Herbert Hurka

Recent speciation associated with the evolution of selfing in Capsella

The evolution from outcrossing to predominant self-fertilization represents one of the most common transitions in flowering plant evolution. This shift in mating system is almost universally associated with the “selfing syndrome,” characterized by marked reduction in flower size and a breakdown of the morphological and genetic mechanisms that prevent self-fertilization. In general, the timescale in which these transitions occur, and the evolutionary dynamics associated with the evolution of the selfing syndrome are poorly known. We investigated the origin and evolution of selfing in the annual plant Capsella rubella from its self-incompatible, outcrossing progenitor Capsella grandiflora by characterizing multilocus patterns of DNA sequence variation at nuclear genes. We estimate that the transition to selfing and subsequent geographic expansion have taken place during the past 20,000 years. This transition was probably associated with a shift from stable equilibrium toward a near-complete population bottleneck causing a major reduction in effective population size. The timing and severe founder event support the hypothesis that selfing was favored during colonization as new habitats emerged after the last glaciation and the expansion of agriculture. These results suggest that natural selection for reproductive assurance can lead to major morphological evolution and speciation on relatively short evolutionary timescales.

Colonization history and introduction dynamics of Capsella bursa-pastoris (Brassicaceae) in North America: isozymes and quantitative traits

Multilocus isozyme genotypic composition for aspartate aminotransferase (AAT), leucine aminopeptidase (LAP) and glutamate dehydrogenase (GDH) was studied for Capsella in the source continent, Europe (9000 plants from 593 populations), and in the colonized continent, North America (2700 plants from 88 populations). North America was depauperate in the number of genotypes (by ≈ 50%), but in terms of frequencies, a few genotypes were common and shared by both continents. Although some, very rare, genotypes were, however, unique for North America, our data provided no evidence to indicate that the introduced gene pools were reconstructed on a multilocus genetic basis after introduction. Instead, they argued for a considerable number of independent introduction events. Geographical distribution patterns of multilocus genotypes in Europe and North America were pronounced and enabled us to trace the colonization history of Californian Capsella back to Spanish ancestral populations and those of temperate North America back to temperate European gene pools. A random‐block field experiment with 14 Californian populations from different climatic regions revealed that variation patterns of quantitative traits reflect ecotypic variation, and the ecological amplitude of Capsella in North America is similar to that in Europe, which can be traced back to the introduction of preadapted genotypes. It appears that certain multilocus isozyme genotypes are associated with certain ecotypes. The variable European gene pool of Capsella was essentially introduced into North America without major genetic changes.

Hybridization and evolution inCardamine (Brassicaceae) at Urnerboden, Central Switzerland: Biosystematic and molecular evidence

Hybridization between two diploid (2n = 2x = 16) species ofBrassicaceae, Cardamine rivularis andC. amara, at Urnerboden, Central Switzerland, resulted in the rather unusual triploid hybridC. insueta (2n = 3x = 24), and later on in the amphiploidC. schulzii (2n = 6x = 48). The hybrid and the neopolyploid species colonized successfully some man-made biotopes. Plants ofC. insueta are mostly functional females with non-dehiscent anthers, but true hermaphrodite individuals with partly sterile pollen grains also occur within the population. Analyses of cpDNA and nuclear DNA permitted to establish the parentage of the hybrid: the maternal parent which contributed unreduced egg cells proved to beC. rivularis whereas the normally reduced pollen originated fromC. amara. The pronounced genetic variability inC. insueta revealed by isozyme and RAPD analyses, at variance with the polarized segregation, heterogamy and strong vegetative reproduction of the hybrid, is possibly influenced by recurrent formation ofC. insueta which party results from backcrosses betweenC. insueta andC. rivularis but may also proceed by other pathways. The amphiploidCardamine schulzii has normally developed anthers but its pollen is sometimes highly sterile. The surprisingly uniform genetic make-up of the new amphiploid species might be related to its possible monotopic origin and/or young phylogenetic age but should be further assessed. Site management seems to be very important to a further development of hybridogenous populations and their parent species. In conclusion, the evolution at Urnerboden is discussed in the context of the traditional concept of multiple plant origins.

Contribution to ITS phylogeny of the Brassicaceae, with special reference to some Asian taxa

Sequence data from the nuclear ribosomal internal transcribed spacer (ITS) region for 189 accessions representing 184 species in 121 genera of Brassicaceae were used to determine monophyly of tribes and genera, tribal boundaries, and component genera. Parsimony analysis and Bayesian inference suggest that the tribes Camelineae and Arabideae are polyphyletic and should be subdivided into smaller tribes. The study also supports the recent recognition of the new tribes Aphragmeae, Biscutelleae, Buniadeae, Calepineae, Conringieae, Dontostemoneae, Erysimeae, Malcolmieae, Megacarpaeeae, and Turritideae. The data argue for the placement of Borodinia in the tribe Boechereae, Litwinowia and Pseudoclausia in the Chorisporeae, Atelanthera and Streptoloma in the Euclidieae, and Megacarpaea and Pugionium in the Megacarpaeeae, and exclusion of Asperuginoides, Didymophysa, and Ptilotrichum from the Alysseae, Macropodium, Pseudoturritis, and Stevenia from the Arabideae, and Crucihimalaya, Irenepharsus, Pachycladon, and Turritis from the Camelineae. Finally, the findings support the expansion of Stevenia to include both Berteroella and Ptilotrichum, Sterigmostemum to include Oreoloma and one species of Anchonium, Crucihimalaya to include Transberingia and several species of Arabis, and Parrya to include Pseudoclausia. The data also suggest that Calymmatium and Olimarabidopsis may be congeneric.

Evolutionary processes in the genusCapsella (Brassicaceae)

Capsella comprises diploid (C. grandiflora andC. rubella) and tetraploid taxa. It is argued that the tetraploidC. bursa-pastoris is of intraspecific origin despite disomic inheritance and “fixed heterozygosity”. It is of considerable age as evidenced by the fossil record and molecular data. Gene duplication by polyploidization and a mixed mating system provided the genetic flexibility for greatest colonizing success. Pronounced variation patterns at a micro- and macrogeographic scale are observed inC. bursa-pastoris for many characters including life history traits, leaf morphology and allozymes. This variation pattern can be explained by several components comprising phylogenetic age, random processes, ecotypic variation and colonization history. The adaptive strategy ofC. bursa-pastoris cannot be assigned to either ecotypic differentiation or phenotypic plasticity alone. It depends on the trait under study.

Systematics and evolutionary history of heavy metal tolerant Thlaspi caerulescens in Western Europe - evidence from genetic studies based on isozyme analysis

Abstract Thlaspi caerulescens is distributed in Europe on metalliferous and not metalliferous soils. Individuals from populations growing on heavy metal contaminated soils are well known as hyperaccumulators of zinc and cadmium. The taxonomical treatment of subspecies of Thlaspi caerulescens is unsettled. We investigated the degree of genetic variation among 28 populations of Thlaspi caerulescens from Europe with isozyme analysis to compare inter- and intrapopulational diversity. British material from heavy metal contaminated environments recognized as Thlaspi sylvestre and T. occitanicum are quite similar to each other on the level of isozyme polymophisms, but they are more closely related to populations from non-contaminated stands from Scandinavia and Middle Europe than to metallophytes distributed in Continental Europe. Our findings indicate that a taxonomical subdivision of T. caerulescens is not possible and, furthermore, heavy metal tolerance might have evolved twice in populations of Thlaspi caerulescens from different areas. The trait of zinc tolerance and hyperaccumulation is frequently found in numerous relatives of Thlaspi caerulescens, and it is suggested that this trait has been established and manifested in populations from metalliferous sites during postglacial colonization. From Scandinavia only non-metallophytes are known. These populations are very similar to each other on the isozyme level. This fits to the hypothesis that Thlaspi caerulescens was introduced to Scandinavia in recent times by human activity. Despite full self-compatibility we estimated varying outcrossing rates up to 0.88 in the metallophytes and 0.658 in the non-metallophytes depending on population size and structure.

Molecular systematics ofCardamine and allied genera (Brassicaceae): Its and non-coding chloroplast DNA

Representatives of the generaCardamine, Dentaria, Nasturtium, Rorippa andArmoracia (Brassicaceae) were analyzed to elucidate their phylogenetic relationships based on nuclear (ITS) and non-coding chloroplast (cp) DNA sequences.Dentaria seems to be polyphyletic. The two studiedDentaria species group with differentCardamine clades, and it is argued thatD. bulbifera is an allopolyploid originating from a hybridization between aCardamine and aDentaria species. In the ITS tree,Nasturtium andRorippa form well supported clades but their relationship toCardamine andArmoracia remains unresolved. In the cpDNA tree,Nasturtium groups together withCardamine. Hybridization events apparently played a role in the evolution ofNasturtium. TheCardamine/Nasturtium clade is separated from a clade placingRorippa andArmoracia together.Armoracia is closely related toRorippa. Analyses of the 19Cardamine species studied revealed three main groupings, a northern hemispheric and two southern hemispheric groups. Within the northern hemisphere taxa theC. pratensis complex forms a well supported clade which seems to be closely related toC. amara, C. raphanifolia andC. flexuosa. The positions ofC. hirsuta andC. impatiens are uncertain. The two southern hemisphere clades consist of New Guinean species and south-eastern Australian/Tasmanian and subantarctic species, respectively. They may reflect migration routes from the northern to the southern hemisphere, but further studies are necessary to fully understand the evolution of the bihemispheric distribution pattern ofCardamine.

Variation of development time until flowering in natural populations ofCapsella bursa-pastoris (Cruciferae)

Seed samples were collected from wild populations ofCapsella bursa-pastoris along a transsect from Northern to Southern Europe. Progeny was grown in (a) open-field random block experiments (47 populations) and (b) in growth chambers under five to seven controlled temperature regimes (18 populations). Beginning of flowering was recorded, and great differences between and also within populations are documented. Some populations are extremely heterogenous whereas others are homogenous in this respect. Some biotypes react positively when exposed to lower temperatures, others are inhibited. In many cases specific effects of day- and/or night-temperatures can be inferred. In some progenies begin of flowering is independent of temperature as long as this exceeds the 5:10°C regimen. Altogether,Capsella bursa-pastoris displays definite intraspecific variation in time required until flowering. Adaptations to local ecological conditions are obvious. In addition to a genotypic component pronounced environmental interactions provide the plants with a component of phenotypic plasticity. The degree of modificability apparently varies itself and seems to be controlled by selection; the phenotypic plasticity, therefore, displays adaptive variation patterns, too.

Molecular phylogenetics of Cochlearia (Brassicaceae) and allied genera based on nuclear ribosomal ITS DNA sequence analysis contradict traditional concepts of their evolutionary relationship

The systematics and phylogeny of the genusCochlearia and allied genera are unsettled. There are no clearly defined genera and subtribal structures to determine subtribeCochleariinae in respect to subtribeThlaspidinae. The use of morphological data, such as fruit form or embryo characters have resulted in contradictory taxonomic concepts in the past due to their homoplastic nature. We investigated all sections of genusCochlearia recognised in the most common concepts, as well as some genera such asIonopsidium, Bivonaea, Pastorea andThlaspi s. l. pro parte. Previous studies based on molecular data and morphological studies have shown close relationships between taxa from subtribeCochleariinae andThlaspidinae. The Internal Transcribed Spacer regions of the nuclear encoded ribosomal DNA operon and the plastidictrnL intron were sequenced from a number of genera. A molecular phylogeny was derived and compared to traditional classification systems. These data grouped sections ofCochlearia outside theCochlearial Ionopsidium core group and integrated them either closely to genusNoccaea in subtribeThlaspidinae (sect.Pseudosempervivum) or positioned them outside both theCochlearia core group and theThlaspi s. l. clade (sect.Hilliella). The molecular data indicate that subtribal arrangements in tribeLepidieae are artificial and do not reflect evolutionary history. The genusCochlearia is represented by sectionsCochlearia andGlaucocochlearia and the genusIonopsidium should be integrated intoCochlearia.

Intrageneric phylogeny of Capsella (Brassicaceae) and the origin of the tetraploid C. bursa-pastoris based on chloroplast and nuclear DNA sequences

Polyploidization, often accompanied by hybridization, has been of major importance in flowering plant evolution. Here we investigate the importance of these processes for the evolution of the tetraploid crucifer Capsella bursa-pastoris using DNA sequences from two chloroplast loci as well as from three nuclear low-copy genes. The near-absence of variation at the C. bursa-pastoris chloroplast markers suggests a single and recent origin of the tetraploid. However, despite supporting a single phylogeny, chloroplast data indicate that neither of the extant Capsella diploids is the maternal parent of the tetraploid. Combined with data from the three nuclear loci, our results do not lend support to previous hypotheses on the origin of C. bursa-pastoris as an allopolyploid between the diploids C. grandiflora and C. rubella or an autopolyploid of C. grandiflora. Nevertheless, for each locus, some of the C. bursa-pastoris accessions harbored C. rubella alleles, indicating that C. rubella contributed to the gene pool of C. bursa-pastoris, either through allopolyploid speciation or, more likely, through hybridization and introgression. To our knowledge, this study is the first of a wild, nonmodel plant genus that uses a combination of chloroplast and multiple low-copy nuclear loci for phylogenetic inference of polyploid evolution.