Self-compatible Research Articles

What causes mating system shifts in plants? Arabidopsis lyrata as a case study

The genetic breakdown of self-incompatibility (SI) and subsequent mating system shifts to inbreeding has intrigued evolutionary geneticists for decades. Most of our knowledge is derived from interspecific comparisons between inbreeding species and their outcrossing relatives, where inferences may be confounded by secondary mutations that arose after the initial loss of SI. Here, we study an intraspecific breakdown of SI and its consequences in North American Arabidopsis lyrata to test whether: (1) particular S-locus haplotypes are associated with the loss of SI and/or the shift to inbreeding; (2) a population bottleneck may have played a role in driving the transition to inbreeding; and (3) the mutation(s) underlying the loss of SI are likely to have occurred at the S-locus. Combining multiple approaches for genotyping, we found that outcrossing populations on average harbour 5 to 9 S-locus receptor kinase (SRK) alleles, but only two, S1 and S19, are shared by most inbreeding populations. Self-compatibility (SC) behaved genetically as a recessive trait, as expected from a loss-of-function mutation. Bulked segregant analysis in SC × SI F2 individuals using deep sequencing confirmed that all SC plants were S1 homozygotes but not all S1 homozygotes were SC. This was also revealed in population surveys, where only a few S1 homozygotes were SC. Together with crossing data, this suggests that there is a recessive factor that causes SC that is physically unlinked to the S-locus. Overall, our results emphasise the value of combining classical genetics with advanced sequencing approaches to resolve long outstanding questions in evolutionary biology.

Pollination and sterility in horseradish

AbstractHorseradish (Armoracia rusticana, Brassicaceae) is a crop that seldom produces viable seed and is propagated exclusively via vegetative (or clonal) procedures for commercial production. Male sterility and self‐incompatibility (SI) have been suspected as possible mechanisms underlying sterility in horseradish. Therefore, greenhouse and laboratory analyses were conducted to assess pollen viability, pollen recognition in pistils, and to quantify success in self‐ and cross‐pollination events using various horseradish clones. Pollen viability was high (98–100%); consequently, male sterility was an unlikely explanation for low seed set. Self‐pollination in some horseradish clones produced seed, although most clones showed some level of SI. Significant numbers of pollen tubes were found in ovaries of self‐compatible (SC) individuals soon after stigmas were saturated with pollen; furthermore, between three and six ovules with pollen tubes were found in the micropyle which correlates with the number of seed typically found in mature fruits from outcrossed or SC individuals. The results of this study suggest variation in the breeding system of horseradish, as some clones exhibit complete SC, while other clones are SI that recognize and reject self‐pollen.

Variation in floral morphology and plant reproductive success in four Ipomoea species (Convolvulaceae) with contrasting breeding systems.

This study tested the hypothesis that self-compatibility would be associated with floral traits that facilitate autonomous self-pollination to ensure reproduction under low pollinator visitation. In a comparison of two pairs of Ipomoea species with contrasting breeding systems, we predicted that self-compatible (SC) species would have smaller, less variable flowers, reduced herkogamy, lower pollinator visitation and higher reproductive success than their self-incompatible (SI) congeners. We studied sympatric species pairs, I. hederacea (SC)- I. mitchellae (SI) and I. purpurea (SC)-I. indica (SI), in Mexico, over two years. We quantified variation in floral traits and nectar production, documented pollinator visitation, and determined natural fruit and seed set. Hand-pollination and bagging experiments were conducted to determine potential for autonomous self-pollination and apomixis. Self-compatible Ipomoea species had smaller flowers and lower nectar production than SI species; however, floral variation and integration did not vary according to breeding system. Bees were primary pollinators of all species, but visitation rates were seven times lower in SC than SI species. SC species had a high capacity for autonomous self-pollination due to reduced herkogamy at the highest anther levels. Self-compatible species had two to six times higher fruit set than SI species. Results generally support the hypothesis that self-compatibility and autonomous self-pollination ensure reproduction under low pollinator visitation. However, high variation in morphological traits of SC Ipomoea species suggests they maintain variation through outcrossing. Furthermore, reduced herkogamy was associated with high potential for autonomous self-pollination, providing a reproductive advantage that possibly underlies transitions to self-compatibility in Ipomoea.

Non-self- and self-recognition models in plant self-incompatibility.

The mechanisms by which flowering plants choose their mating partners have interested researchers for a long time. Recent findings on the molecular mechanisms of non-self-recognition in some plant species have provided new insights into self-incompatibility (SI), the trait used by a wide range of plant species to avoid self-fertilization and promote outcrossing. In this Review, we compare the known SI systems, which can be largely classified into non-self- or self-recognition systems with respect to their molecular mechanisms, their evolutionary histories and their modes of evolution. We review previous controversies on haplotype evolution in the gametophytic SI system of Solanaceae species in light of a recently elucidated non-self-recognition model. In non-self-recognition SI systems, the transition from self-compatibility (SC) to SI may be more common than previously thought. Reversible transition between SI and SC in plants may have contributed to their adaptation to diverse and fluctuating environments.

Cloning of the Self-incompatibility SFB Gene from Chinese Apricot ‘Xiaobaixing’ and Construction of the SFB Expression Vectors

Three kinds of expression vectors of a pollen-S determinant were constructed to provide a reference for molecular breeding of self-compatible (SC) Prunus species. An S-haplotype-specific F-box (SFB) protein gene from the ‘Xiaobaixing’ apricot (Prunus armeniaca) was cloned by reverse transcription polymerase chain reaction (RT-PCR) and 3′-rapid-amplification of cDNA ends (3′-RACE). A 1136-bp sequence complementary to the 3′-end of the cDNA (GenBank accession number KP938528.2) with a 912-bp complete open reading frame (ORF) was obtained. The deduced amino acid sequence contained an F-box domain, two variable regions, and two hypervariable regions with structural characteristics similar to SFB in other Rosaceae plants. Sense, antisense, and RNA interference (RNAi) vectors for SFB were constructed by enzyme restriction. The target fragment was restricted using the corresponding restriction enzyme and then directionally inserted between the 35S cauliflower mosaic virus promoter and the nopaline synthase terminator (NOS-ter) of the expression vector pCAMBIA-35S-MCS-NOS-NPTII. The intron-containing hairpin RNA (ihpRNA) was obtained by fusion PCR. The constructed vectors were transferred into Agrobacterium tumefaciens strain LBA4404 by freezing/thawing. The RNAi vector of SFB was also transformed in tobacco (Nicotiana tabacum). The successful construction of these three expression vectors provides a basis for transforming ‘Xiaobaixing’ apricot and the breeding of SC Prunus cultivars.

Mating system transitions in Solanum habrochaites impact interactions between populations and species.

In plants, transitions in mating system from outcrossing to self-fertilization are common; however, the impact of these transitions on interspecific and interpopulation reproductive barriers is not fully understood. We examined the consequences of mating system transition for reproductive barriers in 19 populations of the wild tomato species Solanum habrochaites. We identified S.habrochaites populations with self-incompatible (SI), self-compatible (SC) and mixed population (MP) mating systems, and characterized pollen-pistil interactions among S.habrochaites populations and between S.habrochaites and other tomato species. We examined the relationship between mating system, floral morphology, interspecific and interpopulation compatibility and pistil SI factors. We documented five distinct phenotypic groups by combining reproductive behavior with molecular data. Transitions from SI to MP were not associated with weakened interspecific reproductive barriers or loss of known pistil SI factors. However, transitions to SC at the northern range margin were accompanied by loss of S-RNase, smaller flowers, and weakened (or absent) interspecific pollen-pistil barriers. Finally, we identified a subset of SC populations that exhibited a partial interpopulation reproductive barrier with central SI populations. Our results support the hypothesis that shifts in mating system, followed by additional loss-of-function mutations, impact reproductive barriers within and between species.

Genetic variation for pseudo‐self‐compatibility in self‐incompatible populations of Leavenworthia alabamica (Brassicaceae)

Self-incompatibility (SI) promotes outcrossing, but transitions to self-compatibility (SC) are frequent. Population genetic theory describing the breakdown of SI to SC suggests that, under most conditions, populations should be composed of either SI or SC individuals. Under a narrow range of conditions, theory suggests that SI may persist alongside reduced expression of SI (pseudo-SI, PSI) in mixed-mating populations. We studied genetic variation for PSI segregating in four SI populations of Leavenworthia alabamica by measurement of the heritability of pollen tube number after self-pollination. We tested for the role of the S-locus in this variation by sequencing seven S-alleles from plants with high pseudo-SC (PSC) and testing for the co-segregation of these alleles with PSC. We found a continuous distribution of PSC in all populations and 90% of plants exhibited PSC. The heritability ranged from 0.39 to 0.57. All seven S-alleles from plants with high PSC exhibited trans-specific polymorphism, and no stop codons were observed within the c. 600-bp region sequenced. One of these S-alleles was directly associated with the inheritance of PSC. We conclude that heritable variation in PSC is largely a result of genetic variation in the signaling cascade downstream of the S-locus reaction, together with the presence of one leaky S-allele.

Recent selection for self-compatibility in a population of Leavenworthia alabamica.

The evolution of self-compatibility (SC) is the first step in the evolutionary transition in plants from outcrossing enforced by self-incompatibility (SI) to self-fertilization. In the Brassicaceae, SI is controlled by alleles of two tightly linked genes at the S-locus. Despite permitting inbreeding, mutations at the S-locus leading to SC may be selected if they provide reproductive assurance and/or gain a transmission advantage in a population when SC plants self- and outcross. Positive selection can leave a genomic signature in the regions physically linked to the focus of selection when selection has occurred recently. From an SC population of Leavenworthia alabamica with a known nonfunctional mutation at the S-locus, we collected sequence data from a ∼690 Kb region surrounding the S-locus, as well as from regions not linked to the S-locus. To test for recent positive selection acting at the S-locus, we examined polymorphism and the site-frequency spectra. Using forward simulations, we demonstrate that recent selection of the strength expected for SC at a locus formerly under balancing selection can generate patterns similar to those seen in our empirical data.

Dissect style response to pollination using metabolite profiling in self-compatible and self-incompatible tomato species

Tomato style is the pathway for pollen germination and pollen tubes growth from the stigma to the ovules where fertilization occurs. It is essential to supplying the nutrients for pollen tube growth and guidance for the pollen tubes. To our knowledge, style also regulates gametophytic self-incompatibility (SI) in tomato species. This study identified the metabolites and monitored the metabolic changes of self-incompatible and self-compatible tomato with self-pollinated or unpollinated styles by gas chromatography-mass spectrometry (GC-MS). A total of 9 classes of compounds were identified in SI and self-compatibility (SC) self-pollinated and unpollinated styles which included amino acids, sugars, fatty acids/lipids, amines, organic acids, alcohols, nitriles, inorganic acids and other compounds. The contents of d-Mannose-6-phosphate, Cellobiose, Myristic acid, 2,4-Diaminobutyric acid, Inositol and Urea were significantly decreased and the rest did not significantly change in SI styles. But change of metabolites content significantly happened in SC styles. In addition, among the total 9 classes of compounds, the different metabolites accounted for a different proportion in amino acids, sugars, amines, organic acids and alcohols compared SC and SI. The result indicated that the physiological changes of styles existed differences in SC and SI after self pollination.

Convergent evolution at the gametophytic self-incompatibility system in Malus and Prunus.

S-RNase-based gametophytic self-incompatibility (GSI) has evolved once before the split of the Asteridae and Rosidae. This conclusion is based on the phylogenetic history of the S-RNase that determines pistil specificity. In Rosaceae, molecular characterizations of Prunus species, and species from the tribe Pyreae (i.e., Malus, Pyrus, Sorbus) revealed different numbers of genes determining S-pollen specificity. In Prunus only one pistil and pollen gene determine GSI, while in Pyreae there is one pistil but multiple pollen genes, implying different specificity recognition mechanisms. It is thus conceivable that within Rosaceae the genes involved in GSI in the two lineages are not orthologous but possibly paralogous. To address this hypothesis we characterised the S-RNase lineage and S-pollen lineage genes present in the genomes of five Rosaceae species from three genera: M. × domestica (apple, self-incompatible (SI); tribe Pyreae), P. persica (peach, self-compatible (SC); Amygdaleae), P. mume (mei, SI; Amygdaleae), Fragaria vesca (strawberry, SC; Potentilleae), and F. nipponica (mori-ichigo, SI; Potentilleae). Phylogenetic analyses revealed that the Malus and Prunus S-RNase and S-pollen genes belong to distinct gene lineages, and that only Prunus S-RNase and SFB-lineage genes are present in Fragaria. Thus, S-RNase based GSI system of Malus evolved independently from the ancestral system of Rosaceae. Using expression patterns based on RNA-seq data, the ancestral S-RNase lineage gene is inferred to be expressed in pistils only, while the ancestral S-pollen lineage gene is inferred to be expressed in tissues other than pollen.

Dissection of the style's response to pollination using transcriptome profiling in self-compatible (Solanum pimpinellifolium) and self-incompatible (Solanum chilense) tomato species.

BackgroundTomato (Solanum lycopersicum) self-compatibility (SC) is defined as self-pollen tubes that can penetrate their own stigma, elongate in the style and fertilize their own ovules. Self-incompatibility (SI) is defined as self-pollen tubes that are prevented from developing in the style. To determine the influence of gene expression on style self-pollination, a transcriptome-wide comparative analysis of SC and SI tomato unpollinated/pollinated styles was performed using RNA-sequencing (RNA-seq) data.ResultsTranscriptome profiles of 24-h unpollination (UP) and self-pollination (P) styles from SC and SI tomato species were generated using high-throughput next generation sequencing. From the comparison of SC self-pollinated and unpollinated styles, 1341 differentially expressed genes (DEGs) were identified, of which 753 were downregulated and 588 were upregulated. From the comparison of SI self-pollinated and unpollinated styles, 804 DEGs were identified, of which 215 were downregulated and 589 were upregulated. Nine gene ontology (GO) terms were enriched significantly in SC and 78 GO terms were enriched significantly in SI. A total of 105 enriched Kyoto Encyclopedia of Genes and Genomes (KEGG) pathways were identified in SC and 80 enriched KEGG pathways were identified in SI, among which “Cysteine and methionine metabolism pathway” and “Plant hormone signal transduction pathway” were significantly enriched in SI.ConclusionsThis study is the first global transcriptome-wide comparative analysis of SC and SI tomato unpollinated/pollinated styles. Advanced bioinformatic analysis of DEGs uncovered the pathways of “Cysteine and methionine metabolism” and “Plant hormone signal transduction”, which are likely to play important roles in the control of pollen tubes growth in SI species.Electronic supplementary materialThe online version of this article (doi:10.1186/s12870-015-0492-7) contains supplementary material, which is available to authorized users.

Transitions between self-compatibility and self-incompatibility and the evolution of reproductive isolation in the large and diverse tropical genusDendrobium(Orchidaceae)

The evolution of interspecific reproductive barriers is crucial to understanding species evolution. This study examines the contribution of transitions between self-compatibility (SC) and self-incompatibility (SI) and genetic divergence in the evolution of reproductive barriers in Dendrobium, one of the largest orchid genera. Specifically, it investigates the evolution of pre- and postzygotic isolation and the effects of transitions between compatibility states on interspecific reproductive isolation within the genus. The role of SC and SI changes in reproductive compatibility among species was examined using fruit set and seed viability data available in the literature from 86 species and ∼2500 hand pollinations. The evolution of SC and SI in Dendrobium species was investigated within a phylogenetic framework using internal transcribed spacer sequences available in GenBank. Based on data from crossing experiments, estimations of genetic distance and the results of a literature survey, it was found that changes in SC and SI significantly influenced the compatibility between species in interspecific crosses. The number of fruits produced was significantly higher in crosses in which self-incompatible species acted as pollen donor for self-compatible species, following the SI × SC rule. Maximum likelihood and Bayesian tests did not reject transitions from SI to SC and from SC to SI across the Dendrobium phylogeny. In addition, postzygotic isolation (embryo mortality) was found to evolve gradually with genetic divergence, in agreement with previous results observed for other plant species, including orchids. Transitions between SC and SI and the gradual accumulation of genetic incompatibilities affecting postzygotic isolation are important mechanisms preventing gene flow among Dendrobium species, and may constitute important evolutionary processes contributing to the high levels of species diversity in this tropical orchid group.

Insight into flower diversity inSolanum lycopersicumandSolanum chilenseusing comparative biological approaches

Gao, M.-F., Peng, H.-Z., Li, S.-S., Wang, X.-L., Gao, L., Wang, M.-H., Zhao, P.-F. and Zhao, L.-X. 2015. Insight into flower diversity in Solanum lycopersicum and Solanum chilense using comparative biological approaches. Can. J. Plant Sci. 95: 467–478. Flowers, the defining feature of angiosperms, are reproductive organs that enable the continuation and propagation of flowering plant species. However, intra- and inter-species specific reproductive barriers have often evolved in plants; one such example being in the section Lycopersicum of the genus Solanum. This biological phenomenon is closely related to flower diversity, including visible morphologic features and processes such as pollination and fertilization are influenced by floral morphology, although the exact relationships remains unclear. In this study, two distinct Solanum accessions that exhibit either self-compatibility (SC) (S. lycopersicum, P86) or self-incompatibility (SI) (S. chilense, LA2405), from section Lycopersicum in Solanum were sub...

Breeding systems in Tolpis (Asteraceae) in the Macaronesian islands: the Azores, Madeira and the Canaries

Plants on oceanic islands often originate from self-compatible (SC) colonizers capable of seed set by self-fertilization. This fact is supported by empirical studies, and is rooted in the hypothesis that one (or few) individuals could find a sexual population, whereas two or more would be required if the colonizers were self-incompatible (SI). However, a SC colonizer would have lower heterozygosity than SI colonizers, which could limit radiation and diversification of lineages following establishment. Limited evidence suggests that several species-rich island lineages in the family Asteraceae originated from SI colonizers with some “leakiness” (pseudo-self-compatibility, PSC) such that some self-seed could be produced. This study of Tolpis (Asteraceae) in Macaronesia provides first reports of the breeding system in species from the Azores and Madeira, and additional insights into variation in Canary Islands. Tolpis from the Azores and Madeira are predominately SI but with PSC. This study suggests that the breeding systems of the ancestors were either PSC, possibly from a single colonizer, or from SI colonizers by multiple disseminules either from a single or multiple dispersals. Long-distance colonists capable of PSC combine the advantages of reproductive assurance (via selfing) in the establishment of sexual populations from even a single colonizer with the higher heterozygosity resulting from its origin from an outcrossed source population. Evolution of Tolpis on the Canaries and Madeira has generated diversity in breeding systems, including the origin of SC. Macaronesian Tolpis is an excellent system for studying breeding system evolution in a small, diverse lineage.

Unilateral incompatibility gene ui1.1 encodes an S-locus F-box protein expressed in pollen of Solanum species

Unilateral interspecific incompatibility (UI) is a postpollination, prezygotic reproductive barrier that prevents hybridization between related species when the female parent is self-incompatible (SI) and the male parent is self-compatible (SC). In tomato and related Solanum species, two genes, ui1.1 and ui6.1, are required for pollen compatibility on pistils of SI species or hybrids. We previously showed that ui6.1 encodes a Cullin1 (CUL1) protein. Here we report that ui1.1 encodes an S-locus F-box (SLF) protein. The ui1.1 gene was mapped to a 0.43-cM, 43.2-Mbp interval at the S-locus on chromosome 1, but positional cloning was hampered by low recombination frequency. We hypothesized that ui1.1 encodes an SLF protein(s) that interacts with CUL1 and Skp1 proteins to form an SCF-type (Skp1, Cullin1, F-box) ubiquitin E3 ligase complex. We identified 23 SLF genes in the S. pennellii genome, of which 19 were also represented in cultivated tomato (S. lycopersicum). Data from recombination events, expression analysis, and sequence annotation highlighted 11 S. pennellii genes as candidates. Genetic transformations demonstrated that one of these, SpSLF-23, is sufficient for ui1.1 function. A survey of cultivated and wild tomato species identified SLF-23 orthologs in each of the SI species, but not in the SC species S. lycopersicum, S. cheesmaniae, and S. galapagense, pollen of which lacks ui1.1 function. These results demonstrate that pollen compatibility in UI is mediated by protein degradation through the ubiquitin-proteasome pathway, a mechanism related to that which controls pollen recognition in SI.

The joint evolution and maintenance of self-incompatibility with gynodioecy or androdioecy.

Mating systems show two kinds of frequent transitions: from hermaphroditism to dioecy, gynodioecy or androdioecy, or from self-incompatibility (SI) to self-compatibility (SC). While models have mostly investigated these two kinds of transitions as independent, empirical observations suggest that, to some extent, they can evolve jointly. Here, we study the joint evolution and maintenance of SI and androdioecy or SI and gynodioecy by the means of phenotypic models. Our models focus on three parameters: the unisexuals׳ advantage relative to that of the hermaphrodites due to resource reallocation, inbreeding depression and the selfing rate. We assume no pollen limitation or discounting. We show that SI helps the maintenance of androdioecy, but favors the loss of gynodioecy, and also that androdioecy facilitates the maintenance of SI, whereas gynodioecy does not affect it. We finally investigate how gynodioecy and androdioecy may affect the diversification of SI groups, especially considering an evolutionary pathway through SC intermediates. We show that while androdioecy prevents the increase of the number of SI groups, under certain conditions of inbreeding depression and selfing rates, gynodioecy allows it.

Increased heterosis in selfing populations of a perennial forb.

Quantifying the importance of random genetic drift in natural populations is central to understanding the potential limits to natural selection. One approach is to estimate the magnitude of heterosis, the increased fitness of progeny derived from crosses between populations relative to crosses within populations caused by the heterozygous masking of deleterious recessive or nearly recessive alleles that have been fixed by drift within populations. Self-fertilization is expected to reduce the effective population size by half relative to outcrossing, and population bottlenecks may be common during the transition to selfing. Therefore, chance fixation of deleterious alleles due to drift in selfing populations should increase heterosis between populations. Increased homozygosity due to fixation or loss of alleles should also decrease inbreeding depression within populations. Most populations of the perennial herb Arabidopsis lyrata ssp. lyrata are self-incompatible (SI), but several have evolved self-compatibility and are highly selfing. We quantified heterosis and inbreeding depression in two predominantly self-compatible (SC) and seven SI populations in a field common garden experiment within the species' native range and examined the correlation between these metrics to gauge the similarity in their genetic basis. We measured proportion germination in the lab, and survival and fecundity (flower and seed production) for 2 years in the field, and calculated estimates of cumulative fitness. We found 7.2-fold greater heterosis in SC compared with SI populations, despite substantial heterosis in SI populations (56 %). Inbreeding depression was >61 %, and not significantly different between SC and SI populations. There was no correlation between population estimates of heterosis and inbreeding depression, suggesting that they have somewhat different genetic bases. Combined with other sources of information, our results suggest a history of bottlenecks in all of these populations. The bottlenecks in SC populations may have been severe, but their strong inbreeding depression remains enigmatic.

Gene expression and genetic analysis reveal diverse causes of recessive self-compatibility in Brassica napus L.

BackgroundBrassica napus (AACC) is self-compatible, although its ancestor species Brassica rapa (AA) and Brassica oleracea (CC) are self-incompatible. Most B.napus accessions have dominant self-compatibility (SC) resulting from an insertion of 3.6 kb in the promoter region of BnSCR-1 on the A genome, while recessive SC in B.napus has rarely been observed. Expression and cloning of SRK and SCR genes and genetic analysis were carried out to dissect bases of recessive SC in B.napus.ResultsEleven accessions were screened to identify stable recessive SC and had the S genotype BnS-7 on the A genome and BnS-6 on the C genome similarly to BrS-29 and BoS-15, respectively. In eight SC accessions, BnSCR-7 and BnSCR-6 were nearly undetectable and harbored no structural mutations in the promoters, while SRK genes were expressed at normal levels and contained intact CDS, with the exception of BnSRK-7 in line C32. SRK and SCR genes were expressed normally but their CDSs had no mutations in three SC accessions. In self-incompatible S-1300 and 11 F1 hybrids, SRK genes and BnSCR-1300 transcripts were present at high levels, while expression of the BnSCR-7 and BnSCR-6 were absent. Plants of S genotype S1300S1300 were completely SI, while SI phenotypes of SBnS-7SBnS-7 and S1300SBnS-7 plants were segregated in BC1 and F2 populations.ConclusionsThe recessive SC in eight accessions is caused by the loss of function of BnSCR-7 and BnSCR-6 in pollen. Translational repression contributes to the recessive SC in three accessions, whose SRK and SCR genes were expressed normally and had identical CDSs to BrS-29 or BoS-15. SI in 11 F1 hybrids relies on the expression of BnSCR-1300 rather than SRK genes. Other factor(s) independent of the S locus are involved in recessive SC. Therefore, diverse causes underlie recessive SC in B. napus, yielding insight into these complex mechanisms.Electronic supplementary materialThe online version of this article (doi:10.1186/1471-2164-15-1037) contains supplementary material, which is available to authorized users.

Genetic architecture of inbreeding depression and the maintenance of gametophytic self-incompatibility.

Gametophytic self-incompatibility (GSI) is a widespread genetic system, which enables hermaphroditic plants to avoid self-fertilization and mating with close relatives. Inbreeding depression is thought to be the major force maintaining SI; however, inbreeding depression is a dynamical variable that depends in particular on the mating system. In this article we use multilocus, individual-based simulations to examine the coevolution of SI and inbreeding depression within finite populations. We focus on the conditions for the maintenance of SI when self-compatible (SC) mutants are introduced in the population by recurrent mutation, and compare simulation results with predictions from an analytical model treating inbreeding depression as a fixed parameter (thereby neglecting effects of purging within the SC subpopulation). In agreement with previous models, we observe that the maintenance of SI is associated with high inbreeding depression and is facilitated by high rates of self-pollination. Purging of deleterious mutations by SC mutants has little effect on the spread of those mutants as long as most deleterious alleles have weak fitness effects: in this case, the genetic architecture of inbreeding depression has little effect on the maintenance of SI. By contrast, purging may greatly enhance the spread of SC mutants when deleterious alleles have strong fitness effects.

Effect of enhanced pollination on economic properties of self-compatible loquat (Eriobotrya japonica Lindl.)

Loquat (Eriobotrya japonica Lindl.), a member of the Rosaceae, carries the RNase-dependent gametophytic self-incompatibility (GSI) fertilization system. However, some cultivars, including the Israeli ‘Yehuda’ and ‘Akko 1’, are self-compatible (SC). Since SC cultivars are not dependent on pollen donors, they are usually planted in ‘solid blocks’. Nevertheless, in some SC Rosaceae species, such as apricot, cherry and almond, enhanced self- or cross-pollination increases fruit set. To determine whether enhanced pollination can improve ‘Yehuda’ and ‘Akko 1’ fruit set and fruit size relative to spontaneous self-pollination, the impact of introducing honey bees and the effect of cross-pollination were examined. We found that honey bees are highly attracted to ‘Yehuda’ and ‘Akko 1’ flowers, and fruit size in both cultivars was positively correlated with the number of seeds per fruit. In a net-caged tree experiment, fruit set of ‘Yehuda’ and ‘Akko 1’ benefited from the presence of honey bees. However, fruit set was also relatively high for the caged trees without honey bees which underwent spontaneous self-pollination, and commercial fruit thinning was required for all treatments. For both cultivars, seed set after spontaneous self-pollination was the same as for self-pollination in the presence of honey bees. The impact of cross-pollination between different cultivars moderately increased ‘Yehuda’ fruit set and ‘Akko 1’ seed set. Taken together, the results suggest that spontaneous self-pollination meets the pollination requirements for the SC loquat cultivars ‘Yehuda’ and ‘Akko 1’, and that the presence of pollen donors and/or honey bees confers a minor commercial advantage.