^-W ⁺ minus L⁺W^-) in the antagonistic index selection lines was 0.21 ± 0.01 index units (I = 0.305 P_W - 0.436 P_L, where P _W and P_L are the phenotypic values for 6-week body weight and litter size, respectively.). The h² _R of index units of 0.14 ± 0.02 calculated from divergence agreed with h²_d of 0.14 ± 0.04. Divergences in litter size (-0.19 ± 0.07) and 6-week body weight (0.46 ± 0.10) were in the expected direction. Antagonistic index selection yielded about one-half the expected divergence in litter size, while divergence in 6-week body weight was only slightly less than expected. Realized genetic correlations indicated that litter size, 6-week body weight and index units each showed positive pleiotropy with 3-week body weight, postweaning gain and weight at vaginal introitus and negative pleiotropy with age at vaginal introitus. Sex ratio and several components of fitness (days from joining to parturition, percent fertile matings and percent perinatal survival) did not change significantly in the selected lines.     

Consequences of a low litter birth weight phenotype for postnatal lean growth performance and neonatal testicular morphology in the pig

The consequences of a low litter average birth weight phenotype for postnatal growth performance and carcass quality of all progeny, and testicular development in male offspring, were investigated. Using data from 25 sows with one, and 223 sows with two consecutive farrowing events, individual birth weight (BW) was measured and each litter between 9 and 16 total pigs born was classified as low (LBW), medium (MBW) or high (HBW) birth weight: low and high BW being defined as >1 standard deviation below or above, respectively, the population mean for each litter size. Litter average BW was repeatable within sows. At castration, testicular tissue was collected from 40 male pigs in LBW and HBW litters with individual BW close to their litter average BW and used for histomorphometric analysis. LBW piglets had a lower absolute number of germ cells, Sertoli cells and Leydig cells in their testes and a higher brain : testis weight ratio than HBW piglets. Overall, LBW litters had lower placental weight and higher brain : liver, brain : intestine and brain : Semitendinosus muscle weight ratios than MBW and HBW litters. In the nursery and grow–finish (GF) phase, pigs were kept in pens by BW classification (9 HBW, 17 MBW and 10 LBW pens) with 13 males and 13 females per pen. Average daily gain tended to be lower in LBW than HBW litters in lactation (P = 0.06) and throughout the nursery and GF phases (P < 0.01), resulting in an increasing difference in body weight between LBW, MBW and HBW litters (P < 0.05). Average daily feed intake was lower (P < 0.001) in LBW than HBW litters in the nursery and GF phases. Feed utilization efficiency (feed/gain) was similar for LBW and HBW litters in the nursery, but was lower (P < 0.001) in HBW than LBW litters in the GF phase. By design, slaughter weight was similar between BW classifications; however, LBW litters needed 9 more days to reach the same slaughter weight than HBW litters (P < 0.001). BW classification did not affect carcass composition traits. In conclusion, LBW litters showed benchmarks of intrauterine growth retardation, LBW had a negative impact on testicular development and germ and somatic cell populations, and was associated with decreased postnatal growth during all phases of production; however, no measurable effect on carcass composition traits was established.     

Heterosis is common and inbreeding depression absent in natural populations of Arabidopsis thaliana

The importance of genetic drift in shaping patterns of adaptive genetic variation in nature is poorly known. Genetic drift should drive partially recessive deleterious mutations to high frequency, and inter‐population crosses may therefore exhibit heterosis (increased fitness relative to intra‐population crosses). Low genetic diversity and greater genetic distance between populations should increase the magnitude of heterosis. Moreover, drift and selection should remove strongly deleterious recessive alleles from individual populations, resulting in reduced inbreeding depression. To estimate heterosis, we crossed 90 independent line pairs of Arabidopsis thaliana from 15 pairs of natural populations sampled across Fennoscandia and crossed an additional 41 line pairs from a subset of four of these populations to estimate inbreeding depression. We measured lifetime fitness of crosses relative to parents in a large outdoor common garden (8,448 plants in total) in central Sweden. To examine the effects of genetic diversity and genetic distance on heterosis, we genotyped parental lines for 869 SNPs. Overall, genetic variation within populations was low (median expected heterozygosity = 0.02), and genetic differentiation was high (median F_ST = 0.82). Crosses between 10 of 15 population pairs exhibited significant heterosis, with magnitudes of heterosis as high as 117%. We found no significant inbreeding depression, suggesting that the observed heterosis is due to fixation of mildly deleterious alleles within populations. Widespread and substantial heterosis indicates an important role for drift in shaping genetic variation, but there was no significant relationship between fitness of crosses relative to parents and genetic diversity or genetic distance between populations.     

Effect of average litter weight in pigs on growth performance,carcass characteristics and meat quality of the offspring as depending on birth weight

Offspring born from normal litter size (10 to 15 piglets) but classified as having lower than average birth weight (average of the sow herd used: 1.46 ± 0.2 kg; mean ± s.d.) carry at birth negative phenotypic traits normally associated with intrauterine growth restriction, such as brain-sparing and impaired myofiber hyperplasia. The objective of the study was to assess long-term effects of intrauterine crowding by comparing postnatal performance, carcass characteristics and pork quality of offspring born from litters with higher (>1.7 kg) or lower (<1.3 kg) than average litter birth weight. From a population of multiparous Swiss Large White sows (parity 2 to 6), 16 litters with high (H = 1.75 kg) or low (L = 1.26 kg) average litter birth weight were selected. At farrowing, two female pigs and two castrated pigs were chosen from each litter: from the H-litters those with the intermediate (H_I = 1.79 kg) and lowest (H_L = 1.40 kg) birth weight, and from L-litters those with the highest (L_H = 1.49 kg) and intermediate (L_I = 1.26 kg) birth weight. Average birth weight of the selected H_I and L_I piglets differed (P < 0.05), whereas birth weight of the H_L- and L_H-piglets were similar (P > 0.05). These pigs were fattened in group pen and slaughtered at 165 days of age. Pre-weaning performance of the litters and growth performance, carcass and meat quality traits of the selected pigs were assessed. Number of stillborn and pig mortality were greater (P < 0.05) in L- than in H-litters. Consequently, fewer (P < 0.05) piglets were weaned and average litter weaning weight decreased by 38% (P < 0.05). The selected pigs of the L-litters displayed catch-up growth during the starter and grower–finisher periods, leading to similar (P > 0.05) slaughter weight at 165 days of age. However, H_L-gilts were more feed efficient and had leaner carcasses than H_I-, L_H- and L_I-pigs (birth weight class × gender interaction P < 0.05). Meat quality traits were mostly similar between groups. The marked between-litter birth weight variation observed in normal size litters had therefore no evident negative impact on growth potential and quality of pigs from the lower birth weight group.     

Population Size and Selection Intensity Effects on Long-Term Selection Response in Mice

Long-term response to within full-sib family selection for increased postweaning gain was evaluated in lines having different effective population sizes (N_e) and selection intensities (i). Line designations were I4(4), I8(2), I16(2), M4(4), M8(2) and M16(2), where I and M indicate selection of the top 50% and 25%, respectively; 4, 8 and 16 represent the number of parental pairs per replicate and number of replicates is given in parentheses. Realized within full-sib family heritabilities (h_R²) in the first phase of selection (0-14 generations) were larger in 16-pair lines than in 4- and 8-pair lines. In the second phase of selection (>14 generations), h_R² declined significantly (P<.01) in all lines, and only the I16 and M16 lines had h_R² values significantly (P<.01) greater than zero. Realized genetic correlations involving number born, 12-day litter weight, weaning weight and six-week weight tended to decline in the second phase of selection. The I16, M16 and control (C16) replicates were crossed in all combinations at generation 14. Crosses were then selected within litters for high postweaning gain. The h_R² values in the crossbred lines were all larger than those in the second selection phase for M16-1, M16-2 and I16-1, but not for I16-2. Within each N_e level, total response was significantly (P<.01) less for I lines compared with M lines. Total response increased as N_e increased, within each level of i. Relatively small differences in realized i values among N_e lines could not account for this result. The difference in total response among the N_e lines at a given selection intensity may be due to inbreeding depression and a combination of interactions involving "drift" and selection. By crossing replicates of the M lines with the C16 control, the effects of inbreeding depression were removed. Inbreeding depression and genetic drift, as defined herein, were equally important in accounting for differences among N_e lines in total response.     

Heterosis and outbreeding depression in crosses between natural populations of Arabidopsis thaliana

Understanding the causes and architecture of genetic differentiation between natural populations is of central importance in evolutionary biology. Crosses between natural populations can result in heterosis if recessive or nearly recessive deleterious mutations have become fixed within populations because of genetic drift. Divergence between populations can also result in outbreeding depression because of genetic incompatibilities. The net fitness consequences of between-population crosses will be a balance between heterosis and outbreeding depression. We estimated the magnitude of heterosis and outbreeding depression in the highly selfing model plant Arabidopsis thaliana, by crossing replicate line pairs from two sets of natural populations (C↔R, B↔S) separated by similar geographic distances (Italy↔Sweden). We examined the contribution of different modes of gene action to overall differences in estimates of lifetime fitness and fitness components using joint scaling tests with parental, reciprocal F₁ and F₂, and backcross lines. One of these population pairs (C↔R) was previously demonstrated to be locally adapted, but locally maladaptive quantitative trait loci were also found, suggesting a role for genetic drift in shaping adaptive variation. We found markedly different genetic architectures for fitness and fitness components in the two sets of populations. In one (C↔R), there were consistently positive effects of dominance, indicating the masking of recessive or nearly recessive deleterious mutations that had become fixed by genetic drift. The other set (B↔S) exhibited outbreeding depression because of negative dominance effects. Additional studies are needed to explore the molecular genetic basis of heterosis and outbreeding depression, and how their magnitudes vary across environments.     

Milk yield in lines of mice selected for growth or maternal ability.

Two experiments were conducted to measure milk yield as a correlated response to selection for increased 12-day litter weight (maternal lines) for 13 generations and increased and decreased body weight gain from 21 to 42 days of age (postweaning gain lines) for 12 to 14 generations. Milk yield was measured from day 6 to day 21 of lactation by separating litters from their dams for 6 h, then weighing litters before and after a 1.5 h suckling period. Average total milk production was 30.3 +/- 0.9 g and 28.6 +/- 1.1 g in maternal and control lines, respectively. This difference was not significant (P greater than 0.05). Differences among gain lines were significant (P less than 0.01) in the second experiment with average production of 44.1 +/- 1.4 g for increased gain lines, 22.5 +/- 0.9 g for the decreased gain line and 31.2 +/- 1.0 g for the control. These results indicated that response to selection for increased 12-day litter weight was not associated with increases in milk yield, but sizable changes inmilk yield accompanied bi-directional selection for early postweaning gain.     

Effects of dietary enrichment with a marine oil-based n-3 LCPUFA supplement in sows with predicted birth weight phenotypes on birth litter quality and growth performance to weaning

The effects of a marine oil-based n-3 long-chain polyunsaturated fatty acid (mLCPUFA) supplement fed to the sow from weaning, through the rebreeding period, during gestation and until end of lactation on litter characteristics from birth until weaning were studied in sows with known litter birth weight phenotypes. It was hypothesized that low birth weight (LBW) litters would benefit more from mLCPUFA supplementation than high birth weight litters. A total of 163 sows (mean parity=4.9±0.9) were rebred after weaning. Sows were pair-matched by parity and litter average birth weight of the previous three litters. Within pairs, sows were allocated to be fed either standard corn/soyabean meal-based gestation and lactation diets (CON), or the same diets enriched with 0.5% of the mLCPUFA supplement at the expense of corn. Each litter between 9 and 16 total pigs born was classified as LBW or medium/high average birth weight (MHBW) litter and there was a significant correlation (P<0.001) between litter average birth weight of the current and previous litters within sows (r=0.49). Sow serum was harvested at day 113 of gestation for determination of immunoglobulin G (IgG) concentrations. The number of pigs born total and alive were lower (P=0.01) in mLCPUFA than CON sows, whereas the number of stillborn and mummified pigs were similar between treatments. Number of stillborns (trend) and mummies (P<0.01) were higher in LBW than MHBW litters. Tissue weights and brain : tissue weight ratios were similar between treatments, but LBW litters had decreased tissue weights and increased brain : tissue weight ratios compared with MHBW litters. Placental weight was lower (P=0.01) in LBW than MHBW litters, but was not different between treatments. Average and total litter weight at day 1 was similar between treatments. mLCPUFA increased weaning weight (P=0.08) and average daily gain (P<0.05) in MHBW litters, but not in LBW litters. Pre-weaning mortality was similar between treatments, but was higher (P<0.01) in LBW than MHBW litters. IgG concentration in sow serum was similar between treatments and litter birth weight categories. In conclusion, litter birth weight phenotype was repeatable within sows and LBW litters showed the benchmarks of intra-uterine growth retardation (lower placental weight and brain sparing effects). As maternal mLCPUFA supplementation decreased litter size overall, only improved litter growth rate until weaning in MHBW litters, and did not affect pre-weaning mortality, maternal mLCPUFA supplementation was not an effective strategy in our study for mitigating negative effects of a LBW litter phenotype.     

Selection for nursing ability and adult weight in mice

Three selection treatments were conducted for 12 generations in each of two base populations (P and Q): (1) increased nursing ability of the mother (n12), as measured by mean 12-day weight of eight young within a crossfostering set (M(P) and M(Q) lines), (2) increased adult (42-day) body weight of the offspring (w42) (W(P) and W( Q) lines), and (3) performance combining the two traits (n12 and w42) into a selection index (B(P) and B(Q) lines). Lines C( P) and C(Q) were maintained as unselected controls in each population. In each line-generation subclass, 92 single-pair matings were made and the offspring assigned to balanced crossfostering sets of four dams each. Regression coefficients of mean performance (in grams) on generations were 0.080 +/-0.029 and 0.054 +/- 0.031 for n12 in M(P) and M(Q), and 0.680 +/- 0.039 and 0.868 +/- 0.051 for w42 in W(P) and W(Q), respectively. The B(P) and B(Q) lines showed genetic gains in n12 (0.090 and 0.053, respectively) and w42 (0.576 and 0.696) intermediate between the performance of M(P) and W(P), and M(Q) and W(Q), respectively, except for n12 of B(Q). Realized heritabilities for n12 were 0.16 +/- 0.05 and 0.11 +/- 0.06 and those for w42 were 0.40 +/- 0.02 and 0.43 +/- 0.03 for P and Q, respectively. The realized genetic correlations between n12 and w42 were 0.70 +/- 0.07 and 0.73 +/- 0.08 in P and Q, respectively. The ratios of the predicted to observed responses in M(P), B(P) and B(Q) were 0.99, 1.03 and 0.89, respectively. However, the predicted and observed responses differed in M( Q), W(P) and W(Q); the ratios were 1.29, 0.65 and 0.65, respectively. The observed combined responses for n12 and w42 in the index lines (B(P) and B(Q)) were smaller than the optimum expected from index selection. A possible cause was that the estimated genetic correlations (0.22 +/- 0.16 and -0.17 +/- 0.16 for B(P) and B( Q), respectively) and heritabilities (0.39 +/- 0.03 and 0.28 +/- 0.02, respectively) for w42 that were used to construct the selection index were smaller than the respective realized parameters.     

Sow and piglet factors determining variation of colostrum intake between and within litters

Colostrum intake has a short- and long-term beneficial impact on piglet performance and mortality. Sows’ colostrum production and piglets’ colostrum intake are limited and highly variable. The present study investigated sow and piglet factors explaining the variation of colostrum intake between and within litters. The CV for colostrum intake and birth weight (BW_b) of all piglets within a litter was calculated to evaluate the variation of colostrum intake and BW_b within a litter (colostrum and litter BW_b heterogeneity, respectively). A total of 1937 live-born piglets from 135 litters from 10 commercial herds were included. Colostrum intake per piglet averaged 371±144 g and was affected by breed (P=0.02). It was lower when oxytocin was administered to the sow during parturition (P=0.001) and with increased litter size (P<0.001). It was higher when the interval between birth and first suckling decreased (t_FS, P<0.001). Colostrum intake was positively influenced by BW_b (P<0.001) and this association was more pronounced in piglets from Topigs (P=0.03) and Hypor (P=0.03) sows compared with piglets from Danbred sow breeds. The positive relationship between colostrum intake and BW_b was more pronounced when t_FS lasted longer (P=0.009). Heterogeneity in colostrum intake averaged 31±11%, it increased when oxytocin was applied during farrowing (P=0.004) and when stillbirth occurred (P=0.006). Colostrum heterogeneity was positively associated with litter size (P<0.001) and litter BW_b heterogeneity (P=0.01). The positive relationship between colostrum and litter BW_b heterogeneity was more pronounced when oxytocin was applied during farrowing (P=0.04). The present study demonstrated that oxytocin should be used cautiously in sows during farrowing. Farrowing and colostrum management should prevent or counteract the adverse influences of stillbirth, large and heterogeneous litters on colostrum intake and colostrum heterogeneity. The study also confirmed the expected association between BW_b and colostrum intake and indicated that the impact of BW_b on colostrum intake was different among breeds (Hypor v. Danbred) and dependent on piglets’ latency to first suckling. Hence, colostrum management should focus on low birth weight piglets, especially in some breeds, and low colostrum intake in low birth weight piglets can be counteracted by shortening the t_FS.     

Correlated responses in growth and body composition of replicated single-trait and index selected lines of mice

Summary Correlated responses in growth, body composition and efficiency were evaluated in lines of mice selected in the following ways: W⁺T _i ^o , increased six-week body weight (WT6); W ° T _i ⁺ , increased six-week tail length (TL6); W⁺T _i ^– , increased WT6 and decreased TL6; W^–T _i ⁺ , decreased WT6 and increased TL6; M16, increased three-to six-week postweaning gain (PWG). Each of the first four selection treatments had two replicate lines (i = 1, 2) selected for 13 generations and the fifth treatment had one line selected for 30 generations. All lines were derived from a randombred ICR albino population which served as a control. Additional traits studied were three-week body weight and tail length, postweaning gain in tail length, percent body composition (ash, fat, moisture and protein) at six weeks of age, and three-to six-week feed consumption (CONS) and efficiency (EFF = PWG/CONS). Efficiency of body constituent gains (ash, fat, protein and caloric value) were determined by dividing each constituent by CONS. Relative to selection treatments, replicate variation in the array of traits was small and was primarily attributable to the effects of genetic drift; more frequent significant replicate differences among traits in W⁺T^– were associated with a replicate difference in cumulative selection differentials. Selection for different criteria involving WT6 and TL6 did not change the allometric relationship between tail length and body weight in the three-to six-week age interval. The significant divergence between W⁺T ° and W °T⁺ and between W⁺T^– and W^–T⁺ was as expected for WT6 and TL6. Significant asymmetry of selection response between W⁺T^– and W^–T⁺ for WT6 and TL6 was attributed to maternal effects. In agreement with theory, antagonistic index selection generally yielded smaller genetic responses than single trait selection. Positive correlated responses in CONS and EFF were found for M16 and W⁺T °. Significant correlated changes in CONS (positive in W °T⁺ and negative in W^–T⁺) were not accompanied by a significant change in EFF. In contrast, W⁺T^– evinced an increased EFF and no change in CONS. Percent fat increased significantly in W⁺T ° and M16. For W⁺T^o, W⁺T^– and M16, an increased energetic, fat and ash efficiency was observed, whereas M16 exhibited a positive increment in protein efficiency as well. Among selection treatment means, there were high positive correlations between WT6 and fat weight, protein weight, percent fat, CONS and EFF and a high negative correlation between WT6 and percent protein.Paper No.4916 of the Journal Series of the North Carolina Agricultural Experiment Station, Raleigh, N.C. 27607. The use of trade names in this publication does not imply endorsement by the North Carolina Agricultural Experiment Station of the products named, nor criticism of similar ones not mentioned.     

A general mixture model approach for mapping quantitative trait loci from diverse cross designs involving multiple inbred lines

Most current statistical methods developed for mapping quantitative trait loci (QTL) based on inbred line designs apply to crosses from two inbred lines. Analysis of QTL in these crosses is restricted by the parental genetic differences between lines. Crosses from multiple inbred lines or multiple families are common in plant and animal breeding programmes, and can be used to increase the efficiency of a QTL mapping study. A general statistical method using mixture model procedures and the EM algorithm is developed for mapping QTL from various cross designs of multiple inbred lines. The general procedure features three cross design matrices, W, that define the contribution of parental lines to a particular cross and a genetic design matrix, D, that specifies the genetic model used in multiple line crosses. By appropriately specifying W matrices, the statistical method can be applied to various cross designs, such as diallel, factorial, cyclic, parallel or arbitrary-pattern cross designs with two or multiple parental lines. Also, with appropriate specification for the D matrix, the method can be used to analyse different kinds of cross populations, such as F2 backcross, four-way cross and mixed crosses (e.g. combining backcross and F2). Simulation studies were conducted to explore the properties of the method, and confirmed its applicability to diverse experimental designs.     

ON THE RELATION BETWEEN LITTER SIZE,BIRTH WEIGHT,AND RATE OF GROWTH,IN MICE

We have been concerned with the connection between size of litter and weight of litter at birth, especially in mice. The weight at birth represents, it is to be presumed (at least in mice, and for certain other cases), the weight at a particular developmental stage. The connection between number in litter (N) and weight of litter (W) has been interpreted as due to the partition of nourishment between mother and young, and on an equal basis among the several embryos of a litter. The "heterogonic" relationship which the data exhibit between N and W shows that the constant K, defined by log W = K log N + const., is independent of the species, and has an essentially constant value (0.85±) in all multiparous mammals; it is therefore regarded as a partition coefficient. In the case of power function relationships between masses of components of a single individual, the respective "drawing powers" of the several organs are diverse, and diverse magnitudes of K are encountered. With developing embryos, the intrinsic drawing powers of the tissues concerned in embryos and mothers are in each case of the same general character, at least among mammals; the constancy of K reflects this. A parallel for the case as it appears in the consideration of relative growth rates of organs in a single individual, and in which the varying magnitudes of the heterogonic growth constant K are presumed to reflect diverse drawing powers of the respective tissues, would be given by intrauterine growth of a litter containing individuals with diverse capacities for growth, —that is, individuals differing genetically with respect to the factors determining the magnitudes of w ₁. We have been dealing with the growth of litters in inbred strains. It is to be presumed that in the case of the growth of a litter containing two categories of individuals so far as concerns intrinsic drawing powers with respect to the nourishment provided by the mother, it would be possible to investigate the way in which K is open to modification. Although difficult, from the standpoint of classifying the individual young, it would appear to be distinctly worth while to make such an experiment, and we have planned it for the future. It is pointed out that for genetic purposes the ideal weight of a litter of 1 is obtainable from a series of measurements of N and W, free from disturbances affecting the apparent value of this quantity as observed in single births. This weight of an ideal litter of 1 should be employed to disentangle the effects of heterosis and fertility factors from those having to do with individual weight at birth. During the suckling period the relation ΔW/W = K (ΔN/N) is maintained for young mice, but with modifications in the case of small and large suckling litters due to (1) the time course of milk yield, and (2) the effect of litter size upon this. It is shown that a growth curve can be obtained for an ideal litter of I, under the condition of milk supply that on each day the mother is able to provide a constant fractional increase of milk for each additional young mouse in the litter. The rate of growth then adheres to the time curve of capacity for production of milk.     

Crossbreeding effects after long-term selection for purebred performance: a model experiment with mice

Summary Results are presented from two replicated three-breed cross diallels that were conducted after 20 generations of selection for purebred performance in mice. The selection criteria for the different lines were: litter size at birth (LS), weaning weight at 4 weeks (WW), weight gain from week 4 to week 6 (WG), and body fat content at week 6 (FT). Additionally, a random-mating control line (C) was kept. Significant maternal heterosis was found in litter size and weaning weight. Estimates of maternal heterosis in litter size were very high, ranging from 17 to 50% of the mean of the corresponding single crosses. Maternal heterosis in weaning weight usually was negative and ranged from +9 to -11%. Significant maternal heterosis in feed efficiency and weaning weight could only be found in a few cases. Total performance of three-breed crosses was highly superior to that of single crosses and purebreds. Means of the corresponding purebreds or single crosses were of little use in predicting three-breed cross performance.     

B-branch electron transfer in reaction centers of Rhodobacter sphaeroides assessed with site-directed mutagenesis

Mutants of Rhodobacter (Rba.) sphaeroides are described which were designed to study electron transfer along the so-called B-branch of reaction center (RC) cofactors. Combining the mutation L(M214)H, which results in the incorporation of a bacteriochlorophyll, β, for H_A [Kirmaier et al. (1991) Science 251: 922–927] with two mutations, G(M203)D and Y(M210)W, near B_A, we have created a double and a triple mutant with long lifetimes of the excited state P^* of the primary donor P, viz. 80 and 160 ps at room temperature, respectively. The yield of P⁺Q_A ⁻ formation in these mutants is reduced to 50 and 30%, respectively, of that in wildtype RCs. For both mutants, the quantum yield of P⁺H_B ⁻ formation was less than 10%, in contrast to the 15% B-branch electron transfer demonstrated in RCs of a similar mutant of Rba. capsulatus with a P^* lifetime of 15 ps [Heller et al. (1995) Science 269: 940–945]. We conclude that the lifetime of P^* is not a governing factor in switching to B-branch electron transfer. The direct photoreduction of the secondary quinone, Q_B, was studied with a triple mutant combining the G(M203)D, L(M214)H and A(M260)W mutations. In this triple mutant Q_A does not bind to the reaction center [Ridge et al. (1999) Photosynth Res 59: 9–26]. It is shown that B-branch electron transfer leading to P⁺Q_B ⁻ formation occurs to a minor extent at both room temperature and at cryogenic temperatures (about 3% following a saturating laser flash at 20 K). In contrast, in wildtype RCs P⁺Q_B ⁻ formation involves the A-branch and does not occur at all at cryogenic temperatures. Attempts to accumulate the P⁺Q_B ⁻ state under continuous illumination were not successful. Charge recombination of P⁺Q_B ⁻ formed by B-branch electron transfer in the new mutant is much faster (seconds) than has been previously reported for charge recombination of P⁺Q_B ⁻ trapped in wildtype RCs (10⁵ s) [Kleinfeld et al. (1984b) Biochemistry 23: 5780–5786]. This difference is discussed in light of the different binding sites for Q_B and Q_B ⁻ that recently have been found by X-ray crystallography at cryogenic temperatures [Stowell et al. (1997) Science 276: 812–816]. We present the first low-temperature absorption difference spectrum due to P⁺Q_B ⁻. This revised version was published online in June 2006 with corrections to the Cover Date.