Although reproductive assurance has been suggested to be one of the most important factors shaping the differential distributional patterns between sexuals and asexuals (geographic parthenogenesis), it has only rarely been studied in natural populations of vascular plants with autonomous apomixis. Moreover, there are almost no data concerning the putative relationship between the level of apomictic versus sexual plant reproduction on one hand, and reproductive assurance on the other. We assessed the level of sexual versus apomictic reproduction in diploid and triploid plants of Hieracium alpinum across its distributional range using flow cytometric analyses of seeds, and compared the level of potential and realized seed set, i.e. reproductive assurance, between the two cytotypes under field and greenhouse conditions. Flow cytometric screening of embryos and endosperms of more than 4,100 seeds showed that diploids produced solely diploid progeny sexually, while triploids produced triploid progeny by obligate apomixis. Potential fruit set was much the same in diploids and triploids from the field and the greenhouse experiment. While in the pollination-limited environment in the greenhouse apomictic triploids had considerably higher realized fruit set than sexual diploids, there was no significant difference between cytotypes under natural conditions. In addition, sexuals varied to a significantly larger extent in realized fruit set than asexuals under both natural and greenhouse conditions. Our results indicate that triploid plants reproduce by obligate apomixis, assuring more stable and predictable fruit reproduction when compared to sexual diploids. This advantage could provide apomictic triploids with a superior colonisation ability, mirrored in a strong geographic parthenogenesis pattern observed in this species.
Gametophytic apomixis is a way of asexual plant reproduction by seeds. It should be advantageous under stressful high altitude or latitude environment where short growing seasons, low temperatures, low pollinator activity or unstable weather may hamper sexual reproduction. However, this hypothesis remains largely untested. Here, we assess the reproductive mode in 257 species belonging to 45 families from the world's broadest alpine belt (2800-6150 m) in NW Himalayas using flow cytometric seed screen. We found only 12 apomictic species, including several members of Poaceae (Festuca, Poa and Stipa), Rosaceae (Potentilla) and Ranunculaceae (Halerpestes, Ranunculus), which are families typical for high apomict frequency. However, several apomictic species were newly discovered, including the first known apomictic species from the family Biebersteiniaceae (Biebersteinia odora), and first apomicts from the genera Stipa (Stipa splendens) and Halerpestes (Halerpestes lancifolia). Apomicts showed no preference for higher elevations, even in these extreme Himalayan alpine habitats. Additional trait-based analyses revealed that apomicts differed from sexuals in comprising more rhizomatous graminoids and forbs, higher soil moisture demands, sharing the syndrome of dominant species with broad geographical and elevation ranges typical for the late-successional habitats. Apomicts differ from non-apomicts in greater ability of clonal propagation and preference for wetter, more productive habitats.
- MeSH
- apomixie * MeSH
- ekosystém * MeSH
- fylogeneze MeSH
- fyziologie rostlin * MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Background and Aims: Apomixis or asexual seed reproduction is a key evolutionary mechanism in certain angiosperms providing them with reproductive assurance and isolation. Nevertheless, the frequency of apomixis is largely unknown, especially in groups with autonomous apomixis such as the diploid-polyploid genus Hieracium. Methods: Using flow cytometric analyses, we determined the ploidy level and reproductive pathways (sexual vs. apomictic) for 7616 seeds originating from 946 plants belonging to >50 taxa sampled at 130 sites across Europe. Key Results: Diploid seeds produced by diploids were formed exclusively by the sexual pathway after double fertilization of reduced embryo sacs. An absolute majority of tri- and tetraploid seeds (99.6 %) produced by tri- and tetrapolyploid taxa were formed by autonomous apomixis. Only 20 polyploid seeds (0.4 %) were formed sexually. These seeds, which originated on seven polyploid accessions of four taxa, were formed after fertilization of either unreduced embryo sacs through a so-called triploid bridge or reduced embryo sacs, and frequently resulted in progeny with an increased ploidy. In addition, the formation of seedlings with increased ploidy (4x and 6x) was found in two triploid plants. This is the first firm evidence on functional facultative apomixis in polyploid members of Hieracium sensu stricto (s.s.). Conclusions: The mode of reproduction in Hieracium s.s. is tightly associated with ploidy. While diploids produce seeds exclusively sexually, polyploids produce seeds by obligate or almost obligate apomixis. Strict apomixis can increase the reproductive assurance and this in turn can increase the colonization ability of apomicts. Nevertheless, our data clearly show that certain polyploid plants are still able to reproduce sexually and contribute to the formation of new cytotypes and genotypes. The finding of functional facultative apomicts is essential for future studies focused on evolution, inheritance and ecological significance of apomixis in this genus.
Background and Aims: Rubus subgenus Rubus is a group of mostly apomictic and polyploid species with a complicated taxonomy and history of ongoing hybridization. The only polyploid series with prevailing sexuality is the series Glandulosi , although the apomictic series Discolores and Radula also retain a high degree of sexuality, which is influenced by environmental conditions and/or pollen donors. The aim of this study is to detect sources of genetic variability, determine the origin of apomictic taxa and validate microsatellite markers by cloning and sequencing. Methods: A total of 206 individuals from two central European regions were genotyped for 11 nuclear microsatellite loci and the chloroplast trn L- trn F region. Microsatellite alleles were further sequenced in order to determine the exact repeat number and to detect size homoplasy due to insertions/deletions in flanking regions. Key Results: The results confirm that apomictic microspecies of ser. Radula are derived from crosses between sexual series Glandulosi and apomictic series Discolores , whereby the apomict acts as pollen donor. Each apomictic microspecies is derived from a single distinct genotype differing from the parental taxa, suggesting stabilized clonal reproduction. Intraspecific variation within apomicts is considerably low compared with sexual series Glandulosi , and reflects somatic mutation accumulation. While facultative apomicts produce clonal offspring, sexual species are the conduits of origin for new genetically different apomictic lineages. Conclusions: One of the main driving forces of evolution and speciation in the highly apomictic subgenus Rubus in central Europe is sexuality in the series Glandulosi . Palaeovegetation data suggest that initial hybridizations took place over different time periods in the two studied regions, and that the successful origin and spread of apomictic microspecies of the series Radula took place over several millennia. Additionally, the cloning and sequencing show that standard evaluations of microsatellite repeat numbers underestimate genetic variability considering homoplasy in allele size.
- MeSH
- apomixie * MeSH
- DNA chloroplastová genetika MeSH
- hybridizace genetická * MeSH
- mikrosatelitní repetice * MeSH
- mutace INDEL MeSH
- polyploidie MeSH
- Rubus klasifikace genetika MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
- Geografické názvy
- Evropa MeSH
Biological invasions can be associated with shifts of the species' climatic niches but the incidence of such shifts is under debate. The reproductive system might be a key factor controlling such shifts because it influences a species' evolutionary flexibility. However, the link between reproductive systems and niche dynamics in plant invasions has been little studied so far. We compiled global occurrence data sets of 13 congeneric sexual and apomictic species pairs, and used principal components analysis (PCA) and kernel smoothers to compare changes in climatic niche optima, breadths and unfilling/expansion between native and alien ranges. Niche change metrics were compared between sexual and apomictic species. All 26 species showed changes in niche optima and/or breadth and 14 species significantly expanded their climatic niches. However, we found no effect of the reproductive system on niche dynamics. Instead, species with narrower native niches showed higher rates of niche expansion in the alien ranges. Our results suggest that niche shifts are frequent in plant invasions but evolutionary potential may not be of major importance for such shifts. Niche dynamics rather appear to be driven by changes of the realized niche without adaptive change of the fundamental climatic niche.
BACKGROUND AND AIMS: In ferns, apomixis is an important mode of asexual reproduction. Although the mechanisms of fern reproduction have been studied thoroughly, most previous work has focused on cases in which ferns reproduce either exclusively sexually or exclusively asexually. Reproduction of ferns with potentially mixed systems and inheritance of apomixis remains largely unknown. This study addresses reproduction of the pentaploid Dryopteris × critica, a hybrid of triploid apomictic D. borreri and tetraploid sexual D. filix-mas. METHODS: Spore size, abortion percentage and number of spores per sporangium were examined in pentaploid plants of D. × critica grown in an experimental garden. The sporangial content of leaf segments was cultivated on an agar medium, and DNA ploidy levels were estimated by DAPI flow cytometry in 259 gametophytes or sporophytes arising from the F2 generation of the pentaploid hybrid. KEY RESULTS: The hybrid is partly fertile (89-94% of aborted spores) and shows unstable sporogenesis with sexual and apomictic reproduction combined. The number of spores per sporangium varied from approx. 31 to 64. Within a single sporangium it was possible to detect formation of either only aborted spores or various mixtures of aborted and well-developed reduced spores and unreduced diplospores. The spores germinated in viable gametophytes with two ploidy levels: pentaploid (5x, from unreduced spores) and half of that (approx. 2·5x, from reduced spores). Moreover, 2-15% of gametophytes (both 2·5x and 5x) formed a viable sporophyte of the same ploidy level due to apogamy. CONCLUSIONS: This study documents the mixed reproductive mode of a hybrid between apomictic and sexual ferns. Both sexual reduced and apomictic unreduced spores can be produced by a single individual, and even within a single sporangium. Both types of spores give rise to viable F2 generation gametophytes and sporophytes.
- MeSH
- apomixie * MeSH
- délka genomu MeSH
- DNA rostlinná metabolismus MeSH
- Dryopteris genetika fyziologie MeSH
- gametogeneze rostlin genetika fyziologie MeSH
- genom rostlinný MeSH
- haploidie * MeSH
- klíčení MeSH
- křížení genetické MeSH
- průtoková cytometrie MeSH
- spory cytologie fyziologie MeSH
- tetraploidie * MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
The Hieracium and Pilosella (Lactuceae, Asteraceae) genera of closely related hawkweeds contain species with two different modes of gametophytic apomixis (asexual seed formation). Both genera contain polyploid species, and in wild populations, sexual and apomictic species co-exist. Apomixis is known to co-exist with sexuality in apomictic Pilosella individuals, however, apomictic Hieracium have been regarded as obligate apomicts. Here, a developmental analysis of apomixis within 16 Hieracium species revealed meiosis and megaspore tetrad formation in 1 to 7% of ovules, for the first time indicating residual sexuality in this genus. Molecular markers linked to the two independent, dominant loci LOSS OF APOMEIOSIS (LOA) and LOSS OF PARTHENOGENESIS (LOP) controlling apomixis in Pilosella piloselloides subsp. praealta were screened across 20 phenotyped Hieracium individuals from natural populations, and 65 phenotyped Pilosella individuals from natural and experimental cross populations, to examine their conservation, inheritance and association with reproductive modes. All of the tested LOA and LOP-linked markers were absent in the 20 Hieracium samples irrespective of their reproductive mode. Within Pilosella, LOA and LOP-linked markers were essentially absent within the sexual plants, although they were not conserved in all apomictic individuals. Both loci appeared to be inherited independently, and evidence for additional genetic factors influencing quantitative expression of LOA and LOP was obtained. Collectively, these data suggest independent evolution of apomixis in Hieracium and Pilosella and are discussed with respect to current knowledge of the evolution of apomixis.
- MeSH
- apomixie genetika MeSH
- Asteraceae genetika MeSH
- biologická evoluce * MeSH
- DNA chloroplastová genetika MeSH
- DNA rostlinná genetika MeSH
- genetické lokusy * MeSH
- genetické markery MeSH
- haplotypy MeSH
- konzervovaná sekvence MeSH
- molekulární sekvence - údaje MeSH
- populační genetika MeSH
- regulace genové exprese u rostlin MeSH
- rostlinné geny * MeSH
- semena rostlinná genetika MeSH
- typy dědičnosti MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
New species are generated by many means, among which hybridization plays an important role. Interspecific hybrids can form isolated evolutionary units, especially when mechanisms increasing viability and fertility, like polyploidy and apomixis, are involved. A good model system to study reticulate evolution in plants is Rubus subgen. Rubus (brambles, blackberries), which only in Europe includes 748 accepted species, out of which only four are sexual diploids and all others are polyploid apomicts. We employed two molecular markers (ITS and cpDNA) to shed light on the evolutionary history of European bramble flora and main processes generating such high species diversity. We distinguished just six ancestral diploids (including two extinct ones) for both markers, which gave rise to all European polyploid accessions, and revealed an extreme reticulation in bramble evolution. We furthermore detected hybridogenous origins and identified putative parents for several taxa (e.g. ser. Nessenses), while in other groups (e.g. ser. Discolores) we could also infer the direction of hybridization. By comparing different cp haplotypes having clear geographic patterns, we hypothesize that the origin of European brambles can be attributed to both Holocene species range expansion and Pleistocene climate fluctuations.
- MeSH
- apomixie * genetika MeSH
- diploidie MeSH
- DNA chloroplastová genetika MeSH
- fylogeneze MeSH
- genetické markery genetika MeSH
- hybridizace genetická genetika MeSH
- ledový příkrov MeSH
- mezerníky ribozomální DNA genetika MeSH
- modely genetické * MeSH
- molekulární evoluce * MeSH
- polyploidie MeSH
- Rubus klasifikace genetika MeSH
- sekvenční analýza DNA MeSH
- vznik druhů (genetika) * MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
- Geografické názvy
- Evropa MeSH
Neither the genetic basis nor the inheritance of apomixis is fully understood in plants. The present study is focused on the inheritance of parthenogenesis, one of the basic elements of apomixis, in Pilosella (Asteraceae). A complex pattern of inheritance was recorded in the segregating F(1) progeny recovered from reciprocal crosses between the facultatively apomictic hexaploid P. rubra and the sexual tetraploid P. officinarum. Although both female and male reduced gametes of P. rubra transmitted parthenogenesis at the same rate in the reciprocal crosses, the resulting segregating F(1) progeny inherited parthenogenesis at different rates. The actual transmission rates of parthenogenesis were significantly correlated with the mode of origin of the respective F(1) progeny class. The inheritance of parthenogenesis was significantly reduced in F(1) n + n hybrid progeny from the cross where parthenogenesis was transmitted by female gametes. In F(1) n + 0 polyhaploid progeny from the same cross, however, the transmission rate of parthenogenesis was high; all fertile polyhaploids were parthenogenetic. It appeared that reduced female gametes transmitting parthenogenesis preferentially developed parthenogenetically and only rarely were fertilized in P. rubra. The fact that the determinant for parthenogenesis acts gametophytically in Pilosella and the precocious embryogenesis in parthenogenesis-transmitting megagametophytes was suggested as the most probable explanations for this observation. Furthermore, we observed the different expression of complete apomixis in the non-segregating F(1) 2n + n hybrids as compared to their apomictic maternal parent P. rubra. We suggest that this difference is a result of unspecified interactions between the parental genomes.
- MeSH
- apomixie genetika fyziologie MeSH
- Asteraceae genetika fyziologie MeSH
- chromozomy rostlin genetika MeSH
- genom rostlinný genetika MeSH
- hybridizace genetická MeSH
- partenogeneze genetika fyziologie MeSH
- polyploidie MeSH
- průtoková cytometrie MeSH
- rozmnožování MeSH
- typy dědičnosti MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
Rubus subgen. Rubus includes common European species with highly complicated taxonomy, ongoing hybridisation and facultative apomixis. Out of approximately 750 species recognised in Europe, only 3 diploid sexual species are known, along with numerous apomictic brambles that are highly connected to polyploidy. One exception of a tetraploid taxon is R. ser. Glandulosi, which is known for prevalent sexuality. This taxon highly hybridises with tetraploid members of R. ser. Discolores and leads to the origin of many hybridogenous populations and individuals. In this study, we verify reproduction modes in different diploid, triploid and tetraploid species of subgen. Rubus, with focus on taxa putatively involved in such hybridisation by applying flow cytometric seed screen analysis. We found 100 % sexuality of diploid species, whereas triploid species had obligate unreduced embryo sac development. In contrast, tetraploid plants had varying degrees of sexuality. Additionally, we discovered that R. bifrons has the ability to undergo a reproduction mode switch as a reaction to environmental conditions. These results provide insight into reproductive modes of European brambles and shed light on their reticulate evolution and speciation.
- MeSH
- apomixie MeSH
- délka genomu MeSH
- diploidie MeSH
- druhová specificita MeSH
- endosperm embryologie genetika fyziologie MeSH
- partenogeneze MeSH
- polyploidie * MeSH
- průtoková cytometrie MeSH
- Rosaceae embryologie genetika fyziologie MeSH
- rozmnožování MeSH
- semena rostlinná embryologie genetika fyziologie MeSH
- tetraploidie MeSH
- zeměpis MeSH
- Publikační typ
- časopisecké články MeSH
- práce podpořená grantem MeSH
