DYNAMICS OF POLYPLOID FORMATION IN TRAGOPOGON (ASTERACEAE): RECURRENT FORMATION,GENE FLOW,AND POPULATION STRUCTURE

Polyploidy is a major feature of angiosperm evolution and diversification. Most polyploid species have formed multiple times, yet we know little about the genetic consequences of recurrent formations. Among the clearest examples of recurrent polyploidy are Tragopogon mirus and T. miscellus (Asteraceae), each of which has formed repeatedly in the last ～80 years from known diploid progenitors in western North America. Here, we apply progenitor‐specific microsatellite markers to examine the genetic contributions to each tetraploid species and to assess gene flow among populations of independent formation. These data provide fine‐scale resolution of independent origins for both polyploid species. Importantly, multiple origins have resulted in considerable genetic variation within both polyploid species; however, the patterns of variation detected in the polyploids contrast with those observed in extant populations of the diploid progenitors. The genotypes detected in the two polyploid species appear to represent a snapshot of historical population structure in the diploid progenitors, rather than modern diploid genotypes. Our data also indicate a lack of gene flow among polyploid plants of independent origin, even when they co‐occur, suggesting potential reproductive barriers among separate lineages in both polyploid species.     

Leaf flavonoids of the Achillea Millefolium group part II: Distribution patterns of free aglycones in leaf exudates

Seventy-eight collections of species forming the Achillea millefolium group were studied for the aglycone composition of their leaf exudates. 6-Hydroxyflavones, 6-hydroxyflavonols and their methyl ethers predominate in the exudates in various combinations. Within the polyploid complex, the diploid taxa Achillea setacea and Achillea aspleniifolia appeared to be chemically well defined. Among the polyploids, little differentiation of major accumulation trends was observed. Infraspecific variability was low in most cases, thus allowing systematic interpretations of aglycone patterns. Possible pharmacological activities are briefly discussed.     

Multiple independent formations of Tragopogon tetraploids (Asteraceae): evidence from RAPD markers

Polyploidy is widely recognized as a significant force leading to the formation of new plant species. Estimates of the number of angiosperm species with polyploid origins are as high as ≈ 50%; however, in spite of this prevalence, many aspects of polyploid evolution remain poorly understood. Recent studies have suggested that recurrent origins of polyploid species are the rule rather than the exception. The present study is one of only a few designed to quantify the number of independent origins of a polyploid species. The two tetraploid species Tragopogon mirus and T. miscellus (Asteraceae) arose within the past 50 years in the Palouse region of eastern Washington and adjacent northern Idaho. Previous work using morphology, cpDNA and rDNA restriction site analyses, allozymes, cytology, and flavonoid chemistry established that T. mirus had arisen at least five times, and T. miscellus at least twice, on the Palouse. To assess the frequency of multiple origins of these species more rigorously, seven populations of T. mirus and three populations of T. miscellus that were indistinguishable based on previous markers were surveyed using random amplified polymorphic DNA (RAPD) markers; populations of the diploid progenitor species from the same sites were also analysed. Each tetraploid population had a unique RAPD marker profile, suggesting that each population surveyed originated independently of the other populations in the region. Only two of the tetraploid populations combined the RAPD marker profiles of the diploid progenitors occurring at the same site. Both polyploid species, whose ranges and numbers have greatly increased since their formation in the early part of the twentieth century, have formed repeatedly on a local geographical scale and during a short time frame. Furthermore, each tetraploid species is spreading not primarily by dispersal of propagules from a single population of origin, but through repeated, independent polyploidization events that recreate the polyploid taxa.     

Independent origins of tetraploid cryptic species in the fern <Emphasis Type="Italic">Ceratopteris thalictroides</Emphasis>

Ceratopteris thalictroides (L.) Brongn is a tetraploid fern species that contains at least three cryptic species, the south, the north and the third type. In this study we combined data from both chloroplast DNA (cpDNA) and nuclear DNA sequences of three diploid species and three cryptic species of C. thalictroides to unravel the origin of the cryptic species, particularly of the reticulate relationships among the diploid and tetraploid taxa in the genus Ceratopteris. Of the three diploid species examined, C. cornuta had cpDNA identical to that of the tetraploid third type plants, and this diploid species is a possible maternal ancestor of the tetraploid third type. Analysis of the homologue of the Arabidopsis thaliana LEAFY gene (CLFY1) identified ten alleles in the genus Ceratopteris, with six alleles found in C. thalictroides. The unrooted tree of the CLFY1 gene revealed four clusters. Each cryptic species showed fixed heterozygosity at the CLFY1 locus and had two alleles from different clusters of the CLFY1 tree. Consideration of the cpDNA sequences, CLFY1 genotypes of the cryptic species and CLFY1 gene tree in concert suggested that the cryptic species of C. thalictroides had originated through independent allopolyploidization events involving C. cornuta and two unknown hypothetical diploid species.     

On the embryology and mode of reproduction of selected diploid species of Hieracium s.l. (Asteraceae) from Bulgaria

Three diploid taxa – Hieracium transylvanicum (subgenus Hieracium), Hieracium caespitosum subsp. brevipilum and Hieracium pavichii (subgenus Pilosella) – from five natural Bulgarian populations were investigated embryologically. The peculiarities of the male and female gametophytes, embryo- and endospermogenesis were established in each species. The results suggest that the species propagate sexually as expected from their diploid chromosome number. However, some forms of apomixis have also been observed, e.g. somatic apospory and integumental embryony in H. pavichii. The presence of apomixis in a diploid taxon shows that polyploidy is not an obligatory prerequisite for apomixis and the two phenomena are independent. The embryological plasticity detected in H. pavichii and H. transylvanicum suggests they may have higher opportunities for adaptation and successful reproduction.     

Shaping polyploid wheat for success: Origins,domestication, and the genetic improvement of agronomic traits

Bread wheat (Triticum aestivum L., AABBDD, 2n = 6x = 42), which accounts for most of the cultivated wheat crop worldwide, is a typical allohexaploid with a genome derived from three diploid wild ancestors. Bread wheat arose and evolved via two sequential allopolyploidization events and was further polished through multiple steps of domestication. Today, cultivated allohexaploid bread wheat has numerous advantageous traits, including adaptive plasticity, favorable yield traits, and extended end-use quality, which have enabled its cultivation well beyond the ranges of its tetraploid and diploid progenitors to become a global staple food crop. In the past decade, rapid advances in wheat genomic research have considerably accelerated our understanding of the bases for the shaping of complex agronomic traits in this polyploid crop. Here, we summarize recent advances in characterizing major genetic factors underlying the origin, evolution, and improvement of polyploid wheats. We end with a brief discussion of the future prospects for the design of gene cloning strategies and modern wheat breeding.     

Ancient allopolyploid speciation in Geinae (Rosaceae): evidence from nuclear granule-bound starch synthase (GBSSI) gene sequences

A nuclear low-copy gene phylogeny provides strong evidence for the hybrid origin of seven polyploid species in Geinae (Rosaceae). In a gene tree, alleles at homologous loci in an allopolyploid species are expected to be sisters to orthologues in the ancestral taxa rather than to each other. Alleles at a duplicated locus in an autopolyploid, however, are expected to be more closely related to each other than they are to any orthologous copies in closely related species. We cloned and sequenced about 1.9 kilobases from the 5' end of the GBSSI-1 gene from two diploid, one tetraploid, and six hexaploid species. Each of the three loci in the hexaploid species forms a separate group, two of which are more closely related to copies in other species than they are to each other. This finding indicates that the hexaploid lineage evolved through two consecutive allopolyploidization events. Based on the GBSSI-1 gene tree, we hypothesized that there was an initial hybridization between a diploid species from the ancestral lineage of Coluria and Waldsteinia and an unknown diploid species to form the tetraploid Geum heterocarpum lineage. Backcrossing of G. heterocarpum with a representative of the unknown diploid lineage then resulted in a hexaploid lineage that has radiated considerably since its origin, comprising at least 40 extant species with various morphologies. A penalized likelihood analysis indicated that Geinae may be about 17 million years old, implying that the hypothesized allopolyploid speciation events are relatively ancient. Six of the 22 cloned Geinae GBSSI-1 copies in this study, which all are duplicate copies in polyploid taxa, may have become pseudogenes. We compared the GBSSI-1 phylogeny with one from chloroplast data and explored implications for the evolution of some fruit characters.     

SYSTEMATIC INFERENCES FROM VARIATION IN ISOZYME PROFILES OF ARCTIC AND ALPINE CESPITOSE FESTUCA (POACEAE)

Variation in isozyme patterns was used to assess species boundaries in North American arctic and alpine representatives of the Festuca ovina L. complex. Isozyme profiles, in combination with chromosome number, delimit four discrete entities within the complex: F. brevissima Jurtzev (diploid); F. aggr. auriculata Drobov (diploid); F. brachyphylla Schultes (hexaploid); and tetraploid populations corresponding in morphology to F. baffinensis Polunin (arctic Canada) and F. minutiflora Rydberg (alpine United States). Although no fixed difference was detected between isozyme profiles of the latter two taxa, they are morphologically distinct. Thus variation in isozymes, morphology, and chromosome number delimits five taxa within the F. ovina complex in North America. Some alleles observed in the polyploid taxa were not detected among the diploids, and some observed in F. brachyphylla, the hexaploid taxon, were not detected in either the diploid or the tetraploid species. One possible explanation for these occurrences is that the North American polyploids originated in Eurasia, where many other potential diploid and tetraploid progenitors occur.     

Genetic diversity in the tetraploid sand dune endemic Deschampsia mackenzieana and its widespread iploid progenitor D. Cespitosa (Poaceae)

Electrophoretic variation was examined in 14 populations of tetraploid Deschampsia mackenzieana, an endemic of the Athabasca sand dunes in northern Saskatchewan, Canada, and 20 populations of its geographically widespread diploid progenitor, D. cespitosa. Three of the D. cespitosa populations were sympatric with the endemic on the Athabasca sand dunes. Populations of the endemic were found to have fewer alleles per locus (1.22 vs. 1.52), fewer alleles per polymorphic locus (2.17 vs. 2.70), lower percent polymorphic loci (18.9 vs. 30.5), and lower heterozygosity (0.062 vs. 0.119) than progenitor populations. Species level genetic diversity parameters also indicated that D. mackenzieana was genetically depauperate relative to its progenitor D. cespitosa. Deschampsia mackenzieana had no novel alleles but did share one allele with sympatric progenitor populations that did not occur in populations of D. cespitosa from other habitats. Although both species were found to partition most of their genetic diversity within populations, D. mackenzieana did have more of its limited genetic diversity partitioned among populations than D. cespitosa. The close genetic relationship between D. mackenzieana and sympatric populations of D. cespitosa may suggest the endemic tetraploid evolved from the sympatric diploid gene pool in the Athabasca sand dune region. The low levels of genetic diversity in D. mackenzieana suggest a restricted origin with limited gene flow from the progenitor since speciation.     

Variation among tripsacum populations of Mexico and Guatemala

Seven species of the genusTripsacum, one with a well marked subspecies, are indigenous to Mexico and Guatemala. Two diploids,T. zopilotense andT. maizar, are restricted to western and southwestern Mexico. The typical diploid form of a third species,T. latifolium, occurs in south central and eastern Guatemala; atypical tetraploid forms of this species are found in western Mexico. Four tetraploids,T. dactyloides, T. dactyloides ssp.hispidum, T. lanceolatum, andT. pilosum, have ranges extending from northwestern Mexico southeastward into Guatemala. The seventh species is the highly sterile tetraploid,T. laxum, of doubtful occurrence in the wild. It is readily propagated vegetatively and has been widely distributed in Latin America and the West Indies as a forage plant. Field studies of the range of variation in 80 Mexican and Guatemalan populations included an evaluation of 16 definitive morphological characteristics from an average of approximately six individuals of each population. Seventy-three of the 80 populations were allopatric, and of these, 50 were classed as typical, 13 as atypical, and ten as intermediates. Thirty-nine of the 50 populations classed as typical were tetraploid, and 11 were diploid. Of the 23 atypical and intermediate populations, 17 were tetraploid and six were diploid. Fifty-eight tetraploid populations, which included typical, atypical, and intermediate forms ofT. dactyloides, T. dactyloides ssp.hispidum, T. lanceolatum, T. pilosum, andT. latifolium, comprised an inclusive intergrading series having different combinations of characteristics distinguishing the two very dissimilar diploids,T. mizar andT. zopilotense. This series was interpreted as support for the hypothesis of Randolph & Hernández (1950), that the tetraploid populations originated as alloploid derivatives of these or closely related diploid species. The occurrence in some of the tetraploid populations of characteristics not seen in either of the putative parental species was noted, and the possibility that other taxa also might have been involved in their origin is under investigation. In addition to the 73 allopatric populations included in this survey there were seven sympatric and mixed associations of diploid and tetraploid taxa. Among these were hybrids with various combinations of the characteristics differentiating taxa of the associated populations. These hybrids effectively obscured differences between taxa, thus creating a syngameon complex from which genetic recombinants were observed to have spread into recently disturbed habitats of neighboring areas. From such direct evidence, and the existing interrelationships among allopatric and sympatric populations, it was concluded that the origin of the Mexican and Guatemalan tetraploidTripsacum species and various atypical and intermediate variants, has involved, and is continuing to involve, alloploid recombinants of diploid species and syngameon complexes, of which those encountered in this survey are examples.     

A novel method to infer the origin of polyploids from Amplified Fragment Length Polymorphism data reveals that the alpine polyploid complex of Senecio carniolicus (Asteraceae) evolved mainly via autopolyploidy

Despite its evolutionary and ecological relevance, the mode of polyploid origin has been notoriously difficult to be reconstructed from molecular data. Here, we present a method to identify the putative parents of polyploids and thus to infer the mode of their origin (auto‐ vs. allopolyploidy) from Amplified Fragment Length Polymorphism (AFLP) data. To this end, we use Cohen's d of distances between in silico polyploids, generated within a priori defined scenarios of origin from a priori delimited putative parental entities (e.g. taxa, genetic lineages), and natural polyploids. Simulations show that the discriminatory power of the proposed method increases mainly with increasing divergence between the lower‐ploid putative ancestors and less so with increasing delay of polyploidization relative to the time of divergence. We apply the new method to the Senecio carniolicus aggregate, distributed in the European Alps and comprising two diploid, one tetraploid and one hexaploid species. In the eastern part of its distribution, the S. carniolicus aggregate was inferred to comprise an autopolyploid series, whereas for western populations of the tetraploid species, an allopolyploid origin involving the two diploid species was the most likely scenario. Although this suggests that the tetraploid species has two independent origins, other evidence (ribotype distribution, morphology) is consistent with the hypothesis of an autopolyploid origin with subsequent introgression by the second diploid species. Altogether, identifying the best among alternative scenarios using Cohen's d can be straightforward, but particular scenarios, such as allopolyploid origin vs. autopolyploid origin with subsequent introgression, remain difficult to be distinguished.     

Correlation between distyly and ploidy level in Damnacanthus (Rubiaceae)

Somatic chromosomes were observed in 661 individuals of 14 taxa, nine species and five varieties, of Damnacanthus (Rubiaceae). Chromosome numbers are reported for the first time for 13 taxa. Diploid (2n = 22) and tetraploid (2n = 44) counts were obtained. Distyly is reported for the first time for four species, D. angustifolius, D. henryi, D. labordei, and D. officinarum. A strong correlation exists between chromosome number and occurrence of distyly. Regardless of taxa in Damnacanthus, distylous populations are diploid, and monomorphic populations are tetraploid. Flowers of the monomorphic populations observed have a long style and short stamens with few exceptions. Polyploidization may have caused the breakdown of distylous to monomorphic flowers. In D. indicus, leaves from the tetraploid populations tend to be larger than those from the diploid populations. Populations of tetraploid D. indicus were distributed in more northern areas than those of the diploid. Three types of sympatric distribution were found for the varieties of D. indicus in Japan: diploid and tetraploid, two diploids, and two tetraploids. Based on the present chromosome number study, the taxonomy of the varieties of D. indicus should be revised.     

Integrated taxonomy of the <Emphasis Type="Italic">Asplenium normale</Emphasis> complex (Aspleniaceae) in China and adjacent areas

The Asplenium normale D. Don complex comprises several taxa that are either diploid or tetraploid. The tetraploids are assumed to have originated from diploid ancestors by relatively recent autopolyploidization or allopolyploidization. Some of the diploids are readily recognized morphologically but most of the taxa have until now been placed into a single species. However, phylogenetic studies have challenged this treatment and emphasized the notion that the taxonomic treatment of this complex needs to be revised. An integrative taxonomic approach was employed to delimit species in the complex using cytological, morphological, and DNA sequence data. Initially, we employed a diploid first approach to establish a robust taxonomic framework. Special efforts were made to collect and identify the diploid progenitors of each polyploid lineage identified in the plastid DNA based phylogenetic hypothesis. A total of six distinct diploid species were identified. The distinctive nature of the six diploids is strongly supported by sequence differences in plastid DNA and nuclear loci, as well as by the results of morphometric analysis. Diagnostic morphological characters were identified to distinguish the six diploid species, resulting in their revised taxonomy, which includes two novel species, namely, Asplenium normaloides and A. guangdongense. Further studies to strengthen the taxonomic classification of all of the tetraploid taxa are warranted.     

Using principle component analysis to compare genetic diversity across polyploidy levels within plant complexes: an example from British Restharrows (Ononis spinosa and Ononis repens)

The investigation of genetic diversity between related plant populations which differ in ploidy levels is problematic, with common statistical methods developed for diploids being inappropriate for polyploid species. Studies into gene flow in such complexes are critical and can shed light on the mechanisms that generate and maintain populations of different polyploidy levels. We have investigated the use of principle component (PCO) analysis as one approach to elucidate population structure within British Restharrows (Leguminsoae, Ononis spp). Restharrows were common agricultural weed species until the advent of mechanical ploughing and both diploid (2n=2x=30; O. spinosa and O. intermedia) and tetraploid (2n=4x=60; O. repens and O. maritima) taxa exist. Patterns of genetic diversity were investigated among British Restharrows using 10 microsatellite loci with 21 Restharrow populations analysed (411 individual plants) from Central and Eastern Britain. PCO analysis revealed clear genetic differentiation of the sampled plants into two groups, corresponding to O. spinosa/O. intermedia (diploid) and O. repens/O. maritima (tetraploid) plants. Evidence of genetic differentiation by distance was also revealed for O. repens/O. maritima, but not for O. spinosa/O. intermedia. The data suggest the presence of strong reproductive barriers between diploid and tetraploid Restharrows in Britain, but not within ploidy levels. This genetic isolation between ploidy levels is confirmed by a detailed analysis of a sympatric site (Harton Down Hill). These results demonstrate that PCA analysis is a suitable general tool for comparing related species of different ploidy levels.     

Plastid DNA variation in the Dactylorhiza incarnata/maculata polyploid complex and the origin of allotetraploid D. sphagnicola (Orchidaceae)

To obtain further information on the polyploid dynamics of the the Dactylorhiza incarnata/maculata polyploid complex and the origin of the allotetraploid D. sphagnicola (Orchidaceae), plastid DNA variation was studied in 400 plants from from Sweden and elsewhere in Europe and Asia Minor by means of polymerase chain reaction-restriction fragment length polymorphisms (PCR-RFLPs) and sequencing. Allotetraploid taxa in Europe are known have evolved by multiple independent polyploidization events following hybridization between the same set of two distinct ancestral lineages. Most allotetraploids have inherited the plastid genome from parents similar to D. maculata sensu lato, which includes, e.g. the diploid D. fuchsii and the autotetraploid D. maculata sensu stricto. D. sphagnicola carries a separate plastid haplotype different from the one found in other allotetraploid taxa, which is in agreement with an independent origin from the parental lineages. Some of the remaining allotetraploids have local distributions and appear to be of postglacial origin, whereas still other allotetraploids may be of higher age, carrying plastid haplotypes that have not been encountered in present day representatives of the parental lineages. Introgression and hybridization between diploids and allotetraploids, and between different independently derived allotetraploids may further have contributed to genetic diversity at the tetraploid level. Overall, the Dactylorhiza polyploid complex illustrates how taxon diversity and genetic diversity may be replenished rapidly in a recently glaciated area.     

Genetic diversity and population structure in Tunisian Lavandula stoechas L. and Lavandula multifida L. (Lamiaceae)

The genetic diversity and population structure of 20 Tunisian Lavandula stoechas L. and Lavandula multifida L. populations, from different bioclimates, were analysed by starch gel electrophoresis using seven isozymes. The genetic diversity within populations varied according to species. Variation in L. multifida was higher than that observed for L. stoechas, and exclusive alleles were detected for taxa.

A high differentiation among populations, for each species, estimated by Wright's F-statistics was revealed. The genetic structure of populations from the same bioclimate was substantial. Nei's, R. [1978. Estimation of average heterozygosity and genetic distance from a small number of individuals. Genetics 89, 583–590] genetic distance among pairs of populations was low. The UPGMA cluster analysis of genetic distance values revealed that populations for each species were not strictly clustered together according to bioclimate or geographic proximity.

For each species, the low genetic divergence among populations and their substantial structure indicate their recent fragmentation due to anthropic pressures. The dendrogram generated from pairwise genetic distance among all populations showed two distinct clusters each corresponding to one species. The high genetic divergence between the two species, based on isozymes, corroborates their taxonomic status, as previously reported using morphological traits. The strategy for the management and conservation of populations should be made for each taxa according to its level of diversity and bioclimate.     

Ancestry influences the fate of duplicated genes millions of years after polyploidization of clawed frogs (Xenopus)

Allopolyploid species form through the fusion of two differentiated genomes and, in the earliest stages of their evolution, essentially all genes in the nucleus are duplicated. Because unique mutations occur in each ancestor prior to allopolyploidization, duplicate genes in these species potentially are not interchangeable, and this could influence their genetic fates. This study explores evolution and expression of a simple duplicated complex--a heterodimer between RAG1 and RAG2 proteins in clawed frogs (Xenopus). Results demonstrate that copies of RAG1 degenerated in different polyploid species in a phylogenetically biased fashion, predominately in only one lineage of closely related paralogs. Surprisingly, as a result of an early deletion of one RAG2 paralog, it appears that in many species RAG1/RAG2 heterodimers are composed of proteins that were encoded by unlinked paralogs. If the tetraploid ancestor of extant species of Xenopus arose through allopolyploidization and if recombination between paralogs was rare, then the genes that encode functional RAG1 and RAG2 proteins in many polyploid species were each ultimately inherited from different diploid progenitors. These observations are consistent with the notion that ancestry can influence the fate of duplicate genes millions of years after duplication, and they uncover a dimension of natural selection in allopolyploid genomes that is distinct from other genetic phenomena associated with polyploidization or segmental duplication.     

GENETIC VARIATION IN TRAGOPOGON SPECIES: ADDITIONAL ORIGINS OF THE ALLOTETRAPLOIDS T. MIRUS AND T. MISCELLUS (COMPOSITAE)

Genetic diversity in the introduced diploids Tragopogon dubius, T. porrifolius, and T. pratensis and their neoallotetraploid derivatives T. mirus and T. miscellus was estimated to assess the numbers of recurrent, independent origins of the two tetraploid species in the Palouse region of eastern Washington and adjacent Idaho. These tetraploid species arose in this region, probably within the past 50–60 yr, and provide one of the best models for the study of polyploidy in plants. The parental species of both T. mirus and T. miscellus have been well documented, and each tetraploid species has apparently formed multiple times. However, a recent survey of the distributions of these allotetraploids revealed that both tetraploid species have expanded their ranges considerably during the past 50 yr, and several new populations of each species were discovered. Therefore, to evaluate the possibility that these recently discovered populations are of recent independent origin, a broad analysis of genetic diversity in T. mirus, T. miscellus, and their diploid progenitors was conducted. Analyses of allozymic and DNA restriction site variation in all known populations of T. mirus and T. miscellus in the Palouse and several populations of each parental diploid species revealed several distinct genotypes in each tetraploid species. Four isozymic multilocus genotypes were observed in T. mirus, and seven were detected in T. miscellus. Tragopogon mirus possesses a single chloroplast genome, that of T. porrifolius, and two distinct repeat types of the 18S-26S ribosomal RNA genes. Populations of T. miscellus from Pullman, Washington, have the chloroplast genome of T. dubius; all other populations of T. miscellus have the chloroplast DNA of T. pratensis. All populations of T. miscellus combine the ribosomal RNA repeat types of T. dubius and T. pratensis, as demonstrated previously. When all current and previously published data are considered, both T. mirus and T. miscellus appear to have formed numerous times even within the small geographic confines of the Palouse, with estimates of five to nine and two to 21 independent origins, respectively. Such recurrent polyploidization appears to characterize most polyploid plant species investigated to date (although this number is small) and may contribute to the genetic diversity and ultimate success of polyploid species.     

Breeding systems,hybridization and continuing evolution in Avon Gorge Sorbus

Background and Aims

Interspecific hybridization and polyploidy are key processes in plant evolution and are responsible for ongoing genetic diversification in the genus Sorbus (Rosaceae). The Avon Gorge, Bristol, UK, is a world ‘hotspot’ for Sorbus diversity and home to diploid sexual species and polyploid apomictic species. This research investigated how mating system variation, hybridization and polyploidy interact to generate this biological diversity.

Methods

Mating systems of diploid, triploid and tetraploid Sorbus taxa were analysed using pollen tube growth and seed set assays from controlled pollinations, and parent–offspring genotyping of progeny from open and manual pollinations.

Key Results

Diploid Sorbus are outcrossing and self-incompatible (SI). Triploid taxa are pseudogamous apomicts and genetically invariable, but because they also display self-incompatibility, apomictic seed set requires pollen from other Sorbus taxa – a phenomenon which offers direct opportunities for hybridization. In contrast tetraploid taxa are pseudogamous but self-compatible, so do not have the same obligate requirement for intertaxon pollination.

Conclusions

The mating inter-relationships among Avon Gorge Sorbus taxa are complex and are the driving force for hybridization and ongoing genetic diversification. In particular, the presence of self-incompatibility in triploid pseudogamous apomicts imposes a requirement for interspecific cross-pollination, thereby facilitating continuing diversification and evolution through rare sexual hybridization events. This is the first report of naturally occurring pseudogamous apomictic SI plant populations, and we suggest that interspecific pollination, in combination with a relaxed endosperm balance requirement, is the most likely route to the persistence of these populations. We propose that Avon Gorge Sorbus represents a model system for studying the establishment and persistence of SI apomicts in natural populations.