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Text S1. Details on methodology. Table S1. Details on provenance of Pilosella rhodopea material used for particular analyses. Table S2. Dataset for 63 plants of Pilosella rhodopea included in the greenhouse experiment. Figure S1. Well-developed and aborted seeds of Pilosella rhodopea. Figure S2. Habitus and vegetative growth characteristics of diploid and autopolyploid plants of Pilosella rhodopea. Figure S3. Percentage of euploid and aneuploid seed progeny produced by diploid, autotriploid and autotetraploid plants of Pilosella rhodopea in natural primary contact zones. Figure S4. Photomicrographs of somatic metaphases in two aneuploid Pilosella rhodopea progeny produced by two triploid (2n = 27) mother plants. Figure S5. Flow cytometric screening analyses of seeds produced by diploid and autopolyploid plants of Pilosella rhodopea.

Apomixis, the asexual formation of seeds, is a potentially valuable agricultural trait. Inducing apomixis in sexual crop plants would, for example, allow breeders to fix heterosis in hybrid seeds and rapidly generate doubled haploid crop lines. Molecular models explain the emergence of functional apomixis, i.e., apomeiosis + parthenogenesis + endosperm development, as resulting from a combination of genetic or epigenetic changes that coordinate altered molecular and developmental steps to form clonal seeds. Apomixis-like features and synthetic clonal seeds have been induced with limited success in the sexual plants rice and maize by using gene editing to mutate genes related to meiosis and fertility or via egg-cell specific expression of embryogenesis genes. Inducing functional apomixis and increasing the penetrance of apomictic seed production will be important for commercial deployment of the trait. Optimizing the induction of apomixis with gene editing strategies that use known targets as well as identifying alternative targets will be possible by better understanding natural genetic variation in apomictic species. With the growing availability of genomic data and precise gene editing tools, we are making substantial progress towards engineering apomictic crops. Open-Access-Publikationsfonds 2020 peerReviewed

Projections indicate that current plant breeding approaches will be unable to incorporate the global crop yields needed to deliver global food security. Apomixis is a disruptive innovation by which a plant produces clonal seeds capturing heterosis and gene combinations of elite phenotypes. Introducing apomixis into hybrid cultivars is a game-changing development in the current plant breeding paradigm that will accelerate the generation of high-yield cultivars. However, apomixis is a developmentally complex and genetically multifaceted trait. The central problem behind current constraints to apomixis breeding is that the genomic configuration and molecular mechanism that initiate apomixis and guide the formation of a clonal seed are still unknown. Today, not a single explanation about the origin of apomixis o er full empirical coverage, and synthesizing apomixis by manipulating individual genes has failed or produced little success. Overall evidence suggests apomixis arise from a still unknown single event molecular mechanism with multigenic e ects. Disentangling the genomic basis and complex genetics behind the emergence of apomixis in plants will require the use of novel experimental approaches benefiting from Next Generation Sequencing technologies and targeting not only reproductive genes, but also the epigenetic and genomic configurations associated with reproductive phenotypes in homoploid sexual and apomictic carriers. A comprehensive picture of most regulatory changes guiding apomixis emergence will be central for successfully installing apomixis into the target species by exploiting genetic modification techniques. Open-Access-Publikationsfonds 2020 peerReviewed

With ever-rising demand for food, forage breeding for intensification of cattle production is also taking a leap. In South America, cattle production systems are displaced to marginal areas poorly exploited with cultivated pastures yet with high potential for growing stocking rates. This places the need for using native genetic resources to breed locally adapted plant genotypes that benefits from better forage quality, yield, and lesser threat to the local biodiversity. Paspalum modestum Mez is a grass species that produces quality forage and grows in marginal areas like estuaries and floodplains, suitable for introduction in breeding programs. In this study we characterize the species' reproductive biology and ecological preferences needed beforehand any improvement. P. modestum plants found in nature are commonly diploids, rarely triploids, and tetraploids. Chromosome associations during meiosis in polyploids indicate they are autopolyploids. While diploids are sexual self-sterile, analyses of embryology, gamete fertility and experimental crossings show tetraploids are self-compatible facultative apomicts, highly fertile and have a high proportion of sexuality compared to other apomictic species. Ecological niche analysis and species distribution modelling show mean annual temperature and precipitation as main ecological drivers and a wide geographical area of climatic suitability where P. modestum can grow and be exploited as a forage grass. Our study points to P. modestum as a native plant resource appropriate for breeding waterlogging tolerant ecotypes and genotypes of high biomass production adapted to low flow areas in the Subtropics of Brazil, Paraguay, Uruguay and Argentina.

Model of mutation elimination under constant recombination rate and selection on female gametophytes. The graph visualizes the model for the first generations. (DOCX 264 kb)

Background In the absence of sex and recombination, genomes are expected to accumulate deleterious mutations via an irreversible process known as Muller’s ratchet, especially in the case of polyploidy. In contrast, no genome-wide mutation accumulation was detected in a transcriptome of facultative apomictic, hexaploid plants of the Ranunculus auricomus complex. We hypothesize that mutations cannot accumulate in flowering plants with facultative sexuality because sexual and asexual development concurrently occurs within the same generation. We assume a strong effect of purging selection on reduced gametophytes in the sexual developmental pathway because previously masked recessive deleterious mutations would be exposed to selection. Results We test this hypothesis by modeling mutation elimination using apomictic hexaploid plants of the R. auricomus complex. To estimate mean recombination rates, the mean number of recombinants per generation was calculated by genotyping three F1 progeny arrays with six microsatellite markers and character incompatibility analyses. We estimated the strength of purging selection in gametophytes by calculating abortion rates of sexual versus apomictic development at the female gametophyte, seed and offspring stage. Accordingly, we applied three selection coefficients by considering effects of purging selection against mutations on (1) male and female gametophytes in the sexual pathway (additive, s = 1.000), (2) female gametophytes only (s = 0.520), and (3) on adult plants only (sporophytes, s = 0.212). We implemented recombination rates into a mathematical model considering the three different selection coefficients, and a genomic mutation rate calculated from genome size of our plants and plant-specific mutation rates. We revealed a mean of 6.05% recombinants per generation. This recombination rate eliminates mutations after 138, 204 or 246 generations, depending on the respective selection coefficients (s = 1.000, 0.520, and 0.212). Conclusions Our results confirm that the empirically observed frequencies of facultative recombination suffice to prevent accumulation of deleterious mutations via Muller’s ratchet even in a polyploid genome. The efficiency of selection is in flowering plants strongly increased by acting on the haplontic (reduced) gametophyte stage. Open-Access-Publikationsfonds 2019