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Use of computerized mate selection programs to control inbreeding of Holstein and Jersey cattle in the next generation.

The expected role of computerized mate selection programs with regard to inbreeding and lifetime profitability of Holstein and Jersey cattle was examined using data from 25 large registered herds of each breed. Sire selection and mate allocation were carried out using linear programming with the following objectives: 1) minimum inbreeding, 2) maximum net merit subject to a fixed inbreeding threshold, and 3) maximum expected lifetime profit after adjustment for inbreeding depression. Inbreeding of actual matings was similar to inbreeding from random matings, indicating that current inbreeding avoidance programs in these herds are ineffective. Inbreeding was reduced by 1.6 and 1.9% in Holsteins and Jerseys, respectively, when a mate allocation program was applied with service sires and usage levels fixed at the actual values. Benefits of mate selection programs increased when both sire selection and mate pair allocation were considered. Maximization of mean net merit with inbreeding restricted to a fixed level (5% in Holsteins and 8% in Jerseys) led to decreases in inbreeding of 0.9 and 1.4% and increases in lifetime profit of $16.66 and $26.86 in Holsteins and Jerseys, respectively, relative to programs that ignored inbreeding. Maximization of mean expected lifetime profit after adjustment for inbreeding depression decreased inbreeding by 1.8 and 2.8% and increased lifetime profit by $37.37 and $59.77 in Holsteins and Jerseys, respectively. Inbreeding coefficients estimated with pedigree traced to 1985 were inadequate predictors of inbreeding coefficients estimated with pedigrees traced to 1960. Mate selection programs cannot function optimally unless extensive historical pedigree data are available, particularly for service sires. Computerized mate selection programs can reduce inbreeding in the next generation, which will lead to an increase in farm profitability. However, if genetic diversity is to be maintained in the long term, procedures for selecting parents of AI sires must also be considered.

Algorithms↗

Inbreeding depression and mixed mating in Leptosiphon jepsonii: a comparison of three populations.

BACKGROUND AND AIMS: Inbreeding depression is thought to play a central role in the evolution and maintenance of cross-fertilization. Theory indicates that inbreeding depression can be purged with self-fertilization, resulting in positive feedback for the selection of selfing. Variation among populations of Leptosiphon jepsonii in the timing and rate of self-fertilization provides an opportunity to study the evolution of inbreeding depression and mating systems. In addition, the hypothesis that differences in inbreeding depression for male and female fitness can stabilize mixed mating in L. jepsonii is tested. METHODS: In a growth room experiment, inbreeding depression was measured in three populations with mean outcrossing rates ranging from 0.06 to 0.69. The performance of selfed and outcrossed progeny is compared at five life history stages. To distinguish between self-incompatibility and early inbreeding depression, aborted seeds and unfertilized ovules were counted in selfed and outcrossed fruits. In one population, pollen and ovule production was quantified to estimate inbreeding depression for male and female fitness. KEY RESULTS: Both prezygotic barriers and inbreeding depression limited self seed set in the most outcrossing population. Cumulative inbreeding depression ranged from 0.297 to 0.501, with the lowest value found in the most selfing population. Significant inbreeding depression for early life stages was found only in the more outcrossing populations. Inbreeding depression was not significant for pollen or ovule production. CONCLUSIONS: The results provide modest support for the hypothesized relationship between inbreeding depression and mating systems. The absence of early inbreeding depression in the more selfing populations is consistent with theory on purging. Differences in male and female expression of inbreeding depression do not appear to stabilize mixed mating in L. jepsonii. The current estimates of inbreeding depression for L. jepsonii differ from those of previous studies, underscoring the effects of environmental variation on its expression.

Biomass↗

Environmental conditions affect the magnitude of inbreeding depression in survival of Darwin's finches.

Understanding the fitness consequences of inbreeding (inbreeding depression) is of importance to evolutionary and conservation biology. There is ample evidence for inbreeding depression in captivity, and data from wild populations are accumulating. However, we still lack a good quantitative understanding of inbreeding depression and what influences its magnitude in natural populations. Specifically, the relationship between the magnitude of inbreeding depression and environmental severity is unclear. We quantified inbreeding depression in survival and reproduction in populations of cactus finches (Geospiza scandens) and medium ground finches (Geospiza fortis) living on Isla Daphne Major in the Galápagos Archipelago. Our analyses showed that inbreeding strongly reduced the recruitment probability (probability of breeding given that an adult is alive) in both species. Additionally, in G. scandens, first-year survival of an offspring with f = 0.25 was reduced by 21% and adults with f = 0.25 experienced a 45% reduction in their annual probability of survival. The magnitude of inbreeding depression in both adult and juvenile survival of this species was strongly modified by two environmental conditions, food availability and number of competitors. In juveniles, inbreeding depression was only present in years with low food availability, and in adults inbreeding depression was five times more severe in years with low food availability and large population sizes. The combination of relatively severe inbreeding depression in survival and the reduced recruitment probability led to the fact that very few inbred G. scandens ever succeeded in breeding. Other than recruitment probability, no other trait showed evidence of inbreeding depression in G. fortis, probably for two reasons: a relatively high rate of extrapair paternity (20%), which may lead to an underestimate of the apparent inbreeding depression, and low sample sizes of highly inbred G. fortis, which leads to low statistical power. Using data from juvenile survival, we estimated the number of lethal equivalents carried by G. scandens, G. fortis, and another congener, G. magnirostris. These results suggest that substantial inbreeding depression can exist in insular populations of birds, and that the magnitude of the inbreeding depression is a function of environmental conditions.

Animals↗

Inbreeding depression for global and partial economic indexes, production, type, and functional traits.

The objective of this research was to examine the effects of inbreeding in the population of Holstein cattle in the Walloon region of Belgium. The effects of inbreeding on the global economic index and its components were studied by using data from the genetic evaluations of February 2004 for production, somatic cell score (SCS), computed from somatic cell counts and type. Inbreeding coefficients for 956,516 animals were computed using a method that allows assigning an inbreeding coefficient to individuals without known parents. These coefficients were equal to the mean inbreeding coefficient of contemporary individuals with known parents. The significance of inbreeding effects on the different evaluated traits and on the different indexes were tested using a t-test comparing estimated standard errors and effects. The inbreeding effect was significantly different from zero for the vast majority of evaluated traits and for all of the indexes. Inbreeding had the greatest deleterious effects on production traits. Inbreeding decreased yield of milk, fat, and protein during a lactation by 19.68, 0.96, and 0.69 kg, respectively, per each 1% increase in inbreeding. The regression coefficient of SCS per 1% increase in inbreeding was +0.005 SCS units. The inbreeding depression was thus relatively low for SCS, but inbred animals had higher SCS than non-inbred animals, indicating that inbred animals would be slightly more sensitive to mastitis than non-inbred animals. Estimates of inbreeding effects on evaluated type traits per 1% increase were small. The most strongly affected type traits were chest width, rear leg, and overall development on a standardized scale. For several type traits, particularly traits linked to the udder, the estimates suggested a favorable effect of inbreeding. The global economic index was depressed by around 6.13 euro of lifetime profit per 1% increase in inbreeding for the Holstein animals in the Walloon region of Belgium.

Animals↗

Effects of inbreeding on milk and fat production, stayability, and calving interval of registered Ayrshire cattle in the northeastern United States.

Inbreeding coefficients of 30,794 registered Ayrshire cows were calculated from relationships between sire and maternal male ancestors. Average inbreeding coefficient of all cows was less than 1% and of the 5,202 inbred cows was 5.4%. Fewer than 2% of inbred cows had coefficients greater than 15%. Percent of inbred cows increased from 23% of cows born in 1972 to 43% in 1980. Over the same period average inbreeding coefficient of all cows increased from 1.2 to 2.0%, but average coefficient of inbred cows decreased from 5.6 to 4.7%. Effects of inbreeding on first lactation, 305-day, 2X, mature equivalent milk and fat production (kg), 48-mo stayability (proportion of cows surviving to 48 mo of age), and first calving interval (days) were estimated by a model that included fixed effects for herd-year-seasons, sire-maternal grandsire groups, inbreeding, and random effects for sires and maternal grandsires within groups. Inbreeding was included in the model as a classification (six classes according to inbreeding coefficient: 0, 0+ to 5-%, 5 to 10-%, 10 to 15-%, 15 to 25-%, and 25 to 35-% and one class for cows with indeterminate inbreeding). Estimates of differences between inbreeding classes 0+ to 5-% through 25 to 35-% and the zero inbreeding class indicated that milk and fat production decreased with increased inbreeding. Effects of inbreeding on stayability and calving interval were small. Inbreeding was fitted also as a linear covariate. Regressions of milk, fat, stayability, and calving interval on inbreeding coefficients were -23, -1, -.008, and -.095 per 1% increase of inbreeding coefficient.

Animals↗

Inbreeding avoidance in an isolated indigenous Zapotec community in the valley of Oaxaca, southern Mexico.

We analyzed inbreeding using surname isonymy in an indigenous genetic isolate. The subjects were residents of a rural Zapotec-speaking community in the valley of Oaxaca, southern Mexico. The community can be classified as a genetic isolate with an average gene flow of < or = 3% per generation. Surnames were collected for individuals in each household in pedigree form using the culturally traditional patronym-matronym naming. Estimation of inbreeding from surname isonymy is facilitated by the traditional patronym-matronym name assignment among indigenous Mexican populations. A total of 2,149 individuals had valid surname patronym-matronym pairings, including 484 deceased ancestors. Surname isonymy analysis methods were used to estimate total inbreeding and to segregate it into random and nonrandom components. The surname isonymy coefficient computed from 119 isonymous surname pairings (119/2,149) was 0.0554. The estimated inbreeding coefficient from surname isonymy was 0.0138 (0.0554/4). The random and nonrandom components of inbreeding were F(r) = 0.0221 and F(n) = -0.0091, respectively. The results suggest that consanguinity is culturally avoided. Nonrandom inbreeding decreased total inbreeding by about 41%. Total estimated inbreeding by surname isonymy was 0.0138, which is similar to inbreeding estimated from a sample of pedigrees, 0.01. Socially prescribed inbreeding avoidance substantially lowered total F through negative nonrandom inbreeding. Even in the situation of genetic isolation and small effective population size (N(e)), estimated inbreeding is lower than may have otherwise occurred if inbreeding were only random. However, among the poorest individuals, socially prescribed jural rules for inbreeding avoidance failed to operate. Thus the preponderance of inbreeding appears to occur among the poor, economically disadvantaged in the community.

Choice Behavior↗

Effects of inbreeding on reproduction and wool production of Rambouillet, Targhee, and Columbia ewes.

Because of a traditional interest in inbreeding as a tool for breed improvement, it was desirable to evaluate the effects of inbreeding on the most important products of the sheep industry, lamb and wool production. The data for this study were based on 13,807 ewe and 16,470 lamb records from Rambouillet (R), Targhee (T), and Columbia (C) sheep collected over 9 or 10 yr from 54 inbred lines. Average inbreeding of the lambs and ewes involved was near 25 and 20%, respectively. The combined effects of lamb's and dam's inbreeding reduced litter weight weaned at 120 d approximately .5 kg for each percentage of increase in inbreeding in every breed. For ewes and lambs of average inbreeding, the reductions relative to noninbred ewes amounted to 12.4, 10.3, and 11.3 kg for R, T, and C, respectively. The reductions constituted declines in weight weaned per ewe of 34, 25, and 28%. Similarly, net reproduction rate (lambs weaned per ewes put into breeding) was reduced more than one percentage point for each percentage of increase in inbreeding, with net declines at average inbreeding of 31.7, 23.5, and 25.7 percentage points for each breed, respectively. These reductions were equivalent to reductions in net rate of 29, 20, and 23%. The combined effect of lamb and dam inbreeding reduced weaning weight by 3.5, 2.6, and 2.2 kg, which constituted reductions of 10, 7, and 6%, respectively. Reduced weaning weight was approximately 30% as important as reduced net reproduction rate in contributing to the decline in litter weight weaned. Effects on fleece weight were curvilinear and amounted to reductions of .35, .18, and .00 kg, respectively, at levels of average inbreeding; however, Columbia fleece weights declined rapidly at levels exceeding 20% for ewes. Potential selection differentials were reduced 16% at inbreeding levels of 25 and 20% for lambs and dams, respectively, and 62% at levels of 55 and 50%. The average economic loss per ewe in value of production was estimated at $17 for average inbreeding and as high as $36 for inbreeding approaching 50%. It seems that the use of inbreeding as a tool for improving productive merit in sheep is much more certain to be a detriment, economically and genetically, than an advantage.

Animals↗

Changes in inbreeding of U.S. Herefords during the twentieth century.

Genetic diversity in the U.S. Hereford population was characterized by examining the level and rate of inbreeding and effective population size. Pedigree records for 20,624,418 animals were obtained from the American Hereford Association, of which 96.1% had both parents identified. Inbreeding coefficients were computed and mean inbreeding (Fx) calculated by year from 1900 to 2001. Inbreeding increased rapidly between 1900 and 1945. From 1946, inbreeding increased linearly to a maximum of 11.5% in 1966. Throughout the 1970s and 1980s, mean inbreeding decreased to mid-century levels. Several alternatives were investigated to explain this decline. The average relationship between prominent sires fell from 20 to 12% during the time that the level of inbreeding decreased, which reflects an increase in the popularity of certain less fashionable sire lines that would have temporarily decreased inbreeding. Pedigrees were constructed for animals born after 1990. This subsample of animals with no missing ancestors in at least 12 generations did not exhibit a decrease in inbreeding. Missing ancestral information therefore contributed to the apparent decline. One cause of missing ancestry results from outcrossing to imported animals. The effect of missing ancestry was investigated by simulating the missing ancestors. In 2001, Fx was 9.8%, and approximately 95% of individuals were inbred. The maximal inbreeding coefficient was 76%. The annual change in mean inbreeding (DeltaFx) was estimated for Herefords born during five time periods from 1946 to 2001, where inbreeding was changing at different linear rates. The DeltaFx for the most recent generation (1990 to 2001) was 0.12%/yr. Assuming a generation interval of 4.88 yr, the estimated effective population size was 85. This study provides a benchmark of current genetic diversity in the Hereford population. Results indicate that inbreeding is accumulating linearly and below critical levels. Increases in the adoption of reproductive technologies could decrease genetic diversity, and in the future, we may need to consider strategies to minimize inbreeding.

Animal Husbandry↗

Maternal and fetal inbreeding depression for 70-day nonreturn and calving rate in Holsteins and Jerseys.

Inbreeding depression for 70-d nonreturn rate was estimated in 50,613 Holstein and 47,673 Jersey cows with five-generation pedigrees using an animal model. Heritabilities of 70-d nonreturn rate were very low for both breeds (1 to 2%). Maternal inbreeding depression was small (3% reduction for 10% inbreeding) and significant only for Jerseys. Fetal and maternal inbreeding depression was not significant for individual parities in Holsteins, but maternal inbreeding depression was significant in first parity only in Jerseys. Maternal and fetal inbreeding depression of calving rate (verified by a subsequent calving) was estimated on separate datasets by parity from 13,229 to 26,876 Holstein and 7374 to 11,742 Jersey cows. First-parity estimates for heritability of calving rate were 1% or less, whereas estimates for later parities varied from 1 to 6%. Significant inbreeding depression in first-parity Holsteins reduced calving rate by 4% per 10% maternal or fetal inbreeding, but effects, while undesirable, were not consistently significant in other parities. In Jerseys, maternal inbreeding significantly reduced calving rate by 6% per 10% inbreeding in first parity, and was undesirable but not significant for second through fourth parities. Fetal inbreeding depression was not significant in Jerseys. Maternal inbreeding depression of 70-d nonreturn and calving rate was small, undesirable, but not consistently significant across breeds and parities. The cumulative economic impact of maternal or fetal inbreeding on lifetime reproductive performance of Holstein or Jersey cows would be more dramatic than results for a single breeding.

Animals↗

The transition to social inbred mating systems in spiders: role of inbreeding tolerance in a subsocial predecessor.

The social spiders are unusual among cooperatively breeding animals in being highly inbred. In contrast, most other social organisms are outbred owing to inbreeding avoidance mechanisms. The social spiders appear to originate from solitary subsocial ancestors, implying a transition from outbreeding to inbreeding mating systems. Such a transition may be constrained by inbreeding avoidance tactics or fitness loss due to inbreeding depression. We examined whether the mating system of a subsocial spider, in a genus with three social congeners, is likely to facilitate or hinder the transition to inbreeding social systems. Populations of subsocial Stegodyphus lineatus are substructured and spiders occur in patches, which may consist of kin groups. We investigated whether male mating dispersal prevents matings within kin groups in natural populations. Approximately half of the marked males that were recovered made short moves (< 5m) and mated within their natal patch. This potential for inbreeding was counterbalanced by a relatively high proportion of immigrant males. In mating experiments, we tested whether inbreeding actually results in lower offspring fitness. Two levels of inbreeding were tested: full sibling versus non-sib matings and matings of individuals within and between naturally occurring patches of spiders. Neither full siblings nor patch mates were discriminated against as mates. Sibling matings had no effect on direct fitness traits such as fecundity, hatching success, time to hatching and survival of the offspring, but negatively affected offspring growth rates and adult body size of both males and females. Neither direct nor indirect fitness measures differed significantly between within patch and between-patch pairs. We tested the relatedness between patch mates and nonpatch mates using DNA fingerprinting (TE-AFLP). Kinship explained 30% of the genetic variation among patches, confirming that patches are often composed of kin. Overall, we found limited male dispersal, lack of kin discrimination, and tolerance to low levels of inbreeding. These results suggest a history of inbreeding which may reduce the frequency of deleterious recessive alleles in the population and promote the evolution of inbreeding tolerance. It is likely that the lack of inbreeding avoidance in subsocial predecessors has facilitated the transition to regular inbreeding social systems.

Animals↗

The influence of variable rates of inbreeding on fitness, environmental responsiveness, and evolutionary potential.

We manipulated experimental populations of the housefly (Musca domestica L.) under three inbreeding schemes (fast, slow, and punctuated) to partition out the influences of different means and variances in the rate of inbreeding, per generation, while controlling for the final level of inbreeding as a constant. One treatment used constant fast inbreeding (11% per generation; Ne = 4 for 4 generations), for a comparison to one that was consistently slow (3% per generation; Ne = 16 for 14 generations). The third followed a model for serial founder-flush events. Each founder-flush episode involved a one-generation pulse of fast inbreeding (Ne = 4) followed by two generations of very low (or no) inbreeding, yielding high intergenerational variation (i.e., for an average inbreeding rate of 4% per generation). Allozyme assays showed that we achieved the intended final inbreeding coefficient of about 37%. All inbreeding schemes decreased fitness levels in terms of egg-to-adult viability, development time, and male mating success relative to the outbred control. The consistently fast inbreeding protocol had more pronounced reductions in fitness, relative to the other two inbreeding schemes. In comparison to the fast and punctuated regimes, the consistently slow protocol preserved evolutionary potential (as assayed by the genetic divergence of subpopulations exposed to different environments) in egg-to-adult viability, and (albeit anecdotally) reduced the extinction probabilities, especially in a novel environment. The punctuated treatment did not optimize the potential for purge as predicted, but instead reduced fitness, evolutionary potential, and environmental responsiveness (as measured by genotype-by-environment interactions). This founder-flush treatment also had the highest extinction probabilities. Longer periods of population flush might be necessary to purge effectively in a punctuated scheme. We conclude that the rate of inbreeding, independent from the final level, can have important effects on population fitness, environmental responsiveness, and evolutionary potential.

Animals↗

On the expected relationship between inbreeding, fitness, and extinction.

We assessed the expected relationship between the level and the cost of inbreeding, measured either in terms of fitness, inbreeding depression or probability of extinction. First, we show that the assumption of frequent, slightly deleterious mutations do agree with observations and experiments, on the contrary to the assumption of few, moderately deleterious mutations. For the same inbreeding coefficient, populations can greatly differ in fitness according to the following: (i) population size; larger populations show higher fitness (ii) the history of population size; in a population that recovers after a bottleneck, higher inbreeding can lead to higher fitness and (iii) population demography; population growth rate and carrying capacity determine the relationship between inbreeding and extinction. With regards to the relationship between inbreeding depression and inbreeding coefficient, the population size that minimizes inbreeding depression depends on the level of inbreeding: inbreeding depression can even decrease when population size increases. It is therefore clear that to infer the costs of inbreeding, one must know both the history of inbreeding (e.g. past bottlenecks) and population demography.

Adaptation, Physiological↗

Inbreeding trends and application of optimized selection in the UK Holstein population.

Important increases in the rates of inbreeding have recently been observed in dairy cattle populations, and methods have been proposed to address these increases. The aims of this study were to estimate the current level and rates of inbreeding in the UK Holstein population and to investigate the potential of applying optimized selection to manage the rates of inbreeding. Inbreeding coefficients were calculated for the entire UK Holstein population using 1940 as the base year. Rates of inbreeding were obtained for 3 time periods by regressing mean inbreeding coefficients on the year of birth of the animals. The expected average pedigree index and expected inbreeding of offspring using optimized contributions for a given set of selection candidates was compared to the expected pedigree index and inbreeding of offspring for the same set of selection candidates using observed contributions. The rate of inbreeding in the UK Holstein population has increased substantially since 1990 when compared to previous time periods. This increase is most likely due to the large influence of a few related sires on the breed in the mid- to late 1980s. The introduction of the individual animal model in the early 1990s may also have contributed to increased inbreeding. Optimized selection appears to represent a promising selection tool, not only to manage rates of inbreeding, but also to increase genetic gain at the same rate of inbreeding.

Animals↗

Effects of inbreeding in the dam on dystocia and stillbirths in US Holsteins.

Dystocia scores were recorded by producers on 120,434 Holsteins (218,213 records) from 1985 through 1996; dystocia scores 3 to 5 were coded as difficult births. Stillbirths were recorded for deaths within the first 48 h after birth. Data were restricted to registered cows for pedigree completeness, and inbreeding coefficients were calculated using 5-generation pedigrees. Computational restrictions required that subsets of the data be created by choosing herds at random but using all records from selected herds. Effects of inbreeding in the dam were estimated in a sire-maternal grandsire (of the calf) threshold model using Gibbs sampling. The model included fixed effects of calf sex and inbreeding of the dam and random effects of herd-year-season of birth, additive genetic, and residual effects. First, second, and third parities were analyzed separately. Solutions for sex of calf and inbreeding from different parities were converted to expected change in probability of dystocia or stillbirth per 1% increase in inbreeding. Inbreeding effects were largest for first-parity cows giving birth to male calves at a 0.42% increase in probability of dystocia/1% increase in inbreeding. Effects of inbreeding for first-parity dams giving birth to female calves were smaller, 0.30%/1% increase in inbreeding. Incidence of stillbirths increased 0.25 and 0.20% for male and female calves/1% increase in inbreeding for first parity births. Effects of inbreeding on dystocia and stillbirths declined with parity. Effects of inbreeding were small, especially in later parities, but were consistently unfavorable.

Animals↗