S-RNases and sexual incompatibility: structure, functions, and evolutionary perspectives

S-RNase-based gametophytic self-incompatibility appears to be the most phylogenetically widespread form of self-incompatibility found in the angiosperms, having been reported in the Solanaceae, Scrophulariaceae, and Rosaceae. This intraspecific breeding barrier is controlled by a single genetic locus termed S. Rejection of self-pollen has been shown to be mediated in the pistil by a highly polymorphic series of ribonucleases, but as yet the pollen component of this recognition system has not been identified. Here we review our present knowledge concerning the structure, functions, and evolution of S-RNases and the S-loci in which they reside. In addition we present two new phylogenetic analyses of S-RNases which suggest that (1). sequence variability between S-alleles is spread across the whole gene and is not as clustered as is generally believed and (2). there is evidence of recombination and/or diversifying selection in two distinct regions of S-RNases. The implications of these findings are discussed.     

The effects of population size limitation on fecundity in mosaic populations of the clonal macrophyte Scirpus maritimus (Cyperaceae)

The clonal macrophyte Scirpus maritimus (Cyperaceae) propagates locally by rhizomes and reproduces sexually by achenes. The purpose of this paper was to examine whether in size-limited habitats in patchy and discrete marshes in two Mediterranean wetlands in southern France natural populations may suffer from a reduced maternal fecundity due to a deficit in outcross pollen. We first verified that S. maritimus suffers from a reduced fecundity when self-pollinated. At a site in the Camargue, mean fecundity (mean number of achenes per centimetre of spikelet) measured in 1995 and 1996 in seven and nine populations, respectively (surface area from 50 to 4500 m) increased significantly with population surface area in 1995 but not in 1996. In the second wetland at Roquehaute, which is composed of small ponds, fecundity was very low in all 12 local populations studied in 1996 (1.1 achenes per spikelet, SD = 1.2) and was not correlated with the population surface area (from 10 to 400 m). We performed a pollen supplementation experiment in five local populations at Roquehaute to determine whether this low fecundity may be due to a pollen limitation. A significant increase in fecundity after among-pond pollinations compared to within-pond pollinations indicated that local populations suffer from a deficit in outcross pollen, since each pond appears to contain one or a few number of clones (or incompatibility types). In S. maritimus, clonal spread may have a cost in terms of reduced fecundity in small habitats because each habitat is colonized by very few clones.     

Antagonism between local dispersal and self-incompatibility systems in a continuous plant population

Many self-incompatible plant species exist in continuous populations in which individuals disperse locally. Local dispersal of pollen and seeds facilitates inbreeding because pollen pools are likely to contain relatives. Self-incompatibility promotes outbreeding because relatives are likely to carry incompatible alleles. Therefore, populations can experience an antagonism between these forces. In this study, a novel computational model is used to explore the effects of this antagonism on gene flow, allelic diversity, neighbourhood sizes, and identity by descent. I confirm that this antagonism is sensitive to dispersal levels and linkage. However, the results suggest that there is little to no difference between the effects of gametophytic and sporophytic self-incompatibility systems (GSI and SSI) on unlinked loci. More importantly, both GSI and SSI affect unlinked loci in a manner similar to obligate outcrossing without mating types. This suggests that the primary evolutionary impact of self-incompatibility systems may be to prevent selfing, and prevention of biparental inbreeding might be a beneficial side-effect.     

Pollination and late-acting self-incompatibility in Cyrtanthus breviflorus (Amaryllidaceae): implications for seed production

Background and Aims

Animal pollination is typically an uncertain process that interacts with self-incompatibility status to determine reproductive success. Seed set is often pollen-limited, but species with late-acting self-incompatibility (SI) may be particularly vulnerable, if self-pollen deposition results in ovule discounting. Pollination is examined and the occurrence of late-acting SI and ovule discounting assessed in Cyrtanthus breviflorus.

Methods

The pollination system was characterized by observing floral visitors and assessing nectar production and spectral reflectance of flowers. To assess late-acting SI and ovule discounting, growth of self- and cross-pollen tubes, and seed set following open pollination or hand pollination with varying proportions of self- and cross-pollen, were examined.

Key Results

Native honeybees Apis mellifera scutellata pollinated flowers as they actively collected pollen. Most flowers (≥70 %) did not contain nectar, while the rest produced minute volumes of dilute nectar. The flowers which are yellow to humans are visually conspicuous to bees with a strong contrast between UV-reflecting tepals and UV-absorbing anthers and pollen. Plants were self-incompatible, but self-rejection was late-acting and both self- and cross-pollen tubes penetrated ovules. Seed set of open-pollinated flowers was pollen-limited, despite pollen deposition exceeding ovule number by 6-fold. Open-pollinated seed set was similar to that of the cross + self-pollen treatment, but was less than that of the cross-pollen-only treatment.

Conclusions

Flowers of C. breviflorus are pollinated primarily by pollen-collecting bees and possess a late-acting SI system, previously unknown in this clade of the Amaryllidaceae. Pollinators of C. breviflorus deposit mixtures of cross- and self-pollen and, because SI is late-acting, self-pollen disables ovules, reducing female fertility. This study thus contributes to growing evidence that seed production in plants with late-acting SI systems is frequently limited by pollen quality, even when pollinators are abundant.     

Dominance Relationship between Two Self-Incompatible<Emphasis Type="Italic">Brassica campestris</Emphasis> Haplotypes in Response to CO<Subscript>2</Subscript> Gas

InBrassica, self-incompatibility (SI) can be overcome by CO₂ application, an effective method for obtaining numerous inbred lines for F, commercial seed. We previously reported two different S-alleles ofBrassica campestris, S⁷³³ and S⁷³⁴, with extremely different degrees of susceptibility to this gas. In the current study, we raised a cross-population between those two genetic lines, and analyzed their reaction level of self-incompatibility to CO₂ (RLSICO₂). Here, all 40 of our progeny from the F₁ cross-population were susceptible, maintaining high values of RLSICO₂. This suggests that the susceptible line, S⁷³⁴, is dominant to the insusceptible line, S⁷³³. We also generated an F₂ selfing-population of each crossed progeny, S⁷³³♀ S⁷³⁴♂ and S⁷³³♂ S⁷³⁴♀, to assess the RLSICO₂ of each individual. PCR-RFLP analysis was performed to determine the S-genotype of the F₂ population. The S⁷³⁴ allele segregated in a theoretical ratio of the dominant trait, and the RLSICO₂ was consistent with the dominance relationship. Therefore, we have now demonstrated that high RLSICO₂ in β.campestris is controlled by a dominant gene. Both authors contributed equally to this work     

Identification of <Emphasis Type="Italic">S</Emphasis>-haplotype-specific F-box gene in Japanese plum (<Emphasis Type="Italic">Prunus salicina</Emphasis> Lindl.)

Self-incompatibility has been studied extensively at the molecular level in Solanaceae, Rosaceae and Scrophulariaceae, all of which exhibit gametophytic self-incompatibility controlled by a single polymorphic locus containing at least two linked genes, i.e., the S-RNase gene and the pollen-expressed SFB/SLF (S-haplotype-specific F-box/S-locus F-box) gene. However, the SFB gene in Japanese plum (Prunus salicina Lindl.) has not yet been identified. We determined eight novel sequences homologous to the SFB genes of other Prunus species and named these sequences PsSFB. The gene structure of the SFB genes and the characteristic domains in deduced amino acid sequences were conserved. Three sequences from 410 to 2,800 bp of the intergenic region between the PsSFB sequences and the S-RNase alleles were obtained. The eight identified PsSFB sequences showed S-haplotype-specific polymorphism, with 74–83% amino acid identity. These alleles were exclusively expressed in the pollen. These results suggest that the PsSFB alleles are the pollen S-determinants of GSI in Japanese plum. Nucleotide sequence data reported are available in the NCBI database under the accession numbers DQ849084–DQ849090 and DQ849118.     

Physical size of the <Emphasis Type="Italic">S</Emphasis> locus region defined by genetic recombination and genome sequencing in <Emphasis Type="Italic">Ipomoea trifida</Emphasis>, Convolvulaceae

Sporophytic self-incompatibility (SSI) in the genus Ipomoea (Convolvulaceae) is controlled by a single polymorphic S locus. We have previously analyzed genomic sequences of an approximately 300 kb region spanning the S locus of the S ₁ haplotype and characterized the genomic structure around this locus. Here, we further define the physical size of the S locus region by mapping recombination breakpoints, based on sequence analysis of PCR fragments amplified from the genomic DNA of recombinants. From the recombination analysis, the S locus of the S ₁ haplotype was delimited to a 0.23 cM region of the linkage map, which corresponds to a maximum physical size of 212 kb. To analyze differences in genomic organization between S haplotypes, fosmid contigs spanning approximately 67 kb of the S ₁₀ haplotype were sequenced. Comparison with the S ₁ genomic sequence revealed that the S haplotype-specific divergent regions (SDRs) spanned 50.7 and 34.5 kb in the S ₁ and S ₁₀ haplotypes, respectively and that their flanking regions showed a high sequence similarity. In the sequenced region of the S ₁₀ haplotype, five of the 12 predicted open reading frames (ORFs) were found to be located in the divergent region and showed co-linear organization of genes between the two S haplotypes. Based on the size of the SDRs, the physical size of the S locus was estimated to fall within the range 34–50 kb in Ipomoea.     

24份甜樱桃材料自交不亲和S基因型鉴定

甜樱桃品种绝大部分自交不亲和,限制了甜樱桃的正确评价和合理利用,因此自交不亲和基因型的鉴定对于生产具有重要意义。以24个甜樱桃主栽品种为材料,用5对蔷薇科李属引物组合对24个甜樱桃品种进行了S等位基因的PCR扩增,克隆S基因的扩增片段,用核酸序列在Gen Bank上搜索,确定了5种S基因的核酸序列和大小。结果表明:Pru C2+Pru C4R引物组合扩增效果最好;在琼脂糖凝胶上位置相同的扩增带其核酸序列相同,是同一种S基因;5种S基因扩增片段的大小分别是S1为800 bp,S3为762 bp,S4为962bp,S5为300 bp,S6为456 bp,S9为650 bp;24个甜樱桃S基因型是红手球、早红宝石为S1S3,拉宾斯S1S4',红宝石S1S6,布鲁克斯S1S9,那翁S3S4,秦林、泰安大紫、先锋、早大果、丽珠、美早、5-106、左滕锦、桑提娜为S3S6,黑珍珠、红灯、萨米脱、秦樱为S3S9,胜利为S5S9,明珠、红蜜、雷尼、滨库为S6S9。