Divergent selection for uterine capacity in rabbits. II. Correlated response in litter size and its components estimated with a cryopreserved control population

Our objective was to evaluate the correlated responses to selection for litter size and its components after 10 generations of divergent selection for uterine capacity (UC). A total of 294 intact females from the 11th and 12th generations of divergent selection for high and low UC and from a cryopreserved control population was used (139, 112, and 43 females, respectively). Uterine capacity was assessed as litter size in unilaterally ovariectomized females. Traits recorded on females for up to five parities were litter size (LS) and number born alive (NBA). Laparoscopy was performed in all females at d 12 of their second parity, and the ovulation rate (OR) and number of implanted embryos (IE) were recorded in these females. Embryo survival (ES = IE/OR), fetal survival (FS = LS/IE), and prenatal survival (PS = LS/OR) were computed. Correlated responses in LS and in its components were inferred using Bayesian methods. Correlated responses in LS were asymmetric. The divergence between high and low lines was 2.35 kits, mainly because of a higher correlated response in the low line (1.88 kits). The lower LS in the low line was associated with a lower PS (control - low = 0.14), because of decreases in ES and FS.     

Divergent selection for uterine capacity in rabbits. I. Genetic parameters and response to selection

A 10-generation divergent selection experiment for uterine capacity (UC) measured as litter size in unilaterally ovariectomized females was carried out in rabbits. A total of 2,996 observations on uterine capacity of does (up to four parities) was recorded. Laparoscopy was performed at d 12 of their second gestation, and ovulation rate (OR) and number of implanted embryos (IE) were recorded in 735 does. Prenatal survival (PS) was assessed as UC/OR, embryo survival (ES) as IE/OR, and fetal survival (FS) as UC/IE. Genetic parameters and genetic trends were inferred using Bayesian methods. Marginal posterior distributions of all unknowns were estimated by Gibbs sampling. Heritabilities of UC, OR, IE, ES, FS, and PS were 0.11, 0.32, 0.22, 0.04, 0.14, and 0.09, respectively. Genetic and phenotypic correlations between FS and ES were low, suggesting different biological mechanisms for the two periods of survival. After 10 generations of selection, the divergence was approximately 1.5 rabbits, or approximately 1% per generation. Approximately one-half of this response was obtained in the first two generations of selection, which may suggest the presence of a major gene segregating in the base population.     

Selection for ovulation rate in rabbits: genetic parameters and correlated responses on survival rates

The aim of this work was to evaluate the correlated responses on survival rates after 10 generations of selection for ovulation rate (OR). Selection was based on the phenotypic value of ovulation rate estimated at d 12 of second gestation by laparoscopy. Traits recorded were litter size (LS), estimated as total number of rabbits born per litter in up to 5 parities; OR, estimated as the number of corpora lutea in both ovaries; the number of implanted embryos (IE), estimated as the number of implantation sites; the number of right and left IE (RIE and LIE); ovulatory difference (OD), defined as the difference between the right and the left OR, expressed as an absolute value; implantatory difference (ID), defined as the difference between RIE and LIE, expressed as an absolute value; embryonic survival (ES), calculated as IE/OR; fetal survival (FS), calculated as LS/IE; prenatal survival (PS), calculated as LS/OR. A total of 1,081 records were used to analyze ES, and 770 were used to analyze FS and PS. The number of records used to analyze the other traits ranged from 1,079 for ID to 3,031 for LS. Data were analyzed using Bayesian methodology. Genetic parameters of OR, OD, and LS were estimated in a previous paper. Estimated heritabilities of IE, ID, ES, FS, and PS were 0.11, 0.03, 0.09, 0.24, and 0.14, respectively. Estimated repeatabilities of IE, ID, and ES were 0.22, 0.12, and 0.20. Estimated phenotypic correlations of OR with ES, FS, and PS were -0.07, -0.26, and -0.28, respectively. Their estimated genetic correlations with FS and PS were negative (probability of being negative 1.00 and 0.98, respectively). Nothing can be said about the sign of the genetic correlation between OR and ES. Ovulation rate was phenotypically uncorrelated with ID. Their estimated genetic correlation was positive (probability of being positive 0.91). The genetic correlation of ID with PS and LS was not accurately estimated. Phenotypic and genetic correlations between LS and survival rates were positive (probability of being positive 1.00). In 10 generations of selection, FS decreased around 1% per generation. No correlated response in ES was observed. In summary, the decrease in FS in rabbits selected for OR seemed to be responsible for the lack of correlated response observed in LS.     

Selection for ovulation rate in rabbits

An experiment of selection for ovulation rate was carried out. Animals were derived from a synthetic line first selected 12 generations for litter size, then 10 generations for uterine capacity. Selection was relaxed for 6 generations. Selection was based on the phenotypic value of ovulation rate with a selection pressure on does of 30%. Males were selected from litters of does with the highest ovulation rate. Males were selected within sire families in order to reduce inbreeding. Ovulation rate was measured in the second gestation by a laparoscopy, 12 days after mating. Each generation had about 80 females and 20 males. Results of three generations of selection were analyzed using Bayesian methods. Marginal posterior distributions of all unknowns were estimated by Gibbs sampling. Heritabilities of ovulation rate (OR), number of implanted embryos (IE), litter size (LS), embryo survival (ES), fetal survival (FS), and prenatal survival (PS) were 0.44, 0.32, 0.11, 0.26, 0.35, and 0.14, respectively. Genetic correlation between OR and LS was 0.56, indicating that selection for ovulation rate can augment litter size. Response to selection for OR was 1.80 ova. Correlated responses in IE and LS were 1.44 and 0.49, respectively. Selection for ovulation rate may be an alternative to improve litter size.     

Correlated response in litter size components in rabbits selected for litter size variability

A divergent selection experiment for the environmental variability of litter size (Ve) over seven generations was carried out in rabbits at the University Miguel Hernández of Elche. The Ve was estimated as the phenotypic variance within the female, after correcting for year‐season and parity‐lactation status. The aim of this study was to analyse the correlated responses to selection in litter size components. The ovulation rate (OR) and number of implanted embryos (IE) in females were measured by laparoscopy at 12 day of the second gestation. At the end of the second gestation, the total number of kits born was measured (TB). Embryonic (ES), foetal (FS) and prenatal (PS) survival were computed as IE/OR, TB/IE and TB/OR, respectively. A total of 405 laparoscopies were performed. Data were analysed using Bayesian methodology. The correlated response to selection for litter size environmental variability in terms of the litter size components was estimated as either genetic trends, estimated by computing the average estimated breeding values for each generation and each line, or the phenotypic differences between lines. The OR was similar in both lines. However, after seven generations of selection, the homogenous line showed more IE (1.09 embryos for genetic means and 1.23 embryos for phenotypic means) and higher ES than the heterogeneous one (0.07 for genetic means and 0.08 for phenotypic means). The probability of the phenotypic differences between lines being higher than zero (p) was 1.00 and .99, respectively. A higher uterine overcrowding of embryos in the homogeneous line did not penalize FS; as a result, this line continued to show a greater TB (1.01 kits for genetic means and 1.30 kits for phenotypic means, p = .99, in the seventh generation). In conclusion, a decrease in litter size variability showed a favourable effect on ES and led to a higher litter size at birth.     

Selection for ovulation rate in rabbits: Direct and correlated responses estimated with a cryopreserved control population

The aim of this work was to evaluate the response in 10 generations of selection for ovulation rate in rabbits using a cryopreserved control population. Selection was based on the phenotypic value of ovulation rate estimated at d 12 of second gestation by laparoscopy. To produce the control population, embryos from 50 donor females and 18 males, belonging to the base generation of the line selected for ovulation rate, were recovered. A total of 467 embryos (72-h embryos) were vitrified and stored in liquid N(2) for 10 generations. The size of both populations was approximately 10 males and 50 females. The number of records used to analyze the different traits ranged from 99 to 340. Data were analyzed using Bayesian methodology. A difference between the selected and the control populations of 2.1 ova (highest posterior density interval (HPD(95%))[1.3, 2.9]) was observed in ovulation rate (OR), but it was not accompanied by a correlated response in litter size (LS; -0.3; HPD(95%) [-1.1, 0.5]). The number of implanted embryos (IE) increased with selection in 1.0 embryo (HPD(95%) [-0.6, 2.0]), but this increase was not relevant. Prenatal survival, embryonic survival, and fetal survival (FS) were calculated as LS/OR, IE/OR, and LS/IE, respectively. Prenatal survival was reduced with selection (-0.12; HPD(95%) [-0.20, -0.04]), basically because of a decrease in FS (-0.12; HPD(95%) [-0.19, -0.06]). Embryonic survival could have slightly decreased (-0.05; HPD(95%) [-0.12, 0.02]). In summary, comparison with a control population showed that ovulation rate in rabbits increased with selection without any correlated response in litter size, basically because of a decrease in fetal survival.     

Boar sperm quality in lines of pigs selected for either ovulation rate or uterine capacity

Selection for 11 generations in swine for ovulation rate (OR) or uterine capacity (UC) resulted in significant changes in component traits of litter size. Our objective was to conserve the unique germplasm for the future and to characterize sperm quality as a correlated response to the selection criterion imposed compared with an unselected control line (CO). Boars representing genetic diversity available in all 3 lines were produced in 2 farrowing seasons. Season 1 was born in September 2005 and was sampled for semen characteristics in October 2006. Season 2 was born in March 2006 and was sampled for semen characteristics in February and March 2007. Each boar (n = 60) was collected twice. The sperm-rich fraction was obtained, and volume and concentration of sperm cells were measured to estimate total sperm production. Each ejaculate was extended 1:3 (vol/vol) with Androhep Plus (Minitube, Verona, WI) and was packed for shipping to the National Animal Germplasm Program laboratory for processing into frozen straws. Semen quality was measured by computer-assisted semen analysis at 3 semen processing points: fresh (FR), 24 h after extender added (E), and postthaw (PT). A mixed model ANOVA was applied to the data. Fixed effects of farrowing season, line, and 2-way interactions were fitted. The random effect of boar (n = 60) within farrowing season and line was used to test line differences. Sperm concentration was not different (P = 0.18) among the lines (0.594 × 10(9), 0.691 × 10(9), and 0.676 × 10(9) cells/mL for CO, OR, and UC lines, respectively). However, significance (P = 0.04) was detected for the volume of the sperm-rich fraction, greatest for OR (86.4 mL), intermediate for UC (75.5 mL), and least for CO (70.2 mL). Line differences were thus detected (P = 0.02) for total sperm production per ejaculate, greatest for OR (54.9 × 10(9)), intermediate for UC (48.7 × 10(9)), and least for CO (40.5 × 10(9)). A larger percentage of progressively motile sperm and greater estimates of sperm velocity only at processing point E (P < 0.01) were detected in favor of CO. Estimates of motility, velocity, and other parameters of sperm movement measured on E processing points were positively correlated with the same estimates obtained PT, but the magnitude was low to moderate (r range -0.03 to 0.23). Thus, selection for component traits of female reproduction had a favorable effect on total sperm production of boars.     

Selection for components of reproduction in swine

The response per generation to 10 generations of mass selection for ovulation were 0.49 ova, ?1.6% in embryo survival and 0.06 piglets per litter at birth. Line differences (select-control) in generation 9 and 10 gilts and sows ranged from 3.4 to 5 ova. Control line gilts and sows had 5.4 to 10.6% higher embryo survival to days 30 and 70 of gestation than did select line females. One generation of random selection followed by four generations of litter size selection, selection for decreased age at puberty or relaxed ovulation rate selection in the high ovulation rate line has resulted in lines that differed from the control line in litter size at birth by 0.78 ± 0.22, 0.37 ± 0.39 and 0.84 ± 0.52 pigs per litter at first, second and third parity, respectively. These results were used to derive a selection index to increase litter size by selection for its components (ovulation rate, OR, and embryo survival, ES). A technique of selection based on laparotomy to increase the number of females tested with a given set of farrowing places is presented. Rate of response in LS from use of the selection index, I = 10.6 OR + 72.6 ES, in a population of 40 farrowing females and 15 males per generation, is expected to increase litter size 2.5 times faster than selection on LS due to higher selection intensity and optimum emphasis on the component traits.     

Relationships between uterine and fetal traits in rabbits selected on uterine capacity

The aim of this study was to investigate whether uterine capacity (UC) in rabbits was related to uterine horn length and weight and whether these uterine traits and vascular supply were related to fetal development and survival. Data from 48 unilaterally ovariectomized (ULO) does of the High and 52 ULO does of the Low UC lines of a divergent selection experiment on UC were used. Does were slaughtered on d 25 of fifth gestation. The High line showed higher ovarian weight (0.08 g, P < 0.05) linked to a higher ovulation rate (1 ovum, P < 0.05) and greater length of the empty uterine horn. There were no differences between lines in the remaining doe traits. The number of implanted embryos and live fetuses, fetal survival, and uterine weight and length were positively associated and explained most of the observed variation. Average weights of the live fetuses and their fetal and maternal placentae were not related to uterine weight and length. The linear regression coefficient of full uterine horn length on the number of live fetuses was 2.43 +/- 0.21. The weight of the full uterine horn showed a small quadratic relationship (P < 0.05) with the number of live fetuses. Full uterine horn length, after adjusting for the number of embryos, was negatively associated (P < 0.001) with the number of dead fetuses. The linear regression coefficient of average fetal placental weight of the live fetuses on number of implanted embryos was higher (P < 0.10) in the Low line (-0.23 +/- 0.04 vs. -0.12 +/- 0.04). The linear regression coefficient of average weight of the live fetuses on the average weight of their fetal placentae was higher (P < 0.10) in the High line (2.56 +/- 0.47 vs. 1.27 +/- 0.57). The High line was more efficient, most likely because an increase in intrauterine crowding has a lesser effect on the development of fetal placentae and fetuses. The fetal position within the uterus did not affect the proportion of dead embryos. Fetuses with placentae receiving a single blood vessel had a higher probability of death (P < 0.001) and the lowest weight. There was no difference between lines for individual weight of the live fetuses, but the High line showed higher individual weights of fetal (P < 0.01) and maternal placentae (P < 0.10). Live fetuses in the midportion of the uterus were lighter in weight (P < 0.05) than in the oviductal and cervical regions (20.3 vs. 21.6 and 21.7g). Increasing uterine capacity increases uterine length and decreases weights of fetus and fetal placenta in rabbits.     

Direct responses to six generations of selection for ovulation rate or prenatal survival in Large White pigs

Effects of selection for ovulation rate or prenatal survival were examined using data from 3 pigs lines derived from the same base Large White population. Two lines were selected for 6 generations on high ovulation rate at puberty (OR line) or high prenatal survival corrected for ovulation rate in the first 2 parities (PS line). The third line was an unselected control line. Genetic parameters for ovulation rate on the left, right, and both ovaries at puberty (ORPL, ORPR, and ORP, respectively) and at fertilization (ORFL, ORFR, and ORF, respectively), total number of piglets born (TNB) per litter, prenatal survival (PS = TNB/ORF), and PS corrected for ovulation rate (CPS = PS + 0.018ORF) were estimated using REML methodology. Responses to selection were estimated by computing differences between OR or PS and control lines at each generation using least squares and mixed models methodology. Average genetic trends were computed by regressing line differences on generation number. Realized heritabilities were estimated using standard procedures. Heritability estimates were 0.17, 0.11, 0.34, 0.13, 0.09, 0.33, 0.14, 0.11, and 0.17 (SE = 0.01 to 0.03) for ORPL, ORPR, ORP, ORFL, ORFR, ORF, PS, CPS, and TNB, respectively. Realized heritabilities were 0.37 +/- 0.08 and 0.10 +/- 0.09 for ORP and CPS, respectively. The different measures of ovulation rate had strong genetic correlations (r(g) > 0.7). The ORF had midrange negative genetic correlations with PS and CPS (-0.45 +/- 0.07 and -0.42 +/- 0.08, respectively). The ORP also had an antagonistic genetic relationship with PS (-0.26 +/- 0.07) but was almost independent from CPS (-0.02 +/- 0.11). The TNB was moderately correlated with ORP and ORF (r(g) = 0.41 +/- 0.09 for both traits). Average genetic trends in OR and PS lines were, respectively, 0.49 +/- 0.10 and 0.11 +/- 0.10 for ORP, and 0.43 +/- 0.11 and 0.11 +/- 0.11 for ORF. Responses to selection were slightly superior in the left than in the right ovary. No significant difference was found for PS or CPS in any of the lines. The TNB did not change in the OR line but significantly improved in the PS line (0.24 +/- 0.11 piglets/generation).     

Number of fetuses and conceptus growth throughout gestation in lines of pigs selected for ovulation rate or uterine capacity

Selection for 11 generations in swine for ovulation rate (OR) or uterine capacity (UC) resulted in 19.6% greater prenatal survival at term in UC compared with OR. Our objective was to characterize the number of fetuses throughout gestation in each line, including an unselected control (CO) line. Five hundred ninety-three gilts produced over 4 farrowing seasons were subjected to unilateral-hysterectomy-ovariectomy at 160 d of age and mated within line at 280 d of age. Gilts were assigned within sire family to be slaughtered (+/- 2 d) at d 25, 45, 65, 85, or 105 of gestation. Ovulation rate and number of live and dead fetuses were recorded for each pregnant gilt (n = 402). Fetal and placental weights were also recorded. Ovulation rate of OR line gilts (18.0 +/- 0.3 ova) exceeded (P < 0.001) CO and UC lines (15.0 +/- 0.3 and 14.0 +/- 0.3 ova, respectively). Line and gestational age interacted to affect number of live fetuses (P < 0.001). Least squares means for CO were 10.1, 8.3, 7.2, 6.7, and 7.3 live fetuses for d 25, 45, 65, 85, and 105, respectively (average SE = 0.46 fetuses). Corresponding means for OR were 13.4, 8.3, 7.9, 6.5, and 6.7 live fetuses, respectively (average SE = 0.44 fetuses). Means for UC were 10.2, 9.0, 8.5, 7.5, and 8.0 live fetuses, respectively (average SE = 0.47 fetuses). In each line, number of live fetuses at d 25 was approximately 72% of ovulation rate. Mortality to d 45 was greatest in OR, intermediate in CO, and least in UC. Reductions in live fetuses continued to occur from d 45 to 105, but line differences at d 45 were essentially maintained to d 105. Number of live fetuses in gilts at d 114 was estimated from each of the survival curves and predicted values of 7.0, 5.9, and 7.8 per uterine horn for CO, OR, and UC lines, respectively. Selection for uterine capacity improved fetal survival primarily during the time period between d 25 and 45. Relative growth rate coefficients throughout gestation for placental tissue indicated a change in rank of the line means, implicating a relative later growth pattern of placental tissue in the UC line.     

Correlated responses for litter traits to six generations of selection for ovulation rate or prenatal survival in French Large White pigs

Effects of selection for reproductive traits were estimated using data from 3 pig lines derived from the same Large White population base. Two lines were selected for 6 generations on high ovulation rate at puberty (OR line) or high prenatal survival corrected for ovulation rate in the first 2 parities (PS line). The third line was an unselected control line. Genetic parameters for age and BW at puberty (AP and WP); number of piglets born alive, weaned, and nurtured (NBA, NW, and NN, respectively); proportions of stillbirth (PSB) and survival from birth to weaning (PSW); litter and average piglet BW at birth (LWB and AWB), at 21 d (LW21 and AW21), and at weaning (LWW and AWW) were estimated using REML methodology. Heritability estimates were 0.38 +/- 0.03, 0.46 +/- 0.03, 0.16 +/- 0.01, 0.08 +/- 0.01, 0.09 +/- 0.01, 0.04 +/- 0.01, 0.04 +/- 0.02, 0.19 +/- 0.02, 0.10 +/- 0.02, 0.10 +/- 0.02, 0.36 +/- 0.02, 0.27 +/- 0.01, and 0.24 +/- 0.01 for AP, WP, NBA, PSB, NW, NN, PSW, LWB, LW21, LWW, AWB, AW21, and AWW, respectively. The measures of litter size showed strong genetic correlations (r(a) >/= 0.95) and had antagonistic relations with PSB (r(a) = -0.59 to -0.75) and average piglet BW (r(a) = -0.19 to -0.46). They also had strong positive genetic correlations with prenatal survival (r(a) = 0.67 to 0.78) and moderate ones with ovulation rate (r(a) = 0.36 to 0.42). Correlations of litter size with PSW were negative at birth but positive at weaning. The OR and PS lines were negatively related to PSW and average piglet BW. Puberty traits had positive genetic correlations with OR and negative ones with PS. Genetic trends were estimated by computing differences between OR or PS and control lines at each generation using least squares and mixed model methodologies. Average genetic trends were computed by regressing line differences on generation number. Significant (P < 0.05) average genetic trends were obtained in OR and PS lines for AP (respectively, 2.1 +/- 0.9 and 3.2 +/- 1.0 d/generation) and WP (respectively, 2.0 +/- 0.5 and 1.8 +/- 0.5 d/generation) and in the PS line for NBA (0.22 +/- 0.10 piglet/generation). Tendencies (P < 0.10) were also observed for LWB (0.21 +/- 0.12 kg/generation) and AWW (-0.25 +/- 0.14 kg/generation) in the PS line. Selection on components of litter size can be used to improve litter size at birth, but result in undesirable trends for preweaning survival.     

Correlated responses of pre- and postweaning growth and backfat thickness to six generations of selection for ovulation rate or prenatal survival in French Large White pigs

Correlated effects of selection for components of litter size on growth and backfat thickness were estimated using data from 3 pig lines derived from the same base population of Large White. Two lines were selected for 6 generations on either high ovulation rate at puberty (OR) or high prenatal survival corrected for ovulation rate in the first 2 parities (PS). The third line was an unselected control (C). Genetic parameters for individual piglet BW at birth (IWB); at 3 wk of age (IW3W); and at weaning (IWW); ADG from birth to weaning (ADGBW), from weaning to 10 wk of age (ADGPW), and from 25 to 90 kg of BW (ADGT); and age (AGET) and average backfat thickness (ABT) at 90 kg of BW were estimated using REML methodology applied to a multivariate animal model. In addition to fixed effects, the model included the common environment of birth litter, as well as direct and maternal additive genetic effects as random effects. Genetic trends were estimated by computing differences between OR or PS and C lines at each generation using both least squares (LS) and mixed model (MM) methodology. Average genetic trends for direct and maternal effects were computed by regressing line differences on generation number. Estimates of direct and maternal heritabilities were, respectively, 0.10, 0.12, 0.20, 0.24, and 0.41, and 0.17, 0.33, 0.32, 0.41, and 0.21 (SE = 0.03 to 0.04) for IWB, IW3W, IWW, ADGBW, and ADGPW. Genetic correlations between direct and maternal effects were moderately negative for IWB (-0.21 +/- 0.18), but larger for the 4 other traits (-0.59 to -0.74). Maternal effects were nonsignificant and were removed from the final analyses of ADGT, AGET, and ABT. Direct heritability estimates were 0.34, 0.46, and 0.21 (SE = 0.03 to 0.05) for ADGT, AGET, and ABT, respectively. Direct and maternal genetic correlations of OR with performance traits were nonsignificant, with the exception of maternal correlations with IWB (-0.28 +/- 0.13) and ADGPW (0.23 +/- 0.11) and direct correlation with AGET (-0.23 +/- 0.09). Prenatal survival also had low direct but moderate to strong maternal genetic correlations (-0.34 to -0.65) with performance traits. The only significant genetic trends were a negative maternal trend for IBW in the OR line and favorable direct trends for postweaning growth (ADGT and AGET) in both lines. Selection for components of litter size has limited effects on growth and backfat thickness, although it slightly reduces birth weight and improves postweaning growth.     

Early embryonic survival and embryo development in two lines of rabbits divergently selected for uterine capacity

The aim of this work is to study early embryo survival and development in 2 lines divergently selected for high and low uterine capacity throughout 10 generations. A total of 162 female rabbits from the high line and 133 from the low line were slaughtered at 25, 48, or 62 h of gestation. There were no differences in ovulation rate and fertilization rate between lines in any of the 3 stages of gestation. Embryo survival, estimated as the number of normal embryos recovered at a constant ovulation rate, was similar in both lines at 25 and 48 h. Embryo survival was greater in the high line [D (posterior mean of the difference between the high and low lines) = 0.57 embryos] at 62 h of gestation. There was no difference in embryonic stage of development at 25 h, but at 48 and 62 h of gestation, the high line, compared with the low line, had a greater percentage of early morulae (83 vs. 72%) and compacted morulae (55 vs. 38%). Divergent selection for uterine capacity appeared to modify embryo development, at least from 48 h of gestation, and embryo survival from 62 h.     

Divergent selection for immune responsiveness in chickens: estimation of realized heritability with an animal model.

With the aim of improving general disease resistance, chickens were divergently selected for their antibody titers 5 d after immunization with sheep red blood cells for nine generations. Selected and control lines differed significantly for primary and secondary responses after three generations. Heritability of the antibody titer was estimated by REML fitting an animal model using a derivative-free algorithm. The heritability estimate using data on all lines simultaneously was .31. Realized heritability of the antibody titer in the selected lines was estimated by using either the phenotypic cumulative response as the deviation from the control line or the mean breeding values obtained with an animal model. Values from the two methods were consistent, giving a realized heritability of .21 and .25 in the high and low lines, respectively. The genetic trend was not linear and the response to selection tended to accelerate over generations.     

Changes in fetal organ weights during gestation after selection for ovulation rate and uterine capacity in swine

We hypothesized that the ability of the fetus to alter nutrient shunting and organ growth might be associated with uterine capacity. White crossbred gilts from a randomly selected control line, a line selected for ovulation rate, and a line selected for uterine capacity (UC) were unilaterally hysterectomized-ovariectomized at 160 d of age, mated at estrus, and slaughtered at 45, 65, 85, and 105 d of gestation (9 to 18 gilts for each line x day combination). Analysis of the data revealed that heart weights and fetal weights were decreased in the ovulation rate line. No significant differences were obtained in fetal, placental, or fetal organ weights between the control and UC lines. Allometric growth of organs was assessed by examination of the slopes of the relationships between fetal weights and fetal organ weights after natural log transformation. Only the relative growth of the liver differed between selection lines and was greater (P = 0.01) in the UC compared with the control line during early pregnancy (d 45 and 65). Allometric growth of the fetal brain, liver, and heart differed with day of gestation. A brain-sparing effect was greater (P < 0.01) on d 85 and 105 compared with d 45 and 65. By contrast, a heart-sparing effect was present during early gestation and disappeared in later gestation. Fetal liver weights were hypersensitive to differences in fetal weights on d 45, possibly associated with placental effects on fetal liver weight. Fetal spleen weights were proportional to fetal weights throughout gestation. These results indicate that selection for ovulation rate decreased total fetal and fetal heart weights, and that selection for UC altered the relationship between total fetal and fetal liver weights during early gestation. Results further indicate significant changes in allometric growth of organs during gestation.     

Direct responses to selection for increased litter size, decreased age at puberty, or random selection following selection for ovulation rate in swine

Nine generations of selection for high ovulation rate were followed by two generations of random selection and then eight generations of selection for increased litter size at birth, decreased age at puberty, or continued random selection in the high ovulation rate line. A control line was maintained with random selection. Line means were regressed on generation number and on cumulative selection differentials to estimate responses to selection and realized heritabilities. Genetic parameters also were estimated by mixed-model procedures, and genetic trends were estimated with an animal model. Response to selection for ovulation rate was about 3.7 eggs. Response in litter size to selection for ovulation rate was .089 +/- .058 pigs per generation. Average differences between the high ovulation rate and control lines over generations 10 to 20 were 2.86 corpora lutea and .74 pigs (P less than .05). The regression estimate of total response to selection for litter size was 1.06 pigs per litter (P less than .01), and the realized heritability was .15 +/- .05. When the animal model was used, the estimate of response was .48 pigs per litter. Total response in litter size to selection for ovulation rate and then litter size was estimated to be 1.8 and 1.4 pigs by the two methods. Total response to selection for decreased age at puberty was estimated to be -15.7 d (P less than .01) when data were analyzed by regression (realized heritability of .25 +/- .05) and -17.1 d using the animal model. No changes in litter size occurred in the line selected for decreased age at puberty. Analyses by regression methods and mixed-model procedures gave similar estimates of responses and very similar estimates of heritabilities.     

Long-term responses, changes in genetic variances and inbreeding depression from 122 generations of selection on increased litter size in mice

Data on mice selected for litter size over 122 generations have been analysed in order to reveal the effect of long‐term selection on responses and changes in variances over a long selection period. Originally, three lines were established from the same base population, namely an H line selected for large litter size, an L line selected for small litter size and a K line without selection. In generation 122, the mean number of pups born alive (NBA) was 22 for the H line and 11 for the K line. Phenotypic response to selection is reduced over generations, but crossing of plateaued lines increased responses and realized heritabilities. Both realized heritabilities and heritabilities from residual maximal likelihood (REML) analyses were, in general, calculated from generation (?1)–44 (period 1), 45–70 (period 2) and 71–122 (period 3) separately. Realized heritabilities were in general smaller than heritabilities estimated from mixed model analysis. An overall estimate of heritability for NBA was found to be 0.19 (±0.01) by REML analysis. Additive variance is constant over all periods in the high line and the control line, but is reduced over periods in the low line. The reduction of additive variance in the low line could probably be explained by changes in gene frequencies. In all lines, environmental variances increased over periods. Inbreeding reduced the mean litter size by 0.72 (±0.10) pups per 10% increase in inbreeding, with substantial variance between periods and lines.     

Alternative methods of selection for litter size in mice: II. Response to thirteen generations of selection

Selection was conducted on an index of components of litter size (I = 1.21 x ovulation rate + 9.05 x ova success; ovulation rate measured by number of corpora lutea and ova success measured as number of pups born + number of corpora lutea), on uterine capacity (measured as number of pups born to unilaterally ovariectomized dams) and on litter size concurrent with an unselected control for 13 generations. Selection criteria (IX = index, UT = uterine capacity, LS = litter size and LC = control) were applied in each of three replicates. In an evaluation after five generations, IX and LS each exceeded LC by about .5 pups, with no response in UT. After 13 generations, mean ovulation rate, ova success and litter size (measured as number of fetuses at 17 d gestation in intact females) were, for IX, 14.25, .84, 11.95; for LS, 14.15, .82, 11.64; for UT, 12.61, .86, 10.77; and for LC, 12.27, .82, 9.98. The regression of number born (litter size in IX, LS and LC; uterine capacity with only a functional left uterine horn in UT) on cumulative selection differential across 13 generations was .12 +/- .01, .09 +/- .02 and .08 +/- .02 for IX, LS and UT, respectively. The regression of breeding value for litter size on each selection criterion, estimated as response in the generation-13 evaluation divided by cumulative selection differential, was .11 +/- .02, .08 +/- .01 and .05 +/- .03 for IX, LS and UT, respectively. Regression of response in number born on generation number was .17 +/- .01, .15 +/- .04 and .10 +/- .02 for IX, LS and UT, respectively. Selection in IX was promising relative to LS, and selection in UT changed number born.     

A genomewide association study in divergently selected lines in rabbits reveals novel genomic regions associated with litter size traits

Uterine capacity (UC), defined as the total number of kits from unilaterally ovariectomized does at birth, has a high genetic correlation with litter size. The aim of our research was to identify genomic regions associated with litter size traits through a genomewide association study using rabbits from a divergent selection experiment for UC. A high-density SNP array (200K) was used to genotype 181 does from a control population, high and low UC lines. Traits included total number born (TNB), number born alive (NBA), number born dead, ovulation rate (OR), implanted embryos (IE) and embryo, foetal and prenatal survivals at second parity. We implemented the Bayes B method and the associations were tested by Bayes factors and the percentage of genomic variance (GV) explained by windows. Different genomic regions associated with TNB, NBA, IE and OR were found. These regions explained 7.36%, 1.27%, 15.87% and 3.95% of GV, respectively. Two consecutive windows on chromosome 17 were associated with TNB, NBA and IE. This genomic region accounted for 6.32% of GV of TNB. In this region, we found the BMP4, PTDGR, PTGER2, STYX and CDKN3 candidate genes which presented functional annotations linked to some reproductive processes. Our findings suggest that a genomic region on chromosome 17 has an important effect on litter size traits. However, further analyses are needed to validate this region in other maternal rabbit lines.