A general stochastic model for sporophytic self-incompatibility

Disentangling the processes leading populations to extinction is a major topic in ecology and conservation biology. The difficulty to find a mate in many species is one of these processes. Here, we investigate the impact of self-incompatibility in flowering plants, where several inter-compatible classes of individuals exist but individuals of the same class cannot mate. We model pollen limitation through different relationships between mate availability and fertilization success. After deriving a general stochastic model, we focus on the simple case of distylous plant species where only two classes of individuals exist. We first study the dynamics of such a species in a large population limit and then, we look for an approximation of the extinction probability in small populations. This leads us to consider inhomogeneous random walks on the positive quadrant. We compare the dynamics of distylous species to self-fertile species with and without inbreeding depression, to obtain the conditions under which self-incompatible species could be less sensitive to extinction while they can suffer more pollen limitation

Reproductive success through high pollinator visitation rates despite self incompatibility in an endangered wallflower

PREMISE OF THE STUDY: Self incompatibility (SI) in rare plants presents a unique challenge—SI protects plants from inbreeding depression, but requires a sufficient number of mates and xenogamous pollination. Does SI persist in an endangered polyploid? Is pollinator visitation sufficient to ensure reproductive success? Is there evidence of inbreeding/outbreeding depression? We characterized the mating system, primary pollinators, pollen limitation, and inbreeding/outbreeding depression in Erysimum teretifolium to guide conservation efforts. METHODS: We compared seed production following self pollination and within- and between-population crosses. Pollen tubes were visualized after self pollinations and between-population pollinations. Pollen limitation was tested in the field. Pollinator observations were quantified using digital video. Inbreeding/outbreeding depression was assessed in progeny from self and outcross pollinations at early and later developmental stages. KEY RESULTS: Self-pollination reduced seed set by 6.5× and quadrupled reproductive failure compared with outcross pollination. Pollen tubes of some self pollinations were arrested at the stigmatic surface. Seed-set data indicated strong SI, and fruit-set data suggested partial SI. Pollinator diversity and visitation rates were high, and there was no evidence of pollen limitation. Inbreeding depression (δ) was weak for early developmental stages and strong for later developmental stages, with no evidence of outbreeding depression. CONCLUSIONS: The rare hexaploid E. teretifolium is largely self incompatible and suffers from late-acting inbreeding depression. Reproductive success in natural populations was accomplished through high pollinator visitation rates consistent with a lack of pollen limitation. Future reproductive health for this species will require large population sizes with sufficient mates and a robust pollinator community

Genetic Control of Self-Incompatibility in Centromadia (Hemizonia) Pungens Subsp. Laevis (Madiinae, Asteraceae)

The presence of self-incompatibility was tested in Centromadia pun gens subsp. laevis and the genetic basis of the self-incompatibility response was explored using crossing studies. We performed full diallel crossing experiments among 10 individuals from one natural population and four F1 families. We observed a strong self-incompatibility response in all individuals tested, with a significant difference in seed set between selfed and outcrossed matings. Most pairwise matings among parental plants were compatible, with some nonreciprocally incompatible matings (i.e., the matings were successful in one direction, but not the other), and only one reciprocally incompatible mating. The full diallel crossing studies among sibs in the four F1 families showed two major compatibility classes. These results are consistent with a single-locus sporophytic self-incompatibility system in this species

The effect of sheltered load on reproduction in Solanum carolinense, a species with variable self-incompatibility

In previous studies, we have investigated the strength of self-incompatibility (SI) in Solanum carolinense, a highly successful weed with a fully functional SI system that inhabits early successional and other disturbed habitats. We have found that the SI response in S. carolinense is a plastic trait—its strength being affected by the age of the flowers, and the presence of developing fruits and that there are genetic differences among families in their self-fertility. However, in species with a fully functional SI response, selfing would not be that common. As a result, deleterious recessives scattered though the genome of horsenettle are only occasionally exposed to selection. It has been suggested that deleterious recessives accumulate near S-alleles in strong SI species because the S-locus is located in a non-recombining region of the genome and because strong S-alleles are never in the homozygous state, thus sheltering some of the genetic load near the S-locus from selection. We performed a series of laboratory and greenhouse experiments to determine the extent to which sheltered load adds to the overall magnitude of inbreeding depression in horsenettle. Specifically, we amplified and sequenced the S-alleles from 16 genets collected from a large population in Pennsylvania and performed a series of controlled self-pollinations. We then grew the selfed progeny in the greenhouse; recorded various measures of growth and reproductive output; and amplified and sequenced their S-allele(s). We found that the heterozygous progeny of self-pollinations produce more flowers and have a greater ability to set both self and cross seed than S-homozygous progeny. We also found evidence of variation in the magnitude of load among S-alleles. These results suggest that sheltered load might slow the fixation of weak (partially compatible) S-alleles in this population, thus adding to the maintenance of a mixed mating system rather than leading to the fixation of the selfing allele

Depresi Silang dalam Biji dan Semai Melaleuca Alternifolia

Tea tree (Melaleuca alternifolia) is known as an essential oil producer. This species is commercially important as a source of essential oil especially in Australia. To improve its oil quality, research on artificial pollination between plus trees were carried out. The information about self-incompatibility and inbreeding depression due to self-pollination should be ascertained to support the pollination success. This study was aimed at examining the self-incompatibility of tea tree by conducting controlled self-pollination and cross-pollination. Controlled cross and self-pollination were carried out at a seedling seed orchard of tea tree, using four mother trees as experimental samples. Natural self-pollination was undertaken by bagging un opened flowers, without emasculation and counting the number of capsule set. Open pollination was used as a control treatment, of which the number of unopened flowers and the number of capsule set were counted. Observations revealed that no capsule was found from controlled self pollination (with the index self-incompatibility = 0), even though natural self-pollination produced low number of capsules (with the index self-incompatibility = 0.24). Therefore, it can be assumed that high level of self-incompatibility was took place in tea tree. Inbreeding depression also existed in this species, revealed by the decreased number of capsul set, lower seed germination rate of self-pollinated seeds than cross pollinated seeds. The slower growth of self-pollinated seedlings than cross pollinated seedlings in the nursery also indicated that inbreeding depression occured

Expression and trans-specific polymorphism of self-incompatibility RNases in Coffea (Rubiaceae)

Self-incompatibility (SI) is widespread in the angiosperms, but identifying the biochemical components of SI mechanisms has proven to be difficult in most lineages. Coffea (coffee; Rubiaceae) is a genus of old-world tropical understory trees in which the vast majority of diploid species utilize a mechanism of gametophytic self-incompatibility (GSI). The S-RNase GSI system was one of the first SI mechanisms to be biochemically characterized, and likely represents the ancestral Eudicot condition as evidenced by its functional characterization in both asterid (Solanaceae, Plantaginaceae) and rosid (Rosaceae) lineages. The S-RNase GSI mechanism employs the activity of class III RNase T2 proteins to terminate the growth of "self" pollen tubes. Here, we investigate the mechanism of Coffea GSI and specifically examine the potential for homology to S-RNase GSI by sequencing class III RNase T2 genes in populations of 14 African and Madagascan Coffea species and the closely related self-compatible species Psilanthus ebracteolatus. Phylogenetic analyses of these sequences aligned to a diverse sample of plant RNase T2 genes show that the Coffea genome contains at least three class III RNase T2 genes. Patterns of tissue-specific gene expression identify one of these RNase T2 genes as the putative Coffea S-RNase gene. We show that populations of SI Coffea are remarkably polymorphic for putative S-RNase alleles, and exhibit a persistent pattern of trans-specific polymorphism characteristic of all S-RNase genes previously isolated from GSI Eudicot lineages. We thus conclude that Coffea GSI is most likely homologous to the classic Eudicot S-RNase system, which was retained since the divergence of the Rubiaceae lineage from an ancient SI Eudicot ancestor, nearly 90 million years ago.United States National Science Foundation [0849186]; Society of Systematic Biologists; American Society of Plant Taxonomists; Duke University Graduate Schoolinfo:eu-repo/semantics/publishedVersio

Recent and Ancient Signature of Balancing Selection around the S-Locus in Arabidopsis halleri and A. lyrata

Balancing selection can maintain different alleles over long evolutionary times. Beyond this direct effect on the molecular targets of selection, balancing selection is also expected to increase neutral polymorphism in linked genome regions, in inverse proportion to their genetic map distances from the selected sites. The genes controlling plant self-incompatibility are subject to one of the strongest forms of balancing selection, and they show clear signatures of balancing selection. The genome region containing those genes (the S-locus) is generally described as nonrecombining, and the physical size of the region with low recombination has recently been established in a few species. However, the size of the region showing the indirect footprints of selection due to linkage to the S-locus is only roughly known. Here, we improved estimates of this region by surveying synonymous polymorphism and estimating recombination rates at 12 flanking region loci at known physical distances from the S-locus region boundary, in two closely related self-incompatible plants Arabidopsis halleri and A. lyrata. In addition to studying more loci than previous studies and using known physical distances, we simulated an explicit demographic scenario for the divergence between the two species, to evaluate the extent of the genomic region whose diversity departs significantly from neutral expectations. At the closest flanking loci, we detected signatures of both recent and ancient indirect effects of selection on the S-locus flanking genes, finding ancestral polymorphisms shared by both species, as well as an excess of derived mutations private to either species. However, these effects are detected only in a physically small region, suggesting that recombination in the flanking regions is sufficient to quickly break up linkage disequilibrium with the S-locus. Our approach may be useful for distinguishing cases of ancient versus recently evolved balancing selection in other systems

DNA-based S-genotyping of Japanese plum and pluot cultivars to clarify incompatibility relationships

Diploid japanese plum (Prunus salicina Lindl.) cultivars are commonly self-incompatible. To date, 14 incompatibility alleles (S-alleles) have been identified and labeled with alphabetical (S-a-S-n) and 5 with numeric codes (S-1, S-3-S-6). We applied polymerase chain reaction amplification of the S-RNase alleles with degenerate and allele-specific primers in 10 japanese plum cultivars and two pluots of unknown incompatibility alleles. Besides DNA sequencing, an additional method for the exact length determination of the first intron region was used for the first time for S-genotype japanese plums. The S-3-allele was shown to correspond to S-k in the alphabetic nomenclature, S-4 to S-c, S-5 to S-e, and S-6 to S-f. The S-5-allele-specific primer can be used as a reliable marker for self-compatibility in japanese plum. 'Black Amber', 'October Sun', 'TC Sun', and 'Super Giant' share the SbSc genotype, which was confirmed by test crosses. These cultivars belong to the widest incompatibility group currently known in japanese plum. An additional incompatibility group (ScSh) was established, including 'Green Sun' and 'Queen Rosa', a cultivar formerly known as a universal donor. By incorporating all previous and recent results, a table was assembled including 49 cultivars assigned to I-VII incompatibility groups, to the self-compatible group and to the group O of unique genotypes. These data may considerably contribute to further growing and breeding activities

Towards F1 Hybrid Seed Potato Breeding

Compared to other major food crops, progress in potato yield as the result of breeding efforts is very slow. Genetic gains cannot be fixed in potato due to obligatory out-breeding. Overcoming inbreeding depression using diploid self-compatible clones should enable to replace the current method of out-breeding and clonal propagation into an F1 hybrid system with true seeds. This idea is not new, but has long been considered unrealistic. Severe inbreeding depression and self-incompatibility in diploid germplasm have hitherto blocked the development of inbred lines. Back-crossing with a homozygous progenitor with the Sli gene which inhibits gametophytic self-incompatibility gave self-compatible offspring from elite material from our diploid breeding programme. We demonstrate that homozygous fixation of donor alleles is possible, with simultaneous improvement of tuber shape and tuber size grading of the recipient inbred line. These results provide proof of principle for F1 hybrid potato breeding. The technical and economic perspectives are unprecedented as these will enable the development of new products with combinations of useful traits for all stakeholders in the potato chain. In addition, the hybrid’s seeds are produced by crossings, rendering the production and voluminous transport of potato seed tubers redundant as it can be replaced by direct sowing or the use of healthy mini-tubers, raised in greenhouses