Inbreeding and heterosis effects on quantitative traits in a White Leghorn population under long-term reciprocal recurrent selection

1. Inbreeding and heterosis effects for quantitative traits were estimated in two White Leghorn lines selected with reciprocal recurrent selection (RRS) since 1950. 2. Pedigreed progeny were produced from full-sib, half-sib and unrelated pure line as well as reciprocal cross line matings to estimate inbreeding and heterosis effects "within sires". 3. Inbreeding effects estimated as linear regressions agreed well with estimates considering inbreeding as fixed effect. The inbreeding depression per 10% inbreeding was highest for hatchability (6-7%), intermediate for production-related traits (3%), and about 1% for shell quality, body weight and egg weight. 4. Heterosis effects were estimated by comparing pure line and cross line progeny of the same sires. 5. Results indicate considerably less heterosis for egg production than found earlier, which is interpreted as a consequence of pure line selection.     

Genetic and phenotypic parameters for monthly egg production in White Leghorn hens

This article reports genetic and phenotypic parameters of monthly egg production and the influence of Box‐Cox transformation on the parameters from a population of White Leghorns, selected for feed efficiency. A total of 6450 daughters of 180 sires and 1335 dams were analysed by restricted maximum likelihood (REML) using a multivariate animal model. The traits considered were monthly egg productions, cumulative production of the first 5 months (S5), cumulative production of first 10 months (S10), and survivor egg production in the first cycle (S12). Two sets of data were analysed: the original data and with the Box‐Cox method transformed data. The results indicated that there were no great differences in the estimates between untransformed and transformed data. The estimates of heritability for monthly egg production were high for the first period, decreased to reach the lowest during peak production, and increased to the end of lay. The estimates of heritability for cumulative records were generally higher than monthly records. Genetic and phenotypic correlations among monthly egg production totals were generally high for contiguous periods and then decreased as the interval between months increased. The highest genetic correlation between monthly records and S5 was for the second month of production, whereas the correlations between monthly production totals and S10 and S12 reached their peak at the sixth and eighth months of production, respectively.     

Implications of genetic selection on yolk proportion on the dry matter content of eggs in a White Leghorn population

1. The responses to genetic selection on yolk proportion as a technique for increasing egg dry matter content, an important criterion for the egg-product industry, was investigated in a pedigree flock of White Leghorn hens.

2. Parents were preselected on high and low yolk proportion from a base population. The absolute estimated breeding value for yolk proportion of both groups differed by 3%. The realised selection difference in dry matter content of eggs between groups was more than 1% in the analysed offspring population.

3. Heritability estimates were moderate and dry matter had a lower heritability (h² = 0.39) than yolk proportion (h² = 0.44).

4. The genetic correlation between yolk proportion and dry matter content was highly positive (r_g = 0.91). Genetic correlations with egg weight were negative and would have to be compensated for in a breeding programme (r_g = ?0.76 with yolk proportion and r_g = ?0.64 with dry matter content). The genetic correlation between the laying performance and yolk proportion was r_g = 0.28 and close to zero (r_g = ?0.05) for dry matter content.

5. Easy recording and lower undesirable correlations make yolk proportion more suitable for commercial selection compared with egg dry matter content in layer breeding.     

Genetic study of embryonic mortality in White Leghorn lines selected for egg production traits

The present study compares embryonic mortality between lines selected for different production traits, assesses the effects of inbreeding of the hen and embryo on embryonic mortality, and estimates genetic parameters of embryonic mortality. The experiment covered 10 generations of selection for increased egg number (EN), egg weight (EW), egg mass (EM) and a control line (C). The data included age at 1st egg, egg number and egg weight. Percent fertile eggs (PF), percent hatched of fertile eggs (PHF) and percent dead chick at hatch (PDH) were also recorded for the selected parents. PDH was higher in the selected lines than in the control line. Among the selected lines, the EW line had the highest embryonic mortality. Inbreeding of the hen and embryo had no significant effect on PDH in any of the lines. Estimates of heritability for PDH were 0.10+/-0.05, 0.02+/-0.02, 0.03+/-0.02 and 0.02+/-0.02 for lines EN, EW, EM and C, respectively. There was a positive genetic correlation between egg weight and PDH in line EW indicating that selection for increased egg weight was associated with high embryonic mortality. A negative genetic correlation between PDH and reproductive traits in line EN was observed, which is favourable.     

One-generation divergent selection on large and small yolk proportions in a White Leghorn line

1. A striking change in egg consumption from shell eggs toward egg products is taking place. Consequently, internal egg quality traits must be seen in a different light, as they may affect the profitability of the egg processing industry. In order to study the possibility of using breeding as a means to alter yolk production, a one-generation divergent selection experiment was set up. The trait was yolk proportion. The effect of hen's age on egg composition was also studied. Genetic parameters for yolk proportion, yolk weight, egg weight and albumen weight were estimated, using a multiple trait animal model, and restricted maximum likelihood. 2. Selection response was obtained for yolk proportion and correlated selection responses for yolk weight and albumen weight. In the 'high line' a decrease in egg weight and an increase in yolk weight was established, whereas in the 'low line', egg weight remained more or less the same and yolk weight decreased slightly 3. Yolk proportion, yolk weight, egg weight and albumen weight all increased with increasing age of the hen in both selection lines. The difference between selection lines in yolk proportion, yolk weight and albumen weight persisted throughout the laying period. 4. Estimated heritabilities for yolk proportion, yolk weight, egg weight and albumen weight were 0.38, 0.22, 0.34 and 0.41 respectively. The genetic correlation between yolk proportion and yolk weight was 0.52, whereas the genetic correlations with egg weight and albumen weight were -0.51 and -0.74 respectively. 5. We conclude that selection can be used to alter yolk production. Selection can be at an early age with a sustained effect throughout the laying period.     

Single-step divergent selection for male comb shape in two White Leghorn lines.

1. Divergent selection for comb shape (SH, the way that the cockerel bears the comb) was performed in 2 White Leghorn lines in a study aimed at assessing possibilities for improving SH. 2. Line A, selected for large comb size (CS) at 29 weeks of age, had great SH problems whereas Line H, selected for high hyaluronic acid concentration in the comb (HA), had minor SH problems. Multivariate analyses were used to estimate genetic parameters for SH, CS and HA in lines A and H and in a control line (C). 3. Significant direct selection responses for SH were achieved in both lines. There were significant unfavourable correlated responses for CS in both lines. 4. The correlated responses for HA were not significant and were unfavourable in Line A and favourable in Line H. 5. Heritabilities for SH and CS were high in both lines and relatively low for HA. Most of the genetic correlations were in agreement with the correlated responses obtained.     

Estimation of direct and correlated responses to selection using univariate animal models.

Analytic results obtained using simple models show that estimates of selection response of univariate experiments using animal models are completely dependent on the heritability used as prior when fixed effects are nested within generations, and both on the prior and on the true heritability parameter when fixed effects overlap across generations. Univariate animal model estimators of correlated changes of a trait not selected directly are usually biased. The absolute value of the estimate of the correlated response is smaller than the true value when the traits are only genetically correlated and larger than the expected value of zero when they are only environmentally correlated. The validity of the results derived from the analysis of simple models is confirmed using computer simulations, which illustrate the magnitude of the bias. It is emphasized that use of univariate animal models to estimate response in breeding programs whose breeding objectives include several correlated traits may lead to erroneous conclusions.     

Genetic basis of the increase in egg weight with pullet age in a White Leghorn flock

1. In a closed White Leghorn flock over a 6-year period the average increase in egg weight from the start of lay to 37 weeks of age was 12.6 g. 2. The heritability of this egg weight increase was estimated at 0.36 +/- 0.06. 3. The genetic correlations of egg weight increase with other characteristics were: age at first egg -0.50, early egg weight -0.51, 37 week egg weight 0.42 and egg production to 273 d 0.44. 4. It is predicted that with unit selection intensity the direct response in egg weight increase will be -1.30 g, with correlated responses in age at first egg of 2.1 d, early egg weight 0.64 g, 37 week egg weight -0.51 g and egg production to 273 d -1.83 eggs.     

Within- and between-strain genetic variability in White Leghorn population detected through RAPD markers

1. Five White Leghorn (WLH) strains, namely G, H, I, J and C, differing in selection history, were screened for randomly amplified polymorphic DNA (RAPD) polymorphism using 50 random decamer primers. 2. Only 12 primers detected polymorphism between the strains. Out of 96 bands scored using these primers, only 21 (21.9%) were found to be polymorphic. 3. Strains differed considerably for within-population genetic similarity, estimated either by band sharing (WS=0.836 to 0.968) or band frequency (WF=0.820 to 0.969). 4. Between-strain genetic similarity estimates, based on band sharing (BS) as well as on band frequency (BF) ranged from 0.756 to 0.958 and from 0.830 to 0.996, respectively. Strains C and H were most closely related to each other, while J appeared to be more distant from other strains. 5. The between-strain genetic distance estimates also revealed similar trends. 6. A phylogenetic tree, derived using Nei's coefficient of similarity revealed similar relationships between the WLH strains.     

Sarcomas and myelocytomas induced by a retrovirus related to myeloblastosis-associated virus type 1 in White Leghorn egg layer chickens

An outbreak of subcutaneous sarcomas in commercial White Leghorn egg layers was observed in the northeastern United States during late 2004. Subcutaneous tumors were confined to three flocks distributed in two locations and belonging to the same company. The tumors were first observed grossly by farm personnel at approximately 7 wk of age and persisted throughout the economic life of the flocks. Most of the tumors observed during the growing period were present on the facial region or around the head, wings, and legs. There was no gross evidence of bursal or visceral involvement. Microscopically, most tumors were undifferentiated sarcomas and myxomas. There was no microscopic evidence of Marek's disease or lymphoid leukosis. Reticuloendotheliosis virus proviral DNA was not detected by polymerase chain reaction either in tumors or in cell cultures. Egg production and mortality rates were within normal limits in the affected flocks and many of the chickens exhibiting tumors seemed healthy otherwise, albeit approximately one-half of the daily mortality exhibited tumors. Avian myeloblastosis-associated virus type 1 (MAV-1) was isolated from tumors, plasma, and serum. Upon initial virus neutralization, the viruses isolated seemed at least partially related antigenically to avian leukosis virus (ALV) subgroups A and B but not to subgroup J (ALV-J). Sequencing of the variable and hypervariable regions of gp85 in the envelope gene revealed that the viruses involved are closely related to MAV-1. Attempts to reproduce subcutaneous sarcomas with MAV-1 isolated from White Leghorn chickens in the case herein reported produced exclusively myelocytomas indistinguishable histologically from those induced by ALV-J in meat type chickens.     

Direct and correlated responses to two-stage selection for ovulation rate and number of fully formed pigs at birth in swine

Our objectives were to estimate responses and genetic parameters for ovulation rate, number of fully formed pigs at birth, and other production traits following two-stage selection for increased ovulation rate and number of fully formed pigs. Eight generations of selection were practiced in each of two lines. One selection line was derived from a line that previously selected eight generations for an index to increase ovulation rate and embryonic survival (the IOL pigs). The other selection line was derived from the unselected control line of the index selection experiment (the COL pigs). The control line (C) was continued with random selection. Due to previous selection, Line IOL had greater ovulation rate (4.24 +/- 0.38 and 4.14 +/- 0.29 ova) and litter size (1.97 +/- 0.39 and 1.06 +/- 0.38 pigs) at Generation 0 of two-stage selection than did Lines COL and C. In Stage 1, all gilts from 50% of the largest litters were retained. Approximately 50% of them were selected for ovulation rate in Stage 2. Gilts selected for ovulation rate were mated to boars selected from the upper one-third of the litters for litter size. At Generations 7 and 8, differences in mean EBV for ovulation rate and litter size between Lines IOL and C were 6.20 +/- 0.29 ova and 4.66 +/- 0.38 pigs; differences between Lines COL and C were 2.26 +/- 0.29 ova and 2.79 +/- 0.39 pigs; and differences between Lines IOL and COL were 3.94 +/- 0.26 ova and 1.86 +/- 0.39 pigs. Regressions of line mean EBV on generation number were 0.27 +/- 0.07 ova and 0.35 +/- 0.06 pigs in Line IOL; 0.30 +/- 0.06 ova and 0.29 +/- 0.05 pigs in Line COL; and 0.01 +/- 0.07 ova and 0.02 +/- 0.05 pigs in Line C. Correlated responses were decreased age at puberty and increased number of pigs born alive, number of mummified pigs, prenatal loss, and individual and litter birth weight. Two-stage selection for ovulation rate and number of pigs per litter is a promising procedure to improve litter size in swine.     

Direct and correlated responses to selection for hatching time in the fowl

Selection over eight generations for a shorter (fast hatching) or longer (slow‐hatching) incubation time in fowls resulted in a difference of 42 h in incubation time. Only minor asymmetry of response was observed since incubation time decreased by 19.8 h in the fast‐hatching line and increased by 22.4 h in the slow‐hatching line. The predicted response was less than that observed, probably because of underestimation of the true selection differentials and a low estimate of heritability used for prediction.

The realised heritability estimates decreased significantly from 0.222 to 0.179 between the first four and the last four generations. The estimate based on all eight generations was 0.199. These values are probably slightly overestimated because of the use of underestimated selection differentials in their calculation. The realised heritability in the original stock was probably between 0.20 and 0.25.

No evidence of correlations between hatchability or egg weight and hatching time was obtained. A significantly lower egg weight in the fast‐hatching line could be accounted for on the basis of random genetic drift.     

Inbreeding and inbreeding depression on reproduction and production traits of White Leghorn lines selected for egg production traits.

1. The rate of inbreeding and its effect on reproduction and egg production traits were studied in White Leghorn lines selected for egg production traits. The experiment was carried out for 10 generations in a control line (C) and in lines selected for increased egg number (EN), egg weight (EW) and egg mass (EM). 2. Data were available on reproduction traits, such as percent fertile eggs (PF), percent hatched of fertile eggs (PHF) and percent hatched of total eggs set (PHT), and on egg production traits such as age at 1st egg (AFE), egg number and egg weight. 3. The rate of increase in average inbreeding per generation was 1.50, 1.24, 1.14 and 0.18% for the line EN, EW, EM and C, respectively. The effect of inbreeding on reproduction and production traits was estimated by including the inbreeding coefficient of the hen (Fh), embryo (Fe) and mate (Fs) as a partial linear regression in the model. 4. There was a significant effect of inbreeding on reproduction traits in line EW attributable to the inbreeding of the hen, embryo and mate. No such effect was observed in the other lines. 5. In all lines inbreeding tended to reduce egg number and delay sexual maturity. In general, all lines reacted differently to inbreeding.     

Renewed selection for heat loss in mice: direct responses and correlated responses in feed intake, body weight, litter size, and conception rate

Divergent selection in mice was renewed in 3 independent replicates for high (MH) and low (ML) heat loss. An unselected control (MC) was maintained in all replicates. Heat loss was measured for individual male mice for 15 h, overnight in direct calorimeters. After 16 initial generations of selection followed by 26 generations of relaxed selection, divergent selection resumed for 9 generations. The realized selection applied was very close to the maximum possible selection according to the criteria and protocol. Selection differentials were greater for high than for low selection due to greater variation in the MH line. When corrected for SD, standardized selection differentials were similar for MH and ML selection. Unintended selection in MC was negligible. Realized heritability for divergence was 0.14 +/- 0.01, which was considerably less than that realized during the initial generations of selection (0.28 +/- 0.03). Realized heritabilities for MH selection (0.16 +/- 0.05) and for ML selection (0.07 +/- 0.06) were less, especially for ML selection, than were observed in the earlier generations. The difference in heat loss between MH and ML males was 55.7% of the MC mean at generation 51, compared with a difference of 53.6% in generation 15; this difference had decreased to 34.4% at the end of the relaxed selection (generation 42). For feed intake between 8 and 11 wk, MH and ML males differed by 34.0% of the MC mean by the end of the selection process. Body weight at 12 wk for MH and ML males was less than for MC males. Litter size response was positively related to the heat loss response. Conception rate was poorer in MH matings than in MC and ML matings.     

Relationship between delayed onset of egg production and involution of the bursa of Fabricius in White Leghorn chickens

The relationship between delayed onset of oviposition and involution of the bursa of Fabricius is described. Chickens studied were 26 weeks of age and had not come into production. The bursae of nonlaying chickens were compared with those of age-matched chickens that had laid the first egg 1, 9 to 11, or 24 days before. The bursae from the nonlaying chickens were similar in weight and morphology to the functional bursae of immature pullets. Bursae of laying chickens, even on the day of the first egg, were markedly involuted. Age of the chicken was not a factor in determining bursal involution. The onset of egg production was directly related to bursal involution.     

Direct and correlated responses to selection for efficiency of lean gain in mice

Improvement in feed efficiency when selection is based on gain:feed ratio has often been accompanied by a reduction in feed intake. The following four criteria were used in mass selection for improved lean gain efficiency in mice with an objective of evaluating changes in lean gain and intake: 1) gain deviation, animals selected had the greatest gain in fat-free mass (FFM) after adjustment to a constant intake; 2) intake deviation, mice selected had the least feed intake after adjustment to a constant gain in FFM; 3) intrinsic efficiency, similar to the second criterion except that adjustment was also made for average weight maintained during the period; and 4) a positive control that used the ratio of gain in FFM: feed intake as the selection criterion. A fifth line, in which a male and a female were selected at random from each litter, served as a negative control. Experimental animals were outbred mice of the CF1 strain. Two replicates of the five lines were included in the study. Twelve males and females were pair-mated within each line-replicate combination each generation. Feed disappearance was measured from 25 to 42 d. Mice were scanned to obtain an electrical conductivity measurement for prediction of FFM. After six generations of selection, realized heritabilities for gain:feed, gain deviation, intake deviation, and intrinsic efficiency were .00 +/- .04, .04 +/- .29, .35 +/- .08, and .28 +/- .06, respectively. There were no differences among lines for gain:feed ratio. The correlated response in feed intake reduction was significant in the intake deviation and intrinsic efficiency lines (-.17 +/- .05 and -.21 +/- .04 g x d(-1) x generation(-1), respectively). The realized genetic correlations between the ratio and gain deviation, intake deviation, and intrinsic efficiency were .83 +/- .15, .01 +/- .04, and .21 +/- .12, respectively. Litter size was depressed in all selected lines.