Inbreeding Depression Values Research Articles

Mitosis, stature and evolution of plant mating systems: low-Phi and high-Phi plants.

There is a long-recognized association in plants between small stature and selfing, and large stature and outcrossing. Inbreeding depression is central to several hypotheses for this association, but differences in the evolutionary dynamics of inbreeding depression associated with differences in stature are rarely considered. Here, we propose and test the Phi model of plant mating system evolution, which assumes that the per-generation mutation rate of a plant is a function of the number of mitoses (Phi) that occur from zygote to gamete, and predicts fundamental differences between low-Phi (small-statured) and high-Phi (large-statured) plants in the outcomes of the joint evolution of outcrossing rate and inbreeding depression. Using a large dataset of published population genetic studies of angiosperms and conifers, we compute fitted values of inbreeding depression and deleterious mutation rates for small- and large-statured plants. Consistent with our Phi model, we find that populations of small-statured plants exhibit a range of mating systems, significantly lower mutation rates, and intermediate inbreeding depression, while large-statured plants exhibit very high mutation rates and the maximum inbreeding depression of unity. These results indicate that (i) inbred progeny typically observed in large-statured plant populations are completely lost prior to maturity in nearly all populations; (ii) evolutionary shifts from outcrossing to selfing are generally not possible in large-statured species, rather, large-statured species are more likely to evolve mating systems that avoid selfing such as self-incompatibility and dioecy; (iii) destabilization of the mating system-high selfing rate with high-inbreeding depression-might be a common occurrence in large-statured species; and (iv) large-statured species in fragmented populations might be at higher risk of extinction than previously thought. Our results help to unify and simplify a large and diverse field of research, and serve to emphasize the importance that developmental and genetic constraints play in the evolution of plant mating systems.

GENETIC PARAMETERS OF SOME AGRONOMIC TRAITS IN YELLOW MAIZE UNDER TWO PLANTING DATES

Six population’s seeds of four yellow maize crosses were formed during 2001 and 2002 growing seasons. Their plants were evaluated during 2003 growing season under two planting dates (14th May and 29th June) for six agronomic traits at the Agric. Res. Stat. of Fac. of Agric., Ain Shams Univ., Shalakan, Kalubia Gover-norate, Egypt. The present work aimed to determine the genetic parameters and their interactions with planting dates for grain yield per plant, 100-kernel weight, number of kernels per row, ear length, ear diameter, and days to silking in the six popula-tions (P1, P2, F1, F2, BC1 and BC2). Mean values of the six populations for all studied traits in all crosses were higher under normal planting date than those under late one. Therefore, normal planting date seemed to be non-stress environment. The potence ratio for all traits in the four crosses exceeded (+1) except days to silking where it was less (-1). The highest heterosis percentage relative to mid and better parent reached 192.06% and 152.01% in cross 1 for grain yield per plant under late plant-ing date. Inbreeding depression values were not-significant in all studied traits ex-cept ear diameter and 100-kernel weight in crosses 1 and 3 as well as grain yield per plant in all crosses also, it was positive for most studied characters in the four cross-es except days to silking trait. Dominance occupied the first rank, additive or domi-nance type of epistasis occupied the second or the third contributor to the genetic ef-fects in order of importance according to cross with exception of days to silking, where additive and additive occupied the first and the second ranks. Narrow sense heritability was relatively high for yield attributes whereas it was low for grain yield per plant. Meantime, expected genetic advance was relatively moderate or low for all traits. Therefore, it could be suggested that selection for most studied traits in the subsequent generations will be relatively more effective than in the early genera-tions.

Evaluation of the extent of among‐family variation in inbreeding depression in the perennial herb Scabiosa columbaria (Dipsacaceae)

Significantly different maternal line responses to inbreeding provide a mechanism for the invasion of a selfing variant into a population. The goal of this study was to examine the extent of family-level variation in inbreeding depression in the mixed-mating, perennial herb Scabiosa columbaria. Plants from one population were raised, and hand-pollinated to produce selfed and outcrossed progeny, and the effects of inbreeding depression on life-cycle traits were analyzed. Inbreeding depression significantly affected early life cycle traits. The pollination treatment by family interaction was significant for almost all traits, indicating a high family-level variation in inbreeding depression. The correlations between inbreeding depression values (e.g., percentage germination and flowering date, and flowering date and aboveground biomass) exhibited alternate signs, illustrating the type of association between inbreeding depression loci for different traits across the life cycle. Overall, it is concluded that the extent of among-family variation in inbreeding depression might allow a selfing variant of S. columbaria to invade an outcrossing population, though the pattern of correlations between inbreeding depression values might prevent effective purging of the deleterious genetic load.

Depressão por endogamia em populações de milho-pipoca

Oito populações de milho-pipoca (CMS-42, CMS-43, Zélia, RS-20, Catedral, Zaeli, UEM-J1 e UEM-M2) foram avaliadas com o objetivo de obter estimativas de depressão por endogamia, componentes genéticos de média e comparar essas estimativas por grupos de populações. O delineamento experimental utilizado foi o de blocos ao acaso em esquema de parcelas subdivididas, cujos tratamentos primários foram os níveis de endogamia (S1 e S0) e os tratamentos secundários, as populações, com três repetições, em dois anos agrícolas (1998/99 e 1999/00). Foram avaliadas várias características, com ênfase no rendimento de grãos (RG) e na capacidade de expansão (CE). O método de GARDNER (1965) foi empregado para obter as estimativas dos componentes genéticos de médias. Observaram-se menores valores de depressão por endogamia e predominância de efeitos gênicos aditivos para CE em relação ao RG. A depressão endogâmica para rendimento de grãos foi maior no grupo dos compostos em relação às populações não melhoradas e à população melhorada. A probabilidade de sucesso na obtenção de linhagens vigorosas e de alta capacidade de expansão é baixa para todas as populações.

Inbreeding avoidance through kin recognition: choosy females boost male dispersal.

Inbreeding avoidance is predicted to induce sex biases in dispersal. But which sex should disperse? In polygynous species, females pay higher costs to inbreeding and thus might be expected to disperse more, but empirical evidence consistently reveals male biases. Here, we show that theoretical expectations change drastically if females are allowed to avoid inbreeding via kin recognition. At high inbreeding loads, females should prefer immigrants over residents, thereby boosting male dispersal. At lower inbreeding loads, by contrast, inclusive fitness benefits should induce females to prefer relatives, thereby promoting male philopatry. This result points to disruptive effects of sexual selection. The inbreeding load that females are ready to accept is surprisingly high. In absence of search costs, females should prefer related partners as long as delta<r/(1+r) where r is relatedness and delta is the fecundity loss relative to an outbred mating. This amounts to fitness losses up to one-fifth for a half-sib mating and one-third for a full-sib mating, which lie in the upper range of inbreeding depression values currently reported in natural populations. The observation of active inbreeding avoidance in a polygynous species thus suggests that inbreeding depression exceeds this threshold in the species under scrutiny or that inbred matings at least partly forfeit other mating opportunities for males. Our model also shows that female choosiness should decline rapidly with search costs, stemming from, for example, reproductive delays. Species under strong time constraints on reproduction should thus be tolerant of inbreeding.

Overdominant epistatic loci are the primary genetic basis of inbreeding depression and heterosis in rice. II. Grain yield components.

The genetic basis underlying inbreeding depression and heterosis for three grain yield components of rice was investigated in five interrelated mapping populations using a complete RFLP linkage map, replicated phenotyping, and the mixed model approach. The populations included 254 F(10) recombinant inbred lines (RILs) derived from a cross between Lemont (japonica) and Teqing (indica), two backcross (BC) and two testcross populations derived from crosses between the RILs and the parents plus two testers (Zhong413 and IR64). For the yield components, the RILs showed significant inbreeding depression and hybrid breakdown, and the BC and testcross populations showed high levels of heterosis. The average performance of the BC or testcross hybrids was largely determined by heterosis. The inbreeding depression values of individual RILs were negatively associated with the heterosis measurements of the BC or testcross hybrids. We identified many epistatic QTL pairs and a few main-effect QTL responsible for >65% of the phenotypic variation of the yield components in each of the populations. Most epistasis occurred between complementary loci, suggesting that grain yield components were associated more with multilocus genotypes than with specific alleles at individual loci. Overdominance was also an important property of most loci associated with heterosis, particularly for panicles per plant and grains per panicle. Two independent groups of genes appeared to affect grain weight: one showing primarily nonadditive gene action explained 62.1% of the heterotic variation of the trait, and the other exhibiting only additive gene action accounted for 28.1% of the total trait variation of the F(1) mean values. We found no evidence suggesting that pseudo-overdominance from the repulsive linkage of completely or partially dominant QTL for yield components resulted in the overdominant QTL for grain yield. Pronounced overdominance resulting from epistasis expressed by multilocus genotypes appeared to explain the long-standing dilemma of how inbreeding depression could arise from overdominant genes.

Comparison among Inbreeding Systems in Maize

The use of less severe inbreeding systems than self-fertilization has been suggested as a means of developing more vigorous maize lines (Zea mays L.). To compare different inbreeding systems, nine traits were evaluated in inbred lines developed by full-sib mating (F=0.5), self-fertilization (F=0.5), half-sib mating (F= 0.305), from the following populations: BR 105, a synthetic variety, and ESALQ-PB 4 , which is a composite of wide genetic base. The means and genetic parameters comparisons showed no important advantages of full-sib crosses over selfing. The inbreeding depression values for grain yield were similar between these groups of lines and inferior for half-sib lines. The inbreeding depression rates / 1% of expected homozigosity were similar among the three inbreeding systems. We conclude, in view of large inbreeding depression showed by tropical populations of maize in Brazil, that it is more reasonable to allocate resources and time in breeding source populations than to use milder forms of inbreeding.

The mutation load under tetrasomic inheritance and its consequences for the evolution of the selfing rate in autotetraploid species

Single-locus equilibrium frequencies of a partially recessive deleterious mutation under the mutation–selection balance model are derived for partially selfing autotetraploid populations. Assuming multiplicative fitness interactions among loci, approximate solutions for the mean fitness and inbreeding depression values are also derived for the multiple locus case and compared with expectations for the diploid model. As in diploids, purging of deleterious mutations through consanguineous matings occurs in autotetraploid populations, i.e. the equilibrium mutation load is a decreasing function of the selfing rate. However, the variation of inbreeding depression with the selfing rate depends strongly on the dominance coefficients associated with the three heterozygous genotypes. Inbreeding depression can either increase or decrease with the selfing rate, and does not always vary monotonically. Expected issues for the evolution of the selfing rate consequently differ depending on the dominance coefficients. In some cases, expectations for the evolution of the selfing rate resemble expectations in diploids; but particular sets of dominance coefficients can be found that lead to either complete selfing or intermediate selfing rates as unique evolutionary stable state.

Late-acting inbreeding depression in both male and female function of echium vulgare (Boraginaceae)

We present data on late-acting inbreeding depression in pollen performance, siring success and seed production in Echium vulgare. Pollen viability and rate of pollen-tube growth were both lower for pollen from plants derived from selfing than for pollen from plants derived from outcrossing. Pollen tube numbers within the styles did not differ for pollen from plants derived from selfing or outcrossing. A pollination experiment with two mixtures of pollen from plants derived from selfing or outcrossing, revealed a significant decline of 55% in siring success for pollen from plants derived from selfing. A second experiment with a complete diallel design revealed inbreeding depression for both siring success of the offspring (32.8%) and a decline in seed production of the offspring (34.8%-40.6%). In addition, results indicated a heritable component for seed number per flower. Offspring fitness, measured as seed production and siring ability, can be severely affected by late-acting inbreeding depression. Inbreeding depression values for male and female functions were not correlated. Both functions must therefore be considered when calculating inbreeding depression.

Inbreeding depression and partial selfing: evolutionary implications of mixed-mating in a coastal endemic, silene douglasii var. oraria (Caryophyllaceae)

Recent studies have found moderate to high levels of selfing in plants despite high inbreeding depression. Because both factors influence the evolution and persistence of rare plants, we conducted glasshouse and field studies of pollination and inbreeding in Silene douglasii var. oraria, a perennial tetraploid endemic to coastal prairies. We detected: (i) variation in reproduction or inbreeding depression among life stages, years and maternal families; (ii) partial selfing yet higher relative fitness in outcrossed than selfed progeny; (iii) differing values of selfing and inbreeding depression using population means vs. matched maternal families. Fruit and seed production varied significantly with pollination treatment and year in flowers manipulated in situ during three seasons of growth. Hand-pollinations providing pollen in excess of ovule production in 1996 yielded more seeds than marked, open-pollinated flowers, implying pollen limitation of seed production. However, among-year differences in reproductive success after open-pollination (i.e. values equivalent to autogamy, selfing or outcrossing) suggest that pollination levels also vary temporally. In pollinations matched by maternal family, selfing yielded significantly fewer seeds than outcrossing. Fitness differences between inbred and outbred progeny were significant (P < 0.05) for seed production, percentage germination, and biomass or fecundity, but not for survival. Maternal family data gave selfing rates intermediate between obligate outcrossers and predominant selfers (S = 0.34-0.51), but population-wide means gave unusually high values (S = 1.1-1.6). Cumulative inbreeding depression was 76% for maternal families, and 70-85% using population means; in all cases, inbreeding depression values were high in early and late life stages, and lowest for survival. Thus far, reproductive assurance offers the most cogent explanation for the coexistence of moderate selfing and high inbreeding depression in this strongly protandrous Silene once thought to be highly outcrossing. This possibility merits further study in other rare plants with mixed-mating systems, where inbreeding depression and pollinator scarcity may both compromise population persistence and raise the threshold below which selfing is favoured by evolution.

Testcross Evaluation of Soybean Germplasm

The F1 generation of a testcross previously was not used to evaluate soybean [Glycine max (L.) Merr.] germplasm, primarily because production of large amounts of hybrid seed has been impractical and because evaluation of quantitative traits with one or a few plants does not give reproducible results. Recently, a method for efficiently producing testcross seed was developed by using the male sterility locus, Ms6, and a genetically linked seedling marker locus, W1. The objectives of this research were (i) estimation and comparison heterosis and inbreeding depression values for agronomic traits of testcross lines; and (it) determination of the utility of per se and testcross evaluation for selecting germplasm to be included in an existing cultivar development program. Two testers were used to evaluate six lines in three‐row plots. Significant F1 and F2 midparent heterosis and inbreeding depression were observed for nearly all traits, including grain yield. The choice of tester was ascertained to be important in determining parental value of germplasm. Parental lines were evaluated by using a combination of per se performance, heterosis values, and a recently developed testcross statistic, Ti. The Ti value was better than F1 midparent heterosis for identifying lines suitable for use in direct crosses. F1 midparent heterosis was better than the Ti value for identifying valuable lines requiring pre‐breeding effort. A combination of F1 and F2 midparent heterosis values identified lines likely to have genes with dominance effects; although, because of inbreeding depression, use of F2 heterosis alone failed to identify valuable lines.

Population improvement in lucerne (Medicago sativa L.): components of inbreeding depression are different in original and improved populations

Summary. Inbreeding depression, the lowered fitness of inbred individuals compared with their non-inbred counterparts, is an important concept in lucerne improvement; but is poorly understood. Two-allele autotetraploid populations are suitable for studying inbreeding depression, especially when the population improvement strategy involves inbreeding, because they are derived from chromosome-doubling of hybrid diploid plants. They have a maximum of 2 alleles and a single allelic interaction per locus. Inbreeding depression was compared in original 2-allele autotetraploid populations and populations that had undergone inbreeding and selection. The original and improved (selected) populations were produced by intercrossing 2 single-cross lines from the original and improved 2-allele autotetraploid populations respectively. Herbage yield of the S1 and intercrossed generations derived from these populations was evaluated in field trials at Arlington, Wisconsin, USA, and used to estimate inbreeding depression. Herbage yield of the S1 and intercrossed generations derived from the improved population were significantly (P&lt;0.01) higher, by 13.3 and 24%, respectively, than those derived from the original population. Selection during inbreeding probably decreased the frequency of deleterious alleles and accumulated favourable alleles. Inbreeding depression values were higher in the improved compared with the original population. Genetic load of deleterious alleles may account for much of the inbreeding depression observed in the original population, whereas, in the improved population, loss of heterozygosity or non-additive gene interactions between favourable alleles on linked chromosome segments may account for the substantial inbreeding depression. Therefore, in a population improvement program, the causes of inbreeding depression seem to be more important than their estimated value.

Outcrossing rates and inferred levels of inbreeding depression in gynodioecious Cucurbita foetidissima (Cucurbitaceae)

Sex-specific outcrossing rates and the inbreeding coefficient of adults in two populations of gynodioecious Cucurbita foetidissima were estimated using progeny array data from four allozyme loci to compare the frequencies of self-fertilization and the estimated levels of inbreeding depression to predictions from sex ratio theory. The frequencies of self-fertilization by hermaphrodites in both populations were similar and averaged 73 per cent. The outcrossing rate for females in one population was not different from unity whereas the estimated rate of outcrossing by females in the second population (t = 0.593, SE 0.178) indicated the occurrence of biparental inbreeding. Despite considerable self-fertilization by hermaphrodites, inbreeding coefficients of adult plants in both populations were not different from zero and thus inferred values of inbreeding depression were not different from one. Electrophoretically inferred levels of inbreeding depression are somewhat in excess of the value of 0.71 (SE 0.07) obtained in a previous field experiment which tested first-year survival of selfed and outcrossed seeds in this long-lived perennial. The high frequency of self-fertilization by hermaphrodites combined with severe inbreeding depression provides a strong selective force which, along with increased seed-set by females, is sufficient to maintain observed frequencies of females in natural populations of C. foetidissima.

The effect of linkage and population size on inbreeding depression due to mutational load

Using a stochastic model of a finite population in which there is mutation to partially recessive detrimental alleles at many loci, we study the effects of population size and linkage between the loci on the population mean fitness and inbreeding depression values. Although linkage between the selected loci decreases the amount of inbreeding depression, neither population size nor recombination rate have strong effects on these quantities, unless extremely small values are assumed. We also investigate how partial linkage between the loci that determine fitness affects the invasion of populations by alleles at a modifier locus that controls the selfing rate. In most of the cases studied, the direction of selection on modifiers was consistent with that found in our previous deterministic calculations. However, there was some evidence that linkage between the modifier locus and the selected loci makes outcrossing less likely to evolve; more losses of alleles promoting outcrossing occurred in runs with linkage than in runs with free recombination. We also studied the fate of neutral alleles introduced into populations carrying detrimental mutations. The times to loss of neutral alleles introduced at low frequency were shorter than those predicted for alleles in the absence of selected loci, taking into account the reduction of the effective population size due to inbreeding. Previous studies have been confined to outbreeding populations, and to alleles at frequencies close to one-half, and have found an effect in the opposite direction. It therefore appears that associations between neutral and selected loci may produce effects that differ according to the initial frequencies of the neutral alleles.

Multilocus models of inbreeding depression with synergistic selection and partial self-fertilization

SummaryMean fitness and inbreeding depression values in multi-locus models of the control of fitness were studied, using both a model of mutation to deleterious alleles, and a model of heterozygote advantage. Synergistic fitness interactions between loci were assumed, to find out if this more biologically plausible model altered the conclusions we obtained previously using a model of multiplicative interactions. Systems of unlinked loci were assumed. We used deterministic computer calculations, and approximations based on normal or Poisson theory. These approximations gave good agreement with the exact results for some regions of the parameter space. In the mutational model, we found that the effect of synergism was to lower the number of mutant alleles per individual, and thus to increase the mean fitness, compared with the multiplicative case. Inbreeding depression, however, was increased. Similar effects on mean fitness and inbreeding depression were found for the case of heterozygote advantage. For that model, the results were qualitatively similar to those previously obtained assuming multiplicativity. With the mutational load model, however, the mean fitness sometimes decreased, and the inbreeding depression increased, at high selfing rates, after declining as the selfing rate increased from zero. We also studied the behaviour of modifier alleles that changed the selfing rate, introduced into equilibrium populations. In general, the results were similar to those with the multiplicative model, but in some cases an ESS selfing rate, with selfing slightly below one, existed. Finally, we derive an approximate expression for the inbreeding depression in completely selfing populations. This depends only on the mutation rate and the dominance coefficient and can therefore be used to obtain estimates of the mutation rate to mildly deleterious alleles for plant species.

INBREEDING DEPRESSION IN SCHIEDEA GLOBOSA AND S. SALICARIA (CARYOPHYLLACEAE), SUBDIOECIOUS AND GYNODIOECIOUS HAWAIIAN SPECIES

Progeny produced by inbreeding were compared to progeny derived from outcrosses for gynodioecious Schiedea salicaria and subdioecious S. globosa to assess fitness consequences of breeding system on parental fecundity (seeds per capsule) and progeny measures of fitness (germination, survival, biomass, and number of flowers). Results from both species indicated that inbreeding depression occurred at all measured stages of the life history. In both species, different females showed different levels of inbreeding depression. Multiplicative fitness functions of the ratio of values for selfed and outcrossed progeny in S. salicaria resulted in inbreeding depression values of 0.62–0.94. Within‐ vs. between‐family crosses of S. globosa also resulted in inbreeding depression values as high as 0.49. These values suggest that inbreeding depression may promote the evolution of dioecy within S. globosa and S. salicaria, depending on the levels of natural outcrossing.

Inbreeding Depression in Schiedea globosa and S. Salicaria (Caryophyllaceae), Subdioecious and Gynodioecious Hawaiian Species

Progeny produced by inbreeding were compared to progeny derived from outcrosses for gynodioecious Schiedea salicaria and subdioecious S. globosa to assess fitness consequences of breeding system on parental fecundity (seeds per capsule) and progeny measures of fitness (germination, survival, biomass, and number of flowers). Results from both species indicated that inbreeding depression occurred at all measured stages of the life history. In both species, different females showed different levels of inbreeding depression. Multiplicative fitness functions of the ratio of values for selfed and outcrossed progeny in S. salicaria resulted in inbreeding depression values of 0.62–0.94. Within- vs. between-family crosses of S. globosa also resulted in inbreeding depression values as high as 0.49. These values suggest that inbreeding depression may promote the evolution of dioecy within S. globosa and S. salicaria, depending on the levels of natural outcrossing.

Dynamics of correlated genetic systems. II. Simulation studies of chromosomal segments under selection

The dynamics of chromosomal segments under selection are investigated by comparing experimental data to simulations of simple models of selection. The simulations assume 93 loci distributed evenly along an entire chromosome. The two issues addressed in this paper concern rates of decay of linkage disequilibria for chromosomes under selection and rates of gene frequency change after perturbation of gametic frequencies to states near the edge of the gametic frequency simplex. The findings are: (1) If reasonable values of inbreeding depression are assumed, linkage disequilibria decays to zero but at a rate nearly twice that expected from neutral theory. Experimental results also show accelerated decay rates. The acceleration of decay seems to be a simple consequence of the increased heterozygosity produced by selection. It is, therefore, argued that massive linkage disequilibria, of the kind found by Franklin and Lewontin (1970) in their simulations, are unlikely to characterize the genetic structure of natural populations of random mating organisms. (2) It is possible to distinguish between two time-honored models of multilocus selection, known as the symmetric overdominant and classical models, on the basis of gene frequency change near the edge of the gametic frequency simplex assuming linkage disequilibria is intense. (3) Examination of experimental data from perturbation experiments shows that neither of these elementary models adequately account for observed rates of gene frequency change, although the symmetric overdominant model does provide the better fit. Instead the experimental data suggest a markedly nonuniform distribution of selective effects along the chromosome. The data also suggest that these selective effects combine in markedly nonadditive ways in determining joint fitness.

Selection among synthetics

The effect of the number of parents and their level of inbreeding on the mean and the variance of synthetic varieties was studied for diploids and autotetraploids. The number of parents and their level of inbreeding act in opposing ways. Maximum mean requires a high number of parents and no inbreeding. Maximum variance requires a small number of highly inbred and unrelated parents. When the number of parents increases (k ≥ 2) the coefficients of the components of variance decrease, and the decrease is more rapid for variances associated with increasing order of interactions between genes. The coefficients of components of variance increase as the level of inbreeding of the parents increases and the increase is greater for components associated with increasing order of interactions between genes.Consequently, according to the values of inbreeding depression and the components of genetic variance and heritability, an optimum genetic base may exist, i.e., an ideal combination of the number of parents and the level of their inbreeding. With no inbreeding, selection among synthetics uses mainly additive variance. By increasing the level of inbreeding of the parents, the effects of dominance and of additive X additive variances on genetic advance when selecting among synthetics increase. One cycle of selection among synthetics appears more efficient than individual selection within populations. The problem of population improvement before selecting among synthetics is discussed.