The benefits of interpopulation hybridization diminish with increasing divergence of small populations

Interpopulation hybridization can increase the viability of small populations suffering from inbreeding and genetic drift, but it can also result in outbreeding depression. The outcome of hybridization can depend on various factors, including the level of genetic divergence between the populations, and the number of source populations. Furthermore, the effects of hybridization can change between generations following the hybridization. We studied the effects of population divergence (low vs. high level of divergence) and the number of source populations (two vs. four source populations) on the viability of hybrid populations using experimental Drosophila littoralis populations. Population viability was measured for seven generations after hybridization as proportion of populations facing extinction and as per capita offspring production. Hybrid populations established at the low level of population divergence were more viable than the inbred source populations and had higher offspring production than the large control population. The positive effects of hybridization lasted for the seven generations. In contrast, at the high level of divergence, the viability of the hybrid populations was not significantly different from the inbred source populations, and offspring production in the hybrid populations was lower than in the large control population. The number of source populations did not have a significant effect at either low or high level of population divergence. The study shows that the benefits of interpopulation hybridization may decrease with increasing divergence of the populations, even when the populations share identical environmental conditions. We discuss the possible genetic mechanisms explaining the results and address the implications for conservation of populations.     

Microgeographic genetic structure and intraspecific parasitism in the ant Leptothorax nylanderi

1. Genetic colony structure of the small central European ant Leptothorax nylanderi is affected strongly by ecological constraints such as nest site availability and intraspecific social parasitism. 2. Although L. nylanderi is generally monogynous and monandrous, more than a quarter of all nests collected in a dense population near Würzburg, Germany, contained several matrilines. As shown by microsatellite analysis, the average nest‐mate relatedness in these nests was 0.20. Genetically heterogeneous nests arise from nest take‐over by alien colonies or founding queens, a result of severe competition for nest sites. 3. In summer, more than one‐third of all colonies inhabited several nest sites at a time. Polydomy appears to be rather limited, with two or three nests belonging to a single polydomous colony. 4. Queens appear to dominate male production; only a small fraction (8%) of males was definitively not progeny of the queen present but might have been worker progeny or offspring of another queen. 5. Strong evidence for heterozygote deficiency was found and a total of nine diploid males was discovered in two colonies. These findings suggest deviation from random mating through small, localised nuptial flights.     

Inbreeding under a cyclical mating system

Summary General recursion formulae for the coefficient of inbreeding under a cyclical mating system were derived in which one male and one female are selected from each of the n families per generation (population size N = 2 n). Each male is given the family number of his sire in each generation, while his mate comes from another family, varying systematically in different generations. Males of the r-th family in generations 1, 2, 3,..., t = n–1 within each cycle mate with females from families r+1, r+2, r+3,..., r+t to produce generations 2, 3, 4,..., t+1=1, respectively. The change in heterozygosity shows a cyclical pattern of rises and falls, repeating in cycles of n–1 generations. The rate of inbreeding oscillates between <-3% to >6% in different generations within each cycle, irrespective of the population size. The average rate of inbreeding per generation is approximately 1/[4 N-(Log₂N+1)], which is the rate for the maximum avoidance of inbreeding. The average inbreeding effective population size is approximately 2 N–2.     

Phyletic and evolutionary relationships ofBrachyscome lineariloba (Compositae)

A comparison of karyotypes ofBrachyscome breviscapis (2n = 8),B. lineariloba cytodemes E (2n = 10), B (2n = 12) and C (2n = 16) suggests that these species have a homoelogous basic set of four chromosome pairs, two large pairs and two small, and that theB. lineariloba cytodemes E, B and C are related toB. breviscapis by successive additions of small chromosomes. A pronounced asynchrony of chromosome condensation between these large and small chromosomes has been observed. In the artificial hybrids betweenB. dichromosomatica (2n = 4) ×B. breviscapis, and theB. lineariloba cytodemes, theB. dichromosomatica chromosomes are similar in size and condensation behaviour to the small chromosomes ofB. breviscapis and ofB. lineariloba cytodemes E, B and C. Meiotic pairing in these hybrids also demonstrates the strong affinities between these chromosomes. It is suggested thatB. breviscapis may be of amphidiploid origin between a species with two large early condensing chromosome pairs and another,B. dichromosomatica-like species with two small late condensing pairs. It seems most likely that the additional small and late condensing chromosomes inB. lineariloba cytodemes E, B and C are derived from theB. dichromosomatica-like parent, and that each addition increases vigour, fecundity and drought tolerance, allowing these cytodemes to colonize more open and arid environments. Transmission of the univalents in the quasidiploidB. lineariloba cytodeme E was verified as being via the pollen, and not via the embryo sacs.The cytology ofBrachyscome lineariloba (Compositae, Asteroidae), 10.     

Mating system of Bracon hebetor (Hymenoptera: Braconidae)

Abstract.

• 1 We report on the mating system of a field population of the parasitic wasp, Bracon hebetor, on a corn pile infested by the Indian meal moth, Plodia interpunctella. We demonstrate that the mating system is based upon male scramble competition polygyny with male aggregations on high places on the corn.
• 2 The sex ratio among adults was greater than 80% males on the surface of the corn, whereas below the surface the sex ratio was less than 45%. Males actively courted females on the surface, but there were no aggressive interactions among males during courtship or mating.
• 3 Approximately 20% of the females found on the surface of the corn had no sperm in their spermathecae, regardless of age, but the numbers of unmated females decreased later during the day.
• 4 In laboratory studies we showed that females from this population oviposit a female biassed sex ratio, and that only 14% of females were mated before dispersing from their place of emergence.
• 5 Thus sib-mating is unlikely in this gregarious parasitoid. This outcrossing mating system probably arose because of severe inbreeding depression that B.hebetor suffers via a sex locus: diploids that are heterozygous at the sex locus develop into females, but homozygous diploids are male and are generally inviable. The female biassed sex ratio may have evolved in B. hebetor in response to males being the more expensive sex, females dispersing more frequently from the population than males, or a fraction of females remaining unmated in the population.

     

Modeling problems in conservation genetics using captive Drosophila populations: Consequences of equalizing founder representation

Equalizing founder representation is a recommended practice for maintaining captive populations. However, this procedure has not been subject to controlled experimental evaluation. The effects on inbreeding, genetic variation, and reproductive fitness of maintaining small captive populations by equalizing founder representation (EFR) versus randomly choosing parents (RC) were compared. Ten replicate lines were created with unequal founder representations, split into EFR and RC lines, and maintained for a further eight generations. Founder representations computed from pedigrees were closer to equality in the EFR lines than in the RC lines or the base population, most of the changes being evident after one generation. Significant benefits of EFR were found in lowered inbreeding (mean inbreeding coefficients of 0.35 and 0.41, respectively, for EFR and RC lines) and average heterozygosity (0.141 for EFR, 0.084 for RC, compared with 0.216 in the base population). However, EFR was not significantly better than RC in moving allele frequencies towards equalized founder representation. No significant difference was found in reproductive fitness between EFR and RC (relative fitnesses compared to the base population were 0.179 for EFR and 0.182 for RC). The use of equalization of founder representation for a few generations can be recommended in the genetic management of captive populations derived from a small number of founders that contribute unequally. © 1992 Wiley-Liss, Inc.     

FIXATION PROBABILITIES OF SELFING RATE MODIFIERS IN SIMULATIONS WITH SEVERAL DELETERIOUS ALLELES WITH LINKAGE

The fixation rates of selfing rate modifiers were found by stochastic simulation in an infinite site model, including effects of several deleterious alleles with variable effects, which were randomly distributed in the genome without assuming any pollen discounting. Previous results on the evolution of selfing obtained by more precise methods were in this study further validated, and it was concluded that the effect of genetic associations on the evolution of mating systems is small except in the case of full pollen discounting. Furthermore, attention was given to the uneven distribution of the genetic load in the population, and the accompanying large among-genome variation in fixation rates. This among-genome variation will be of significance for the evolution of mating systems.     

DROUGHT STRESS AND INBREEDING DEPRESSION IN LYCHNIS FLOS-CUCULI (CARYOPHYLLACEAE)

Interactions between drought stress and inbreeding depression were studied in Lychnis flos-cuculi. Four inbreeding levels (F = 0, 0.25, 0.50 and 0.75), and three watering treatments were used. Performance was scored for germination rate and proportion, survival, plant size, proportion of plants flowering, flowering date, stem height, number of flowers, flower size, anther weight, fruiting proportion and number of capsules. Multiplicative fitness values were estimated from these traits. Inbreeding affected most of the traits studied, and a severe inbreeding depression was found for the combined fitness estimates. The higher inbreeding depression found here relative to the same family groups in a former experiment may reflect greater dominance and suppression in the present experiment at higher density.     

Inbreeding depression in Lychnis flos-cuculi (Caryophyllaceae): effects of different levels of inbreeding

In a controlled crossing experiment on Lychnis flos-cuculi plants in the greenhouse, outbred and selfed maternal plants were each treated with pollen from unrelated plants, siblings and selves. The seeds thus obtained had expected inbreeding coefficients of 0, 0.25 and 0.5 for the outbred maternal plants, and 0, 0.5 and 0.75 for the selfed maternal plants. Seed abortion rate, seed weight and germination rate were estimated. Seedlings were transplanted to an outdoor garden, and monitored for survival, probability of flowering, number of capsules and area of capsules next spring. Inbred seeds germinated slower and in lower proportions than those less inbred, and seedlings had lower survival, flowering, fruit set and area of capsules if inbred. Combined fitness values were estimated from the survival and fecundity components, and severe inbreeding depression was detected for these estimates (0.51 and 0.56 for one generation of selfing). The fitness function decreased linearly with the increase in inbreeding coefficient, which is as expected if the inbreeding depression is additive among loci.